THE MECHANISM OF LIFE " Of philosophy I will say nothing, except that when I saw that it had been cultivated for many ages by the most distinguished men, and that yet there is not a single matter within its sphere which is not still in dispute, and nothing, therefore, which is above doubt, I did not presume to anticipate that my success would be greater in it than that of others; and, further, when I considered the number of conflicting opinions touching a single matter that may be upheld by learned men, while there can be but one true, I reckoned as well-nigh false all that was only probable." DESCARTES: The Discourse on Method. " When I wrote my paper on the thymus gland, I was very con- scientious about the literature on the subject. I found that many memoirs had been written and published, and I looked at them all or, at least, at all of them that I could obtain. There were many German works, not many French and Italian ones, and a number of English papers. I collated and made abstracts of them , and discussed all the results and conclusions, and, generally, rounded off our knowledge with regard to the matter. Altogether I found afterwards that there were fifty-two memoirs on the development of the gland. My paper only made the number fifty - thrv> !" Unpublished Letter from a Young Zoologist. THE MECHANISM OF LIFE IN RELATION TO MODERN PHYSICAL THEORY BY JAMES JOHNSTONE, D.Sc. PROFESSOR OF OCEANOGRAPHY IN THE UNIVERSITY OF LIVERPOOL LONGMANS, GREEN & CO LONDON: EDWARD ARNOLD IQ2I All rights res i- wed B1C PRINTED IN GREAT BRITAIN. PREFACE IT is possible that the title of this book may be misleading to some readers, and so an explanation may, very appropriately, form the subject of this introduction. Well, then, by " the mechanism of life " is meant nothing more than the results of a scientific analysis of the activities of living animals. First, we must define what is meant by " scientific method," and this is not at all difficult now that Einstein, Eddington, and the other relativists, have persuaded us to think about what we do when we investigate something " scientifically." What we do, in that case, is to observe space-time coincidences in a four- dimensional manifold that is really and actually our procedure, though it seems rather dreadful ! It would be very inconvenient, also, to sustain oneself in this plane all the while, and so we proceed to let ourselves down to earth, so to speak. From the space coincidences that we observe (for instance, the coincidences of the top of a column of mercury in a barometer tube with certain marks on the adjoining scale) we infer space measurements, and from the coincidences of the hands of a clock with marks on the dial we infer time measurements. That simplifies the method a good deal. Then it is only the relations between series of space-time measurements that form the data of science (its differential equations), but that, again, is very trying, and so we assume that there are things in nature. These things are separated from each other, at the same instant of time, by intervals of space, while they are separated from each other, in the same space, by intervals of time. Thus we have something to lean up against and sustain ourselves in this rather difficult process of apprehending nature. The things that we regard as existing apart from each other in space and time are electrons. But just yet that is rather inconvenient, and so we regard our natural things as atoms and molecules in motion in an arbitrary three dimensional space and an arbitrary one- dimensional time. V 460727- vi PREFACE Thus there are atoms and molecules which exist and move and form configurations that is, constitute physico-chemical systems in space and time. When we speak about a " mechan- ism," we mean the motions and configurations of material particles. Already we have gone a long way, via inferences, from our " real and actual " observations of the passage of nature, which observations are space- time coincidences; but never mind that : let us stick to our notion of mechanism systems of material particles and their motions and configurations. The descriptions of such systems by making use of space measure- ments, and the devising of mathematical relationships between them (the differential equations), are the method of science. What we call " space " may be measured in terms of x and y and z, the old space dimensions, and t the time one ; and so the equations that we make involve the four " variables," x, y, z, and t. That is what physiology does; whatever its particular methods may be, they involve the observations of space- time coincidences the readings of the pointers, scales, etc., of instruments. It observes systems of material particles (the atoms and molecules making up tissues) in certain configurations, and then, after intervals of time, in other configurations. Sometimes the differ- ences between the configurations can be thrown into mathe- matical forms, but more often they cannot. This, therefore, is what is called mechanism, and it is the method of physiology. It is the study of the successive phases of a material energetic configuration or system. Note that it is not necessarily the study of an organism. Usually what is investigated is a part of an organism, or even the dead material of the latter. And in all cases it is the study of the physico- chemical activities of the organism that is the object of physiology. In the very act of investigation these activities are necessarily dissociated from each other, and the result is a number of partial views of the whole organic activity. Of course, all this is in- dispensable, and so the greater part of this book is really a sum- mary of the main results of physiological science, and is intended to give the reader an attitude (for he must supplement what is said here) in his attempt to understand life. It would be inconvenient, and even pedantic, to state these results in terms of the fundamental space-time concepts, and so our analyses of the activities of the living organism must continue PREFACE vii to use the familiar ideas of atoms, molecules, colloids, chemical and physical states of equilibrium, energy-transformations, potentials, radiation, and so on. In the light of modern physical theory, however, most of these concepts are derived ones, and if we use them in speculations upon the nature of life, there may be some crudeness in our statements. Thus, quoting a very good modern statement as to the aims of biology * 1. " Scientific biology is strictly deterministic. It admits the possibility of only one result from a given set of antecedents." 2. " Scientific biology endeavours to explain organic pheno- mena on the basis of antecedent physical conditions, though admitting that our knowledge of cause and effect is in the last resort empirical, to the extent that much which happens could not have been predicted in advance." 3. " Scientific biology declares that vital phenomena are chemico-physical in the sense that they are the inevitable out- come of the particular material aggregations which we call organisms." Now whether our knowledge can be regarded as proving the above theses is the subject of the following chapters. We must be very clear as to what is meant by " determinism," " antecedent physical conditions," " cause and effect," and " particular material aggregations." We have really nothing to do with determinism because the concepts that we employ in discussing our results are those of functionality, correlation, and proba- bility. Determinism is a logical category, or perhaps convention, and it is strict only in mathematics, where, since we make the rules, the strictness of result is to be expected. We assume determinism because it is our mental postulate, and also because we find that it works more or less approximately in chemistry and physics, and even in physiology, so that we can construct and use machines and cure some oliseases. But it never works out exactly, and our results always have the form V e, where V is the value we adopt for something or other as the result of experiment, and e is a " probable error." W^e never get a unique biological result from a " given set of antecedents " : thus the bodily characters of an individual animal may surely be regarded as the result of the characters of its ancestry (which are the " antecedents " ; but this individual result is only one of * F. B. Sumner, The American Naturalist, vol. lii., 1919, pp. 193-217;. vol. liii., 1919, pp. 338-369 viii PREFACE a number of such (the combinations of bodily characters of the brothers and sisters of the individual), and all of these combina- tions differ from each other though they have the same antece- dents. Sometimes we say that the individual results " ought to be " the same if only we could experiment or observe with suffi- cient accuracy ; but in saying so, are we not simply dogmatising ? Further, it is said that " vital phenomena are chemico-physical in the sense that they are the inevitable outcome of the par- ticular material aggregations that we call organisms." But they are not " inevitable," and are they the outcome of " material aggregations " ? It is quite certain that it is not material aggre- gations that our method of science observes in nature, but rather space-time coincidences. We do not know about things, but only about relations which are differential equations between dx, dy, dz, and dt, which symbols are, after all, " ghosts " of space and time. No doubt it is very difficult to think in this way, and one naturally leans up against a mentally constructed world of atoms mathematics is so tiresome ! But when we would speculate about the nature of life and so on, we must not lean up against anything, and pur analysis should at least be as penetrating as that of the physicists. On the whole, one prefers the conclusions of some of the latter our knowledge of the world is a knowledge of form and not of content. We know relations only, and the unknown stuff of the world may just as likely be the stuff of our consciousness as something consisting of electrons. Anyhow, life is, after all, mainly an affair of organisms acting individually and as entire, undecomposable entities. It is mind, feeling, perception, memory, emotion, pleasure, pain, and so on. To the vast majority of men and women (to say nothing of all the " lower " animals) these states are life, and it would be very stupid not to recognise that in our philosophy. Tropisms and " concatenated reflexes " and colloids and enzymes, and so on, are all very well in their way, and we have to investigate them if we are to get on in the world, and be comfortable, and live long (quite legitimate objects of scientific research); but are these notions anything else than the terms in our description of how the living (or dead) organism cuts up, so to speak ? Is not this common sense ? If we do recognise that mind and intuition of living are to count in our speculations, what becomes of determinism (and PREFACE ix prediction, which surely goes with determinism if we cannot predict, why say that events are determined) ? For mind and memory and feeling and perception are certainly not measur- able in terms of space and time (despite the Weber-Fechner " psycho-physical law "). So the problems of free-will and necessity and determinism are meaningless when they are con- sidered with reference to the mind, for the very essence of these notions is measurement, and we cannot measure mind. Such, then, are the lines on which the phenomena of life are discussed in this book, and the reader is asked to take the arguments " on their merits," and without conscious clinging to either mechanism or vitalism. Vitalism holds that there is something in the living organism which is not present in an inorganic thing. This may be " spirit," or " soul," or perhaps some hitherto unrecognised " biotic energy- form," or some factor which is not energetic in nature, but which confers direction upon the energy-transformations that occur in the living organism. Most people take one or other of these attitudes; thus some may confess to sympathy with those who would " remove organisms from the domain which includes the stars and precious stones," but may not think that mechanism " exhausts the reality of earth and heavens, still less that of the flower in the crannied wall " ; others like to think that " the sun and moon and all the little stars are a glorious mechanism." Either feeling is, of course, permissible, provided it does not influence our judgment. Also, no one can think about these questions without becom- ing " metaphysical," even if he does not know it. There is no harm in that either, provided that one does it nicely. So I have taken care that anything " transcendental," or otherwise objec- tionable, has been discreetly relegated to the Appendices. J. J. LIVERPOOL, 1921. CONTENTS , CHAPTER I THE NATURE OF ANIMAL LIFE PAGE The organism as a structure Structural differences Structure and function Integration of activities Co-ordination of activities The organism and the State ... -I CHAPTER II THE SENSORI-MOTOR SYSTEM The meaning of organic movement Mobility patterns The sensori- motor system The motor organs The sensory mechanisms The nervous links An example of sensori-motor activity CHAPTER III THE PRINCIPLES OF ENERGY The nature of a material body Modern theories of matter The capacity for doing work Energy The diminution of available energy Energy in the abstract Potential and kinetic energ}' Releasing transformations The principle of becoming - CHAPTER IV THE SOURCES OF ENERGY The inanimate engine The animate engine Digestive process in animals The foodstuffs Absorption and distribution The respiratory interchange The sources of energy The removal of waste Glandular activity - 55 CHAPTER V ON VITAL PRODUCTION Animal metabolism The fate of the proteid residues Plant meta- bolism The balance of life Production and consumption - 76 xii CONTENTS CHAPTER VI BRAIN AND NEEVE PAGE The general scheme The arrangement of the ganglia The spinal cord The brain The development of the nervous system The human brain Connections within the central nervous system 87 CHAPTER VII THE SPECIAL NERVOUS MECHANISMS The sensory mechanisms The motor mechanisms The " simple reflex" mechanism The cranio-spinal reflexes The mechanism of co-ordination Cortical control - 106 CHAPTER VIII THE ANALYSIS OF BEHAVIOUR Stimulus and response in general The lower brain activities Nervous inhibition The cortex cerebri The meaning of behaviour The nervous system as a whole - 130 CHAPTER IX THE MECHANISTIC CONCEPTION OF LIFE The Cartesian mechanism The influence of chemical and physical investigations Modern mechanisms of life - 152 CHAPTER X THE MEANING OF PERCEPTION The nature of sensation Perception The problem of free-will In- determination in acting Memory and habit The " categories of the understanding " Space and time - - 164 CHAPTER XI THE NATURE OF LIFE The laws of thermodynamics The condition of the universe Dysentropic phases in the universe Digression on orders of magnitude and duration The improbability of life in the uni- verse The physical nature of life Life and energy - - 192 APPENDICES I. A .M MA 1-HYSICAL DISCUSSION . 222 I'ACE IN THE MODERN THEORY OF RELATIVITY - - 235 III. THE SUB VI AXILLARY GLAND AN INSTANCE OF ORGANIC FUNCTIONING - 237 INDEX -.--.--- 241 THE MECHANISM OF LIFE CHAPTER I ON THE NATURE OF ANIMAL LIFE The Organism as a Structure. The student of biology usually begins his studies by dissecting the body of a warm- blooded animal; and, first of all, he notices a division of this into well-marked regions: head and neck, trunk, limbs, and perhaps a tail. Then, taking a knife, he slits the skin over a limb and finds beneath it fleshy masses (the muscles) attached to rigid supports (the bones). He sees that the latter are movable on each other by articulations, or joints, and from his general knowledge of the animal in the living state he recognises that bones and muscles are parts that move in ways that depend on the nature of the articulations. Looking more closely, he sees white, glistening threads beginning in the muscles, joining together and passing into the central parts of the body. These are the nerves. There are also two kinds of bloodvessels: one kind which are stiff and white and apparently empty, and another kind which are rather thicker and softer, and which contain blood. The former are arteries, and the latter veins. All these parts bones, muscles, nerves, arteries and veins are wrapped up in a loose kind of material called connective tissue, and this must always be separated in order to disclose the forms of the muscles and other parts, and the ways in which they are joined together. Then, opening the cavities of the body, he discovers the viscera that is, the lungs and heart-, which occupy the cavity of the thoracic region of the trunk and the stomach, liver, alimentary canal, kidneys, digestive glands, and reproductive organs, all of which are contained in the abdominal cavity. Here, too, attentive study shows the existence of bloodvessels and nerves which ramify among the visceral parts. Lastly, breaking open the bones of the skull, he finds a white, 2 ' ITE -MECil-AlSM ' OF LIFE soft mass in the cavity of the head. This is the brain, and further examination shows that it is connected, by means of nerves, with the great sense organs of the head that is, the eyes, ears, nose and tongue but also with a similar mass of nervous tissue lying inside a tubular cavity in the backbone. This is the spinal cord, or marrow. Looking at it more closely, he finds that the spinal cord gives origin to nerves which can be traced out into the muscles of the trunk and limbs, and also into the skin. In short, he finds that the animal body is a structure : a series of parts arranged in a definite way. Structural Differences. Now, extending his study to other examples of the animal kingdom, he finds other structures which are analogous, or similar in a general way, to that which he has already examined. There are, however, very notable differences. The fore and hind limbs of the mammal are replaced by the wings and legs of the bird, or by the little side fins of the fish. He will find that all backboned animals possess these two pairs of limbs in some form or other, but that they may differ remarkably in structure. Examining the animals below the vertebrates, he will find still greater differences: thus there are about twenty pairs of limbs (or appendages) in a crab, shrimp, or lobster, but many more in a centipede, and such a creature as a starfish has apparently no limbs at all (though one speaks about its radial " arms "), but beneath the body he will find some thousands of mobile parts which serve for locomotion, and are known as the " tube- feet." The viscera also vary very strikingly. Thus there are distensible lungs in the quadrupeds, but these are relatively compact and inelastic in the birds, and attached to them are a number of long " air-sacs." There are no lungs at all in the fishes, but he will find gills there, organs which are apparently wanting in the warm-blooded animals. The heart, again, differs greatly in the various kinds of animals; thus it has four chambers in the mammals and birds, three in the frog and some other amphibians, but only two in the fishes. The eyes are large and well- developed in all backboned animals, but quite different in structure in insects and shellfish, like the crab and lobster; different, again, in the cuttlefish, rudimentary in the snail and in most worms, and wanting altogether in such creatures as the oyster, mussel and cockle. Thus the student of comparative anatomy finds very great
  • AND LIMBS IN A BACKBONED , , . , , . . , ANIMAL. behind, and are joined together in front at the pubis. The skeletons of the hind-limbs are jointed into the pelvic bones, but the latter, together, also act as a kind of basin-shaped support for the lower bowel, the urinary bladder, and the reproductive organs. Note that the articulation of the limbs with the limb girdles means that the former are set well out from the axis of the body, so that the feet rest on the ground on as wide a base as possible. The Limb Skeleton. All the higher animals that is, the vertebrates, the Crustacea (such as the lobster, crab, or prawn), the insects and spiders possess true limbs, which are organs of locomotion, and are also bodily weapons. The nature of these limbs is, of course, very different in the various groups of higher animals, but in all those that we have mentioned they are jointed appendages of the body, freely movable on the latter, and as a rule furnished with weapons. The limb may be a walking leg (as in the case of the fore and hind legs of most quadrupeds), a fin (as in the case of a fish), a paddle (in seals, dolphin, and 18 THE MECHANISM OF LIFE whales), a wing (in birds or insects), a " swimmeret " or paddle- 1 like limb (as in many Crustacea), a mobile weapon or tool fur- I nished with claws and cutting organs (as in the case of the fore- j limbs of a cat, the large chela?, or claw limbs of a crab or lobster, I or the arms and hand of a man), and so on. Sometimes its I structure and degrees of freedom of movement are relatively I simple (as in the side fin of the ordinary fish), but in general the I limb is a freely movable appendage of relatively complex struc- ture. The reader may easily verify all that we are about to say by I looking at a human skeleton in a museum, and by observing the I modes of motion of the parts of his own body. The fore-limb, or I arm, then, contains a skeleton that consists of a number of bones I connected together in various ways. The bone of the upper arm 1 (the humerus) articulates with the shoulder-blade by a ball- and- 1 socket joint, so that it is freely movable in every direction, and I the shoulder-blade itself can be moved (as in " shrugging "), so I that the arm has thus additional freedom. The skeleton of the I forearm consists of two bones (the radius and ulna), and the latter I is articulated to the humerus by a hinge-joint, so that the forearm I can be bent on the upper, or extended into the same straight line 1 with the latter, giving 1 degree of freedom of movement. The radius, however, has a peculiar twisting movement on the 1 ulna, and as the wrist and hand are articulated with it they canl be turned round through a half- circle, the elbow- joint remaining immovable. Thus the hand can be turned palm up (supinated) ] or palm down (pronated). The hand itself is capable of] 3 degrees of movement that is, it can be*turned in each of the] three directions of space. There are eight wrist bones arranged! in two rows, all articulating with each other, one row being! jointed with the radius and the other with the long bones of the] hand. Thus the hand, as a whole, can move up and down andj from side to side on the wrist, so that it has 3 degrees ofl freedom. Each of its bones, the metacarpals, articulates withj the bone of a finger that is to say, with the first, or proximal* phalanx, which articulates with the middle one, which finally; articulates with the terminal phalanx (the one that carries the] claw, or nail). The articulations between the terminal and middle phalanges and the middle and proximal ones are hinge- j joints, so that the fingers can only bend on each other in one] plane; but the 'articulation between the proximal phalanx and' THE SENSORI-MOTOR SYSTEM 19 its corresponding metacarpal is, to some extent, an universal joint. Thus the thumb can be bent in the same way as the other digits, but it can also be rotated or swung round in the same way that the whole arm can. The fi^st finger has the same kind of motion, but less of it, so to speak, and so also with the others. The hind- limb of a quadrupedal animal, or the leg of a man, is built on the same plan, and there may be very little difference between the two limbs (as in a chimpanzee, for instance) with regard to the degrees of mobility of the parts. In man. however, the freedom of movement of the hind-limb is restricted, first by Shoulder bla.de (fixed bone) FIG. 3. MOVEMENTS OF THE BONES OF THE FORE-LIMBS. the rigid attachment of the pelvis to the vertebral column, and second by the rigidity of the bones of the ankle relatively to those of the wrist. Such an analysis of the skeleton as we have just indicated (and which can easily be made more precise by the reader himself) shows that the degree of mobility of any part of the body of the higher animal is determined in general by the configuration of the skeleton, and particularly by the shapes of the joints. The latter make it possible for one bone to move on another in one or more ways; thus we have the hinge- joints between the skull and the atlas, the peg- and- socket joints between the atlas and the axis, 20 THE MECHANISM OF LIFE the ball-and-socket joints between humerus and scapula, and femur and pelvis, and so on. There is always a checking action of some kind which restricts the extent to which one segment of the skeleton can move on another ; thus the movement of the head is checked by muscles (and the relaxation of these when a man goes to sleep causes the head to drop, or fall to one side); the olecranon process (or " funny bone ") in the elbow checks the movement when the whole arm comes straight; the patella (or knee-cap) has the same function with regard to the movement of the lower leg on the thigh, and the slipping aside of the patella is the cause of the knee " going out of joint." In other cases the movement is checked by tendons, or ligaments (as in the case of the separate vertebrae). The Motor Organs. Now clothe the skeleton with muscles, which are so arranged as to pull on the bones in the ways that the latter are free to move because of the nature of the joints. This arrangement of muscle and bone is the subject matter of anatomy, and it can be treated in enormous detail ; but the reader will easily see that, from a mechanical point of view, it is all very simple. We have, in fact, systems of levers, with the various powers : thus w w t W v/ IF F FIG. 4. THE VARIOUS LEVER MOVEMENTS OF THE FOOT. Here 1 is a lever of the first order, 2 of the second, and 3 of the third order, and, with various modifications, these are typical of the mechanisms of movement of the parts of the limbs and body. THE SENSORI-MOTOR SYSTEM 21 One example may be given here, that of the bending of the forearm on the upper arm. Four bones are included in this mechanism the scapula, humerus. radius, and ulna. When we consider only the movement of the forearm on the upper by the elbow- joint, the scapula and humerus are the fixed, and the radio- ulna (considered now as one segment) is the movable part. -Shoulder blade. Jlead of humerus merus \J* 7 /ex o r muscle, (biceps) Ex. tensor muscle ( Triceps) Tendons _ of insertion --Ulna. Radius FIG. 5. DIAGRAM OF THE MUSCLES THAT MOVE THE FOREARM* Two muscles, we see. are necessary for the movement the biceps, which is the bending (or flexor) muscle, and the triceps, which is the straightening (or extensor) one. In each case the muscle consists of the tendons of origin (those which are attached to the fixed bone), the belly or contractile part, and the tendons of insertion (which are attached to the movable bone). When the radio- ulna is to be bent, or flexed, on the upper arm the biceps contracts, or shortens (that is, it becomes thicker and shorter), and simultaneously the triceps relaxes, or lengthens (that is, it becomes thinner and longer). By the pull of the tendon of insertion of the biceps the radio-ulna is bent on the upper arm, and by the pull of the tendon of insertion of the triceps it is drawn back again into a straight line with the humerus. It is important to'note that the two muscles are antagonistic ones, and that the flexion of one of them is always associated with the 22 THE MECHANISM OF LIFE extension of the other. Extension is not merely a passive relaxa- tion, but is the result of a nervous impulse just as much as is contraction. Note also that the humerus and scapula are fixed bones with regard to the flexion and extension of the forearm, but the humerus is itself a movable part, and the fixed points to which its muscles are attached are in the scapula. The latter is also movable, and then the fixed bones are the vertebra and ribs. The hand is a movable part, and the fixed bone to which its muscles are attached is the radio- ulna. Thus " fixed " and " movable " bones are relative to the parts that are to be moved. The reader must also note that the movement of any part of a limb is never so simple as we have indicated, for several pairs of i antagonistic muscles are generally in action at the same time. - Circular muscles, Circular muscles >f the Tine ^.Longitudinal muscles L ongitudina. I muscles contracted. FIG. 6. DIAGRAM OF THE MUSCLES OF THE INTESTINAL WALL. Non-Skeletal Mechanisms. There are many movable parts in the body for which there are no skeletal supports. Thus the heart consists of a very complicated series of muscle bundles that contract and expand automatically. Simpler cases are those of the intestine and bloodvessels. In the former we have two series of muscles, one which is made up of bundles of fibres running circularly in the wall of the bowel, and the other being made up of fibres that run lengthways. When the former contract the diameter of the intestine is decreased, and when they relax and the longitudinal muscles contract the length of a segment of the bowel is decreased and the diameter is correspondingly increased. In this way waves of contraction of calibre of the intestine are set up, and these propel the food contents from place to place. The mechanism of contraction of an artery is very similar, except that the longitudinal muscles are not present. The wall of the artery is elastic, and when the circular muscles contract the calibre is diminished, but when they relax the elasticity of the wall, and THE SENSORI-MOTOR SYSTEM 23 the pressure of the blood within, cause the vessel to regain its former calibre. There are other special mechanisms of this kind; thus the actions of closing and opening the eyes involve the contractions and relaxations of two antagonistic muscles which are present in the eyelids. One of these, the closing set, consists of fibres that form a kind of ring in the upper and lower lids. When these contract, the opening between the latter is diminished, or obliterated. The closing muscle consists of fibres which originate in^the sheath of the optic nerve, and which are inserted into the cartilage of the upper lid ; when they contract the latter is raised, the ring-like closing muscle then becoming relaxed. Ca.frilla.ry FIG. 7. MUSCULAR TISSUE. HIGHLY MAGNIFIED. The Minute Structure of Muscle. Just a word about this. Examination of a piece of muscle beneath the microscope shows that it is composed of a very great number of minute fibres. Each of the latter is about If inches in length, and it tapers away to a fine point at each end. Numbers of fibres are bound together by connective tissue to form muscle slips, which are similarly joined up to form bundles which constitute the muscle itself. A nerve fibre terminates in each muscle fibre, as shown in Fig. 7. In addition, arteries carry blood to the muscle, where it is distributed through the capillaries and from which it is carried away by the veins. The muscle fibres act at the same time, all of them thickening or contracting when the muscle pulls on its movable bone, or simultaneously lengthening when the muscle relaxes. As the figure shows, the fibres are cross-striped, and this stria- tion is, in some way, part of the mechanism of contraction and relaxation. But the muscle fibres that are present in the intestinal or arterial wall are not cross-striped. This presence or absence of striation goes along with a difference in the control over the muscular'actions ; in^most orthe cases^'where the activity of a muscle is controlled by the will (voluntary muscles) the fibres 2 \ THE MECHANISM OF LIFE are striated, while in those over which we have no control (the muscles of the heart, alimentary canal, and bloodvessels chiefly the involuntary muscles) the fibres are " plain," or non-striated, or are otherwise different from the ordinary striated fibres. The Sensory Mechanisms. So much for the apparatus of mobility. The reader can easily supplement the above very general summary from the textbooks on physiology. Next we have to consider how this apparatus is set in motion, and that leads us to the study of sensation. There are certain mechanisms, mainly those of the heart and respiratory organs, which are automatic that is, they work in the absence of any external cause (though their rate of working is affected by outer events) but in most of the muscular mechanisms of body and limbs a stimulus is necessary in order that they may be set in motion that is, something analogous to the pressing of the spring in our imagined model must occur. Sensation, then, is the preliminary to movement, and here we mean by " sensation " only the physical and chemical events that occur in the organs of sense and in the nerves and brain, and not any affection of consciousness; when the latter occurs we have to do with perception, and this we consider later. Things that happen in the environment, then, affect the organs of sense : the movements of material bodies are known to us by vision, for the light reflected from those things enters the eyeballs and forms a picture on the retina, which is the essential organ of vision ; vibrations in the atmosphere are set up by other material move- ments outside ourselves, and these vibrations are received by the auditory organs ; the particles of chemical substances floating in the air are drawn into the nose or mouth, and so affect the olfactory and gustatory organs; while contact of material sub- stances with the skin similarly affects the nerves that terminate there. These sensations we call sight, hearing, smell, taste, and touch. Exteroceptive and Proprioceptive Sensation. Changes that occur outside the body thus affect the organs of sense, or, as we shall call the latter, the receptors. The eyes and auditory organs are therefore exteroceptive organs, or distance receptors, for they are affected by events that may happen at considerable THE SENSORI-MOTOR SYSTEM 25 distances away from the body. The olfactory, gustatory, and touch organs are near receptors, for they can only be affected by substances that actually come, into contact with the body. Our sensation of temperature may be a near affect (as when a cold or hot substance touches the skin), but it may also be a dis- tance affect (as when we are warmed by the radiation from the sun). Now there are receptor organs which are stimulated, or affected, by changes that occur within the body itself; thus when we lift anything we have a feeling of effort (the muscular sense) ; when we walk we have also sensation arising from pressure set up in the joints; when the body is in different postures there is sense of equilibrium, which is something quite apart from seeing or hearing (for it can arise when a man is blindfolded and has his ears plugged); and there is also general sensation from the viscera, for a man may experience stomach-ache. In these cases receptor organs in the muscles, tendons, joints, ears, and viscera are stimulated, producing what we call proprio-sensation. Emotion (that which moves us), blushing, the pallor of fear, trembling, rigidity, even the " Rabelaisian effect of fear on the bowels " (" affective tone or feeling," as it has been called), are the results of the stimulation of the proprioceptors. Here, in spite of the warning given above, we are speaking of sensation as it results in " feeling," or consciousness, because it is very difficult to avoid doing so ; but, nevertheless, the activity of the receptor organs, in so far as we are about to relate it to the initiation of movement, need not be accompanied by any affection of the mind. The Nervous Links. Certain mechanisms intervene between the receptor organs on the one hand and the motor organs on the other; these are the peripheral and central nervous systems. Proceeding from a receptor organ there is an afferent nerve (" afferent " because it conveys something to the central nervous system), and pro- ceeding from the motor organ there is an efferent nerve (" efferent " because it conveys something from the central nervous system). The reader must understand clearly that a motor organ is (in general) not directly stimulated to act by something that occurs outside the body ; it is stimulated via the central nervous system. Receptors and afferent nerves are therefore " the way into " the brain and spinal cord, while efferent nerves and motor organs 26 THE MECHANISM OF LIFE are " the way out." When the animal does anything in response to some change that occurs in its environment (1) a receptor organ is stimulated by the external event, (2) an impulse is propagated along an afferent nerve into the central nervous system, (3) some change occurs in the latter, (4) an impulse is propagated out along an efferent nerve, and (5) the motor organ is stimulated, and responds by some kind of movement. The Minute Structure of the Nervous System. Complicated as this is, its general scheme is easily understood. In a receptor organ there is always an essential part which we call the nerve NodA FIG. 8. FORMS OF NEURONES. ALL HIGHLY MAGNIFIED. 1, Transverse section of a nerve fibre; 2, a nerve fibre; 3, nerve cell from the spinal cord; 4, nerve cell from the cerebellum; 5, nerve cell from a sympathetic ganglion. termination; this is the retina in the eye, the auditory hairs in the ear, and so on. Between the receptor and the brain or spinal cord there are stretched a series of nerve fibres, and it is along these that the impulse passes. In the central nervous system there are collections of nerve cells and nerve tracts. Between the centre and the motor organ there are other fibres, and in tl former there are. again, nerve terminations. All this structural detail is built up of nervous elements called neurones. Neurones are structures that vary greatly in appearance and size, but they have all the same general form. Each of them consists of a nerve cell that is, a minute fragment of specialised THE SENSORI-MOTOR SYSTEM 27 protoplasm containing a nucleus. From one side of such a cell there arise one or more prolongations of the protoplasm, and these branch repeatedly to form a plant-like growth an arborisation which is known as the dendritic system of the cell. From some other part of the cell there arises a delicate filament which does not usually branch, but is prolonged out into a long thread, the nerve fibre. As Fig. 8 shows, this fibre, which is called the axon of the cell, becomes invested in a sheath, and between the sheath and the axon there is a sort of packing, the medulla. The axon may be very long; thus the cells of the grey matter of the spinal cord send out axons into the sciatic nerve, and some of these may be about 3 feet in length, though the nerve cell itself is only about ^-Q-Q to ^ inch in diameter. The axons, of STIMULUS Cell Jtferve terminations Motor plate J$EURONE FIG. 9. DIAGRAM OF THE WAY IN WHICH NEURONES ARE CONNECTED. course, constitute the nerves. At the extremity away from the nerve cell the axon always breaks up into a second series of dendrites, or terminations. The essential part of a sense organ such as, for instance, the retina of the eye, or the auditory cells and hairs in the organ of Corti in the internal ear, always consists of the proximal part of a neurone, or even of several neurones, each of which has usually a very short axon. It is always the dendrites of a nerve cell that receive the stimulus, and the latter is always conducted along the axon from the nerve cell to the other terminal arborisation. Thus nerves invariably conduct impulses in one direction only. Neurones are always connected together by means of synapses. At a synapse the distal dendrites of one neurone come into close proximity to the proximal dendrites of another neurone. As 28 THE MECHANISM OF LIFE Fig. 9 shows, the two series of dendrites do not touch each other, but the branches of the arborisations " interdigitate." They approach each other very closely, but there is always a space between them, and this space is, of course, filled by other tissue or by liquids. This mode of joining up of successive neurones to form a chain is universal throughout the nervous system, both in the central and the peripheral regions. An Example of Sensori-Motor Activity. We shall take what is, perhaps, the simplest and most convenient common action, that of " winking." When something unexpectedly and rapidly approaches the face, the eyes " instinctively " close in order that they may be protected. Now this action of closing and subsequent reopening involves a receptor organ, an afferent To cortex. SgQSfL otor Centres oculomotor Jrorn cortex Closing muscle- -3 FIG. 10. SCHEME OF THE NEURONES CONNECTING THE RETINA WITH THE MUSCLES OF THE EYELIDS. nerve, a nerve centre, an efferent nerve, and a motor organ. The receptor is the retina of the eye. Light reflected from the moving object forms a minute picture on the retina, and in some way the proximal dendrites (which in this case, however, are not dendrites in the usual sense, but rather the rods and cones of the retina) are affected by the variations in light and shade (that is, by differences in the intensity and quality of the radiation), and nervous impulses are set up which travel along the axons of the retinal nerve cells in the optic nerve to the brain. There they are received by a nerve centre, and the afferent impulse is con- verted into an efferent one, which travels out from the centre, via the axons of the nerve cells there, through two nerves. One of these is the third cranial, or oculo-motor nerve, and the other is a branch of the seventh cranial, or facial nerve. The former THE SENSORI-MOTOR SYSTEM 29 goes to the lifting muscles of the eyelids, and the latter goes to the closing muscles. The same event in the centre gives rise to both impulses simultaneously, and that travelling along the oculo- motor nerve causes the opening muscles to relax, while that which travels along the branch of the facial causes the closing muscles to contract. The eyelids thus close. Im- mediately afterwards the series of events is reversed because of a second pair of efferent impulses; the opening muscles now contract, and the closing ones relax, and so the eyelids open. Representing this series of action in a very schematic way, we get the diagrams given on pp. 28 and 29. ^Retina Eyelid. enmef muscle) FIG. 11. -SCHEME OF THE CONNECTIONS BETWEEN THE RETINA, BRAIN, AND EYELIDS USED IN THE ACT OF WINKING. According to the figure, there is only one proximal set of den- drites in the retina ; in reality, there are three (as is represented in Fig. 30, p. 107). Further, only one centre is shown, but really the optic nerve ends in three centres in the mid- brain (see Fig. 32, p. 115). From these centres other neurones make connection with the oculo-motor centre, and from the latter another set of neurones pass out along the oculo-motor nerves to the opening muscles, while yet another traverses the nucleus (or centre) of the facial nerve, and go out through the latter to the closing muscles of the eyelids. In any activity of the sensori-motor system, then, a rather complicated mechanism is involved. Some change occurring in 30 THE MECHANISM OF LIFE the environment acts upon a receptor organ by stimulating the proximal dendrite of a nerve cell, and thus setting up a nervous impulse which travels up the axon of this cell into a nerve centre, or nucleus, or ganglion, in the brain or spinal cord. The axon ends in the centre as a series of distal dendrites, which form a synapse with the proximal dendrites of another neurone. The axon of the latter leaves the brain or spinal cord in an efferent; nerve, which goes to the muscles concerned. The impulse travelling out is distributed to the muscle fibres by the distal dendrites of the final neurone, and, entering the muscle, it releases energy and causes the latter to contract or relax, thus causing the movement. This, it must be remembered, is only a scheme of the mechan- ism, and we elaborate it in Chapters VL to VIII. Meanwhile, it will suffice to give the reader a preliminary notion of what are the essential activities of the sensori-motor system. CHAPTER III THE PRINCIPLES OF ENERGY So far our description of the animal body has been that of a mechanism which can be actuated, or made to "go." How it is actuated that is, what are the sources of its energy, and how these sources are utilised is the subject of the following chapter. Meanwhile, however, something must be said about the general principles 01 energy in so far as they concern us in our study of life. First, then, we ought to consider what is meant by The Nature of a Material Body. A material thing or body is something that is heavy ; that has shape, or occupies space; that coheres and is dense in varying degrees; that has heat, also in varying degrees; that has texture, lustre, colour, smell, and taste. In short, a material body is a massive substance which has physical " properties." Coherence. Material bodies may be solid, liquid, or gaseous. When they are solid they can be disintegrated by mechanical means that is, they can be crushed, broken, powdered, filed, etc. and therefore they must consist of smaller parts that cohere together, but which can, nevertheless, be separated. There appears to be a limit (in the practical sense) to the degree to which a solid material can be pulverised, though we can reduce it to exceedingly fine particles. By-and-by, however, our mechanical means of disintegration fail to make the particles any finer, but in imagination we can still divide them. The chemist can show that the finest particles to which a body can be reduced are still aggregates of molecules, and thus he regards the latter as the very finest particles to which a material body can be reduced without losing its specific properties. Molecules are therefore the ultimate particles of which bodies are made up, and they cohere together more or less strongly, or not at all. When the degree of cohesion is such that the body preserves its shape irrespective of anything that contains it, we say that it is solid; but when the molecules still cohere, but slip on each other, we call the body viscous (as in the cases of pitch or 31 32 THE MECHANISM OF LIFE syrup), or liquid (as in the case of water). When they do not cohere at all, so that the material can take any shape and expand to any extent, we call it a gas. Thus in ice the molecules cohere strongly, but not nearly so much when the ice is melted, and hardly at all when the water is converted into steam. Temperature and Heat. Now all bodies of which we have any experience possess some heat. Heat is a " mode of molecular motion " a form of energy and the intensity of heat (but not its quantity) depends on the rapidity of motion of the molecules. At a temperature of 273 C. below the freezing-point of water all movements of the molecules themselves (but not of the parts of the molecules) cease, and this temperature is the absolute zero, and is approximately that of cosmic space. When the tempera- ture rises, the molecules move more rapidly until they become loosened from, but still attract, each other ; then the body melts. Rising in temperature still more, the molecules finally cease to attract each other, and each of them moves freely so that they tend to fly apart; then the body becomes a gas. The tempera- tures at which melting and vaporisation occur are, of course, different ones in different materials, and depend on the nature of the molecules of which the body is composed. When bodies are at different temperatures, heat tends to flow of itself from the warmer to the colder body. If, when we touch a body, heat flows from it to our skin, we say that the body is warm, and if the converse happens, we say that the body is cold. Density. The more molecules there are in the same bulk the denser, we say, the body is. Thus a square inch of ice contains a certain number of molecules of H 2 0, and the substance has a certain density ; but when the ice is melted and the temperature of the water rises to, say, 65 F., the same number of molecules now occupies a greater volume (for they are further apart), and so the density becomes less. When the water is raised to, say, 220 F., it is converted into steam, and it now occupies over 1,700 times the volume it had in the liquid state. Therefore its density is very much less. This means that the density of a body depends on the closeness with which the molecules are packed, or cohere together. But it also depends on the relative weights of the molecules, for some are heavier than others; thus the molecules of quicksilver are, each of them, much heavier than those of water. THE PRINCIPLES OF ENERGY 33 Texture and Lustre. This depends on the ways in which the particles are, so to speak, laid alongside each other ; thus polished steel has a smooth texture, but that of a fractured piece of steel is rough and crystalline. Lustre is a kind of texture, the body exhibiting it being very smooth, so that it reflects the light that falls on it. Colour. This depends on the chemical nature of the mole- cules, and on the ways in which they are arranged together. Gold, for instance, when highly polished, has bright, metallic lustre, and is yellow in colour, but when it is finely divided it may be purple. Sunlight is a mixture of light of different colours, or wave-lengths, and material bodies can act differently on this mixture. When the proportions of the lights of different wave-lengths reflected from a body are the same as the proportions in sunlight, the body appears white (or grey) to our eyes. When all the light that falls on a body is absorbed by it, none being reflected, we say that the latter is black. When some of the wave-lengths are reflected and others are absorbed, we say that the body is coloured. Thus rouge absorbs all the light falling on it except that which has the wave-length associated with what we call scarlet; it reflects this kind of light, and so it appears coloured. In monochromatic light that is, light of one colour all bodies look as if they had the same hue, only brighter or duller. Smell and Taste. Most bodies give off fine particles into the air, and these become inhaled into our nostrils. If such particles can dissolve in the liquid bathing the olfactory mucous mem- brane, they can react upon or stimulate the nerve terminations in the latter, and so they give rise to the sensation of smell. If a substance placed on the tongue can dissolve and affect the termination of the gustatory nerves, we have the sensation of taste. Different Kinds o Molecules. The molecules, or ultimate particles of which material bodies are composed, are not all the same; for instance, quicksilver consists of molecules of the chemical substance mercury, and water consists of molecules, each of which consists of three atoms (H 2 0), two of hydrogen and one of oxygen. Molecules are therefore made up of atoms, and there are about 100 different kinds of the latter. All material bodies, of whatever nature they may be, are therefore composed 3 34 THE MECHANISM OF LIFE of molecules, each of which contains a relatively small number of one or more kinds of chemical atoms. Now colour, smell, taste, lustre, texture, temperature, and density, are dependent on the chemical nature of the molecules, and on the ways in which the latter are arranged and on their motions. Thus water may be warm or cold to our sense, but it consists in each case of the same molecules moving relatively slowly when the water is cold, and relatively quickly when it is warm. Phosphorus may be yellow or red, and in the former case it is odoriferous, poisonous, and combustible; while in the latter case it has no smell, is not poisonous, and is non-inflammable in the conditions in which yellow phosphorus is inflammable. Yet the substance phosphorus is chemically the same in both cases, only the atoms are in different configurations. The same number of molecules of H 2 may be dense in the form of ice, less dense in the form of water, and less dense still in the form of steam, accord- ing to the distances that its molecules are apart from each other. Something, however, is the same in the case of the red and yellow phosphorus, or the solid, liquid, and gaseous water that is, the mass of the chemical substance itself. Apparently colour, taste, smell, density, temperature, etc., may be variable, while mass remains invariable. Weight and Mass. Weight itself is something that may vary,] while mass remains the same. A material body weighs more at^ the earth's poles than it does at the equator, and if it could be* transported to the sun it would be much heavier, or if to the moon much lighter, than it is on the earth. If it could be removed to several millions of miles away from any cosmic body its weight would be almost nothing. Weight depends on the mass of the body, on its distance from some attracting body (such as the sun, earth, or moon), and on the mass of the latter. The mass of our material body would be the same everywhere (we are neglecting some physical results included in the theory of relativity), but its weight would vary. Mass. Something, then, seems to be invariable, or nearly so, and this is the mass or quantity of matter in a body. The mass of a cubic inch of iron is so much in all circumstances, and that of a cubic inch of gold is more than that of the same bulk of iron, but is also invariable. By the quantity of matter, or mass of a body, we therefore mean the number of molecules in the body rnulti- THE PRINCIPLES OF ENERGY 35 plied by their molecular weight. (The molecular weight of the lightest molecule, that of hydrogen, H 2 , is taken as the standard to which all other molecular weights are applied.) Mass and Inertia. Mass is, then, the most fundamental thing in our notion of materiality, but it is not an irreducible concep- tion, and we must seek for some way of defining and measuring it. Note, first of all, that we have a bodily intuition of mass: let there be two exactly similar stoneware, corked bottles, and let one be filled with water and the other with mercury. We cannot say which is which merely by looking at them, but we can distinguish if we lift them, for a greater degree of effort of our muscles is necessary to lift the mercury than to lift the water, and we have the " feeling " of this effort. Nevertheless, such an intuition would be mostly useless to us, for it would not apply to great masses which we cannot lift, nor to very small ones, nor to masses which were nearly the same. A\ e must have some way of measuring mass by taking some dimension of space, or by counting (or numbering) something. Now there is a constant and universal property of masses that of gravitation; all material bodies, whatever their nature, or -mass, fall to the surface of the earth when they are free to move. If they are sufficiently far away they will fall 16 feet in the first second, 64 feet in the first two seconds, 144 feet in the first three seconds, and so on, their rate of fall being accelerated by 32 feet per second per second during the time that they are falling. "We do not know in the least what the force of gravitation is, but we know precisely what it does it accelerates the rate of motion of a body free to fall. Consider what this means : there are three factors mass, space, and time; a mass, when it is attracted by the earth, falls with increasing velocity that is, it falls through a greater space in the third second than in the second one, and through a greater space in the second one than in the first. This acceleration of the motion of a mass during a certain time we shall call the work done by the force (whatever the latter may be). Place a mass of metal (say a pound weight) in a scale pan ; it will fall, and the other scale pan will rise. Now place another pound weight in the other scale pan, and the two will balance each other so that neither falls. The work done by gravity on the one is equal to the work done on the other: neither is accelerated; the space and time factors are the same, and therefore the masses are 36 THE MECHANISM OF LIFE the same. Take a piece of wire of uniform diameter and density, and (say) 10 inches long, and put it in one scale, and put a piece the same wire 1 inch long in the other. The mass of the 10-in< length will be greater than that of the 1-inch piece, for the scale) pan in which it is placed will fall, and the other one will rise.j Put a 1-inch length in one pan and another 1-inch length in the] other, and it will be found that the work done by the earth's] gravity is the same in each case, and the masses are therefore equal. Finally, put the ten 1-inch pieces in one pan and the] 10-inch piece in the other, and again it will be found that the work done is the same in each case, so that the mass of each 1-inch length is one- tenth of that of the 10-inch length. Thus we can find the mass of a body by measuring the world done upon it, when it is free to fall, by the earth's gravity, and comparing this with some standard amount of work done. In whatever way we measure this work done it will be found that we always measure a space. Even when we measure time it is really a space that we determine. A material body that is in a state of rest will continue in aj state of rest, or if it is in a state of uniform motion it will continue , in that state of motion unless work is done upon it. Of it-sol t it is inert. Its inertia varies according to its mass, so if a greater mass that is at rest is to be moved, or if a greater mass that is in I uniform motion is to be stopped, a greater amount of work must be done. Inertia means the tendency of something to remain as it is, unless some external agency acts upon it. Modern Theories of Matter. We must say a few words about these. Not so long ago it wa* thought that all material bodies, or kinds of matter, were built up of chemical atoms which were the ultimate particles. An atom was regarded as indivisible. But some atoms were known to be heavier than others, and so their masses were not the same ; thua the mass of an atom of platinum is about 194 times greater than that of an atom of hydrogen. Therefore, the heavier atoms ha< more of something in them than the lighter ones, but more o what ? There might be some universal kind of matter containe< in greater quantity in the heavier than in the lighter atoms, but if so the former could hardly be thought about as indivisible which they must be, according to chemical theory. The way out from this paradox was found by the discovery o THE PRINCIPLES OF ENERGY 37 radio-activity. As the result of the physical research based on this, it became very probable that an atom (which is still the smallest particle of matter as such that can exist) is really com- plex, and consists of a system of electrons. An electron is not material, but is an unit charge of electricity, the smallest charge that can exist. In the centre of the atom there is an electron, and revolving round this there are others, much in the way the planets are revolving round the sun in the solar system. Now one or more of the electrons can be expelled from the system, and then the latter becomes a new kind of chemical atom. Different atoms contain different numbers of revolving electrons. What is an electron ? All we can say about it is that it is elec- tricity, and not material. Thus, materiality dissolves into energy, and the latter is the fundamental physical reality. Mass disappears, but inertia remains, only the latter is now electro-magnetic inertia. The Capacity for Doing Work. Things, then, remain as they are unless something is done to them. A train remains at rest unless work is done upon it, causing it, for instance, to attain a velocity of thirty miles per hour three minutes after it has started to move. Conversely, the train will continue to run after the steam has been shut off unless work is done upon it, causing it to stop, and such work may be done by the application of the brakes, or by the friction of the air, or that of the wheels on their bearings or upon the rails. The mechanical work that is done in both these cases is measured in horse-power. 1 h.p. being the amount of work done by raising a weight of 33.000 pounds 1 foot high in one minute. The water in a steam boiler will remain at one temperature unless work is done upon it. Heat must be supplied to the water by the combustion of coal in the furnace. Now we can express the amount of heat in units, which are called Calories, each Calorie generated in a second being 5-62 h.p. Thus to raise the temperature of the water in the boiler that is, to change its state in a certain way so much work must be done. An electric tram will stand still upon the rails for ever unless work is done upon it. When the switch is closed electric current enters the motors, and the latter revolve propelling the car and giving it a speed of, say, ten miles per hour in one half- minute. We measure this work done by the current in units, called kilo- watts, each of the latter being 1-341 h.p. 38 THE MECHANISM OF LIFE A dynamo will remain at rest and yield no current unless work is done upon it, and this also we measure in h.p. Supply motive power equal to so many h.p. to the dynamo, and the latter, with its conductors, changes its state and develops so many kilowatts of current. Finally, a man may climb a mountain, say 5,000 feet high, in six hours, but not unless certain substances in his body become | oxidised, yielding muscular power capable of raising his body against the resistance of the earth's gravity. But, again, he cannot supply this muscular power and do work unless he takes food, and a certain quantity of the latter that is, so much proteid, fat, and carbohydrate estimated in equivalent heatj units, called Calories, must be supplied before his body can do the specified work. In all these cases something called the capacity for doing work is added to the thing that changes or does work. Chemical sub- stances (the fuel and oxygen) are burned in the steam boiler, i electric current is fed into the motor, mechanical motion is im- parted to the dynamo, and chemical substances are assimilated into the muscles of the mountaineer. All these things represent the capacity for doing work, and the latter we define more shortly j as available energy. Obviously there are different forms of available energy, and' these can be transformed one into the other. There is mechanical energy, that of the motions of material bodies for instance, the motion of the parts of the locomotive engine; heat energy that is, the enormously increased velocity of movement of the mole- cules of something or other ; electric energy, which is the flow of I electrons through a conductor; muscular energy (about which we know very little) ; gravitational energy ; the energy of radiation, I etc. We know little or nothing of what these various forms of] available energy are, although we recognise them as different, for they affect OUT sense organs differently; thus we recognise mechanical energy because we exert muscular power in opposing j it ; we recognise heat through the stimuli of certain sense organs ; I radiation usually by the stimuli of the retina (and also of thej skin); electric energy mainly by its stimulation of the muscles,] and so on. But all forms of available energy can be converted one into the other, and all of them are the capacity for doing work, or more] generally the state of something or other. But, again, in what- j THE PRINCIPLES OF ENERGY 39 ever form this capacity for doing work may exist, it can always be measured, directly or indirectly, as a certain quantity of mechanical work performed. That means that we do not know what available energy is, we only know what it does. Energy Transformations. We may profitably expand the result stated in the last para- graph, and give some further examples. There is a heap of coal, and this we shall call available chemical energy. It contains certain substances carbon and hydrocarbons and these can combine chemically with the oxygen of the air to form carbonic acid gas and water (with some other substances). In this act of chemical combination heat is generated. If the burning fuel is confined in a furnace, a large fraction of the heat may be commu- nicated to the water contained in a steam boiler, and its effect is to increase the motions of the molecules of the water. By-and- by these motions become so rapid that the molecules fly apart and the water is converted into steam. Later on the velocities of the molecules of the steam are increased by the further flow of heat from the furnace, and they collide with, and rebound from, the walls of the boiler in which they are contained, thus setting up steam pressure. The chemical energy of the fuel is therefore transformed into the kinetic energy of the molecules of steam under pressure. The steam is then allowed to expand into the cylinders of the engine, and in so doing it pushes out the pistons and communi- cates a rotatory motion to the crank shafts. Thus the kinetic energy of the molecules of steam is trans- formed into the kinetic energy of the large, moving parts of the engine that is, into mechanical energy. The motion of the engine is next communicated, by means of shafting or belts, to the armature of a dynamo. When the latter revolves it generates an electric current, and this uses up the mechanical energy; for if the switch of the dynamo is open, relatively little power is required to rotate the latter, and no current -passes. But when the switch is closed, relatively much power is required, and a current is generated. Mechanical energy therefore transforms into electric energy. 40 THE MECHANISM OF LIFE The current generated can now be used in various ways; for instance, it can be sent through lamps, when the latter glow; or it can be made to decompose clay, producing aluminium; or it can be fed into motors, generating mechanical energy; or passed through electric fires, generating heat. Thus electric energy transforms into radiant (light) energy; into available chemical energy; into mechanical energy; and into heat energy. And so on ; summarising the example we have given, and many others that might be quoted, we get our first principle of energetics : Any form of available energy can be converted into any other form . . (1) The Diminution of Available Energy. We come now to a very important result. Returning to the series of examples just given, we note that a certain quantity of coal contains chemical energy, and that the latter may be con- verted into heat. The quantity of heat which can be generated by the combustion of the coal can be estimated in the following manner: A small weighed piece is powdered, mixed with some substance which yields free oxygen, and is placed in a kind of bomb, which is put into a vessel containing a known mass of water at a known temperature. The mixture of coal and oxygen- yielding substance is fired electrically, and precautions are taken that all the heat generated goes to raise the temperature of the water. The result is that a certain quantity of water, say 1 kilogram (=2-2 pounds), is raised in temperature, say 1 C. That quantity of heat is called a Calorie, and so we estimate the " calorific value " of the coal that is, the quantity of heat that is generated when an unit mass is burned. Now let the coal be burned in a steam-boiler furnace, and let the steam produced work an engine. The work developed by the ' latter can easily be estimated, and so we can find the number of h.p. given per ton of coal, or per Calorie. Next, use the engine to drive a dynamo, and use the current generated by the latter to drive an electric motor. Suppose that we utilise the entire power of the engine to drive the dynamo, and the entire current given by the latter to drive the motor. Now estimate the h.p. developed by the motor, and compare it with the h.p. developed by the engine; it is much less. THE PKINCIPLES OF ENERGY 41 Or use the entire power of the current to work a series of electric fires, and estimate the quantity of heat generated by the latter; it is very much less than the quantity of heat originally generated by the burning of the coal that raised the steam, that worked the engine, that drove the dynamo, that produced the current, that generated the heat of the electric fire; or, again, estimate the h.p. required to drive the dynamo, use the current given by the latter to drive a motor, and then estimate the h.p. developed by the latter; it is about 5 to 10 per cent, less than that required to drive the dynamo. Finally, estimate the number of kilowatts of current required to work a motor, and then employ the latter to drive a dynamo, and measure the current generated by the latter; it is less (5 to 10 per cent.) than the current originally used to drive the motor. That means that there is waste incurred whenever there is an energy transformation, and it is quite easy to see how this waste occurs. When coal is burned in the furnace of a steam boiler, a quantity of heat is lost by radiation from the boiler and furnace doors, and another fraction is carried away through the flues into the atmosphere ; there is radiation from the steam pipes and hot cylinders, and still another fraction of heat is given up to warm the condenser water. Thus all the heat generated by the com- bustion of the coal does not transform into the mechanical energy of the engine; in fact, only about 10 to 20 per cent, does so. Further, there is friction in the bearings, slides, etc., of the engine, and so mechanical energy is lost. In communicating the engine power to the dynamo more friction is incurred. In the dynamo itself there is friction, and some of the current developed is wasted on heating the parts of the mechanism, while more is lost by imperfect insulation. These latter sources of loss also exist in the means whereby the current is transmitted and then converted into radiant energy, or heat, or mechanical energy. When the current is used to generate light there is great loss, for heat must first be generated until the intensely hot filaments, or arc, or vapour, glow. If light could be generated directly from chemical action as it is by a glow-worm there would be great economy. Thus there are a host of ways in which available energy is lost in the course of its transformations. There is friction, imperfect heat conduction and insulation, imperfect electric conduction and insulation, materials which are not perfectly elastic or per- fectly rigid, etc. In all cases this lost energy reappears as low- 42 THE MECHANISM OF LIFE temperature heat, but the latter is unavailable energy, and we cannot transform it. Thus we get our second principle : In all transformations of energy some of the latter becomes unavailable. Thus the capacity for doing work continually diminishes . . . . . . . . . . . . (2) Now consider another notion which is really involved in our second principle, but which it is useful to discuss separately; that is, the notion of Reversible and Irreversible Transformations. Take a suitable form of dynamo and cause it to revolve, thus generating a current of, say, K kilowatts, by the expenditure of, say, H horse- power. Then take K kilowatts of current, and supply it to the dynamo, when the latter will begin to revolve and will act as a motor. Thus the same machine is reversible; when current is supplied it will generate mechanical energy, and when mechanical energy is supplied it will generate electricity. But notice particularly that we supplied K kilowatts of current, and got H horse-power. Now if we supply H horse-power, we only get about 95 per cent. K of current. Therefore our rever- sibility is not quite perfect; if it were we should have got K kilowatts of current, instead of 95 per cent. K. Take the case of a steam engine and consider its theory. We have a " working substance," the water in the boiler. Now let this working substance be heated; it expands enormously, does work by moving the pistons, and finally escapes into the con- denser, where it heats up the circulating water. Therefore, the working substance takes heat from the steam boiler, converts some of this heat into mechanical work, and gives up some to the condenser. Now let the water in the boiler cool down, and try to reverse the process by actuating the engine in the opposite direction. If the mechanism were reversible, heat would be taken from the condenser, the mechanical work done on the engine from outside would transform into heat, and these two quantities of heat would be transferred to the water in the boiler, heating up the latter to boiling-point and beyond, and so establishing steam pressure. This cannot be done (in practice), and so the steam engine is an irreversible mechanism. Take a very famous experiment and try to reverse it : Joule, in 1843, caused a paddje-wbeel to revolve in water con- THE PRINCIPLES OF ENERGY 43 tained in a heat-insulated vessel. He took all possible precau- tions, measured the amount of mechanical work done (by causing a falling weight of known mass to actuate the paddle), and measured the mass of water and the rise of temperature produced by the friction of the paddle. He found what is now called the " mechanical equivalent of heat " that is. he found that when a weight of 1 kilogram falls through 427 kilometres, 1 kilogram of water is raised in temperature 1 C. This amount of heat communicated to the water is called 1 (large) Calorie. Now imagine the water contained in the mechanism to be heated up 1 C. ; if the latter were reversible the paddle would revolve. But it does not, and the apparatus is therefore irreversible. Finally, make the end of a poker red-hot in the fire, and then take it out and let it stand ; it will cool down, and in half an hour or so it will have attained the temperature of the air in the room. Its heat has been lost by radiation and convection, and has gone to raise the temperature of the air, furniture, and walls of the room ever so little. Imagine the experiment to be reversed, so that the heat of the room would flow into the end of the poker and raise its temperature to redness ; such an effect has never been observed to occur (though if it did occur physicists would not be incredu- lous !). Therefore the flow of heat, of itself, is irreversible. These examples will enable us to formulate two more statements : Some energy transformations are approximately reversible and others are irreversible . . . . . . . . . . (3) The flow of heat (of itself) is irreversible, only going from a hot [ to a cold body . . . . . . . . . . . . (4) Thus we can, quite easily, cause some energy transformations to occur, but not so easily, or not at all, some others. We can easily cause all kinds of energy to transform into heat, and, in fact, they do transform into heat of themselves. Mechanical friction always generates heat, chemical action generally produces heat and sometimes chemical energy com- pletely transforms into heat ; the flow of electricity through a con- ductor, the straining and bending of materials (internal friction), the reception of light by substances in short, all physical reac- tions transform, or tend to transform, into heat. Therefore All forms of available energy tend to be transformed into heat, but heat is not at all, or it is only with difficulty and loss, transformable into other forms of available energy . . (5) 44 THE MECHANISM OF LIFE And that is why there is always waste, or loss of available energy, and it is why the capacity for doing work tends to diminish indefinitely in the universe as we know it. Quantitative Energy Transformations. The following state- ments are the actual results of experience: (a) 1 horse-power = 33,000 foot-pounds per minute. That is to say, the amount of mechanical work that would be done against the earth's gravity when a mass of 33,000 pounds is raised 1 foot in one minute, or conversely the amount of work done by the earth's gravity when a mass of 33,000 pounds falls 1 foot in one minute, is called a horse-power. Long ago British engineers found, by actual experiments, the strength of an average cart- horse as measured in the above ways, and the quantity so found was, later on, adopted as a mechanical unit. (b) 1 h.p. =0-178 Calorie per second. This means that when the work done by a mass of 33,000 pounds in falling 1 foot in one minute is completely changed into heat (which it can be experimentally), 10-8 kilogram of water is raised 1 C. in temperature. (c) I gram of fat =$-3\ f . .j . ..Calories, when burnt in 1 gram of proteid =4-1 V J f , , , the body. 1 gram of caroohydrate=4:'l ) These are statements of the convertibility of chemical sub- stances into heat. They mean that when certain quantities of combustible materials are burned in oxygen, certain quantities of heat are generated. Many more instances might be given. The transformations are complete ones that is, all the available chemical energy transforms into heat energy. (d) 1 kilowatt=0-23S Calorie per second. That is to say, when a current is sent through a conductor against a resistance it transforms into heat. Some of the current (=1 kilowatt) disappears, and a certain quantity of heat (=0-239 Calorie) is developed. The heat developed is energetic- ally equivalent to the current that disappears. Now, since 1 kilo watt =0-239 Calorie per second, and 1 -h.p. =0-178 Calorie per second, it follows that (e) 1 kilowatt= 1-341 h.p. THE PRINCIPLES OF ENERGY 45 And by reversing statements (b) and (d) and (e) we get (/) 1 Calorie per second=5-62 h.p. (g) 1 Calorie per second=4:-18 kilowatt (h) 1 h.p. =0-746 kilowatt. But equations (e), (/), (g), and (h) are only true in theory, and they are not true in practice. They would be if equations (b) and (d) were reversible, which they are not. And that shows that we must make a very clear distinction between available energy (which is the capacity for doing work) and energy in the abstract, which is understood when we disregard experience and consider all transformations as reversible ones. Energy in the Abstract. When an energy transformation occurs, one kind of available energy is converted into other kinds ; for instance, so many cubic feet of coal gas are supplied to a gas engine, which then does mechanical work. Now the " calorific value " of the gas can easily be determined that is, we can find the quantity of heat into which the chemical energy of the gas is completely trans- formed when it is burned. So, also, the heat equivalent of the work done by the engine could be determined; for instance, the latter could be employed to lift a mass against the earth's gravity, and then this work can be represented as heat by using the mechanical equivalent in heat of this work done [equation (6), p. 44]. There will be a balance, for the heat equivalent of the gas is greater than the heat equivalent of the work done by the engine, and so available energy is lost. But we know very well that this available energy is really lost as heat radiated away from the engine, lost in the products of combustion (which are still hot when they are blown out into the air), and by other " leakages." And so the energy is not really destroyed, but simply dissipated. In many cases it can be traced, and where it cannot be traced we assume that it might be. This leads us to consider energy in the abstract without considering whether or not it is available, or possesses the capacity for doing work. First of all, however, we must make the concept of an isolated system. Let there be a large island which has absolutely no commerce or other communications with the rest of the world. It produces all that its inhabitants require, and it utilises itself all that it produces. It is self-sufficient, and is economically an 46 THE MECHANISM OF LIFE isolated system. So we might imagine an apparatus, or experi- mental plant, containing its own source of energy and absorbing itself all the energy given off, or wasted, or dissipated. That would be an isolated, energetic system. It is true that such an isolated physical system is a fiction, for there must always be some interchange of energy between any apparatus that we can devise and work and its surroundings; the only really isolated physical system is the entire universe. But let it be possible to take account of the energy lost by the system to without, or received by it from without; then we approximate to our concept of physical isolation. Making all allowances, then, for experimental error, friction, loss of heat by radiation, convection, and conduction, imperfect rigidity and elasticity of materials, etc., we can, very approxi- mately at least, trace the quantity of available energy that becomes unavailable, or is dissipated. Thus we arrive at our fundamental principle of energy: In all the transformations undergone by an isolated system the total quantity of energy contained is neither increased nor diminished . . . . . . . . . . (6) This is the law of conservation of energy, and it means that although the capacity for doing work that is, available energy diminishes, energy in the abstract does not diminish. When available energy diminishes, unavailable energy always appears in its place. Very often we can prove this. If we cannot do so we assume the law, and our assumption is always justified by other experimental results. But we are compelled, anyhow, to make the assumption, because the law of conservation is an a priori mode of our thought. To this matter we return later. We may now state the principal result of our work in the form of a quotation: " In all the transformations of a material system considered in this book, there is a certain entity which (1) remains constant in quantity, and (2) is capable, under certain conditions, of assuming the forms of kinetic and potential energy, which are dealt with under the study of Rational Dyna- mics. This Entity is Energy."* Several points arising from this definition must now be dis- cussed: first of all the nature of " Rational Dynamics." In this * G. H. Bryan, Thermodynamics, Teubner's Lehrbuchen der Mathe- matischen Wissenschaften, xxi., Leipzig, 1907. THE PRINCIPLES OF ENERGY 47 department of science we deal with, the behaviour of material bodies that " obey " Newton's laws of motion. There is no friction; bodies are perfectly rigid or perfectly elastic, and everything happens in an ideal world where all events can be described strictly mathematically. We do not consider heat at all, and energy is always mechanical, and is either potential or kinetic. There is no dissipation, and all transformations are reversible ones. In such circumstances " energy " is an entity that can be denned as above. Potential and Kinetic Energy. Sometimes available energy seems to vanish, although it is not converted into unavailable energy, or dissipated. Thus a grand- father's clock has stopped because its weights have run down, and we proceed to wind up the latter, but do not start the pen- dulum. In doing so we have expended muscular work, which is measured by the masses of the weights and the distance through which they have been raised. But the clock doesn't " go " until the pendulum is given an initial swing, and we have thus to account for the energy which we have expended. Now the system, clock and weights, and the earth, is in a different state from what it was before the weights were wound up, for the latter are now some 5 feet further away from the centre of the earth than they were when the clock had run down, and they are therefore free to fall. So long as the clock is not started, the weights possess potential energy that is, energy of position relative to something else. When the clock is started the weights begin to descend, and the potential energy begins to transform into kinetic energy, which is the energy of a mass in motion. Call M the mass of the weights and V the velocity with which they fall, and we get |MF 2 =the kinetic energy developed. During the time of fall the potential energy decreases, and the kinetic energy passes into the unavailable form, for the friction of the wheels, etc., generates heat, which is radiated away. When the weights have reached the bottom of the case they are no longer free to fall, and the system contains no more potential energy that is available for doing work at least, so far as the clock itself is concerned. Think about a mixture of oxygen and hydrogen in the pro- portions of one molecule of the former and two of the latter ; this system contains potential chemical energy. It will remain a 48 THE MECHANISM OF LIFE mixture of oxygen and hydrogen for an indefinite period of time, nothing happening in it. But let a spark pass through the gases, and they will combine together with an explosion and generation of heat, and the potential energy which they contained trans- forms into the kinetic form. Prior to the explosion the mixture might be represented thus : H 2 , H 2 , 2 , etc., the molecules of the gases (each of them consisting of two atoms bound together) being apart from, and not influencing each other ; but after the spark has fired the mixture we have H 2 0, H 2 0, etc., the molecules now occupying a different position relative to each other. Thus chemical energy is energy of position. Now before the explosion the mixture of hydrogen and oxygen possessed kinetic energy (as a mixture and apart altogether from the chemicaT^ature of its constituents). Each molecule was moving with a certain velocity, and from the temperature and pressure of the gas the average velocity of all the molecules can] be determined. The mixture can do mechanical work, for if it were allowed to rush into a vacuum it could drive in a piston just as steam drives in the piston in a cylinder. But as the] explosion occurs, the work that the gases can do is enormously increased (thus the explosion can propel a projectile if it is fired in a suitable vessel), and this energy that appears is the increased kinetic energy of the molecules of steam at a very high tempera- ture that is, the latter are now moving with much greater velocities than were the molecules of hydrogen and oxygen, and so they exert a greater pressure. The energy therefore becomes manifest, or visible, in its! mechanical effects; but where was it before the explosion occurred ? We cannot answer this question, and we assume; that it was not in the molecules of hydrogen and oxygen, but was in the medium between them that is, in the ether of space. The hypothesis is one that does not lead us astray, and there- fore we assume its validity or " truth." Available chemical energy, the energy of a weight raised above the surface of the earth, and free to fall when it is released, and that of a coiled spring these are examples of potential energy which transforms into kinetic energy, or the motion of bodies, when a releasing transformation occurs. THE PRINCIPLES OF ENERGY 49 Releasing Transformations. A word about this matter. The grandfather's clock, with its weights wound up, is at rest, but a very small expenditure of muscular effort will cause the pendulum to make a swing, and then the mechanism starts. The mixture of hydrogen and oxygen is also at rest, and nothing happens until a very small spark (involving a very small expenditure of heat) causes several molecules of H 2 and 2 to combine, thus liberating heat, which starts off other molecules, and so on, until the whole mixture fires in a very small period of time. A stone poised on the summit of a hill remains there until a very small push liberates it, and it starts to roll down, acquiring a high velocity, and becoming capable of doing much mechanical work. The coiled spring of a gramophone remains coiled until the touch of a little lever allows it to uncoil and actuate the mechanism. These are examples of releasing transformations. A relatively large quantity of energy is potential, and because of some condition of " false equilibrium," due to friction of some kind, it remains potential; but a relatively small expenditure of kinetic energy suitably applied upsets the equilibrium or overcomes the friction, and releases the energy which was potential. Such releasing transformations are of much importance in the discussion of organic activity. The Principle of Becoming. Taking a general survey of the results discussed so far, we may ask the question, Why does anything at all happen ? The enquiry is not so foolish a one as it may appear to a " prac- tically-minded " person, for a little consideration will convince such that if anything happens if there is " becoming " there must be an energy transformation. This is true, whether the event that happens is something very great and important from our point of view for instances, the formation of a huge sun- spot with accompanying magnetic storms, a cyclonic disturbance which wrecks a number of vessels, or a stellar collision and it would also be true of quite trivial things, such as the con- densation of a man's breath on the surface of a shaving mirror. Therefore our question becomes this, Why does an energy transformation occur ? And as such it is a legitimate subject for enquiry. 50 THE MECHANISM OF LIFE Now in every transformation three things are involved : 1. The intensity of the energy (intensity factor). 2. The quantity of the energy (capacity factor). 3. The direction of the transformation (the sign). These factors may be illustrated as follows : The System in which the Transformation occurs. Intensity Factor. Capacity Factor. A steam engine A water motor An electric motor or lamp A source of heat Pressure of steam Head of water Electric potential (volts) Temperature Quantity of steam Quantity of water Quantity of current (amperes) Quantity of heat In all these cases there is a system in which an energy trans- formation may occur; thus there may be a steam engine and boiler actuating some mechanism, a water reservoir and wheel, etc. Will a transformation occur in the system, and, if so, what will it do ? To be able to predict these occurrences we must be able to specify the conditions under which the system exists. The steam engine will " go " if the pressure of steam in the boiler is considerably greater than that of the atmosphere, and ' it will continue to go if this pressure is maintained that is, if new steam is generated as quickly as steam is drawn from the boiler. So, also, with the water mill : the motor will revolve if there is a sufficient head of water, and to keep it revolving the head must be maintained, or the quantity of water in the reser- voir must be very great, or must be renewed as fast as it is : depleted. If an electric current flows, there must be some way of raising electric potential, for a difference of this is the reason why the current flows. But the quantity of current is also a factor; thus the voltage of an electric stove might be the same as that of a lamp, but a greater quantity of current would be necessary in the former case (for heavier wires are employed). Finally, the temperature of a steam radiator 1 foot square might be the same as that of a radiator 6 feet square, but the quantity of heat distributed by the latter would be much greater. So there must be a difference of intensity if there is to be a transformation. If a hot-water radiator were at the .same temperature as that of the room in which it were placed, there THE PRINCIPLES OF ENERGY 51 would be no transference of heat, no matter what quantity of hot water were in circulation. Heat of itself flows from a hotter to a colder region that is, there must be a difference of intensity in order that the flow may occur. Why does a chemical reaction occur ? Why, for instance, do coal gas and oxygen combine, of themselves, to form carbonic acid gas and water ? The reaction is (assuming that the gas is CH 4 that is, marsh gas; it really contains hydrogen and other inflammable substances): CH 4 +20 2 =C0 2 +2H 2 0, and the reaction occurs as if the equation were written from left to right. Why from left to right, and not from right to left ? Marsh gas and oxygen combine to form carbonic acid and water, because in doing so a large quantity of heat is evolved; but if CO, and H,0 are to combine, a large quantity of heat (or more generally of energy) must be given to them, or be absorbed by them. Therefore they do not combine of themselves, although they can be made to do so. Chemical reactions occur, as a rule, only if heat is evolved, and so we may predict the occurrence if we know that heat may be generated. We cannot enter into the qualifications of this statement, but it may be made more general by saying that a chemical transformation will occur if in the occurrence work will be done. That is " why " it occurs, and if no work can be done by the chemical substances by reacting with each other they will not react of themselves. And if work is done as the result of the occurrence of a trans- formation energy will be dissipated, for all work done tends to Tansform into heat. But, again, all heat generated tends to listribute itself by conduction, radiation, and convection. It annot of itself accumulate in one place, as it tends to become uniformly diffused throughout the system the room, the world, and, finally, the whole universe. And so our experience is that n all transformations energy becomes dissipated that_Jis, ntensity differences become levelled down. Therefore we say hat an energy transformation occurs of if sp-lf _if f - energy can be issipated, or if intensity differences can Ke abolished. This is the sign of the transformation. Marsh gas and oxygen eact with the evolution of heat and the production of carbonic cid and water. But C0 2 and OH 2 do not react with the pro- uction of marsh gas, because heat would be absorbed in this 52 THE MECHANISM OF LIFE case; or work is done by the system (CH 4 +0 2 ) in the first case, but work would be done on the system (CO., and H 2 0) in the second case. The latter transformation does not occur unless there is a compensatory energy transformation, a matter to which we return later. Thus we obtain another principle: Energy transformations will only occur of themselves if energy r becomes dissipated that is, if intensity differences are diminished . . . . . . . . . . . . (7) The principles stated in the course of this chapter are, to some extent, repetitions, and they may be summarised and included in the two fundamental laws of science the first and second laws] of thermo-dynamics. These are: ^^ 1. The Energy of the Universe is Constant. 2. The Entropy of the Universe tends to a maximum* Now we may, very shortly, examine these statements. Matter, we have seen, can only be defined in terms of energy,; so the first law includes the old statements of the conservation matter and " force." But energy was defined on p. 46 as the entity that was con- stant in all transforming systems, and so there is something in universe that is constant. This something is energy. Th< fore the first law is a definition of energy in the abstract. Thinking about all this critically, the reader cannot fail to that the law of conservation is not true without some qualifica- tion. It says that there is an entity an existence in thej universe that can neither increase nor diminish. He notice that we have experience of nothing but energy trai formations, to which, it appears, all existences must reduce. Bi there are existences which arise from nothing, and are annihilal These are dreams, hallucinations, " spooks," etc. Our dm may be vivid and convincing, and " true while they last," am apparitions, if we may fully believe those who have seen them, can have all the appearances presented to us by material t Obviously disordered sense organs may mislead us, and we ma] hear noises and smell odours which we may be pretty sure d< not come from outside our own bodies. Then there is telepathy, mediumistic control, table-rapping, the planchette, etc., all * The concept of entropy is discussed in Chapter XI. THE PRINCIPLES OF ENERGY 53 them phenomena about which one hesitates to dogmatise. These things may be, as William James said, " the wild beasts of the philosophical desert," but still the^ are beasts ! We do not get rid of them by saying that they are purely subjective, for we do not seem to be quite clear as to what is subjective and what objective, and whether there is any difference ! But this is quite clear : dreams, apparitions, spooks, telepathy, and the like, are phenomena that are a nuisance, for they are existences that are not conserved, and so we cannot investigate them. They are not energy transformations, for if they Were they would continue in some other form after they had dis- appeared, and then we could count and measure, and describe and weigh them. We cannot do so, of course, and so most scientific men simply do not " believe " in spooks. They are existences, but they are not real existences. Real things are the things that are conserved. And so the law of conservation applies to some things and not to others, and the things to which it does not apply are unreal. That seems to be the best way out of the difficulty. Nothing that science can possibly do will disprove the law of conservation. Energy seems to vanish, but, if so, we only say that it has become potential. It may appear to come from nothing, and then we say that potential energy becomes kinetic, and we invent an ether of space to give us an abiding-place for potentialities. The discovery of radio-activity is a good example, and we may refer to it. A fragment of diamond can be made to burn, giving off heat, and transforming almost immediately into carbonic acid gas, which can no longer give off heat. A fragment of radium of the same size, however, gives off heat spontaneously, and continues to do so ; and a physicist who could observe it for a thousand years would find that it was still giving off heat, but that, like the bush that Moses saw, nee tamen consumebatur. To a chemist of the middle of the nineteenth century such a pheno- menon would have seemed to be almost miraculous, yet it is doubtful if he would have distrusted the law of conservation. He would, like Sir J. J. Thomson and Sir Ernest Rutherford, have invented a new kind of " bound energy." He would, like the physicists of our own time, have been justified in his faith. His ether and electrons and bizarre atomic systems are pure inventions, but they are real; they continually lead us back to order and not to chaos, and they are the means 54 THE MECHANISM OF LIFE whereby we make discoveries and obtain increased power over material things. They are concepts that can be investigated, inasmuch as they include the notion of conservation. All this goes to show, does it not, that the law of conservation is an a priori one it is a form of our thought ? Compare it with Kant's " first analogy of experience " : "In all changes of phenomena substance is permanent, and the quantum thereof in nature is neither increased nor diminished." Pheno- mena are, in so far as they are real, energy transformations, and their " substance " that which underlies them is energy in the abstract. If Kant's transcendental logic is valid, the first law is a mode of operation of the mind; it is something that makes experience by arranging sensations. And so we are not surprised when a physicist invents a potential energy to account for something that appears to arise out of, or passes into, nothing, and when his invention works out and leads to discoveries and practical applications. He acts upon nature because he has sensory data and " categories of the under- standing " mental operators, we shall say that deal with sensory data. It is because he acts that he has a mind, for the latter is the expression of his action. Since the law of conser- vation is a means of action, it follows, of course, that operations based upon it that is, potential energies, ether, electronic systems, etc. must " work " or have " pragmatic value." The second law is quite different : it has nothing at all a priori in it, as it is simply the resume of our experience. It tells us that entropy always increases when anything happens, and that means, for our present purpose at least, that water does not run up hill, a cold poker in a cold fireplace does not become red-hot, chairs and tables do not " levitate," etc. We can imagine all i these things happening, and if we think about it we do not see why they should not happen, except that they never have done so in our experience. With this observation, however, we defer the discussion of the second law till a later chapter. CHAPTER IV THE SOURCES OF ENERGY IT is clear that the animal has a double relationship with its environment. On the one hand it becomes aware, by means of its sense organs, of the changes that occur in the outer world. Events may happen there that menace it, and so there are places and things that are to be avoided; while other things present advantages food and shelter, for instance and these things it must seek and utilise. As the result of sensation and perception it acts upon the external world : its second relation to the latter. This acting involves mobility of body and limbs that is, the performance of mechanical work and we have next to enquire into the source of this kinetic energy, or vis viva, of the animal body. Common experience shows that it is obtained from the food that the animal eats, and so we regard this food as a store of potential energy which becomes transformed into the movements that make up the locomotory, defensive, and aggressive actions. The whole series of energy transformations falls into three stages : (1) The digestion, assimilation, and distribution of food material; (2) the oxidation of the assimilated food substances, with the liberation of the bound energy contained in it ; and (3) the excre- tion of the products of the process of oxidation. All this train of changes undergone by the substances that are taken into the animal body, built up to form its tissues, oxidised and excreted, is called metabolism. The Inanimate Engine. Now it will help us greatly in our study of the animal mechanism if we discuss first the way in which energy is transformed in the inorganic engine. In the most common form of the latter, the steam engine, there is a working substance, the water, which is heated in the boiler until it forms steam under pressure. This steam is admitted into a cylinder, where it is allowed to expand, thus forcing out the piston, and so doing mechanical work. It is again expanded in intermediate and low-pressure cylinders, thus doing more work, and by the time it has entered the COD denser, and attained 55 56 THE MECHANISM OF LIFE the temperature of the circulating water there, it has parted with all the energy which is available under the circumstances in which the steam engine works. The condensed steam is then returned to the boiler, reheated, and the cycle of operations recommences. Note that there is a source of energy contained in the coal, and that this energy is converted into heat, which is the kinetic energy of the molecules of steam. With each quantity of steam that leaves the boiler and enters the cylinders a certain quantity of energy is taken from the source (or coal) by the working substance (or steam), and some of this is given up to the condenser (for the water circulating in the latter becomes heated). But much more energy is taken from the source than is imparted to the condenser, and the difference is represented by the me- chanical work done by the engine. Thus, of the total heat generated in the furnace a certain fraction becomes transformed into the kinetic energy (or mobility) of the engine. The Animate Engine. Life in general that is, plant and animal life presents a series of events similar to that just described. There is a working substance which is represented by the very simple chemical compounds, water, carbonic acid, and certain mineral nitrogenous salts (which we shall call " nitrate " for short). These things correspond to the water employed in the steam engine. There is a source of energy corresponding to the heat generated in the steam boiler; this is the energy radiated by the sun (solar radiation we shall call it). Just as the burning coal imparts energy to the working substance of the steam engine, so the solar radiation enables green plants to manufacture carbohydrates, fats, and proteids from the water, carbonic acid, and nitrate supplied to them by the atmosphere and soil. Solar radiation acting through the green plants imparts energy to the working substance of life. The latter, in the form of carbohydrates, fats, and proteids, all of which substances contain large quantities of chemical energy, is then eaten by the animal organism, and, after the digestive changes, incorporated in the tissues of its body. In the course of the metabolic changes which it undergoes its energy becomes transformed. Steam at a high temperature (and with high intensity of energy) enters the cylinders of the inanimate engine, and leaves the condenser at a relatively low temperature (and thus with low intensity of energy). In much the same way the life working substance enters the animal body while con- THE SOURCES OF ENERGY 57 taining much free chemical energy, and leaves it in the form of the excretions while containing almost no free chemical energy. In the steam engine the difference between the quantities of energy possessed by the working substance when entering and leaving the mechanism is represented by the mechanical work done and the heat lost by radiation, etc. Similarly, the difference between the energy of the working substance (as the food) when it enters the animal body and when it leaves it (as the excretions) is represented by the mechanical work done by the animal, and by the heat radiated away from its body and lost in the excretions. The working substance, now degraded in respect of the energy it contained, is taken up again by the green plants, reconverted into carbohydrate, fat, and proteid, and the cycle of operations recommences. The Digestive Process in Animals. In the meantime, we consider only the animal part of the life energy cycle. The food matters exist, and the animal obtains these by the exercise of its sensori- motor organs. But these food matters must be digested, distributed to the tissues of the body, assimilated, oxidised, and then excreted. It is this train of events that we have now to study. Why must they be digested ? The foodstuffs must enter into the blood-stream of the animal that eats them, and in order that this may happen they must be dissolved. Now, as a rule, the food of an animal is a complex and heterogeneous collection of substances, how complex every healthy man and woman knows. In the animal living in the wild it is even more complex, since such creatures usually hunt down, kill, and devour other animals, and often ingest flesh and bone, fur, feathers, scales, hair the edible as well as the inedible parts, which latter civilised man tries to reject in the processes of the preparation of his food. But even when the latter is carefully prepared and cooked it still contains inedible parts the cellulose or woody fibre of vegetables, fruits, cereals, peas, and other plants, and the fibrous substance of meat and certain fats which are not easily dissolved. All these inedible constituents of the food are excreted in the faeces, and, of course, they vary in different animals: thus her- bivores can digest cellulose, while man cannot as a rule. Trituration, or mechanical disintegration of the substance of the food, occurs mainly in the mouth. Finely divided particles 58 ' THE MECHANISM OF LIFE are thus produced, so that the digestive juices may act all the more easily. These digestive juices play the principal part in the reduction of the food into such a form that it can enter the blood- stream, and we must say something about them. There are certain glands in connection with the alimentary canal (the mouth, oesophagus, stomach; and intestine), and their function is to elaborate liquids which issue from them through ducts, and are conveyed into the cavities of the mouth, stomach, and intestine. The salivary glands in the mouth secrete saliva, the gastric glands of the stomach manufacture an acid liquid called gastric juice, the liver prepares the bile, and the pancreas the pancreatic juice. All along the intestines there are glands which also secrete a digestive juice. These various juices are active in virtue of certain mysterious substances called enzymes (or ferments) which they contain. The saliva contains ptyalin, the gastric juice pepsin, the pan- creatic juice trypsin, and the intestinal juices erepsin, etc. The bile secreted by the liver is active mainly in virtue of the alkaline substances that it contains, and the juices of the intestine contain a substance called enterokinase, which " activates " the erepsin. What these enzymes are we do not know in the least, for they have never been prepared in a pure condition, and they are pre- sent in their respective juices in very small quantity. But we know a very great deal about what they do, and there are many inorganic substances which have very much the same properties, so that we cannot say " for certain " that the digestive enzymes are, in any way, substances that are exclusive to the living organism. The Foodstuffs. To understand their modes of operation we must consider the chemical composition of the foodstuffs. Anything that we eat consists of a mixture of substances called proteids, carbohydrates, and fats. White of eggs (albumen) is a nearly pure form of proteid containing much water. The lean flesh of all animals is mainly proteid, while the substance of peas, beans, lentils, cheese, and parts of the cereals and of the solids of milk are proteid in nature. Carbohydrates are represented by cane-sugar and glucose, the starchy foods, such as rice, the substance of potatoes, the main constituents of cereals, the sugar of milk, etc. Fats are the fatty parts of flesh meat and the massive fat that occurs in most animals, the cream of milk, butter, margarine (which is now mainly " hardened " or " hydro- THE SOURCES OF ENERGY 59 genated " vegetable oils, plant oils such as olive, cotton- seed, palm, etc.). Proteids are innumerable, and the flesh of almost every species of animal contains distinct kinds. They are the most complex of all chemical substances, and it is only during the last quarter of a century that they have been successfully investigated (mainly by Fischer and the German chemists of his school). All of them contain the chemical elements carbon, hydrogen, oxygen, and nitrogen, united together in a very complex way. They are now known to be built up of peculiar substances called amino- acids, and one of the simpler examples of the latter has the following formula and name: '>CH CH 2 CH(NH 2 ) COOH. Leucine or a-amino-iso-caproic acid. Very many of these amino-acids are known, but many more still have yet to be investigated. Now -imagine a number of these united together in this way : R' R" R'" NH CH CO - NH CH COOH. (2) (3) Here we have a chain of members, each of which has the form : R -NH CH CO ; the N, H, C, and each represent an atom of nitrogen, hydrogen, carbon, and oxygen respectively; and R represents a complex " radicle " or group of atoms such as (in a simple case) : FCH 3 CH 2 CH 2 . In the example we have given the chain consists of only three members, but there may be very many more. It may be bent round to form a ring, and the ring may be joined to other rings, and there may be " side chains " attached in various ways. Now compare all this with the chemical formula of urea (which is the characteristic substance present in the urine): NH 60 THE MECHANISM OF LIFE and we get an idea of the exceedingly complex chemical structure of the proteids a structure upon which, in some way or other, the phenomena of life depend. Proteids are therefore combinations of amino-acids, and they contain, generally, about 16 per cent, of nitrogen. Carbohydrates contain carbon, hydrogen, and oxygen, but no nitrogen. They are also very complex chemical substances; thus " grape- sugar " (dextrose, or glucose) has the formula: CH.OH CH(OH) CH(OH) CH(OH) CH(OH) CHO. Many other sugars are more complicated, and the starches and gums are more complicated still. Fats are still more complicated in chemical structure than the carbohydrates. What we usually call an " oil " or a " fat " is a combination of one or more fatty acids with glycerine. A fatty acid consists of carbon, hydrogen, and oxygen atoms united together to form a chain; thus stearic acid is: CH 3 CH 2 15 other CH 2 links COOH, and may be written in short by R COOH, where the R stands for the chain CH 3 CH 2 to seventeen terms. Now three molecules of stearic acid combine with glycerine to give us R.-CO CH 2 I R CO CH R_CO CH 2 which is tristearin, the principal constituent of mutton fat. Since there are a great number of different fatty acids there must also be many kinds of fats. Now all this seems very technical, but it must be clearly under- stood that no one can hope to get even a slight acquaintance wi- the mechanism of life without a knowledge of at least such detai as we have set out above. And, after all, it is not very difficult ! What we eat is therefore a mixture of proteids, carbohydrates, and fats. The proteids, especially when cooked, are insoluble in water, and so are the fats. We can take cane- sugar, and other sugars, into the alimentary canal, but they must be converted into dextrose before they can be used by the tissues. The starches which form the bulk of potatoes, rice, and similar foods, as well as a large part of cereals and the breads that are made from these : ail THE SOURCES OF ENERGY 61 things, are also insoluble, and they must be converted into dextrose. Digestion consists, therefore, of the solution of the proteids and fats, and the conversion of the starches and other carbohydrates into dextrose. As soon as food enters the mouth digestion begins. The salivary enzyme, ptyalin, acts on the starchy substances, con- verting them into sugar, and in the stomach the pepsin of the gastric juice acts upon the proteids. The fats are not touched at all until the food enters the duodenum, and then the main diges- tive operations begin. The trypsin of the pancreatic juice and the erepsin of the intestinal juice act energetically on the proteids, carbohydrates, and fats, and the bile secreted by the liver aids in a way that we cannot explain here. The result of the action of all these enzymes is that the proteids are split up into their constituent ammo-acids, the starches are converted into dex- trose, and the fats and oils are split up into fatty acids and glycerine. Now all these substances are soluble in water (or at least in the liquid bathing the internal walls of the alimentary canal), and so they can soak through into the blood. These substances amino- acids, fatty acids, dextrose, and glycerine are what the animal organism wants they are its proximate food principles. Absorption and Distribution. They must get into the blood- stream and be carried to the muscles, glands, and other tissues of the body. How ? That leads us to consider the organs of circulation. Everywhere in the body there are minute blood- vessels, called capillaries, forming a close network, and along with these there is a separate system of vessels called the lymphatics. We do not consider the latter here, except to say that ultimately they communicate with the bloodvessels. We regard the bloodvessels (from our point of view of the distribution of nutritive matter) as beginning in the walls of the alimentary canal. There the capillaries form a very close network just within the internal lining (the mucous membrane), and the liquids in the alimentary canal are separated from that in the capillaries by the mucous membrane, some very loose connective tissue, and the very delicate walls of the capillaries themselves. So it is easy for the digested, soluble food substance to soak through the mucous membrane, the loose submucous layer, and the walls of the capillaries. Now the " soaking " through is not the same process as the soaking of a few drops of 62 THE MECHANISM OF LIFE soup through a piece of porous blotting-paper; in the latter case the liquid soup goes through unchanged, except for the solid particles, which are kept back by the blotting-paper, while the living cells of the mucous and other membranes act on the digested food matters. We have seen that the digestive enzymes change the composition of the proteids thus: Proteids > amino- acids. But the same enzymes are contained in the cells of the mucous and other membranes, and as the amino-acids pass through External layer , (Serous coat) y Artery Jtfi/COUS membrane Capillary network in submucous /ayer Inferior Canal (ein Muscular layers FIG. 12. PART OF A TRANSVERSE SECTION THROUGH THE WALL OF THE INTESTINE. VERY DIAGRAMMATIC. they are again acted upon by the enzymes thus : Amino-acids > proteids. Thus the activity of the enzymes is reversible. Now it is no use presenting all this to the reader as a finished picture of the processes of digestion and absorption of the food substances, for we have only the vaguest notion of what is the nature of the reversibility; one thing is, however, certain the proteids are split up into amino-acids in the cavity of the alimentary canal, and as these substances are soluble they can pass through the intestinal wall into the capillary blood-stream. In passing into THE SOURCES OF ENERGY 63 From , Fo re- Ifm b s head and. neck The Caval veins iver the tissues which utilise them, the same enzymes now reconvert the amino-acids into proteid. But the proteid is now a different one (or ones), for the amino-acids have been rearranged. Somewhat similar rearrangements take place in respect of the fats. The blood circulating in the intestinal wall thus becomes charged with proteid, fat, and carbohydrate, all three classes of substances being now thrown into forms which are " native " to the body of the animal in which the digestive pro- cess occurs. One eats the proteids characteristic 70 of beef, mutton, pork, fowl, fish, etc., but all these become converted somewhere into human proteid, and all the different fats that one eats are similarly con- verted into human fat. That is, the processes of digestion, absorption, and assimilation are a laborious and round- about series of chemical conversions which make up a large portion of our vital activity, and which might be shortened with much advantage. Little by little the capillaries in the walls of the stomach and intestine unite together to form veins, and the smaller veins unite further until one large vessel the portal vein drains away all the blood circulating in the alimentary canal from the stomach backward. This great vein carries the intestinal blood, laden with nutritive substance, to the liver, and there it divides up again into small and smaller veins, and these finally break up into capillaries which ramify among the cells of FIG. . 'a.//menTa.ry Canal From other Viscera. Prom hind limbs and- Trunk 13. DIAGRAM OF THE GREAT VEENS IN THE BODY OP A MAMMAL. 64 THE MECHANISM OF LIFE the liver. Something is done to the blood there, and then the' capillaries reunite into veins and then into another great vessel] the hepatic vein. At the same time the blood which has been circulating in the: capillaries of the muscles of the hind-limbs and trunk, in thd kidneys, reproductive organs, spleen in short, ia all the body ; behind the upper limbs is gathered up into a vein called the^ posterior vena cava, and this is joined by the hepatic vein. All the blood that has been circulating in the fore-limbs, the chest, head, and neck is collected by two veins called the anterior vena- cavce. Finally, all three caval veins unite and pour their blood into the heart. , To muscles of- From muse let / of trunk FIG. 14. DIAGRAM OF THE BLOODVESSELS OF A FISH. i, ii, iii and iv are the gills. Thus there is a vascular system, which consists of a series of veins returning all blood that has been circulating anywhere in the body to the heart. Clearly there are at least two kinds of blood in the veins: (1) That which has been circulating in the muscles, and which has, therefore, been depleted of some of its nutritive properties; and (2) that which has been circulating in the intestine, and which has become enriched with substances of nutritive value. Now we have to consider the other half of the blood vascular system the arterial vessels which distribute this nutritive substance received from the alimentary canal (via the' liver) to the body at large. First of all we look at the arterial vessels of a fish (for here the conditions are very simple). Much the same kind of venous system is present in the fish as in man (or at least we need not worry about the minor differences), but the arteries are not complicated by the presence of lungs. THE SOURCES OF ENERGY 65 The fish respires by taking oxygen into the blood through the gills, and also by giving out carbonic acid in the same way (we return presently to this respiratory process, and only consider just now the path of the blood). In the fish, then, the heart is a fairly simple organ consisting of two main chambers, the auricle and the ventricle. Each of these is a muscular, hollow organ which expands and contracts rhythmically. The blood which has returned from the body via the caval veins is poured into the auricle, which then contracts. There are valves at the entrance to the auricle and others at the opening of the latter from the Proper right. To body Lung Proper left. FIG. 15. THE CONNECTIONS BETWEEN THE HEART AND THE LUNGS IN A WARM- BLOODED ANIMAL. into the ventricle, and these structures so act as to prevent the blood from being forced back into the caval veins, but they allow it to flow into the ventricle. There are valves at the opening of the ventricle into the aorta (that is, the great artery that springs from it), and these act so as to allow the blood to pass through the aorta when the ventricle contracts. Thus the contractions and dilatations of the heart send the blood in one direction only from the caval veins into the aorta. From the latter vessel it goes to the gills, where the arteries break up into networks of minute capillaries. Then the latter vessels reunite until they form several large arteries, two of 6 66 THE MECHANISM OF LIFE which go to the head, several to the " shoulders " of the fish, and one runs down the body just underneath the back- bone and supplies the viscera and the muscles of the body and tail. Thus there is a fairly simple and complete circulation in th< fish. The blood is propelled by the heart into and through th( gills, and then it is distributed through the arteries to all parts of the body. Having traversed every part of the latter, 11 returns to the heart again via the great veins. The reader will now easily understand the main scheme o: circulation in the warm-blooded animals, including man. Here we have a double heart, one half of which (the right one) is connected with the lungs, and the other (left) half with the resi of the body. Tracing out the paths taken by the blood-stream, we will see that the caval veins discharge into the right auricle, which con- tracts and sends the blood into the right ventricle. From there it goes through the pulmonary arteries into the right and lef lungs, and having traversed the capillaries in these organs, it is returned to the left auricle by the pulmonary veins. The lef auricle forces it into the left ventricle, and from there it is propelled all over the body, being distributed by the great aorta and the arteries that branch out from the latter. The first complete demonstration of the circulation of the blood must have appealed to physiologists as a perfect proof o the mechanical conception of life. This proof, however, was slow in coming, and many men contributed to it. Servetus, scholar of the early sixteenth century, seems to have discoverec the pulmonary circulation, and, curiously enough, he announce it in 1553 in a theological work called De Restitutio Christianismi for the publication of which he was hunted out from Spain b] the Inquisition, only to encounter the intolerance of Calvin a Geneva, where he was burned. The English physician Harve; discovered the other, or systemic circulation, and gave a com plete and formal demonstration of the whole scheme in 162 Fabricius had already discovered the valves in the veins, an< in 1661 Malpighi found the connection between arteries ancl veins the innumerable, minute, hairlike vessels called thej capillaries and that completed the proof. But even before thej latter discoveries the great mind of Descartes had made use oi the incomplete demonstration of Harvey to establish his conJ THE SOURCES OF ENERGY 67 ception of an automatic, mechanical, animal body, and no such momentous contribution to a working hypothesis of life had before then, or has since, been made. The Respiratory Interchange. Thus food materials are taken into the alimentary canal, digested, and transferred to the blood- stream in such a form that they can be assimilated by the tissues. They are then distributed in the blood-stream to all parts of the body. But simultaneously there is an interchange of something between the blood itself and the air that is inspired by the lungs. Here we mention, for the first time, the arterial and venous kinds of blood. That which issues from a cut vein, and in general from a slight wound, is blood that is rather dark red in colour, and which oozes out from the cut vessels, whereas that Artery X7ab///ary network Air Vesicle FIG. 16. AN ALVEOLUS, OR AIR SAC, FROM THE LUNGS, WITH ITS BLOODVESSELS. HIGHLY MAGNIFIED. which comes from an artery is brighter red in colour and it flows ut in pulsations. Examining these bloods, physiologists have ound that that flowing in the arteries contains more oxygen and ess carbonic acid gas than that flowing in the veins. Examining tie air that is inspired, they find that this contains the usual 1 per cent, of oxygen and about -^ per cent, of carbonic acid, while the expired air contains only about 16 per cent, of oxygen nd about 4 per cent, of carbonic acid. Both the composition of the blood and that of the air are hanged in the lungs. Now look at the structure of the latter: ach consists essentially of an immense number of air vesicles urrounded by capillary bloodvessels. This is a scheme of the essential structure of the lung. The rachea divides to form the two bronchi, and the latter divide nd subdivide again and again until they terminate in multitudes of sacs, or vesicles, one of which the greatly simplified figure 68 THE MECHANISM OF LIFE represents. The pulmonary artery (carrying venous blood) enters the lung, and also divides and subdivides again and again until, in the end, each ultimate twig of the artery breaks up into a network of capillaries on the outer wall of an air sac. Air is drawn up into the lungs and expelled out from them by the upward and downward movements of the diaphragm (the muscular partition between the chest and abdominal cavities) in men, or by upward and downward movements of the ribs and chest wall in women. Thus the sacs are full of air which is continually being renewed (we do not really draw air right down to the " bottom " of the lungs, but into the upper parts only). The air of the smallest sacs renews itself, however, by diffusion into the fresh air that is continually entering the larger passages. Now the venous blood that enters the air-sac capillaries contains (in solution) less oxygen and more carbonic acid than that which ^ FIG. 1 7. BLOOD-CORPUSCLES. 1, White; 2, human; 3, human in " rouleaux "; 4, human, seen from the side; 5, from a frog; 6. from a fish. is contained in the air of the sacs, and therefore an interchange occurs such that oxygen goes through the very delicate membrane of the sac and the equally thin wall of the capillaries from the air into the blood, while carbonic acid gas passes from the blood into the air. In respiration, then, we take oxygen from the outer air anc transfer it to the blood-stream, and we take carbonic acid froi the blood-stream and transfer it to the air. And so the venoi blood, which goes to the lungs and comes from the body, contains more carbonic acid and less oxygen than the arterial blood, which comes from the lungs and goes to the body. Just a word as to how the oxygen and carbonic acid are carried by the blood. The latter, when seen under the microscope, becomes a clear, colourless liquid in which there are enormous numbers of minute, biscuit-shaped bodies called the red blood- corpuscles. THE SOURCES OF ENERGY 69 These contain a chemical substance called haemoglobin, which carries the oxygen, while the carbonic acid is carried by the clear liquid part of the blood, or plasma. In all animals there are other corpuscles which are colourless, and some of these are called phagocytes (devouring cells) because they have the power of ingesting foreign substances, such as bacteria, which may enter the blood. The pus which forms when an abscess gathers consists largely of phagocytes. They are a protection against infection. We have now seen how the nutritive materials the proteids, fats, and carbohydrates and the oxygen which is to " burn " them, are obtained, and how they are carried to the tissues. The Sources of Energy. These proteids, fats, and carbo- hydrates, digested, dissolved, split up, and recombined, are the immediate sources of energy (the ultimate source is, of course, the solar radiation, which enables the green plants to synthesise water, carbonic acid, and nitrate into proteid, fat, and carbo- hydrate). They contain potential, chemical energy which is at a high intensity, and is available for doing work. How ? In the inanimate engine there is a working substance, the steam, and we regard this as something different from the engine, which may exist even if there is no steam in it. But in the animate engine the mechanism (muscle and nerve) are the same as the working substance (the proteids, fats, and carbo- hydrates, which are the muscle and nerve). It may be the case that a " living " muscle contains " non-living " substances which are oxidised and yield energy, but since we only define the " life " of the muscle by its irritability (that is, its ability to contract- when it is stimulated), we cannot be sure that there are parts of it w^hich are not alive. It is probable that the substances carried to the muscle by the blood-stream are actually built up into the living tissue they become alive and are then oxidised, or die. Further, the inanimate engine steam, gas, or petrol is a thermodynamic machine. It contains a working substance steam at a temperature, say, 120 C., or a mixture of gases resulting from the explosion of gas and air, or petrol vapour and air at a temperature of, say, 700 C. to over 1,000 C. At these high temperatures the molecules of the gases are moving with enormous velocities, and so they exert pressure. Therefore a heat engine is a mechanism which converts the kinetic energy of 70 THE MECHANISM OF LIFE a gas at a relatively high temperature into the kinetic energy of a material structure (pistons or turbines) at a relatively low temperature. It may be that the muscle of an animal is a heat mechanism of this kind a thermodynamic machine but it is not likely, and it is far more probable that the muscle contracts because the chemical energy of the proteids, fats, and carbohydrates is trans- formed into mechanical work without passing through the stage of heat. There are physical mechanisms that do this. A galvanic battery may produce a current of electricity, and the latter may work an electric motor and no heat at all (or only a trace of it) may be generated. Here chemical energy (that of the zinc and acid in the battery) transforms into electricity, and the latter does mechanical work in the motor. Now all that we can state positively about the events that happen when a muscle contracts are these: there are complex, chemical compounds in the protoplasm of the muscle which break down (or " explode "). Very complex molecules thus dissociate into relatively simple ones, and the energy that held these complex molecules together is liberated and is transformed into the mechanical work done by the muscle when it contracts against a resistance. But it is quite certain that heat is also generated that every- one knows who becomes warm when he does hard muscular work. Now, is this generation of heat a necessary step in the doing of bodily work ? In cold-blooded, wholly aquatic animals it is not certain that heat is produced, or, if it is, it is only in very small quantity. A warm-blooded animal preserves a constant tem- perature, which is usually considerably higher than that of its environment, and so it must generate heat. And it is probable that this heat is produced by the oxidation of the relatively ! simple molecules into which the complex protoplasmic substance breaks down. These products of dissociation are oxidised into water and carbonic acid. Protein- containing tissues break down into the same products, and also into simple nitrogenous substances that later on become urea in the liver and kidneys that is, into such substances as can easily be removed from the muscle in solution in the circu- lating blood. The Removal of Waste Glandular Activity.- In the heat engine the working substance steam in the steam engine, or the products of the explosion in the internal combustion motor THE SOURCES OF ENERGY 71 loses its available energy and must be removed. The steam is passed out from the condenser, and is returned to the boiler to be reheated, and the cylinder gases in the gas engine are blown out into the atmosphere. So also in the animate engine the working substance loses its available energy: protoplasmic substances disintegrate into simpler ones, which are then oxidised to form carbonic acid and water. These latter compounds, with the urea, which is the end product of the disintegration of the nitro- genous part of the protoplasmic substance, must be removed from the animal body. So we must say a word or two about excretion. The water which appears in the muscles as the result of the disintegration ~Glandu/a.r pa.rtoF kidney ,- Cavity FIG. 18. THE KIDNEY OF A MAMMAL. A, The organ with its bloodvessels and duct; B, the kidney cut through longitudinally. of the protoplasm, or of the oxidation of the products of dis- integration of the latter, simply oozes through into the blood- stream, and the carbonic acid similarly produced is also taken up by the blood. The effect of work done by the body is there- fore to add water and carbonic acid to the blood, and so the latter becomes changed, " impure," or venous. As it streams through the capillaries in the lungs the carbonic acid is excreted, passing out into the expired air, while, at the same time, some of its excess of water also passes out in the same way. But most of the water, and also the urea, is excreted by the kidneys. Consider first of all the urea. The proteid substance of the muscles is, as we have seen, continually disintegrating into simpler substances, giving up energy as it thus breaks down. These products of metabolism are not yet urea, but they are 72 THE MECHANISM OF LIFE carried to the liver, where they are converted into this latter substance. The blood circulating in the body therefore contains a certain small proportion of urea. This is the waste product of nitrogenous metabolism ; it has no useful function, and it must be removed from the system. We consider the kidneys in some detail, because they may be taken as types of glands, structures which we have not, so far, described. Each of them, then, is an organ provided with (1) an aiteiy carrying blood into it from the aorta ; (2) a vein carrying blood Surface oF fridn* v Cavity of the kidney FIG. 19. ONE OF THE SECRETORY UNITS (OR URINIFEROUS TUBULES) OF THE KIDNEY REPRESENTED IN A VERY DIAGRAMMATIC WAY. MAGNIFIED. away from it into the posterior vena cava ; (3) a duct, the ureter, which carries away the water and other sub- stances taken from the blood that flows into the gland ; and (4) nerves. When in action, blood contain- ing urea and other waste substances, as well as an excess of water, is continually flowing into the kid- neys through the renal arteries, while the same blood deprived of these waste substances and of a certain quantity of water is continually flowing out through the renal veins. Water containing the urea, etc., gathers in the cavities of the kidneys and slowly trickles down the ureters into the urinary bladder, from which it is periodically expelled. The nerves that enter the kidney go to the arteries, and they act by exciting the muscles in the walls of these vessels to expand or contract, thus altering their calibre, and so increasing or diminishing the quantity of circulating blood. If the calibre of the renal veins remains the same, the blood-pressure of the kidneys increases, and so the secretion of THE SOURCES OF ENERGY 73 urine increases, and vice versa. Thus the secretory activity of the kidneys can be regulated. Similar regulatory mechanisms exist everywhere over the animal body, and the nerves mainly responsible belong to what is called the sympathetic nervous system. Now look at the minute structure of the kidney. Here we represent, very simply, and on a much bigger scale than the natural one, one of the tubules that excrete the water, urea, etc. First of all the blood in the renal artery becomes distributed through a multitude of branches or twigs of this bloodvessel, and we may regard each twig as going to a peculiar structure called a glomerulus. Capillary knot x Outer (Jill of glomerulus FIG. 20. THE GLOMERULUS OF A URTNIFEROUS TUBULE, WITH ITS BLOODVESSELS. HIGHLY MAGNIFIED. This glomerulus is like a bulb at the end of a urinif erous tubule. )ne part of the bulb is pushed in, and a twig of the renal artery enters and breaks up into a little complex knot of capillaries, from which a twig of the renal vein gathers up the blood and takes it away. This little twig of the renal vein again breaks up into capillaries, which surround the rest of the urinif erous tubule, as shown in Fig. 19. Thus we see a rich blood-supply carried by capillaries surrounding the parts of the gland that secrete. AVater is secreted from the blood in the capillaries of the glomeru- lus, and this water oozes through the inner wall of the bulb, and so goes into the uriniferous tubule. Then the cells of the latter take urea, etc., from the blood in the capillaries that sur- 74 THE MECHANISM OF LIFE round them, and these waste substances go with the water into the cavity of the kidney, and so down the ureter into the bladder. Note that the essential things in this mechanism are the cells of the walls of the glomerulus and tubules. It is these cells that have the power of taking water, urea, hippuric and uric acids, etc., from the blood; but they do not take sugar, albumen, or other things. If they do (as in diseased kidneys), that is not their proper function, and the organ assumes an individuality that is harmful to the organism a disharmony is established. How exactly the cells of the normal kidney function in removing waste substances and nothing else we do not know. This is a type of glandular activity, and there are numerous other organs in the body that do analogous things. The gastric glands secrete hydrochloric acid and pepsin into the stomach; the pancreas secretes a mixture of ferments or enzymes called trypsin; the glands of the mouth secrete saliva; those of the skin secrete sweat, water, oil, etc. (all waste substances), and so on. In all these cases there are tubular mechanisms (as a rule much simpler than those of the kidney), and these are provided! with nerves and arteries and veins in much the same way. There! are also ducts, tubes which carry the products secreted by thd glands to the places where they are wanted, or via which they are] eliminated from the body. But there are ductless glands also. Such are the thyroid, the thymus, the pituitary, and pineal glands (see p. 99), and the adrenal glands. In such cases we have glandular cells surrounded by capillaries, but there is no duct and there is! apparently no secretion. We know, however, that there is a secretion, and that this goes into the blood itself, and is then carried to the rest of the body. These ductless glands are of! enormous importance in many ways. The reader will miss the true understanding of vital activity] if he does not note the character of unification of activity that is implicit in our conception of the animal mechanism. This chapter and the last one deal with mechanisms: the sensori-motol system that is, sensory organ connected with motor organ; the mechanism of alimentary canal and glands that converts the] crude food substances into the specific proteids, fats, and carbo- hydrates that are required by the animal tissues as sources of energy and means of growth; the apparatus of circulation that^ distributes these substances; the respiratory mechanism that THE SOURCES OF ENERGY 75 obtains the oxygen that must be distributed along with the food- stuffs; and the excretory organs that remove the degraded food substances and expel them from the body. It has been necessary to consider these things separately, but that is only our method of analysis of a single phenomenon, and it must be repeated that all these activities are co-ordinated and are really one. Nothing in the way of mechanisms that we can make or devise exhibits the unification of activities expressed in the life of a normal animal. The modern State which has, rather foolishly, been called an organism is said to represent the greatest achievement of men, but no one can attempt to analyse its activities dis- passionately, as we have analysed those of the living animal, and see in them anything other than a tissue of disharmonies. The scientifically organised State has still to come into existence. Think of the distribution of foodstuff to the functioning animal body. In severe manual work the bloodvessels of the muscles dilate so as to permit of a more abundant supply of nutritive material and oxygen, and a rapid removal of waste products. In intensive mental work the blood-supply to the brain is similarly speeded up. In digestion the same thing occurs with relation to the intestinal circulation. Such regulations are as yet hardly at all evident in the activities of the modern State. CHAPTER V ON VITAL PRODUCTION WE have now traced the working substance of life through the transformations which it undergoes in the animal body. These are as follows: Animal Metabolism. (1) Certain food substances contain the working substance in the form of a mixture of proteids, fats, and carbohydrates, derived from the tissues of animals and plants. This mixture is taken into the alimentary canal, where it is digested and dissolved. As the result of the processes of diges- tion and absorption the proteids, fats, and carbohydrates of the food are made similar, or assimilated, to those substances such as they occur in the body of the animal that eats them. (2) They are then distributed by the blood-stream and incor- porated in the protoplasm of the muscles and other tissues. (3) These protoplasmic substances are disintegrated into simpler chemical compounds, with the result that energy is liberated. (4) The products of the disintegration of the substances of the tissues are finally oxidised and excreted. The result of these metabolic processes is that the working substance becomes degraded energetically and chemically. Let us take the energy transformations first. An ordinary day's diet is likely to contain about 125 grams of dry proteid, 125 grams of dry fat, and 400 grams of dry carbohydrate,* and if the heat value of these quantities of dry foodstuff be found it will add up to about 3,500 calories. Now the result of the chemical transformation undergone by this average quantity of food is that a certain quantity of carbonic acid and water is excreted by the lungs and kidneys, and about 42 grams of urea by the kidneys. The water and carbonic acid contain no free (or available) energy, but the urea contains about 105 calories. Therefore about 3,390 calories of the energy contained in the food is utilised by the body. About 12 to 30 per cent, of this * In British weights about 4 ounces of dry proteid, 4 ounces of dry fat, and 14 ounces of dry carbohydrate. 76 ON VITAL PRODUCTION 77 energy is transformed into mechanical work done by the muscles of the body and limbs, and the rest is transformed into heat. That is the energy transformation. Assume an adult animal in perfect health and of stationary weight; then the working substance enters the tissues in the form of proteid, fat, and carbohydrate, and leaves them in the form of water, carbonic acid, and urea. That is the chemical transformation. Looked at both from the point of view of available energy and that of chemical structure there is degrada- tion. First, in that a high intensity of chemical energy becomes a low intensity; and, second, inasmuch as complex chemical compounds become simple ones. The working substance of life is now water, carbonic acid, and urea. Let us trace its further history. The water and carbonic acid undergo no further change (just yet, at all events). But the urea (and the other proteid degradation products) do. The Fate of the Proteid Residues. The urea (and other nitrogenous substances) contained in the urine find their way, through drains and by other means, into the water of rivers and of the sea, or on to the land, and then it is at once attacked by micro-organisms. Now a few words about the latter. Micro-organisms are (in the present connection) bacteria, moulds, yeasts, and infusoria. The bacteria are exceedingly minute, single-celled organisms which cannot be said to be either animals or plants, since their modes of generation are altogether special. Many of them (but still a small minority) are called pathogenic organisms, and are the causes of certain infectious and epidemic diseases, such as cholera, enteric fever, pneumonia, " influenza," diphtheria, catarrh, septicaemia, etc. Being exces- sively minute, capable of living in water, and even of being partially dried, they may be distributed in liquids, in dust, on infected clothing, etc. When they enter a suitable " soil " that is, a liquid containing certain organic substances in solution they multiply at an incredibly high rate. The human body possesses certain defences against these pathogenic bacteria that is, the phagocytes of the blood can ingest and destroy them, or the plasma can form certain substances which can neutralise their poisonous activity. If the infected animal fails to set up such adequate defences, the invading pathogenic bacteria multiply with great rapidity and form toxins which are injurious tojme or more tissues, and so instigate a condition of disease. 78 THE MECHANISM OF LIFE Other micro-organisms have, on the whole, beneficial effects.! Yeasts are the causes of the fermentations in brewing and analo- gous processes. Moulds and bacteria cause the ripening of cheese. Fermentative and putrefactive bacteria act upon and break down organic matter, and moulds and infusoria behave in \ similar ways. Many substances that we meet with everyday "go bad": milk curdles; broths, soups, etc., "go sour"; and meat of allj kinds becomes tainted and then putrefies. Beers and wines may sour. In the case of meat and fish, the " ripening " (as in thej case of game) and the tainting are the result of the activity of I bacteria and of enzymes that are naturally present in the flesh; itself. When the flesh is alive, the activity of these enzymes are] inhibited in some way, but after general death of the tissues they ; begin to digest the flesh in which they occur. This process is| called autolysis, or self-digestion. At first it produces savoury,! and then noxious substances. By-and-by the meat putrefies, with the production of offensive odours, and this is the result of the action of bacteria which) infect the decaying substance. If this activity is allowed to go on unchecked, the offensively smelling meat begins to disappear, I and by-and-by the smell itself passes away. Any organic substances exposed to the air or contaminated with dust, " dirt," or impure water, will " go bad," ferment, ori putrefy, and this is because fermentative and putrefactive: bacteria gain admittance, multiply, and break down the organic matter, using the latter as a source of food. If rigorous and successful means be taken to exclude all micro-organisms from the organic matter, the latter will not go bad. Thus if milk be " pasteurised " (that is, heated for several days in succession in closed bottles to a temperature of about 80 to 90 C.), it will remain fresh and sweet for an indefinite period. If meat be sterilised by heating to over the boiling temperature of water in sealed tins for a sufficient period, it will be preserved in good condition for years. If meat be frozen to well below zero centi- grade it will remain good. If salt, boracic acid, formaldehyde, chlorinated water, etc., be added to the organic substance, putrefaction will not occur. Heating to a sufficiently high temperature kills all micro-organisms, as also does antiseptic substances like salt and boracic acid. Freezing arrests the multiplication and vital activities of bacteria, but does not kill ON VITAL PRODUCTION 79 them, so that frozen meat will putrefy if it thaws. This is the rationale of all processes of preservation of meat and other food substances: heating destroys the "germs"; salting and drying and freezing arrest their activities; and rigorous exclusion of air, dust, and other media which contain the organisms may delay or prevent the putrefaction. Filtration of water through porous earthenware keeps back the germs, and exposure to strong sun- light, or, better still, to the rays from an electric mercury vapour lamp, is said to destroy them. The Mode of Action of Micro-Organisms. In fermentation and putrefaction much the same chemical processes occur as when food substances are digested in the alimentary canal of an animal. In fact, the micro-organisms form enzymes similar in effect to the enzymes that are found in the stomach and intestine. The enzymes split up proteids into amino-acids and ferment carbo- hydrates, but the processes of disintegration of proteids and carbohydrates go much further in bacterial action than in the digestive operations. These processes are very complicated, and are far from being fully understood. We call the breaking-down of fats and carbohydrates by bacteria, yeasts, and moulds fermentation, and that of proteids putrefaction. Many species of bacteria, etc., are concerned in each process, and the rapidity with which the latter occurs depends on the temperature and other conditions. The final results are quite clear and well known. In all cases, and if there is time enough, fats, carbohydrates, and cellulose, such as the vegetable substances in grains, grass, leaves, fruits, woody fibre, etc., are decomposed, with the result that their chemical substance transforms into carbonic acid and water. Putrefaction of proteid substance is accompanied or succeeded by what is called nitrification that is, other organisms, called " nitrifying bacteria," also play their part. In the end the proteid putrefies to form ammonia compounds, and then the nitrifying bacteria oxidise the ammonia, converting this into nitrous acid and the nitrous into nitric acid. The lime, soda, and potash in the soil or in streams, rivers, etc., combine with the nitric acid to form nitrate, and this is the way in which Chili saltpetre and other natural stores of nitrate have been formed. Thus the action of micro-organisms on all dead organic matter is to convert the latter into carbonic acid, water, and nitrate. This is, in general, the fate of the excretions of animals and of all 80 THE MECHANISM OF LIFE dead animal and plant bodies, " offal," or remains. All organic substances whatever are susceptible to bacterial action, and since micro-organisms are universally distributed in nature, ini the air, soil, and in fresh and salt water, organic matter thus* suffers resolution into innocuous mineral substances of very. simple chemical constitution. There are certain conditions im which this bacterial decomposition of organic matter is delayed. At the bottoms of deep oceans the temperature of the sea is very* 1 little above freezing-point (or is even below the freezing-point of! fresh- water), and there putrefactive action goes on very slowly., In extreme northern climates the air temperature is also very low, and dead bodies of animals are sometimes frozen in ice on in the soil, and so escape decay. In very dry climates organic: matter also remains in a relatively stable condition, since the 1 putrefactive bacteria are unable to function in the absence of! water. Such is the mode of origin of many forms of guano. But, in general, all organic matter is ultimately resolved into carbonic acid, water, and nitrate, and these substances tend to be distributed all over the earth. Carbonic acid is contained to the extent of about 0-04 per cent, in the atmosphere, and it is present in rather larger proportions in fresh and salt water. Water itself is distributed everywhere except over desert land areas. Mineral nitrogenous substances, such as nitrites, nitrates, and salts of ammonia, are also contained everywhere in the soil and in fresh and salt water, and oxides of nitrogen, capable of forming nitric acid, are formed in the atmosphere by the combination of nitrogen and oxygen that occurs whenever there are lightning discharges. But mineral nitrogenous compounds which are the indispensable materials of life occur everywhere in exceed- ingly small quantities,* and the abundance of life depends on the quantity of these substances that is available. It is therefore convenient roughly to classify all living tilings into animals, bacteria, and plants. The animals consume proteids, fats, and carbohydrates, oxidising these substances into water, carbonic acid, and certain nitrogenous residues such as urea, uric acid, hippuric acid, etc. The bacteria act upon the nitrogenous residues, converting them into nitrate, and they also act upon dead animal and plant tissues, and upon the excreta * Except under quite special conditions, as when deposits of " Chili saltpetre " and analogous substances are formed. Here the dry atmo- sphere and soil inhibit plant growth, with the result that nitrate accumulates. ON VITAL PRODUCTION 81 of animals, resolving those substances into water, carbonic acid, and nitrate. The plants then utilise the latter substances as crude foodstuffs. Plant Metabolism. We have now to consider the further history of the working, substance of life after it has undergone the chemical and energetic degradations that are the results of animal and bacterial metabolism. Returning to our inanimate engine, it may be recalled that the working substance, or steam, expands and does mechanical work on the pistons, and actuates the mechanism. Then it passes through the condenser, having lost its available energy. It is returned to the boiler and is heated, and so takes up fresh available energy, and the cycle of operations recommences. We take the same general view of the animate engine. The working substance, which is a mixture of fats, proteids, and carbohydrates, passes through the animal body, undergoing chemical transformations, doing mechanical work, and (it may be) heating the body. Then it passes out from the body as the excretions, having lost most of its available energy, and it is further acted upon by bacteria, when it loses the re- mainder. It must now be transformed so as to reacquire avail- able energy, just as the cold water entering the steam boiler again takes up energy in the form of heat. This absorption of energy by the life working substance is effected in the tissues of green plants, and we must now refer briefly to the metabolism of the latter. A green plant is an organism that is, something that trans- forms energy of itself, grows, and reproduces its individual form and mode of behaviour. Its growth is an obvious thing, and so it is clear that it absorbs material from the medium in which it lives. Now, with rare exceptions* the plant organism does not take in visible foodstuff that is, we can see no obvious process of feeding, mastication, digestion, etc. We know that it can only grow under certain conditions a plentiful supply of water to its roots and air to its leaves and it must also have free access to sunlight. In the failure of such conditions the plant does not grow. Its foodstuff is necessarily contained in the air surrounding its green leaves and in the water of the soil that contains its roots. Examining the air contained in an enclosed space, we find that * Insectivorous plants. 82 THE MECHANISM OF LIFE this consists of about 79 per cent, of nitrogen, 21 per cent, of oxygen, and a trace of carbonic acid. If we next examine the air of an isolated space which has been used by plants, we shall \ find that its percentage of carbonic acid has been greatly dimin- ished, while that of oxygen has increased. Evidently the effect of plant life is to rob the atmosphere of its carbonic acid and to enrich it with oxygen just the opposite to that of animal -, life. If, further, we proceed to examine the water w T hich is taken up by the roots, we shall find that this is not pure, but always con-1 tains mineral substances like nitrates, chlorides of potash and| magnesium, salts of iron and lime, phosphates, silica, sulphates,] etc., and it can be proved that the effect of plant metabolism is to take such substances from the soil water. The materials from which the green plant builds up its tissues are therefore water, carbonic acid, nitrates, and other simple mineral substances. These materials it converts into proteids, oils, and waxes, and carbohydrates such as starch, sugar, and; cellulose. Now all of the latter substances contain much poten- tial energy, for, if we dry them, we can burn them and so obtain heat. But the water, carbonic acid, and mineral salts cannot be oxidised further, and they contain no available energy. In building up its tissues the green plant must therefore obtain energy from somewhere, and it can be proved that it is obtained from light ; if the plant is kept in the dark, there can be no growth.* But it can grow in the light radiated from an electric arc It is, in fact, fairly easy to show that a green leaf exposed to sunlight is continually forming starch, while, if it is kept in the dark, no such thing happens. Here, then, we have the source of energy of the green leaf. Outside the latter is water (OH 2 ), carbonic acid (C0 2 ), and solar] radiation; and inside it is the mixture of pigments called chlorophyll. The solar radiation is not heat, although it isj transformed into heat when it impinges upon most material] objects. When it falls upon certain substances, such as luminous paints, it is transformed into light, and in certain conditions it can be transformed directly into mechanical work. When it- impinges on the green leaf and is absorbed by the chlorophyll pigments, it is immediately transformed into chemical energy;] for it can be proved that there is no starch in a leaf which is kept 1 * No increase in mass, although a seed a potato, for instance may germinate in the dark. ON VITAL PRODUCTION 83 in the dark for'some time, while within a few minutes of its exposure to sunlight starch accumulates in the cells. It is difficult to convince the non-chemical reader what a very extraordinary thing this process of photo-synthesis of starch by the green plants must be. Let him note that on the one hand there is water and carbonic acid, and on the other there is dex- trose and finally starch. The chemical equation is (probably) : 6C0 2 + 6H 2 = C 6 H 12 6 -f 60 2 . Carbonic acid Water Dextrose Oxygen Then the dextrose is converted into starch (C 6 H 10 5 )n, and the latter gathers in the cells of the leaf till it is required by the plant, when it passes into solution as dextrose, and is removed from the leaf by the circulating sap juices. This is the process of photo-synthesis. Look, however, at the first equation and read it from right to left. There is dextrose and oxygen. Dextrose is a highly combustible substance, and when it burns it combines with oxygen to form C0 2 and OH 2 , with the libera- tion of a large quantity of energy in the form of heat. What actually occurs in the plant, however, is represented by reading the equation from left to right, and this shows that when the C0 2 and OH 2 are synthesised in the plant to form dextrose, the same quantity of energy must be absorbed as is liberated when dextrose burns in oxygen to form C0 2 and OH 2 . The former kind of reaction an endothermic one can be made to occur. A chemist can decompose C0 2 into C and 0, as, for instance, when a burning piece of magnesium ribbon is placed in a jar containing C0 2 . But the C0 2 is only decomposed into carbon and oxygen if a large quantity of energy is supplied in the form of the heat liberated by the burning magnesium, and so the decomposition only occurs at a relatively high tempera- ture. The following statement is very important in all such discussions as these : reactions like that in which C0 2 is decom- posed into C and 2 only occur when a compensatory energy transformation is brought about that is, when at least as much energy is supplied from outside as would be yielded if the reaction went in the opposite direction. Such reactions an endothermic \ process coupled with a compensatory one do not occur of \ themselves. When they do occur, the coupling is the consequence V of some outside agency. In the green plant, however, it is precisely this coupled reaction 84 THE MECHANISM OF LIFE that occurs. C0 2 and H 2 are combined together, and the energy necessary to bring about the combination is taken from the sunlight. A chemist could cause the combination to take place at a very high temperature and in quite special conditions, but it occurs in the green plant of itself, and at ordinary temperatures. All the above is very important, and may not be neglected when we are considering the nature of the living process. We return to the matter in a later chapter. The Balance of Life. We see now in what way the energy degraded by animal life becomes rebuilt up again. The animal takes proteids, fats, and carbohydrates into its body, disin- tegrates and oxidises these compounds, makes use of their con-' tained energy to obtain heat and do mechanical work, and then excretes the products of disintegration and oxidation in the form of water, carbonic acid, and urea. The bacteria convert the urea (and other nitrogenous residues) into nitrate. The green plants take energy from solar radiation, and rebuild water, carbonic acid, and nitrate into proteid, fat, and carbo- hydrate, when the cycle of operations recommences. Now it is easy to see that all animal life depends upon plant life. Animals are carnivorous (feeding upon flesh), or her- bivorous (feeding upon vegetable substances), or omnivorous (eating both flesh and vegetable substance). There are also some animals which are called saprozoic, with regard to their manner of nutrition, and these can utilise as food liquids or debris containing broken-down organic matter. Here we need not consider the saprozoic organisms, and it is enough for our present purpose to regard animals as either carnivorous or herbivorous, or both. Since the carnivores eat other carnivores or herbivores, or both, and since the herbivores eat vegetable tissues, it is easy to see that all animal life on the earth ultimately depends upon vegetable life. On the other hand, vegetable life depends for its continuance on a supply of water, carbonic acid, and nitrate. The quantity of i water on the earth is (for our purpose) unlimited, but not so the ; quantity of carbonic acid and nitrate, and upon the supplies of the latter substances the abundance of vegetable life hangs. What, then, are the sources of carbonic acid and nitrate ? Home quantity of each substance comes from the earth in the exhala- tions from volcanoes, and possibly as the results of the disintegra- tion of certain mineral substances. There is probably far more ON VITAL PRODUCTION 85 CO., in the atmosphere and dissolved in fresh and sea water than the plants are able to utilise that is to say, there is a surplus of both water and carbonic acid on the earth. But this is not the case with nitrates and other inorganic nitrogen compounds, which are quite indispensable for the nutrition of plant life. There is an enormous quantity of ele- mentary nitrogen in the atmosphere, but this is unavailable, for, in order to utilise it, plants must have it in combination with oxygen (as nitrous and nitric acids), or with hydrogen (as ammonia). Some nitrogen is always being combined with oxygen as the result of electric discharges in the atmosphere, and even under the influence of solar radiation ; but, on the other hand, some nitrate, nitrite, and ammonia are always being decomposed into elementary nitrogen by certain bacteria present in water and soil. Therefore the total amount of nitrogen in the forms of nitrate, nitrite, and ammonia, and thus available for the nutrition of plants, is nearly constant, and certainly does not change appreciably during very long periods of time. That means that the total average quantity of vegetable life on the earth is practically the same from year to year. And that being so, the total average quantity of animal life on the earth is also practically the same from year to year over a great lapse of time, for all animal life depends for its continu- ance on vegetable life. There is a certain balance between the two kingdoms of life. In restricted areas of land and sea, and for restricted periods of time, either the plants or animals may predominate; but in the long-run there is a relation between the two masses of animal and plant substance, and this relation is a nearly constant one. If animal life becomes temporarily very abundant, vegetable life must decrease, because it is being used up to a greater extent than is normal, and this condition will in turn lead to a diminution of animal life. Production and Consumption. The reader will now see what is meant by " vital production." All animals eat organised substance fats, proteids, and carbohydrates contained in the tissues of other animals and plants. They reduce these sub- stances to the forms of water, carbonic acid, and certain nitro- genous residues, utilising the contained available energy for the production of mechanical work and heat. Then the working sub- stance is thrown out of circulation and is no longer available for the sustenance of the animal organism. The latter is a consumer. All the time solar energy is, so to speak, running to waste. 86 THE MECHANISM OF LIFE Falling upon the sea, it evaporates water, which rises up into the air as vapour, falls down as rain and snow, and returns to the sea in rivers, having done nothing of itself but wear away the land and generate heat by friction. The heat is radiated away into space, and is for ever lost to the earth. Falling upon rocks, stones, gravel, sand, soil, etc., and upon the atmosphere, the solar energy heats up all these substances to a slight degree ; butj again this heat is radiated away, and is therefore irretrievably lost. Apart from vegetable life, there is therefore a continual dissipation of solar energy. But the green plants intervene. They absorb the solar radia- tion, utilising this to recombine the water, carbonic acid, and nitrate which result from animal metabolism. Apart from the activity of the plants, this energy would be dissipated and the products of animal life would remain unavailable for further life. They are, however, recombined by the plants, and the solar energy which would otherwise be wasted is thus fixed. The plants are producers. In the past plant life has, on the whole, been more active than animal life, and the result is the accumulations of coal (and perhaps oil) upon which modern industrial civilisation is based. This civilisation we must, however, regard as merely an episode in the history of terrestrial life, for the enormous increase on human population during the last few centuries has only been possible by the utilisation of the materials produced by a former surplus of vegetable life. With the depletion of this surplus the balance will be restored. It would, of course, be very rash to| regard the future reduction of the human population of the earth as inevitable, for there may be other immense accumulations of i energy in that now bound in the atoms of some material sub- stances. Also some supply of energy may be obtained from the tides, winds, and rivers. But, again, it would be very foolish to^ count upon this possibility, for all that we know so far about radio-active transformations suggests that the process is one] which we cannot initiate or control, and that the energy of Corpus obfic striatum Calamus FIG. 25. FURTHEB STAGE IN THE DEVELOPMENT OF THE BRAIN. The primitive lower brain in a fish, such as the cod, is very simple (Fig. 26). The mid-brain, with its great optic lobes, is the most important part. The fore-brain is the part lying between the two corpora striata. The cerebellum is not greatly developed. In its general characters such a brain as that which we have just described represents what we shall refer to hereafter as the " lower brain " of the higher mammal. Imagine the thin membranous " pallium " which covers the corpora striata to become enormously thickened and to grow backwards over all the rest of the brain, and suppose the cerebellum greatly to increase in mass relatively to the other parts. Then we should have the brain of the mammalia" n animal. BRAIN AND NERVE 97 The Human Brain. The human brain is represented in a very diagrammatic way in Fig. 27. It is supposed that the whole organ has been cut through in a mid-vertical plane, and a number of details that are rather difficult to understand have been omitted from the diagram. The parts actually cut through are cross-hatched, and we see the central canal of the spinal cord widening out to form the " fourth ventricle " of the anatomists that is, the primary hind-brain vesicle. Between this and the " third ventricle " that is, the I Olfactory Sbinal ' cord. K.Optic(eyes) JJf Oculomotor (muscles ofeyesj )Y,Trochlea.r ( M uscles of eyes) -Yf.Abducens , (muscles of eyes) Y- Trijgeminal (facet-Jan*) yM Auditory (ear) facial .ossof)ha.ryn$ eal (Mouth ( pharynx., X. Vagus (Gills, heart**) ifia/ nerves FIG. 26.- THE BRAIN OF A FISH (COD) SEEN FROM THE UPPER OR DORSAL SURFACE. primary fore-brain vesicle is a narrow passage which is the " aqueduct of Sylvius," and this represents the primary mid- brain vesicle. The floor, sides, and roof of the hind-brain vesicle are enormously thickened to form the cerebellum and the peduncles which connect it with the other parts of the brain. Similarly the roof, sides, and floor of the mid-brain vesicle have become thickened to form the corpora quadrigemina, and the great crura, or peduncles of the cerebrum, and some other parts. The roof and floor of the " third ventricle " remain thin, but the 7 98 THE MECHANISM OF LIFE lower parts of the sides are thickened to form the optic thalamij one of which is represented in the lateral wall in the figure. The enormous lobes, called the cerebral hemispheres, are the roofs of the primary lateral brain vesicles thickened to form the cerebral cortex with its systems of projection nerve fibres. In one corner; of the third ventricle a little opening, called the foramen of MonroA is shown, and dotted lines indicate how this leads into a cavity in each cerebral hemisphere, called the " lateral ventricle." The] cross-hatched part, called the corpus callosum, is the junction of Junction of R. and L hemispheres Corpus , Sthatum " n ng ^Kl. inTo " 1st. ventricle L atera I ventricle _ _ ^Pineal body - Cor bora. --Cerebellum -Pituitary ', body I 7eduncle of the cerebrum cord. FIG. 27. THE HUMAN BRAIN: AN IMAGINARY SECTION ALONG THB I MIDDLE PLANE. The cross-hatched areas represent the cut surfaces; the numbers I, lit and IV, show the lateral, third and fourth ventricles respectively. the two (right and left) hemispheres. Just round the foramen] of Monro the corpus striatum of the right-hand hemisphere is shown. This ganglion appears to bulge into the third vontricloj but it really belongs to the lower part of its cerebral hemisphereJ Because of the enormous growth backwards of the latter orgaM the relations of the various parts of the brain become difficult to visualise. Thus the corpus callosum appears to form thl roof of the third ventricle, but this is not really the case. The- reof consists of a delicate vascular membrane, which is now easily representable in a diagram, and the corpus collosmn, with; some other parts which are omitted, are the sections of the BRAIN AND NERVE 99 of the two cerebral hemispheres which have coalesced by their internal faces. The reader must not omit to note the two little bodies that are attached to the roof and floor of the third ventricle. That one attached to the roof is called the pineal body, and it was here that Descartes placed the seat of the soul. It is really a " ves- tigial organ " which has acquired a new function. Early in the history of the primitive vertebrates there were either one or more " cyclopean " eyes, and even in some of the lizards such eyes exist, though they are never functional organs of vision. In modern mammals the pineal body has become a ductless gland that is, an organ that forms some substance which is discharged into the blood-stream. What the substance is we do not know, but it is said that it exercises an inhibitory in- fluence, restraining precocity of growth and a too early maturity of the reproductive organs. The organ on the floor of the third ventricle is called the pituitary body, and it also is a vestigial structure consisting of two parts, one of which appears to have formed the primitive vertebrate mouth. The pituitary body is a ductless gland secreting some substance into the blood which inhibits or controls the growth of the skeleton, particularly the bones of the face. Removal of the gland is always a fatal operation, and disease or hypertrophy produce curious exaggerations of growth, some of which, it has been noted, recall in bizarre fashion the charac- teristics of the extinct Neanderthal human race. Connections within the Central Nervous System. The white tracts in the spinal cord, in the peduncles of the cerebellum, and in the parts called the cerebral peduncles or crura in Fig. 27 are the great paths along which nervous im- pulses travel within the central nervous system. -It is known that there are, in the white matter of the cord, two main cate- gories of nerve fibres: (1) such as convey impulses to the brain ascending tracts; and (2) those that transmit impulses from the brain to the grey matter of the cord these are the descending tracts. Starting, then, with the white matter of the cord, we may next consider the main paths along which impulses travel in the central nervous system. The Sensory Tracts. First we take the ascending paths, those along which impulses originating as the results of stimula- tion of the receptor organs (see p. 106) reach the lower brain. 100 THE MECHANISM OF LIFE All sensory stimuli from the skin, the muscles, joints, and viscera pass into the spinal cord via the dorsal (or sensory) roots ofi the spinal nerves, and either go directly up to the brain or they enter the grey matter and undergo " relays "* there that is, they are shunted on to new paths. Some of them pass out from the cord via relays and the motor roots, when reflex actions occur' (see later), but the others travel up to the brain along tracts in the white matter. The great ascending tract in the cord is the! bundle marked " posterior tracts " in Fig. 23, and this is made up mainly of nerve fibres which start from the cells in the grey! matter round which the sensory root fibres terminate. The upward path of these bundles is represented very diagram-; matically in Fig. 28 as " great sensory tracts from cord to medulla." The individual fibres run up into the medulla, and end] there as synapses round cells in two centres or nuclei, thus enter- j ing into a second relay. From these centres two new tracts,- called the lemnisd, start. Immediately after leaving the nuclei 1 the lemnisci cross or decussate that is, the fibres starting from> the right-hand nuclei cross over to the left-hand side, and! vice versa. Kunning up through the crura of the lower brain, they end in the corpora quadrigemina and optic thalami, ancfl this is their third relay. From these mid-brain nuclei new tracts of fibres start again, and proceed upwards into the cortex. Thus the stimuli originating in the sense organs of the skin on the trunk and limbs, and in the muscles, joints, and viscera of] those body regions, pass into the spinal cord, ascend the latteri to end in the medullary ganglia. From there they travel to the! mid-brain ganglia on the opposite sides of the brain, and from] the mid-brain they proceed to the cerebral cortex. It is mostly the impulses giving rise to the sensations of touch, heat and cold, j and pain originating by stimulation of the skin that take this very complicated path. The impulses coming from the receptor organs in the muscle and joints mostly take a different path. They enter the greyl matter of the cord via the sensory spinal nerve roots as before, J and end in cells from which new ascending fibres start, or they may travel up in the white matter of the cord without first undergoing a relay in the grey matter. But in either case thefl take an independent course, and proceed to the cerebellum, e * A " relay " is an interruption in a nervous tract. One set of neuroneH join on to a new set by means of synapses. 102 V1J.S MECHANISM OF LIFE entering the latter via the inferior peduncles. These are the tracts shown in Fig. 28, and called " afferent cerebellar tracts." Thus the receptor organs in the trunk and limbs are connected with the ganglia contained in the mid-brain and cerebellum. This is also the case with the great receptors in the head the visual, auditory, and gustatory organs; with the touch, heat, and cold receptors in the skin of head and face; and with the muscular and articular receptors in the same region, but not with the olfactory receptors. With the latter exception, all the nerve fibres starting from the sense organs in the head and face pass into the medulla via the cranial nerves, and end in separate nuclei. The latter are then connected in complex ways with the corpora quadrigemina, the optic thalami, and the cerebellum. The fibres carrying impulses from the olfactory organs are con- nected directly with the cerebrum ; later on we shall consider the connections of the great sense organs in more detail. Connections of the Cerebellum with the Other Parts of the Central Nervous System. The cerebellum of a higher mammal is connected with the cord, mid-brain, and cortex by three great pairs of tracts contained in the cerebellar peduncles. The inferior peduncles are connected with the white and grey matter of the cord in the way just suggested, the superior peduncles are connected with the mid-brain (mainly corpora quadrigemina and thalami), and the opposite sides of the cerebellum are joined together by the middle peduncles which form what is called the pons varolii. Thus the cerebellum has a grip, so to speak, on the nuclei or ganglia which receive all the sensory impulses coming from every part of the body whatever. Further, since the grey matter of the cord and the nuclei of the cranial nerves are ganglia from which motor fibres go out to the muscles of body, limbs, and head, the cerebellum has also a grip on the centres controlling movements. The Connections of the Cortex Cerebri with the Rest of the Central Nervous System. The cerebral hemispheres are, we have seen, parts of the central nervous system which are super- added to the lower brain, inasmuch as each of them is to be regarded as a great lobe growing out on each side of the original fore-brain vesicle, expanding enormously, and arching over all the other parts of the brain. The base of each hemisphere is formed by the great ganglia called the corpora striata, and these come into close relationship with the ganglia of the fore- brain BRAIN AND NERVE 103 that is, the optic thalami. These two pairs of basal ganglia are present in the brains of all vertebrates, and we are regarding them here as parts of the " lower brain." The roofs and sides of the lateral fore-brain vesicles are either not present at all as nervous grey matter, or are imperfectly developed in the lower vertebrates, and they become very important only in the mammals. There they form a relatively thin sheet of grey matter which constitutes the superficial part of the hemispheres, and this increases so greatly in area that it becomes crumpled and folded in a complex way, so that it may be contained within the cranial cavity. The crumpling leads to the formation of the cerebral convolutions. There is therefore a thin layer of grey matter the cortex cerebri on the surface of each hemisphere, and beneath this there are great bundles of nerve fibres running in various ways. Be- neath these bundles of white matter, again, are the basal ganglia. The white matter of FIG. 29. A SECTION THROUGH A CEREBRAL HEMISPHERE NEAR THE MIDDLE PLANE TO SHOW THE MAIN COMMISURAL TRACTS. the cerebral hemispheres form two main categories of bundles the commissural tracts and the projection tracts. We have dealt with the latter in Fig. 28. The commissural tracts are represented in Fig. 29. One system of commissural tracts, represented chiefly by the corpus callosum, connect together the right and left halves of the cerebrum. Probably corresponding parts of the two hemi- spheres are joined by the fibres of the corpus callosum, but it is also possible that any one part of one hemisphere is connected with most parts of the other. In addition to these transverse commissural tracts there is an elaborate system of internal commissural or *' association " tracts in each hemisphere, and some of these are represented in Fig. 29. They connect together the various convolutions, or groups of convolutions, by short paths, while longer tracts connect distant parts of the cortex with each other. More and more, as cerebral physiology de 104 THE MECHANISM OF LIFE velops, do the interactions of one part of the cortex with other parts by means of the commissural and association tracts come to possess significance in the development of the manifestations of intelligence and higher mental faculties.J From all parts of the cortex bundles of fibres radiate inwards towards the cerebral peduncles; these form the "projection tracts " those that connect the cortex with the lower brain and spinal cord. Some of the projection tracts are represented in Fig. 28 in a very schematic way. One, which is called the " great sensory tract," connects the parts of the cortex lying behind the Rolandic fissure (see Fig. 38) with the mid-brain ganglia; another tract of fibres, starting from the cortical region well in front of the Rolandic fissure, travels downwards and ends in the grey matter of the pons varolii, where it forms a series of relays with the fibres coming from the two sides of the cerebellum via the middle peduncles of the latter. This is the " tract from the cortex to the pons and cerebellum " of Fig. 28. Thus the cortex is connected with the lower brain on the one hand by a direct tract, and with the cerebellum on the other hand by two indirect tracts, one via the mid-brain and the other via the pons and the middle peduncles. The most important of all the projection tracts in man and the higher mammals is the pyramidal tract, that represented in Fig. 28, and called " the motor tract from the cortex to the cord." The fibres composing it start in the peculiar pyramidal cells of the cortical region lying in front of, and immediately round and in the depths of, the Rolandic fissure that is, in the " motor area " of the cortex. These fibres are gathered up into two bundles which travel down in the crura of the brain towards the medulla, where they decussate, or cross, to the other side of the body. The crossing is not, however, complete, as the figure shows, and some of the pyramidal fibres remain on the same side of the body as that in which they originate. But most of them cross over. Running down in the white matter of the cord are therefore two descending tracts of fibres, " the direct and crossed pyramidal tracts." All these fibres pass into the grey matter of the cord, and end there as synapses round the cells that give off the fibres of the motor roots of the spinal nerves. The pyramidal cortical tracts, with their nuclei in the cortex, are the typically " higher " parts of the mammalian brain. They are relatively small in such animals as the mole or the rabbit, BRAIN AND NERVE 105 and more developed in the dog, and still more in the monkeys. In the anthropoid apes and in man they attain the maximal development, and are to be regarded as the paths along which those impulses travel that are the stimuli to what we call " willed " or spontaneous actions and movements. This is a very summary account of the main features of the gross anatomy of the central nervous system, and we deal in greater detail later with the special nervous mechanisms. The reader should now be able to visualise the whole as a series of parts developing progressively in the course of the evolution of the vertebrate animals. Primarily there was a double series of ganglia, one pair in each segment of the body, and all of them were connected together by transverse and longitudinal com- missural tracts. Nerves issuing from the ganglia were dis- tributed to the sensory and motor organs of the body, conducting inwards impulses originating in the stimulation of the receptors, and conducting outwards impulses setting the muscles in move- ment or causing glands to function. With increasing complexity of bodily structure and greater freedom of movement the ganglia increased in mass and began to coalesce, thus forming the con- tinuous core of grey matter of the cord, the hind, mid, and fore brain. The great sense organs became concentrated in the head, and so the cranial ganglia increased in functional importance, finally assuming the conditions that have been described. There has been a progressive complexity in the movements included in locomotion as we ascend the series of evolutionary phases represented by the fishes, amphibia, reptiles, birds, and mammals. The need for precise adjustment and co-ordination of the impulses issuing from the spinal cord and setting muscular organs in motion therefore led to the development of specialised ganglia carrying out such functions, and so the cerebellum assumed the anatomical importance that it obviously has in the human brain. And in the later stages of this evolution that is, in the mammals a kind of activity, connoted by the terms " intel- ligent," " spontaneous," and " volitional," became the charac- teristic one exhibited by these animals. This, we shall see, involved a nervous mechanism other than those of the mid- brain and cerebellum, which are to be regarded as the ganglia controlling movements and activities that are largely " auto- matic." This mechanism, the latest one to be evolved, is contained in the cortex cerebri and its connections. CHAPTER VII THE SPECIAL NERVOUS MECHANISMS THE very general survey that we have just made of the rough anatomy of the central nervous system will enable the reader to study in greater detail the more special mechanisms into which we, rather arbitrarily, decompose the whole. These mechanisms are those of sensation, of motor control, and of co-ordination. The Sensory Mechanisms. " Sensation " involves the stimulation, by some physical agency, of receptor organs distributed everywhere in the body. We must suppose that all bodily tissues are irritable that is, that they react in some way to stimuli, which may be chemical or physical. From what we know of the irritability of the sur- face tissues of the lower organisms, we may also conclude that this is a general irritability, and that the same tissue is potentially susceptible to light, electric, chemical, and mechanical stimuli. But in the higher animals the general irritability of the tissues of the lower organisms becomes modified by the evolution of the special organs of sense. We must think of this generalised susceptibility of the skin, say, as being restricted, so that any one kind of stimulus must be intense enough to pass over a "threshold"; if it is too feeble, the receptor is not stimulated at all. A special sense organ, therefore, is a part of the skin or other tissue where the height of the threshold is reduced for stimuli of some particular nature, and raised for all others. Thus the retina is a highly specialised part of the embryonic outer surface, which is extremely sensitive to the stimulus of light, but is relatively insusceptible to changes of atmospheric pressure, while its situation is such that it is not usually stimu- lated chemically, electrically, or mechanically. Similarly, the auditory hairs in the cochlear part of the internal ear are highly susceptible to sound vibrations occurring in the atmosphere outside, while they are so sequestered that they are not readily exposed to stimuli of any other kind. The nerve terminations of the olfactory and gustatory nerves in the mucous membranes of the nose and mouth are so placed that they are very readily 106 THE SPECIAL NERVOUS MECHANISMS 107 exposed to stimulation by chemical substances floating in the inspired air or dissolved in the liquids taken into the mouth, and they are highly sensitive to such chemical stimuli. On the other hand, they are not at all affected by light or sound vibra- tions, and mere physical contact with them of some solid in- soluble substance only evokes a sensation of touch. All sensory surfaces are stimulated by electric discharges, as one may find by placing the electrodes from a battery cell on the tongue, but in such cases the sensation has not the quality of that which is evoked by the special agency appropriate to the sense organ. A special sense organ consists, therefore, of the terminations (dendrites) of a nerve cell. The axon of this nerve cell forms one of the fibres in the sensory or afferent nerve which conveys into the central nervous system the impulses generated by the stimulation of the dendrites. As a rule the essential or special part of the sense organ is not so simple as we have indicated. Thus we have in the retina, or essential part of the organ of vision, the following structures at least : /-' irfi; Inferior of Gan fcell& \ e^- ball Thickness of me retina FIG. 30. A DIAGRAM OF THE NERVOUS ELEMENTS OF THE RETINA. ["he latter is supposed to be seen in section, the concave surface being that receiving the light from the pupil and the convex surface being that turned to the back of the eye. The actual nerve terminations that receive the rays of light are the nerve cells, called the rods and cones (1). From these elements axons go off and make synapses with the dendrites of another nerve cell (2), which gives off an axon that enters into a synapse with a third nerve cell (3). The axon of this is pro- longed out into the optic nerve, and so passes up into the brain. None of these nervous elements, or neurones, is in actual physical contact with another. 108 THE MECHANISM OF LIFE This is a very complicated instance, but it is the type of an essential sense receptor. There is always a terminal nerve cell, and the dendrites of this receive the physical stimulus and trans- mit it through synapses to other neurones, and, finally, via the sensory tract, to the cerebral centres. These terminal dendrites, or sensory nerve terminations, are peculiarly modified in each case, and their threshold is lowered, so that they are susceptible to one form of physical stimulus rather than others. The non-nervous parts of the sense organ are concerned in transmitting the particular stimulus and in excluding others. Thus the retina is enclosed in the ball of the eye, and the front part of the latter (the cornea) is transparent, so that light can pass through it. Behind the cornea is the lens with the mechanisms for altering its focus, and the changing diaphragm, or iris, which regulates the amount of light that penetrates it. These dioptric parts of the eye are in every way comparable with a camera, lens, and diaphragm, which focus the external light upon a sensitive plate, making a picture there. In the same way the ear consists of the outer, middle, and internal parts. Externally there is a stretched membrane (the drum), which is set in vibration by sound waves in the outer atmosphere. The drum communicates its motions to a chain of three small bones (the auditory ossicles), which again transmit the vibrations to the liquid contained in a cavity in the bone of the skull. This vibrating liquid then stimulates the nerve terminations of the cochlear (auditory) nerve in a most complex structure, called the organ of Corti, and the stimuli are trans- mitted along the cochlear nerve to the auditory centres. The organ of hearing thus excludes light, chemical and mechanical stimuli, but allows the periodic variations of atmospheric pressure that we call " sound " to reach the nerve terminations. The nerve terminations in the mucous membranes of the mouth and nasal cavities are " bare " that is, they are exposed to all physical stimuli: variations of temperature, some light, mechanical pressure, and chemical activity. But they are highly sensitive to the stimuli of different kinds of chemical substances, so that they can distinguish between the latter. And they are very susceptible in this way, so that, for instance, one easily tastes the difference between the flesh of, say, cod and haddock, or plaice and sole, a distinction which cannot yet be made by chemical analysis. The nerve terminations of the THE SPECIAL NERVOUS MECHANISMS 109 olfactory nerve are still more delicate even in man, in whom the sensation of smell is degenerate and chemical substances existing in such small quantity in the air that they cannot be detected by any known methods of analysis are easily distin- guished by smell. Still more incredible is the delicacy of the sense in such an animal as a bloodhound. Heat receptors in the skin are nerve terminations that are stimulated by a rise of temperature above a certain limit, but are not affected by a fall below that. Cold receptors are, con- versely, stimulated by a decrease, but not by an increase of temperature. Pressure receptors, or muscular sense receptors, are stimulated mechanically that is, by something pressing on the skin, or by the degree of tension of a muscle, but not (or at least not much) by chemical changes in the skin or muscle, and not by changes of temperature. Equilibrium receptors, which are present in the " vestibular " part of the internal ear, are stimulated by changes in the position of the body with respect to its surroundings. Thus a man who is blindfolded has his ears and nose stopped, and who is lying immobile on a turn-table, can appreciate a noiseless, Motionless change in the position of his body, and can even roughly estimate the magni- tude of the angle through which he is turned. Thus the first step in the development of sensation consists in a refinement of the general irritability, or susceptibility to external changes, which we regard as one of the essential pro- perties of living tissues. The " refinement " means that speciali- ties of reaction are evolved: one kind of tissue, or arrangement of nerve terminations, becomes more sensitive to one kind of physical stimulus and less sensitive to all others. These specialised receptors then localise themselves in appropriate parts of the body, and become served by separate nerves. It is improbable that the fibres themselves are different in different sensory nerves that is, could we transplant the auditory nerve fibres to the optic tracts, they would probably conduct optic stimuli just as well as they conducted auditory ones, and vice versa. The nerve fibres are conductors, and nothing more. The next step is to place the fibres that transmit impulses from a receptor in connection with fibres that transmit impulses to a muscle or other effector organ, and that is done in the spinal cord, the lower brain, and the cerebellum. The sensory (or afferent) fibres enter into synapses with motor* (or efferent) 110 THE MECHANISM OF LIFE fibres, and thus an impulse arising from the stimulus of some receptor owing to a change in the environment becomes con- ] verted into another kind of impulse that stimulates muscles to contract and relax, and so enables the animal to make an appro- priate (or adaptive) response. But why " appropriate " ? We discuss this question (but do not answer it) in a later chapter. The lower brain, cerebellum, and spinal cord, are therefore centres (or ganglia) where afferent impulses become converted into efferent ones. They are simply the loti of synapses. The last development of sensation is the becoming aware of the external changes that stimulate the receptors. It must not be thought that the development of consciousness of the environ- ment, or of the body itself, is the sole, or even the main, function of the nervous system. What the latter does in the lower animals almost entirely, and what it mainly does, even in man, is to convert a sensory stimulus set up by some change in the environment into an appropriate response. Consciousness and psychical life doubtless accompany this conversion in all animals, though their intensity is the dimmer the lower in the scale of evolution the organism is placed. In ourselves this final develop- ment of sensation is the function of the cortex cerebri. General Body Sensation. Remembering, then, that the impulses arising from the stimulation of a receptor organ need not, and usually do not, give rise to changes of consciousness, we may con- sider, first, the paths by which the afferent impulses coming from the skin, muscles, and joints of the limbs and trunk reach the brain. All such impulses enter the cord by the dorsal (or posterior) or sensory roots of the spinal nerves, but the paths along which they afterwards travel depend upon their nature. Those that arise as the stimulation of receptors in the deeper muscles and joints enter the grey matter of the cord, and are received by the synapses of nerve cells there. From these nerve cells axons pass out into the white matter, and these form nervous tracts on each side that go up through the medulla, enter the inferior peduncles of the cerebellum, and end in the grey matter of that part of the brain. At the same time other nerve fibres, carrying similar im- pulses, enter the cord by the sensory roots, and travel directly up in the white matter without first forming synapses in the grey matter. Thus a large number of nerve fibres coming from the receptor organs in the deeper muscles and joints deliver their impulses into tracts of fibres in the cord, which then run up through the THE SPECIAL NERVOUS MECHANISMS 111 medulla and the inferior peduncles of the cerebellum to end in the grey matter of the latter part of the brain. But a large number of fibres entering the cord via the sensory roots take a very different course. These do not form synapses in the grey matter, but turn into the white matter at once, and form the two great tracts of fibres represented in Fig. 28 as the " great sensory tracts in the cord." These tracts end in the medulla in four prominent nuclei that is to say, the fibres form synapses with the nerve cells in these ganglia. The axons passing out from the latter cells are collected together to form the two great sensory tracts called the lemnisci, and the latter, after crossing, as indicated in Fig. 28, run up into the corpora quadri- gemina and optic thalami, and end by forming synapses with the cells in those nuclei. There they come into relation with nervous tracts passing out from the brain, and serving as the avenues along which motor impulses go out to the muscles of the body. So far, then, as we have studied it, the great sensory tract from the cord to the brain is one which carries impulses arising in the muscles and joints up into the mid-brain centres, via the ganglia in the medulla and the lemnisci tracts, and which, again, is mainly or solely a means of muscular co-ordination. In order that consciousness may be affected by these impulses, another link must be added to the chain of paths, and this is, we shall see, the tract of fibres passing up from the mid-brain to the cortex cerebri. Lastly, impulses arising from the heat, cold, touch, and pain receptors in the skin pass into the cord through fibres in the sensory roots of the spinal nerves. These go into the grey matter directly, and form synapses round the nerve cells there. The axons of such cells go out again into the white matter of the cord, but they do not form a continuous tract of fibres. Instead of that, they run upwards only a short distance, turn back into the grey matter, form other synapses, and then re-enter the white matter to form another short tract. By a series of such linkages the impulses reach the medulla, and then pass up into the lower brain by a devious route. Finally, they become connected with tracts passing up into the cortex, when they undergo full development into psychical affections. Thus the paths or tracts " mediating " general sensibility of the limbs and trunk lead up through the sensory roots of the spinal nerves and through the grey and white matter of the cord in in halami and mid- brain 3-1 Cord and medulla FIG. 31. DIAGRAM OF THE MAIN PATHS OF NERVOUS IMPULSES FROM THE BODY INTO THE BRAIN. Three spinal nerve fibres (I, II, III) are shown entering the cord. Each fibre divides into three branches. Left : Impulses resulting from general sensation pass up in the cord as shown, entering into and again emerging from the grey matter ; theconnection isdirectly with the thulamua, Centre: The paths taken ly impulses coming from the muscles and joints ; the COM nection is with the thalamus, but indirectly through nuclei in the medulla. lit : The paths taken by impulses coming from the muscles and joints, nnd entering in the cerebellum, but also ascending to the cortex via the thalamus and returning to the cord via the red nucleus. THE SPECIAL NERVOUS MECHANISMS 113 into the medulla. The greater number of the impulses passing along these paths are of the nature of stimuli, which lead to muscular responses only, and they are rearranged and co- ordinated in the cerebellum and mid-brain, so that the actions resulting from them are purposeful and appropriate. Some of them, but apparently only a small fraction of the total, proceed upwards to the cortex after passing through synapses in the medulla and mid-brain, and give rise to the changes of conscious- ness which we recognise as the sensations of pain, heat, cold, touch, pressure, resistance, etc. General Sensibility in the Head. The same kinds of general receptors are found in the skin, the muscles, and the joints in body, limbs, and head. But the impulses coming from the region of the head and face enter the central nervous system through the sensory fibres of the cranial nerves. Some of these are nerves of special sensation, and we consider them separately. Others (the 3rd, 4th, and 6th) are purely motor nerves, and go to the muscles of the eyes. The remainder (that is, the 5th, 7th, 9th, and 10th) are mixed that is, they contain both sensory and motor fibres and they may be compared with spinal nerves. They, with the purely motor nerves, convey efferent impulses from the medulla and mid-brain to the muscles of the eyes, face, jaws, and neck, and they also carry afferent impulses, arising as the result of the stimulation of the general receptors of those regions, into nuclei in the medulla and lower parts of the mid- brain. The internal paths in the brain are rather complicated, and we cannot attempt to describe them here ; but there is the same general scheme as in the case of the special nerves : the afferent impulses coming from the face and head go up to the mid-brain, and through this to the cortex, on the one hand, and into the cerebellum on the other. The Sensation of Smell. This stands quite apart from all the other sensory mechanisms in that the connection of the olfactory organ is with the cortex direct, and not with the medulla and mid-brain. The organs of smell and vision and hearing have also a different embryogenic origin from that of the other sense organs in that they arise as parts of the brain, which become pushed out, so to speak, and come into relation with other parts arising from the embryonic integument. The auditory organs originate in this way from the hind-brain vesicle, the visual organs from the fore-brain vesicle, but the olfactory organs 8 114 THE MECHANISM OF LIFE arise as outgrowths from the lateral fore-brain vesicles that become, later on, the cerebral hemispheres. Therefore the connection of the nerve terminations in the mucous membranes of the nose are with the cortex cerebri direct, and this may be the reason why the sensation of smell is said to be more " reminiscent " than those of vision or hearing: it sets up more immediate " associations,-' because of its place of entrance into the higher brain. Less, however, is known about the internal tracts along which the olfactory impulses travel in man than in the lower animals, because of the degeneracy of the human organ of smell in comparison with that of most other vertebrate animals. In some fishes, for instance, the great development of the olfactory organs and their nervous tracts suggests that the sensation in question plays a very important part in the general behaviour of these animals. The Auditory Organs. Two entirely different sense organs are contained in the " internal ear." " Vibrations of sound " that is, very rapid, periodically repeated alternations of com- pression and rarefaction of the air are made to impinge on the tympanic membranes, or " drums of the ear," and then, by means of the chain of little bones, called the auditory ossicles, these vibratory movements of the drums are transmitted to the fluid contained in peculiarly shaped cavities in bones on each side of the head. Two organs are contained in this cavity, or " bony labyrinth " the organ of equilibrium, and that of true hearing. The former consists of a little membranous sac from which proceed three semicircular canals, which are arranged in the three planes to which we refer all positions in abstract space. That is, one canal is vertical and runs forward and backward, another is vertical and runs from side to side at right angles to the first one, while the third is horizontal and is at right angles to the first and second. The dendrites of one branch of the eighth or auditory nerve project into the fluid contained in the bases of the semicircular canals. Operative interference inj many animals and diseased conditions in man show that destruc- tion of one or other of the canals produces well-marked abnor- malities in locomotion, apparent giddiness, and lack of co- ordination over the muscles of the limbs. The " vestibular " part of the ear is therefore an organ of equilibrium, and its nerve terminates in a nucleus in the medulla, which is connected in ways that are not well known with the mid- brain on the one THE SPECIAL NERVOUS MECHANISMS 115 hand, and the cerebellum on the other. There is, however, no evidence that it has any connections with the cortex. The organ of hearing, called the " cochlea," is essentially a long tube bent on itself like a hairpin, and twisted round spirally like the shell of a periwinkle. Within this tube is a fluid which bathes a very peculiar and complicated structure called the organ of Corti. The fibres of the true auditory or cochlear nerve terminate in cells in the organ of Corti, and delicate auditory hairs project out from these cells into the fluid. When the latter is set in vibration, the auditory hairs are stimulated in such a way that afferent impulses are transmitted along the fibres of the nerve into two nuclei in the medulla, where they are received by synapses. The axons of the cells in these nuclei are now gathered up into certain internal tracts, which pass up along the lemnisci (see Fig. 28) into the mid - brain. Other fibres go to the cerebel- lum. From the cells in the mid-brain round which the axons of the lemnisci end other axons go up, along a very obvious tract, called the auditory radia- tions, into the sensory region of the cortex. The Organs of Vision. We have already said something about the general structure of the eyes. The essential elements are the cells of the retina, which, we have seen, are arranged in layers. Light is received by either the rods and cones, and these are prolonged into axons which form synapses with other " bipolar" cells, the axons of which form other synapses with third, " ganglionic " cells, the axons of which pass out of the retina into the optic nerve. The latter then run up towards the Corpora. CjjUadrijgemina. Thalamus Optic " radiations FIG. 32. THE CONNECTIONS OF THE RETINAS WITH THE BRAIN. Only the tracts on one side are shown. The section of the brain represented dia- gratnmatically passes rather obliquely through the eyes. 116 THE MECHANISM OF LIFE mid-brain as the two optic tracts, which then partially cross each other in the middle line of the head. The further course of the optic tracts in the brain is now fairly well known. Fig. 32 represents these tracts in a very schematic way. Some of the fibres of the right optic nerve pass over to the left-hand side of the brain, while some keep on the same side (and vice versa). Then all the optic fibres terminate by forming synapses with the cells in three nuclei contained in the optic thai ami and corpora quadrigemina. That is their lower-brain termination, .but a very obvious tract of other fibres, the optic radiations, start off as the axons of cells in the lower visual centres or nuclei, and proceed up to the cortex cerebri. Audition and vision are complex sensations. When we hear we distinguish loudness (that is, the amplitude or " intensity " of the sound waves), pitch (which is the frequency of occurrence of the sound waves), and musical quality (which we explain by assuming that the sound waves are complex, and can be de- composed into components). Similarly, in vision we distinguish between intensity of light and quality of light (or colour). By intensity we mean the amplitude of the vibrations of the medium (ether) which transmits that which we recognise as light, and by colour we mean the components of this mixed light as they are separated from each other by the physical media outside the retina itself. How the analyses of sound and visual stimuli are carried out by the auditory and visual organs is, of course, far from being understood, and we cannot discuss the question now. So much, then, for a very summary consideration of the ways by which the stimuli of the receptor organs of the body are transmitted into the central nervous system and brought to bear upon the various ganglia there. We must next consider The Motor Mechanisms. From what has been said in Chapter II. the reader will already know that a motor mechanism includes (1) an afferent nervous path leading into the central nervous system; (2) a nucleus 01 ganglion; and (3) an efferent nervous path leading out from the ganglion to the motor organ. Thus we take the case of a pureh special mechanism. Some receptor organ is stimulated (say a touch spot in th< skin), and an afferent impulse is set up and propagated along t spinal nerve into the grey matter of the cord. This impulse THE SPECIAL NERVOUS MECHANISMS 117 enters the latter via the dorsal or sensory root. In the grey matter it is received by the dendrites of a nerve cell, and is then transferred (after something has been done with it) to the axon of the cell. This axon issues from the grey matter and passes out from the cord through the ventral or motor root of the same, or a different, spinal nerve, and is transmitted by the latter as an efferent impulse to a muscle which it then sets in motion. This is merely a scheme illustrating the simplest conceivable form of nervous motor apparatus. It may be regarded as the unit FIG. 33. DIAGRAM OP THE SIMPLEST POSSIBLE SPDSTAL REFLEX MECHANISM. An afferent fibre is represented as entering the cord, and ending in a synapse round a cell, which then sends out an axon to a muscle fibre. Through an additional cell and synapse another axon goes out to the antago- nistic muscle. The + sign indicates that one muscle contracts, and the sign that the antagonistic one relaxes. or element of the sensori-motor mechanism, but the reader must be very clear in his mind that the very simplest actual nerve-muscle mechanism is very much more complicated than Fig. 33 suggests. So we may now proceed to elaborate it till it approximates to the conditions that may be studied experimentally. First, then, one muscle never, by itself, constitutes a motor mechanism. For there are always two antagonistic organs, one of which contracts while the other simultaneously relaxes. Secondly, the nerve that supplies a muscle always contains fibres that carry 118 THE MECHANISM OF LIFE efferent or motor impulses from the nucleus (or ganglion) to the muscle, and also fibres that convey afferent or sensory impulses from the muscle to the nucleus. Thirdly, the stimulus that starts the nervous impulse and excites the muscular organs to activity nearly always originates in a different segment of the body from that containing the former, so that there must generally be a path, or tract, in the central nervous system itself along which the impulse travels. Lastly, there are hardly ever only two antagonistic muscles concerned in a movement, but rather a muscle system consisting of several or many such pairs. Thus the simplest actually observable action that can occur in the body of a higher animal includes a rather complicated mechanism, and the complexity of this becomes all the greater when we take account of the connections of the nucleus immedi- ately controlling the action with the higher brain centres. When such connections exist we have the possibility that the action may be modifiable to almost any degree by the volition of the animal, or by its " experience." Leaving aside, in the meantime, the factors of volition and experience, we may consider the action as it is performed in an automatic, mechanical manner. Let it be that which may occur when the side of the spinal dog* is tickled and the " scratching reflex " occurs (see pp. 119, 137). Fig. 34 represents nervous mechanisms that are involved in this 'action. These mechanisms (to judge from the figure) appear to be somewhat complicated, and yet we have taken account only of those which must be in action, and we have omitted others that are less essential to our present explanation. The stimulation, then, of the touch organs in the skin of the body sets up impulses that are transmitted to the spinal cord via the afferent fibres of a spinal nerve. Now this segment of the cord is well in front of that from which are given off the motor nerves supplying the hind-limb, and so there must be a nervous tract connecting centres (a) and (6). The afferent impulse, after being received by the synapses of the grey matter in centre (a), is modified in some way, is retransmitted along a tract of fibres in the white matter of the cord, and is received by motor nerve cells in segment (6). We must think about these motor nerve cells as being arranged in some way or other * A " spinal animal " is one in which the brain has either been destroyed altogether (by operation) or has been separated from the spinal cord. Thus the muscles of the body are entirely controlled by the cord. THE SPECIAL NERVOUS MECHANISMS 119 in pairs, some of them being the cells of axons that go to the flexor muscles, while others are the cells of axons that go to the extensors. Now the same impulse breaks upon both these series of axons, but it causes, at the same time, the flexor to contract and the (antagonistic) extensor to relax, thus bending the limb. But the muscle fibres contain receptors which are stimulated by the acts of contraction and relaxation, and thus set up impulses that arc propagated >/>/' na / S final B along the affer- ent nerve fibres coming from the muscle. These afferent fibres pass into the grey matter of the cord, and are received by nerve cells there. From the latter cells axons pass off and enter in- to synapses with the motor cells which control the movements of the muscles. Thus the mus- cular apparatus responds to a stimulus which enters the central nervous system at a place removed some distance from that place from which the motor nerves emerge, so that there are internal paths of communication in the cord. Also the contracting and relaxing muscles " advise " the motor centres (via their own receptors and afferent nerves) what they are doing, so to speak, so that the entire series of actions may be adjusted, or controlled, or accelerated, or inhibited, while they are still in progress. -^x Tens or muscle *- -Relaxor muscle FIG. 34. THE MAIN MECHANISMS INVOLVED IN THE SCRATCHING REFLEX OP THE DOG. An afferent fibre enters the cord, and is connected by various cells, their synapses and axons with two motor cells in a lower segment. The latter send out axons to the two antagonistic muscles, and the muscles send up afferent fibres which come into connection, via synapses, with the motor cells. 120 THE MECHANISM OF LIFE Now add to all this (1) a new series of connections between the nerve cells of the grey matter in segment (6) and the cere- bellum, and note that there must be two sets of paths, to and from the cerebellum, included in these connections. Add (2) a series of connections between the segment (a) and the mid-brain, also to and from paths ; these are, of course, the tracts described on pp. 99-104 and in Figs. 28 and 31. Add, finally, (3), the con- nections between the mid-brain and the cortex on the cne hand, and the mid-brain and the cerebellum on the other these also being to and from paths and the reader may get some idea of the complexity of the machinery of nerve cells and nerve tracts that are involved in a relatively simple action when it is the object of control by will or experience. The " Simple " Reflex Mechanism. The schematic simple reflex mechanism, such as it is figured in p. 117, is to be regarded as a " fiction" in the conventional, legal sense that is to say, it is a " scheme " which is useful in enabling one to understand the maze of nervous tracts and paths along which impulses travel within the nervous system. If we could isolate all the afferent fibres connecting one group of nerve cells in the spinal cord with one group of receptor organs in the skin, these would represent the afferent path of a simple reflex. Then we should have to isolate all the nerve fibres . connecting the same group of nerve cells with some one muscle, and that would be the efferent path. So we should obtain our schematic " reflex arc." Nothing like this exists in the higher animal. The afferent impulses originating in any one small group of receptors do not go to one segment only in the spinal cord, but to several such. Each of these segments is connected with the same muscle, for the fibres going out from the cord through one motor root and ending in one muscle are derived from several segments. Thus one efferent and one peripheral afferent path are connected together by various nuclei, and these nuclei are joined by several intraspinal paths. Then we have the further complications indicated in Fig. 34. The schematic single efferent paths are really double ones, each of them including a nerve branch going to a muscle and to its antagonist. The same impulse which breaks upon the synapse in the nucleus stimulates the nerve going to the muscle, so that THE SPECIAL NERVOUS MECHANISMS 121 the latter contracts, and it also stimulates the nerve going to the antagonist, so that it relaxes. These two antagonistic effects must be regarded as the single functional muscular action that is, their effect is, say, to bend or extend a limb. But the movement of the limb is something that varies to a very great degree: the part may be completely or partially bent (or extended), and it may bend or extend against resistances which also vary immensely. The duration of the movement therefore varies, and so also does the quantity of energy trans- formed more or less work is done according to the circum- stances under which the action is performed. To some extent all this regulation is carried out in the nucleus from which the efferent impulses proceed out to the muscle, but it is also the work of the contracting (or relaxing) muscle itself. There are receptors in the latter which are affected or stimulated by the events taking place in it, and by the circumstances of the action the load, for instance, borne by the contracting muscle. These receptors generate impulses which ascend into the nucleus and modify, if need be, the efferent impulses sent out by the latter. A mechanical analogy may make this very important function j of the muscle proprioceptors clear. Let us think about a small railway system in which the movements of every train are initiated and regulated by operators working in a central telegraphic control station. But as every train proceeds on its journey, moves from block to block, stops at and starts from . a station, it records its position telegraphically (and perhaps auto- matically) on a time and space chart in the control office, so that the operator may see at each movement where it actually is. That would represent the system of efferent nerves going from the spinal cord nucleus to the muscular apparatus, and the receptors of the latter, with their afferent nerves, going back into the nucleus. These, then, are the mechanisms concerned in the simplest spinal reflex arc: several cord segments in receipt of afferent impulses from the skin, and all of them in connection with the same small group of antagonistic muscles and a system of re- ceptors in those muscles in connection with the nuclei from which the efferent impulses start. The Cranio-Spinal Reflexes. The muscular apparatus concerned in movements are, we have seen, under the immediate control of ganglia or nuclei in the spinal cord, or (what are very similar) the ganglia in the medulla 122 THE MECHANISM OF LIFE from which the motor fibres of the cranial nerves issue. Keflexes, or purposeful and useful adaptive movements, may be carried out under such control, and apart altogether from the activities of the brain, although there are probably very few actions into which some amount of brain control does not enter. So far we have supposed that the stimulus to an action is some change in the environment acting on receptor organs situated in the skin, but in the intact, normal animal it is far more likely that the Mfd Infra- cerebral bram ^/pafh, cerebellum to rnid-u Cerebellum -Afferent ftaih. Cord To cerebellum -Afferent f}a1h, cord To lhalamus and m id- bra in ---Efferent fait,, mid- brain fd Cord FIG. 35. AFFERENT IMPULSES FROM THE SKIN ARE SHOWN GOING UP INTO THE THALAMUS, WHERE VISUAL IMPULSES ARE ALSO RECEIVED. Other afferent impulses enter the cerebellum from the muscles. From the cereballum impulses pass to the mid-brain, and from there motor impulses pass down through the cord, and to the muscles concerned in locomotion. receptors stimulated will be one or several of the great cranial organs of special sense. Thus an animal usually responds to something which it sees, or hears, or smells, and so we must include among the afferent paths that take part in the reflexes, or other actions, those going from the visual, auditory, and olfactory organs into the mid-brain. Yet the impulses actually setting the muscles in action must issue from nuclei in the medulla and spinal cord, and must go out to the muscles via the motor THE SPECIAL NERVOUS MECHANISMS 123 roots of the cranial and spinal nerves. That means, obviously, that there must be paths, confined to the central nervous system itself, along which impulses may travel from the nuclei of special sense in the mid-brain to the motor nuclei in the medulla and cord. The principal intracerebral and intraspinal paths, as well as those which pass from cord to brain, and vice versa, are represented in Fig. 28. Therefore our " simple " reflex, or other action, will (in the intact, normal animal) include all the paths, or analogous ones, represented in Fig. 33, and also another series of paths between the organs of special sense and the mid- brain, and between the latter and the medullary and spinal motor nuclei. Thus the main paths in use in the case of a man walking in a crowded street and " mechanically " avoiding other people must be somewhat as indicated in Fig. 35. Here we do not represent the afferent paths between the acting muscles and the nuclei in the cord, and, of course, no details of the very imperfectly known paths between the mid- brain and spinal centres. The Mechanism of Co-ordination. Nor do we ever suggest the all-important apparatus of co- ordination. In such a complex series of actions as those involved in walking a very great number of muscular systems are at work. Practically all the muscles of the legs are active and immediately concerned in the production of the movements, but the body is also carried erect and balanced, and this is the work of antag- onistic muscle systems belonging to the trunk. The arms swing about. At each step, and with every deviation from a straight line, the centre of gravity of the whole body changes its position. Practically every nucleus, or ganglion, in the whole spinal cord must be concerned in the generation of the impulses going out along the efferent nerves to the muscles, and in receiving the afferent impulses generated in the acting muscles and joints, and giving information from instant to instant as to what is going on there. Now it is unlikely that the relatively simple reflex mechanisms constituted by the sensory and other afferent nerves entering into synapses with the motor nerves is competent for this work of co-ordination, and there is much evidence that an additional nervous apparatus is involved. This additional mechanism is the cerebellum and its connec- tions. We have seen that there are tracts of nerve fibres passing 124 THE MECHANISM OF LIFE T halo. us From cortex through the superior peduncles and putting the cerebellum in communication with the mid-brain, and so with the organs of special sense. The vestibular part of the auditory organ that is, the part which is known experi- mentally to be concerned in the maintenance of equilibrium and posture of the body has also direct nervous connection with the cerebellum. The receptors in the deeper muscles and joints are con- nected with the cerebellum by special tracts (see Fig. 31), and there are also descending tracts connecting this part of the brain with the motor nuclei of the spinal cord. Even the nuclei of the cranial nerves are eereb'ellvm ho cord. Skin FIG. 36. THE CONNECTIONS OF THE CEREBELLUM. Impulses pass up from the skin, muscles, joints, etc., into the cerebellum, via I, the inferior, and S, the superior peduncles. From the cerebellum impulses connected with pass to the thalami and red nuclei, and from the latter efferent impulses pass down into the cord. The cerebellum is also connected, via M, the middle the cerebellum. It is known peduncle, with the cortex, and in other ways, not also, from direct shown in the diagram, with the organsof special sense. experiments, that injuries to, or even removal of, the cerebellum leads to no obvious impairment of sensation, and yet the organ has most conspicuous THE SPECIAL NERVOUS MECHANISMS 125 I and direct connections with almost all kinds of receptors. But I such injuries or operative interference do produce serious motor disturbances, so that ordinary, automatic, or customary move- I ments, such as those of gait, locomotion, and posture, are greatly I affected, and the animal when walking, running, swimming, or flying, behaves in an ineffective, incompetent manner. What- ever, then, are the functions of the cerebellum, it is very plain that they must be of considerable importance and are very complex a deduction from the peculiar and most intricate structure of the grey matter of this organ. The general con- clusion attained by the study of all these lines of evidence is that the work of co-ordination of movements performed customarily ! and automatically is carried out in the cerebellum. Impulses arising in the sense receptors are conveyed to their nuclei in the spinal cord and brain, or they may be conveyed directly into the cerebellum, and if the direct path does not exist, there is one connecting the immediate nucleus of the afferent impulses with the latter part of the brain. That is to say, the cerebellum is the recipient of impulses coming from the receptor organs, and particularly from those which have to do with equilibration and those others which come from the acting muscles themselves and from the joints (where the effort involved in the movements of the limbs must be particularly felt). On the other hand, the cerebellum also stands in close connection with the nuclei which give origin to impulses going out to the muscles of the limbs and body. There is, therefore, a mechanism whereby those complex movements which are the means of locomotion may be timed, adjusted, and co-ordinated, and this we suppose to be situated in the grey matter of the cerebellum. Of the nature of the mechanism we have not even a suggestion. Cortical Control. We shall consider in the next chapter the experimental evidence on which our knowledge of the functions of the cortex cerebri is based, and in the meantime we deal only with the nervous mechanisms themselves, so far as these have become known by anatomical research and experiment. The greater part of each cerebral hemisphere, then, is a core of nerve fibres making such connections as are indicated in Figs. 28 and 29 connections, that is, between the various parts of the cortex itself and between it and the other nuclei in the lower brain, cerebellum, and spinal cord. With the exception of the corpora 126 THE MECHANISM OF LIFE striata, the great nuclei of the cerebral hemispheres are situated in the thin sheet of cortical grey matter, and it is the cells of this that are connected with the underlying core of fibres. The latter are either the axons of these cells (in which case they carry impulses out from the cortex) or they end in forming synapses with the cortical cells (in which case they carry impulses to the cortex). Fig. 37 represents, on the same scale, a " pyramidal " cortical cell from the frog's cortex and one from man; the actual cell bodies are very much the same in the two cases, but the dendrites and axons are very much more complex in the human than in Principal axon FIG. 37. Two CORTICAL (PYRAMIDAL) CELLS FROM THE BRAIN OF MAN AND THE FROG. HIGHLY MAGNIFIED. the amphibian cortex. Each cell is pyramidal in general form (as it can be seen in section), and from the apex and lower parts there start off a great number of fine fibres which branch in a complicated way and have a structure peculiar to the cortex. From the middle point of the base of the cell there issues a prominent fibre, which is the axon of the cell, and branches pass off laterally from the axon and again branch. These are the " collateral " axons of the cell. Soon after leaving the latter the axons acquire medullary sheaths (see p. 26), and become the commissural or projection cortical fibres represented in THE SPECIAL NERVOUS MECHANISMS 127 Fig. 28. Other fibres, commissural or projection ones, enter the part of the cortex in question and break up into terminal dendrites, which then come into relation with the apical and basal dendrites of the cell by means of synapses. Thus each cell in the cortex can make a very great number of connections with other cortical cells, or with nerve cells in other parts of the central nervous system. Each of the collateral axons, for instance, can become a commissural or projection fibre, and a number of fibres coming from anywhere else can form synapses with the dendrites of any cortical cell. This is why the latter are so much more crowded in the canine, and still more so in the amphibian, than in the human cortex : there is the increasing tendency, as we raise in the scale of evolution, for the connections between the cells to become more and more numerous and complex, so that the number of paths which an impulse leaving a pyramidal cell may take tends always to increase. The higher is tJie type of brain, the more manifold are the ways in which the various centres may communicate with each other. Localisation o! Function in the Cortex. We must think of the activity of the cortex cerebri in a twofold way: it is one organ in so far as every part of it is connected with every other part, and it is multiple inasmuch as functions are specialised or localised in it. This specialisation has become known to us, partly by experiment upon the brains of the higher mammals (other than man), partly by the extension of these results to the human brain (which is, of course, very similar in structure to that of the anthropoid apes), and partly by observations made on human subjects suffering from disease and accidents. Thus it is possible to distinguish in the human cortex a " motor region," which is concerned with the initiation and elaboration of willed movements; a " sensory region," upon which is dependent the full psychical development of the sensations arising from stimulation of the receptor organs; and a " prefrontal region," about the functions of which we have very little positive knowledge. Fig. 38 represents the approximate positions and boundaries of the regions, and the particular areas into which the latter are divided. The two great fissures those of Rolando and Sylvius serve as lines which enable us to divide up the cortex into these regions and areas. One hemisphere (the left one) is seen from the side, but the reader must understand that the cortex dips down into the great median fissure that separates right and left 128 THE MECHANISM OF LIFE hemispheres, as well as into the Rolandic and Sylvian fissures, so ' that such a view as that of Fig. 38 does not show its entire area. Round about, but mainly in front and in the depths of the fissure of Rolando, is the motor region, and this has been divided up into a number of particular areas, each of which controls the movements of some small part of the body. Thus, directly in front of the fissure are the " centres " or " areas " for the movements of the trunk, arm, hand, head and eyes, face, etc. These areas are not very well defined- (for instance, they do not coincide exactly with the areas of the various " gyri " or con- volutions), and they overlap to some extent. When any one of them is stimulated in a suitable manner, usually by application of -Muscular rum impr* FIG. 38. DIAGRAM OF THE PRINCIPAL CORTICAL CENTRES IN THE LEFT HUMAN CEREBRAL HEMISPHERE. a feeble electric current, certain groups of muscles twitch or con- tract, and this is the evidence that the area in question controls these parts. The movements that are so elicited are not the finished ones that occur in the normal behaviour of the animal, but they indicate none the less the nature of the nervous controls and paths. The cortex is, as we have seen, a thin sheet of grey matter overlying the core of nerve fibres that forms the greater part of the mass of the hemispheres. Directly beneath it we come upon the white matter, and much of this consists of the axons proceeding from the overlying pyramidal cells of the cortex. Tracing these fibres by various means, we find that they run down into the crura or cerebral peduncles towards the medulla. There, as Fig. 28 shows, these " pyramidal tracts " mainly cross each other, that one coming from the left-hand side of the brain THE SPECIAL NEKVOUS MECHANISMS 129 going over to the right-hand side of the medulla, and vice versa. The pyramidal tracts are then continued down the spinal cord, and there their fibres enter into the grey matter and form synapses with the nerve cells which give origin to the axons that go out into the ventral or motor roots of the spinal nerves. Thus the grey matter of the motor region of the cortex is, on the one hand, in connection with fibres that run uninterruptedly down into the spinal cord and control the nervous centres there which set the muscles of the body and limbs in action. On the other hand, it is in connection with the centres in the other parts of the cortex which are the termini of the sensory paths. But it is also in connection with the cerebellum via another specialised cortical region. Figs. 28 and 36 show a prominent tract of fibres descending from the cortex and ending in the region of the pons varolii that is, the middle part of the middle peduncles which unite together the two halves of the cerebellum. Thus our cortical motor cells have communication with the nervous mechanism of co-ordination. And just in the same way as the pyramidal tracts connect together the cortical and spinal nuclei, so do other more com- plicated paths connect the motor cortex with the medullary nuclei which give origin to the nerves that supply the muscles of the head and face. Thus every part of the motor system of the whole body is under the control of the cortex cerebri. This is particularly the case in man. In the fishes the great pyramidal tracts are hardly to be recognised, and they become developed to a progressively greater extent as we ascend the scale of evolu- tion represented by the dog, monkeys, anthropoid apes, and man. In the lower vertebrates the nuclei which mediate between the organs of sense and the spinal ganglia are those in the mid- brain, but in the higher mammals this nervous mechanism becomes more and more replaced by the cortex and its connections. Almost all the cortex behind the motor area is, as Fig. 38 indicates, the seat of sensory functions. Those centres called " visual," " auditory," " music," etc., contain pyramidal and other cells, the dendrites of which form synapses with the tracts of fibres coming up from the mid-brain nuclei in which the nerves coming from the organs of sense terminate. Stimulation of these cortical sensory centres is certainly essential to the development into full consciousness of the impulses arising from the stimulation of the sense organs. This is a matter to which we return in the following chapter. 9 CHAPTER VIII THE ANALYSIS OF BEHAVIOUR OUR analysis of the nervous system has, so far, been that of a very complicated structure built up of superposed reflex arcs. A reflex arc is the series of nervous parts that connect together a receptor and an effector organ; thus the arc that is functional in the simple act of winking connects the retina of the eye with the visual centre in the mid-brain (by the optic nerve), the visual centre with the oculo-motor centre (by an intracerebral tract), and the oculo-motor centre with the muscles of the eye- lids (by the oculo-motor and facial nerves). When an action of any kind occurs, such a reflex arc or arcs become functional. The receptor organ receives a stimulus that is, something happens outside the body: a flash of light, a noise, a current of air carrying odoriferous particles in suspension, etc. and this event causes some change in the receptor organ. The nerve terminations in the retina, the internal ear, or the mucous membrane of the nose are thus stimulated, and they initiate a nervous impulse which is propagated along the sensory nerve to the centres of the brain. There the afferent nervous impulse breaks upon a series of synapses, and so enters a number of nerve cells that constitute the centre (or nucleus, or ganglion). Something now happens to the impulse in these cells, and its physical nature is doubtless changed in some way. At any rate it is transferred from the cells to the axons which come from the latter. Those axons constitute a nervous tract leading to another nucleus, where there is also a series of synapses. The impulse, after further modification in the cells of the second nucleus, is now transferred to an efferent nervous tract which is constituted by the axons passing out from those cells. After traversing this third nervous tract (or effector nerve), the impulse is received by the effector organ. Let us suppose that this is a muscle. The latter there- upon contracts or relaxes. This is a scheme of what happens whenever a bodily action occurs ; a stimulus is initiated and propagated along an afferent 130 THE ANALYSIS OF BEHAVIOUR 131 nerve into the central nervous system. It is there prolonged through an efferent nerve into the effector organ, where it releases energy. Now we may try to make a " mechanical model " of this train of events, premising that whatever it may be that occurs in the actual nervous arc it is sure to be (physically) very different from the things that happen when our model is set in action, but that the energy relations in the actual muscle-nerve structures and the parts of the model will be the same. Suppose, then, that a number of billiard balls, a to g, are suspended by threads so that they almost touch each other; let another ball, h, be poised on the end of a tipless cue. Let the end ball, a. be pulled a little to one side, and then let go so that it hits b very gently. Further, we may assume (as it is generally assumed when logical hypotheses or mechanical models are made) that the balls are perfectly elastic (which they nearly are), and that there is no friction in their suspensions. The ball a will then com- municate momentum to b, and OO OO Q O O cause b to hit c, c to hit d, and f 6 d C b so on; c will communicate as much energy to d as it received from b, and a wave of mechanical displacement will travel alqng the row. The ball g will hit h with as much force in the blow as a has hit b, and the blow will cause h to roll off its perch and drop to the floor. Suppose still further that a has been lifted up J inch; that the weight of each ball is J pound; and that the height of h above the floor is 6 feet. Now, in falling through J inch a does work equal to its weight X the distance through which it falls that is, oVx|=T2 foot-pound. The quantity of energy so represented is " propagated " along the row of balls and com- municated to h, and is just sufficient to push the latter off the end of the cue jon which it is poised. It then falls 6 feet, and so does work equal to 6xJ=2 foot-pounds. Prior to being displaced it had this quantity of potential energy due to its having been lifted from the floor and put in such a position that it was free to fall 6 feet. A small quantity of energy representing T-V foot-pound of work can therefore be propagated without loss, and can release a much larger quantity that is, 2 foot-pounds. FIG. 39. 132 THE MECHANISM OF LIFE What we know about the passage of a nervous impulse along a nerve suggests that the former passes without any of the substance of the nerve being used up. Arriving at the first synapses, it passes into the nerve cells there and transforms into the same quantity of energy, which then passes along the intra-j cerebral tract into the second synapses and cells, where another equal quantity of energy is transformed and is propagated along the efferent nerve (again without loss), and thrown into the! muscle. But there it releases a much greater quantity of energy, I which is represented by the force with which the muscle con- tracts or relaxes. The latter contained energy in the potential (chemical) form, and the minute quantity entering it as a nervous impulse sets free or transforms this chemical energy in the same way as a small electric current can release (or fire) the huge quantity of energy contained in an explosive charge. Now, from the point of view that we have taken so far, every- i thing that happens in the sensori-motor system of an animal conforms to the structural scheme of reflex arcs and to the dynamical scheme of the billiard-ball model. The peripheral and central nervous system is a means whereby the energy received by the stimulation of the receptor organs is transmitted as a nervous impulse to the nerve centres, and is there trans- formed into another kind of impulse which is transmitted to an effector organ, where, finally, potential chemical energy is released, and muscles contract and relax or glano^s secrete. The impulse which results from the stimulation of a sense organ goes to the central nervous system, but there it may pass along one or more of a great number of different paths. Upon the path it takes depends its effect; thus the stimulation of the retina may lead to a reflex act of winking, or the " mouth may water," or the man may start violently, or run, or sit down, or laugh. Nothing, then, is explained by our structural analysis of the nervous system; all that we have studied is the means whereby one of a number of effects that may be produced by a stimulus is produced. Consider now the energetical side of the process of stimulus and response. Just that quantity of energy which is received by the receptor organ is transmitted along the afferent nerve into the centre, and the same quantity again is sent through the central nervous system from centre to centre, and still the same quantity is transmitted along the efferent nerve into the effector THE ANALYSIS OF BEHAVIOUR 133 organ, where it is all expended on releasing the potential energy which is to produce the effect in question. Or suppose (for it does not matter to our argument in the least) that some of the energy entering the sensori-motor system is dissipated, or wasted, or " lost " by " friction " (as some would actually be " lost " in the billiard-ball model). This dissipated energy will therefore appear in the form of heat, and the rest will be transmitted as before to the effector organ. Where, then, does the conscious- ness of having seen something and acted upon the stimulus so received come from ? For we must consider the suggestion that an affection of consciousness is the result of an energy trans- formation ; that it comes from the energy of light that stimulated the retina and transformed into a nervous impulse just in the same way as the heating of a metallic filament is the result of the transformation of an electric current which passes through a lamp. We are not going to accept this suggestion, but we notice it in order that the reader may see quite clearly what are the various possibilities (let us say) : (1) All the energy that comes from without the body and stimulates a sense organ is transmitted without loss through the peripheral and central nervous system, and is physically trans- formed in the effector organ, releasing potential energy there which does work. (2) Some of it is dissipated, and the rest is transmitted as above. (3) Some of it is transmitted as in (1), with or without dissipa- tion, and some of it is physically transformed into " consciousness." Now what we know suggests that there is extremely little or even no dissipation of energy in the transmission of a nervous impulse through a reflex arc. And the more we think about the third of our three hypotheses, the more unlikely it appears to be. At all events, it does not seem possible even to attempt to verify it. We return to this discussion in a later chapter. Meanwhile, let the reader note that all that we have studied so far all, indeed, that it appears that cerebral physiology can study are the ways in which the stimulation of the organs of sense set up nervous impulses, the paths along which those impulses travel in the central nervous system, and the motor and secretory effects that they produce. There is nothing at all about a theory of know- ledge in the results of such an investigation, but very much about a theory of action. Sensation is not something that enables us to contemplate the external world, but is rather that 134 THE MECHANISM OF LIFE which enables us to act upon the latter. It is not so muc prolonged or developed into consciousness of our environmen as into action upon the environment. We must deal further with this aspect of physiology. Stimulus and Response in General. We consider again a " mechanical model." Let there be little compass, NS, placed in the neighbourhood of a wire int which an electric current may be thrown by depressing a switch. So long as the current does not flow through the conductor, the needle remains in the position NS that is, in the mag- netic meridian; but whenever the switch is put down and the current passes in the (conventional) direction indicated by the arrow, the compass needle deviates and takes up the new position WE. When the current is switched off the needle returns to NS, and no matter how often we do this the effect is always the same. Now call the making contact in the switch a "stimulus" and the deviation of the needle the "response"; the latter, we see, is invariable and inevitable. Given that the same quantity of current flows FIG. 40. A COMPASS through the conductor, and that the in- NEEDLE, NS, LAID . ., , ,, ,, , ,. ~ ,, ALONGSIDE A WIRE, 06, tensity of the earths magnetic field THROUGH WHICH A CUB- remains constant, then the needle always KENT CAN BE PASSED. deviates in the same direction and to the same extent. There is strict determinism that is, the " response " (a certain deviation) always follows upon the " stimulus " (a current of a certain value). The Muscle-Nerve Preparation. Now let the prominent thigh muscle (gastrocnemius) in the frog's leg be dissected out, leaving its tendon still attached to the femur and an inch of its nerve (the sciatic) attached to the muscle which carries a weight. Place two electrodes carrying a current in contact with the nerve. Throw a momentary electric current into the circuit containing the electrodes, and the nerve will be stimulated at the place a. The stimulus initiates an impulse which traverses the nerve and releases potential energy in the muscle, whereupon the THE ANALYSIS OF BEHAVIOUR 135 latter contracts, lifting the weight against the resistance of gravity. Do this again and again, and the same effect follows. By-and-by the muscle, or the nerve terminations in it, will become fatigued, but until this happens an electric stimulus will always elicit the same muscular contraction or response. Here, also, there is, perhaps, strict determinism. Tropisms in Green Plants. When a seed germinates, two structures grow out from it. One of these is the original root and the other is the original shoot. The former always grows down into the soil, and the latter grows upwards into the air. When a green plant is placed in front of a window, the leaves tend to turn towards the source of the light. The leaves of a tree tend always to place themselves so that their flat surfaces are perpendicular to the principal direction from which the light comes. All these effects are called " tropisms," or directed growth movements of fixed organisms. The tissues of the plant have the general power of growth, but the stimulus of light falling on them is a directive agency, which causes growth to take place in one direction rather than any others. Such directed growth movements in response to the stimulus of light are called heliotropisms. They are invariable and subject to determinism, just as are the deviation of the compass needle or the contraction of the isolated frog's muscle when the nerve supplying it is stimulated. Taxis in the Lower Animals. Chip off some barnacles (Balanus) from the stones on the beach between tide- marks during the last week of March or the first ones of April, and place them in a soup-plate containing clean sea-water. In a short time the embryos contained in the reproductive organs will hatch out and swim about in the water. Examine the latter in a feeble, diffused light, and it will be seen that the larvae swim about at random and in no particular direction; but place a lamp close to one side of the vessel, and it will be seen that they swim 136 THE MECHANISM OF LIFE towards the point from which most light comes. This response is called a phototaxis, and it may be denned as the directed move- ments of an organism in response to a directed light stimulus. It is invariable and determined, and a logical hypothesis that is, one which is consistent with what we know about light and its general effect on living tissues can be made to account for it. The Spinal Reflex Action. " Pith " a frog that is, cut through the spinal cord immediately behind the brain, and then destroy the latter by pushing a blunt wire into it. The body of the animal is now solely under the control of the spinal cord and its ganglia; to convince oneself that this is the case, the head may be removed. If, now (or, rather, after the effects of the " shock " have passed off), a drop of strong vinegar be placed on the back of the animal, one of the hind-legs will be bent forward and the acid will be wiped off. A reflex arc of some complexity is here involved: afferent impulses pass into the spinal cord from the receptors in the skin irritated by the acid, and these impulses are received by the nerve cells in the segment of the cord which innervates the part of the skin stimulated. From the segment stimulated tracts of fibres convey the impulses received to the grey matter of the segments from which the legs are supplied with motor nerves. The latter are then stimulated, and the antagonistic muscles contract and relax, carrying out the series of movements described (Fig. 42). Now here, again, there is determinism, or, at least, the results of stimulating the skin can be predicted. But there is now a difference between the response in this case and the " mechani- cally " repeated one that occurs in the muscle-nerve preparation: when the drop of acid is placed on the right flank of the headless frog, the right leg is used to wipe it off. Now let this leg be cut away or forcibly held, and the left one is used to make the same kind of response. Something occurs, then, in this case which we have not, so far, observed. Evidently there are two mechan- isms, one for each side of the body, but in what we may call ordinary circumstances one of them is passive. Let, however, the " normal " mechanism be prevented from operating, and then that one which was passive before now responds vicariously. We have to deal here with a " regulation." Such a " spinal reflex " that is, a co-ordinated action carried out by the nervous ganglia of the cord can be elicited from higher animals than the frog. When the cord is severed in the THE ANALYSIS OF BEHAVIOUR 137 dog and the effects of the shock of the operation have passed away, suitable stimulation of the skin of the side just behind the shoulder is followed by the response called the " scratch reflex": the hind-leg of the same side carries out a series of kicking movements somewhat similar to those performed when the normal animal scratches himself as the result of irritation by a flea. Even in man something of the same kind may occur. Thus, in cases of hemiplegia, when the connection of the cortex _ Afferent , fibres s^Area of f ' \ stimulated -/<- 1 r c/s >inal cord ^- ^^x II ^ J ' J^ -- il I - nerves Seg mefl fs _of the cord ~ Supplying I i mbs FIG. 42. THE " SCHEMATIC " SPINAL REFLEX, BEING THE NERVOUS PATHS INCLUDED WHEN THE STIMULUS is THROWN INTO A DIFFERENT SEGMENT FROM THAT INNERVATING THE MUSCLES THAT ACT. with the cord is rendered ineffective by a " stroke," reflexes in the paralysed leg may be obtained, and these are doubtless due to the centres in the spinal cord. Reflex actions, complex and purposeful in character, may thus be carried out by the nerve centres in the spinal cord. In such cases the stimuli (irritation by chemical substance, an electric current, mechanical pinching, pricking, etc.) applied to the skin are simple physical ones, and the response is generally such an action as would be carried out by the muscles concerned, in the normal animal, for some useful purpose. It always has a certain " inevitability " that is, it " comes off," as a rule, when the stimulus is applied. There is determinism, or at least a large measure of determinism. It can usually be predicted. 138 THE MECHANSIM OF LIFE Reflexes in the Normal Animal. Now compare with these j spinal reflexes those that may be observed in the normal, intact animals. Such a response is easily observed in the case of a j dog which is lying sleeping on his side with his legs stretched out. When the skin over the ribs is rubbed vigorously, the hind-leg of the same side will often make a series of foolish little kicks, like incipient scratching movements. We may even study such reflexes in ourselves. We usually start violently on hearing a loud, unexpected noise, and the muscular actions involved have meaning, inasmuch as they seem to be such as would place the body in a posture of defence to meet some sudden menace, i When some object makes an unexpected movement towards the eyes the lids are rapidly closed, and here also the action has j purpose the protection of the eyes. As a general rule these i and similar responses are quite involuntary, and it may even ] require some considerable effort of the will to arrest or prevent them. But that they can be arrested or prevented is the charac- j teristic that distinguishes them from the reflexes which are j carried out by the spinal cord alone in the lower vertebrates or in the higher vertebrates deprived of their cerebral hemispheres, j Even the sleeping dog will not always give the incipient scratch reflex when his side is rubbed, and by strong attention and " will- power " a man may keep his eyes open when someone flicks some object towards the face, or when he puts his head under- neath water, and one may easily arrest the start that he naturally makes when he hears a loud, unexpected noise. This point is very important. The response that may be elicited by a stimulus applied to the animal possessing its integral nervous system is not a " fatal " or " inevitable " one. It may occur, but it may not. It may occur in one of several ways, and it may occur and immediately be arrested. All that means that the strict physical determinism that one sees in the " response " or deviation of a compass needle when a magnet is brought near to it, or the equally determined tropistic and tactic responses of i the lower organisms, or the spinal animal, do not occur in the higher vertebrate possessing its entire central nervous system. ; In the responses that one studies in such cases there is always what we must call indeterminism : they are usually unpredictable. But while this is the case, it is no less clear that we can study a series of responses beginning with the purely inorganic reaction of the compass needle and magnet, and passing through the THE ANALYSIS OF BEHAVIOUR 139 muscle-nerve response, tropisms, taxis, reflexes, in animals sub- jected to various kinds of operative interference, and ending with the behaviour of the normal, intact higher animal. At the beginning of such a series there is determinism, and at its end there is indeterminism ; but it would be rather difficult to suggest any place where determined reaction became replaced by some degree of chosen, deliberated behaviour. In such a series of organic mechanisms we should be able to trace the beginning and the gradual elaboration of a central nervous system. Nothing of the kind is, of course, present in the compass-needle-magnet system, and it is very doubtful if it can be recognised in the typical plant organism. It is present in a very simple form (far simpler than that which we have represented as present in the earthworm) in the barnacle larva, and it becomes progressively more complex as we ascend the vertebrate series. Somewhere, then, in the ascending scale of living things indeterminism of response is developed; we emphasise the word "response" so that it may not appear that we exclude indeterminism of func- tioning in the most general sense from the lower organisms. The Lower Brain Activities. We may next consider how the reactions exhibited by an animal are modified when certain parts of the brain are removed, or isolated from the other parts. Now, in such an animal as the frog, both the cerebellum and the cerebral hemispheres are undeveloped relatively to the medulla and mid-brain. The former organ, may be called rudimentary, and the cerebrum consists of the two corpora striata, each with a rudimentary cortex cerebri. It is an easy matter to remove the two cerebral hemispheres (including the corpora striata), and the animal sur- vives the operation. It is then in possession of a central nervous system, including the parts developed from the three primary brain vesicles, but lacking the fully developed cerebellum which is present in most vertebrate animals higher than the reptiles. Now every action that can be performed by the intact frog can equally well be performed by the decerebrate frog provided that a suitable stimulus be applied. It can swim, leap, and crawl, and it assumes, when motionless, the natural posture of a frog. It can avoid obstacles when it is moving, and it is sensitive to light. Its visceral, respiratory, and nutritive organs function normally. It swallows food which is placed in its mouth. If HO THE MECHANISM OF LIFE it is turned over on to its back it can regain its proper attitude. But it does none of those things spontaneously, and if it is not stimulated it makes none, or very few, movements of its own accord, and it nearly always remains in exactly the same position if it is left strictly alone. When it is stimulated it responds in an almost invariable way, and the effect of any particular stimulus can be predicted. What is wanting in its reactions and movements is the evidence of spontaneity: there is no " intrinsic stimulus," and nothing that indicates the possession of volition. It is " a machine, and nothing more, while the frog possessing its cerebral hemispheres is a machine governed and checked by a dominant volition." Its responses to external stimuli indicate that it possesses the power of completely co-ordinating its movements. That power, we shall see, is dependent in higher vertebrates on the presence of the cerebellum, and on the full connections of this organ with the rest of the nervous system. But in the frog the cerebellum is rudimentary, and we must therefore conclude that the func- tions that it performs in the higher vertebrates are carried out by the mid-brain. Therefore we cannot say that co-ordination is the work of the cerebellum exclusively in all vertebrates. It can be effected by other parts of the brain. Now remove the cerebral hemispheres that is, the corpora striata and pallia from a fish, and the animal survives the operation. But there is no absence, " not even the temporary absence." of apparently spontaneous movements. It is there- fore evident that spontaneity of behaviour, which depends, in the frog, upon the presence of the cerebral hemispheres, can be mediated, in the fish, by the mid and hind brain, just as co-ordi- nated movements, which depend, in the bird, upon the presence of the cerebellum, can be mediated by the mid-brain in the frog. The cerebral hemipheres are much more highly developed in the birds than in the amphibia, but they can be removed and the animal (the pigeon) can be kept alive for a considerable time after the operation. The same general effects observed in the frog can also be seen in the decerebrate bird. It either remains quiet and impassive, or it moves about restlessly and without any apparent purpose. Of itself it does not attempt to fly, but if it is thrown into the air it will fly and avoid obstacles. It does not spontaneously pick up corn, though it will do so if its THE ANALYSIS OF BEHAVIOUR 141 beak is thrust among the grain. Like the frog, it responds to stimuli, and it sees and hears in so far as seeing and hearing are the stimuli to movements; but sight and hearing do not, ap- parently, evoke memories or utilise experience. Its movements are perfectly co-ordinated. There is the same inevitable response to external stimuli, the same automatism of activity, and the same lack of spontaneity and volition (which latter is indicated by spontaneity) that we see in the decerebrate frog. Among the mammals the cerebral hemispheres are much more highly developed, again, than in the bird, and their removal is a formidable operation. Yet it has been accomplished in the dog, and when the animal is nursed with the utmost care and solicitude it may survive for over a year. In the few successful experiments that have been made there has been some little difficulty in exactly describing the condition of the patients. As in the frog and bird, there is no paralysis and no lack of co-ordination so long as the cerebellum is intact. All ordinary movements are carried out, and, as in the bird, there is a tendency to restlessness. The sense organs are active in so far as the motor organs can respond to ordinary sensory stimuli, but things that would, in the intact dog, evoke expressions of terror, dislike, and pleasure, do not appear to affect the decerebrate animal in anything like the same degree. Signs of hunger are exhibited, and food brought near to the nose is eaten. Disagreeable food may be rejected (for instance, one of Golz's dogs would not eat the flesh of another dog). Both in decerebrate dogs and cats painful stimulation elicited expressions of anger (growling, barking, and snarling), but no caressing could evoke any indica- tions of pleasure or affection. There was apparently no dream- ing. Finally (a most curious thing), Golz's dog ate much more than the normal animal did. Nervous Inhibition. We must say more about these results. In the decerebrate mammal there is the same lack of spontaneous activity as in the frog and bird, understanding by this the absence of apparently willed, or deliberated, or intelligent, or chosen activities, and not the aimless, automatic activity to which we referred above as " restlessness." There is the same tendency for a stimulus to evoke an inevitable response the actions of the animal can be predicted. But the inhibitions are still more interesting. 142 THE MECHANISM OF LIFE A nervous inhibition is the arrest, entire or partial, of some activity in progress, and such effects are very common and very important. The rhythmic activity of the heart, for instance, is intrinsic that is, it apparently goes on as the result of some periodic stimulus originating in its own substance. But this rhythmic activity is subject to nervous control, the heart-beat being regulated by two sets of nerves, one coming from the sympathetic system, and the other from the tenth (pneumo- gastric or vagus) nerve. The stimulation of the former accelerates, and that of the latter inhibits, the rate of the heart, so that, by suitable stimulation of the vagus, the beat may be slowed down or may even be arrested altogether. Now nervous " shock " that is, the great prostration arising from an injury, or the effects of a surgical operation is to be regarded as a series of inhibitions. In some way or other im- pulses passing down from the higher brain centres are blocked (as when the spinal cord is severed), or are not initiated (probably as the result of afferent stimulation in the case of a severe injury), and the stopping of these impulses is the main cause of the prostration and other effects called " shock." In the decerebrate mammal, then, impulses that normally issued from the cortex cerebri cease, and the general activities of the animal are affected. The restlessness referred to above may be traced to the cessation of inhibitory impulses which in normal life arrest or modify aimless, random movements. The abnormally large consump- tion of food may also be traced to the cessation of cortical impulses regulating the metabolism of the tissues and economis- ing energy, and the condition that anger and dislike could be elicited, but not pleasure and affection, may also be traceable to the loss of inhibitions. For one may (taking a general survey of organic behaviour) conclude that the feral animal has normally a " bad time." It struggles for its existence both with inorganic nature and with its organic enemies. It must " eat or be eaten." " Softness of heart," dalliance, pity for others, and the like, are feelings that are wanting, or, if present, are likely to be detrimental or fatal, while their opposites make for self-preservation. What we call loosely the altruistic motives must be regarded as the product of the herd instinct, and they are to be interpreted as inhibitions or checks upon the natural, predatory, and highly individualistic modes of beha/iour. They are opposed to most tendencies that THE ANALYSIS OF BEHAVIOUR 143 are the products of natural selection, as a study of adaptations among wild animals will show. Insanity in man that is not the result of lesions, it has been argued, is to be interpreted as con- flict between the mental complexes that the struggle for existence has engendered on the one hand, and those newer complexes that arise from the development of the herd instinct on the other. As in Mr. Wells' s Island of Dr. Moreau, there is a " law " (a series of inhibitions) which comes into tragic antagonism with the naturally evolved animal propensities. And so, lacking the inhibitory controls acquired by domestica- tion and centred in its cortex, Golz's dog could growl and bark, and manifest anger and displeasure, but not affection which to it was something quite secondary an inhibition of the " currish " nature of the natural dog. And so also a study of social evolution and of mass psychology impels one to the conclusion that what we recognise as " good " is mostly the inhibition of what we may call the lower animal instincts that are in us. The result is, of course, clearly demon- strable in much of our conduct, for our codes of private and public morality are, to a great extent, summaries of the things that may not be done inhibitions and which the natural man would often like to do; while our legal systems supply the sanc- tions for those prohibitions and restrictions. What are called " socialist tendencies " are, of course, attempted inhibitions of individualism. The ruthlessness that characterised German methods of warfare far more than those of their opponents was the result of a slackening of inhibitions, a disregard of the amenities of armed conflict (the things that might not be done), and as such it met with general reprobation. The " war psychoses " of European countries during and after the year 1919 illustrate the same proposition: one cannot help noticing a tendency to increased dislike of people belonging to other nationalities than our own, and a general indifference to suffering borne by other peoples. Those feelings were potential in us before the war, but they were repressed, and the strains set up by the necessity for natural self-preservation loosened the in- hibition and allowed of their expression. The Cortex Cerebri. From what has already been said about the progressive development of the cerebral hemisphere in the vertebrate animals the reader will see that we ought not to speak about " the " 144 THE MECHANISM OF LIFE cortex. It is not the same thing in the mammals as it is in the birds and reptiles, where it is a more complex organ than it is id the amphibians, and it does not occur at all among the fishes. This disparity of development must always be remembered when we discuss the functions of the cerebral hemispheres so far as these can be made out from observations of the effects of opera-j tive interference. Removal has little or no apparent effect in a] fish, where the " cortex " is non-nervous, but the result is very obvious in the frog, and still more so in mammals such as the rat, rabbit, and dog. In the monkeys and apes, and, of course, in man, the operation is an impossible one, for these animals do not survive it long enough to make clinical study profitable. In lower vertebrates, such as the frog, the cortex is relatively unimportant (judged from the anatomical standpoint) when it is compared with the rest of the brain, and the functions that it subserves are also unimportant. When it is removed the change of behaviour is not a profound one, and it is possible that some of the things that were formerly done by the cerebral hemispheres are then done (or partially done) by the mid-brain. To some extent this is also the case with the dog, where vicarious function- ing on the part of the lower brain may be set up when the cortex is removed. That such is the case is suggested by the observa- tion that animals deprived of their cerebral hemispheres tend to act more normally the longer they can be kept alive and in g general health. Now, in the monkeys and anthropoid apes, th development of the means of control over acting by the co: has been carried so far that vicarious functioning by the mid- brain and thalami become impossible, and this is still more the case with man. Reviewing the bare summary of the evidence that has been made here we see, however, that with the removal of the cortex cerebri spontaneous movements tend to disappear, while auto- matic and mechanical activities persist. The animal so trailed reacts to stimuli in a blind, inevitable manner,so that its responses can be predicted. There is physical determinism at any rate, much more of such than in the intact, cerebrate animal. Our only criterion of volition, deliberation, and intelligence is this presence of spontaneous behaviour, and therefore we are justified in placing the immediate expressions of will and intelligence in the activities of the cortical mechanisms. THE ANALYSIS OF BEHAVIOUR 145 The Nervous System as a Whole. Having made this very bare analysis of the structure of the central nervous system, we are now in a position to consider its working as a whole, but, first of all, we may profitably think about it as a series of superposed mechanisms, thus following the natural path of evolution. Fundamentally, then, the nervous system mediates between the stimulation of the organs of sense on the one hand, and the organs of activity on the other. Some physical change in the environment leads to the stimula- tion of a receptor organ, and the initiation of a nervous impulse. The latter is received by the nerve cells in the ganglion, and in its A * Shin Iffector Receptor organ .Muscles FIG. 43. THE SIMPLEST POSSIBLE SENSORI-MOTOR ACTIVITY. A spinal reflex mechanism, the line AB, represents the junction of spinal cord and medulla. turn initiates a number of impulses which pass out from the ganglion along efferent nerves and set up a co-ordinated and purposeful activity in the muscles to which those nerves go. There need not be (and there usually is not) any perception in such a stimulus and response. The latter is usually determined, and can be predicted. Next, the development of the special sense organs in the head, and the concentration of ganglia there, lead to a certain integra- tion of sensory stimuli and their resulting responses. The head ganglia were primarily the centres, or nuclei, of the great organs of sense, but it happens they also become connected with the lower spinal centres, as we have indicated in Fig. 28, and so we get a superposed series of connections. 10 146 THE MECHANISM OF LIFE The mediating mechanism is now greatly complicated, inas- much as the receptor organs which are in primary connection with the spinal ganglia are also in connection with the ganglia of the organs of special sense in the head, and there are also motor paths between the mid-brain and the cord. Obviously actions immediately carried out by the motor centres in the cord can now become more complex, and may be co-ordinated to a greater extent because of the additional stimuli given by the special c Receptors in head Muscles in head m x ot motor % \Mid 1 brain, J lhalami, medulla ^Tracts x " between ~ "\" Mid- brain K J j- cord -T\ : il ~3J iclei ** C cranial "nerves > J?ecefifors ,'- n skin t-c. Muscles of trunk and limbs v.^ ' Sfiinal cord 'Afferent spinal nerve irrerent Spinal nerves FIG. 44. MAIN PATHS BETWEEN SPINAL CORD AND LOWER BRAIN. sense organs. Still, the responses may largely be automatic and inevitable ones, although there must be some subtle difference in them when compared with the purely spinal reactions. Such a structure and relationships are, in general, those of the fish, where the cortex is absent and the cerebellum is rudimentary. Now consider the additional complexity which is brought about by the development of the cerebral hemispheres and cerebellum, organs which have evolved simultaneously, or nearly so. Neglecting the intermediate stages of this evolution (for we THE ANALYSIS OF BEHAVIOUR 147 know relatively little of the nerve tracts in the brains of the amphibia, reptiles, and birds), we may consider the conditions in man. We have already indicated the connections between the cord, cerebellum, mid-brain, and cortex, but a purely schematic figure will be useful. Fig. 44, then, represents the main com- munications between the mid-brain and spinal cord. The parts control CORTEX nerves IfferenT Cranial muscles of head Afferent sfiina I nerves general bodily sensation Efferentl Spina I 1 nerves muscles of body, THALAMl ft I'D -BRAIN MEDULLA B SPINAL CORD \--general sensibility in muscles '-T > ain i heat i cold Touch ~ flu scu la r control FIG. 45. MAIN PATHS IN THE NERVOUS SYSTEM AS A WHOLE. below the line, AB, we regard as the simple, primitive, central, nervous system, such as it probably was in the ancestors of the vertebrates, while the parts above AB represent the addi- tional connections established when the great sense organs of the head have become evolved. In Fig. 45 we add the mechanisms involved in the cerebellum and cerebral hemispheres. 148 THE MECHANISM OF LIFE Now, but for two exceptional paths that of the impulses passing directly into the cerebral hemispheres through the olfactory nerves, and that of the impulses going to the cerebellum from the vestibular part of the auditory organ the parts repre- sented above the line CD in Fig. 45 connect together the various components of the central nervous system they are the integrating mechanisms. Thus the cerebellum receives impulses directly from the muscles and joints of the body and others indirectly (via the mid- brain) from the skin ; it receives impulses indirectly (also via the mid-brain) from the organs of special sense in the head, and it is in direct (to and from) communication with the cortex. And so it has a grip on all the afferent impulses, whether those of general or special sensation, arising anywhere in the body. The cerebral cortex has conspicuous connections with the cerebellum, as we have just said, but the communications with the nuclei of the mid-brain, in which the nerves of sense end, are just as prominent, and the great pyramidal tract leading down from the motor region of the cortex to the nuclei of the motor spinal nerves is more conspicuous still. And so the grip of the cortex cerebri on both receptor and effector organs is as evident as that of the cerebellum on the receptor organs. We may now resume the general working of this whole mechanism. The parts below the line AB constitute relatively simple reflex arcs sufficient in themselves for movements initiated by the stimulation of the sense organs in the limbs and trunk. The parts between AB and CD that is, the mid-brain complex also form reflex arcs, but the distance receptors (visual and auditory organs) are now included, and become incorporated with the spinal arcs by means of the tracts joining mid-brain and cord. The stimuli initiating movements are now much more varied and numerous, and so the latter become more complex than when they depend on the cord alone. Finally, when the parts above CD are fully evolved, something quite new is added to the activities of the central nervous system. The cerebellum mediates between the impulses coming from the receptors in general (but particularly those in the muscles and joints), and the impulses that pass out from the cortex to set muscular mechanisms in action. It is not concerned in the psychical life of the animal to any degree that can be recognised, but it " standardises " and co-ordinates a number of very com- THE ANALYSIS OF BEHAVIOUR 149 plex movements performed in a customary manner those of locomotion and posture. Something additional to this is effected in the cortex. Looking at the main features of the anatomy of the latter, we see the extraordinary development of the motor region and the tract of fibres connecting this with the spinal nerve nuclei, and studying the results obtained clinically (in the case of man) and experimentally (in the case of other mam- malia), we see that the movements most suggestively connected with cortex and pyramidal tracts are of a special kind : they are skilled movements, and the facility to make them is acquired by training and is ino^vidual that is, they are not transmitted by heredity. Further, they are usually spontaneous movements, or at least they are such when they are being learned. Looking again at anatomical, experimental, and clinical results, we see that the cortex is to be associated with sensation that develops into consciousness. The connections between it and the sensory nuclei in the mid-brain are very obvious, and all observational results point to the same conclusions. We return to this matter in a later chapter. The Meaning of Behaviour. We define " behaviour " (premising that we are now dealing with the higher animals) as the totality of the activities of the entire sensori-motor system a definition which, however, must not be strained. It excludes the activities of the viscera (heart and bloodvessels, respiratory, nutritive, and excretory organs) because these are subsidiary to the proper functioning of the sensori-motor system. It excludes the activities of the repro- ductive organs, because it is the individual animal that we are studying, and not the race. In behaviour in general we see the activity of the entire motor system, as in running, walking, swimming, flying, and other modes of locomotion, as well as the partial activities of bodily weapons, as in biting, clawing, etc. It would include the actions of defence, flight, concealment, aggression, the construction of nests and shelters, etc. It must also include the actions carried out in play, courtship, etc., and generally the " behaving " of the animal in the ordinary, non- technical sense of the word. Organic behaviour is to be regarded biologically as constitut- ing a series of adaptations. The things that occur in the outer world cannot, in general, be prevented by the animal living 150 THE MECHANISM OF LIFE there. Thus the succession of day and night, the order of the seasons, changes in weather, such as storms, gradual or catas- trophic geological changes, the vicissitudes of climate, tides, ocean currents, winds, etc. all these things are phases in a " cosmic order," and are beyond the control of the feral animal, or even of man. All that the organism can do is to avoid menace to itself, or to take advantage of external changes in so far as it can do so by some variation of bodily functioning. The real meaning of the evolutionary process, from the biological point of view, is the slow acquirement by the organism of the means of varying its functioning, so that it can evade external changes that are inimical to it or take advantage of other changes. When winter comes, many arctic animals respond to the external changes by a change in the colour of the fur, say, from brown to white. This renders them less conspicuous in a snow- covered country, and the animal obtains an advantage in that it is less easily seen by its enemies, and can the more easily approach other animals that are its natural prey. Many fishes and other animals hibernate during cold weather that is, they seek shelter of some kind, and lie in a passive condition, economising muscular movements as much as possible. The respiratory and heart movements slow down. Oxidation of the tissues is partially inhibited, and the animal lives on its reserves of fat and proteid. The advantage gained in such cases is that the animal is spared the necessity of seeking food at a time when this is very scarce. When a man passes rather suddenly from a tropical into a temperate climate, he excretes much less water from his skin and much more through his kidneys, and he thus economises heat by an inhibition of evaporation of sweat. These are instances of adaptations of the functioning of organs other than those belonging to the sensori-motor system, and they do not involve " psychological factors " that is, they would be possible in the absence of the higher parts of the brain, or at least they need not include the activities of the latter organs. They do not constitute behaviour. Neither do many adaptations that we call instinctive ones. Thus the blowfly lays its eggs in fresh meat, so that when the larvae hatch out they may obtain abundant nutriment. A bird builds its nest in such a place and in such a way that it is difficult to distinguish it from the surrounding objects, and so the con- cealment of the eggs and young is an advantage. A crab casts THE ANALYSIS OF BEHAVIOUR 151 its shell, and for a time it is almost unprotected against many natural enemies. In such cases it seeks shelter in a crevice among the stones or weed on the sea floor, and so conceals itself. These are examples of responses involving the activity of the sensori- motor system, but they are fixed, " mechanical " ones that have become part of the organisation of the animals displaying them ; they are transmissible by heredity, and they do not have to be acquired individually. It is true that they must have been acquired some time or other, but in a different way from the cases which we are aljput to mention. Young chicks learn to distinguish between small stones and grains of corn. Collie dogs find that they ought to leave sheep alone, and soon do so. Retrievers pick up, and bring back, game without crushing or eating it. Cats and dogs learn to open latches, to beg for food, and to recognise people who take notice of them. A man puts on an overcoat when the weather becomes cold, and the master of a ship alters his course when a sudden fall of the barometer in certain latitudes suggests that a cyclonic storm is approaching. These are all true adaptations, and each of them ends in the animal obtaining advantage to itself of some kind or other. They all involve intelligent acting of the sensori- motor system, and they are based upon experience. Some time in the past similar external changes have occurred, and the responses made have been successful or not that is, they have or have not been advantageous. The stimulus is remembered, and also the nature of the responses made, so that when the same external event, or change, recurs the unsuccessful response is avoided and the successful one is made. It does not matter that the master of a ship which is approaching a cyclonic storm may not himself ever have had this experience; he has acquired the experience of others, and knows what response is the advan- tageous one. Behaviour, in the sense that we employ the term, is therefore the functioning of the sensori-motor system which is based upon experience. By " experience " we mean individual acquirements, and not something that is inherited. When experience becomes a factor in the determination of the particular response that is being made to a stimulus, something new that is, something that we have not yet considered is included in the activities of the animal organism, and our conception of mechanistic responses must be re-examined, CHAPTER IX THE MECHANISTIC CONCEPTION OF LIFE THIS seems to be the proper place to say something about the " materialistic " or, as it is now termed, the " mechanistic " view of the living organism, for, although we have assumed such an hypothesis in all that has been said so far, we are about to abandon it, or, at least, we are about to give to " mechanism " a meaning that is rather different from the one that is usually accepted. Our description of the animal body was first of all mechanical, then we had to make use of physical and chemical ideas, and now we must search for some other concept, which must, nevertheless, still be a logical one. The body of the higher animal, then, is a system of muscles and other soft parts which are built up round about, or are supported by a system of rigid parts the skeleton. The latter consists of separate bones immovably attached together, as in the case of the parts of the skull and pelvis, or movably attached, as in the case of the vertebral column and the skeletons of the limbs. In the former cases the skeleton acts as the supports of the soft organs thus the skull contains the brain and the great sense organs while in the latter cases the vertebra and limbs are the apparatus of movement. Where the bones move on each other they are said to be articulated, and the configurations of the articulations (or joints) determine the ways in which the movements occur. Muscles, ending in tendons, are attached to these movable bones, and by their lengthening or shortening the parts are made to extend or bend on each other. Thus the enormous variety of movements that an animal can carry out depend on the shapes and lengths of the bones, the configurations of the joints, and the modes of attachment of the muscles and tendons. Further, the muscles are supplied with nutritive matter by a series of conduits the arteries, capillaries, veins, and lymphatic vessels. At a central point of this vascular system is a propul- sive organ, the heart, which keeps the blood and lymph in motion. The directions in which these fluids move are deter- 152 THE MECHANISTIC CONCEPTION OF LIFE 153 mined by valves, some of which are placed in the heart itself, others at the place of origin of the great arteries and veins, and others, again, in the veins and lymphatics. The velocity with which they stream through the vessels depends upon the propul- sive power of the heart, upon the varying calibre of the arteries and veins, and upon the posture of the body, for it is apparent that more power is expended upon circulating the blood when a man stands erect than when he sits down or reclines. So far, then, the bodily a^aratus is a purely mechanical one levers of various shapes and " orders," elastic parts that pull these levers, and channels along which fluids are propelled by the action of a force pump and are directed by means of valves. It would be possible to construct a model of the human body which would carry out the same kinds of motions that are made by the living organism. We should have to supply to this model some source of hydraulic power, some automatic contractile vessel worked by a spring that would cause the fluids to circu- late in the vessels, and we should have to devise some means of supplying motive force to the muscles say, compressed air from a reservoir contained somewhere in the body itself. There might be springs here and there on the surface of the model, and these could be made to actuate valves in the reservoir of compressed air, so that by touching them the limbs of the model would perform certain motions. All this is quite possible, and. indeed, automata of such a kind have been constructed. It would, of course, be impossible to imitate the extreme minute- ness of some parts of the animal body, or to duplicate the intricacies of the motions of other parts, but such limitations would be due only to our defective craftsmanship. There would, of course, be an essential and very curious difference between the activities of the living animal and those of the mechanical model a difference which is rather hard to describe in simple language. The Cartesian Mechanism. Descartes thought about the animal body as just such a mechanism as that suggested above. The bodies of the lower animals were pure automata, so that if they were touched, or pinched, or otherwise stimulated (as we should say), they would move, or fight, or run away, or cry out, etc., but these responses were as mechanical as would be the movements made by the model when a spring was pressed. The body of a man, he said, was also an automaton, but it contained 154 THE MECHANISM OF LIFE a " rational soul," while the bodies of the lower animals did not. The latter could not feel pain or pleasure, and they acted quite unconsciously; but the rational soul in the human body felt the stimulus and response, and could even modify the latter, or could initiate movements of the body by its own volition. Neverthe-j less, the body of a man or woman was an automaton, and it could do all that the body of a lower animal could do even when the rational soul was inactive. To understand this conception we must try to forget our modern notions of matter and energy, and we must study I Descartes' cosmogony. We shall return to the latter presently, 1 but in the meantime we must consider the physiology that was | current before his time and the ways in which he modified it. Now the science of mechanics was very well developed at the beginning of the seventeenth century, and craftsmanship had j reached a very high level of attainment. Anatomy had long i been studied, so that the general structures of the bodies of man and the lower animals were very well known and had been described in considerable detail. Thus the gross anatomy of the j skeleton, muscles, viscera, bloodvessels, brain, and nerves was known to a degree that is not greatly inferior to our knowledge ] (which excels that of medieval times mainly in its minutia?). j The microscope had not been invented, or at least had not been j improved to such an extent as to render it an aid to anatomy, and therefore such organs as the brain, the muscles, nerves, and viscera had then little of the complexity that we now know them to possess. Our modern chemistry did not exist, so that there is hardly what we should call a single chemical idea in all Descart cs' physiology. Finally, our kinetic energy was then called vi-s viva, \ and there was nothing at all comparable with our essentially modern conception of potential energy. The physiology was, then, largely that of Galen, but it wasj modified very remarkably by Harvey's demonstration of the circulation of the blood. The materials of the food taken into the stomach were supposed to be converted by the agitation of their particles into chyle, a turbid fluid which is present in the small intestine. This was absorbed by the bloodvessels of the gut, and was carried to the liver as the " natural spirits/" In the latter organ it became endued with the " vital spirits/' and the blood, containing this fluid, ascended to the heart, and was poured into the right auricle. There was a " fire " or " innate THE MECHANISTIC CONCEPTION OF LIFE 155 heat " in the heart similar, Descartes says, to that which may be observed in fermenting, damp hay, and this fire was fed by the vital spirits brought to the heart from the liver. As the blood fell into the right ventricle it became expanded, " just as all liquids do when allowed to fall, drop by drop, into a highly heated vessel " in other words, it was made to boil, but not simply to boil, as we should say, for it yielded nutriment to the fire of the heart at the same *ime. From the right-hand side of the heart the blood went to the lungs, where it lost some of its vapours, and became thick again by contact with the respired air, " without which process it would be unfit for the nourishment of the fire " of the heart. Returning then to the left ventricle it became distilled, or highly rarefied, with the production of a " very subtle wind, or, rather, a pure and vivid flame," which was the " animal spirits." This was the most agitated part of the blood, and as it ascended into the narrowing arteries leading to the brain the grosser parts were left behind, while the " very subtle wind " or " flame " entered the brain to become stored in the cerebral ventricles. The more sluggish and thicker parts of the blood, which were nevertheless a hot and nutritive fluid, then became distributed to all the other parts of the body (except the lungs and muscles). Nutrition and the formation of the " humours " (or secretions, as we should say) depended on the existence of sieve-like struc- tures at the extremities of the arteries, so that when the blood was forced through these the larger and more sluggish particles were left behind " in the same way that some sieves are observed to act, which, by being variously perforated, serve to separate different species of grain." The secretions and nutritive parts of the blood were, therefore,* filtered off, in the modern sense. The animal spirits are meanwhile stored in the cavities, or ventricles, of the brain. Now the nerves were described by Descartes as minute tubuli which contained axial threads (our axons). They had a twofold function: on the one hand they transmitted something from the extremities to the brain (our afferent impulses), and on the other they transmitted something (our efferent impulse) from the brain to the extremities. When a sense organ was stimulated, the axial nerve thread was affected in the same way as when a wire is pulled or shaken, and this motion acted upon valves in the walls of the cerebral ventricles, allowing the animal spirits to escape. The latter flowed out- 156 THE MECHANISM OF LIFE wards through the tubuli of the nerves, and so into the muscles. Expanding in the latter, they produced the contractions and relaxations which moved the parts of the body. Now in all this there is nothing but mechanism, in the strict sense. A fluid (the blood) is expanded and rarefied by heat; gross and cold, or slowly moving constituents are separated from finer, more ardent, and more quickly moving constituents merely by the difference of the motions; larger and coarser particles are separated from smaller and finer ones by sieves; liquids flow through tubes, and the thinner parts flow through the narrower tubes more quickly than do the thicker parts; there are valves that are pulled open by stretched organs; there are muscles, which swell or relax according to the way an expansible fluid is thrown into them; these muscles act upon levers, the bones, which thereupon move the limbs and other parts of the body. All this is pure mechanism in our sense. What were the animal spirits ? It is very difficult to avoid reading into Descartes' physiology our present-day notions of energy; still, the spirits were a fluid, but a very rare and subtle one, hot, and thin, and ardent that is, they were kinetic in our sense (like a highly heated gas under pressure). But they could be confined in the thin-walled cerebral ventricles and nerve tubuli, so that their energy must have been repressed in some way in our term, the energy was potential until it was released by the afferent nervous impulse. As Huxley says, a relatively slight change in Descartes' terminology would have brought his physiology into line with ours. Obviously he did very much what we do in our chemical and physical hypotheses of living activities he pushed his specula- tions to their limits, as the mathematicians would say. His analysis of structure stopped at what could be revealed by dis- sections, since there was no microscope, and therefore he assumed that his mechanism held true, in the parts that were beyond his observation, just as it did in the parts that he could see. It is very doubtful, however, whether he would have modified his speculations greatly even if he had known as much about the minute structure of the animal body as we do. For instance, the histology of muscle and nerve, as we know it, would lend itself quite well to his mechanical explanation, and the structure of the kidney, as we have sketched it on pp. 71-3, can easily be assumed (in the absence of chemical data, it must be noted) THE MECHANISTIC CONCEPTION OF LIFE 157 to be such a mechanical filtering apparatus as he postulated. It I is true that he could not see the pores, but neither can we even with the aid of the microscope. In short, Descartes assumed that his pure mechanism was operative beyond the limits of actual observation (and, as we shall see, that is also assumed by us in respect of our chemical and physical mechanism). The Influence of Chemical and Physical Investigations. - Now, apart from the influence of chemical discovery, it is not easy to see how the Cartesian physiology need have become modified. But the great advances made by chemistry towards the end of the eighteenth century changed the point of view entirely. The true theory of combustion, as it developed in the hands of Priestley, Black, and Lavoisier, was very revolutionary in its effect on the conception of the nature of vital activity. It was shown that when a piece of metal was strongly heated in the air a calx (that is, an oxide) was formed, and that when carbonaceous material was also made to glow in air some part of the latter disappeared, and " fixed air " (that is, carbonic acid gas) came into existence. Now the animal body, when dried, was found to consist largely of carbonaceous material, and it was discovered that when air was taken into the lungs some part of it disappeared and was replaced by fixed air, just as in the case of the combustion of carbon outside the body. The inference was soon made that in respiration there was an actual process of combustion, and that this occurred in the lungs, or blood, or tissues, and was the source of animal heat. There was not, therefore, an innate heat of the heart, and the latter became regarded solely as the propulsive organ of the circulation. With these discoveries the Galenic physiology became obsolete. The modern idea of energy came later, and developed naturally from the applications of the motive force of steam that were made by Watt and the engineers, and the theory of the perfect steam engine that was worked out by the French physicist, Carnot. This great man showed that an engine did work by taking heat from a source, and giving it up to a condenser, and that there was a transformation of heat into mechanical energy. His investigation became the foundation of our modern science of thermo-dynamics, which, in its turn, became enormously fruitful in the treatment of chemical problems. Later on Rumford pointed out that mechanical friction generated heat 158 THE MECHANISM OF LIFE (a thing that many people before must have observed, but did not think about), and by-and-by Joule estimated how much mechanical work was transformable into just how much heat, and he measured this quantity of heat by noting the increased temperature it could impart to a known mass of water. Joule's work had enormous influence on physiology, for it became possible to estimate the " calorific values " of known quantities of various kinds of food substances. Now a certain mass of combustible material thus became associated with a certain quantity of mechanical work theoretically deducible from it, and so it could be assumed that when food substances were eaten and oxidised in the animal body, some of their heat became transformed into mechanical work. Thus chemical energy could pass into the form of heat, and heat could be transformed into mechanical work. Apparently the animal body was a thermo- dynamic machine. Let us note, very shortly, the other great ideas borrowed physiology from chemical and physical science. Graham, aboi the middle of the nineteenth century, found that there were t) categories of chemical substances the crystalloids (like commc salt) that could pass through the pores of an animal membrane when they were dissolved in water, and the colloids (like gelatine or albumen solutions) which could not pass through. Later on de Vries showed how to measure osmosis that is, the passage of water through organic membranes. When a vegetable cell is placed in pure water the latter passes through the wall, so that the cell swells up, while, if it is placed in salt solution, which is more concentrated than the sap, it shrinks, because water now passes out from the cell to dilute the stronger salt solution. Graham's research on dialysis and de Vries' work on osmosis have had a profound effect on physiological research so much so that it has been said that the chemistry of life is largely a matter of the chemistry of colloids. Another most fruitful conception was that of catalysis. It had long been known that fermenta- tions in vegetable and animal substances were due to enzymes (or ferments); that an exceedingly small quantity of the latter substances could produce a chemical change which would not otherwise take place; and that the ferment itself need not be used up during the reaction. But during the nineteenth century it was discovered that a great number of purely mineral sub- stances could act in precisely the same way with other mineral THE MECHANISTIC CONCEPTION OF LIFE 159 substances. Thus the same kind of reaction was observed to occur both in organic and inorganic materials, and though we still speak of enzyme action in relation to vital chemistry and catalysis in regard to inorganic reactions, we know very well that the two classes of chemical changes are of the same kind. Along with all this investigation went on the synthesis of " organic " substances by the chemists. It is almost possible, even now, to feel the thrill that physiologists must have ex- perienced in 1828 when Wohler synthesised urea from mineral * substances. Before then there was an apparently hard line drawn between organic and inorganic chemical compounds, and it was thought that there were many substances, such as urea, starch, sugar, albumen, etc., which were only and could only be found in the cells of a living plant or animal. Wohler's syn- thesis of urea broke through that line, and the far more wonderful syntheses of sugars and polypeptides (the constituents of proteids) by Fischer and his pupils obliterated it altogether. To-day we look forward with complete assurance to the time when starches, proteids, and fats capable of assimilation by the animal body will be prepared, from their elements, by the chemists. Finally (and now we come down to a relatively short time ago), Jaccju.es Loeb demonstrated the possibility of artificial parthenogenesis, and thus physico-chemical mechanism seemed to be attacking vitalism in the very citadel itself. Before these famous experiments were made parthenogenesis (that is, virgin generation) was known to occur in a very few of the lower animals, but in most lower and all higher forms it was quite unknown; the ovum formed by the female could only be made to develop when it was impregnated, or fertilised, by the sper- matozoon formed by the male, and apart from the latter there could, apparently, be no reproduction. Now early in the present century Loeb showed that by simply adding certain chemical substances to the water containing the eggs of sea urchins, the latter could be made to undergo normal develop- ment. It is true that the male element in reproduction is not replaced by a chemical substance, for the spermatozoon does much more than simply initiate the process of segmentation and development of the ovum : it adds the paternal characters to the maternal ones. Still, even the initiation of the process of development by a simple mineral salt is a result of the most extraordinary theoretical interest. 160 THE MECHANISM OF LIFE So much for the main ideas and methods which have been introduced into biology from the chemical and physical side.1 Nothing at all has been said here about the impetus which these] ideas have given to what we*may call utilitarian biology; to! processes of further preparation of foodstuffs; to the breeding and rearing of useful organisms; and above all to methods for the prevention and curative treatment of disease. Even now,] and much more in the future, the relative freedom from disease that may be enjoyed by every sane man and woman is to bel traced back to the investigations made by a few chemists and physicists and biologists and medical men during tile last two! centuries. To many people all this work is very disagreeable,! and they do not like to think about it or even to enquire how itj is carried on. Like Tennyson's Princess, they shudder at " Those monstrous males that carve the living hound, And cram him with the fragments of the grave, Or, in the dark dissolving human heart, And holy secrets of this microcosm, Dabbling a shameless hand with shameful jest, Encarnalise their spirits." Throughout their lives they live upstairs, so to speak, knowing that most of the things that make living tolerable are the work of those that labour in the kitchen ; and one likes to think that j some time or other science takes its revenge, and that in the j end they have to go down into the basement and found a pathetic j reliance on the rather sordid toil that goes on there. It has been said by Michael Foster, and the statement can easily be verified, that the periods when mechanistic conceptions of life have dominated biology have also been those when the greatest advances in our knowledge of the activities of the organism were made; while the periods during which vital istic views were generally current were sterile ones from the same point of view. Modern Mechanisms of Lite. Now one may ask what is the difference between the Cartesian mechanism and that which is represented in, say, the writings of Jacques Loeb, or is there really any difference ? To Descartes there was nothing in the activities of the animal body but matter, the configurations of matter, and the motions of matter. Everything was mechanical. There is no doubt at all as to what he meant. " I wish it to be considered," he says in the Discourse on Method, " that the motion which I have now explained " (that of the circulation of THE MECHANISTIC CONCEPTION OF LIFE 161 the blood) " follows as necessarily from the very arrangement of the parts which may be observed in the heart by the eye alone, and from the heat which may be felt with the fingers, and from the nature of the blood as learned from experience, as does the motion of a clock from the power, the situation, and shape of the counterweights and wheels." But to the nineteenth- century biologists there seemed to be no activities in the plant or animal body except those of physical and chemical reactions. What happened in the processes of animal and vegetable meta- bolism was the result of the chemical constitution of the sub stances that compose the living organism. Here we must go back to Descartes' cosmogony. In the extreme generality of his hypotheses, in prophetic anticipation, and in sheer dynamic mentality no one has excelled the great French philosopher. " His imagination," says Clerk Maxwell, himself a man highly gifted with just the same qualities, " knew no "bounds." He made not only a cosmogony, but a cosmic evolutionary process. There was nothing, he said, but matter in motion, but matter itself was only extension. Figure and solidity were not essential to a material body, for the latter could be melted or dissolved or broken down without ceasing to be material, and its colour and smell and other qualities that make appeal to our senses were clearly not essential to its materiality. Nothing was essential but the condition that it occupied space. (This anticipates the theory of relativity.) There could be no void, or vacuum, he said, because empty space could only be conceived in terms of matter, which is extension. There could not be action at a distance (here he anticipated Newton), and the whole universe consisted of the same kind of matter (anticipating modern spectroscopic research). The universe was full, and was a continuum (thus anticipating our modern concept of the ether). Matter originally lay together in closely fitting, angular blocks, but it was set in motion (by God), and so these blocks suffered mutual attrition, grinding each other down and becoming spherical. An exceedingly fine dust or material resulted from this attrition, and this was relatively inert and formed the Cartesian " first element." The rounded particles, which were very small and were in active motion, made the "second element," while there was a " third element " consisting of particula striata that is, particles which had acquired a spiral shape by passing 11 162 THE MECHANISM OF LIFE between the rounded particles of the second element, so that A \ they possessed a rotatory motion as well as a translational one. ] The first matter made up the sun and stars that is, the cosmic bodies that are hot and radiate; the second matter made up the ' heavens that is, the atmosphere and cosmic space, which j convey radiation ; and the third matter was that of the earth and other planets, bodies that are cold and are becoming inert.; By vortices in the first and second matters he tried to explain (but with no more success than Kant and Laplace) the evolution of solar systems, while by the motion of the spiral particles he] sought to explain attraction. The particles of the second ele-j ment correspond to our modern atoms (or molecules, as Clerk j Maxwell calls them). To what do the fine particles of the first ; element correspond in our cosmogony ? Fifty years ago we should have said that the universe was j made up of atoms, of matter which here and there were aggre- j gated to form stars and planets, comets, nebula, and meteoritic j dust in short, cosmic bodies attracting each other with forces depending on their masses and on the distances between them and that these bodies filled only a most insignificant fraction of empty space. But they attracted each other across this space, j and heat and light and electro-magnetic radiation were trans- mitted through it, so that it could not be empty. There was a medium the ether of space and this conveyed the radiation, j So much had to be postulated merely to explain things and give us a " working hypothesis," but what was the nature of the ether ? It was something that was perfectly elastic, a kind of ; jelly, so to speak, across which energy could pass without dis- sipation. It had to be continuous and dense that is, there \ could be no interspaces in it such as there are even in the densest \ kinds of gross matter, which, Sir Oliver Lodge tells us, has a texture like gossamer compared with that of the ether. Now the latter cannot be absolutely continuous and perfectly elastic according to the modern theory of energy as formulated byJ Planck, for if it were it would possess " infinite degrees of freedom," and thus it would absorb all the energy in a system, leaving none at all to the matter of the system, and that we, cannot believe. So quite lately physics has come back again to the notion of a discrete ether of space something which is particulate, but which must nevertheless be continuous in some \ way or other, how it is difficult indeed to see ! THE MECHANISTIC CONCEPTION OF LIFE 163 But that would be Descartes' first matter, the exceedingly fine dust resulting from the attrition of the primitive matter which fills up all tEe interstices between the other molecules. Since the universe was originally quite full, it must have remained] full, even when its motions had evolved the three elements.' Therefore there is continuity and yet a particular structure, and that seems to be what the Planck theory of energy requires. After many vicissitudes, therefore, physics returns to what seems, in essentials, to be the Cartesian cosmogony. And, of course, our physico-chemical mechanism reverts to that of Descartes. There is nothing but matter and its con- figurations and motions, and this is true whether we think about Descartes' matter or the Newtonian matter, or the modern matter of electro-magnetic theory. The Cartesian matter was extension, but what is the matter of to-day ? We are gradually accustoming ourselves to think about it as immaterial. Atoms are merely systems of electrons, but electrons are pure energy, and energy, as we have seen, is something about the " nature " of which we do not know anything all that we know is that it is a measure of causality, and that we can measure it in terms of work done, and that, we have seen, comes in the long-run to measuring a distance or space. So our matter is extension, just as was that of Descartes, and we ought now to call it substance in the philosophic sense of the term. Anyhow, our mechanism of the organism has come again to a crisis. First of all it was a mechanical explanation of life, and that being insufficient biology resorted to a physico-chemical explanation, which was also insufficient, since physics and chemistry are again becoming mechanical. Looking about for the new conception that biology has now again to borrow from physics, we have little difficulty in finding it, and it would appear as if it were really something new. The concept is given to us in the physical notion of statistical mechanics, and to this we shall return presently. CHAPTER X THE MEANING OF PERCEPTION WHAT we have done so far has been to make a summary of the main results of physiology. Now it is obvious that this summary has been a very bare one, but we have so designed it that the reader should have little difficulty in greatly amplifying his knowledge by the study of a good, recent textbook. Assuming, then, that he has done so, it will be seen that the outcome of both medieval and modern anatomical and physiological research has been to give us an analysis of the means of acting of the animal mechanism an analysis that has become more and more refined as our chemical and physical methods of investigation have become more powerful and penetrating. It is quite easy to show this by tracing out the history of i physiology with respect to any one particular mode of function- ing for instance, conduction within the nervous system. Thus Descartes made an explanation of action which was based: entirely upon anatomical studies, and which, as we have just seen, involved only mechanical ideas. The nerve joining a sense \ organ the eye, for example with the brain contained one kind of fibres, which were, in effect, threads stretched between the! receptors and certain valves in the walls of the cerebral ven- j tricles. When these threads were stimulated they were jerked in some kind of way, and this jerk opened the valves and allowed animal spirits to escape from the reservoir in which they wen se Dense l^a. re. TZare. FIG. 47. DIAGRAM OF A SERIES OF SOUND WAVES IN THE AIR. The distance apart of the dots represents the condensation and rarefaction of the air. Thus the older mechanics and the classical chemistry deal with the displacements of bodies or of molecules or atoms that is, they deal with movements of material substance while the newer physics and chemistry now deal with radiation which occurs in a medium (the ether) invented to enable us to describe the radiation. Everything now becomes vibratory motions or waves. What is a wave ? "If you ask a mathematician," says Sir Oliver Lodge, " what he means by a wave, he will probably reply that the most general wave is such a function of x and y and t as to satisfy the differential equation: while the simplest wave is: y=a sin (x vt), and he might possibly refuse to give you any other answer." He would, no doubt, be sure that you ought to know what a differential equation means, anyhow ! Well, all science that is quantitative expresses itself in such statements: the ether is really a set of differential equations. When a scientific result reaches such a form it tells us that there are certain ratios of displacements. In the above expression the d's are differentials THE MEANING OF PERCEPTION 169 that is, differences in measurements of y and x and t that may be as small as we like; y is a measurement of space in a certain (vertical) direction; x is also a measurement of space in a horizontal direction perpendicular to that of y ; t is a time o measurement, and v is a velocity that is, it is a ratio, -. Time, in scientific investigations is always a space measurement; if, for instance, we use a clock, time is the position of the hands relative to their zero position. Thus the equation represents nothing but the ratios of measurements of space; the crest of longitudinal motion of crest (a1~ihe moment t,) Fio.^48. DIAGRAM OF A WAVE IN Two POSITIONS. the wave, which we may take to be an Atlantic roller travelling uniformly, moves with a certain velocity, v that is, at a certain moment, t lt the crest of the wave, which is travelling in the irection ox, is at a certain position (x=o), and the water level s at a certain height (y^. A moment later (t 2 ) the crest is at a different position, x l9 and the water level, which was originally at the position represented by y v has now sunk to the posi- tion y. 2 . Obviously, then, our differential equation tells us how the crest of the wave is displaced along the direction ox, while the water level at any position in the line ox is being displaced along the up and down direction oy. It takes a moment of 170 THE MECHANISM OF LIFE time, t, at which the crest is at a certain position, and then an " interval " of time, dt, after which the crest has attained a new position, dx. But at the moment t the water level was at the position y, and after the interval dt it has dropped to the new level dy. We call the d?s " intervals," but it must be noted that they are " infinitesimal " intervals that is, they may be smaller than any finite lapse of time, no matter how short the latter may be. Clearly, then, our differential equation simply gives us a series of connected displacements. As time changes that is, as some standardised mechanism, such as the hands of a clock, are displaced by so much so the crest of the wave is displaced so much in a horizontal direction, and the water level is displaced so much in a vertical direction. Now all this is not at all irre- levant to the work of this chapter, for our differential equation is a perfect example of a scientific " law." Science measures motions, or, rather, displacements, and its residts must be expressed in measures of space. Next we turn to the study of organic functioning, and we may choose for an example any bodily activity whatever; in the meantime our treatment of the animal is entirely " objective " that is, we are regarding it as a system in the physico-chemical sense, something outside ourselves that we can observe and measure just as we can an ocean wave. We might take the action of the heart, or the interchange of oxygen and carbonic acid between the atmosphere and the bloodvessels of the lungs, or the excretion of urea it does not matter. In all these cases we investigate and express our results as motions, or, more precisely, as measured displacements of something. When we study the circulation we measure the velocity of the blood- stream (that is, the displacement of a blood-corpuscle in an unit interval of time), or the pressure of the blood in an artery (that is, we record the displacement of mercury in a pressure gauge connected with the bloodvessel), and so on. When we investigate respiration we may measure the tension of oxygen in the blood and compare it with the partial pressure of the gas in the air (that is, we observe and measure the volume of a gaseous residue after certain manipulations). When we study the formation and distribution of urea in the blood and kidneys we make quantitative chemical analyses, and in so doing measure the masses of certain substances at certain times and places, and THE MEANING OF PERCEPTION 171 compare them with, the masses of the same substance at other times and places. And in identifying these chemical substances we make use of measurements of space (colour, specific gravity, vapour pressure, boiling-point, chemical properties in general, which are all consequences of the positions of atoms within molecules and of the movements of the atoms and the molecules), and so on. To save time we may assume that the reader is thoroughly conscientious, and desires to verify these statements ; he will find that the results of the investigation of animate, no less than inanimate, activities all reduce down to observation of motions and measurement of displacements. But we shall not, or we shall very rarely, get differential equations like we do when we investigate the ether and elec- tricity and relativity, and so on. We may illustrate this by considering growth. Think of a crystal of salt growing in ideal conditions in a pure solution of sodium chloride: if we know the length of one of its sides we can find its mass, thus : Mass=c (length) 3 , and the differential equation is - =2cZ 2 (where m=mass, dl Z=length of a side, and c is a constant depending on specific gravity). Knowing the length of one side of the crystal, we can predict its mass. We cannot do this with a growing animal. If it grew like the crystal does, we could use the equations just given to find the weight of the animal from a measurement of its length. We can- not find this, as experiment will show, and all that we can do is to make an empirical equation after we have measured both length and mass. This equation will be : Mass=aZ+6Z 2 +cZ 3 -h where Z=the length, and some of the constants, a, 6, and c, may be negative numbers. Go on now to a study of behaviour, and the same contrast emerges. Nothing in the way of differential equations is ever possible, and we cannot, in general, predict how a normal animal will respond when we attempt to stimulate it to activity of some kind or other. Yet we can think of a series of organic responses which begin by showing a high degree of determinism and end by failing to show any. 172 THE MECHANISM OF LIFE Perhaps we had better define physical determinism, or, rather, illustrate it by an example. A compass needle free to move points in London in a certain direction (about 16 degrees to the west of true north), and it does so because the earth's magnetic force is directed along lines which are running in that direction. Now we know this horizontal component of the earth's mag- netism, and we can also find what would be the force exerted by a feeble bar magnet placed at right angles to the magnetic meridian in which the compass needle is situated and (say) a foot away from the pivot of the latter. Then a physicist could calculate the angle through which the needle would deviate when the magnet was placed as we have indicated. This deviation would always be the same provided that the earth's magnetic field and that of the disturbing magnet remained con- stant. Call the effect of the latter the " stimulus." and the deviation of the needle the " response"; then there is physical determinism, and we can always successfully predict to a high degree of approximation that a certain response will follow the application of the stimulus. Now take a muscle-nerve preparation (see pp. 1345) and stimulate the nerve by a feeble electric current; a momentary twitch of the muscle, exerting a certain pull upon a weight attached to it, follows. So long as the muscle remains irritable and unfatigued this response occurs, and we can predict it. There is physical determinism, even although this may not be absolute. Next take the frog in which the brain has been destroyed, leaving the spinal cord intact. When a drop of acid is placed on its back it will wipe this away with one of its hind-legs, and when other stimuli are applied it will respond in a mechanical, automatic manner. We can, in general, predict what it will do when certain stimuli are applied in certain ways; thus there is determinism. Yet this is not absolute for instance, the spinal frog appears to prefer to use one leg rather than the other, but if it is prevented from employing that leg it will use the other. Generally the behaviour, in response to stimuli, of the spinal animal is of this mechanical, automatic nature it can usually be predicted; and this statement is true of frogs, birds, reptiles. and mammals, whatever animals have been the objects of the experiments. It is also true of a certain number of reflex actions in the normal animal that is, it is generally true, for reflexes cannot THE MEANING OF PERCEPTION 173 always be elicited. It is true of habitual, learned, and mechani- cally repeated actions which have, so to speak, become customary to the animal that makes them. And it is true (again in general) of the instinctive actions which make up a considerable part of the behaviour of invertebrate animals in particular, and even of the higher vertebrates. Yet habits can be varied, and instinctive actions are not quite invariable. Thus some animals, such as bees, which build structures of certain materials, may alter their methods when different materials are supplied to them. Clearly the responses made even by intact, normal animals are not rigidly determined ones capable in all circumstances of being predicted. And such behaviour as we have indicated in Chapter VIII. in dealing with the functioning of the nervous system shows that there may be spontaneity of behaviour on the part of all animals that have been studied with sufficient care. They react to stimuli sometimes in one way, sometimes in others, and again they may not react at all, or they may act in unpredictable ways even when there are no apparent stimuli. One can see this in men and women: a slight irritation may cause a man to cough again and again, but he may easily repress the response, even when the same physical stimulus recurs. Now in all such cases of variable behaviour to the same kind of stimulus it is open to us to say that there may have been other unrecognised stimuli which modified the response; that in cases where the behaviour seemed to be absolutely spontaneous there " must have been " some causes which we could not trace, or that there were " physiological conditions " introducing elements of uncertainty, and so on. All this may be so ; still, the fact is that in a great many cases we cannot say that organic actions are preceded by definite specifiable causes, and when we say that a response " must " always have its appropriate stimulus we simply dogmatise. It hardly seems necessary to lay stress on the conclusion that we are about to make; nevertheless, it is well to do so. Inanimate occurrences are always " more or less " determined, and equivocal results invariably suggest defects in the experimental methods employed, or neglect of some condition or other, or a degree of complexity incapable of complete analysis. That a result can be predicted is a kind of test of a successful physical investigation in fact, we disregard all other results; they are not useful to us. To some extent this physical determinism is true of the functioning of the lower animals and of the higher ones that 174 THE MECHANISM OF LIFE have been subjected to operations which destroy certain parts of the central nervous system. But in most animals there is some indetermination and spontaneity of behaviour, and the more highly organised is the central nervous system, the greater seems to be the degree of indetermination that is exhibited. The Nature of Sensation. We are now about to cross over from the " objective " to the " subjective," and the reader is asked to attach no other mean- ings than those that we specify to the technical terms that we shall employ. By " sensation," then, we mean only the train of physical events which occur when a receptor organ is affected or stimulated by some change in the external medium, and when, in consequence of this stimulation, nervous impulses are pro- pagated along the afferent nerve of the receptor organ into a cerebral or spinal centre. Let us examine this train of events in one case ; the reader can easily make similar analyses for others. The Sensation of Hearing. A gramophone begins to play- that is, a disc in which there is a spiral groove of uneven depth revolves, raises and lowers a needle, which presses unequally upon a diaphragm and throws the latter into rapid vibrations. These vibrations set up waves of condensation and rarefaction in the air, and the latter impinge on the drum of the ear, setting this into vibrations which resemble those of the diaphragm of the gramophone. The vibratory movements of the tympanic membrane are communicated to the three little bones of the middle ear, which communicate them to a liquid (the peril ymph) filling the bony labyrinth of the internal ear. Then the vibratory movements of perilymph are transmitted to the endolymph contained in the membranous labyrinth, and the latter vibrations are communicated to the hair cells or other terminations of the cochlear nerve. Nervous impulses are now set up in the fibres of the nerve of hearing, and these are propagated to the auditory centres of the lower brain. There is therefore a chain of dependent events : the mechanism of the gramophone, the atmospheric vibrations, the movements of tympanic membrane and auditory ossicles, the vibrations of perilymph, endolymph, and nerve terminations, the nervous impulses, and, finally, the changes produced by the latter in the cells of the auditory centre. Throughout all this series of changes, or motions, there is physical determinism. THE MEANING OF PERCEPTION 175 Are the changes that occur in the cells of the auditory centre heard ? They may or they may not. If a gramophone starts to play in the house next door just as a man is going to bed he will certainly hear it ! But a man who is intent on some mental work may not hear a knock at the door, while someone who is with him but is not busy may hear it. And, of course, while he walks through a busy street his eyes and ears receive a multitude of stimuli, and, no doubt, transmit these to his brain, but he is not aware of a large fraction of all this stimulation. Sensation, then, may or may not be accompanied by changes of conscious- ness. Now there is a stimulus leading to sensation, but there is no " awareness," and, again, when the same stimulus is repeated, we hear or see. Why in one case and not in the other ? There is a " psycho-physical " law which attempts to relate together the intensity of a stimulus that can be measured and the intensity of the feeling of awareness that a sense organ is being stimulated. Let the stimulus be a weight placed on the hand held out straight, and let the subject be asked to say when he feels the increase of weight, as (unknown to him) increments of mass are added to the latter. Say that the weight is P, and let small mcrements'AP, 2AP, SAP, etc., be added to P ; the subject will not recognise an addition to the weight until this addition has become a certain fraction of the original weight. For instance, he will not recognise that P+ AP is greater than P, nor that P-J-2AP is greater. He witt recognise that P-J-3AP is greater than P, though he will not recognise that P+3AP is greater than P+2AP. Suppose, now, that there is a strict determinism, and that every perception is a function of its stimulus; then it is clear that /(P)=/(P-hAP)=/(P+2AP); also /(P). Kelvin, and Gibbs. Then came the electro-magnetic theory of j Clerk Maxwell and those who followed him, and, in our own time, the French and English work on radio-activity and the new physics] built up by J. J. Thomson, Planck. Einstein, and Nernst. This latter theory, which is still in the making, is obviously the bridge that will lead us across the gulf between matter and the physics of the ether. Apparently, we have yet to find whether or not] physical science has said the last word in its effort to explain life. The Laws of Thermo-dynamics. Go back now to our discus- sion of the laws of thermo-dynamics. The first law, we have seen, ' is really a kind of mental postulate, or convention. We make up our minds that there is something that is permaiient that can neither be created nor annihilated in a.11 the changes that occur in nature. This something we call energy. If it appears to arise out of nothing, or to vanish into nothing, we simply do not believe that apparent result, and we proceed to invent poten- tialities that will account for the appearances or disappearances. Usually we are successful, and we find that our hypotheses of i potential energies work, and we are so led to further discoveries. Then we say that the things we are investigating are real ones, since they are conserved. Or we may find (when we try to investigate spooks) that our hypotheses astral bodies, higher planes of being, telepathy, and the like do not work. They mislead us. The test of their validity is that they should enable us to predict, and we find no good evidence that they do so. Therefore, being useless to us, and rather a nuisance, we say that the phenomena of spiritualism are unreal. That is the first law that something is conserved so that the total quantity of it in our universe remains constant. It is not so much a physical as an a priori " law," or mode of our mentality. The second law is quite different, inasmuch as there is nothing at all a priori about it. It merely describes, in the most com- prehensive way possible, our experience of the universe. It tells us how things happen: that water runs downhill; that a red-hot poker taken from the fire cools down to the temperature of the room; that a cigarette burns away, leaving behind it some ash, water vapour, and carbonic acid gas, and evolving heat as it ON THE NATURE OF LIFE 195 burns. These things seem to us to be so inevitable that we find it difficult to imagine them happening in the opposite way. It appears to be ridiculous to think of water running uphill of itself; of a table rising up of itself from the floor ; of a cold poker becom- ing red-hot by exposure to the air; or of the ash, the water vapour, and the carbonic acid combining together (with absorp- tion of heat) to form a cigarette. But why should not these things happen ? Puzzle over this, and we find we can give no reason other than that no one has seen them happen. Nevertheless, there is no logical reason why they should not happen, and we can easily imagine them to do so. We can even picture them or imagine ourselves so situated that they appear to happen. Take the case of the red-hot poker: it is red-hot because its molecules are moving so rapidly that their atoms radiate and so give off visible vibrations. While they glow they are vibrating much more quickly than are the molecules of air which come into contact with them, and so they accelerate the velocity of the latter just as a quickly moving billiard ball would accelerate the motion of one that is moving less rapidly when the two collide. There is, therefore, an excess of kinetic energy in the motions of the molecules of the poker compared with that in the molecules of the air in the room, and so the excess becomes levelled down, so to speak. When the poker has cooled to the room temperature all the energy which it contained is still in existence, but it is now uniformly distributed between the metal and the air which came in contact with the latter instead of being concentrated in the poker. Imagine that we could actually see the molecules and their motions, and that we could photograph them so as to make a kinematographic record. Working the latter, we should then have a picture of the transfer of molecular motions from the hot metal to the cold air, but if we were to work the record backward just the same motions and. molecules would be thrown upon the screen, but in the reverse order. W^e should first see molecules of air and metal moving at certain rates in equilibrium with each other, and we should then see the air molecules begin to move more slowly and to give up some of their kinetic energy to the molecules of the poker, which would move more and more rapidly, until they become red-hot and give off visible radiation. Or imagine a record of a cigarette smoker to be worked back- wards ; we should then seem to see the smoke and ash solidify to 196 THE MECHANISM OF LIFE form the cigarette, and the latter to grow to its original, unlit, unsmoked size. If the molecules were visible, we should see those of the ash, the water, the carbonic acid gas, and the particles of smoke combine together to make up the molecules of the cellulose of the tobacco and paper. We should see energy taken up from the air and become concentrated, so to speak, in the glowing cigarette end, and then pass into the potential, chemical state in the tobacco and paper. Or, again, take the famous illustrations of Einstein's theory of relativity. A man looking down from a stationary balloon might have seen the explosion of a mine on the Menin Ridge, but suppose that, during the few seconds of that occurrence, the; balloon had been moving away from the earth with the velocity of light; then our observer would have seen nothing happen, although someone in a stationary balloon would have seen the earth, stones, smoke, etc., thrown up into the air. Suppose that the observer had witnessed the explosion from his balloon, and that the latter had then immediately started to move away from the earth with a velocity greater than that of light; then he would (assuming he had a telescope powerful enough) have seen the whole train of events proceed backwards. Smoke, stones, dust, and earth would appear to come together from nowhere and coalesce to form the solid ground. Apparently, then, there is no logical reason why any physical change should not go in either direction. Water runs downhill, but it might run uphill. (Note that we can make it go uphill by forcing it through a closed pipe, but what we are to imagine is it running uphill itself. We can spend energy in forcing it up, and then we find that we have spent more power than can be recovered by allowing the water to run downhill again and work a pump.) We can think of, or imagine, or picture, the water going uphill of itself, and we cannot find any reason why it should not except that it does not. In the abstract, therefore, all physical changes are reversible, but it happens that we are living in an universe in which they are irreversible. This is our experience, and we can generalise it in several ways. The most comprehensive way of doing so is to say that in all things that happen a certain mathematical func- v tion, called entropy, increases in quantity. What is meant by entropy we shall explain presently. Note, in the meantime, that the necessary condition that any phenomenon or physical change ON THE NATURE OF LIFE 197 may happen in our universe is that there must be available lergy, and that this must transform into the inavailable form, [n other words, energy must be concentrated somewhere, and when a phenomenon occurs it becomes dissipated, or spread out. Having become dissipated or spread out, nothing more can happen, for this energy has lost its capacity for doing work. The second law of thermo-dynamics states that in all natural changes energy IX becomes dissipated, or entropy increases. The Condition of the Universe. Next, we must consider this problem, Is the universe finite or infinite in space and duration ? Or we may put the question more clearly. If we were able to travel away from our earth into outer space, like Richter's Dreamer, should we at length enter regions where there were no longer any stars ? Or if we could, like Mr. Wells's Time Traveller, go indefinitely far back into duration, should we come to a time when there were no earth and stars ? It may seem foolish to suggest these questions, but, as a matter of fact, we can, pro- visionally at least, answer the first one. There does seem to be a limit to the number of stars in the sky, and it seems that these form a cluster, or galaxy, which is finite. If the numbers of stars were really infinite, certain consequences would be the result. If there were no absorption of light by dark cosmic bodies the night sky would be an uniform blaze of light, and even if there were absorption of light energy would still be infinite in some form or other. Such is not the case, and so we conclude that the stellar universe which radiates energy is a finite one.* Is its past duration also infinite ? We can easily imagine ourselves travelling further and further away from the earth without limit, and passing out at last from the region of stars, or, more generally, of available energy. But we cannot think of a time in the past, however remote, when the universe did not exist, when there was no available energy. For the quantity of available energy in our universe is decreasing, and in the past it must have been greater than it now is. Was there a time, then, when energy came into existence from nothing ? That implies creation, and, since we hold to the law of conservation as a mode of our thought, we cannot, therefore, think of a time when the uni- verse began. We conclude, then, that its past duration is infinite.f * The theory of relativity helps us here. The universe that we see is finite, but unbounded. Its form of space is four-dimensional and spherical. See Appendix II. t See Appendix II. 198 THE MECHANISM OF LIFE Now we take the plain result of our experience that available energy is continually decreasing in quantity (or that the entropy of the universe is increasing). This is the bed-rock fact of our experience: that everything that happens depletes the universe of its store of available energy. " Every energy transformation that occurs leaves an indelible imprint somewhere on the course of events in the universe considered as a whole." There was an original quantity of available energy in our universe, and this is becoming exhausted as entropy increases. Now this quantity being finite, and duration being infinite, it follows that the time must come when there will be no available energy left (entropy having attained its maximum), and so there must occur a com- plete cessation of all energy transformations, or phenomena. But if past time is infinite, why has not this physical death of the universe already occurred ? No matter how great a lapse of duration is required, we can still imagine this to be possible. Here, then, we have our physical impasse. The second law, solidly based on our experience, says that entropy tends towards a maximum, when universal happening must cease. Past time is without limit, so that, no matter how slowly entropy is being augmented, the maximum ought to be attained. But the uni- verse is still the locus of energy transformations, so that entropy has not attained its maximum value. Obviously the second law cannot be universally true, although .is true of our experience. Somewhere or other, or at some time or other in the universe, it must be reversed or evaded. Now since it is not logically necessary that entropy should always tend towards a maximum value, we must next enquire under what circumstances the second law can reverse itself; under what conditions may water flow uphill of itself, or may heat flow of itself from a region of lower to a region of higher temperature. The Reversal o! the Second Law. In order that such an investigation may be possible we must choose some simple, mathematically manageable case. Consider, then, the physical condition of a small volume of some gas, say hydrogen, at ordinary temperature and barometric pressure; it consists of an enormous number of molecules which are moving very rapidly. and so incessantly colliding with each other. It would be rather like a swarm of midges in the air provided that the insects fl-\v about in straight lines instead of avoiding each other. In a volume of hydrogen equal to 1 decilitre (that is, a cube of nas ^it i ON THE NATURE OF LIFE 199 rather less than 2 inches along each side), there will be 100x2-705xl0 19 molecules. Each of the latter is moving, but the velocities are variable within certain limits, and their average is about 1-65X10 6 cm. per second. No molecule can move very far without colliding with some other one. and the average free path of a molecule is about 0-00000182 cm. We may regard them as small, spherical bodies of about 2-17xlO~ 8 cm. in diameter. The average velocity of movement depends on the temperature (or, rather, what we call temperature is the variable velocity of the molecules; the higher the latter the higher is the temperature, and vice versa). Evidently the molecules will be moving in every conceivable direction, and so they must collide with each other in all sorts of possible ways. Usually the collisions will take place at some ansjle to the directions of movement; sometimes a rapidly moving \ (Z)* * f3*f \x FIG. 50. DIAGRAM OF CHANCE COLLISIONS BETWEEN MOLECULES IN A GAS. molecule will overtake a more slowly moving one travelling in the same direction, and sometimes two, which are moving in the same direction, but towards each other, will collide " end on." Being perfectly elastic bodies, no energy will be dissipated in such collisions. After the collision the directions and velocities will be changed in ways that are easily worked out from the well-known theorems of mechanics. The reader may easily construct these results by making use of vector diagrams, such as those of Fig. 50. Now by certain mathematical formulae (deduced by Clerk Maxwell and others from the theorems of probability) the relative frequencies with which encounters between pairs of molecules moving in all possible directions, and with all the possible velocities lying between the upper and lower limits, can be calculated. An encounter in any possible way (such as in the Cases 1 to 4 of Fig. 50) is equally likely to occur. But there 200 THE MECHANISM OF LIFE is one case which is of particular interest to us, that of two molecules moving end on with the same velocity (Case 4), and at any moment a certain very small fraction of all the pairs of molecules must collide in such a way. It is possible that all, or |, or J, or T ^, and so on, of the total number may collide end on, but it is much more probable that y^ will collide end on than will }, and so on. It is possible, we say, that all the molecules will at the same moment collide in pairs, and end on, as in Case 4, Fig. 50, and the probability that this sort of encounter may simultaneously occur throughout the whole volume of gas can be calculated. Now, though possible, the chance of this occurrence is almost incredibly small; small as it is, however, we must consider it in speculative reasoning. Imagine now our decilitre of hydrogen enclosed in a box divided into two equal parts by a partition, and imagine the walls of the box and the partition to be made of some material abso- lutely impermeable to heat. Let there be a hole in the partition closed by a valve which is also a non-conductor of heat. Let the gas in the right-hand side have a temperature of 20 C., and that on the left-hand side a temperature of 10 C. In such a case the system contains available energy. There is the same mass of gas in either division, but that on the right has a temperature of 20 C., and so its pressure is higher than the gas in the other division. If the valve is now opened the gas at high pressure will rush through the aperture, and it can do work (say, by turning a small propeller) while the pressure is being equalised. But when the latter condition has been attained the system, in itself, can do no more work. Its total energy is still the same, but there is no difference in intensity, and so no transformation (with regard to pressure change) can occur.* Consider, also, what happens while the temperatures and pressures are being equalised. The gas in the right-hand division is at a temperature 20 C., and contains a certain quantity of heat, Q'. After the transformation has occurred, the whole mass of gas attains the average temperature 15 C., because Q units of heat have now flowed from a region originally at 20 C. to a region originally at 10 C. Now replace ordinary temperatures by absolute ones that is, add 273 C. to each of the former; * For simplicity we neglect here the conditions under which the trans- formation must occur to render the calculation that follows applicable. ON THE NATURE OF LIFE 201 Q units of heat, then, have flowed from a mass at 293 (or 273+20) to another mass at 283 C. (or 273+10). When heat flows from a body at high temperature to another body at low temperature, the entropy of the former is diminished and that of the latter is increased. Change of entropy is measured by the simple ex- pression -, Q being the quantity oJ^e^^wMcluDJBes-aiid JT_ being the absolute temperature_. The entropy of the gas at 293 is therefore diminished by the amount 293 But at the same time a mass of gas. originally at 10 C. (or 283 Abs.) becomes raised in temperature to 15 C. (or 288 Abs.) as the result of Q units of heat entering it, and so we get the change due to the receipt of Q units of heat by the colder gas as . Now the total entropy change due to the Q units of heat 288 lost by the hot gas, and the Q units gained by the cold gas is + - . Obviously this expression is positive, and so we get our result that entropy increases as the result of the mixture of a hot with a cold gas. Let, then, all this occur in the case of our specified decilitre of hydrogen. The mixture of gases takes place in a few seconds, and the resulting gas, at a temperature of 15 C., is stable and homogeneous as regards temperature. No further work (with regard to its temperature) can be done by it. But while that is the case, it is still possible that the state of all the molecules in the gas may become such that at a given moment the latter become disposed, purely by " chance," in pairs, the members of which are approaching each other with equal velocity in the same straight line. When they collide their velocities will be the same as before, but the directions in which all the molecules are moving will become simultaneously reversed. A very important result flows from this. It can be shown (though the proof cannot be given here) that when every mole- cule completely reverses its direction of motion, the velocity remaining the same, the gas will retrace its past history. That means that the total quantity of gas at a temperature of 15 C. (or 288 Abs.) will separate into two portions one at a tem- perature of 10 (or 283 Abs.), and the other at a temperature of 20 C. (or 293 Abs.). Therefore Q units of heat will flow from 202 THE MECHANISM OF LIFE a region of 288 to a region at 293, and entropy will decrease by the amount . At the same time Q units of heat will enter 288 a region at a temperature of 293, and entropy will increase by the amount . Therefore the total change is }-- , and 293 288 293 obviously this expression is negative. Now we get the result that when, by reason of the reversal of the direction of motion of all the molecules of the gas, the latter segregates into regions at different temperatures entropy decreases. We have now obtained a result of considerable interest. It has been shown that it is theoretically possible that a gas origi- nally at an uniform temperature can, of itself, separate into two equal portions, one of which is at a higher temperature than is the other. Heat can flow of itself from a region of lower to a region of higher temperature. A gas which in virtue of its temperature alone possesses no available energy can pass, of itself, into a condition in which it does possess available energy. A system can, of itself, decrease its entropy. All this is very surprising, for these statements mean the same thing that is expressed by saying that water can, of itself, flow uphill ! They mean that we can state the second law of thermo- dynamics as follows: The entropy of the universe may tend to a maximum or a minimum. But we must consider the probability of this reversal of the sign of the second law of thermo-dynamics, for our results merely give a theoretical possibility. The condition that it may occur in such a case as we have investigated above is that at the same moment all the molecules of the gas are arranged in pairs, and the members of each pair are approaching each other end on and with the same velocities. Now this condition is only one of an incredibly great number that may exist in the gas, and the probability that it may exist is the ratio, unity to a very large numerical value. Perhaps the reader will best appreciate what is this probability if we put it in Boltzmann's own way. In order that we may observe this reversal of the second law in a decilitre of gas, we might have to wait for a number of centuries repre- sented by unity followed by 10,000 millions of zeros ! It is as probable that it may occur as it is probable that every house in London may catch fire accidentally and independently of all the others on the same day, or that every grown-up person in London ON THE NATURE OF LIFE 203 may commit suicide (also indepeadently of all the others) on the same day. Now an insurance company taking risks of houses being burned down by accident, or of people committing suicide, calculates its possible liability from the application of the theorems of probability, and it would safely ignore such risks as those we have just mentioned. The latter are incredibly small. The chance that a decilitre of gas will separate into two portions of different temperature is also incredibly small, but it is not zero. In the ordinary affairs of life we neglect small risks, and say they are " practically zero," or " infinitesimal," but when we apply such chances in specula- tions concerning the origin and fate of the universe we must not dismiss them unless we are sure that they are really insignificant in the conditions. Now we are not going to extend Boltzmann's results obtained from a study of the kinetic theory of gases to the whole universe that is, we must not suppose that a reversal of the second law depends on the collision end on of every mole- cule. All that we suggest is that it is possible in some way other that entropy may decrease in our universe instead of increasing. This is a logical possibility, and, given certain arbitrary conditions in a very limited system, it is a probability, the numerical value of which can be estimated. Further, we are compelled to postulate that somewhere or other, or some time or other, the second law of thermo-dynamics must reverse itself that is, some time or somewhere entropy must decrease, or have decreased, in our universe, otherwise we shall be compelled (as Sir William Thomson was) to postulate a beginning, or creation. Disentropic Phases in the Universe. Neglecting, in the mean- time, the numerical value of the probability that entropy may decrease (or that unavailable universal energy may become available), we. may now proceed to consider the possible history of our universe, taking as our " conceptual model " the changes that occur in a small volume of gas left to itself. In the latter, then, incredibly great periods of time may pass, and during these the molecules of our gas are moving and colliding in a haphazard fashion. Nothing happens in the system considered as a whole; it does no work, although the total quantity of energy contained in it is conserved. But some time or other, if we wait long enough, the system changes, and its original, segregated condition becomes restored. This condition in which the gas system contains available energy and can do work lasts for an in- 204 THE MECHANISM OF LIFE finitesimal period of time, and the " normal " condition of maximum entropy is again attained. The visible universe that is, our galactic system of radiating stars has, we have reasons for believing, definite boundaries. If we could travel out from our sun in any direction for about 30,000 light years* (that is, 30,000x365x24x60x60 seconds X 186,000 miles) it is possible that we should reach those boun- daries. But beyond this we can still imagine ourselves travelling on indefinitely far.f In this outer space there would, however, be no radiating cosmic bodies, though we might suppose that there are dark, cold suns and satellites, and cold, cosmic dust. Then, after incredibly further voyaging, we might encounter other galactic universes. This means that our picture of the entire universe is one in which the normal condition is physical death the complete cessation of all happening entropy having attained its maximum. But here and there in the whole uni- verse, and occupying regions that are of infinitesimal extent, there are individual universes, of which our galactic system is one. The normal condition of the entire scheme of things is that to which we see all physical changes tending, the complete dissipation of energy. But now and then, and for periods of time that are infinitesimal in duration, infinitesimally small regions of the entire universe blaze up, so to speak, the second law becoming reversed and entropy becoming decreased. After this has occurred, the individual universe then much more slowly sinks back again to the normal condition that in which entropy tends slowly towards its maximum and to which physical death is the limit. The universe, then, in which we are living is an incredibly small fraction of all that exists, and its genesis as an individual, physically active universe was an occurrence essentially similar to that of our gas model. Some time in the past the second law became reversed, and available energy was restored. Then this became slowly dissipated, so that the phase in which we are living is the more probable one that which tends always towards complete degradation of energy. In this way we avoid the physical impasse to which we are brought when we assume the universal validity of the first and second laws of thermo-dynamics. Our whole universe becomes a * A light-year is about six billions of miles. f Not on the theory of relativity. We cannot go beyond the boundary of the universe, for outside the latter there is no space. See Appendix II. ON THE NATURE OF LIFE 205 cyclic order, such that the most probable phases are those in which entropy tends towards its maximum value, and the least probable ones are those in which entropy tends towards its minimum value. As such it is a permanent universe, self- sufficient, without beginning and without end. Throughout its greater extent nothing happens, and this condition we call the normal one. Here and there, however, and for infinitesimally small periods of duration, there is physical activity, and this condition we call the abnormal one. The probability that any- where and at any time there is such physical activity is of the same order of magnitude as that calculated by Boltzmann for the reversal of direction of motion of all the molecules contained in a small volume of gas. A Digression on Orders of Magnitude and Duration. The principal difficulty in appreciating the force of such an argument as the above one lies in the reluctance with which we apprehend extremely small and extremely large magnitudes. We refer all measures, space, and time to those spatial and temporal values that limit our bodily activities, and if something is very great compared with these we are apt to say that it is " infinite," while if it is very small we say that it is " practically zero." Now a magnitude that we can estimate is always finite, no matter how big or how small it may be, and greatness and small- ness are always relative to something or other. The same magnitude may be extremely small compared with some other one, but extremely great when compared with yet another. Thus the radius of the earth (4,000 miles) is to us a fairly familiar magnitude, being a distance that we might traverse during a few weeks by means of a steamship. A micron, however that is> TTiVo millimetre, the microscopist's unit of length is a magnitude that is about I/ 6378X10 8 of the earth's radius, and we may call it an infinitesimal of the first order of smallness.* But, again, a molecule of hydrogen is about 2-17X10' 8 mm. in diameter, so that it is an infinitesimal of the first order when compared with a micron, but of the second order when compared with the radius of the earth. f Starting now with our standard magnitude that is, the earth's radius we may compare it with others that are " infinitely great "; thus it is about one-millionth * It is about i -billionth part of the earth's radius. 2-17 t 2-17 x 10 - 9 mm. = mm., say 1 thousand millionth mm. 206 THE MECHANISM OF LIFE of the radius of the orbit of Neptune that is, the limit of our solar system and so it is itself an infinitesimal. But the diameter of the solar system is also an infinitesimal when compared with the distance of the nearest fixed star (about 3J light years, or 3-25x6 billions of miles). Now let the reader calculate for him- self this series of magnitudes in terms of the same unit (say a kilometre), and he ought to have little difficulty in seeing that any one, however small or great, may have significance with regard to some other one. Consider now intervals of time. It is said that an ordinary person can easily realise, or feel, the lapse of duration represented by A second of astronomical time, and certainly one second may in some circumstances be a rather prolonged period. Now, A second is about r roWv~ i niUu n rth of an ordinary lifetime (that is, it is about 10 ~ n lifetime), and thus, with respect to the lapse of duration which we may call the standard one, 5 V second is an infinitesimal of the first order. But in looking at a red light for that length of time we receive some 400 billions of ether vibrations of a certain length that is to say, 4xl0 14 separate events or things that actually happen. One of these events, therefore, has a duration of 2 X 10 ~ 10 seconds, and this we call an infinitesimal of the second order with respect to the lapse of a lifetime. Probably life itself has existed on the earth for 1,000 million years, so that even a very long lifetime is only about Vo -millionth of the whole life-period of the globe. But the latter (10 7 years) must itself be only an infinitesimal fraction of the period of time during which our solar system has been in existence. The Meaning of Duration. For men and women this period of gV second of astronomical time is to be regarded as an unitary lapse of duration: it is the smallest period for which one may exist as a conscious, sentient being. While it passes, a star moves through an arc of about 1-5 seconds that is, a shift in the sky which can easily be measured by refined astronomical methods. In looking at red light for the same period one gathers up into perception some 400 billions of ether vibrations, each of which has a wave-length of 8xlO~ 4 cm. In listening to the note on the piano which is three octaves above the middle C one combines together about forty-one separate vibrations of the atmosphere to make a sound of a certain quality. Our rhythm of duration (in Bergson's phrase) is therefore such that we synthesise these motions, or changes, to make unitary, ON THE NATURE OF LIFE 207 individualised perceptions. We see the motion of the star through 1 -5 seconds of arc, we see light of a particular colour, and hear sound of a particular pitch. Each of these perceptions is there- fore a synthesis of external events effected during the same small fraction of an individual lifetime. Our sensory mechanisms are such that we can make these syntheses, but not some others ; thus there are ether vibrations which are less rapid than those corre- sponding to the red of the spectrum, but we cannot see them, though we can feel them as heat. Below these, again, are others which are much slower, and which we can neither see as light nor feel as heat, though we can detect them by the receiver of a wire- less telegraphic apparatus. Above the violet of the spectrum are vibrations which are so rapid that we cannot see them, though we can make them act upon a photographic plate. Other animals are certainly different from what we are in respect of these matters; thus a dog can certainly see light of lower wave-length than we can, V and some insects can hear sounds which are too acute for our ears. The reader will easily see, then, that rhythm of duration varies. Obviously it varies even in ourselves ; thus the familiar experience that time passes more quickly when one is fifty years old than when one is ten means that the later ones have fewer events in them that is, the periods of our duration which correspond with a revolution of the earth round the sun do not gather up so much of other external events when one is old as when one is young. To say that young people have a " fuller " life is not figurative, but is strictly scientific language. Now think of how the rhythm of duration varies in different animals. It is probable that the imagines (the fully meta- morphosed insects) of some species of Ephemerida3 which live for only a few hours can appreciate, or feel as distinct lapses of duration, very much smaller intervals of astronomical time than we do. Suppose, for instance, that a period of ^STTOOTJZF second can be felt by them (in which period they would still receive some 8,000 millions of vibrations when looking at red light) ; then their lifetime of a few hours would contain as much (that is, have the same duration) as ours of seventy years. Suppose, again, that some very long-lived animals, such as crocodiles, have unitary periods of duration that are much greater than ours; then their lives would be no longer, for they would contain no more. Note, however, that in such cases the universes con- structed from perception would be different ones. 208 THE MECHANISM OF LIFE Carry such speculations to their limits. Imagine minds which, just as we synthesise in perception billions of ether vibrations to make visible radiation, might synthesise hundreds of lunar revolutions or solar ones in a single perception. Or in such a mind all the changes that make up the origin, growth, and dot-ay of a solar system might be gathered up as a single perception. As the rhythm of duration thus successively slows down the existence in tune (with respect to astronomical, periodic events) would become more and more prolonged, and in the limit it would be without end, or immortal. But note that for such an immortal mind all the details of the universe, and its changes, that exist for us would be unknown, inasmuch as they would be synthesised in other perceptions which can have no meaning for us. To summarise, then, the very important results that we have now obtained: If we regard the laws of conservation of energy and augmenta- tion of entropy as of universal validity, we come to an impasse. For the latter law tells us that the changes that occur in the universe tend towards absolute degradation of energy, and there- fore towards cessation of all physical phenomena. But since there can be no limit to past tune, this ultimate degradation ought to have already occurred, and we know that it has not occurred. Therefore the second law is not of universal application. AVe must postulate that it may be reversed, so that, in certain circum- stances, energy that has become unavailable may again become available. In other words, we are compelled to think about the universe that we know as a mechanism that has been started, and is now running down. There must be some means of winding it up. We can imagine no logical reason why unavailable energy should not pass into the available foim; all that we can say is that, in our experience, this transformation does not occur. Then, since absolute reliance upon our experience of the universe brings us to an impasse, we must conclude that this experience cannot be inclusive of all that happens. There must be two directions taken by energy transformations. In one of these energy is continually degraded and entropy becomes augmented. That such changes occur is highly probable so probable that we know by actual observation of no other direction of change. In the other direction available energy becomes restored, and entropy diminishes. That these changes ON THE NATURE OF LIFE 209 occur is highly improbable so improbable that we have no experience of them, though we can imagine them occurring. This means that, in addition to the physics that we know, another and a transcendental physics is conceivable. The Improbability of Life in the Universe. It follows from what has been said that it is extremely improbable that any- where in the entire universe energy is available for the production of physical phenomena. It is easy to show that such is the condition that we know, and that even in OUT individual, galactic universe, where an initial store of energy is exhausting itself, this availability is improbable. If we were able to travel away from the sun in every direction- and with the speed of light, we should reach the boundaries of our solar system in about four hours. Then we should travel outwards into cosmic space for over three years and a quarter before coming to the nearest fixed star. This sphere of space, containing only our solar system (so far as we know), is not, however, empty, but is " full " of ether. The latter is the " substance " of all energetic changes, but we must regard it as inert, or physically inactive. Nothing happens in it of itself. It can be temporarily strained or modified in some way, as when potential energy is locked up in it, or withdrawn from it, or when radiant gravity or gravitational energy is transmitted across it. It is permanently modified in localised regions as physical gravitating matter, which may or may not be the locus of energy transformations. We must think about it as some- thing real, but physically dead or inert. What fraction of this universal substance is, then, physically active ? Calculating the volume of this sphere of three and a half light years in radius, we find that it is about 240 X 10 35 cubic miles. In it practically the only cosmic body that contains available energy is our sun, and this has a diameter of (say) 870,000 miles. Find- ing its volume, we get the value 15X10 16 cubic miles. It is now easy to find that our sun occupies about 1/2 X 10 21 th of the part of the stellar universe that we have in imagination ex- plored. The probability, then, that we should find physical activity anywhere in the region of space within three and a half light years from our sun is something like 1 in 2,000 trillions. Now consider the probability that living matter exists in this physically active fraction of the cosmic space which immediately surrounds'us. We can hardly think of it as existent in our sun, 14 210 THE MECHANISM OF LIFE nor in most of the planets that surround the latter (unless we extend the definition of life in a way that we do not contemplate just yet). Even upon our own earth living substance forms only a surface film of incredible tenuity when compared with the dimensions of the planet. It is probably most dense in shallow seas near the land, but even there the mass of organised matter is only a few parts per million of water. In oceanic regions, both at the surface and in the depths, the density of life is much less, i and on the land (when we take account of polar, mountainous, and desert areas) the density is still less. All that is, of course, only the surface of our planet ; in its depths life is very nearly absent. From the idealistic standpoint life (in strictness, our own life, or mind) is all that is, for the universe and all the things that science places there are only mental constructions. Man, says Pascal, is only a reed, but he is a thinking reed. He is among the least and most fragile of things. But he is also, among the greatest of things ; he is, indeed, greater than all other things, for he can comprehend the universe. Life is like Pascal's reed. Anywhere in the universe it is highly improbable that energy exists in such a form as to givej rise to physical changes. Even in this fraction of the universe,] where there is physical change it is also highly improbable that among these activities there are some that exhibit what we call the phenomena of life. In other words, the chance that life exists anywhere in our universe is an infinitesimal of the second order, The Physical Nature of Life. If we would try to explain life we must, first of all, be very clear about what we mean by anj " explanation." Consider the motions of the bodies that make up our solar system: the planets revolve round a central sun in elliptical orbits, and the satellites revolve round the planets ; thd sun, planets, and satellites rotate on their own axes, and that latter process, or rotate, while still retaining their inclination to the plane of the ecliptic; the ellipticity of each orbit clumps periodically; the various bodies " nutate " ; each of them perturb^ all the others, and so on through a host of motions. To describe all the latter would be very difficult, and the description would be very hard to understand for anyone who is not an astronomer. But assume that each body has a certain mass, that it had aj certain initial velocity, and that it attracts every other body with a force that depends on the various masses and on the squared of the distances between the bodies. Then all the motions can ON THE NATURE OF LIFE 211 be seen to be consequences of the assumptions that we have just made. We start with certain very simple concepts, mass and time and space and the law of gravitation, which is a relation between mass and space and time. Given these concepts and a know- ledge of the velocities of the various bodies, and we can then deduce all their movements, present, past, and future. Every- thing that happens in the motions of the solar system happens because the various bodies have mass and attract each other in a certain way. Therefore we " explain " these very complicated movements by the concepts of mass and gravitation. The latter are simple or irreducible that is, we cannot (so far) explain them by supposing them to be the consequences of something still simpler and more general than mass and gravity. Knowing the positions of the planetary bodies at any moment, as well as their velocities, knowing also their masses and assum- ing that the law of gravitation holds good in all circumstances, we can then find what will be their positions at any future time and what were their positions at any past time. These pre- dictions and retrospects generally are successful, and when they fail (as they do in rare instances) astronomers assume other simple concepts (as in the theory of relativity), and then their calcula- tions work out true to what can be observed. Thus we have a planetary theory, which is the explanation of a host of complex events by a very simple hypothesis that the cosmic bodies have / mass and attract each other. This theory is verified, inasmuch as it enables us successfully to predict forwards or backwards. Now if we had a theory of life we should also have certain very simple, irreducible concepts, and it would be the case that all organic phenomena growth, reproduction, assimilation, ex- cretion, behaviour, adaptation, evolution, and so on would be the inevitable consequences of these fundamental concepts, just as solar and lunar eclipses, tides, seasons, etc., are the inevitable consequences of the ways in which the heavenly bodies move, which are, again, the consequences of the law of gravitation. Knowing these fundamental factors, we should be able to " explain " life. We should be able to say what an animal will do at any future time, and what it did at any past time. We ought to be able to trace out the past history of the species to which it belongs, as well as its future evolutionary history, just because we know what the animal now is and what are the concepts by means of which we " explain " its activities. 212 THE MECHANISM OF LIFE Now we cannot do any of these things. We cannot predict what an individual man or woman will do at any time in the i future, though we can sometimes say what a population will do. (The reader must note this distinction between individual and statistical effects; it is most important in speculative reasoning.) We cannot say, merely from a knowledge of its structure and behaviour, what was the past of the species to which an animal / , belongs, and what is going to be its future. Let there be no misunderstanding here: we do know a great deal about the j phylogenies (or lines of descent) of some races of animals, but that is because these evolutionary careers have left historical records. Thus we know (or at least we believe) that the existing one-toed horse has descended from a three-toed species which had, in its turn, come down from a five-toed form, but this is because we have fossil remains of the three- and five-toed horses, because the living animal has vestiges of the second and fourth toes, while the skeleton of the three-toed horse has vestiges of the first and fifth digits. Evolutionary careers have therefore left records in the form of fossil remains and vestigial structures, and we are sometimes successful in reading these and so in tracing out a line of descent. Obviously our success is not due to any process of true deduction from a few fundamental concepts, as i is the astronomer's when he calculates backwards to find when eclipses occurred. Eclipses leave no records on the faces of sun, moon, and earth like the evolution of the horse has done. And so the astronomer can calculate forward or predict, while the biologist cannot do so, since the successes of the latter depend upon records which cannot exist in the cases of events which have not yet happened. Therefore we cannot predict what will be the future evolutionary history of the horse or of any other race of existing animals or at least no biologist has yet risked his reputation by making such an attempt ! Now we may as well admit that this argument may not appear to possess much force in itself. It may be that biology cannot predict (and so supply the necessary test of the validity of its theories) just because it has not yet attained the necessary knowledge. No doubt Copernicus could not have predicted the times at which equinoctial spring tides would occur at any particular place in the North Sea, for his knowledge of planetary theory was imperfect, and he had not obtained the concept of gravitation. So it may be that biology cannot yet predict just ON THE NATURE OF LIFE 213 because its knowledge is incomplete and its concepts are not yet formulated. This latter conclusion is, however, all that we seek to make here biology, not having attained the concepts that will enable it successfully to predict, has not yet been " explained." Now let the reader refer back to our discussions of Chapter VIII. It is very clear that organic behaviour (in the cases of the higher animals, at all events) is indeterminate: it cannot be predicted. When anyone says that physical determinism must hold in organic as well as in inorganic functioning, he means that it ought to hold because the concepts by means of which we " explain " life ought to be the same as those by which we explain inorganic happening. It may be that they are the same, and that by-and- by biological predictions will be successful; but the plain fact is that, so far as we know, much of the behaviour of the higher animal is spontaneous, and cannot be predicted, while no satis- factory logical proof can be given of determinism in its application to organic acting. We are not going to argue here that, because physiological investigation discloses no activities in the animal body other than physical and chemical ones, organic happening is necessarily the same thing as inorganic happening. It is quite true that the physiologist finds nothing in the functioning of an organ but physical and chemical reactions, for, because of his methods, these are all that he could possibly expect to find. When he studies the submaxillary gland, say, he finds that a saliva, a liquid of a certain chemical composition, is secreted and poured into the mouth. He finds that this secretion occurs when food has entered the mouth, and when the nerve endings there have been stimulated chemically by the substances of the foodstuff. A reflex occurs and the gland functions. The nature and abund- ance of the saliva depend on various factors blood-pressure, osmosis, hydrostatic pressure, chemical reactions in the cells, etc. and investigation has taught us very much as to the ways in which these various factors act. We have dealt in some detail with this typical example of animal activity in the Appendix, and we may leave the reader to study it further in the textbooks. He will see, however, that all that has been obtained by in- vestigation is a description of the manner of functioning of the gland; there are nervous impulses, changes in the calibre of the bloodvessels, changes in the pressure of the circulating blood, changes in the pressure of the salivary liquid contained in the duct 214 THE MECHANISM OF LIFE of the gland, osmotic changes, chemical reactions, and so on. These tilings do not explain the secretion of saliva : they only describe it. An explanation would place beneath blood-pressure, chemical reactions, etc., some simple irreducible concepts, just as in the planetary theory we place beneath the complex motions of planets and satellites (which correspond logically to the molecular move- ments occurring in the tissues of the gland) the law of gravitation. It is, perhaps, not easy for the non-professional reader to appreciate this point. It seems natural to suppose that, because chemical and physical reactions are all that we can study scien- tifically in the living animal body, these things explain life. They only describe life: they are the physical expressions of the activities of the organism. Investigation itself is not primarily speculative, but is rather something useful. Just as the submaxillary gland and all the other parts of the body that enable it to act function so as to produce a liquid that digests certain foodstuffs, so the mental mechanism that we call the categories of the under- standing functions so as to do something that (like the production of a ferment) is useful to the organism. In the higher races of man it has become speculative also, but even there it is pre- dominantly practical, and (as Bergson shows) is therefore hampered when its aims are purely speculative. Let the reader reflect on all this, and he will certainly see that physiology has never explained life activities, but has only described them, and that indeed is the glory of the science, for the description has given us (or will yet give us) the mastery over inimical nature. Let him examine the attitude of the " ordi- nary " man that is (say), some nine-tenths of all those who are capable by prolonged study of following the methods and TOM ills of scientific investigation. This ordinary man will almost certainly ask, What is it for? He will expect that investiga- tion ought to have some useful result, and he will probably be unable to conceal his disappointment (or even contempt) when he learns that the result is only an increase in abstract knowledge ! Life activity therefore expresses itself in physical and chemical phenomena, but physical and chemical reactions do not " ex-j plain " life that is the result to which we seem to be led. Life and Energy. In seeking for the explanation we Lave, therefore, to find some concept which will be special to the' phenomena which we call vital ones, and which need not be required when we investigate and explain inorganic happenings. ON THE NATURE OF LIFE 215 It must have the " property " of reality that is, it must not be in contradiction to the law of conservation for it is to be a means of investigation. Thus we are compelled to reject any of those concepts upon which the theories of spooks are based. It must work that is, it must lead to the discovery of further workable results or hypotheses and it must therefore avoid the reproach (addressed originally by Bacon to certain philosophies), and extended to Driesch's " psychoids " by a well-known cytologist,- that it is like the vestal virgins, dedicated to God and barren ! Now it ought to be clear to the reader at this stage in what direction we are seeking for our concept. We have throughout this book kept the notion of energy in the forefront in order that we might lead up to our thesis that the concept \ which is special to the organism is one which involves the reversal v of the second law of thermo-dynamics. We must not be troubled by the strangeness of such a concept or by the appearance of paradox that goes with it. It is not more strange than some notions in mathematics that have, nevertheless, been fruitful of result than, for instances, the non-Euclidean postulate that more than one line can be drawn through a point and still be parallel to some other line not going through the point ; that the sum of the angles of a triangle may be less than two right angles ; that a negative number may have a square root. The test is that, like the notions just given as instances, ours shall have pragmatic value may work. So, again, we contrast the inorganic and organic physico- chemical systems. In the former, anything at all that happens of itself happens because energy transformations take a certain direction. This direction is that which leads to a decrease of the differences of intensity in different parts of the system. Available energy is concentrated in this part of the system rather than that, and if it can be levelled down, so to speak, something will happen. If it cannot be levelled down, being equally distributed everywhere, nothing will happen. If water is accumulated in a reservoir at a higher level than that in a lake, it can flow down into the latter, turn a mill-wheel as it flows, and so produce various phenomena. If it is all contained in the lake, if there is no difference of level, there can be no flow, and no energy available for the production of physical phenomena. This, then, is the concept by which we " explain " the fact that 216 THE MECHANISM OF LIFE physical phenomena do occur; there are in the universe differences of intensity of energy, and these differences tend, of themselves, to become levelled down. As the process of reduc- tion of intensity difference takes place, energy transformations occur and entropy increases. But, as we have seen, the concept of the second law proves inadequate as an explanation in the universal sense, and we are compelled, even in the treatment of inorganic, cosmic systems, to postulate another concept, that of the second law of thermo-dynamics reversed in sign. Somewhere or some time entropy must decrease instead of increasing. If this reversal were to happen, the results would be unex- pected, fantastic, and paradoxical. But a physicist would not be incredulous. He would, probably, seek to be very sure that lie was not mistaken, and that his observations were trust- / worthy; then, having satisfied himself upon these points, he would accept the result. But, as a matter of fact, the reversal does not occur, and every energy transformation that the physi- cists and chemists can observe and investigate happen because, in this part of the universe known to us, entropy always tends to increase. Therefore the concept which he utilises in order to explain physical happenings is this energy differences tend, of themselves, to become abolished. Now it is quite clear that this concept (which is, of course, only another way of stating the second law) does not fully explain organic happening, for in such processes energy differences do not tend, of themselves, to become abolished. Again, we must be very precise in our statement of this result; we do not mean to say that the animal physico-chemical system, or body, does not " obey " the second law. We recall here the caution sug- gested in the first part of Chapter II., that we must not seek for absolute distinctions anywhere in nature, since these are logical constructions only. There are no such things as mathematical points, straight lines, or planes, for the points and lines and planes which we observe and measure are only approximations. Let a very small spot become smaller and smaller, and " in the limit " it becomes a mathematical point, and so on. Logically, then, we can construct a conceptual world in which there are absolute distinctions between inorganic and organic, and between processes which tend to entropy increase and others which tend to entropy decrease, but wJuit we do actually observe are only the tendencies which, in reasoning, we carry towards their limits. ON THE NATURE OF LIFE 217 In the activities of life as a whole, what we observe, then, is the tendency for the perpetuation of differences of energy intensity. In purely physical happenings energy tends to become unavailable, but organic changes set themselves against this tendency. When sunlight falls upon desert sand, rocks, raw soil, and upon the surface of the sea, its energy of radiation transforms into heat. Sand and stones become warm, and when the sunlight is withdrawn this heat becomes radiated away into outer space, is dissipated, and is, for us at least, for ever irre- coverable. When it falls on the surface of the sea it heats up the water, which then evaporates, rises up into the atmosphere, is distributed in winds, and is precipitated in rain, etc., returning ultimately to the ocean from which it came. In these changes the motions of the winds and water become transformed by friction into waste heat, which, as before, is radiated away and lost. Sunlight, which is energy of high intensity, thus, of itself, becomes degraded or levelled down, entropy increasing in all the transformations that occur. Let, however,the sunlight fall upon green vegetation, and some- thing very different occurs. Its energy transforms into chemical changes, as the result of which water and carbonic acid (sub- stances which are fully degraded and have no free or available energy) become combined together, with increase of available energy, to form carbohydrate. Trace the entropy change, and it will be found that this is positive, and has a certain value when solar radiation transforms wholly into waste, low-temperature heat. Trace it again when sunlight is absorbed by the green plant and transforms into the potential chemical energy of carbohydrate, and it will be found that the increase of entropy is now much less than it was. In the first case the solar energy is for ever lost to this world, but in the second it becomes fixed, or stored up, as the chemical energy of wood, vegetation, oil, or coal. Vegetable life (that is, the predominant form in which life exists on our world), then, has for its tendency the storing up of available chemical energy. The latter becomes locked up, so to speak, in the form of starches, celluloses, proteids, and oils of fruits, seeds, woody tissues, etc. The vegetative processes of reproduction are the most powerful in the animate world, so that on every available place on the surface of the earth, in swamps and in the shallow water of seas and lakes and ponds, plant life spreads and accumulates to the greatest extent possible. Even 218 THE MECHANISM OF LIFE when, by reason of overcrowding and the absence of the neces- sary mineral food substances, this continued multiplication is no longer possible, dead vegetable substance in the form of woody tissues, mould, seeds, leaves, oil, etc., accumulate and form deposits of material of high calorific value. Some of these, in the form of coal, and perhaps oil, remain throughout long periods of geological time in a permanent, utilisable form. The fraction of the solar energy that is absorbed by green plants, and is afterwards dissipated as waste, irrecoverable heat, is very small. For these organisms are, in general, immobile, and so their energy is not transformed into mechanical work, which would become dissipated by friction into waste, low- temperature heat. But in the animal the latter kinds of energy transformations occur to a far greater extent than they do among the plants. The animal is characteristically a machine for the conversion of potential chemical energy into movement of body and limbs, and this movement inevitably leads to friction, and thus transforms into heat which becomes dissipated. Here we are not considering the processes of reproduction ; if we were, we should see that the tendency is, even in the animal kingdom, for the indefinite multiplication of every species, and for the distribution of the individuals over as wide an area of the surface of the earth as possible. All animals, even the most slowly breeding ones, are enormously prolific, and there seems to be no limit to the numbers to which species may theoretically attain. There is, of course, a practical limit which depends upon the quantity of vegetable food substance available, which depends again upon the area of land and sea which can be occupied by green plants, and upon the quantities of the ultimate foodstuffs (chiefly mineral compounds of nitrogen) that are available for the plants. If there were (say, as the results of volcanic activity) a con- tinued supply of these indispensable food substances (nitrates., nitrites, ammonia compounds, carbonic acid gas, and some other mineral salts), there would seem to be no theoretical reason \\ liy the quantity of available energy upon the earth in the form of the organic substance of plants and animals should not steadily accumulate. As it is, some substances which are the results of organic activity do tend to accumulate; these are peat, lignite, coal, perhaps oil, carbonate of lime in the form of coral reefs ;mtein in retarding the natural degradation of cosmic energy may easily be traced even in the lower animals, and it operates in the highest degree in human activities. Even among the lower forms of life we see it, however, in everything that is called an adaptation that is, in every device which gives an animal greater power over inimical nature. The change of coat colour from brown to white in an arctic mammal as the winter approaches is clearly such an attempt. This is the real meaning of adaptation, and one might discuss it at length if it did not constitute the greater part of biological science. Obviously, adaptation attains a maximum in the human animal when the latter develops and perfects the use of tools. We do not distinguish between the bodily weapon and the tool or machine adapted or manufactured from inorganic materials: these are, in their effects, as much adaptations as is the change of coat colour from brown to white, or the conversion of a fold of skin into a flying membrane in some squirrels. The ball- bearings in the hub of a bicycle, or the use of grease in the axle- boxes of a railway waggon, are adaptations ; they would not exist apart from life ; they are the indications of life, and their effect is to retard the dissipation of mechanical energy into waste heat by minimising friction. We cannot think of any inorganic, physico- chemical system tfiafc does this ; the essence of any such system that is, of any mechanism that " goes " of itself, or of any aggregation of chemical substances that interact with each other is that the system tends as rapidly as possible towards a condition of equilibrium in which the mechanism ceases to " go," and the chemical substances cease to interact. Here we get our first concept by which we " explain " physical happening: when water runs downhill; when combustible sub- stances burn and apparently disappear into the air; when coal burns in a fire and generates heat; when proteids and carbo- hydrates decompose into water, carbonic acid, and mineral nitrogenous salts; when a hot poker becomes cold, and so on when these things happen, energy becomes degraded or entropy increases, and it is because entropy increases that they do ON THE NATURE OF LIFE 221 happen. So water does not rraf uphill ; a piece of brick put into the fire does not generate heat; and water and carbonic acid do not, of themselves, combine to form carbohydrate, because in such processes entropy would necessarily increase. Our inor- ganic concept is, therefore, the entropy of the universe tends towards a maximum value. On the other hand, things do happen in the course of which energy is not degraded, but, on the contrary, becomes elevated, so to speak. Water and carbonic acid react together and utilise the degrading energy of sunlight, producing carbohydrate. Proteid, carbohydrate, and fat, when aggregated in the form of living protoplasm, do not decompose, but increase in quantity by reproduction. In these energy transformations entropy decreases, and unavailable energy becomes available.* Some adaptations among wild animals clearly tend to reduce the waste of available energy (as when change of coat colour or the accumu- lation of blubber beneath the skin minimises the loss of heat), and all adaptations have the same general tendency. Human operations may lead to degradation (say by war, by the cutting down of forests, the extinction of seals and whales, or the de- pletion of coal resources), but they may also have the opposite tendency (as in afforestation, the cultivation of the land, the breeding of domestic animals, the increase of populations, and the invention of antifriction machinery). A man may expend his energy in rolling stones downhill, when he dissipates potential energy in useless friction; or he may by the same expenditure of energy carry stones uphill, so that they are in a position to roll down again, thus accumulating potential energy. In such " vital " processes and tendencies available energy accumulates and entropy decreases. Therefore the concept by which we explain truly inorganic happening fails to explain organic happening, for in the latter the inorganic concept is reversed. In living processes the increase o! entropy is retarded this is our " vital " concept. * The reader must not misunderstand this statement. Starch accumu- lates in the green leaf exposed to sunlight, but the whole system is the green leaf + the C0 2 and OH 2 + the degrading sunlight. In the system thus denned entropy increases very slowly. The system is one in which there are coupled energy transformations, (1) the degrading sunlight: and (2) the photo-synthetic process. If there were no coupling, the solar energy would degrade, with a maximum entropy increase; if there is a coupling, the entropy increase becomes minimal. The coupling is always the mark of life activity. APPENDIX I A METAPHYSICAL DISCUSSION THE argument of Chapter X. may be summarised as follows : " Sensation" is a purely physical process, beginning with the stimulation of a sense organ and ending with the physical affec- tion of a nerve centre. In it are involved only molecular dis- placements of the substance of nerve fibres and cells. It is not accompanied by changes in consciousness. " Pure perception " is the prolongation of this physical process of sensation into appropriate actions. Beginning in the animal body with the stimulation of a sensory organ, it ends with a motor action (that is, the co-ordinated movements of a system of muscles), or a glandular change (that is, a process of secretion, or other form of metabolism). It is unaccompanied by any change of consciousness when it is " pure." That is, of course, a limiting case exemplified by typical reflexes, most habitual, visceral responses (respiration, the heart -beat, peristalsis, " mechanical," " habitual," " automatic," and instinctive re- sponses). In such cases consciousness may be absent or very dim. " Perception " simply means a chosen response one that is accompanied by some degree of deliberation, and exhibits some degree of freedom or novelty. We are compelled to postulate this indeterminism of response in admitting the continued evolu- tion of form and functioning, and we are convinced of it if we reflect upon our conduct and the motives that sway us. Such deliberated, chosen, and free response is again the limiting case, because habit; physical, legal, and moral restraints; heredity and convention, determine largely what we shall do in any circumstances. But not altogether. There is some degree of true freedom of acting. The greater the degree of indeterminism that follows upon a sensation, so much greater is the pitch of consciousness. In strong and sustained mental effort there is just this intensely felt deliberation. The latter may end in a choice (when we take some course of action), or it may end in si solution (as when one, works out some physical or mathematical problem) that is, it ends in an actual or virtual action. Then one feels the strong t possible mental satisfaction. A METAPHYSICAL DISCUSSION 223 Or our mental struggle may Ite equivocal, so that there is no choice or solution, or at least none that is satisfactory. Then we are certainly impressed with the sense of failure and, it may be, of acute dissatisfaction. Or, again, there may be persistent sensation that fails to result in appropriate response, and then we experience pain. Between our consciousness that we call mental dissatisfaction resulting from the failure to find a solution and the consciousness that we call "physical" pain there is no clear and essential distinction. But in pain what we describe as " physical " consciousness attains its maximum of intensity, fading away only when, at last, a response (or healing process, or reaction) occurs, or when there is partial or complete destruction of the receptors and neural tracts that are involved in the physical process of sensation. Life Intuition. Clearly there is something in the living animal that is additional to, or, at least, is not merely the physical process of, sensation and response. Subject the latter to analysis, and we find a mechanism, receptor organ, afferent nervous tract, central nervous connections and tracts, efferent nervous- tract, and effector organ. This may be set in motion, so to speak, and evoke nothing but processes that are certainly energetic ones, although the behaviour that we can thus observe may be appro- priate, purposeful, and complete. But it may set in motion again, and then it may evoke response only after deliberation and more or less acute consciousness. Something, then, is operative which is not merely physical sensation and physically actual or virtual response. This may be the exercise of pure memory, the disciplined deliberation that we call reasoning, " mental " and " physical " pain, pleasure, the unsustained and desultory efforts of reverie and " day-dreaming," and so on. It would be as foolish to deny this as to attempt to characterise it all in terms of energy. Just what to call it we do not know. " Spirit " will be a term that is objectionable to many; mind is only one aspect of the thing that we mean; consciousness is inadequate, for we almost seem compelled to postulate " subconscious " mentality; Berg- son's " vital impetus " indicates a passage of some kind or other, and is not just the term we require. In the meantime, at all events, we may refer to it as simply " life intuition," suggesting nothing more than what is meant here in the context. We deal, then, with something that is expressed in the functioning and behaviour of the animal body. 224 THE MECHANISM OF LIFE Intuition and the Law of Conservation. When the animal dies it does not cease to exist, because its molecules fall into new chemical configurations and its energy is conserved even if it is dissipated. But we cannot apply the law of conservation to life intuition, because the law itself is only a mode of operation, or a category, of mind which is the intuition itself. And the application of the law of conservation is necessarily restricted to entities that are measurable, while, obviously, mind and the qualities of mind and " feeling " are not, in themselves, capable of measurement they are not " in " space and time. It is not a perception that we measure in applying the " psycho-physical " law of Weber and Fechner, but the strength of the stimulus that precedes a change of consciousness, while the mathematical relation between the change of consciousness and the energetic change in the stimulus proves to be a spurious one when sub- mitted to analysis. Intuition is, therefore, non-energetic, non- measurable, and is not conserved. All that we know about it is itself it is us. It manifests or expresses itself in animal behaviour and functioning, and so, when the animal body dies or becomes unable to function we must believe that life intuition ceases to exist. Or, at the very least, we have no reason for believing that it can survive the dissolution of the body. Of course, the latter, and therefore the life that it expresses, is immortal in a certain sense. In reproduction a part of the body becomes detached and grows to form a new individual, which again reproduces, and so on indefinitely. Since the ability of expression in functioning and behaviour is thus transmitted through an endless series of generations, something in the life intuition is immortal, but undoubtedly something is abolished in the act of reproduction. This is the intuition of continuous, personal identity, which is lost, or at least does not pass from individual to individual. That which we call memory comes into existence with the individual, and ceases to exist when the latter dies. With memory there is associated, in some way, mdetermmation of response to sensation, and thus the possibility of evolutionary change. And, of course, with this indeterrnina- tion goes what we call motive, praise and blame, personal respon- sibility, and " sin." All these cease with the death of the individual body, some of the activities of which are their ex- pression. If the expression of the whole intuition does not persist, then, can we say that the whole intuition itself is con- served ? When we say that life intuition is, but expresses itself in functioning and behaviour, we make the statement because of A METAPHYSICAL DISCUSSION 225 our own personal intuition and its expression in our activities, and then we assume that activities in other individual bodies similar to ours are also the expressions of individual life intuitions. So, in an individual, human career, we seem to see the gradual cessation of intuition : first, the loss of creative mental power at a relatively early period of life ; then the slackening of the feelings of initiative and responsibility, and a progressive stagnation of in- tellect with increasing domination of habit. Then follow the loss of reproductive power and senile decay. What remains of life becomes concentrated in the effort to give bodily metabolism the peculiar " vital " directions, and little by little even that fails and ceases. What is transmitted in the act of reproduction is the vital impetus; in reproduction a life intuition dissociates, one part remaining as the parental body, and the other part becoming the offspring. All the powers of life that manifest themselves in the functioning and actions characteristic of the parental organism are conveyed, in reproduction, to the offspring. What is not conveyed is the memory of the former and, in general, the new form of acting which it acquires in the course of its individual experience. Just as death ends the individuality and personal continuity of the parental organism, so in giving birth to a new individual that personality dissociates itself. The Working Out of Intuition. The life intuition of one individual is incommunicable, as such, to another. No process of instruction conveys the knowledge of life in itself; what is conveyed is the materialisation of that life intuition. If there is any way in which feeling and thought can be transmitted from one organism to another otherwise than by an action, we do not know of it. What we are told by modern " spiritualism " as to the possibility of " telepathy" and the like does not come within the purview of science, and, at the very best, those results are grossly materialistic ones, and are their own refutation; it is not " spirit " that the " seances " disclose to us, but matter. And the individual and personal life intuition is not trans- mitted from parent to offspring. Heredity is the communication of motor and chemical habit. Form, the idiosyncrasies of acting, moies of functioning these are transmitted. It is the generalised intuition that passes over in reproduction. In the interpretation of the thoughts and intuitions of other people, and in the interpretation of our own thought and intuition, what we have to do with are the physico-chemical expressions of those thoughts and intuitions. Life manifests itself " objec- tively " in no other way. 15 226 THE MECHANISM OF LIFE The appearance of the sky at dawn . may give one a feeling of strong mental satisfaction. Now to describe that appearance in . terms of form, colour, and intensity of illumination, and to explain it by reference to sunlight falling obliquely through an j atmosphere laden with water vapour and dust, may seem to many to be an imperfect and sordid interpretation of a very beautiful phenomenon. But that is all that we can do. If we speak of the " dawn in russet mantle clad," do we express our intuition in a manner that is essentially different ? Obviously not. In either case the materialism is patent. The same kind of mental satisfaction may be experienced when we listen to music. If we describe this in terms of rhythm, pitch, and the relations between sounds heard simultaneously and sounds heard consecutively, we may be told that the descrip- tion is clumsy and inadequate. But the analysts are no more successful. " Thus Fate knocks at the door," said Czerny. "It is the song of the Yellow Hammer," said Beethoven, with reference to the same melodic phrase. Necessarily, the analogy or description, whatever it may be, makes use of materialism. Sometimes one is seized with a feeling of apprehension, and even quite irrational fear, in walking along a lonely road at midnight. The feeling itself is quite indescribable, and it, like aesthetic ones, is private to the individual that experiences it.' But try to describe it: one listens intently; moves as silently as possible; the body is held tensely and in a posture of preparation; heart-beat and respiration respond to the slightest stimuli. Obviously we translate the emotion of fear in materialistic terms. Our feeling of pleasure on looking at a dawn or sunset, or in listening to music, and the dread that may possess one in a strange situation are certainly experiences sui generis. But our intellectual, communicable description of them is, and must be, a materialistic one. " Sunset and evening star " we know to be an atmosphere of a certain constitution penetrated by the radia- tion from cosmic bodies. The C minor symphony, as we hear it, is an exceedingly complex series of displacements of the molecules of the atmosphere. Irrational fear is the emotional, afferent reflexes from a body that is thrown into a state of preparation to resist something material. And so on. Seek to understand and express intellectual, emotional, or other intuitive experience, and we find that we can do so only in terms of matter and energy. Life passes into materialistic phenomena. Or, at least, that is our " first approximation." A METAPHYSICAL DISCUSSION 227 The Description oi Nature. Now we go a step further. Our life intuition, disappearing as such, inserts matter and energy into nature. Or, rather, it does not create matter, so much as pass over into it. That is what we should have said twenty years ago, but can we say it now ? Obviously not. For we do not really observe matter and energy in nature. We observe space-time coincidences, and from these we infer' space and time measurements. It is not matter, nor even energy, that we infer it is only space-time measurements or intervals. That is our " second approximation," but we can go further still, thanks to the modern, generalised theory of relativity. That which we know as " nature " is not matter and energy, nor even space and time, but the relations between space-time coincidences. What is the "weather" ? Intuitively it is a feeling of dis- comfort, but we " know " it and describe it, and act upon it (or are acted upon by it, when the " sign " of the action is changed) in strictly materialistic ways: it is rain and wind and a low temperature and mist. Looking into the matter more closely, we find, for instance, that the low barometer is one of the terms in our description, and the quantity of water in the rain gauge is another. But these are space measurements, for the low baro- meter is a column of mercury (a linear dimension) of a certain magnitude, and the rain-gauge observation is a volume of water (a cubic dimension). The temperature is also a linear dimension the length of a mercury, capillary column. Coupled with these observations there are time ones: we read barometer, rain gauge, and thermometer at a certain instant when the hands of the clock had moved through a certain arc (since the last twelve noon or midnight). Obviously the time is a space measurement. So, also, the material water, nitrogen, and oxygen in the environment are time and space measurements. Whatever they feel to us that is, whatever our intuition of them may be we describe them as molecules which are made up of atoms which we regard as electrons in movement. And of the electrons we can know nothing but space measurements : a positive nucleus with several electrons and a surrounding swarm of negative ones; relative velocities and stresses, repulsions and attractions. The " actual " molecule, or atom, or electron, moves; thus we have, coupled with positions, instants in time. That is, our knowledge of nature is space and time measure- ment, and, indeed, we can see no distinction between the measure- ment of space and that of time. We do not know space and time intellectually, for space is the intuitive knowledge of our freedom 28 THE MECHANISM OF LIFE of mobility, and time is our duration as conscious, remembering individuals. What we know and deal with intellectually are points in space and instants in time. There is nothing between the space points but a mathematical relation and the intuition of a possible actual or virtual bodily movement, and between the time instants there is a similar relation and duration, which is, perhaps, the " stuff " of our intuition. Proceed a little further, and note that in "reading the baro- meter ' ' we observe a space-time coincidence. There is an instant at which the top of the mercury column coincides with a mark on the adjoining scale. That instant itself is a coincidence of the hand of the clock with a mark on the scale or dial. There is, therefore, a double space coincidence. In the barometric reading we define a point y l in reference to another point = / , the scale zero. In the time measurement we find an arc by defining a point x, y in reference to the " zero " of the dial its centre. This latter point we, however, call t, the time. Thus our measure- ments are defined by reference to some co-ordinate system, and their statement is always an arbitrary or conventional one, and depends on the choice of the scale zeros or co-ordinate systems. Proceed with such an analysis in relation to any transforma- tion, or event, or object, or phenomenon whatever. The " elements " of our knowledge the perceptions with which we construct the universe are space-time coincidences. In any such perception we generally observe the coincidences of four points (three space points and one time instant), x l7 y^ z ]7 ^, with other four, #,, y 2 , z 2 , t. z . Let us state this in a quite dogmatic way (for it can be amply demonstrated, if necessary) all the data of our knowledge of nature are space-time coincidences and the relations of such. Nature a System of Relations. We do not deal with even the space and time points, for these have no meaning except with relation to a " frame of reference," or co-ordinate system. The latter is always an arbitrary (though convenient) one, and obvi- ously the position of any point depends on the zero from which it is measured. We take as an illustration (for a clear under- standing of the matter is very desirable) the trajectory of a material body falling freely in vacuo. We do not deal with " spaces " and " times " here, but with " differentials," ds and dt. The symbol ds is not an infinitesi- mally small space, but the limit to a space that becomes smaller and smaller without ever becoming zero. It can be so small that the error involved in regarding it as zero will be less than any A METAPHYSICAL DISCUSSION 229 standard error, no matter how^ small this is taken to be. So also with the symbol dt : it is not an infinitesimally small dura- tion, but the limit to a duration which always diminishes. Our falling body, therefore, is at certain points s , s^ s 2 > etc. when the time is ? , ^, Z 2 ? etc. The s-points coincide with the ^-points, the former being read from a measuring rod, and the latter from a clock. Representing these coincidences graphically, we get the following " one-to-one correspondence " of s-point with -point. Obviously we might remove the origins, or zeros, of the time and space scales further to the left without altering the character of the diagram: t = 1 second might just as well be t = 10, and s 100 feet might as obviously be t = 1,000 the inclinations of the dotted lines showing the coincidences would not be changed. >OO SOO -4-OO ^OO 600 ' -7OO / 77 Q. //near space cL ' m e " -S" / on FIG. 51. THE UPPER POINTS REPRESENT OBSERVED TIMES, THE LOWER ONES THE CORRESPONDING SPACE. What, then, we obtain from such an analysis is the relation between s-points (which have no magnitudes) and ^-points (which have no duration). In other words, it is a naked relation that we obtain, and nothing else there are no space or time regarded as entities. Space and time are not entities, as we have seen already. The relation is the ratio between the differentials ds and dt\ thus - ds and this is what we find when we observe the fall of a stone in wcuo. Such differential equations are the " la^s of nature." As a rule, however, we try to avoid using the space-time equa- tions in their differential forms, and we express them, when possible, otherwise. Thus the above relation is more easily recognisable in its integral shape We have now reached the conclusion to which we have been tending. Nature, as we know it intellectually that is, the Nature of Science is a system of relations, and nothing more. It is, however, our practice to speak of these more fundamental rela- tions as " matter," " fields of force," " energy," " atoms," " electrons," " radiation," and so on, all these things " occurring " 230 THE MECHANISM OF LIFE in space and time. It would be highly inconvenient in the practical affairs of life to do otherwise, and it would be clumsy even in scientific investigations. Just now, however, we are not concerned with practice, but with the attempt to search into the meaning of things. These meanings we can grasp in a fumbling kind of way. We postulate relata between which we make differential equations or relations. But the relata are only space-time coincidences, ds's and dt's, and these have no " reality " apart from co-ordinate frames to which we refer them, and the choice of such co-ordinate frames is quite arbitrary. Something, however, is an absolute element of our knowledge, and this is the relation itself which exists, and is invariant even if the frames of reference vary. Obviously, then, " our knowledge of nature is a knowledge of form and not of content,"* and what that content is we do not know. We are assuming (though the assumption cannot be proved " scientifically ") that there is a content of nature some- thing apart and independent of the intuition of life. The latter works itself out in a description of the manner in which it acts upon the content, but it does not describe the content. That is the ordinary, " natural " way in which we think about things here is life, in us as well as in a multitude of other animals, and there is an " environment " upon which life acts. That assumption is contained in everything that an animal does, and it is implicit in all our civilisations, and so one cannot but make it a part of our philosophy. We know imperfectly how we act upon nature (the environment), but we do not know what it is that we act upon. The elements of our scientific knowledge are the forms of the actions, but not the stuff acted upon. The " Passage of Nature."t Assuming, then, this nature that exists independently of our form and power of action upon it, we note that it " passes " that it is a progress or career. It has a tendency in a particular direction, that tendency being described by the second law of energetics. This law we must regard as the fundamental thing in our experience, the concept that is un- shaken by any recent development of scientific theory and specu- lation. And, of course, its fundamental character suggests that the law is in us in our mode of acting upon nature. First, we note the phenomena of radio-activity. Certain ex- ceptional substances the atoms of uranium, radium, actinium, * Eddington, Space, Time and Gravitation, Cambridge University Press, 1920, p. 200. t A. S. Whitehead, The Concept of Nature, Clarendon Press, 1920. A METAPHYSICAL DISCUSSION 231 thorium, polonium, etc. disintegrate, giving off large quantities of radiant energy. We know that this energy is not created in the atom, but is a store that diminishes by reason of its emission. Therefore, the radio-activity of a disintegrating atom comes to an end sooner or later. Uranium passes into radium, and radium, after a long series of intermediate steps, passes into lead. The atomic energy of lead is bound, since this substance shows no sign of radio-activity. Now, the great majority of the chemical atoms are in the same condition as lead, and we know that the few that are still capable of emitting radiant energy will pass into the condition of lead, when their energy will exist entirely in the bound state. Extra- polating the process far into the future, we predict the time when all chemical atoms in the earth, sun, and the luminous stars, will have ceased to emit energy that is, the substance of the cosmic bodies will have passed into the inert material stage. So, also, extrapolating the process far back into the past, we may be sure that we pass through stages in which the rate of emission of energy from radio-active elements was progressively greater and greater, and was at a maximum at some remote stage in the evolution of our universe. In that stage the substance of the latter existed in the pre-material state. With regard, then, to this generation of available energy by radio-activity, the passage of nature is one from a pre-material to an inert material stage ; from the condition of free energy to that of bound energy. This, doubtless, accounts for the greater fraction, or perhaps all, of the energy that takes part in universal transformations, or physical phenomena. But consider further the free or available energy emitted by radio-active bodies, or free energy in general, apart from any question as to its origin. This energy degrades. Whatever its nature, and whatever the nature of the trans- formations which it undergoes, it ultimately becomes heat at low temperature. Further, this heat tends always to become equally distributed everywhere throughout the universe, so that the time will come when there will no longer be any difference of intensity, or potential, in the unique form (heat) in which energy will then exist. That will be the condition of universal physical death (" Warm- Todt," in Clausius' term). The universe, therefore, tends from a condition of physical activity to one of physical death. It runs down. Free energy becomes completely dissipated, and unbound energy becomes bound. From a condition of minimum entropy the universe 232 THE MECHANISM OF LIFE attains a condition of maximum entropy. From a pre-material state it passes into an inert material state. This is the tendency, or direction, of the passage of nature. The Relativity of the Passage. Now one thinks about the life passage in just the same way. If we are right in our inter- pretation of life intuition as something indescribable (though perfectly well "known" to us) which "runs down" in some way, expressing itself in inert material manifestations, then the life passage exhibits the same tendency, or direction, as does the natural passage.* In Bergson's terminology it defends. Note that all recent work interprets " matter," just as Descartes did, as extension, and note that the quality of life, mind, memory, intuition (call it what one likes), is that space measurements cannot be made with regard to it. There is, therefore, some quality in life that we express as intensitiveness, meaning the opposite of extension. By detending this quality becomes something extensive, and so capable of space measurements, and therefore of physical investigation. That which Bergson puts in this way we have tried to put in another way, but the idea involved is the same one. Life, then, passes always into the inert material state, and the passage is our intellectual description of it. Assume, now (if for no other reason than to see where the assumption leads), that there is a stuff of nature, an environ- ment of life, something other than life which exists as well as life. Assume that our superficial investigation of nature tells us that this environment does not remain the same, but passes into an inert material state. Life, acting on this stuff of nature, retards the progress of the latter towards the inert material state (when life could no longer utilise it). It retards the augmentation of entropy, but does not (cannot, in fact) prevent the ultimate attainment of maximum entropy. As the temperature of the sun falls life must (as we know it) become the less and less able to persist, and must ultimately cease. That is our ordinary conception of the relation of life to its environment. But it seems (since one has been compelled to think in another way by the stimulus of the modern relativity theory) that we may just as easily regard nature as something that is at rest, and which does not pass, and hold that the apparent passage is due to the passage of life. If the atmosphere is at rest, and if one * Remember the (easily ignored) fact, that of living substance we literally know nothing. We study the behaviour only of a living organism. When- ever we study organic substance, it is necessarily dead, inert material that \vc investigate. A METAPHYSICAL DISCUSSION 233 moves quickly through it, one sets up an apparent current of air, and this current may apparently blow in either of two opposite directions, according to the way one moves. It would not be difficult to devise conditions in which it would be impossible for a man to ascertain whether he was at rest and the air were blowing past him, or whether he was being carried through air which was at rest. So, instead of the environing nature changing as we have suggested, it may be that it is " at rest " ; that it is all there, so to speak, though locally different; and that life changes, or moves, so that it encounters the local modifications, just as a man in a railway carriage is carried through a "changing" landscape which is " really " at rest. There is an apparently formidable difficulty to our thinking anything of the kind : we seem to be convinced that nature passes whether we are there or not. We believe that the sun shone and its energy became degraded before we were born, and that it will shine and dissipate its heat after we are dead. Certainly these processes occur while we are asleep and uncon- scious (though we are still there in those conditions, it should be noted). It is worth considering whether we are not there also before we were born, and will be there after we are dead; and to those that accept the doctrine of personal immortality the apparent objec- tion we have suggested ought not, of course, to exist. Now, we do suggest that we were there, and shall be there, before and after individual life; indeed, that is really the case. We (that is, our life) actually and really existed, in the strictest scientific sense, before our individual lifetime as a fragment of germ plasm in the parental body, and in the grand-parental body before that, and so on indefinitely. And, potentially at least, all the future generations of life are contained in us in our actual, physical bodies. Therefore, life is continuous, unitary, and always there, just as the hypothetical environment is there simultaneously that is the plainest and most easily grasped result of biological science. We ignore it just because we are obsessed by the notion of individual forms, individual bodies, species, genera, families, and so on. But the forms are surely continuous, flowing into and out from each other in an evolutionary process. And we are also obsessed by the notion of our personal con- tinuous memories, which are most certainly discontinuous careers, begininng with the awakening of the " categories of the 234 THE MECHANISM OF LIFE understanding " by experience, and ending with the death of the matured " somatoplasm " (not necessarily the "body," note, for the latter was both somatoplasm and germ plasm). This remembered experience, with the indeterminism that is asso- ciated with it, and the morality, immorality, virtue, and " sin " that it is also associated with in virtue of its indeterminism, is discontinuous, comes into existence and ceases to exist. We have no reason to say that it is conserved, and the very notion of conservation is inapplicable to it. Yet that personality is only a little of life, and the greater fraction of the latter is continuous, and all the life of the world is one. That is just what one means by reproduction and heredity. What one means by evolution (or transformism) is that life changes, that it undergoes passage. There is, of course, the other difficulty, felt by those who profess " solipsism " : there is, literally, nothing in nature but the thinking mind, or rather the thought. That, we hold, is negatived by every life action. It is absurd to ' ' common sense ' ' (and surely no philosophy can disregard that !). It is a pretence in anyone who merely says or writes that he believes it (for in speaking and writing he assumes that there are other minds that listen to his words or read his writings), and we even suggest that there is a kind of intellectual dishonesty in talking about it. So the objection fails, for life is always and everywhere there (for we cannot say that thought is in space; a man is in a room, but is his mind there ? Obviously not when he thinks of the seaside holiday or of his boyhood). Therefore, life being always and everywhere, we are free to suggest that its passage and that of environing nature are relative to each other, and that we cannot say " which is which." But, again, it seems quite possible to hold that this question of the relativity of the life passage and the environmental nature passage is one that has no meaning. We do not know what is the stuff of nature, since all that we discover by investigation of any kind is a system of naked relations. There may be an homo- geneous stuff (as Bergson says), and it may be that the foim of this is just what our possibilities of action make it. Atoms, and electrons, and energy, are just the ways, it may be, in which our life intuition cuts up this homogeneous stuff. The latter may be the actual stuff of our consciousness, so that there is, then, nothing in the universe but this: the stuff of life which continually SPACE IN MODERN THEORY OF RELATIVITY 235 appears to itself to degrade itself into inert-material that is, into nothing, for the qualities of inertia are negative ones. Here it is obvious that our discussion is becoming very frankly metaphysical , and must stop. APPENDIX II SPACE IN THE MODERN THEORY OF RELATIVITY SOME of the leading ideas in the theory of relativity are not at all difficult to grasp, and they are of extreme importance. First, then, we note that the Euclidean system of geometry is insufficient for speculation upon the nature of the universe. It is based largely on the parallel postulate that no more than one straight line can be drawn through a point that lies outside a given straight line and still be parallel to that given straight line. Euclid assumed this, but could not prove it, and no one since him has been more successful. We can assume the contrary that more than one straight line can be drawn parallel to a given straight line. Then we can prove a number of propositions that seem to be absurd, but are really quite consistent and free from contradiction. The non-Euclidean geometries describe the ordinary things that we see quite satisfactorily, but they are not so easily worked as the classic system, so we use the latter in our everyday affairs. Second. The Euclidean geometry of three dimensions does not describe events. A thing is not simply there, so to speak it always happens. A molecule of water is not really a thing it is things in motion (electrons). Therefore, we need the idea of time in describing nature. We say that a thing is somewhere it is so much distant from the plane of X, so much from the plane of Y, and so much from the plane of Z. Its space description involves three variable numbers X, Y, and Z but since its description also requires a statement of the time at which it happens, we want a fourth variable, T. (We want the when as well as the where.) Thus our universe must, at least, be four-dimensional. Things are really and actually "specified" by four variables: X, Y, Z, T. This is the space-time continuum of Minckowski, adopted by Einstein. Third. The four-dimensional (but still Euclidean) geometry is insufficient. The latter is to be thought about as a three- dimensional one of three co-ordinate planes^, Y, Z which are 23(5 THE MECHANISM OF LIFE continued somehow in a straight line, T. Now this does not describe the universe when we take account of relative positions and motions. Somehow or other the co-ordinates must be curved, and to allow for this we have to introduce a fifth dimen- sion. This is " quite all right," and the mathematics of a five- dimensional geometry are as straightforward (though immensely more difficult) as are those of three dimensions. The trouble is that we cannot visualise five dimensions. Still, the mathe- matical results are there a fact of very great significance, for an abstract result of this kind always means possible action. Given any abstruse mathematical result, and we may be pretty certain that by-and-by it will mean something " real " that is,, something that practical science, and by-and-by even industry > will do. SD we live in a non-Euclidean universe of four dimensions which is "in" a continuance of a fifth dimension. Leave it at that and see what are the consequences. The consequences (for our speculations of Chapter XL) are that space cosmic space, that is is " finite but unbounded." Think about this by analogy: the surface of the earth to us is finite,, but unbounded. One can go on travelling over it in " straight "" lines (geodesies) without being brought up anywhere ; there would be no end to our journeying if we could live for ever. We might come back to the same place from which we started, though we should be travelling in a " straight " line, but even then we could still go on in the same direction. Now extend that to cosmic space, and suppose that it, too, is- finite and unbounded. We are stepping out of non- Euclidean,, two-dimensional, curved space into non-Euclidean, four-dimen- sional, curved space. This means that the universe is finite but unbounded. We can go on through it with the velocity of light for ever, and always in the same direction. There is no end ta our journey. Perhaps we might come back to the starting-place, perhaps not; if we did, we should still be facing the same way. It means that the galactic universe is finite (and Einstein even tells us how big it is). Outside it there is nothing, not even space. Light travels in what we call curved lines, and never goes outside the universe from which it originated. The notion is of immense speculative importance. Suppose, though, that the time dimension is also curved t Einstein does not seem to have worked out that. Perhaps the reader may see to what extraordinary speculations this may lead. THE SUBMAXILLARY GLAND 237 APPENDIX III THE SUBMAXILLARY GLAND AN INSTANCE OF ORGANIC FUNCTIONING THREE pairs of glands open into the mouths of mammalian animals: the sublingual, submaxillary, and parotid salivary glands. Their function is to secrete saliva. Saliva consists mostly of water which contains some mineral salts, some soluble proteids, some mucin, and, in some animals, an enzyme called ptyalin, which has the power of dissolving starch and converting the latter into sugar. Saliva mixes with the food, so that the latter can be worked up into a " bolus," which is then swallowed; it cleans the mouth in that it removes dirt in the " spittle," and it may be a digestive ferment, or enzyme. FIG. 52. A DIAGRAM OF A SALIVARY GLAND WITH ITS VESSELS, NERVES, AND DUCT. The gland itself consists of a very great number of alveoli which are connected with ductules, which unite to form the duct which opens into the mouth. The saliva is formed, or secreted, in the cells which constitute the walls of the alveoli. Leaving the details for a moment, note the structures connected with the gland. An artery conveys blood to it. The latter circulates round the alveoli, and leaves the gland by means of a vein. Lymphatic vessels are also connected, and through these a watery fluid, called lymph, leaves the gland and finally enters the circulating blood of the body. A ducb leaves the gland, and through this the saliva that is secreted in the alveoli reaches the mouth. 238 THE MECHANISM OF LIFE Two nerves enter the gland. One of these comes from the medulla, and is called the chorda tympani. The other comes from the sympathetic nervous system. These are efferent nerves conveying stimuli to the gland from the central nervous system. The duct' divides into (or, rather, is formed by the union of) a great number of smaller ductules or tubes. The ends of these tubes may be thought about as swelling out to form bulbar enlargements. The walls of the ductules are thin, but those of the enlargements, or alveoli, are thick, being made up of large cells. It is in these cells that the actual formation of saliva occurs. The artery which enters the gland divides up into a great number of smaller vessels, or arterioles, and the latter again FIG. 53. A, Diagrammatic section through an alveolus (or secreting unit) of the gland, with its capillary circulation; B, diagram of the terminations of a secretory nerve among the cells of an alveolus. break up into capillaries. The capillaries are distributed over the alveoli, as shown in Fig. 53. After circulating through the capillaries, the blood leaves the gland by means of the vein. The nerves similarly break up into smaller branches. Some of these divide further into very fine fibrils, which are distributed as a fine network over the alveoli on the outside, or even in between, the cells. Other branches of both nerves break up into fibres which end in the muscles of the walls of the artery and arterioles. Such is the structure. How does it function ? Generally saliva flows into the mouth when savoury food substances stimulate the gland to act. This is a reflex, and the afferent nerves are the gustatory and olfactory ones that is, those conveying the taste and smell stimuli to the brain. But this is not all by any means, for if meat be shown to a hungry THE SUBMAXILLARY GLAND 239 dog there will be a reflex flow of saliva from the mouth, even though the animal does not s-mdl the food. The afferent paths are now the optic nerves. But, again, this is not all, for " when the dog realises that he is being played with . . . the psychical secretion of saliva ceases." And yet again, it is said that the thought, or memory, of savoury food may cause a secretion of saliva in ourselves. When small pebbles are put into a dog's mouth, the animal shifts them about with his tongue and then spits them out. When the same substance, crushed into sand, are put in his mouth, thin watery saliva is secreted, and the sand is washed out. So, also, with dry biscuits. When meat is put in his mouth, thick, viscid saliva is secreted. Thus, although the secretion of saliva is a reflex, it is one that is controlled and modified (or even arrested) in a variety of ways. There is no simple, invariable, mechanical response. How is the control effected ? It was thought at one time that the control was effected by the change of calibre of the arterioles. The saliva comes ulti- mately from the blood that circulates through the gland. Now the fibres of the chorda tympani dilate the vessels and permit a more abundant flow of blood through the gland. Accordingly, there is a greater flow of saliva. The fibres of the sympathetic contract the arterioles, and so a decreased flow of blood takes place and there is a decreased flow of saliva. The inference appeared to be that the secretion of saliva was a mere filtration from the blood, and depended on the pressure of the latter the greater the pressure, the greater the flow. But the saliva has not the same composition as the liquid part of the blood, so there is more than mere filtration to be accounted for. Further, when the gland is poisoned with atropin, the dilating action of the chorda tympani is not affected; yet when the latter nerve is stimulated in such a poisoned gland, there is no increased flow of saliva. On the other hand, when it is poisoned with pilocarpin there is an increased flow. The explanation of these experiments is that there are secre- tory nerve fibres in addition to those that dilate or contract the arterioles. Poisoning these secretory fibres in one way or another affects the secretion of saliva. We see that there are nerve fibres coming into connection with the cells of the alveoli. It is, then, these secretory nerve fibres, acting directly on the cells, that stimulate the latter to secrete the liquid. The other nerve fibres which act on the bloodvessels control the supply of 240 THE MECHANISM OF LIFE blood to the cells as the circumstances require. How do the secretory fibres act on the cells ? When the latter function, it can actually be seen that granules of some material taken from the blood have been formed and deposited in their substance. The stimuli of the secretory fibres seem to cause these granules to disintegrate, absorb water, and swell, finally discharging their contents into the cavity of the alveolus. This is the thing, then, that has to be " explained." What is the substance of the granules ? How is it taken from the blood ? How is it liberated into the alveolus when the cell is stimulated ? . How is it that the nature of the substance varies according to the kind of food ? How can the sight of food produce the same effect as the taste or smell of food ? How can the memory of food do the same thing as the taste of food that is, how does an immaterial, non-energetic stimulus produce the same response as an energetic, material one ? It will be seen that what we have attained so far is a rather imperfect description of the mechanism of control of the secretion of saliva. So far as the act of secretion itself is concerned, physiology has succeeded in substituting a biochemical descrip- tion for a mechanical one, but even the biochemical description involves at the present time hypotheses that have still to be verified. INDEX ABSORPTION, 61, 62 Acceleration, 35 Acquirements of behaviour, 151 Acting, functional, 180; indetermin- ism of, 180; unconscious, 178 Action, creative, 181; difficult, 178; forms of, 178; habitual, 13, 180; and psychical processes, 178; studied objectively, ISO Activation of enzymes, 58 Activity, individualistic, 8; spon- taneous, 9, 182 Adaptations, 149; concealing, 150; functional, 150; and intelligence, 151 Aggression, organs of, 3, 11 Air, fixed, 157; sacs in birds, 2; vesicles in lungs, diagram, 67 Algae, 9 Alimentary canal, 163 Altruism, 142 Amino-acids, 59, 62 Ammonia in nature, 79 Analysis of activities, 12, 193; of sensation, 174 Anatomy, 20; comparative, 2; and function, 125 Animal automata, 13, 153, 178, 192; behaviour, 192; combustion, 56; definition of, 4; and energy, 156; feral, 142; saprozoic, 84; sedentary, 9; spirits, 155; structure, 1; and vegetable, 85; warm-blooded, 1 Animate engine, 81; mechanism, 69 Anthropoids, brain of, 127 Antiseptics, 78 Appendages and locomotion, 12 Aqueduct of Sylvius, 97 Arteries, muscles of, 22 Articulations in forelimb, 18; in skeleton, 14 Asphyxiation, 6 Assimilation, 57, 63 Association paths, diagram, 103 Atmosphere, composition of, 82 Atomic weight, 36 Atoms, 33, 36; indivisibility of, 36; rearrangements of, 167 Auditory organ, 108; ossicles, 12; sensation, 114; sense and equili- brium, 114; tracts in brain, 114 Autolysis of flesh, 78 Automata, artificial, 153; decerebrate, 141 ; Organic, 13, 153, 178, 192 Available energy, 38 ; decrease of, 197; in universe, 209; waste of, 197 Axons of nerves, diagram, 26 Bacteria, action of, 219; fermentative, 78; nitrifying, 79; oceanic, 80; pathogenic, 77; putrefactive, 78 Barnacles, taxis in, 135 Becoming, principle of, 49 Behaviour, animal, 9, 12, 149; grading of, 187; measurement of, 171; pre- diction of, 172 Bell and nervous impulses, 164 Bergson and duration, 206; .and in- telligence, 214; and matter, 185; and spirit, 185 Biology, the modern impasse, 193; utilitarian, 160 Black and combustion, 151 Blood, arterial and venous, 67; circu- lation of, 63; corpuscles, diagrams, 68; gases of, 68; plasma of, 69; venous, 70 Blood flow, regulation of, S8 Blood, vascular svstem, diagram, 63, 64 Bloodvessels, 61 ; calibre of, 22 Bodily action, scheme of, 130 Boltzmann and entropy, 202 Bones, fixed and movable, 22 Brain, 94; alternative paths in, 183; development diagrams, 95 ; diagrams of, 98, 100; earthworm, 92; evolu- tionary stages in, 105; fish, 96; general scheme of, 89; lower, 103, 139, 146; of mammal, 96; vesicles of, 94; working of, 148 Burrowing animals, 11 Calorie, 37; definition of, 40, 44 Calorific values, 158 Calorimeter, 40, 44 Cancer, 7 Capacity factors of energy, 50 Capacity for doing work, 37, 33, 42 Capillaries, 61 Carbohydrates, 58, 60; decomposition of, 79 241 16 242 THE MECHANISM OF LIFE Carbonic acid in lungs, 67; in nature, 84 Carnivorous animals, 84 Carnot, 194; and the steam engine, 157 Catalysis, 158 Categories of locomotion, 11 Categories of the mind, 54, 185, 188; biological view of, 191 ; are mental constraints, 189; mental operators, 188; and virtual action, 191 Cause and effect, 179 Cerebellum, 95, 123; connections of, diagram, 124; connections of, 102; importance of, 125; injuries to, 124; and standard movements, 148; structure of, diagram, 26 Cerebral hemispheres, 94; functioning of, 125 Cerebral physiology, 133 Chemical combustion, 48 Chemical configurations, 34 Chemical energy, 48 Chemical reactions, 51 Chemical reactions, types of, 167; endothermic, 83; and energy trans- formations, 167 Chemistry of eighteenth century, 157 Chemistry and modern biology, 193 Chlorophyll, 82 Chyle, 154 Circulation in body, 152; Circulation of the blood, diagram, 65; discovery of, 66; in fish, 64; Civilisations,industrial and pastoral, 86 Clausius, 231 Claw- like organs, 8 Coal and plant life, 86 Cochlea, 115 Coherence, 31 Cold, sensation of, 109 Colloids, 158 Colour change in animals, 156; nature of, 33 Combustion in animal bodj r , 44, 56; nature of, 39, 40; theory of, 157 Commissures in cortex, 128 ; in nervous system, 92 Compensatory transformations, 83 Compounds, chemical, 34 Consciousness and colloids, 184; and cortex cerebri, 110; and energy, 133, 184; not measurable, 184; and sensation, 110, 176 Conservation, law of, 46; and action, 54; a convention, 179, 194: a law of thought, 46, 53; qualifications of, 52 Consumption by animals, 85 Convolutions of brain, 103, 128 Co-ordination of activities, 87, 182; and cortex cerebri, 129, 140; mus- cular, 123 Cord, spinal, tracts in, 11 1 Corpora striata, 95; quadrigemina, 95 Corpuscles of blood, diagram, 68 Correspondences, one to one, 229 Cortex cerebri, 143; and cerebellum, 129; connections of, 102, 127; and control of body, 120; diagram of, 128; in evolution, 127, 144; and learned actions, 149; localisa- tion of function in, 127; motor area of, 128; prefrontal area, 127; sen- sory area of, 129; and spontaneous actions, 149; structure, diagram, 126; and volition, 105 Corti, organ of, 108, 115 Cosmic order and behaviour, 150 Cosmic space and matter, 209; finite, 236 Cranial nerves, 89, 113 Cranium, the vertebrate, 16 Crura of brain, 128 Crustacea, locomotion in, 11 Crystal growth, 171 Crystalloids, 158 Death, meaning of, 225 Death, universal, 231 Decerebrate bird, 140; dog, 141; fish, 140; frog, 139; mammal, 141 Decussation of nervous tracts, 100 Defensive organs, 3, 11 Degeneration of nerve fibres, 165 Dendrites of nerves, diagram, 26 Density and temperature, 31 Descartes and animal heat, 155; cos- mogony of, 161; and Harvey, 66; and mechanism. 153; and modern vitalism, 193; and physical specula- tions, 161 ; physiology of , 154 Detension, 232 Determinism, physical, 134, 172; and free-will, 179; in organic re- sponse, 173; in spinal frog, 136 Dietaries, energy of, 76 Differentials, 228 Differentiation of structure, 2 Digestion, 57, 58, 61 Digestive juices, 58 Dimensions of space, 189; of time and space, 235 Dinosaurs, skeleton of, 15 Discourse on method, 160 Disease, biology of, 6; epidemic, 77 Disentropic phases, 203 Disharmonies of functioning, 6, 7, 74 Dissipation, law of, 97,217; life retards, 221; in nervous pystem, 133 Dog, scratch reflex of, 119 Dreams. 171> Driesch and logical categories, ISO Ductless glands, 99 Duration, ni'-aiiingof, 206: not infinite, 197; orders i.f. :.'<>.">: and perception, 207; rhythms of, 206; and time, 207 INDEX 243 Dynamics, 46 Dyspepsia and secretion, 7 Ear, internal, 103 Effort, bodily, 35 Einstein, 194, 235 Electricity and heat, 44; measure- ment of, 37 Electrons, 37; and modern cos- mogony, 162 Elements, Cartesian, 161 ; chemical, 36 Embryo brain, diagram, 96 Emotion, absent in decerebrate ani- mals, 141 ; nature of, 25 Endothermic reactions, 83 Enererj', abstract, 45; available, 38; bodily sources of , 69 ; bound, 53 , 23 1 ; ' conservation of, 46; definition of, 46, 52; dissipation of, 51, 86, 231; electric, 38; equations, 45; factors of, 50; of food, 69; forms of, 38; gravitational, 38; history of, 157; kinetic, 39; leakage of, 45; loss by friction, 41; muscular, 38; radiant, 40; of space, 209; of state, 48; of steam, 39; unavailable, 42 Energy transformations, 39, 41, 42; alternative, 108; and becoming, 49; compensatory, 83; conditions of, 215; coupled, 221; examples of, 49; irreversible, 42; quantitative, 44; release of, 30; reversible, 42; sign of, 51 ; and work, 51 Engine, animate, 56; inanimate, 55 Entelechy, 193 Enterokinase, 58 Entropy, 52, 196; and becoming, 216, 220; changes, 200, 202; increase of, 221 ; retarded by life, 220 P^nvironment, action on, 55; of life, 55: stimuli of, 13 Enzymes, action of, 62; nature of, 58 Epiphenomena, 184 Equations, differential, 168, 171, 229 Equilibrium and auditory sensation, 114; mechanical, 40 Erepsin, 61 Ether of space, 162; seat of potential energy, 48 Evolution and adaptations, 186; cos- mic, 181 ; creative process, 181 ; historic proofs, 212; and structure, 2; unpredictable, 211 Excretion, 70; vicarious, 6 Exercise, physiology of, 87 Existence, real and unreal, 52 Exoskeletons, 15 Experience and acting, 182; and adaptations, 151 ; in reflexes, 118 Explanations, scientific, 210 Extension, 232 Extensor muscles, 21 .Eye a camera, 108; cyclopean, 99; of invertebrates, 2 ; of vertebrates, 2 Eyelids, movements of, 23 Fabricius, 66 Fats, 58 Fats, decomposition of, 79 Fatty acids, 60 Fear, analysis of, 226 Ferments, digestive, 58 Fibres, muscular, 23 Fischer, 59, 159 ! Fish, bloodvessels of, 64; brain, diagram, 97; cerebellum of, 95; locomotion of, 11 i Flagellates, 11 ! Flexor muscles, 21 i Food, composition of, 58; distribution of, 61; and energy, 158; inedible parts of, 57; of plants, 81 ; prepara- tion of, 57 ; substances, 57 Foodstuffs, categories of, 58 i Food substances, ultimate, 218 | Foot, movements of, 20 i Forearm, diagram, 21 ; movements of, 18,21; muscles of, 21 l Forebrain, 94 1 Forelimb, diagram, 19; of mammals, 18 i Foster and vitalism, 160 j Freedom, degrees of, 18, 19 Free-will, 179 Friction and heat, 43 Frog, decerebrate, 139 Functioning, organic, 213 ; Galen, physiology of, 154 ! Ganglia, *in brain, diagram, 101; medullary, 100; nervous, 91; seg- mental, 91, 92; structure of cells of, 26 i Gases, heat transference in , 200 ; kinetic theory of, 198; in lungs, 67 Gastric glands, 58 Genetics, 8 : Geometry, 235; Euclidean, 90; four- dimensional, 190; non-Euclidean, 215,235 Gibbs, 194 Gills of fish, 64 Gland action, 70, 74; control, 239; ductless, 74; nature of, 58; pineal, 74; pituitary, 74; structure of, 238; thyroid, 74 Goitre, exophthalmic, 7 Golz, 141 Graham and dialysis, 158 Gravitation energy, 35; an explana- tion, 211 Green plants, 87 Grey matter of brain, 91, 128 ; structure of, 126 244 THE MECHANISM OF LIFE Growth, directive. 135; movements, 9; organic, 171; orientation of, 9 Guano, origin of, 80 Habits, 179; variable, 173 Haemoglobin of blood, 69 Hallucinations, 179; and energy, 52 Hand, articulation of, 18 Harmony of activities, 6 Harvey and circulation of blood, 66, 154 Head, structure of, 94 Heat of body, 70; economy of, 150; engines, 69; and entropy,' 201; flow of , 43 ; a form of energy, 3 1 ; measure- ment of, 37 ; mechanical equivalent of, 43 Hearing analysed, 174; organ of, 114 Heart-beat, regulation of, 88 Heart of vertebrates, 2; action of, 65; inhibition of, 142; innate heat of, 154; muscles of, 22 Heliotropism, 135, 183 Hemiplegia in man, 137 Hemispheres, cerebral, 98; connections of, 102; functions of, 125; removal of, effects, 139 Herbivorous animals, 84 Herd instinct, 142 Heredity, not explained, 193; what is transmitted, 225 Hibernation, 150 Horse-power, 37, 44 Hunger, 6 Hydraulics of body, 156 Idealism, 176 Immortality, physical, 208 Inanimate engine, 55 Inco-ordination, 6 Indeterminism, 222 Indeterminism of response, 138; evolution of, 139 Individualism, 142 Inertia and mass, 35 Inevitability of response, 137 Infusoria, 9, 11 Inhibition, 119, 141; of conduct, 143; of heart, 142 ; and shock, 142 Insanity and herd instinct, 142 Insects, locomotion of, 11 Instinct, herd, 142 Integration of activities, 4, 88; of functioning, 74 Intelligence and adaptations, 151; and spontaneity, 144 Intensity factors of energy, 50 Intestine, muscles of, 22 Intuition not communicated, 225; not conserved, 224 Irrevereibility, physical, 195 Irritability, 69; the basis of sensation, 106 Isolated systems, 45 Joints, ball-and-socket, 21 ; and de- grees of freedom, 19; elbow and knee, 20; Joule and theory of heat, 42, 158, 194; and ligaments, 20 Kant and a priori ideas, 54, 186; and space and time, 188 Kelvin and mechanism, 4, 194 Kidney, diagram of, 72, 73, 76; func- tioning of, 6 Kinetic energy, 39, 47; and vis viva, 154 Knowledge, elements of, 230; theory of, 186 Labyrinth, auditory, 114 Lavoisier, 157 Law of conservation and intuition, 224; qualifications of, 52 Laws, scientific, 170; of thermody- namics, 54 Legal inhibitions, 143 Lemniscus tracts, 100, 111 Levers in the skeleton, 20 Life, balance of, 84; concept of, 221 ; continuity of, 233; degradation to materiality, 226; duration of, 207; energy, cycle of, 57; and energy, 2 14; improbability of, 209; and indeter- minism, 213; intuition, 223; mani- festations of, 225; passage into material, 233; physical nature of, 210; and senile decay, 225; sub- stance, 233; tendencies of, 216 Light and plant growth, 82, 135 Limb appendages, 18 Limb girdles. 17 Limbs, modifications of, 17; skeleton of, 17 Liver of vertebrates, 03, 04 Locomotion in animals, 11; and cere- bellum, 125; ciliary, 11 ; in Crustacea, 2; in earth worm, *1 5; in jelly-fishes, 15; in molluscs, 15; in starfishes, 2; in vertebrates, 2, 17; in water, 11 Lodge and matter, 162; and waves, 168 Loeb, 159; and consciousness, 184 Lungs in man, 65; structure, 67; in vertebrates, 2 Lymphatic vessels, 61 Magnitude, orders of, 205 Malpighi, 66 Mass, nature of, 34 Materialism, 226 Materiality, 31; analysis of, 227; and energy, 37 Matter, conservation of , .")_' : Descartes' theory of, 161 ; and extension, 161; theory of, 36; universal, 36 Maxwell, J. C., and Descartes, 101; and gas theory, 199 INDEX 245 Measurements, microscopic, 206 Meat, tainting of, 78 Mechanical energy, unit of, 43 Mechanical models or theories, 4 Mechanics, the Cartesian, 160; classi- cal, 193 Mechanism, the animal, 4, 152 Mechanism, physico-chemical, 158 Mechanisms, automatic, 24 Mechanisms, bodily, integrating, 148; of joints, 20; sensori-motor, 116; non-skeletal, 22 Mechanistic conception of life, 152, 160; insufficient, 162 Medical research, 160 Membranes, animal, 61 ; synaptic, 108 Memory not conserved, 225; a factor in action, 178; and habit, 182; and morality, 224; a multiplicity in unity, 183 ; pure, 182 ; how stored, 183 Mental satisfaction, analysis of, 226 Mentality, subconscious, 223 Metabolism, 55, 56, 70; animal, 76; plant, 81, 219 Micro-organisms, activity of, 79; occurrence of, 77 Mid-brain, 104; and cord, 122, 146; and sensation, 122; and sensory tracts, 111, 115 Minckowski, 235 Mind, continuity of, 192 Mobility in animals, 9; and bodily environment, 55; freedom of, 190; inorganic, 10; organic, 10; patterns of, 11; in plants, 9 Molecules, 31, 33; collisions among, 199; dimensions of, 199; energy of, 48; velocities of, 199; weight of, 35 Molluscs, locomotion in, 11 Motion, cosmic, 10; Newton's laws of, 47; relative to the body, 189 Motor actions, complexity of, 118; organs, 20 Motor habits, 182; inherited, 183; mechanism of, 182; nerves of, 117 Movements, adaptive, 10; apparatus of, 13: control of, 121 Moulting in Crustacea, 15 Muscle-nerve preparation, 135, 172 Muscle tissues, 23 Muscles, antagonistic, 21, 22, 25, 29, 117, 119; contraction of, 21 ; control of, 121; disintegration of, 70; extensor, 21 ; fibres of, 23; flexor,21 ; of intestine, 22; involuntary, 24; receptors of, 121, 165; and skeleton, 14; striped, 23; trunk, 123; un- striped, 23; voluntary, 23 Muscular power, 38, 70 Muscular sensation, nerves of, 119 Mutations, 187 Nature, its form and content, L'^O; laws of, 229; materialistic descrip- tion of, 227; passage of, 230; as space-time data, 227; stuff of, 233 Neanderthal man, 99 Nebular hypothesis, 162 Neovitalism, 193 Nerast, 194 Nerve fibres, 26, 89; secretory, 239 Nerve-muscle preparations, 117 Nerves, 89; afferent, 25; Cartesian cells, 26; centres, 87; conception of, 155; cranial, 89: dendrites of, 27; efferent, 25; and energy, 133; respiratory, 88; sciatic, 27; spinal, 90; sympathetic, 88; terminations, 106, 107 Nerve tracts, 100; ascending, 100; association, 103; commissnral, 103; projection, 103; pyramidal, 104; sensory, 100 Nervous arborisations, 27 Nervous co-ordination, 88 Nervous impulse, 166; Cartesian con- ception of, 164; and energy, 131; model of, 131; paths of, 112 Nervous releases, 131 Nervous system, 90, 130, 145; central, 91; development of, 94; divisions of, 89; of earthworm, 91; and hi- de terminism, 174; mechanism of, 25; peripheral, 91; primitive, 147; scheme of, 26;structure of, 26, 91 Neurones, 27 Newton, 188 Nitrates in nature, 79, 84 Nitrification, 79 Nitrogen, fixation of, 85 Nitrogenous minerals, 80; residues, 80 Noumena, 176 Nutrition, organs of, 2 Nutritive materials, 69 Objectivity, 176 Oceanic bacteria, 80 Oil and animal life, 86 Olfactory paths, 113 Optic radiations, 116: thalami, 95^ tracts, 115 Organic and inorganic concepts, 215; movements, 10; reactions, 170 Organisms, remains of, 219; and the State, 7 Ossicles, auditory, 108 Oxygen in the lungs, 67 Oyster, shell of, 14 Pain, mental, 223; physical, 223 Pallium, cerebral, 96 Pancreatic gland, 58 Paralysis, motor, 93; sensory, 93 Parasites, animal, 9 THE MECHANISM OF LIFE Parthenogenesis, artificial, 159 Pascal's reed, 210 Passage of nature, 230; its relativity to life, 232 Pasteurisation, 78 Paths in nervous system, 101, 147 Patterns of locomotion. 11 Pelvis, 17, 19 Pepsin, 58 Perception, 24, 222; and dissipation, of energy, 184; and indeterminism, 181; kinds of, 178; nature of, 176; persistence of, 185; pure, 178; a sensori-motor process, 178 Personality, 234 Phagocytes, 69, 77 Photosynthesis, 82 Phototaxis, 136 Physico-chemistry of life, 214 Physiology an analysis, 164, 166; Cartesian, 192; and medicine, 166; the modern impasse, 193; and mind, 185 Pineal gland, 99 Pituitary gland, 99; and goitre, 7 Planck, *162, 194 Plants, characters of, 8; and entropy, 217; motions of, 9 Pleasure, analysis of, 226 Plexus, limb, 93; solar, 90 Poison glands, 12 Postures, animal, 11 Potential energy, 47 Prediction in science, 212; the test of theory, 173 Preservation of meat, 78 Priestley, 157 Probability and entropy, 203 Production by plants, 85 Proprioceptors, 121 Proteids, chemistry of, 59; foodstuffs, 58; putrefaction of, 79; residues, 77 Psychoids, 215 Psycho-physics, 175 Ptyalin, 58 Pulmonary circulation, 65 Putrefaction, 78 Pyramidal cells, 128 Pyramidal tracts, 104, 129; evolution of, 129 Pyramids, decussation of, 104 Radicles, chemical, 59 Radioactivity, 53, 86, 230 Radium, 53 Rankine, 194 Rational dynamics, 46 Reactions, coupled, 83 Real and unreal existence, 179 Reality and law of conservation, 53, 215 Reason, evolution of, 186 Receptors, 24; cold, 109; connections of, 102; distance, 24; equilibrium, 25, 109; gustatory, 25; heat, 25; joints, 110; muscle, 110, 112, 119, 124; olfactory, 109; touch, 25; vision, 107 Reference, frames of, 189 Reflex action and cerebellum, 120; control of, 120; and cortex, 120; craniospinal, 121; diagram, 117; gustatory, 238; inhibition of, 138; in normal animal, 138; olfactory, 238; predictable, 172; schematic, 120; simple, 120; scratch, 119 Reflex arcs, diagram, 29, 120; units of nervous system, 130 Regulations of activities. 136 Relations, 228 Relativity, 235; and Descartes, 161; and a finite universe, 197; and re- versibility, 196 Relays in nervous system, 100 Representations, mental, 176 Reproduction, 3 Respiration, automatic, 87; and carbonic acid, 87; gaseous inter- change, 67; mechanism, 17, 65; movements, 68; regulation, 87; theory of, 157 Responses, adaptive, 110; animal, 10, 13; and heredity, 150; indeter- mined, 138, 175; and stimuli, 139; unpredictable, 173 Restlessness and decerebration, 141 Retina, 107, 115 Reversibility, physical, 43, 195 Ribs, skeleton of, 16 Rolandic fissure, 104 Roots of plants, 9 Rotifers, locomotion in, 11 Rumford and heat, 157 Russell, 185 Rutherford, 53 Salivary gland, 58 Saltpetre, 79 Saprozoic nutrition, 84 Science, method of, 166, 170; and utility, 214 Scratch reflex, 137, 138 Second law of thermodynamics, 194; reversal of, 198, 216 Secretion, Cartesian theory of, 155; chemistry of, 239; psychical, 239 Segments in nervous system, 92 Sensation, 24, 106, 222; and action, 133; chemical, 108; complex, 116; disordered, 52; exteroceptive, general, 110; gustatory, 25; and knowledge, 133; muscular, 109; nature of, 174; olfactory, 108; paths of, 111; and perception, 175; phvsical, 176; proprioceptive, 24; tactile, 25; visual, 107; INDEX 247 Sense organs, stimulation of, 24, 107; accessory, 108 Sensibility, general, 1 1 1 Sensori- motor activity, 28 ; mechanism, 117; system, 13, 145; Sensory mechanisms, 24 Servetus, 66 Sherrington, 165 Shock, nervous, 142 Skeleton, 152; absence of, 15; analysis of, 19; axial, 16; of birds, 16; con- nections of, 14; in Crustacea, 15; internal, 15; joints in, 16; and mechanics, 14; of mollusca, 14; muscles and, 20; plan of, 13; size of, 16; of snakes, 16; of sponges, 14 Skull, structure of, 16 Smell, 33; associations of, 114; deli- cacy of, 109; sensation of, 113 Snaring devices, 12; Social complexes, 7 Socialisation of activities, 7 Solar radiation, 69, 86 Soul, rational, 154; sensitive, 192 Sound, analysis of, 116; waves, 168 Space, abstract, 191; Euclidean, 189; a form of sensibility, 191; measure- ment of, 169; non-Euclidean, 235 Space time, 228, 235 Spermatozoa, 11 Spinal animal, 118 Spinal cord, 92, 93; autonomy of, 94; connections of, 110 Spinal frog, 136, 172 Spinal nerves, 93; roots of, 93, 165; sensory, 111 Spinal reflex, 136, 137 Spontaneous actions, 105 Spooks and energy, 52 Spirits, animal, 155; natural, 154 Spiritualism, 193 Starch in foods, 60; in plants, 83 State and organism, 75 Statistical and individual concepts, 212 Steam engine, 42, 55 Stearic acid, 60 Sterilisation, 78, 79 Stimuli of environment, 13; intrinsic, 140; physical, 13; and sensation, 106 Stimulus and response, model of, 132, 134 Structure of animals, 2; Descartes' analysis of, 156; and function, 3; modifications of, 12 Sweating, regulation of, 88 Swimmerets in Crustacea, 18 Swimming, 11 Subjectivity, 53, 176 Submaxillary gland, 237 Sugars, chemistry of, 60 Suggestion, mass, 180 Sunlight, energy of, 218 Sympathetic nervous system, 88, 90, 165 Synapses, 27 Syntheses, organic, 159 Systems, physical, 45 Taste, 33, 108 Taxis in animals, 135 Techniques, origin of, 182 Telepathy, 225 Temperature, 31 Tendencies, logical, 216 Tendons, 21 Texture, 32 Theory, astronomical, 210 Thermodynamics, 194; history of, 157; laws of, 52; and life, 69; and mechanism, 69 Thinking and action, 188 Thomson, Sir J. J., 53, 194 Thomson, Sir W., 203 Thorax, movements of, 17 Threshold of sensation, 106 Thumb, mobility of, 19 Time, astronomical, 191; dimension, 236; measurement of, 191, 206; a reversible series, 196 Touch organs, Il8 Toxins of bacteria, 77 Tracts, nervous, 99 Transformations, releasing, 48 Transformism, 234 Trial and error, 188 Tropisms, 9, 13, 135, 180 Trypsin, 58 Unavailable energy, 42 Universe, condition of, 97; cycles in, 205; dimensions of, 204; and dissi- pation, 204; finite, 197, 236; in passage, 231; running down, 203; structure of, 162 Urea, chemistry of, 59; excretion of, 70, 76; nitrification of, 84 Urine, 73 Variability and determinism, 181 Variations, organic, 181 Vascular system, 152 Vegetative processes, 217 Veins, 63 Ventricles of brain, 97 Vertebrae, articulations of, 16 Vertebrates, locomotion of, 2; skele- ton of , 17 Vestal virgins, 215 Vestibular sensation, 1 14, 124 Vicarious functioning, 144 Vision, organs of, 108 Vis viva, 55 Vital impetus, 223, 225 Vivisection, 160 248 THE MECHANISM OF LIFE Volition and cortex cerebri, 140; and motion, 118 Voluntary muscles, 23 Vortices, Cartesian, 162 Vries, de, 150 Walking, 11; nervous mechanism, 123 Waller and nerves, 165 War psychoses, 143 Waste matter of body, 70 Watt, 157 Waves, 168, 169 Weather, analysis of, -'21 Weber, 224 Weight, 34; and mass, 36 Weismann and germ plasm, 193 White matter, 91 Winking, 2,8, 29 Wohler, 159 Work, 37; capacity for doing, 37 Working substance, 42; in life, 56, 77 Wrist, 18 Yeasts, 78 b PRINTKD IN GREAT BRITAIN BY BIU.IXO AND 80N3, LTD., OUILDFOnD AND ESHER THIS BOOK IS DUE ON THE LAST DATE STAMPED BELOW AN INITIAL FINE OF 25 CENTS WILL BE ASSESSED FOR FAILURE TO RETURN THIS BOOK ON THE DATE DUE. THE PENALTY WILL INCREASE TO SO CENTS ON THE FOURTH DAY AND TO $1.OO ON THE SEVENTH DAY OVERDUE. MAR 1 4 1949 FEB 271951 APR 2 3 1951 APR 2 1951 L-. ^rc 1 * iasq "1 3LfLfc 33 Vu^Wjttc V ^ ( ^ *""'' S '- 195b LD ui -lOOm-12, 81