AMERICAN~ GRAPE C1ULTURE.                 279
Renew old vineyards of Isabellas, Concords, Catawbas and othel
grapes, unfortunately set, is the rule; and is, I am pleased to say, in
the reach of all who have theln. Of course no man will renew by grafting an Isabella or other vineyard in a place where ti8. grape ripens
well, for the sake of getting a better grape, recommended to him by
any one.




280            )VL1 V YORKi STATE AG6RICU'LTURAL SOCIETY.
CONTRIBUTION TO THE BIOLOGY AND HISTORY OF
THE DEVELOPEMENT OF THE USTILAGINE 1t.
BY DR.'. FrSCHER VON' WALDHEIM.
[Translated from the German in Pringshcim's Jahrbucher fur Wissenschaftliche Botanik, vol. vii,
part 1-2, 1859.]
I.-IIISTORICAL  INTRODUCTION.
As it is known, the  Ustilagineae  which, as endophytic fungi, have
been  frequently examined  and with so little success, have  been  from
early times the objects of attention, not only of the agriculturist, but
also of many naturalists.   Yet, as  to  the  nature  of these  parasites,
there were entirely diverse and  mostly erroneous opinions, even until
the  second  quarter of our century.   They were, even  by later investigators, considered first as symptoms of sickliness in  the  supporting
plant, Losana,' Turpin,2  Schleiden;s  then  as  disease, Unger,4  also
Wiegmann;5 or as the product of the plant, Fries,6 IMeyen.7  On the
other hand, Bulliard,s Prevost,9 Link,l~ De Candolle," Philippar,l2 and
others, and especially in modern times, Tulasne,l3 Leveille,'4  de Bary,D'
Kuhn 16 and Hoffmann 17 attained only the right perception of the fungi
as parasites, and as the cause of sickness in the supporting plant, and
sustained it with irrefutable proof.
1 Losana, Delle Malattie del Grano. Carmagnola, 1811, p. 92.
2 iurpin, Memoir sur la Globuline (according to De Candolle's description).
s S&cleiden, Grundz. d. wissensch. Botan., III, Aufl., 1850, P. II, p. 34.
4 Unger, Die Exantheme d. Pflanzen, Wien, 183.3, p. 356.
5 WTiegmann, Die Krankheiten nnd krankhaft. Missbild. d. Gewachse, Braunschweig, 1839, p.
116, sqq.
6 ~Fies, Syst. mycol. Gryphisw., 1832, III, p. 456, etc.
7 Me/en, Ueber die Entwickelung des Getrcidebrandes in der IMais-Planze ( Wiegm. Arch. f. Naturg,.,
1837, p. 419-421).
8 Bdlliard, Hist. d. Champignons de la France. Paris, 1791, I. p. 90.
9 Prevost, Mcm. sur la cause immediate de la Carie. Montauban, 1807.
10 Link, Observ. in ord. plant. nat. Dissert. I, 1809, and in other publications.
11 De Candolle, Physiol. Vegetal. Par. 1832, III, p. 1435. Also in earlier publications.
12 Philippar, Traitc organogr. et physiol.-agric. sur la Carie, etc. Versailles, 1837.
13 Tulasne, Memoir sur les Ustillg. comparees aux Ured., Ann. d. sc. nat. 3, ser. t. vii, 1847. Second
Mem. s. 1. Uredinees et les Ustilag., Ann. d. sc. nat. 4, ser. t. ii, 1854.
14 Leveille, Recherchcs s. 1. developp. des Ured. Ann. d. sc.-nat., 2 ser., t. xi, 1839. Etudes sur les
champign. de la fam. d. Ured., Diet. univ, d'hist. nat., par d'Orbigny.
15 De Bary, Unters. uber d. Brandpilze, 1853. Recherches sur le developpement de quelque champ.
paras., Ann. d. sc. nat., 4 ser., t. ii, 1863. Also in other publications.
16 Kuhn, Die Krankh. der Kulturgew, 2 Aufl., 1859.
17 Iloffmann, Ueber den Flugbrand, Bot Unters., pr Alis'ed by Karsten, 1866, p. 192, seq., et,




FISCHER  VoN JtALDIIEJ —    I   UST'LAGI.FiE.            28 1
In order to save repetition, I refer the reader directly to the works
of Tulasne1 and  de Bary,2 where the details of the different historical periods regarding this subject will be found defined and illustrated
up to a period later than 1840.  We will, therefore, pass over these,
and  only consider the  most important facts which were observed
mostly by later naturalists.
1. The localization of the Ustilagineae (or to express it more clearly,
the place of their development) became an object of the earliest investigations, as well as such changes in  the supporting plants as were
caused by them and most easily perceived.
The work of Tulasne, just mentiored, regarded the parasites in
this connexion as worthy a detailed observation.  Especially are  the
sickly changes in the affected plant particularly specified, which are
produced  by Tilletia  Caries Tul., and Sorghi vulgaris Tul., Ust.
Carbo  Tul., Maydis Lev., and  antherarum   Fr.  The observations
regarding Ustilago Vaillantii Tul., Candollei Tul., Utriculosa Tul.,
and Duriaeana Tul., are incomplete;  still less specific  are those
regarding the remaininlg species mentioned in the systematic part of
that work.  Accurate delineations of the parts of the plants affected
by this parasitic disease add to the value of Tulasne's treatise.
Subsequently we find also in this connexion, many accounts of
Ustilago  secalis  Rabh.  (Rabenhorst).3    Urocystis  occulta  Rabh.
(v. Schlechtendal),4 Ust. Ilaydis (de Bary)5 (Kiihn),6 longissima LeV-.,
hypodytes, antlherarum  (de Bary),7 Candollei (de Bary),8 Sorisporium
Saponariae Rud. (de Bary),9 Tilletia Caries, destruens Dub., Urocystis
occulta (Kiihn),l~ and Ustilago Carbo (Kuiihn 11 and Hoffman'2).
2. Early naturalists had also given attention to the mode and nature
of the propagation of Ustilagineae.  First, however, Prevost, that,
indefatigable observer of parasites, made the surprising discovery that
the fungus spores germinate.  Hie observed the germination development of Tilletia Caries and Ustilago Carbo.  In the former, he discov1 Premier Mem. s. 1. Ustil.
2 Brandpilze.
3 Rabenhorst, in Flora, allgem. bot. zeit. 1849, p. 209 sqq.
4 Schlechtendal, Bemerkungen zu einer Decade fur d. Flora v. Halle n. Pilze. Bot. zeit. v. MloAs
and Schlechtendal, 1852, p. 602 sq.
5 Brandpilze, p. 4-8.
6 Ibid p. 70 sq. Unfortunately I am unacquainted with Kuhn's treatise on Ust. MIaydis in Rabcnkorst's Hedwigia, 1863, and in the Zeitschrift d. landw. Centralver. d. Prov. Sachsen, 1866, p. 108.
7 Ibid, p. 8-12.
s See official report of the meeting of the German Naturalists and Doctors in Gottingen, 1854
9 Ibid. Later likewise in Champign. Paras. p. 96.
10 Ibid, p. 54 sqq.
11 Ibid, p. 64. ib. p. 51-53, a short list is given of the supporting plants upon which Ustilagine.e have
been observed.
1r Ibid.




282        NEW  IYORIK STATE AGRICULTURAL SOCIETY.
ered on the exterior of the promycelium, the so-called primary sporidium and the spores detached from it.'  In Ust. Carbo he perceived
only the germ tube, provided with two or three branches.2
The germinating power of the Tilletia spores was agaiL verified by
De Candolle,3 Carm  and Vandenhecke.4  Even Tulasne failed in his
experiments upon the germination of this Tilletia until 1853, wihen
lie communicates his observations of their more or less capricio.us
germination.5  IIe discovered lollg and short, sterile  germ  tubes
crowned with clusters of eight or ten primlary sporidia (Tulasne's secondary spores), and described also the till then unlknown II-formed
connexion of the sporidia which appears only out of water, some of
the united spores germinating, but a greater number of them producin_, secondary sporidia which also germinate, or are capable of producing tertiary sporidia.
Kuhn gives a yet more particular account of the germination of
this Tilletia.G   According to him, the mode of germination of Tilletia lolii Auersw.7 is very similar.
De ]3ary, whom  I here follow, carried out this termiinology for
these, as Well as also for Ustilagineae, by designating the first tube
proceeding from the spore as promZyceliumn, the bodies first detached
firom this as sporidia, the next as secondary sporidia, and so on.8  This
terminology was already employed by Tulasne for YT. Caries and Ul.?2ecep)ta culo rum.9:Besides Prevost, Tulasne observed again the germination of lUstil(ago Carbo.  Hie saw, in like manner, upon the promycelium  one or
two little lengthened branches (probably sporidia) as well as their
separation by contraction.'0  Bonorden also succeeded at this time in
germinating the U. Carbo."l  Miore complete observations were published by Kuhn.1  He menltions the formation of primary and secondary
sporidia, knee-formed connexion of the promycelia, and the germination of separate portions after breaking up.
In  his first memoir Tulasne describes also the germination of'Ibid.
2 Ibid, p. 20 sq.
3 Physiol. veget. 1832, III. p. 1436.
t In 1835. See Tulasne ir Mem. s. les. Ustil. p. 35.
5 2e Mem. s. i. Ured. pp. 161-163.
6 Ibid., pp. 57-60.
7 Ibid., pp. 61, 62.
8 De Bary, Morphol. und Physiol. d. Pilze (I-andb. d. phys. Botan. Yon Ieofmeister II. B.), 1866, p.
150 sq.
9 2e MIem. pp. 159 sqq.
o0 Ir Mem. pp. 32-34.-2c Mem. p. 157.
I Bonorden, Handb. d. allg. Mykologie, 1851, p. 39.
12 Ibid.,p. 66-67.




FISCHER VON WALD UEIJ-   - USTILAGINE3.           283
[Jstilago antherarum.1  The promycelium  attains twice the diaimeter of the spores, towards the middle is of greater circumference,
and divides into two or three parts.  Later,2 Tulasne adds, that the
promycelium, during its attachment to the spore, divides and,-separated from  it, attains a circumference three times larger than originally, but without increasing its length.  According to de Blary,5
the germination of this species is similar to that of Ust. Carbo.
Tulasne also describes in his second memoir, the germination of
the Ustilago receptaculorum of Tragopogon pratensis and Scorzonera
humilis.4  Ite observed in the first the cross division of the promycelium into from two to four cells, out of which sprout, singly or in
pairs, almost stemless "reproductive bodies" (sporidia). Among those
broken off, many are connected in pairs by a bridge. Ust. receptaculorum of Scorzonera humilis showed smaller dimensions of the promycelia and sporidia, and a quicker falling off of the latter as well as a
greater number, after two or three days.  Very true to nature are
the accompanying illustrations; also those of de Bary, who remarks
of this Ustilago, that "as it appears the sporidia are successively
detached."5
KIuhn was the first to describe and delineate Ustilago destrnens, Maydis and Urocystis occulta.6  In Ust. destruens Kuhn saw
very similar and also partly different phenomena of germination
from that of the Ust. Carbo, a partly hyaline production and granulation of the promycelium, the formation of stemmed egg-shaped
sporidia on the side, or unstemmed terminal ones, as well as lengthening and ramifying single germ  tubes.  The knee-formed lateral
branch of the promyceliumr  he thinks sometimes becomes separated.
Ustilago Maydis germinates by forming a straight, wavy or bent
promyceliumn (during which process the end may shoot out in a
thread), which seldom has a lateral branch, and still more seldom
shows the knee-formed bending.  Sporidia appear on the side and.
point of the proimycelium.
Urocystis occulta, according to Kuhn, developes from a skein of
spores, at the most, from two to three germ tubes of different thickness.  On the stronger promycelium  only, from  two to six primary
sporidia appear, mostly of different lengths.  The primary sporidia
frequently separate, grow then further, and only seldom detach secondary sporidia.  In two cases, weak promycelia occurred on the side
or point instead of sporidia. In only one case an H-formed connexI P. 34 sq.                      4 P. 158-160.
2 2e Mem., p. 158.               5 Ibid., p. 150.
3 Morphol. und Phys. d. Pilze, p. 151.  6 Ibid. page 69, sqq.




284        YEWV YORKI STATE AGRICULTURAL SOCIETY.
ion of the primary spores was shown, and that in the upper part.
On the contrary, according to de Bary,1 such connexion of the Uroc.
occulta and pompholygodes does not happen.
Hoffman was the first to mention the germination of the. Ust.
longissima.2 - De Bary 3 gives the first accurate delineation of its promycelium, together with its sporidia.  The promycelium, according
to him, is a thin cylindrical tube, from two to three times the length
of the diameter of the spore; from its point is detached a spindleformed cylindrical sporidium; but whether generally single, or
several in succession, is still to be determined.4   Moreover, Hoff;
mann  communicates a few facts regarding the influence of various circumstances5 upon their germination; also Kuhn6 and de
Bary.7
3. Concerning the spore form of the Ustilagineo, as well as the
mycelium, Meyen was the first to describe the spore-forming threads
(according to my terminology), and the formation of the spores from
them.8  He made observations, though by no means complete, of the
Ustilago Mlaydis.  According to his description, in the maize smut
balls, are seen little transparent slimy deposits, from which spring
thread-formed, unjointed, ramifying " little plants," which are repeated,
ramifying and becoming corymbiated, from which little irregular
globular bodies detach, until at last they all break up into such
globules.
Leveille describes the latest conditions of Ust. Maydis, as seen by
him, as short, knotty, articulated threads, and simply adds that the
spores spring from them.9
Meyen soon after confirmed his earlier investigations on the spore
formation of Ustilago Maydis, and added some new discoveries on
that of Ustilago hypodytes.10  The latter appeared to him  as a compound mass of slime-covered threads, on the surface of which appear
the " brand pustules," by the detaching of those threads at the point,
until the whole mass was converted into such "pustules."
Tulasne knew the latest condition of the Ustilagineae only as a
1 Ibid., p. 152.
2 Hoffmann, IJntersuch ub. d. Keimaug d. Pilzsporen, Pringsh. Jahrb. f. wiss. Botan., 1860, II,
Bd., p. 273.
3 Ibid., p. 151.
4 Ibid.
5 Ibid., p. 297, sqq. Compare Ueber Pilzkeim. Bot. Zeit. v. Mohl nd Schlechtend, 1859, p. 2,09, sqq,
6 Ibid., pp. 46-48.
7 Ibid., p. 209, sqq.
8 Ibid., p. 419, sq.
9 Recherches s. I. developpement des Ured., pp. 5-16.
10 Xreyen, Pflanzenpathologie, 1841, p. 103.




FISCITER VON WALD1EZI — STILA GINVEH.                28 5
gelatinous slimy mass, which in Ustilago Maydis was sometimes
found in the inter-cellular spaces, and even in the cells.' These
masses separate (in Ustilago Ilaydis) into little parts, around which
is formed from the surrounding slime, a double membrane, the endosporium and episporium.  The spores ripen from  the centre of the
attacked parts of the supporting plant toward the periphery.  Tulasne
also discerned in the slimy mass, short, ramifying threads, with nitrogenous contents, evidently spore-forming threads, whose import
remained obscure.  In like manner he made the same observation
in regard to the spore formation of the Ustilago antherarum.2
Tulasne3 first observed the formation of the Tilletia spores upon
the ends of short branches springing from a common stem; for Prevost4 and also Philippar5 had only a dim insight into the manner of
the formation of Tilletia spores.
De Bary succeeded in showing the formation of the spores not only
of Ust. Maydis, but also of Ust. longissima, hypodytes, and antherarum, out of distinct spore-producing threads.'  Ite designated, however, these threads by the name of "Mycelium."  The position of
these in Ust. Maydis is supposed to be always in the inter-cellular
spaces, as had been previously,7 and was afterwards,8 mentioned by
Unger also; they appear colourless, somewhat dark, as it were
opalescent, and change into a slimy gelatinous mass, from which the
spores form from the point of the branches backward.  The account
of the effect of different reagents on these filaments,9 leaves no doubt
that they were only spore-forming, threads. The latest spore conditions of this species appear as circumscribed, roundish masses, like
beads, strung on an indistinct thread.  Then a membrane formed
around these bodies, and the cells almost doubled in size.  When the
spore became ripe, this thread had disappeared.  De Bary also
examined the latest mass in the intercellular spaces of U'stilago
longissima, in the form of short thin threads, which, for the greater
part, had already detached spores, and, in fact, frequently in bead-like
branched groups.10
As in Ust. longissima, so  de Bary perceived only spore-forming
threads in Ust. hypodytes.1t  This is indeed proved by his opinion
I lr Mem,, p. 20, sqq.                    9 Ibid., pp. 13-15.
2 Ibid., p. 25.                           lo Ibid., pp. 8, 9.
8 Ibid., p. 30,                           11 Ibid., pp. 10, 11.
4 Ibid., p. 10.
5 Ibid., pp. 1S and 21.
6 Brandpilze, pp. 5-14.
7 U7iger, Beitrage zur.i vergl, Pathologie, 1840, p. 24.
8 U7nger, Anat. u. Physiol. d. Pflanzen, 1855, p. 129..




286        NEW YORK STATE AGRICULTURAL SOCIETY.
at that time regarding this ustilago to be an epiphyte that only
existed on the epidermis; an opinion which was also defended by
MAeyen.1 The ripening of the spores takes place here on the outside
toward the epidermis.
De BaIly found, in the male blossoms of the Silene Otites, a whitish
mass of the Ust.  antherarluln  composed  of  delicate, colourless,
tufted, ramified and confused threads, from  the points of which are
formed the bead-like spores.2
De Bary soon after treats of the spore formations of Ust. Candollei and Sorisporium  Saponariae.s  In the latter the spores are
formed out of a granulous slime, surrounded by clustered "Mycelium" (also spore-forming threads).  The spores of Ust. Candollei
are formed in the same manner as inl the other species of ustilago,
only that at the same time they form  in the ovary a columnella,
together with peridia, also mentioned by Tulasne.4
According to Kiihn's description he knew about the mycelium of
of Ust. Maydis only imperfectly, and if his remarks regarding the
threads designated by him as mycelium are correct, then it was the
same with Ustilago destruens and Tilletia Caries.5 Although in the
introduction6 he speaks in general of the mycelium of the smut
fungi, he refers, without doubt, afterwards only to Tilletia Caries and
Ust. destruens, which alone are used as examples, especially as later
true mycelium  is specified only for Ust..  laydis. The mycelium,
according to him, developes from  the germinating threads which
penetrate the supporting plant  abundantly and  spreading in all
parts of the plant, if many germinating threads penetrate;  in
the contrary  case, growing up  with  it more  sparingly.  The
under, older part of the mycelium threads soon dies away,
becomes empty of protoplasm, clear as water, and disappears; they
only continue growing on the point. This is the reason its observation is so difficult in the developed supporting plant.  Kiihn refers to
the plate, which, however, to me, has rather the appearance of sporeforming threads; while Kuhn, as seems probable, much oftener
considered these threads as the true mycelium; consequently he was
not able to arrive at any correct conclusion. This is shown in the
following definition: "' The mycelium threads of the smut fungi are
1 Pflanzenpathol, pp. 122 and 124.
2 Ibid., pp. 11-13.:3 Official report of the meeting of German naturalists in Gottingen, 1854. Also Flora, allg. b3t.
zeit., 1854, pp. 647, 648.
4 1r. Mem. pp. 94, 95.
5 Ibid., pp. 49-50, and p. 73.
G Ibid., pp. 49-50.




FrISCHER VON TALD HEJ:r — STTrL GINVEhE.        287
very delicate and of a very small diameter, and easily known by their
denYse, blueisll-coloured protoplasm contents."
(iiKhn gave a more copious description of the spore formation of
Tilletia Caries, which confirmed Tulasne's investigations,2 and of Ust,
Carbo, Maydis, and Uroc. occulta. The latest condition of the Ust.
Carbo Kiihn observed as " crum.by, soft, glittering" masses, composed
of numberless tender ramified threads.  Through the entire length
of the branches, protuberances developed, which, at the end of the
developement, separate into spores.
In the Ust. Maydis it is thought the spores are formed on especially
fine, tender threads, which grew  from  the "principal mycelium
threads."4  The ends of these, swelling for the purpose, produce
granules which increase in size, whilst the threads themselves, at the
same time, empty and branch into corymbiated forms.  The grannies
become separated one fiom another by a cross-wall, and the resulting
cells, assuming a roundish shape, become spores.
Kiihn was the first to describe the peculiar spore formation of the
Uroc. occulta.5  Bladder-like swellings, which become cells, originate
in the spore-forming threads.  Each such cell remains single, or
divides into two or three, and by side shoots forms several secondary
spores, together constituting the spore-skein. The spores which come
first commonly occupy the middle of the little pile of spores; sometimes the secondary spores are only partially joined, but always have
their own episporium.
De Bary remarked, in this connexion,6 that the little clear cells are
not sister cells of the dark spores, but limbs of the young hyphen,
which appear on the outside of the spore while yet young. De Bary's
opinion appears to me to be correct, because an entirely analogous
process takes place in the formation of the spore-balls of the Sorisporiumn Saponarice.
That Hoffinann made no real distinction between mycelium and
spore forming threads, is to be seen from  his treatise on Ust. Carb.7
No real mycelium is mentioned during the description of the spore
formation. All the thicker and thinner branched and bent threads
have been nothing but spore forming threads in different degrees of
development.8
This is plain, from Hoffmann's statement, that they were;" crowded
full of plasm," as well as from their shape and appearance. For even
IIbid., p. 50.                  5 Ibid., pp. 78, 79.
2 Krankh. d. Kuturg., p. 56, sq.  6 Morph. und Physiol. d. Pilze, p. 125.
3 Ibid., pp. 65, 66.            7 Ueber den Flugbrand.
4 Ibid., pp. 73, 74.            8 Ibid., p. 201.




288        NEWr  YORiK STA TE' A GRICULTURAL SOCIETY.
the "fine mycelium'; which sometimes is called the scarce myceliuin3
and which is something thought to spin completely around the young
ear as a mesh-net (later in my observations I will return to this latter
circumstance) can only be considered as a transition form of the true
inycelium, which is formed entirely different in spore forming threads,
or as a superfluous, so to speak, wasted residue of it (only a conjectnre).  The investigation of the spore formation during the time of
the gelatinous condition of the threads in which only the glittering
parts of the cont'ents are still to be seen, makes Ioffmann's conjecture,
among others, that in the origin of the plasm  grains there is a similarity to an independent cell fo.rnation.1
The perennial character of the imycelium (of the Sorisporiunn Saponarfie), was first made known, some years ago, through de Bary's successful examinations.2  He found it in all parts of the supporting
plant, and in order to demonstrate its vitality planted a saponaria
branch in which mycelium  existed.  In the course of a few months
two shoots appeared covered with leaves. An examination of one of
the branches showed  mycelium  very plainly; the other developed
blossoms later, whose flowers were all infected with the parasite. By
this experiment the correctness of Aymen's investigationswas proved,
who mentioned as early as 1760, that in his garden for five successive
years the same plants of wild pinks and saponaria officinalis had only
smutted flowers.3  Besides, de Bary was the first to refer to the suckers (see my observations) of the Sorispor. Saponarie, in that he imentioned that the delicate intercellular mycelium threads send stronger
branches into the cells, which generally have an irregular contour.
and by braiding together form little various-shaped skeins resemblingvery small membranes.4
4. After the spores of the Ustilaginem were discovered, the idea
became common that it was only through them  that the parasite
could be in the supporting plant; but in what manner has been
explained in different ways up to the present time.  First it was supposed to be the granules contained in the spore (De Candolle,5 Link);6
after the discovery of germination, it was the germ tubes which penetratedthe supporting plant and transplanted the parasite (Prevost,'
Le'veille).8  I omit the still less remarkable opinions of Bulliard,'
Sylvestre,lo and Philippar," and mention only the opinion of Tulasne,
Ibid., p. 202.                     6 Ibid.
2 Champign. paras., p. 96.         7 Ibid.
3 Tulasne, Ir Mem. sur leos Uslil., p. 42.  8 Dict. univ. d'hist. natur d'Orbigny, ibid.
4 Ibid., p. 97.                    9 Ibid., I, p. 91.
5 Ann. du mus. d'hist. natur. de Paris. 1836, t. 10 Annales de Fromont, t. iv., p. 145.
&i. p. 56 sq. Physiol. veget. III, p. 1437.  11 Ibid., pp. 37-38.




FIscH ER VON WTA LDHEIr — US'z.IA GIANE'A.    28 9
to whom it seemed most probable that the germ tubes of the Tilletia
spores penetrated either the young stems or root of the wheat, by
piercing the epidermis of these parts.'
De Bary, relying on the similarity to the Uredines and on his awn
observations, believed that the germ tube enters the stomata of the
supporting plant and is further developed in the interior of it (as, for
example, Ust. iMaydis); or it developes only on the epidermis, as
Leveille and Decaisne observed in Ust. receptaculorum  and Meyen
and himself in Ust. hypodytes.2  The mycelium'of Ustilagineae,
according to the view of de Bary at that time, was newly generated
each year from the spores, and at each time through the spore formation became completely used up and disappeared.3  At that time,
de Bary conceded the possibility of its perennial character, but
considered it hardly probable.
Kuihn mentions having directly investigated the penetration of the
germ threads of Tilletia Caries in wheat, and even in the root nodes
of the little germinating plants.4 IHe germinated, for this experiment,
grains of wheat sprinkled with numerous Tilletia spores, placed tliem
in earth near the surface, and kept them in a sufficiently moist condition:.
Sections through the germinating wheat, even before the death of the
rootlet and the complete development of the first leaf, showed him the
gerim threads penetrating directly through the cells of the root nodes.
The penetrating germ threads arise from the sporidia, or are an immediate attenuation and ranmification of the point of the germ tube.5
De Bary mentions having repeated Kuhn's observations on the
penetration and vegetation of the Tilletia Caries with success.6  In
another workl, however, he supports the statements of the penetration of the "germ tubes" only from the investigations of Kuhn and
Hoffmann. We may conclude from  this that his earlier approbations
of Kiihn's experiment related only to the vegetation of the Tilletia
Mlycelium, and not to its inoculation of it.
Ioffmnann considered the inoculation of the Ust.Carbo, demnonstrated
by his own experiments.8  After the consideration of several places
of penetration, Hoffinann regarded the space between the inner
side of the split root sheath and the young root as the normal and
most suitable place for that purpose.  Spores deposited at this point
developed an abundant my6elium which grew inward and upward
until near the vegetation point.  Whence this nmyceliumn originated,
I lr Mem., p. 41.               5 Ibid., p. 61.
2 Brandpilze, p. 123.           G Champign. paras, p. 99.
3 Ibid., p. 129.                7 Morph. u. Physiol. d Pilze, p. 186.
4 Ibid, pp. 48, 49.             8 Ibid., pp. 202-206.
[AG.1  19




290        NEW  YORK ST  TE  GRICUVLTUIRA:A    SOCIETY'
froln the germ  tube or out of the sporidia, is not mentioned, nor its
outward appearance. The accompanying delineation resembles little
my representation of the myceliuln of Ust. Carbo.  Moreover, the connexion of this mycelium with single spores, described by Hoffmann,
might have been accidental; that is to say, merely a juxtaposition or
superposition of tile spores having taken place. Only the undeniable
connexion of the observed threads with ustilago spores would be a
sure proof of the mycelium  originating from them, if the latter had
not possessed air entirely characteristic stamp of its own.
Besides, Hoffmann regards a penetration through the primary nodes
as possible, because after direct application of the spores he could
perceive, at this place, inward penetrating germ-tubes.' IHoffmann
appears also to concede that the penetration could take place through
the stomnata. He likewise tried inoculation by applying the spores
on the burst root-sheath or to the uncovered primary nodes or in
the region of the point of vegetation; in this way, in a few  cases
out of several hundred failures, lie produced  smutted ears.  The
sprinkling with spores of other parts of the growing barley-plants
resulted only in failure.
Hoffmann also states2 that he placed spores of Ust. destruens, with
millet seeds moistened with spittle, and suceeeded in bringing them
to germination in boiled earth.  From  six of these seeds Hoffmann
produced three plants with smutted panicles.3
5. It remains to mention, in conclusion, the observations of spore
structure, which belong also to the process of the development of the
parasites.
Tulasne first discerns a double membrane in the ustilago spores:
endosporium  and  episporium.  The first, the inner, delicate and
only perceivable when the episporium bursts; the latter is colored and
smooth, or provided with various protuberances, which in Tilletia
Caries, after the application of sulphuric acid, became visible as a protruding net formed of little polygonal meshes.4  The contents of the
spores are pale, and composed of oleaginous grains. The yellow
iodine reaction shows the presence of nitrogenous substances.
De Blary agrees with Tulasne in regard to the membrane, but finds
[ Ibid., pp. 204-205.
2 Ibid., 206.
3 That the infection with spores sometimes produced smutted flower stems, showed only the possibility in these particular cases of a previous presence of the parasites in favorable conditions for
propaLgation, which, hlowever, had escaped his examination, especially as all parts of the plant cannot be examined wheln their further development is desirable.
1 ir irnem., p.'1.




FISCHER VON WALDHEIM — [USTrLAGIN-Ee.                            291
in  the contents a grain, apparently  identical with a nucleus.l  Later,
however, de Bary refutes these latter views, and considered the grain
only  a protoplasm   body.2   Sugar and  sulphuric  acid  coloured  the
grain rosy red.   The protuberances of the episporium  of Ust. Maydis
and antherarum  did not disappear by boiling in a solution of potash;
according  to   de Bary's opinion  at that time, the  episporiilmin  may
originate before the inner skin.3
Kiihn, on  the  contrary, considers the  episporiumn   a result of the
secretion of the endosporium, and analogous to the cuticle of the
pollen  grains.4   Later  de Bary expresses the same view.5   Various
chemical details concerning the spores are also mentioned at the
same place by  de Bary,6 and at other places by Hoffmann.7
6. A  review  of the  foregoing  historical data, in which  the  so far
known results of many excellent experiments have been briefly stated,
renders it possible to decide on the questions yet to be answered.
ThCe JIyceliqzIm  of the  Ustilaginece  is the least known of all parts
of the parasite.  We only find it mentioned for Sorisporium  Saponarie,
partly also for Ust. M3aydis, and if, in the inoculation experiment of
Kiihn and Hoffmann, a myceliuml has been found peculiar to the
investigated parasite (which I may not doubt), also for Tilletia Caries
[Kiihn], Ust. destruens [Kiihn], and  Carbo  [Hoffinann].   But notwithstanding  these accounts, we possess no  thoroughly characteristic
description  of the mycelium, but merely intimations which  require
further developement.
Also  spore-forming parts, although  multifariously  observed, are
not sufficiently investigated, to allow a clear view of the changes proceeding by degrees in them from the beginning of the formation of
the spores, until they become fillly ripe.   And then the spore-forming
1 Brandpilze, pp. 14, 15.
2 Morph. u. Physiol. d. Pilze, p. 125.
3 Brandpilze, p. 15.
4 Ibid., p. 50.
5 Morph. u. Physiol. d. P., p. 127.
6 Pringsheim's Jahrb., 1860, II, p. 308, sqq.
7 Soon after the conclusion of this historical introduction, I received two new works on Ustilaginee
by Rallier, Mycolog. Unters, von E. Hallier, part iv, Entwichlungsgeschichte des Staubbrandes,
Ustilago Carbo; and part vi. Entwg. des Steinbrandes, Till. Caries. Landw. Versuchs-Stat., 1867,
pp. 260-72 and 355-363. According to Hallier, the Ustilago developes from Micrococcus, and can also
be produced from Aspergillus, and probably also from the Eurotium, representing an interposed
Oidinm between both. The Tilletia spore grains [!] either divides into many, and at last changes into
Micrococcus, etc. (as far as can be seen from the text and drawing, it refers here only to the very oily
contents, which appears in the form of a large drop of oil in the Tilletia spores, which can divide
into many smaller, like the others); or, according to the ordinary germination (with a promycelium),
produces finally penicillium; from the latter Tilletia again, etc. I confess, that the "facts" mentioned
by Hallier are presented in a form not so convincing to me as:to compel entering, unwillingly, by a
more extended consideration of them, upon a longer discussion, for which this is not the appropriate
place. I content myself, therefore, with referring the interested reader to the works of Hallier above
cited.




292       NE W YORK STATE A GRICULTJRAL SOCIETY.
threads have been recognized only for a few of the Ustilaginee; but
in the most of them a very advanced condition is known, in which
that threads appear as a gelatinous mass with shiny particles.
T/ie spore formation itself has been more generally investigated;
those of the Tilletia Caries the most extensively. For the Ustilago,
Urocystis and Sorisporium species more complete, correct and comprehensive investigations are yet very desirable.
/hie structure of the spores requires a revision in conf'ormity to the
present state of knowledge, especially a thorough investigation of the
structure of the episporium.
She germination of the Ustilago spores has been established for
Tilletia Caries and Lolii, Ustilago Carbo. destruens, Maydis, antherarum, receptaculorum, longissima, Urocystis occulta and pomplhologodes; though in comparison more accurately investigated in the
first three, in which, also, the sprouting of the sporidia has been
observed.
5The nzanner and 2mode in which the parasites come uipon the supporting plant, is still wrapped in complete darkness. For besides
the inoculation experiments with Tilletia Caries and Ust. Carbo and
destruens, we know at present nothing in relation to all the remaining Ustilagineae, and how many repetitions are needed of Kiihn's
and Hoffmann's inoculation experiments, notwithstanding all the confidence which is due them, will be seen hereafter.
The localization of the parasitefor the purpose of its spore formation, has been most fully investigated. However, as to the occurrence of the parasite in the different parts of the supporting plant, we
have but little knowledge beyond the observations already mentioned.
If I, for my part, notwithstanding the excellent works, especially
of Tulasne, de Bary and Kiilln, did not hesitate to take up again the
laborious investigation of the Ustilagineae, it was only upon the suggestion of Professor de Bary, my honoured instructor, and in the hope
that my own limited observations might contribute to the advance of
the knowledge of the biology and course of developement of these
endophytes.
My observations, which are communicated in the following pages,
I consider by no means complete, particularly as regards the inoculation experiments. They contain also many already anticipated discoveries; in the latter case they can also serve as confirmation, and
lave, therefore, not been excluded.
The most of these observations were made in the Freiburg' botanical laboratory during the summer of last year, where Professor de




FISCHER VON WTALDH1EIM- USTILA GEGI.E.                293
Bary kindly assisted me with advice and material, foI which I take
occasion to express my deepest obligation.  For kind contributions of
Ustilaginem I feel bound also to offer my grateful acknowledgments
to Professor A. Braun and Dr. L. Rabenhorst, as well as Messrs. A.
Millardet and P. Magnus.
II. PERSONAL OBSERVATIONS.
1. i[yceliunm.
Upon looking at the tissue of the supporting plant under new accumulations of different Ustilagineae, as, for instance, Ustilago hypodytes, Sorisporium  Saponarike, etc., it will be found  penetrated
with  threads, which are in immediate connexion with the sporeforming part.  Investigation shows that the spore-forming parts grow
as branches from  these threads; these latter must, therefore, be
regarded as the Miycelium.
The mycelium  is the vegetative part of the parasite.  In  it no
spore formation takes place. It appears well defined in shape, often
ramifying, mostly in double contoured threads (i, 2; ii, 9,1 etc.), often
with watery and not unfrequently globular contents.2
The branching may frequently be ramified and laced into skeins,
especially if they occur in thick walled cells. The direction of the
thread is mostly parallel to the lengthl of the organ, for example,
upright in the stalk; oftener, however, especially when much ramified, winding or bent and irregularly curved.
In the same species the mycelium threads run in and between the
cells of the supporting plant (ii, 2 and others). On the contrary,
the short, often coiled, little branches originate in the cells (ii,
3, 4, 10). The intercellular occurrence of the latter, and their resemblance also in form to the suckers of peronospores, entitle us to regard
them likewise as such in the Ustilagineoe. Their operation is likely
similar to those of the suckers in the phanerogamous parasites. The
oftener these suckers appear, the shorter is the long-extended mycelium
part. The thicker the walls of the cells through which myceliuni
passes, the more plentiful are the suckers.
It is an especial characteristic of the mycelium, that it has strong
walls, which are particularly distinguished by the sometimes very
mlarked double contour of the membrane, and the power of resisting
1 The Roman figures denote the tables, the Arabic the figures.
2 Also in immature parts of the plant, mycelium penetrating them and developing with them.s
characterized in the same way, and presents later the marks cited. It is, therefore, by no means an
old or exhausted part of them which presents these characteristics; but the true mycelium (conste
quently no transition form) is always the same whether young or old.




294.NEwr YORK STA1TE AGRICULTURAL SOCIETI.
3ome -eagents.  A solution of potash produces only a rising of tle
thread;' concentrated sulphuric acid dissolves the thread, thouglh
remarkably slowly.   Treated with iodine and sulphuric acid, or
chloride of zinc and iodine, its membrane does not sh'ow tl;e blue
reaction of cellulose.  The interior, which, as mentioned, is very
watery, and often filled with vacuoles, shows none or only a very
weak protoplasm reaction.
In different species cross walls appear in the mycelium, and in some
there are more numerous vacuoles than in others.  Ust. hypodytes
(ii, 7) possesses numeronus cross walls and vacuoles; and also Tilletia
endophylla de Bary, (in the mycelium of the leaves, but less in those
of the stalk).  This appearance is manifestly in connexion with the
degree of waste of the plastic contents.
Up to the present time I have found the mycelium in the following
species: Tilletia Caries Tul., endophylla  de Bary,  de Baryana
milhi, Ustilago Carbo Tul., Mlaydis Lev., hypodytes Fr., antherarum
Fr., flosculorum  Fr., receptaculorum  Fr., longissima Lev., Candollei
Tul., utriculosa Tul., urceolorun,  Tul., and Sorisporium  Saponarise Rud.
The form  and dimensions of the mycelium  are more unvarying
than the spore-forming threads, but change, not inconsiderably, according to the species.
In Tilletia (T. endophylla and de Baryana) the mycelium  appears
as comparatively long extended threads, which are especially distinguislled by the frequent formation of worm  or coil-formed suckers
(especially in the rhizoma and nodes), as well as by many round or
globular sprouts appearing near one another.  The free mycelium
ends have a blunt, rounded point.  The thickness of the threads
amounts to fiom.002 to.005amm; their length from one branch to
another, or fiom  the suckers to the branch, changes materially; in
some threads it is hardiy.002mm, in others ten times or more.
The Ustilago species possesses mnore variously formed mycelium.
UJst. longissima has many long extended threads with a diameter of
from.003 to.004mm, with  conspicuously deficient and imperfect
suckers.  Ust. AIaydis has a similar mycelium  though with more
numerous suckers.  On the contrary, in Ust. Carbo it is much
thinner (that is in Arrhenatherum  elatius), hardly.002mm thick; in
the same species upon oats it appears imore wavy and twisted, which
1 When the influence of the reagent is not too strong, the supporting tbsune becolnes more transparent, and the thread comes out plainer, and the potash can therefore be employed with advantage
for the clearer observation of an existing myceliumn. After some hours' influence of thinned glycerine,
the mycelium became also more distinctly visible in the preparation.




FISCHER VOAN i TVALDHEIM- USTILA GINVEA.         29t
however depends much upon the nature of the penetrated supporting
tissue.
Generally there appears in the nodes, especially in their thick walled
cells, partially thinner mycelium, very much twisted, and furnished
with suckers, for example, in Ust. hypodytes, in the rhizoma, and in
the nodes of the supporting plant.  The mycelium  of Sorisporium
Saponaria is particularly distinguished by very numerous coiled suckers, especially in the thick walled parenchyma tissue of the root and
the nodes, between the bark and fibrous tissue, among the clusters of
crystals (ii, 10). On the contrary, where the tissue is tender, wider
celled and long extended, as in the internodes, suckers seldom appear,
and the threads are twisted less and thicker, especially in Ust. Mlaydis, longissima and CGarbo.
In the greater number of the ustilagoineoe, the mycelimn is found
most firequently in the cells themselves.  Relativ'ely, Till. endophylla,
Ust. longissima and Maydis appear oftener than other species to have
long extended intercellular mycelium.  So, for instance,. Tilletia
endophylla is often seen in the interior of the cells (especially of the
leaves) with several rows of straight running mycelium threads, boring
through the cell walls in the same direction, and extending througli
many cells (i, 1). UTst. MIaydis also forms long, straight running
mycelium skeins, often going in a long row through many (tflat is,
eight or more) cells, especially in the ovary wall and the woody
parenchyma cells of the rhachis nearer to the surface.
The long, and for the most part straight, mycelium threads hlave a
remarkable appearance in some ustilaginee, especially often in Ust.
Maydis (i, 6), and Sorisporium Saponarioe (ii, 12), less frequently in
Ust. Carbo (ii, 1), which, according to unpublished observations of
(e Bary, appear also, although seldom, in uredos.  The mycelium
threads become covered by a cellulose membrane, which often becomes
very thick, entirely enveloping the thread, rendering it invisible; in
other cases some of its parts or branches are still visible (i, 6 and G6).
This cellulose membrane belongs, without doubt, to the membrane of
the cells of the supporting plant; it represents an immediate continuation of their layers, as I could distinctly perceive in a few cases. It
tppears in layers, and refracts light considerably. By applying iodine
and sulphuric acid, after treatment with potash, it quickly turns a briglht
violet: and afterwards a deep blue. By warming the threads enveloped
in cellulose in a solution of potash, and afterwards applying iodine,
the cellulose sheath becomes transparent, and allows the double con
toured mycelium threads to be distinctly seen in the interior (i, 7).




29i6      NEw YORK STATE A GRICULTURAL SOCIETY.
The threads inclosed in cellulose resemble little straight ridges which
enlarge at their point of contact with the cell wall; they can be
traced through many cells (i, 6). Their thickness varies very much,
and depends chiefly on the circumference of the mycelium. I found
these threads all over in Ust. Mlaydis, where the mycelium generally
occurs in the greatest bulk in the ovary wall, and in the woody
parenchyma of the rhachis.
The mycelium  passes through the supporting plant, at the beginl
ning, probably uninterruptedly, proceeding from the root or rhizonla
to the place of spore formation. In the further growth of the supporting plant, and when the spores are foimed, it is very possible
that the mycelium establishes itself only in the further growing and
developing parts of the new organs; similarly in the roots and
nodes. Nevertheless, there appears to be here an important difference
in the supporting plants, namely, in monocotyledons the mycelium
can be traced through the entire supporting plant; in the dicotyledons
I could only detect it with certainty (in many species of Ustilagineue)
in those organs, or in their immediate neighborhood, where the spore
formation took place; for example, in Ust. Carbo I found the mycelium in the root itself, and from there through the entire axillary part
of the supporting plant, up to the blossom  (ii, 3, 4, 2, 5), and more
abundant in the nodes and the rhachis than in the internodes.  The
mycelium of Ust. Carbo was likewise spread through Hordeum distichum, Triticum  vulgare, and Arrhenathernm  elatius, as well as in
Avena sativa. It showed itself most distinctly all over in growing
specimens of the supporting plant, and in abundant spore formations.
Repeated investigations of the same supporting plant, as well as of
different ones, always gave the same result of universal spreading
of the Ust. Carbo. It is necessary to remark, in order to prevent
misapprehension, that in-the statement of the spreading, those organs
are excepted, through which it is not necessary for the mycelium to
pass in order to get from the bottom of the stalk-, through the tissue
of the supporting plant, to the place of the spore formation. So in
Ust. Carbo the mycelium is found neither in the leaves nor in the
root points, although outgrowths of them, which have no relation to
the spore formation, are unmistakably present, as in cases where the
mycelium also passes over in the awns. In the axillary parts we find
this ustilago only within the fibrous tissues.  In the panicle it
spreads more or less. In the former case it penetrates in all parts of
it, not excepting the glumes (ii, 6); in the latter not only a few of the
blossom parts remained, intact, but generally a few blossoms and




FISCHER vON WALDHEI —      USTILA GINEx3.        297
spikelets. The same happens in the side shoots of the supporting
plant. Commonly, also, the shoots are affected by the ustilago as
well as the principal stem; though not seldom variously modified
exceptions occur. I mention only, that among the side shoots,
some are entirely free fiom  mnycelium, in others of them  it exists,
witl or without succeeding spore formation. There are also cases
known of spore development in the side shoots, exclusive of the
principal stem  (but whether the latter contained mycelium  is not
stated).l
Ust. hypodytes also has a mycelium easily traced. I found it in the
rhizoma nodes with very strong walled coils; in the parenchyma,
between the outside and the fibrous tissue. In the uppermost part
of the lrhizoma, and in the nodes, it is less coiled, nevertheless it is
strong walled, and has numerous cross walls, narrow lumen, and is
fiom.0025 to.0035 mm thick. In the internodes (ii, 7) the mycelium
is found.005 mm in thickness, which even passes into the leaves and
continues to grow further, but mnore sparingly than in proximity to
the spore formation.
In Tilletia de Baryana we have a further example of the spreading
of the mycelium  in the monocotyledons.  Here it pervades the
rhizoma and the nodes (i, 4) and internodes of the stem. For spore
formation it goes into the leaves (ii, 2, 3). The presence of the
Tilletia mycelium in young stalk internodes is easier to be observed
in the neighborhood of the leaf sheath and nodes; here it encircles the
fibrous tissue mostly, even reaching to the parenchyma of the pith.
The myceliumn of this Tilletia more frequently passes between the
cells; that of Tilletia endophylla, on the contrary, chiefly throughl
the cells.
The presence of the Tilletia myceliumn in the rhizoma and nodes
of HIolcus mollZis is simply and irrefutably proved in the following
experiment: I planted in earth rhizomes, as well as different parts of
the stalk, of several specimens of Holcus which were attacked by
Tilletia.. After a few days I not only perceived a mycelium in the
newly developed parts, but also spore forming threads from which
afterwards normal spores came. A like result was given in de Bary's
experiments on the presence and spreading of the mycelium in Sonisporium Saponarioe.
Investigations of the occurrence of the mycelium of Ust. Maydis,
showed a very great spreading of it also. I not only found it in the
ovary wall, which was filled with spores, but also in the woody parenI Hoflmann on the Flugbrand, p. 193.




298        VETW YORK STATE A GRICULTURAL SOCIETY.
chyma of the rhachis, on the outside of the pith tissue, and also some
internodes lower; in the nodes, almost near the base of the Zea Mays
(mostly nearer the surface of the stalk amid the fibrous tissue); it
appeared also with many suckers in the parenchyma of the leaves
covered with brand pustules.
Ustilagos occurring on dicotyledons show a spread mycelium principally near the place of spore formation. In other parts of the supporting plant, for instance in the root and stalk, it can be found only
with the greatest difficulty, because it appears so sparingly at the
time when spore forming threads already exist, or the spore formation has commenced. I have had no opportunity to examine it in
earlier conditions. I refer particularly here to the multitudes of
mycelia I have examined, of Ust. flosculorum, receptaculorum, and
partly of Ust. antherarum. Besides, there are, first, strong, thick,
woody cells of some tissues; for instance, the rhizoma and nodes of
the Knautia arvensis (with  Ust. flosculorum), whose cell walls,
when cut, have a deceiving resemblance to mycelium threads; and
again, the rich protoplasmatic contents, and the not easily removed
chyle in Tragopogon (with Ust. receptaculorum), which greatly
impede the discovery of the myceliunm.
The mycelium makes its appearance in very early conditions of the
attacked plant. It grows with the supportingplant and spreads with
and in it, in order to arrive at the place of spore formation. This
was shown very distinctly in experiments with rhizomes cut from
Hrolcus rmollis, in which mycelium of Tilletia de Baryana existed.
Planted in earth they took root easily and produced shoots, in which,
at the same time, mycelium from the rhizoma made their appearance,
which kept pace with the further growth of the shoots until the spore
formation began in the young leaves.
In regard to the duration of the mycelium, de Bary's observations
of the reappearance of the parasite in the same plant many years in
succession, speak  for its being perennial in perennial supporting
plants. According to de Bary's investigations (verbally communicated), a large plant of Saponaria oficinalis growing entirely isolated
in the Freibnrg botanical gardens, was attacked each summer. for
more than ten years, by Ust. antherarum, in all the anthers. Without
being willing to draw without exception any over-hasty conclusions
from these investigations, I can also mention that I found in the same
manner Ust. flosculorunl  for the second time in the same meadow.
mostly in the fullest luxuriance, where it had also appeared in tilhe
spring; that is after the first mowing of the Knautia arvensis, as soon




FISCHER VON WALD EIMJ — USTILA GIaEE. 2..99
as the new blossoms had developed.  In like manner i observed Ust.
Carbo on the after-growth of Arrhenatherum  elatius, after plants in
the same place, attacked by the parasite in their first growth, hl:ld
been mnowed (I cannot say with certainty whether it was on the salne
specimen of the supporting plant) tlle after-growth not only contained
mycelium  spreading through the entire stock, but also spores in
different stages of development.'
2. Spore Formation.
In those parts of the supporting plant, where the spore formation
takes place, are found the spore-forming threads.  They come imnmediately from the mycelium, as branches, and are distinguished by an
entirely different appearance and action.
The spore-forming threads commonly have a thickness of from.002 to.004mm.  In Ust. antherarum, of Saponaria oflic., nevertheless
they are scarcely.0001mm thick; on the contrary, Ust. Maydis, in the
largest spaces between the cells, attains a thickness of.0012mm.  In
places, where the spore formation has already begun, they are.008mm
and upwards.  Their membrane is distended with gelatine; the contents are sparingly fine grained, lustrous, sometimes plainly oleaginous.
The difference in the lustre depends on the extent to which the thread
has become gelatinous; the more this happens the thread, the more
indistinct become the contents.  The membrane which has become
gelatinous and the granular, or sometimes distinctly oleaginous and
abundant, contents shining through it, distinguish it also in its less
indistinctly marked form fromn the mycelium. This difference is very
marked in Sorisporiumn  Saponariae (iv, 7); the mycelium  thread
changes immediately to the spore-forming, part, and the latter is distinguished at first sight by its extensively gelatinized membrane, and
the granular contents. In all cases, however, the change is not so
abrupt.  In Ust. Carbo, Tilletia  de B3aryana and others, tllere is a
transition form of the mycelium in the spore-forminfg threads, which
is nevertheless always distinguished from  the mycelitum by thle more
delicate, or already somewrllat gelatinized, membrane, and by the more
plastic contents.
Shortly before the beginning of the spore formation, the membrane
3f the thread swells perceptibly, and the lumen consequently becomes
narrower (iii, 20; iv, 2, 8).  On the contrary, where this process has
begun, the lumen enlarges and the diameter of the thread increases
1 De Bary examined Ust. Vaillantii Tul. (in Scilla bifolia L.) two years in succession in the same
place, and found at the bottom of the bulb (lccus). in the stalk and blossoms of the supporting
plant a mycelium similar to that of Sorisporium Saponarie (from unpublished investigations).




300       NEW  YORK STATE AGRICULTURAL SOCIETY.
to.008rm, and even more(iii,30-32, 40; iv, 5). This feature is shown
very distinctly in Ustilago and Sorisporium; in Tilletia neither such
an extensive swelling of the membrane, nor such a contraction of the
lumen takes place (iii, 1, 6, 9).
The threads are found in the interior of the supporting tissue, or on
its exterior surface - everywhere where spore formation takes place.
Thus, in Till. endophylla, de Baryana, Ust. longissima, and partly also
in Ust. Maydisand Urocystes pompholygodes, in the parenchymna of
the leaves, between the fibrous tissue; in Till. Caries, Ust. fiosculorum.
receptaculorum, antherarum, Carbo, ilaydis, urceolorum and Sorisporium Saponarioe, in different part of the blossoms; in Ust. hypodytes on the outer surface of the internodes; in Ust. MIaydis likewise
in the surface tissue of the stalk, and of the male panicle stems, etc.
In some ustilagos the threads occur in very great masses, especially
in Ust. urceolorum and Sorisporium Saponariae.
The application of reagents indicates nitrogenized substances in the
contents of the spore forming threads. Iodine colors them  a deep
yellow; chloride of zinc and iodine a brownish yellow. The membrane remains colorless. In water it swells somewhat, resists weak
sulphuric acid, and dissolves only through longo influence of concentrated acid. Numerous oil drops are only occasionally found in the
contents-the oftenest in Ust. Maydis-especially in the wall of the
young ovary of the Zea of nearly 1 centimetre in length (iii, 19). The
oil appears the plainest at the beginning of the spore formation, and in
that part of the thread where the spore formation will occur next (iii,
20), consequently it serves as a material for the spore formation.
The threads just described form the spores in different ways: either
singly on their ends (Tilletia), or they break up into spores themselves
(Ustilago); besides, the spore formation of Sorisporium and Urocystis
occurs in a peculiar form, which will be further mentioned later.
The spore forming threads of Tilletia appear in Till. Caries (in the
ovarium tissue of wheat) as principal stems (of nearly.002 mm thick),
and secondary stems. The latter are thinner, less ramifying, and the
spores form on their ends first. The spore formation occurs in succession, and begins by slender pear-shaped branches of.0015 mm thick,
appearing on the side of the principal stem (iii, 1). These outgrowths
extend themselves in length from the bottom. The pear-shaped part
increases in circumference and becomes more round, by which the
under end becomes a little stem, or plainly marked branch, of about.001 mm thick (iii, 1, 3 underside at the right). The latter, at this
time, is firom.003 mm to.0035 mm long, and fiom.0025 mm to.003 mm




FISCHER  -ON WALDHEIi- UST'ILA4 GINE.               301
thlick.  The contents coming out of the principal stem shine through
the somewhat gelatinized membrane of the little branch. The upper
part shows itself distinctly as a young spore in form of a little pustule,
which is transparent, roundish, and filled with bright granular contents.  Shortly before the completion of the growth this obtains a
double contour,   indicating the formation of the episporium  (iii,
3).  By this time the little branch is so exhausted by the spore formation that it commonly appears as a delicate, thin, little winding
thread of from.0005 mm to.001 mm in diameter, containing vacuoles
(iii, 5).  Later. the episporiumn becomes gradually darker and uneven,
and the contents become especially rich in oil.
Tilletia endophylla shows a very similar process of spore formation,
and needs, therefore, no fuirther description.
Vacuoles-are seen in young spores of both species of Tilletia while
still transparent, which through the influence of water become perceptibly larger and more numerous (iii, 11). On many of the completely formed spores are seen the remains of the thread.
The spore formation of the Tilletia de Baryana shows a deviation
worthy of mention.  On the one hand it forms its spores at the end
of the threads, like Till. Caries and endophylla; but on the other
these threads are larger in circumference and their gelatinized membrane envelopes the spores until their maturity.  The free end of the
thread which has already changed into a spore, is, at the beginning,
compressed to a beak shape (iii, 14), but later becomes round.  The
gelatinous membrane attains a considerable thickness; but, however,
as soon as the spore contents in it separate from it by the firm contour (iii, 15, 16), it begins to disappear more and more until at full
maturity of the spores there is no trace of it remaining.'
Ustilago shows a different process of spore formation. The sporeforming threads increase in circumference; their lumen contracts and
appears filled with the contents, as a shiny narrow stripe (iii, 20; iv,
1, 2); they form numerous shorter or longer branches (iii, 20) and
run closer together. The threads swell at several places, more or
less near together (iii, 30, 67), in succession, so that the spores may
often be seen in all stages of developement at the same time (iii,
32, 36, 39).  These roundish or oval swellings occur in connexion
with a considerable gelatinizing of the membrane, and an enlargement
at the same time of the lumen, which, filled with the contents,
1 In Till. de Baryana the beginning of spores are found also in the form of small, clear, transparent blisters, with homogeneous watery contents; but on account of their small size and number
they may be considered as abnormal conditions.




302       NrEW YORK STATE AGRICULTURAL SOCIETY.
mostly appears somewhat pear-shaped or more elongated (iii, 41, 44;
iv, 3, 4).
As the result of the local swellings, the contents become broken
up into a corresponding number of parts or little lumps(iii, 30, 32,
37). At the same time the contour of the entire thread becomes
undulating, and by reason of many lateral and terminal sprouts
generally gets a tangled irregular appearance (iii, 36, 37, 39, 40.)
Often a swelling on the thread is scarcely apparent, when the contents
are already distinctly divided into several parts (iii, 20; iv, 3).
The occurrence of a very large number of spore-forming threads,
together with their branches, causes such crowding, that the gelatinous
membranes pressing against one another, glue together, and in the
further development their contours almost entirely disappear (iv, 5).
Under these circumstances the entire bulk of complex threads, intertwisted and glued together, have the appearance of a gelatinous mass,
in which are perceived many clear, shiny parts of the contents of the
threads, shining through like little short stripes, little lumps, etc. (iii,42).
The individual parts of the thread soon begin to differentiate (iii, 31,
33). The swellings become rounder and their contents likewise take
a rounder form.  There follow  multitudinous sproutings of the
branches, as well as a further separation of the contents of the
swellings themselves (iii, 34, 42). Around each of such parts of the
contents, the gelatinous membrane becomes larger in circumference.
Notwithstanding this irregular production of the spores, however,
some threads generally show a spore formation downwards from the
point (iii, 31, 39). At this time the circumference of the thread varies
greatly; at some places it is from.003 to.004mm thick, at others,
where there are larger swellings, from.006 to.008mm. However the
thickness and greatness of the threads, together with their spore
beginnings, depends upon their species; Ust. longissima, hypodytes,
Carbo and flosculorum, show, for instance, smaller dimensions than
Ulst. receptaculortln, Maydis and urceolorum.
During the later process of the spore formation, the entire mass of
the threads always separates more distinctly into single parts, which,
however, remain together until the completion of the spores, and,
according to their developement, show a more or less thick gelatinous membrane surrounding the clear, shining lumps of the contents.
The membrane, together with these lumps, has angular sides throughout, where they were contracted through pressure against the adjacent
parts during their formation iii, 34:; iv, 6).
The contour of the gelatinous membrane becomes more distinct on




FISCHER VON WA lDHEf I~ — USTILA GIiEAE.          303
the exposed side, as well as that of the contents(iii, 23, 45). At the
same time the beginnings of spores are seen completely separated
froml one another.  The interior enlarges and contains a multitude
of oil drops;  the  gelatinous  membrane  shlrinks  perceptibly.
Around the contents a distinct contour is perceived; it corresponds
with the outer spore membrane, which assumes a deeper colour.  The
irregularities of the episporium  are formed while still surrounded
with a thin gelatinous layer(iii, 35, 46).  At the same time the entire
mass of spores become perceptibly larger in circumference, and the
developed spores separating, take their typical, mostly spheroidal
form. Ripe spores show no trace of the gelatinous membrane, or
any remains of threads attached to them.
Sorisporium  Saponarike is distinguished by an entirely different
spore formation, which, however, by close observation, shows some
resemblance to Ustilago.
Extraordinarily abundant mycelium  (with threads from.002 to.004mm thick,. at the time of the spore formation),penetrates all parts
of the blossom. In the immediate neighborhood of the free surfaces cr
the blossom, not excepting the interior of the ovary, the myceliui.
changes directly into spore-forming threads (iv, 7), which like thoe
of Ustilago, appear gelatinous, from.004 to.007mm thick, and fillel
with shining contents.  Running on the free surface, these threads
have a feature different from  Ustilago, that the friee ends curv,
and bend inwards, as if to roll up the point (iv, 10). At the same
time branches appear at different places on the threads, which alao
roll up (iv, 8, 9, 11). The threads, with one or several such branches,
incline distinctly toward one another (iv, 9), and twist themselves
into a little ball. The membranes of the thread become much more
gelatinous; there appear in them  hardlyperceptible divisions. At
the same time other threads appear on the ball, encircle it, and twist
themselves into one another, or with the branches of the earlier ones.
The whole twisting appears not unlike the formation of the lichen
thallus. The gelatinous condition of the membrane, and the great
confusion of the threads, cause here also an almost entire disappearance of the contours, which finally are only discerned in a few ca'ses,
or at some places on the ball. But it is often possible to trace the
lumina filled with shining protoplasm, within the apparently homogeneous gelatinous substance, the contents appearing here also broken
into little strips or lumps (iv, 12, 15).
The spore formation in the ball is limited solely to the middle. At
the beginnning, where the spores are indistinctly separated from one




304       NEW  Y7ORK STATE AGRICULTURAL SOCIETY.
another, it is difficult to get an accurate view of them. Then, clear
spots better defined, the beginnings of spores, are usually to be
descerned in the interior of the ball. Soon after they separate one
from  another, showing a distinct contour; they assume a faint light
brown color, and appear in one plane of view as from four to sixteen
more parts or cells (iv, 12 to 14). Seemingly these spores which
now have begun to show themselves, and in fact by repeatedly separating form the real spores, may frequently reach sixty or even over
one hundred in one ball (iv, 22). Such spore beginnings, which show
angular formations in every place where there is pressure, are already
contained in the spore balls of.05 mm in their greatest diameter.
In the further development of the spores, the ball becomes larger
in circumference, the gelatinous threads swell greatly, surrounding
the young spores like a gelatinous zone (iv, 15, 16). The contour
of the threads can still only be partly perceived near the spore (iv,
15). At a certain stage of the developement, as soon as the spores
take a more intense brown color, the gelatinous zone has begun to be
absorbed (iv, 15, 16, 21) by being applied to the developement of the
spores. Spore balls of.07 mm in diameter have a gelatinous zone
of only from.004mm to.006 mm thick. In ripe spores it entirely
disappears (iv, 22).
The spore ball masses show the same peculiarity in regard to
the course of their developement as the single thread spores of
the ustilagos. For if there exist compact layers, mostly six to
eight, of spore balls on the exterior surface of the supporting
plant, then the outermost row is composed of the most developed
spore balls; those which follow inwardly are made up  of less
developed spore balls (iv, 20). Probably the first formed layer of
spore balls is pressed out by those following, and so on, and can then
either develope further independently, or, what is more probable, by
many spore-forming threads which have become very much lengthened,
remain in connexion with the supporting plant.
Besides the spore balls described, there is found a second form of
Sorisporium Saponariae spores, solitary. Their origin is similar to
Tilletia, inasmuch as one spore only forms on single threads. These
are strong gelatinized threads of faint lustre (iv, 16), which are
observed as branches, or perhaps only the pressed out ends of threads
wound round the ball. Or are they threads, which appear in the
immediate neighborhood of the ball, coming firom the twisting of the
threads generally forlning spores? I have not been able to ascertain
by direct observation.




FISCHER VON tVALDHEDI-M- USTILAGINEE.            305
The solitary spores surrounded by the gelatinous membrane of the
thread are developed similarly to the Tilletia de Baryana. At maturity this membrane disappears entirely, or, although seldom, coveis
the spore as a very thin layer, hanging together with the remainder
of the spore-forming thread. The solitalry spores have a roundish
form; they appear seldom, and almost always only singly on the
b)all. Application of iodine, or chloride of zinc and iodine, renders
them quite distinct.
Here may be mentioned the oval little bodies of.006mm long and.004:mm thick, which are very often fastened to the Sorisporium balls
by a small stem generally.006mm long and.001mm thick. In experimental germination these are found to be spores of Pleos.pora herbarum Rabh.
The spore formation of Urocystis8 (about which there is mention of
one or two cases in the historical introduction) I have had no opportunitv to examine.
The application of reagents shows nitrogenous substances in the
contents of the young spores, the same as in the spore-forming threads.
Iodine colors it yellow; chloride of zinc and iodine, brownish yellow.
The gelatinous membrane dissolves in sulpllhuric acid, and is not colored by the above reagents.  On the contrary, a solution of potash
colors it yellow. In water it expands more in the younger conditions
than latev-. Alcohol causes a great shrinkage not only of the sporeforming threads, but also of the almost completely formed spores.
The cited reagents remain the same, without any remarkable distinction, in the different stages of the development of the spore. Sulphnric acid only shows a great quantity of oil drops in the developed
spores whose gelatinous membrane has ahlmost disappeared, which
then shine through the episporium, or by the bursting of it come out.
3. Place of ta1  Spore Formation and Developement of tAe Spore
Jlasses.
In comparing the different conditions of the spore formation in connexion with the supporting plant, many variations appear.
Tilletia Caries chooses for the spore formation, the ovaries of some
of the graminaceae, especially of wheat (Triticum vulgare). The
beginning of the spore formation can be discovered from  exterior
indications; the ears (of wheat) appear of a darker green, the spikelets stand somewhat away from the stem. If this spikelet is examined,
the ovary appears rather swollen, though still oval, by which means
it can be distinguished from the round swelling one, infested by the
[AG.1  20




*306      NE IV IYORK STATE XARICULTURAL SOCIETYI
Anguillula Tritici Ktihn. In the ovary only, a light grey  pappy
mass is found. It consists of spore-forming threads and many young
spores, which are mnixed with more developed ones, provided with the
scarcely coloured episporium.  At this time, a distinct myceliurm pervades the wall of the germ, which later, can be followed to, and in
the stein.  Generally, the shrunken anthers rest on the ovary, in which
appear a few complete, but withered grains of pollen.
The presence of the parasite is marked more conspicuously in the
further perfection of the spores; the growth of the affected ear, is
slower than the normal ones; its colour is still green, when the others
have become yellow; the spikelets appear sparingly, and the entire
plant has a stunted growth.  In the ovary the spore mass is of a
greyish black color, of less moist consistency, and has a characteristic
odour. Among the clear brown ripe spores, many are found that are
limpid, although already of from.016 to.02 mm in diameter. Entirely
ripe spores appear as a brownish black powder.  They often burst the
ovary..The ear turns yellow.
The Tilletia endophylla forms its spores in the leaf parenchymna of
Brachypodium pinnatum.  The youngest conditions are best found in
the side shoots of an affected stock, whose leaves are still rolled up.
The innermost of them, contain the youngest stages, which are
indicated on the outside of the leaf, by rather pale yellowish or brownish
stripes. A long incision of tender leaf shows spore-forming threads
plainly, particularly between the cells, with oil drops in the contents.
Spores are here also to be seen, later, as clear pustules; they are distinctly separated from one another, and not piled together, as in ustilagos. Later the spore beds appear as reddish-brownish, much lengthened stripes, more developed, and mostly containing light colored spores.
Here the spore formiation also takes place from the exterior, towards
the interior of the tissue.  At last the epidermis bursts, and in the
openings is seen a brownish blackl powder, which consists only of ripe
spores. The spore masses possess the same disagreeable odour, as
Tilletia Caries.
The spores of tle Tilletia de Baryana are found in the parenchyman
of the leaves, and in the sheath of the leaves of holcus mollis (velvet
grass). The spore masses appear, according to their ripeness, as more
or less long narrow stripes of a brownish-black colour, often extending
from the sheath of the leaf, to the point between the leaf-nerves,
mostly covered only by the epidermis. At  maturity, the surrounding tissue of the sulpportitngo plant, appears to witller and be.omnes
l)rownisi!-yellow or red; it is busteld open by the spore nmasses, upon




FISCHER  VON WALDHEIM- USTILAGINE-.                    307
the upper and under side of the leaf and leaf sheath. The edges of
the leaves retain their green colour the longest. Exterior appearances
correspond with the different stages of the spore formation, as in thlo
Tilletias already mentioned. The spore masses possess no odour.1
The spore formation of the Ust. flosculorum  occurs only in the
anthers of the Knautia arvensis.   The beginnings of the spores
are found already in the youngest blossoms which admit of a close
examination, when scarcely the size of a pin head. The space in the
anther-chamber is already filled with the gelatinous spore-forming
thread texture, sometimes mixed with the pollen mother cells and
developed pollen grains.  Abundant mycelium threads penetrate the
wall of the anther, and are evidently in connexion with the sporeformingt texture; I have found none in the stalk.
In the interior of the closed blossom the anthers are found sprung
up with normal splits in them, and covered with a whitish mass,
which generally is formed of half-ripe spores, still surrounded with
the gelatinous membrane and very young spores. In the full grown
blossom  all the spores of the parasite are ripe.  The parts of the
blossom, with the exception of the already withered anthers have, on
the whole, a structure differing scarcely from  the normal one, the
ovule especially, some of the pollen grains even coming to a
regular developement.  The blossom  remains closed for a very surprisingly long time, and the points of the corolla segments become
often early faded and withered..The entire colour of the blossom
crown is changed by the plentiful sprinkling of the violet-coloured
spore powder; the head has a bright violet tinge.2
Uat. antherarum in like manner forms its spores in the anthers, and
only in the different species of Silenese. Mycelium and spore-forming
threads are often found in very young anthers; Saponaria blossoms of
only 1.-2 millim. long, with anthers S — of a millim. long, contained
already a gelatinous mass of indiscriminate spores and spore-forming
threads; at the same time, pollen, which afterwards disappeared. Blossoms of from 3-4 millim. long already appear somewhat swelled, and
in them is seen the characteristic violet-coloured spore mass coining
from the opened anthers. It consists mostly of ripe spores, furnished
with a coloured episporium, with which are mixed many younger
gelatinous spores. This causes the'entire mass to have a soft moist'
1 This new Tilletia I found last summer, in the greatest luxuriancy, in the ditch on the right of the
highway near Freiburg (Baden), in the direction from St. Loretto, towards Guntersthal. Prof. de Bary
and myself, found isolated species of the attacked Holcus, also, on the south side of the Schlossberg.
The spores ripen in Jnne. See Rabenhorst, Fungi cur. exsicc. Cent. x, No. 1073.
2 Compare Fischer v. Waldheim, contribution to the knowledge of the Ustilago. Botan. Zeft. v.
Mohl. and de Bary, 1867, p. 39:3, sq.




308       NEIW YTORK STATE AGRICULTURAL SOCIETY.
appearance. Later the blossom opens, the bursted anthers extend on
the stalks over the flower leaves, covered with a violet spore
powder. Spores coming from here cover here and there, as little
powdered spots, single parts of the blossom.
Ust. antherarum  shows the same characteristics in Lychnis Flos
cuculi, diurna, Dianthus Carthusianorum and Silene inflata. Only
tile proportionate size of the blossoms in which a certain stage of
spore development is met With, is evidently different in comparison
with Saponaria officinalis.
Ust. receptaculorum developes its spores on the outer surface of the
receptacle and of the blossoms of Tragopogon pratensis, porrifolius,
and Scorzonera humnilis.  My observations refer to Tragopogon
pratensis.
The presence of the parasite can be detected also here by the
swelled up appearance of the young blossom yet enclosed in the calyx.
If such an one of from 4-5 millim. in diameter is opened, all of the
receptacles as well as the blossom appear covered with a bright rosy
violet, doughy layer. There are already, at this time, spores with
small perfected episporium, however they almost all have yet a gelatinous membrane. A cut, vertical to the receptacle of the flower,
shows how the spore formation proceeds from the exterior toward
the interior, the  ripest conditions lying nearest the exterior,
while down toward the upper surface of the receptaculum the spores
are by degrees less developed. Later, when the blossom buds have
become not quite double in size, the calyx still remains closed.
Receptaculum and blossom appear then covered with a dark violet
powder. A horizontal cut through the first shows, below the almost
perfect spores, others whose gelatinous membrane has' already begun
to be absorbed.  At last the still green and strong calyx leaves
separate, but retain their former position, in which they bend together.
Through the openings comes a rich, dark, brownish violet powder of
ripened spores.
4st. urceolorum forms its spores exclusively on the upper surface
of the ovary of different species of Carex. I found them alive while
examining the Carex pilulifera. The mycelium of the fungus is found
spread through the entire ovaryv, as well as in the ovary wall, and in
the tissue of the shrunken ovule, as also in the hollow spaces, which
are caused by the shrinkage of the ovule, between its upper surface
and the ovary wall, and between the kernel and integuments. On the
contrary, I did not find the mycelium in the pericarp surrounding the
ovary.




FISCHER VON UALDHDEIr — USTILA GINES.            309
The spore-forming threads, pass ons the outer upper surface, for the
purpose of spore formation, and are here developed into a thick layer,
overspreading the ovary wall, in which the spore formation begins,
on the outside, and proceeds inwardly towards the ovary wall. Here
are found still early stages, while the outer surface consists of dark,
completed spores. Such a spore bed, when cut horizontally, commonly shows two layers of spores, distinctly separated firom  one
another, one dark and glossy, which, with the exception of the fastening place of the stem, uninterruptedly surrounds the ovary. The
former consists of almost ripe spores, and is from.48 to.56 mm thick;
and the latter, contiguous to the ovary, of gelatinous spores, is from.04
to.06 mm thick. With the perfection of the spore-forming fungus bed,
the volume of the body inclosed in the urceolus is evidently increased
even to double the normal size of the carex fruit. The skin itself
bursts longitudinally, and the fruit, covered with the fungus, appears
as a dusty ball.
In one and the same ear, either all the seed may be affected with
the parasite, or only a few, often even only one, while the remainder
appear normal and sound.
Ust. Cacrbo spore formation takes place exclusively in the blossoms
of some Graminace.e.  This parasite usually attacks all the organs of
the spike, although cases of a more partial infection are not infrequent.
Oat blossoms of 11-2 millim. long (the covering-leaves being
omitted) already contained mycelium. On the outer surface, or generally on the upper surface of some spikelets, about, cent. long, appear
roundish, encircled, gelatinous masses. These latter, contiguous to the
supporting tissue, cause small holes in it, and consist of spore-forming
threads run  together, with spore beginnings.  Larger blossoms,
approaching 1 centimetre in length, contained brownish-coloured
spores, which, as a brownish-black powder, covered a few organs;
besides spores still with their gelatinous membrane. The undermost
spikelets, nearest the stem, are the least developed, and contain
younger spore conditions, than the upper ones.
In the further development of the spores, the involucre is destroyed,
and the brown-black powdery spore mass is spread over the naked
blossom organs. These conditions are already found in panicles, which
are still completely enclosed by the sheath of the leaf.
Hoffmann mentions a fine mycelium of Ust. Carbo of barley and
oats, which in some cases spins around the still wrapped up ear like a
spider-web, and binds it to the sheath of the leaf.' I have often had
I On Dust Brand, p. 200.




310       NEW  YORK STATE A GRICULTURAL SOCIETY.
opportunities to examine similar conditions of barley and oat brand,
but in all my cases the kind of cobweb mycelium proved to be
entirely heterogeneous, not having the least generic connexion with
Ust. Carbo. It originated mostly from luxuriantly developed Aero&
talagmus or Verticillium. I never found mycelium of Ust. Carbo ofn
the young spore beds.
Ust. Carbo shows similar relations during the spore formation in
other supporting plants; a description may therefore be omitted.
The commonest place of the spore formation of Ust. BMaydis is the
interior of the ovary of Zea iMays. The ovary appears then of an
abnormal, spongy form, also with a length of 1 centimetre. In the
interior of its wall, but nearer the outer surface, are found separated pale
brownish spots, formed of gelatinous spore masses. Those spore-forming threads which are not run together, appear of from.003mm to.007mm thick, sometimes even up to.012mm, and have rich oleaginous
contents. They pass mostly between the cells, but often penetrate
them, themselves or with their shoots. At the same time there
come not unfrequently, long, somewhat gelatinous-appearing, isolated and septate threads on places where spore formation can take
place. It remains to be decided whether it is not a certain transition
state of the spore-forming threads (iii, 17).
A cut along the ovary shows the single spore formation mass as
dirty brown spots (iii, 24), which, by the further developement of the
spores, increase in numbers, become larger and darker colored. The
texture of such an ovary already becomes abnormally large, appears
very loose, spongy and porous. The latter itself is of a changed
form, swelled up and hanging over towards the outside. Such conditions are found on an ovary of about two inches long and half an inch
broad. They are by no means confined to a definite size of it, for the
appearance, number, etc., of the spore-formation masses depend on
the abundance in which the latter appear and the degree of their
developement.
The peculiar manner of the spore formation of Ust. Maydis explains
the origin of isolated spore masses. The spore-forming threads form
very numerous iamifications (influenced perhaps by the large tender
cells of the Zea tissue), which, however, develope in a more tufted
form on certain places of the thread. In each tuft, or in the ramifications arising fiom it, begins a spore-formation  process isolated from
the rest. So originate the isolated spore masses; the spores ripen
An them in a centripetal direction. In consequence of the large-celled
supportin, tissue, the single spores suffer a slight mutual pressure,




FISCnER  VO  T WALDHEIM- US'ILAGINER.            311
and on that account appear of a spheroidal form, and are hardly distinguishable from one another (a somewhat advanced stage is shown
in iii, 21, 22).
As the ramifications become more iunieq'ous, and later the incipient
principal threads separate into spores, the spore-formation nlasses
increase extraordinarily, and, luxuriantly developing, entirely fill
the texture of the ovary which more and more diminishes. Only
tlhe cells of the outer wall remain complete. The developed spores
appear like a greasy, brownish black mass, which  breaks tllroughl
the ovary when ripe.
As to the ovary itself, it should be mentioned that at the beginning
of the spore formation the ovule becomes hypertrophied and appears
filled with spore-forming threads. Later, it is destroyed, the same as
the rest of the texture.
Ust. Maydis forms its spores also in other parts of the supporting
plant; for instance, often in the internodes of the male and female
axes; more seldom  in the surface tissue of the nodes (where I have
observed brand protuberances as large as a lien's egg), at the bottom
of the leaves, and in the texture of the leaf laminse. The attacked
places, under these circumstances, likewise appear hypertrophied and
swelled  up, filled  with a multitude of spores and growing to a
large size.
The spores of Ust. Ebypodytes occur on the outer surface of the
main stem (except the nodes); also on the inner surface of the leafsheathll, and the under parts of the leaves, especially on Elymus arenarius. The attacked plants may be distinguished by the following:
Backward growth, the stem remaining unusually long rolled up in
the leaves, which are shorter and blunter than usual, and appear
swelled in consequence of the developing spores between them and
the stem. In the specimens, I have examined, the ears did not
de velope.
If the younger conditions of the spores are investigated, a horizontal cut, in an affected internode, shows the mycelium in the cells
lying nearest the outer suface (iii. 26). Its threads are distinguished
by very many cross walls, and by their undulating course, especially
in the inner wall of the cells. A  transition form, in the sporeforming threads of a slightly gelatinous nature, is found in the two
outer rows of the stem cells. When these threads have penetrated
the free surface of the stem, they become much more gelatinous,
although only measuring.002 mm, or, on places where the spore formation begins,.004mm thick (iii. 25). In consequence of such a small




312       NEyW  YORK STATE A GRICULTURIIAL SOCIETEY
diameter of the intertwisted threads, the lumen can scarcely be perceived. The spore formation of this Ustilago also-proceeds from the
exterior towards the interior —towards the surface of the stein and
leaves. As soon as the otter layer is composed of ripe spores, and
amounts to a thickness of.2m, the later layer lying nearest the sirface of the supporting plant contains mostly unripe spores, clear as
water, and measures.04mm in thickness.
Ripe spore beds surround the internodes, generally excepting the
two lowest ones, as a compact, greenish-brown, pulverized covering.
and, from the bottom to the top, where the youngest stages are found.
Thus, for instance, eleven internodes, counted from the top, are
covered with spores, and only the two lowest ones remain free.
Ust. longissimca forms its spores in the parenchyma of the leaf ofl
some Glycerias. The youngest stages are seen in the form of clear
pale yellow spots or stripes on the not yet unrolled leaf. A proper
cut, at such places in the texture of the leaf, between the woody tissue, shows a rich texture of spore-forming threads.  They pass
through the cells in several rows; the principal threads are from.006
to.01mm thick, the branches are from.003 to.004mm thick. The
latter appear in this stage mostly as already run together, and formini
a gelatinous mass, with little lumps of the contents shining through.
The spots and stripes enlarge during the further developement of the
spores, chiefly in length. At the same time they become dlarker
coloured. Stch conditions are seen on leaves of about four or five
inches in length, already out of the sheath. Here the most of the
spore beginnings are plainly different, surrounded with a thin gelatinous membrane, and with yellow shining lumpy contents (iii, 27, 28).
The interior of the spore masses contains the youngest conditions.
When the spores become ripe, the epidermis of both surfiaces of the
leaf bursts at the stripes, and the spores fill up the opening as a dirty
greenish-brown powder.
So8ispo~riun Saponarice spore balls originate on all parts of the
free surfaces of the blossom inclosed by the calyx of Saponaria officinalis (sometimes also on the inner surface of the bottom of the calyx).
Blossoms of one millimetre in diameter, as soon as they contain the parasite, appear somewhat swelled at the bottom. In them is found merely
an abundant gelatinous skein of spore forming threads, which pass
along in the immediate neighborhood of the free surfaces of the several
parts, and with numerous ends come to the surface itself.  Myceliumn
also occurs there. Blossoms of two millimetres in diameter show very
many formed spore balls in various stages of developement, and young




FIscHER voN WALDHEII —   USTILAGINE-E.           313
solitary spores, but only seldom, as is the case also later. The Soris
porium  characteristics of the blossoms at this time are roundish
inflated forms, and closed calyx. The blossoms, already filled with
the ripe spores, retain this exterior form; they have a globularpointed appearance; their greatest diameter is from  six to nine
millimetres. At the middle, more seldom they are oval or pear shaped,
the greatest diameter being ten millimetres at the base. The calyx
leaves are mostly pale green, at the point a brownish violet; the
flower leaves thickened and short. The threads attain a thickness of
one millimetre. The anthers appear white, and with pollen grains;
the ovary thickened (to three millim. in diameter), and with an undeveloped style. All these parts, except the outer surface of the calyx,
are covered with a brownish rust-coloured spore powder. The spores
also fill the inner space of the ovary. Each ripe spore ball appears
to the naked eye like an atom.
When the spore mass becomes ripe, the calyx becomes irregularly
torn open at the top and sides. It also appears pale green, partly
dark violet (especially at the point), or often brownish, with a bright
surface and dry skinned. The attacked saponaria scarcely reaches
half the height of the sound one, and often has a decumbent stem.
The foregoing observations on the way and manner of the spore
formation in reference to the supporting plant allow some general
results to be briefly stated.
1. The spore formation takes place only in definitely fixed parts of
the supporting plant. I refer especially to Tilletia Caries, endophylla
de Baryana, Ust. hypodytes, longissima, Carbo and Sorisporium
Saponarioe.  Ust. MIaydis is an exception, forming its spores on the
different parts above the ground, even on the same individual, supporting plant. Probably Urocystis pomnpholygodes should be included,
as its spores originate in different parts of the stem and leaf of Itelleborns, Ranunculus and Anemone.
2. The spore formation begins in the youngest conditions of the
supporting plant. As example, I mention Ust. flosculorum, Carbo,
Sorisporium Saponarike, etc. But the spore-forming threads appear
already at the time when the developement of the part of the supporting plant relating to them first begins (for example, Ust. Carbo).
3. With few casual exceptions the parasite is spore forming in all
its parts which allow spore formation, or, to express it more distinctly,
ill those parts of the supporting stock which are destined for it, as
well in those cases where the remnants of the mycelium may be found,
as where it cannot -be found. In order to illustrate these conclusions.




314        NEW  YORK STATE AGRICULTURAL SOCIETY.
I briitg forward the following facts. The spores of Tilletia Caries
form,  according to their individual capacity, in the ovary of wheat.
Generally all the sprouts of the attacked stock contained the parasite,
and all the ovaries of a spike its spores. The same happened in Ust.
Carbo, and all other Ustilagineae, so far as their spore formation is
known.  In some of themn (for instance, Ust. Carbo and Till. de
Baryana) I succeeded in following the mycelium  through the entire
supporting plant, from the base to the place of spore formation, and
thereby showing that the mycelium grows radiating from the root or
root stock of the supporting plant, through all the sprouts toward the
places of spore formation, and may be discovered here when the
spore formation takes place.  In other cases, as in Ust. flosculorum,
receptaculorum, etc., this was not possible; in these species, and only
seldom, there were scarcely perceivable remnants of the mycelium at
places not immediately bordering the place of the spore-forming
masses.  ZNotwithstanding which, the spore formation took place
here, as before, not only in all the proper places but in all in which it
was possible.  Exceptions often occurred; they were, however, caused
by insufficient and delayed developement of the parasite, or by many
otherl yet uninvestigated circumstances.
4. Not only those parts of the supporting plant, above the ground,
contain mycelium, but also those under it; the spore formation, however, takes place only in parts above ground, both in the interior and
on the exterior of the texture.  Only two species are exceptions; one
of them completes its entire life cycle under the earth (Ust. hypogaea,
Tul., in the root of Linaria spuria, Mill.'), the other, under water (Ust.
marina, Dur., in the texture of Scirpus parvulus, Rom. et Schult2).
5. The spore-forming masses of all Ustilagineoe have a defined succession of spore formation. As in the single thread of a Ustilago,
so also in its entire texture, a centripetal process of formation is distinctly recognized. The spores on the exterior are the ripest; those
on the interior are the least so. This is plainest in Ustilago; not less
so in Tilletia (only in relation to the entire spore masses); likewise in
Sorisporium, only here spore balls represent the single spores.
1 lmasne, Fungi hypogaei, 1862, p. 196.
2 ultasne, Super Friesiano Taphrinarum gencre. Ann. d. Sc. Natur. V. ser., t..v, 1867, p. 133-136.




FISCHER VON WALDHHEI — USTILAGINEx.               315
S YNOPSIS OF THE USTILAGINEAE IN RELATION TO
THEIR SUPPORTING PLANTS AND THE PLACE OF
THEIR SPORE FORMATION.1
[Those personally observed are marked *.]
1. USTILAGO HYPODYTES, FR.
SUPPORTING PLANTS.           PLACE OF THE SPORE FORMATION.
Agropyrum repens, P. B.           *Exterior surface of the main
*Elymus arenarius, L.               stem and sheaths of the leaves.
Glyceria fluitans, R. Br.
*Lygeum sp. (Ust. Lygei, Rabh.=
Ust. hypodytes).
Panicum repens, L.
Phragmites cominunis, Trin.
Triticum vulgare, (?) Vill.
2. UST. LONGISSIMA, LEV. (Ust. fusco-virens, Ces.)
Glyceria aquatica, Prsl.         *In the lamina of the leaf, next
"C   **fluitans, R. Br.         the outer surface.
4C   *spectabilis, M. K.
2a. UST. LONGISSIMA VAR. MEGALOSPORA, Riess.
Dactylis sp. Riess; with spores  *In the leaf parenchyma.
*Poa sp.       of Tf"'
3. UST. CARBO, TUL.
Aira caespitosa, L.               *On the free surface of some parts
Andropogon hirtus, L.               of the blossoms (excepting the
*Arrhenatherum elatius, P. B.       glumes).
Avena flavescens, L.
"  pubescens, L.
" *sativa, L.
Brachypodium ciliatum, P. B.
Cynodon Dactylon, L.
Festuca pratensis, Huds.
Hordeum distichum, L.
cc"   murinum, L.
"    *vulgare, L.
1 From personal observations and materials contained in the Herbariums of De Bary, Fwuckel,
Klotzch and Rabenhorst, and in writings mostly cited in the introduction.




316        NEW YIORK STATE AGRICULTURAL SOCIETY.
Loliun perenne, L.
"   temulentum, L.
AMelica sp.
Triticnm scabrum, R. Br.
"9   *vulgare,  Vill.
3a. UST. CARBO VAR. COLUIELLIFERA, TUL.
Panicurn colonum, L.               In the interior of the blossoml at
Pennisetum cenchroides, Rich.        the place of the ovary.
4. UST. DIGITARI.E, Rabh.
Digitaria sp.                      *0On the outer surface of the blossom and the *axis of the spike
(destroys the spike).
5. UST. TYPHOIDES, B. a. Br. (Ust. grandis, Fr.)
*Phragmites communis, Trin.       In the *stems and *sheatlls of the
Typha minor, L.                      leaves.
6. UST. CANDOLLEI, Tul.
Polygonum alpinum, All.            *In the interior of the globular
Bistorta, L.             formed, swollen ovary (with a
"'*Ilydropiper, L.         columella and peridium).
c"    viviparurn, L.
6a. UST. CANDOLLEI VAR. a. BERKELYANA, Tul.
Polygonum sp. (southern Asiatic In the interior of the ovary (sarne
species).                          as above).
7. UST. PHCENIcIS, Cord.
Phcenix dactylifera, L.            In the fruit.
8. UST. FIOUUM, Rchdt.'
Ficus Carica, L.                  In the peduncle (coenanthium);
destroys the fleshy parts.
9. UST. GRAMMICA, B. a. Br.
Aira caespitosa, L.                On the stems.
Glyceria aquatica, Prsl.
10. UST. SALVEII, B. a. Br.
Dactylis glomerata, L.             On the upper surface of the leaf.
1 See Verh. d. zool., botan. Gesellsch. in Wien; 1867, p. 335.




FISCJHER VON WALDHEI.1- USTILA GI.VE.E.         317
11. UST. M-ARNA, DIur.
Scirpus parvulus, R. Sch.         *In the roots.
12. UST. ISCHAEMI, Fckl.
*Andropogon Ischaemum, L.         *On the outer surface of the
spikelet (destroys the blossom)
parts.
13. UST. HYPOGAEA, Tul.
Linaria spuria, Mill.             In the upper part of the roots.
14. UST. ERCEOLORUM, Tul.
Carex *brizoides, L.              *On  the outer surface of the
"  capillaris, L.                 ovary.
" clandestina, Good.
" digitata, L.
ericetornm, Poll.
ferruginea, Scop.
" firma, L.
fliava, L.
"  *glauca, Scop.
" Michelii, Host.' montana, L.
"  ornithopoda, Willd.
panicea, L.
paniculata, L.
"  pilosa, Scop.
"C *pilulifera, Scop.
prsecox, L.
"  Pseudo-Cyperus, L.
pulicaris, L.
" rupestris, All.
Schreberi, Schrk.
" stellulata, Good.
" "supina, Whlbg.
"  *sylvatica Huds.
Elyna spicata, Schrd.
15. UST. MONTAGNEI, Tul.
*Rhynchospora alba, Vahl..        In the ovarly.
Schoenus sp.




318        NEW  YORK STATE AGRICULTURAL SOCIET:.
15a. UST. MONTAGNEI VAR. MAJOR, Desm.
*Rhynchospora alba, Vahl.          *In the ovary.
16. UST. VINOSA, Tul.
Oxyria reniformis, Hook.           In the receptacle of the flower.
17. UST. VAILLANTII, Tul.
Muscari comosum, Mill.
Scilla anthericoides, Poir.        In the *anthers and *pistils (de'  *bifolia, L.                    Bary).
4' maritima, L.
18. UST. OLIVACEA, Tul.
*Carex riparia, Curt.              *In the interior of the ovary.
19. UST. MAYDIS, Lev.
*Zea Mays, L.                      In the *ovary, *stems, *axis of
the spike, masculine blossoms,
and in the lamina of the leaf.
20. UST. SCHWEINITZII, Tul.
Zea Mays, L.                       In the spikes.
21. UST. DREGEANA, Tul.
South African grasses.             On the branches of the panicle.
22. UST. destruens (Schltd.), Duby.1
Dactyloctenium segyptiacum,        In the ovary.
Willd.
*Panicum miliaceum. L.
"     repens, L.
*Setaria glauca, P. B.
23. UST. SECALIS, Rabh.
Secale cereale, L.                 In the ovary.
24. UST. BRO3IIVORX MA. (Ust. Carbo. var. broinivora, Tul.)
*Brolnus secalinus, L., and other  *On the ovary and Xespecially in
species.                           the blossom.
1 Later addition: Us!. neglecta, Niessi. MIpt. (Bot. Zeit., 1868, p. 560), according to the characte,
1stics specified, is nothing else but Ust. destruens.




FISCHER VON WALDHEIJI —   USTILA GINEX:.         819
25. UST. FLOSCULORUM, Fr.:'Knautia arvensis, Coult.            *1In the anthers.
26. UST. ANTHERARUM, Fr. (UST. VIOLACEA [Pers.] Fckl.')'Dianthus Carthusianorum, L.           In the anthers-and, according to
Gagea lutea, Schult. (according to    lTlasne, also  in  the  stigmaUnger and Kuiihn).                     papillae, as well as in the paren-:Lychnis diurna, Sibth.                  chyma  of the  point of the
Flos. cuculi, L.             ovary, the filaments and blosvespertina Sibth.             som leaves
*Saponaria officinalis, iL.
*Silene infilata, Sm.
" nutans, L.
"    *Otites, Sm.
" *rupestris, L.
Stellaria graminea, L.
27. UST. RECEPTACULORUM, Fr.
*Scorzonera humilis, L.                On the outer surface of the *blosTragopogon porrifolius, L.               som receptacle and the *blossoin.
"c     *pratensis, L.
28. UST. UTRICULOSA, Tul.
Polygonum  Ilydropiper, L              At the bottom  of the inner sur"' lapathifolium, L.            face of the  perianth, in  the
c c    minus, I-Iuds.              ovary and at the base of the
c"      *Persicaria, L.            stamens.
iRumex acetosella, L.
29. UST. DURIAEANA, Tul.
Cerastium glomeratum, Thuill.    In the capsule.
30. UST. CARDUT, M. (n. sp.)
*Carduus acanthoides, L.               In the ovary.
31. TILLETIA Dr BARYANA, Mi. (n. sp.)
*Holcus mollis, L.                     *In the lamina of the leaf, near
the outer surface.
I Later addition: VUt. Holostei, de By. (n. sp.); spores from.011 to.013mm. in diameter, rolund,
transparent rose-violet, net-formed thickenings, mostly six-cornered, not entirely regular squares.
Spore powder dark violet. Discovered by de Bary, at IIalle (on the Saar), in April, 1868, on the anthers
and ovary of the Holosteum unbellatum, and very kindly communicated to me by hinm.




30         VE1V YORKi' STATE AGRZICULTURAL SOCIsETY
32. TILL. ENDOPHYLLA, de By. (Uredo olida Riess M. spt.)
*Brachypodium pinnatum, P. B.  *In the parenchyma of the leaf;
next the outer surface.
33. TILL. CARIES, Tul.
Aira caespitosa, L.              In the ovary.
Bromus secalinus, L.
Hordeum murinum, L.
Poa pratensis, L.
*Triticum monococcum, L.
sativum, Lam.'    Spelta, L.
C'   *vulgare, Vill.
34. TILL. SORGHI VULGARIS, Tul
Sorghum vulgare, Pers.           In the ovary.
35. TILL. LOLII, Auersw.
Lolimn  perenne, L.              In the ovary.
*"'temulentum.
36. TILL. SPIIAEROCOCCA (Rabh.) M. (Till. Caries, Tul., var. # Agrostidis, Awd.)
*Agl'ostis vulglris., WVith. var. In the ovary.
pumila, L.
Apera Spica-venti, P. B.
37. SORISPORIUIT SAPONARIs, Rud. (Sorosporiuln Sap. Rud.)
Dianthus Carthusianorumn.        *0Xo the free surfaces of all parts
Gypsophila, sp.                    of the blossom, excepting the
Lychnis dioica, L.                 surface of the calyx.
*Saponaria officinalis, L.
Silene inflata, Sin.
" velutina, Pourr.
3S. USTILAGO CAPSULARUM, Fr. (Thecaphora hyalina, Fingerh.)
Calystegia sepium, R. Br.        In the seed involucre and on the
"   Soldanella, R. Br. (Ann.    outer surface of the anther
Mag. nat hist., 3 ser., v. 18,    wall.
p. 121..)




FISCfHER voN WA LDIEII- USTILA GIaNE.                  321
39.  TTROCCYSTIS PARALLELA (BJ. a. Br.) M.
Carex sp.                             The stems, leaf sheaths and leaves.
Secale cereale, L.
40. UROC. POMPHOLYGODES, Rabh.
Anemone sp.                           In  the %stem, *leaf stalk  and
Ficaria ranunculoides, Roth.             *laulina.
~"Helleborus viridis, L., var. atrovirens among others.
*Rannnculus bulbosus, L.
(("     *repens, L., amnong
others.'
41. URoc. OCCULTA, Rabh.
*Secale cereale, L.                  In the *stern, *leaf sheath, and
Triticuim vulgare, Vill.                *ovary.
42. Urtoc. AGROPYRI M. (Uredo Agrop. Pers.)
*Carex acuta, L.                     In thle  leaves and *stem.
43. UROC. COLCIICI, Rabh.
*Colchieum autumnale, L.             *In the leaves.
Scilla sp.
44. UTnoc. VIOLE (B. a. Br.) M. (Sorosporium schizocaulon Violhe, Ces.)
*Viola odorata, L.                   In the leaves and *petiole.
The following species are knowni besides I (whether they belong to
the Ustilagineoe, and represent independent species, I am  unable to
decide, as I only know them  by name.
USTILAGO axicola Berk., Caricis Fuck., fetens Rlaxen., Haesendonckii
Westendorp, leucoderma Berk., macrospora Desmaz., Persicarise
ienz.,  Tritici (Westend. et Wall. herb. crypt. belge).
SORosPoRIUM Seleranthi (Klotzschii herb. viv. mryc., Nr. 1481).
POLYC~STIS (TUrocystis Rabh.), Filipendule Tul., HIolci Wstd., Lolii
(Bull. de l'Acad. de Belg., ser. xxi, 2, p. 246), opaca Strauss. Polyc.
Ahemones, Ficarite and Ranunculace-aruml (see Fries, Summa. veg.
Scand.) is, without doubt, none other, than as Uroc. pompholygodes, Rablh.
I The Uroc. Pompholygodes, pretended as occurring in the leaves of Tulipa sylvestris (Rabh
Fungi eur. exs. C. X, Nr. 1099), also known under the name of " Caeoma Tulipme, Heufl. in sched."), is
a new species of endophyte, but by no means a Urocystis, as I was able to see on fresher leaves of
the Tulipa, kindly sent to me by Baron von Hohenbuhel-Heufler.
2 Compare Hoffmann, Index Fungorum, 1863.
[AG.]   21




322      1NEW ITORK STATE AGRICULTURAL SOCIETY.
DOUBTFUL USTILAGINEE.
USTILAGO PILULAEFORMIS, Tul.
SIIPORTING PLANTS.         PLACE OF THE SPORE FORMATON.
J uncuns sp., south African species. In the ovary.
THECAPHORA ATERRIMA, Til.
Carex Linkii, Willd.            In the axil of the scale of the
"      praecox, L.            male spike.
*Euphrasia sp. (the parasite known In the *stem of Euphrasia.
as " Sorosporium schizocaulon
Euphrasise, Ces.)
THECAPHORA DEFORMANS, Dur. et Mlntgn.
Medicago tribuloides, Lam.       In the cavity of the ovary.
THECAPHORA DELASTRINA, Tul.
Veronica praecox, All.          In the fruit.
THECAPHORA (.) AURANTIACA, Fingh.
VTrtica dioica, L.               On the under side of the leaf.
THTECAaPORA (?) PALLESCENS, Fingh.
Fragaria collina, Ehrh.          On the leaves.
USTILAGO CYANEA, Rabh.
Balsamia vulgaris, Vitt.  (See  In the interior cavities.
Botan. Zeit., 1855, p. 287.)
USTILAGO (?) CISSI, Tul.
-Cissus sicyoides, L.            In the ovary.
USTILAGO SCLERIME, Tul.
Scleria sp. (from Guiana).      In the pedicle, and scales of the
spikelets.
Ustil. Monotropme and Orobanches, Lev., are at present added to
the species Tuburcinia, Fr., by Tu7casne (Fungi hypogaei, page 196).




FISCHER  VTON WALD/,rfEIi — USTILAGINE.E.           323
STRUCTURE AND APPEARANCE OF THE RIPE SPORES.
The ripe Ustilaginee spores have an endosporium  and an episporhim.  The former surrounds the contents as a colourless, delicate
homogeneous mnembrane, and comes out during the germination, in
the form of a tube, as the promycelium.  It is often invisible -witlhoult
the application of sulphuric acid, but by the use of this acid the episporiumn becomes more transparent, or bursts open, and the inner skin
becomes visible. Iodine and sulphuric acid, or a solution of chloride
of zinc and iodine, causes no blue cellulose reaction; potash causes a
swelling up.
The examination of the structure of the episporiumn shows a conformity with the episporium of other cryptogamous spores.  Here,
also, is found either a smooth, homogeneous episporiullm, or it is provided with various thickenings in form of papillae, nets, etc., which
seldom or never project over the level of it; its greatest part, viewed
in the direction of the radius of the spore, is imbedded within the
episporiim, and it shows only the imperfection of the microscopes of
former observers, who describe the thickenings as projecting over it in
almost all cases.  This is only seldom  the case, and when it is so, it
occurs in Tilletia de Baryana, Ustilago Olivaccea, Maydis, Vaillantii,
and, according to Tulasne, also in Ust. Dregeana and Schweinitzii.
The so-called net-formed thickenings appear as connected, narrow.
more solid, waterless places of the episporium, and surroullnd many
angled, mostly five or six sided,planes or parts, which are watery and
often lie deeper than the thickenings.  As in other cryptogamous
spores, the action of water and various reagents, on the entire episporium and upon a section of it,shows it to be more watery in one place
than another.'
On the outside, the episporium is enveloped by a cuticle, while by
the application of concentrated sulphuric acid the exterior episporium
layer not only does not dissolve, but comes out plainer.
The episporium is either colourless, or much oftener of a dirty yellow, brownish, or rosy-violet colour.
The size of the spores, in the largest diameter, amounts to from.004:mm (Ust. hypodytes) to.03mm (Urocystis Agropyri). Their form
is spheroidal, or elongated spheroidal, or pressed more or less angular
and flat.
I Compare Nageli, Verhalt. d. polar Licht. Sitzungsber. der bayer, Acad. 1862, I. p. 293, sqq.; and
//ofmwter, Pflanzenzelle, Handb. d. physiol. Botan. I. 1867, p. 199, 200.




324         NEW  YORK STATE AGRICULTURAL SOCIETY.
Their contents are composed of protoplasm, wth many drops of
fatty oil.
The ripe Ustilaginese spores appear (1st) mostly, as isolated spores,
unconnected, and (2d) much more seldom, as spore balls or skeins, in
which several spores are united, making a ball or skein.  At the same
time the spores can be classified according to the structure of their
elisporium  and form as follows' 
A. ISOLATED SPORES.
I. WITH FLAT HO'MOGENEOUS EPISPORIUMs.
a. Globular or oval Spores.
Ustilago hypodytes, Fr. (Ust. Lygei, Rabh.)
"    longissima, Lev. (Ust. fusco-virens, Ces.)
Carbo, Tul. (except the var. d. bromivora, Tul.)
Digitarise, Rabh.
"    typhoides, B. a. Br.
Candollei, Tul.,   belong also to the following division.
marina, Dur.,
Ficuum, Rchdt. (according to Reichardt).
"    Phcenicis, Cord.,'    Phcnicis, Cord.,   }- (according to Tulasne.2
Tilletia Sorghi-vulgaris, Tul.
Ustilago grammica, B. a. Br.,. (according to Cooke.s)
Salveii, B. a. Br.
b. Round and long-angular Spores.
Ustilago Ischaemi, Fckl.
hypogaea, Tul. (according to Tulasne)
II. EPISPORIUM WITH AREOLA FORMATION.
a. With a granulous Episporium.
Ustilago urceolorum, Tul.
"    iMontagnei, Tul.
"6 64  var. major, Desm.
vinosa, Tul. (according to Tulasne).
b. Withpapillose Episporium.
Ustilago Vaillantii, Tul.
1 More copious details may be found in my published work, " Sur la structure des Spores des Ustili
ginees," Bul. de la Soc. Imp. des Natur. de MIoscou, 1867, i, p. 242-261, with a chronological table
2 ir. Mem. s. i, Ustilag. p. 76-117.
3 Cooke, Rust, Smut, Mildew and Mould, 1865, p. 203.




FISCHER  VON WALDEIM —  USTILAGINEt'.                   325
c. Episporium  with isolated point.o
Ustilago olivacea, Tul.
"    Mlaydis, Lev.
Tilletia de Baryana, m.
Ustilago Dregeana, Tul.,   (acording to Tulasne).
C    Schweinitzii, Tul.
d. Episporiun    with net-formned thickenings.
Ustilago destruens, Dub.
"    bromivora,  m.
" Secalis, Rabh. (according to Kiihn).
"    antherarum, Fr.
4    flosculorum, Fr.
4'   receptaculorum, Fr.
"    utriculosa, Tul.
Tilletia endophylla, de By.
6" Caries, Tul.'    Loiji, Awd.
sphaerococca (Rabh.) rn.
Ustilago Cardui, m.
D' Duriaana, Tul. (according to Tulasne).
B. SPORE BALLS.
[. THE SPORES COMP(OSINTG Ti EM UNIFORM.
Sorisporium  Saponarihe, Rud. (Ulst. Rudolphii, Tul.)
1I. THE SPORES NOT UNIFORM.
Urocystis pormpholygodes, Rabh.
4"    occulta, Rabh.
"     Agropyri, m.
"C    Colchici, Rabh.
4' C  Viole (B. a. Br.) in
"4    paralella (B. a. Br.) m. (according to Cooke).
5. GERMINATION OF THE SPORES.
Ustilago  spores germinate in many different ways; for the imnost
part still unknown.'  The spores of some species only germinate after
many  hlours, and  under certain  precautions  (Tilletia, Urocystis);
I At present (including my investigations) the germination of the following twelve varieties is
established: Till. Caries, Lolii, Ust. antherarum, flosculorum, Carbo, destruens, Maydis, receptacu.
lorum, longissima, Vaillantii, Urocystis pompholygodes and occulta.




3"26       NEW  YORK STATE AGRICULTURAL SOC1EI'Y.
others, easily, and after a few hours (Ustilago).  The duration of the
capacity of germination varies likewise according to the species.'
For the normal germination of the different species of Ustilago, a
certain quantity of water or moisture is usually necessary.  For this
purpose the spores need only be placed in a drop of water, or upon
mnloistened earth, or even merely in an atlnosphere kept moist; foi
instance, under a glass globe placed over a dish of water. But Tilletia
and Urocystis germinate only in damp air (for instance, under the
glass globe mentioned), and their germinating spores coming in contact with water only show (abnormal appearances.
1 have used in the following germinating experiments freshly
gathered spores, and seldom  those gathered fi'om  one to one and a
half months before.
Ustila/o Jfosculorum  germinates in warmn weather (16-20~ R. in
June), under the most favourable circumstances, after 5-6 hours.2
At the commencement of the germination the endosporiumn comes
through a small opening of the bursted episporium  like a tube.  At
the base this tube - the promycelium - is somewhat smaller than in
the other, almost uniform, part, and rounded at the point (v, 47); the
contents appear slightly glossy, in some places more grumnous.  The
promycelia in a short time lengthen perceptibly, and increase little in
thickness.  In from fifteen to eighteen hours from  the sowing, they
are from.018 to.022mm long, and from.002 to.004mm thick, and
filled with coarse grained contents (v, 48 a).  At the same time
sporidia —  branches of the promycelium — likewise  appear, filled
with granular contents (v, 48 b, c).  The sporidia begin as little
projections with a very small base, coming fiom  the promycelium,
whose protoplasm  comes partly over in them.  After they attain a
certain length they become separated at the base, and complete.  At
the time mentioned generally a terminal sporidiunm appears (of.OO4mm
long and.002mm  thick), and one or two lateral sporidia (of.006 to.007mm long and.002 to.00-mm thick). Secondary sporidia frequently
grow from  the primary of.0035mm long and.0015mm thick.  The
spore itselt is emptied, and collapses.  Sowings made under the
same  germination  conditions  showed, hlowever, great variations
as regards  size.   A  sowing  of the  19tll June, after twentyfour hours, had promycelia of.02mm long, and from.003 to.004mo
thickl (toward the point) as well as primnary sporidia of from.004 to
I See Hqffrann, Unters. uber d. Keim, d. Pilzporen. Pringsh. Jahrb. ii, p. 321, sqq.; and de Bary,
MIorph. und Physiol. d. Pilze, p. 209, sqq., where this subject is treated more copiously.
2 They germinate most favourably ii warm, damp, cloudy weather. Fresh spores in most cases
^rerni nat e the quickest.




FISCHER VON WA LDHEIM- USTILA JNEEr.              327.006'" long, and.003mm thick. But a second so-vwirng, made un(le:
the same conditions, and after the same number of hours, showed
prorryeelia mostly of.036mm long, and at the point of.0025-m thick
(at the base.0005mm thinner) with primary sporidia of.01mm long,
and.004mm thick, whose secondaries were.004mm long, and.002mtnh
thick.
In Ust. flosculorum the formation of the secondary sporidia fronl
the primary is very easily traced, while the latter is still fastened
to the promycelium; also the vactioles which originate still later.
After twenty-four hours, and in other sowings a few hours earlier,
four large, nearly round, bright glossy vacuoles are seen in the promycelium. Between each two of them cross walls of the promycelium
soon distinctly appear (v, 49 a, b). However these cross walls appear
here, as also in other Ustilagos, before the formation of the sporidia
and vacuoles, but are often invisible without the application of
reagents. But if the promycelium is treated with iodine, the contents,
which usually have separated into four parts, show a division which
has already taken place. More seldom  a greater number than five
or six vacuoles is shown (v, 50). The formation of the sporidia precedes the vacuoles, and continues later.  The complete vacuoles take
up nearly the entire space inside of each of the four parts of the promycelium.  Such promycelia remain connected with their spores
until the seed decays.
The sporidia appear mostly on the side of the promycelium, at the
places of division, either singly or in pairs. Sometimes they occur
(also when single) immediately after the germination, in from six to
seven and a half hours from  the seed, on promycelium  of hardly.008mm long. Later their number and dimensions become larger.
Sporidia also occur not only on the point, but also on the side of the
pronyecelium, often some of them  opposite to one another, or irregularly between each of the two divisions of the promycelium, or several
(three or four) surround it in the same plane (v, 52).
The number of sporidia is variable.  Generally, after twenty-four
hours, there are found on the promycelium. from two to four lateral
and one terminal. which later detach secondary ones, and remain
attached to it still one or two days. In other sowings they sprout
again much more abundantly - to the number of from five to seven
lateral, and two to four terminal, to be seen at the same time on the
promyceliumn (v, 52).
Generally the dimensions of the sporidia are the largest wh-en only
a few of them have separated from the promycelium. Besides the




328        NEW. YORK STATE A GRICULTCURAIL SOCIETY.
above mnentioned, there occur cases (for instance, thirty-nine hours
after the seed) where the primary, lateral sporidia attain half the length
and nearly the thickness of the full grown promycelium, and at the
same time detach secondary and tertiary. sporidia, of slnaller dimiensions (v, 51).  Terminal sporidia also appear of considerable length
rand more oval; but do not attain the size of the lateral sporidia.  The
-arious sizes (for instance, of the terminal sporidia) which, at the same
time, are observed on the promycelium, indicate a successive origination of the sporidia (v, 53).
Later the sporidia fall from the promycelium, but remain connected
together, that is, primary with secondary and even tertiary.  Generally a large vacuole is seen in their interiors, near the basal end.
After a few days the appearance of yeast cells, infusoria, etc., obstructs
the observation.  But even in clearer germinations, only a gradual
disappearance of the contents in the sporidia and the promycelium,
and a disorganization following it, is perceivable.
Only in one seeding I observed several cases of copulating sporidia.
The copulation showed itself only on the fifth day after the seed,
between pairs of isolated sporidia lying in water.   Their length
amounted to.004mm, and their thickness to.002mm; they appeared
of a blunt oval forom, with similarly fine grained contents, both of
almost the same size.  The union was effected by a narrow (.0015mm
long',.0005mm thick) cylindrical crooked tube, which was in connexion
with both sporidia lumina.  These copulating sporidia showed no
perceptible changes, and at last, after a few days, perished with the
first seedinog.
Ust. antherarum  germinates generally in from  six and a half to
eight hours.  Spores which were taken from different species of supporting plants showed some differences in this process.
Ust. antherarum  of Diantllhus Carthusianorum  germinates in the
same manner as above.2  The promyceliuml  reaching a length of
nearly the diameter of the spore, has a narrower base, oval point, and
fine grained contents (vi, 1 a).  In later growth it becomes larger in
circumference (vi, 1 b, e), appears of the same thickness through the
entire length, only at the bottoln it is a very little narrower. Worthy
of notice in this Ustilago, is the existence, on the side of the promnycelium, of a pointed knob, at an almost equal distance from its
ends (vi, 1 c).  Commonly on the opposite side of the promycelium
1 Compare my "Beitr. z. Kenntn. d. Ustil," Bot. Zeit., 1867, p. 394, sq.
2 I omit here, and later, the description of the first germination appearances, in so far as they are
similar to those of Ust. flosculorum.




FISCHER VON WALDHEI~-I- USTILA GINEA,.             229
is an imperceptible indentation. The knob projects above the level
of the promycelium, nearly.0005 to.0006mm, and remains later
unchanged.  It is seen eighteen hours from the seed, on the promycelium, wiech has fallen off, or which is still connected with the spore.
From the latter part of its contents passes over through the knob.
Whether the knob merely represents a projection of the promycelium
is difficult to decide, inasmuch as after the application of iodine, or a
solution of chloride of zinc and iodine, the promycelium is divided by
a cross wall at the place of indentation, the middle and both halves are
held together by the knob (vi, 3 c). In this case it suggests a resemblance to a copulation which is effected by it. Proinycelia with distinct
knobs are.014 mm long, and from.002 to.0025mm thick.  In eighteen
hours from the seed, promycelia of half the lenrlth are seen; in the
large ones the contents are already separated into from two to four
parts, with a clear shining oil drop in the centre of each of them.
The partings correspond to indistinct indentations of the promycelium.
Promycelia of from.008 to.01mm long separate sporidia (commonly
two lateral and one terminal).006mm long and.005mm thick. Promycelia and sporidia readily fall off; in from  twenty-four to thirty
hlours most of them have sunk down in the water. At the same time
there appear many sporidia in copulation by pairs (vi, 5, 6), or promycelium in copulation with a sporidium  (vi, 7, 9). Here, also, the
copulation is accomplished by a tube which, according to de 3Bary's
precedent,1 I will designate as a bridge. The latter proceeds from
various places, but most frequently on the side of the promycelia and
sporidia (vi, 7), in the latter, also, often from the point itself (vi, 5).
The bridge is from.003 to.004mm long, and.0015 to.002mm thick,
and is straight (vi, 6, 7), or curved (vi, 8), after several days sometimes bent in the form of " S."  The connexion is firm, because the
bridge does not get broken loose by rolling the sporidia with the
needle, or by infusoria swimming past, etc. Iodine colors it yellow;
weak sulphuric acid causes a little swelling in the middle part, also
the appearance of fine granular contents. Neither iodine nor chloride
of zinc and iodine causes a separation of sporidia in copulation (vi, 7 a).
Their length amounts to from.008 to.Olmm, the thickness from.002
to 004mm. After a few days vacuoles appear in them.
In the promycelia themselves there appear, during the formation
of the sporidia, or shortly before, transverse indentations (vi, 2 a),
and after forty hours (instead of the earlier small ones), as many large
vacuoles as there are divisions (vi, 3 b).
1 Morphol. u. Physiol. d. Pilze, p. 152 & 161.




330       NEW YORK STA:'E A GIRICULTURAL SOCIETY.
The cases where numerous and large sporidia are developed appear
worthy of remark. In a seeding of June 11, in which, however, the
water had dried up unperceived, no germination was shown until the
13th. On this day, fresh water was added, and on the 14th (therefore
after nearly twenty-four hours) promycelia were seen of from.01!m to.012mm long, and.005mm thick. Some of them produced terminal
sporidia of from.005m' to.006mm long. Among them I found a case
where, besides the promycelium with two lateral sporidia, there were
two shorter promycelia on the opposite side of the spore, one of which
produced lateral sporidia (vi, 12).  On the 15th no perceptible
changes occurred. On the 16th almost all the promycelia had produced sporidia at different places, and besides, out of the same spores
from one to two new short promycelia grew. Where the formation
of sporidia had taken place, several vacuoles and indentations showed
themselves in the proumycelium  (vi, 13). On the 17th there were
produced from the primary sporidia, still connected with the promycelium, secondary and tertiary sporidia, of friom.01 to.014mm long, and.006mm thick (vi, 14 to 17). Short promycelia growing on the opposite
side of the spore showed almost the same length as their sporidia,
namely,.005m". In all these organizations vacuoles appeared. On
June 22 (the eleventh day) there were first seen some sporidia (vi, 19)
and promycelia (vi, 20 and 21) with threads, which I will designate as
germinating threads. They were from.014mm to.024mm long, part
straight, part crooked and winding (especially the longer ones). In
this condition the promnycelia were frequently connected with the
spores. In the germinating threads appeared vacuoles (vi, 19 to 21).
In the following days no further particular change showed itself, until
June 26, when the entire seeding became very indiscernible on account
of the too numerous yeast cells and infusoria.
In Ust. antherarum of Saponaria officinalis, a more abundant germination of the promy. celium could be perceived. After twenty-four
hours, many of them had produced germinating threads.02mm and
over in length, and.0015mm thick, to which the greater part of the contents of the promycelium was transferred (vi, 22, 23). After three days,
such promycelium was seen distinctly divided into two nearly equal cells
-the germinating threads growing from the place of separation, or from
the point of the promycelium, appeared.028mm long and.002mm thick.
Some threads had a beak-shaped end (vi, 24). On the 5th day the
promycelium was empty, but the germinating threads were filled with
contents (vi, 25). There were also cases of dichotomous separation
of the germinating threads, with a kidney shaped swelling at the




FISCIHER VON WI'LDHEIm- USTILA GINEN.            331
point of one of the two branches, as it were in order to form a spori.
dium (vi, 26).
I did not discover in these Ustilagos, originating from the Saponaria, either a copulation or so frequent an appearance of the knob.
Ust. antherarum  of Lychnis diurna germinated earlier-in fromn
four to five hours. The other changes were almost the same as those
already described in the first.
Ust. receptaculorum spores (of Tragopogon pratensis) under favourable circumstances germinate in from seven to eight hours. The promycelium increases rapidly in length and circumferencee. It is completely
full of homogeneous, scarcely granular contents. Toward the point the
promycelium  becomes round and appears thicker than at the base.
After twenty hours its length amounted to from.034mm to.036mm,
with a thickness at the middle part and toward the point of.0006mm,
n1md at the base.005mm (vi, 27). It appears straight, sometimes somewhat crooked and winding. At this time sporidia also are found, tubes,
with a smaller base and fine granular contents (vi, 28 to 30). Their
formation seldom follows earlier than thirteen or fourteen hours after
the seed.  They appear, at first, as oval projections, on the side
or at the end of the promycelia whose contents partly pass over into
them, and they increase principally in length. After twenty hours,
sporidia of from.014 to.02mm long, and from.003 to.004mm thick
are perceived. Their number generally amounts to from three to six,
of which one or two are terminal.  The lateral ones occur most commonly at that part of the promycelium, where cross divisions are to be
seen. These divisions and vacuoles aie seldom found until twenty hours 
from the seed. After forty-three hours the promycelia are often from.04m: to.044mm long, and from.006mm to.007mm thick, and at the same
time have four distinct cross divisions, also just as many vacuoles (vi,
31, 32). In Ust. receptaculorum the formation of sporidia precedes
also the cross divisions (vi, 28 to 30). However, by the application
of reagents, it becomes apparent here also that a division has taken
place, which also agrees with Tulasne's' investigations. After the
cross divisions have distinctly appeared, the proinycelium still goes on
to produce sporidia, though their number is in no case considerable,
in proportion to that of the spores germinated. A successive appear.
ance of the sporidia is here, as also in other Ustilagos, not to be mis
taken.
The dimensions of the sporidia are not uniform. Their length increa.
ses rapidly, and attains to from.02mm to.024mm, with a thickness of from
1 2d Mom. sur les Ured. p. 159.




332        NE'w YORK STA TE A GICULTURAL SOCIETY.,004~" to.0025mm; in many seedings (after forty-eight hours), even.04mm by from.005m  to.0055mm thick.
Their t'orm also changes according to the degree of developement.
At the beginning they are rather roundish, but they soon become
nuchl longer.  The free end is the thickest (vi, 30); at the base, on
tihe contrary, they become smaller as they grow further, until, as if
supported on a little short stem, they fall with it from the promycelium.
Becoming free, they grow further and their ends become rounder (vi,
34 to 36). Vacuoles appear in the contents, which at last fill the
entire sporidiumn. Kidney formed sporidia,'as represented by Tulasne,'
did not occur in my experiments.
After eighteen or twenty hours, in many cases even earlier, sporidia are found, not only detached from  the promycelium, but also
still attached to it, in the process of copulation.  The copulation
bridge, similar to that earlier described, occurs at the time when the
sporidia are still attached to the promycelium, which agrees with
de Bary's observations as communicated to me by himself. The points
of connexion are very ununibform, and therefore also the starting
place of the bridge.  As an example, I mention two extreme cases,
between which all others only represent transitions.
In the first the promycelium  showed, on one and the same side,
two longish, somewhat crooked and pointed sporidia, which, coming
firom different heights, almost touched the promycelium, but at the
same time, came in contact, the upper end of the one with the base of
the other (vi, 29).  In the other case two, also lateral, sporidia (with
rounrded points), were observed, which, bending toward each other
crosswise, came to lie with their middle parts one upon the other
(vi, 30).
Between that terminal and this lateral connexion appeared various
modifications.  The most common are those where the bridge passes
from the end of one sporidium into the middle part of the otlher (vi, 33
d); and those in which a lateral connexion between two takes place
close to the ends (vi, 33 b). It is difficult in all cases to see the
bridge at the first; later it becomes thicker and longer, whereby
the connected sporidia likewise enlarge.   Commonly it appeal'rs
straight (vi, 33 a, b, d), often also bent (vi, 33 c).  One of the coniiected sporidia is often still attached to the promycelium (vi, 32);
t!tey are almost always of different sizes, the difference mlay eren
1 1. c. Tab. xii, fig. 26.
2 Compare Morphol. u. Physiol. der Pilze, p. 161, where this is mentioned as conjecture, in con
nexion with the observation, that the sporidia occasionally appear connected three together.




FISCHER VON WALDHEIrM — USTILA1 GINEE.           333
be considerable (vi, 37). For instance, after forty-eight hours one of
the connected sporidia was.04mm long and.0055"m thick, the other
was only half the length and.004mm thick.
After from thirty-eight to forty-three hours, or even somewhat
earlier, in each of the connected sporidia, a cross wall appears.  The two
cells of almost equal size, arising thereby, remain connected together,
and resemble the teleutospores of the. Pucciniae.  In each of them
there is a vacuole.
The sporidia also germinate in these Ustilagos. I observed in one
very successful seeding, spiral tubes after forty-three hours. Each
of them belonged to a pair of connected sporidia which could only be
distinctly perceived by a short length of the tube or germ thread
filled with semi-glossy, granular contents (vi. 34). In these sporidia
also a cross wall was seen, but much less uniformly. Threads of from.04mm to.05mm long appeared also still in distinct connexion with the
sporidia, but these had become so transparent and tender, that they
might have escaped a less attentive observation (vi, 35 and 36). Some
sporidia, with still longer germ threads, vanished almost completely
during the observation. From a pair of sporidia, in whatever way connected, always only one generating thread originates. By a considerable
lengthening, the part of the thread nearest the sporidium becomes transparelit, the granular contents gather together toward the point (vi,
35 and 36); thus, for instance, in a thread of.136mm, one part of it to
an extent of.046mm fiom the base, was empty, and at the same time
cross walls had occurred. Later, the contents of the threads disappeared in places more and more; after six days, no trace of them
could be discovered, although other sporidia and promycelia still
remained distinctly visible.
Germination experiments, by withholding light, gave no anomalous
appearances, excepting the germination occurring some hours later.
LUst. Carbo  spores germinate easily  and quickly, those from
Avena sativa and iordeum distichum in from four and a half to five
hours (at 20" R., v, 7). The number of the germinated spores (of the
supporting plants mentioned) only becomes considerable after seven
or eight hours from the seed. Later, the promycelium often bends,
and a little rounded projection arises on the outer curved side (v, 8
to 11).  After eighteen hours, terminal sporidia are perceived on
many promycelia, on others merely a tapering point, which afterwards
swells up oval and separates as a sporidium. After forty-eight hours
these promycelia mostly appear, as it were, bent in the shape of a
knee, they are similar to the straight ones, and are from.014"m to




3834      NEW YORK STATE A GRICULTURAL SOCIETY..026mm long, and from.0025mm to.003mm thick; the granular contents
have disappeared here and there. Sporidia are mostly found as three
lateral and one terminal (v, 14 and 18). The lateral ones appear at
first as a conical protuberance of the promycelium.  Sometimes a
second promycelium grows out of another place of the spore (v, 12, 13).
In the germination without water in contact with air, the short
promycelium is divided commonly into two or three parts or branches
of unequal length.  The length of such promycelia amounts, for
instance, to.01mm, the thickness.0025mm; while one of the branches
is.054mm long, the other.134mm, with a thickness, when the granular
contents can be seen, of.002am, and at those places which have
become hyaline and are provided with cross walls.0015mm thick.
Besides from such spores grew one or two more promnycelia near to
the earlier one, or more frequently opposite; their granular contents
disappeared in a short time and cross walls appeared in them.
The germination of the Ust. Carbo of Arrhenatheruml elatius proceeds differently from the above. Its promycelia are much longer,
more remarkably pointed, and seldom ha-re the knee-formed bend (v,
24 to 28). Seventeen hours from the seed, most of them possess a
length of.032mm, and a thickness of.002mm; also longish oval sporidia of
the same thickness, and.005mm long, which often contract at the base
in a sterigma. After twenty hours the length of the promycelium
amounts to from.04mm to.066mm; the longest appear, however,
without sporidia, and transparent here and there.
The sporidia occur also later. After forty-eight hours the sterigmas
appear much extended) and secondary sporidia may appear (v, 31 to
33); also promycelia, with one or two lengthened, curved, crooked
branches, whose front ends come mostly in near connexion with the
promycelium (v, 35 to 37), and become hyaline at the point (v, 39).
Some of the fallen off sporidia lengthen in a beak-form at one end.
Occasionally after seventy-two hours this beak-formed end lengthens
into a germ thread, of from.008"m to.02mm long.
Those promycelia coming in contact with the air lengthened
directly into a long thread (v, 41); for instance promycelia of.036mm
long, and.0025mm thick, developed a thread of.15m  long and.001mm
thick.
Experiments by withholding the light caused here also a retarding
of the germination for several hours, and especially a frequent occurrence of the promycelium bent in the knee shape. At the sane time
only few sporidia were produced, in some seedings none at all. A
deviation from the spore formation worthy of mention is shown (v, 30).




FISCHER VON WALDHEIM — USTILA GINE.              8335
Ust. Carbo of Triticum vulgare developes mostly straight, or bowtbrmed, promycelia, the latter from.04mm to.07mm long. The disorganization of the promycelia followed earlier than in those mentioned
above, often after four days.
Ust. iaycli.s germinates best in damp air, and in water with much
nore difficulty. In both cases many ungerminated spores remained
back in the seeding. After twenty-two hours the prolnycelia are.01mm long and.002mm thick.  They are straight (v, 4) or bent in the
upper part (v, 5), and end with a thin point.  Their contents are fine,
granular, and glisten through.  After iorty-eight hours the length of
the straight promycelia amounts to from.036mm to.04mm, their thickness at the base to.0015Tmm, and at the point (wllich now is already
thickened, and contains vacuoles).003nm.  Crooked promycelia do
not show any such considerable enlargenlent.
Tst. longissimna germination is entirely different from the before
llentioned. Fresh spores germinate in warm weather, in summer, in
three hours.   Before the germination, the spore swells perceptibly, its contents appear as a clear shining central drop.  Soon after
a very thin straight tube, the prornyceliumn, comes through a small,
scarcely perceptible opening of the episporium. It swells at the free
end; this swelled part — or sporidium - increases in length, and separates visibly from the thinner part - the promycelium - which also
becomes longer (v, 42 and 43). In an hour after the germination,
the promycelium is.004mm long, and from.0005mm to.001mm thick,
the sporidium from.006mm to.008mm long and.0015mm to.002mm
thick.  Very soon the latter separates, whereby the promycelium,
together with the spore, change their position somewhat. The promiycelium shortens perceptibly soon after; in an hour a new sporidium
is formed in the same manner. The separation of the sporidia follows
thus successively, and only at the point of the promycelium; it continues also until the spore has emptied, commonly after about twentyfour hours. How great altogether the number of the sporidia are
which separated from one promycelium, I could not exactly decide,
notwithstanding the repeated use of Recklinghauseu's moistening
chamber.'  The great number of fallen off sporidia in a twentyfour hour  seeding, which  surpass the number of the spores
several times, shows, without doubt, that the separation is often
repeated.  The separating spores increase in length perceptibly, for
after seventy-two hours they are mostly from.008mm to.014mm long;
there are also a few vacuoles to be seen in them. Sporidia are also
1 See Virchow's Archiv., Bd. 28, 1863, p. 162.




336       YNEW YORKI STr,4TE AGRICULTURAL SOCIETY.
found, though but seldom,.03nm long, twenty-four hours from  the
seed. The fallen off sporidia sometimes separate secondary sporidia.
No other changes occurred, worthy of mention, until the seeding
perished.
In other seedings the germination occurred in from four to seven
1hours. Spores which have laid one or two months, germinate slower,
sometimes in not less than eighteen hours.
If the promycelium in germinating comes in contact with the air
(for instance, by the evaporation of the water), it divides in several
branches of unequal length, and in that way resembles the sporidium
of Tilletia Caries (v, 44 to 46).
That case where, after twenty-four hours, a lateral sporidium occurs
at the base of the promycelium, cannot be considered otherwise than
as abnormal.
De Bary investigated the germination of Ust. Vaillantii (from
unpublished observations).  To judge from the drawings, they are
very similar to Ust. longissima.
~Tilletitz Caries belongs to the late and more difficult germlinating
species. Their germination only occurs after at least fifty hours and
more, and that only by strewing the spores upon water, or in damp
air (v, 1). At the length of.004mm the promycelium  is alr eady.00}25mm thick.  The course of the germination shows the same
appearances as have been represented by Kiihn (see the historical
introduction), and requires here, therefore, no further description.
T remark here that the length of the primary sporidia amounts to.08mm. Those promnycelia which occur in contact with water are
only.24inm long, and have protoplasmatic contents only in the upper
half:
I succeeded only a few times in observing the germination of Urocystis pomphiolygodes (of various species of Ranunculus).  The promnycelium  springs exclusively from the middle, darker coloured spores
of the ball. If the germination ensues in a very little water, so that
the forthcoming promycelium  comes in contact with the air, in
twenty-four hours they become.004mm long and.002m  thick, but
send from  their front end three or four fork-formled branches, or
sporidia, similar to Tilletia Caries (vi, 38). All these formations are
filled with fine granular contents.  The form of the sporidia is longish
oval, broader in the upper part than at the base. The point is
mostly rounded, seldom long drawn out. The length of the sporidia
amounts to from.01mm to.014mm, the thickness to from.003mm to.0035mm. After twenty-four hours the sporidia were.022 mm long and




FISCHER VON WALDHEIM- USTILA GIrNE..    33 7.004'm thick (vi, 41). At the same tinle vacuoles, quite small or else
two large ones, appear in each of them (vi, 42). No further changes
took place before the disorganization.
If the germination takes place in water, a promnycelium appears
which forms no sporidia,'it lengthens immediately (vi, 39), or divides,
though seldom, in two branches, which likewise become long (vi, 40).
After twenty-four hours the undivided promyceli urn, as well as the other,
together with the branches, attains a length of from.14mm to.16mm
and a thickness of from.004mm to.006m. The contents retrograde
to the point; the rest appears hyaline and septate.  Some branches
still show a fork-like division at the point. After four days the tubes
appear entirely transparent and shrunk, break up (vi, 43), and become
indiscernible.
Notwithstanding many repeated attempts, I did not succeed in
getting the following to germinate: Tilletia endoophylla, de Baryana,
Ustilago Candollei, utriculosa, urceolorum, hypodytes and Sorisporium Saponarioe.  My experiments with these species were not only
with fresh spores, but also with such as had been lying a few weeks
and even from one to two years. The seeding was tried in and on
water, as well as in damp air. In order to modify the germination,
and suit the natural conditions more, the spores were sown on very
thin transparent slices of elder pith, moistened in different degrees,
but in all cases without any satisfactory results. I could not investigate the germination of the remaining Ustilagos for the want of fresh
material.
6. EXPERIMENTS ON THE PENETRATION OF THE USTILAGINE29 IN THEIR
SUPPORTING PLANTS.
In order to demonstrate the penetration of the germinated spores
in the supporting plants, I experimented with Ust. receptaculorum,
fiosculorum and Carbo.
The germinating seeds of Tragopogon pratensis were sufficiently
sprinkled with water and spores of Ust. receptaculorum (Tragopogonis)
sowed on different places of the embryo, and the seeding was kept
under a glass globe in a moist condition.  Some days thereafter promycelia and sporidia were visible, but only lying on the surface of
the embryo. Neither penetration nor germination of the sporidia
took place any where then or. afterward. Spores placed upon the
surface of the root and leaves, as well as of the stalk, and observing the
same mode of proceeding as in the former experiments, germinated
with more difficulty than in a drop of water on an object plate, and
[AG.]        22




338       N7.EW YORE STATE AGRICULTURAL SOCIETY.
there  was no  penetration whatever of the germination products
perceptible in the supporting tissue.
Sowirg the spores of Ust. flosculorum  on different parts of the
growing Knautia arvensis gave the same negative results.
The most attention was given to sowing Ust. Carbo on oat and barrey
(eb!lbryos. As in the former species experimented upon with reference
to their germinating capacity, fresh spores originating on the oat and
barley plants were placed upon the germinated seeds, and even on the
entire surface of the rootlets, between them and the radicle, and, in
more developed embryos, on the base point and entire surface of the
leaves. The quantity of the spores was likewise not uniform. In this
way infected embryos grew on earth, clean sand, or on the object plate.
In each of these cases sufficient damp air was provided.  Other
embryos were sprinkled in the same manner with germinated spores.
In other cases I sowed the seed with spores before their germination,
so that in sprouting the rootlet not only came in contact with spores,
but also met promycelia and sporidia. In all the above cases the
result was the same-negative. Notwithstanding multitudinous germinations of the spores in the form of promycelia and sporidia, not a
single case occurred of a penetration into any of the cells of the supporting plant. These experiments were repeated many times, but
without successful results in any case. They are therefore in direct
contradiction to Kuiihn's Tilletia investigations (which I was unable
to repeat, therefore this case is not to be considered here). Further,
they are in contradiction to Hoffmann's statements of Ust. Carbo,
but he says himself that out of a hundred infection experiments, only
very few were successful. In view of this statement, and as in my
investigations all precautions were taken, by means of which the penetration of other parasites, such as Uredos and Peronospora, had been
easily accomplished, it appears to me that these negative results surely
indicate that for the penetration of the Ustilago germs, peculiar conditions are necessary which at present are not yet discovered. The
nature of these conditions, and whether a di-morphism is perhaps connected with it, is to be discovered through further observations.
CONCLUSION.
All Ustilagineoe are exclusively endophytic fungi, inasmuch as
their vegetation is accomplished only in the interior of the supporting
plant. No matter whether the particular part inhabited by them
grows above the ground or under it, or even under water.




FISCHER VON WALDHEI —         USTILA GIVNE.        339
The formation of their fruit takes place always in certain organs of
the supporting plant, and according to their species, either in the
interior of the tissue or on the surface. All Ustilaginee formnn spores,
mostly in large massive beds, in which the spore formation is centripetal.  With some exceptions, there is only one kind of spore knowr
at present formed upon the supporting plants.  Sorisporium S&cponarice, Ustilago marina' and caypsuarum' make these exceptions.
In germination all Ustilagineme produce promycelia which form
sporidia, though in many species the sporidia development proceeds
mtolre sparingly than in others.  As the sporidia are to be considered
as a particular species of spore, thus to all Ustilagineoe belong two
kinds of spores, and to Sorisporium  Saponarie, Ustilago maria and
capsularum probably three.
The penetration of the embryos of the Ustilagineoe into the supporting plant, and cases of di-morphism, etc., are to be demonstrated by
further experiments. Two facts make it highly probable that the
myceliumrn in some way enters at the base of the young plant, first,
that in all positively ascertained cases of plants a year old, the mycelium passes through the entire attacked plant from  its base to the
place of spore formation; and, second, that in the numerous completely known parasitic fungi, the mycelium grows through the
attacked plant from the place of its entrance to the place of the spore
formation (Uredos, Peronospora).
The relation of the Ustilagines to the growth and developement
of the supporting plant is diverse and often very strange.  The?emycelium injures the supporting plant the least.  So it is met with
in the cells which are still filled with chlorophyll (Ust. Carbo), or
which contain a primordial utricle (Ust. Maydis). The developement
of the pollen still takes place, so long as the mycelium only, or at the
most only the first commencement of the spores are present, in the
anthers.
On the contrary, the spore formation itself exerts a much more
injurious influence. Ust. longissima and Sorisporium Saponariae affect
the growth of the supporting plant in this relation most unfavourably.
Others hinder the growth of the supporting plants less than the formation of some of the organs, and therefore cause abnormal conditions. Thus Tilletia Caries destroys the ovulum, Ustilago Candollei
and Maydis which also affects other parts of the maize, the entire ovary,
as in part also does Ust. urceolorum.  Ust. flosculorum and antherarum
prevent the formation of the pollen, and therefore cause sterility of the
1 7Tdasne, Super. Fries. Taphr. g. Ann. d. sc. nat., iv ser., t. v, 1867, p. 133 to 136.




340       NEW  YORK STATE AGRICULTURAL SOCIETY.
blossoms. Ust. Carbo and destruens destroy the entire blossom. Sorisporium Saponarise is also followed by sterility cf the blossom. Ustilago
hypodytes, Tilletia de Baryana and endophylla cause a backwardness
in the developement of the main stem, together with the leaves,
spikes, etc.
To conclude, the biological appearances of the Ustilagiie2e show a
great conformity with the known facts of other endophytes, and
especially with those of the Uredos, with which latter the Ustilaginee
are generally associated in their system.
Moscow, December; 1867.
REVIEW OF THE SUPPORTING PLANTS AND THE
USTILAGINE.zE OCCURRING  ON THEM.
CRYPTOGAMS.
Fungi: Tuberacei.
Balsamina vulgaris, Vitt.-Ust. cyanea, Rabh. (doubtful species).
MONOCOTYLEDONS.
Typh/acece.
Typha minor, L.-Ust. typhoides, B. a. Br.
Graminece.
Loliim perenne, L.
olim  permentne, L.  J -Ust. Carbo, Tul.; Till. Lolii, Awd.
"   temulentum, L. 
Brachypodiumrn ciliatum, P. B1. —Ust. Carbo, Tn].
- c "   pinnatum, P. B.-Till. endophylla, de By.
Agropyrum sp. (?)-Uroc. Agropyri m.===Uredo Agrop., Pers. (.)
repens, P. B.-Ust. hypodytes, Fr.
Triticum monococcun, L   -Till. Caries Tl.'"   sativum, Lam.
scabrulm, R. Br.-Ust. Carbo, Tul.
vulgare, Vill.-Till. Caries, Tul., Ust. Carbo, Tul., Uroc.
occulta, Rqbh. (in Australia), Ust. hypodytes, Fr. (? See
Westend. et Wall. herb. crypt. beige, Nr. 1161).
Secale cereale, L. —Uroc. occulta, Rabh.; Ust. Secalis, Rabh.; Uroc.
parallela (B..a. Br.) m.
Elymus arenarius, L.-Ust. hypodytes, Fr.
Hordeum distichum, L. —Ust. C rbo, Tul.
"    mlurinum, L.-Ust. Carbo, Tul.; Till. Caries, Tul. (Durieu)
C"   vulgare, L.-Ust. Carbo, Tul.




FISCHER VON T7WALDHEIF — USTtrsA GINEA.            341
Bromns secalinus, L., and other species.-Ust. bromivora m. (-Ust.
Carbo var. d. bromiv., Tul.), Till. Caries, Tul. (Philippar).
Festuca pratensis, Huds.-Ust. Carbo, TuL
Dactylis goloinerata, L. —Ust. Salveii, B. a. Br.
sp. —Ust. longissima, var. megalospora, Riess.
Melica sp.-Ust. Carbo, Tul.
Poa pratensis, L.-Till. Caries, Tul. (Philippar).
"  sp.-Ust. longissima, var. megalospora, Riess.
Glyceria aquatica, Prsl.- Tst. longissima, L6v.; Ust. grammica,
B. a. Br.
fluitans, R. Br.-Ust. longissima, Lev.; Ust. hypodytes,
Fr. (Tulasne).
spectabilis, M. K.-Ust. longissima, Lev.
Hiolcus mollis, L.-Tilletia de Baryana m.
Avena flavescels, L.
ipbeseens L.  — Ust. Carbo, Tul.
"   sativa, L.
Arrhenatelcrunl elatius, P. B.-Ust. Carbo, Tul.
Aira caespitosa, L.-Ust. Carbo, Tul.; Ust. grammica, B. a. Br.;
Till. Caries, Tul. (Philippar).
Dactyloctenium aegyptiacum, Willd.-Ust. destruens, Dub.
Agr ostis vulgaris, With. var.pumila,L.  — Till. sphaeroeocca (Rablh.) mi.
Apera Spica-venti, P. B. 
Phragmites communit, Trin. —Ust. typhoides, B. a. Br.; Ust. hypodytes, Fr.
Sorghum vulgare, Pers.-Till. Sorghi, Tul.
Andropogon hirtus, L. —Ust. Carbo, var. columellifera, Tul. (a. transfissa).
Ischaemum, L.-Ust. Ischaemi, Fekl.
Cynodon Dactylon, L. —Ust. Carbo, Tul.
Digitaria sp. —Tst. Digitariae, Rabh.
Pennisetum cenchroides, Rich.  — Ust. Carbo, var. r columellifera,
Panicum colonum, L.                  Tul. (b. tricophora).
9"    miliaceum, L.-Ust. destruens, Dub.
repens, L. —Ust. l ypodytes, Fr. (with  spores of from.0032mm to.0048mm in diameter); list. destruens, Dub.
Setaria glauca, P. B. —Tst. destruens, Dub.
Lygeum sp. —Ust. hypodytes, Fr. (=Ust. Lygei, Rabh.)
Zea Mays, L.-Ust. Maydis, Lev.; Ust. Schweinitzii, Tul.
Soutll African grasses. —Ust. Dregeana, Tul.




342.Nw YORR STATE A GRICULTURAL SOCIETY':
yyperacews.
Carex acuta, L.-Uroc. Agropyri m. (==Uredo Agrop., Pers).
"  brizoides, L.
"   capillaris, L.
clandestina, Good.
"   digitata, L.
" ericetorum, Poll. t -Ust. urceolorum, Tul.
"  ferruginea, Scop.
firma, Host.
flava, L.
"  glauca, Scop.
"  limosa, L.
Linkii, Willd.-Thecaphora aterrima, Tul. (doubtful species).
Michelii, Host.'  montana, L.
"  ornithopoda, Willd.
"  panicea, L.          _ —Ust. urceolorum, Tul.
"  paniculata, L.
"  pilosa, Scop.
"  pilulifera, L.
"  praecox, L.-Ust. urceolorum, Tul.; Thecaphora aterrima:
Tul. (doubtful species).
P seudo-yperus, L.   Ust. urceolorum  Tul.
" publicaris, L.
riparia Curt.-Ust. olivacea, Tul.
rupestris, All.     1
"   Schreberi, Schrk.
stellulata, Good.   ~ —Ust. urceolorum, Tul.
"  supina, Whlbg.
"   sylvatica, HIuds.
"   sp.-Uroc. parallela (B. a. Br.) m.
E yna spicata, Schrd.-Ust. urceolorum, Tul.
Seeria sp.-Ust. (?) Sclerise, Tul. (doubtful species).
Scirpus parvulus, R. Sch. —Ust. marina, Dur.
Rhyncllospora alba, Vahl. —Est. Montagnei, Tul.; Ust. Montagnei,
var. major, Desm.
Schcenus sp.-Ust. IMontagnei, Tul.
Juncacece.
Juncus sp. —Ust. pilulaeformis, Tul. (south Africa; doubtful species).




FISCHER VON WALDDHEIM- USTILA GINElm.            349
Colchicacece.
Colchicum autumnale, L.-Uroc. Colchici, Rabh.
Liliacece.
Muscari comosuin, Mill.
Scilla antherieoides, Poir.  {Ust Vaillantii Tui
"  bifolia, L.          j
"  maritima, L.         J
"  sp.-Uroc. Colchici, Rabh.
Gaogea lutea, Schult.-Ust. antheraruin, Fr. (?)
Parlme.
Phoenix dactylifera, L.-Ust. Phoenicis, Cord.
DICOTYLEDONS.
Artocarpece.
Ficus Carica, L.-Ust. Ficuum, Rchdt.
Utricacece.
Urtica dioica, L:-Thecaphora (?) aurantiaca, Fingh. (doubtful speaies).
Polygoneac.
Rumnex Acetosella, L.-Ust. utriculosa, Tul.
Oxyria reniformis, Hook.-Ust. vinosa, Tul.
Polygonum alpinum, All.   -Ust. Candollei, Tul.
"     Bistorta, L.
"     i Hydropiper, L.-Ust. Candollei, -Tul.; Ust. utriculosa,
Tul.
t"   lapathifolium, L. }
t"    minus, Huds.      -Ust. utriculosa, Tul.
cc    Persicaria, L.
t"    viviparuin, L. —Ust. Candollei, Tul.
sp, (south Asiatic species)-Ust. Candollei, var. a. Berkeleyana, Tul.
Dipsacea.
Knautia arvensis, Coult. —Ust. floscnlorum, Fr.
Comnpositce.
Carduus acanthoides, L. —Ust. Cardui m.
Tragopogon porrifolius, L.
"     pratensis, L.    — Ust. receptaculorum, Fr.
Scorzonera humilis, L.




344       NEw YORK STATE AGRICULTURAL SOCIETY.
Convolvulacece.
Convolvulus arvensis, L.    )
Calystegia sepium, R. Br.     — Ust. capsularum, Fr.
Soldanella, R. Br.
Srophkularineae.
Linaria spuria, Mill.-Ust. hypogaea, Tul.
Veronica  praecox, All.-Thecaphora Delastrina, Tul. (doubtful
species).
Euphrasia  lutea, L.- Thecaphora aterrima, Tul. (-= Sorisporium
schizocaulon Euphrasike, Ces.; doubtful species).
Rosoace.
Fragaria collina, Erhr.-Thecaphora (.) aurantiaca, Fingh. (doubtful
species).
Papilionacece.
Medicago tribuloides, Lam.-Thecaphora deformans, Dur. et Montgn.
(doubtful species).
Ampelidece.
Cissus sicyoides, L. —Ust. (?) Cissi, Tul. (doubtful species).
Alsinece.
Stellaria graminea, L.-Ust. antherarum, Fr. (Sowerby).
Cerastium gloineratum, Thuill.-Ust. Druiaeana, Tul.
Silenece.
Gypsophila sp.-Sorisporium Saponarise, Rud.
Dianthus Carthusianorum, L. —Ust. antherarum, Fr.; Sorisp. Saponarie, Rud.
"    Seguierii, Vill.-Sorisp. Saponariae, Rud.
Saponaria officinalis, L.-Ust. antherarum. Fr.; Sorisp. Saponariae,
Rud.
Silene inflata, Sm.-Ust. antherarum, Fr.; Sorisp. Saponariae, Rud.
"   nutans, L.
"   Otites, Sm.           -Ust. antherarum, Fr.
"   rupestris, L.
-" velutina, Pourr.  -Sorisporium Saponarie, Rud.
Lychnis dioica, L.
diurna, Sibth.
"    Flos cuculi, L.  -Ust. antherarum, Fr.
vespertina, Sibth.




FISCHER VON WALDHEI — USTILAGINE..345
Violaceca.
Viola odorata, L.-Uroc. Violae (B. a. Br.) m. ( —Sorosp. schizocau.
ion Violhe, Ces.)
Ranunculacece.
Anemone sp.
Ranunculus bulbosus, L.                     -Uroc. pompholy.
44    repens, L., and others.    -        godes, Rabh.
Ficaria ranunculoides, Roth.                    godes, Rab
Helleborus viridis, L., var. atro-virens and others.




346        i2V  w  YORK S'A TE A GRIC ULTURAL SO CIET'Y;
EXPLANTATION OF THE PLATES.
PLATE I-MYCELIUM.
FIG. 1-TILLETIA ENDOPHYLLA D B Y (900-1).
Fig. 1. Long, extended threads, with vacuoles extending through several cells.
Cut lengthwise through a young leaf.
FIG. 2-4-TILLETIA DE BARYANA M. (900-1.)
iYg. 2. Isolated, exposed threads, with large vacuoles, from the parenchyma cells
of the base of the leaf. Cut lengthwise.
Fig. 3. Threads with the commencement of the ramification and a few vacuoles.
A young leaf sheath which has sprouted from the rhizoma cut lengthwise.
Fig. 4. Threads penetrating through the cells. Cut across one of the nodes.
FIG. 5-8-USTILAGO MAYDIS LEV.
Fig. 5. Long, extended threads beginning to ramify, running through a pith cell (in
which a nucleus is still to be seen). From the second internode below a blighted
rhachis (500-1).
Fig'. 6. A long, extended thread, covered with a cellulose sheath and passing
through many cells. At a the thread shines through; at b is plainly seen the lumen
of the cut sheath and of the thread. Cut lengthwise through the rhachis at the bottom
of the ovary (250-1).
F-ig. 6a. The previous figure at a, a conspicuous piece of thread enlarged 900 times;
in the thread is seen the contents, the cellulose sheath is strongly marked.
Fig. 7. Threads which have become distinctly visible in the cellulose sheath by the
application of potash and iodine. Shown detached from a cell of the ovary wall (500-1).
Fig. 8. Thin walled thread (like the transition form in the sporeforming threads),
with many shoots and fine grained contents. Parenchyma of ovary wall cut lengthwise (500-1).
PLATE II-MYCELIUM.
FIG. 1-4 —USTILAGO CARBO TIL. OF OATS.
Fig. 1. A thread contained in cellulose of shining appearance in the node cells.
Cut lengthwise (500-1).
Pig. 2. Mycelium running partly in and between the cells, with the commencement
of branches within the cells. A node from the upper half of the stem cut lengthwise (900-1).
Fig. 3. Mycelium, with a sucker in the upper part. The stem of a very young hothouse root cut lengthwise (500-1).
Fig. 4. A piece of a thread passing through two cell walls; cut lengthwise in the
node of the above hot-house root (500-1).




.FISCHER    VON WALDHEIM- USTILA GINEE.                  3447
FIG. 5 AND 6-UsTILAGO CARBo  TUL. OF ARRHENATHERUM ELATIUS.
Fig. 5. Mycelium from the uppermost node. Cut lengthwise (900-1)..ig. 6. Branched threads in the cells of the base of the glumes; transition form in
the spore-forming threads. From a young blossom still unrolled (500-1).
FIG. 7-USTILAGO  HYPODYTES FR., FROM ELYMUS ARENARIUS.
Fig. 7. Threads with cross walls and watery contents. The main stem cut lengthwise (900-1).
FIG. 8-USTILAGO LONGISSImA LEV.,FROM GLYCERIA FLUITANS.
Fig. 8. Threads filled with a watery fluid, running in the cells and through the
cell walls. Prarenchyma of the leaf cut lengthwise (900-1).
FIG. 9-UsTILAGO ANTHERARUM FR., FROM SAPONARIA OFFICINALIS.
Pig. 9. Threads running in the anther walls (900-1).
FIG. 10-12-SORISPORIUM SAPONARizE RUD.
Fig. 10. A crooked thread furnished with a sucker, penetrating through the cell
walls. Node cut lengthwise near the rhizoma (500-1).
Fig. 11. A straight thread having partly plastic and partly watery contents and
vacuoles. Cut lengthwise through the under part of the ovary (900-1).
Fig. 12. The upper part of the thread in a cellulose sheath, the remainder ramifying. The main stem cut lengthwise (500-1).
PLATE III - SPORE-FORMING THREADS AND SPORE FORMATION.
NOTE.-The dark, shaded lumps and stripes in the interior of the spores and threads represent the
glossy contents.
FIG. 1-5-TILLETIA CARIES TUL., FROMi WHEAT.
Fig. 1. Spore-forming threads with the commencement of branches, on the end of
which occur the spores. From an ovary of 1 cent. long (900-1).
_Fig. 2. A young spore still in connexion with the thread (900-1).,Fq. 3. Young spores in various stages of developement, mostly on the branches
of the principal thread. The spores contain oil drops, the smallest a vacuole (500-1).
Fig. 4. Separation of the thread into two branches, on the end of each of which are
spores of different degrees of ripeness (500-1).
Fig. 5. Clear transparent, single contoured spore, with the remains of the sporeforming thread. In the contents, oil drops (500-1).
FIGS. 6-13. TILLETIA ENDOPHYLLA, D BY.
All conditions of the young leaves of the Brachypodium pinnatum. (Figs. 6 to 10,
500-1; the remainder, 900-1.)
Fig. 6. Swelling of the oil-containing threads preliminary to spore formation.
Fig. 7. Commencement of spores of a longish form, and with an oil drop. In figs
7 to 9, a denotes the boundary of the cell wall from which the thread projects.
Figs. 8, 9. Later conditions; the spore is rounder, but has not yet attained the
limits of the thread attached to it. In the contents an oil drop.
Fig. 10. A developed spore which has come to full size.
Fig. 11. Developed spores, with beak-shaped lengthening, and partly still in con
nexion with the spore-forming thread. In the spores, oil drops and vacuoles. Cut
lengthwise.




348         NEW YORK STATE A GRICULTURAL SOCIETY.
Fig. 12. The endosporium is distinctly tobe seen in the beak-formed continuation;
except at this place, the episporium covers the entire periphery.
Fig. 13. An incompletely formed transparent spore, whose contents appear like a
large oil drop.
FIG. 14-16. TILLETIA DE BARYANA M. (900-1).
Fig. 14. Spores distinctly occurring on the branches, one of them still connected,
the other, its limits attained, has become rounder.
Fig. 15. The gelatinous layer of the upper end of the thread still surrounding the
spore.
Fig. 16. Completed condition; the spore still surrounded with the gelatinous layer.
FIG. 17-24. USTILAGO MAYDIS, LEv.
Fig. 17. Form of the threads as they sometimes occur in the cells of the ovary, in
which there are already ripe spores; their use is still unknown (900-1).
Fig. 18. Thread become gelatinous; the contents are indistinctly seen. From a
gelatinous spore mass (900-1).
Fig. 19. Spore-forming threads, with bright oleaginous contents, running between
the cells. Ovary cut lengthwise (500-1).
Fig. 20. Gelatinous thread with a distinct breaking up of the contents for spore
formation; the upper part represents the same conditions as fig. 19. Its course
between the cells was still, in part, distinctly perceivable (900-1).
Fig. 21. More transparent spores forming in the interior of the gelatinous thread
(900-1).
Fig. 22. The same; however a great number of the transparent spores are seen
lying in a row, lengthwise in the stem. The latter appears more transparent (900-1).
Fig. 23. A spore become brownish, surrounded by the gelatinous membrane of the
thread. From the ovary wall (900-1).
Fig. 24. An affected ovary cut lengthwise; the single mass of ripe spores appear
as black spots or stripes arranged between the vessels (natural size).
FIG. 25, 26. USTILAGO HYPODYTES, FR., FROM ELYMUS ARENARIUS (900-1).
Fig. 25. Gelatinous threads, in which spores are already formed; isolated from a gelatinous spore mass.
Fig. 26. Stem cut crosswise, on the surface a bed of ripe spores; in the cells mycelium,
which enters the exterior layer of cells.
FroIG. 27-29. USTIL.aGO LONGISSIMA, LEV., FROM GLYC. FLUITANS (900-1).
Fig. 27. Gelatinous thread, in which spores have already developed. From a spore
mass.
Fig. 28. Part of the edge of a young spore mass; the gelatinous thread breaking
up into spores of glossy gelatinous appearance.
Fig. 29. Young transparent spores with glossy oleaginous contents.
FIG. 30-35. USTILAGO ANTIERARUM, FR., FROM DIANTHUS CARTHUSIANORUM.
Fig. 30. Gelatinous threads, with swellings at the beginning of the spore formation;
t the 1border of the epidermis cell. An isolated anther wall cut lengthwise (500-1).
Fig. 31. Isolated gelatinous thread with distinctly breaking up glossy contents,
whose lumen at the under end is still filled up as earlier (900-1)
Rig. 32. Gelatinous thread, of which the greater part has already distinctly broken
up into spores; at its end riper conditions (500-1).




FISCHER  vON  WALDHEIM- USTILAGINER E.                  349
Fia. 33. A number of gelatinous threads run into one another, in which spore
f)ormation is distinctly going on. The round, mostly contoured contents shine distinctly through. From an anther (900-1).`Fiq. 34. Distinct breaking up of the thread into spores; at a the contents have divlded into three parts for spore formation; b, remains of the thread (900-1).
Fig. 35. In the upper part of' the thread a coloured spore, in the loker part the
commencement of a spore (900-1).
FIG. 36. USTILAGO ANTiLrLALtM FJ'S., FROM LYCHNIs DIURNA (900-1).
Fig. 36. Piece of a gelatinous thread with very irregular ramifications, and spores
in different stages of development. F'rom a young anther.
FIG. 37. USTILAGO ANTHERARIUM Fin., FROMI SAPONARIA OFFICINALIS (900-1).
Fig. 37. Distinct contraction of the gelatinous thread and with the contents breaking up.
FIG. 38. USTILAGO UTRICULUSA TUL., FROM POLYGONUM PERSICARIA (900-1).
Fig. 38. Fork formed separation of the gelatinous thread, and therefore of the contents which pass in to the branches.
FIG. 39-42. USTILAGO FLOSCULORUM FR. (900-1).
Fig, 39. In the upper part the distinct remains of a gelatinous thread; further
down the spores begin to develope.
Fig. 40. Gelatinous thread, in whose branches spores also develope.
FPig. 41. Gelatinous thread with still very young commencement of spores and with
ramifications.
FIig. 42. Gelatinous mass of threads running in and out of one another, with contents
breaking up and completed spores. From a part of the edge of a young spore mass.
FIG. 43-40. USTILAGO RECEPTACULORUM FR., FROM TRAGOPOGON PRATENSIS.
(Fig. 45. From an unpublished drawing of de Bary 500-1.)
Piq. 43. A thread already become slightly gelatinous with distinct ramifications.
From a young spore mass.
Fig. 44. The upper part of a thread with a strong gelatinous membrane, and with
a contents mass.
Fig. 45. Later conditions, in the branch the spore membrane appears distinct, the
gelatinous membrane becomes thinner; only in the thread itself is seen the first breaking up of the contents.
Fig. 46. Young, but considerably developed spores, still surrounded by the gelatinous membrane of the thread.
PLATE IV - SPORE-FORMING THREADS AND SPORE FORMATION.
FIG. 1-6. USTILAGO URCEOLORUM TUL., FROAM CAREX PILULIFERA.
Fig. 1-4, 900-1;`Fig. 5-6, 500-1.
Fig. 1. Gelatinous threads, whose contents in the narrow lumen only glimmeil
through in a few of them. From a tress of threads from the interior of the ovary
and somewhat spread apart with a needle..Fig. 2. Separated thread with the contents distinctly passing over in the side branch
Fig. 3. Separated piece of thread, the contents breaking up into shining stripes.




350         NEW YORK STATE A GRICULTURAL SOCIETY.
Fig. 4. The same, but with a broader lumen; the contents are only visible in places
Fig. 5. Spore formation out of threads run into one another, whose contour is
still perceivable; the spores themselves are in different degrees of developement.
Separated from a young spore mass.
Fig. 6. Unripe spores, still surrounded with the gelatinous membrane; the'contour
of the threads from which they came indiscernable.
FIG. 7-23. SORISPORIUM SAPONARILE RUD.
(Figs. 13, 14 and 16 to 23 from unpublished drawings of de Bary's.)
Fig. 7. Mycelium threads boring through the cell walls and immediately changing
to spore-fbrming threads (900-1).
Fig. 8. A gelatinous thread, with a very narrow lumen in one of its branches,
coming from the anther tissue (500-1).
Fig. 9. The same but with the branches bending together (900-1).
7ig. 10. The same, the end curving in for the formation of the ball (500-1).
Fig. 11. Separated gelatinous thread with numerous ramifications (900-1).
Fig. 12. Young spore ball with the beginnings of four spores, visible in a plane,
surrounded by gelatinous threads run into one another, with small parts of the
contents glimmering through. The young spores have been separated by the application of a strong solution of iodine (500-1).
Fig. 13. Young spore balls with a few spore beginnings (200-1).
Fig. 14. The same, but with a greater number of spore beginnings (200-1).
Fig. 15. Developed spores, surrounded with a thinned layer of the gelatinous
thread (900-1).
Fig. 16. The same, at the same time two separated spores are seen in connexion
with the ball (300-1).
Fiqs. 17 to 19. Separated spores in different degrees of developement (300-1).
Fig. 20. Part of the anther filament cut lengthwise, with distinct centripetal spore
formation (70-1).
FPig. 21. Various ripe spore balls; the gelatinous layer has almost disappeared. from
the ripest (200-1).
Fig. 22. Almost ripe, somewhat long spore ball, from which the gelatinous layer
has disappeared (200-1).
Dig. 23. Blossom stalk of the Saponaria officinalis, with ripe spores in the three
large, closed blossoms, m tea others, spores of different degrees of ripeness (natural sized.
PLATE V.1- GERMINATION OF THE SPORES.
FIG. 1, 2. TILLETIA CARIES, TUL., FROM WHEAT.
Fig. 1. Promycelium at the commencement of the germination; fifty hours from
the seed (500-1).
Fig. 2. Separation of the promycelium; fifty-two hours2 (450-1).
FIG. 3-6. USTILAGO MAYDIS, LEV. (500-1).
Fig. 3. Promycelium with a projection; twenty-four hours.
Fig. 4. Spore germinated later with a straight promycelium; forty-eight hours.
Fig. 5. The same with the promycelium bent in a knee form.
Fig. 6. Formation of a lateral sporidium (after twenty-four hours).
1 In plates v and vi, the thickness of the episporium is not shown.
2 Indicates in all cases from the seed.




FISCHER vo   J' WALDH~EIU --  STILA GI:NEf.              351
FIG. 7-15. USTILAGO CARBO, TUL., FROM OATS.
Fig. 7. Germinating spore; four hours (500-1)
Fig. 8. Promycelium with a projection on the side; eighteen hours (900-1).
Fig. 9. The same, with a bending of the promycelium  immediately upon its
coming from the spore (900-1).
Figs. 10 and 11. Other forms of bent promycelia; eight hours (900-1).
Fig. 12. The promycelium divided in two at the point, on the side a sporidium is
developed, and another promycelium coming from the spore; forty-eight hours (900-1).
Frig. 13. Two promycelia, the longest with two sporidia; forty-eight hours (900-1).
Fig. 14. Promycelium with three lateral sporidia and one terminal; forty-eight
hours (900-1).
FPig. 15. Thread formed lengthening of the promycelium; a spore germinated partly
in contact with air (500-1).
FIG. 16-20. USTILAGO CARBO, TUL., FROM BARLEY.
Fig. 16. Crooked promycelium with two lateral sporidia, and the beginning
of a third; forty-eight hours (900-1).
Fig. 17. Promycelium with a crooked sporidium; forty-eight hours (900-1).
Fig. 18. Promycelium with three lateral sporidia, one of which is on the projection.
F1ig. 19. Promycelium with a terminal sporidium, and coupled with one that has
fallen off; fifth day (900-1).
Fig. 20. Crooked promycelium with a lateral sporidium bending inwards, and the
beginnings of two others; fifth day (900-1).
FIG. 21-41. USTILAGO CARBO, TUL. FROM ARRHENATHERUM ELATIUS.
(Fig. 21-40, 900-1; fig. 41, 500-1.)
FRig. 21. Germination, twenty-four hours, with sufficient water.
Fig. 22. The same, with an insufficient amount of water.
Fig. 23. The same; from the thin promycelium proceeds a thread-like utricle.
Fig. 24. The same; with two lateral sporidia bending toward each other.
Fig. 25.'The same; with a longer promycelium, and with a projection and lateral
sporidium.
Fig. 26. Developement of a secondary sporidium; thirty hours.
bFig. 27. Promycelium with a lateral and terminal sporidium; thirty hours.
Fig. 28. Promycelium with a terminal and two lateral sporidia and thinned end;
thirty hours.
Fig. 29. Unusual form of a forked promycelium; thirty hours.
Fig. 30. Peculiar formation of the sporidia,.which seldom happens; thirty hours,
in darkness.
Fig. 31 Promycelium with secondary sporidium formation; forty-eight hours.
Fig. 32. Considerable lengthening of the lateral sporidia; forty-eight hours.
Fig. 33. Secondary sporidium formation from a considerably lengthened lateral sporidium; forty-eight hours.
Fig. 34. From the spore comes a second promycelium with a terminal sporidium;
forty-eight hours.
Fig. 35. Promycelium, with a long, serpentine, lateral sporidium, which comes in
contact with the projections; forty-two hours.
Fig. 36. Bow-shaped, lateral sporidium, whose end has grown to the promycelium;
forty-two hours.
Fig. 37. Promycelium, with two such sporidia; forty-two hours.




352         NEw  YORK S TATE AGRICUL TURAL SOCIETY.
Pig. 38. Serpentine sporidium vertical to the promycelium, with a secondary sporidium on the point; forty-two hours.
PFg. 39. Another form of a bow-shaped sporidium; fourth day.
Fig. 40. Spore with three promycelia, on two of them spore formation; fifth day.
Fig. 41. Spore germinated in contact with air; seventy-two hours.
FIG. 42-46. USTILAGO  LONGISSIMA LEV., FROM GLYCERIA FLUITANS.
Fig. 42. Promycelium with a developed sporidium; eight hours (300-1).
Fig. 43. Promycelium, with a spindle-shaped sporidium; twenty-four hours (400-1).
Fig. 44, a, b, c. Various forms of lateral spdridia which, in connection with the
promycelium originated in air; forty-eight hours (300-1).
Fig. 45, a. A second promycelium grows from  the spore; b, sporidium, with a
secondary on the point. Both cases, in connection with the promycelium, originated
in air; forty-eight hours (300-1.)
Fig. 46. Sporidium formation, resembling the sporidia of Tilletia, from the previous seeding (300-1).
FIG. 47-53. USTILAGO FLOSCULORUM[ Fi.
Fig. 48, 450-1; the remainder 900-1.
Fig. 47. Germination; six hours.
Fig. 48. Different stages, twenty hours; a, promycelium without sporidium; b,
promycelium with a terminal and two lateral sporidia; c, promycelium, with a terminal and three lateral sporidia.
Fig. 49. Different stages, second day; a, promycelium  divided into four distinct
parts, with four small vacuoles and a terminal sporidium; b, the same, with four distinctly marked divisions and large vacuoles.
FTg. 50. Promycelium with secondary sporidium formation; twenty-four hours.
Fig. 51. Promycelium with secondary and tertiary sporidia formation; thirty-nine
hours.
Plg. 52. Promycelium with numerous sporidia; third day.
Fgiy. 53. Promycelium with numerous secondary sporidia; thirty-nine hours.
PLATE VI — GERMINATION OF THE SPORES.
FIG. 1-21. USTILAGO ANTHERARUM, FR., FROM DIANTHUS CARTHUSIANORUM.
Fig. 2-21,900-1.
Fig. 1. Germination; a, seven hours (500-1); b, promycelium with bright little oil
drops; eighteen hours (300-1); c, somewhat later conditions with a distinct lateral
knob (500-1).
Fig. 2. Different conditions: a, promycelium with a contraction and lateral sporidclium; b, promycelium with a contraction and two lateral sporidia; c, promycelium
with four vacuoles and two sporidia.
Fig. 3. Separated promycelia; forty hours. All show a distinct knob; a, with four
small vacuoles; b, with two large and two small vacuoles; c, after the application of
a solution of chloride of zinc and iodine, the promycelium is broken up, except at
the place of the knob.
Fig. 4. Promycelium fallen off with an unusually well-developed knob; seldom the
case; forty hours.
Fig. 5. Two sporidia in terminal copulation; forty hours.
Fig. 6. Two sporidia in lateral copulation; forty hours.




FISCHER  VON WVALDHEIE  - USTILA GINVE~.                 35 3 
Fig. 7. Promycelium in lateral copulation with a sporidium; forty hours.
Fig. 8; Lateral copulation of the promycelium with the point of the sporidium;
third day.
Eig. 9. The same, but the copulation of the sporidia is lateral, the promycelium still in connection with the spore.
Fig. 10. A promycelium which has fallen off, with a gerinm thread.
Fig. 11. Two copulating sporidia with a long, crooked bridge; one of them clear
as water.
Fig. 12. Besides a pronlycelium with three sporidia, two others come out of the
spore (one of them with a sporidium); second day.
Fiqg. 13. The same with some modifications; fourth day.
PFig. 14. The same with secondary formations; fifth day.
Fig. 15. The same; another modification; fifth day.
Fig. 16. Two promycelium grown from the spore on opposite sides, with tertiary
spore formation; fifth day.
PFig. 17. Luxuriant sporidium formation on a promycelium which has fallen off; fifthl
(lay.
F-ig. 18. Promycelium in copulation with a sporidum out of which a secondary sporidium has developed itself; sixth day.
Fig. 19. Germinating sporidium still in connexion with the promycelium; eleventh
day.
Fig. 20. Germination of the promycelium; eleventh day.
Fig. 21. The same, but with two other promycelia grown out of the spore.
FIG. 22-26. USTILAGO ANTHERARUM, FR., FROMI SAPONARIA OFFICINALIS (900-1).
Fig. 22. Germinated promycelium; its upper part hyaline; twenty-four hours.
Fig. 23. Germination of the promycelium at the point, at the opposite end a sporidium. Vacuoles in the contents.
Fig. 24. Another form of lateral germ-thread, with a swelling at the point. In the
lower part of the promycelium, watery contents; forty-eight hours.
Fig. 25. The plastic contents of the promycelium have passed over into the side
tube; fifth day.
Fig. 26. Dichotomous division of the promycelium; at the end of the longer branch.
apparently a sporidium. The promycelium partly hyaline; fifth day.
FIG. 27-37. USTILAGO RECEPTACULORUM, FR., FROM TRAGOPOGON PRATENSIS.
(Fig. 27-30, 450-1; fig. 31-36, 900-1; fig. 37, 500-1.)
IFg. 27. Promycelium; twenty hours.
Fig. 28. Promycelium with two lateral sporidia; second day.
Fig. 29. Promycelium with three lateral sporidia, two of them in copulation at
their ends; second day.
Fig. 30 The same, but the sporidia lie cross-wise over one another in lateral copulation.
Fig. 31. Promycelium divided in four parts, and hyaline; fourth day.
Fig. 32. The same, but with three lateral sporidia, the under one of which is in
copulation with a sporidium which has fallen off.
Fiq. 33. Different conditions of copulating sporidia; fourth day; a, side copulation;
b, side copulation, but nearer to the point; c, side copulation, with a long, crooked
bridge; d, side-terminal copulation.
[AG.]         23




354         NEW  YORK STATE A GRWICULTURAL SOCIETY.
Fig. 34. Two hyaline sporidia in lateral copulation, one of which has germinated;
fourth day.
Fig. 35. The same, the sporidia, however, are in terminal copulation; the germthread is longer, and hyaline at the bottom.
Fig. 36. The same; the germ-thread very long, partly hyaline, and with cross walls.
Fig. 37. Two sporidia in lateral copulation, of considerable and unequal length;
fourth day.
FIG. 38-43. UROCYSTIS POMPHOLYGODES, RABH. FROM RANUNCULUS REPEINS (500-1).
Fig. 38. Promycelium with four sporidia; twenty-four hours.
Fig. 39. Out of one spore has grown a promycelium with three sporidia; out of
the other, two long tubes, through contact of the germ with water; twenty-four
hours.
Fig. 40. Promycelium dichotomously divided into long branches, which are partly
hyaline and partly septate, occurring through contact with water during the germination; twenty-four hours.
Fig. 41. Promycelium with three distinct sporidia; twenty-eight hours.,ig. 42. The sporidia are close together, and contain vacuoles; the promycelium
invisible; twenty-eight hours.
Fig. 43. From standing long in water, instead of there being promycelia, tubes
have appeared, which are scarcely to be seen; the under one iS already breaking up,
and is disorganized; fourth day.




t~~i,~  j.
4.~~~~~
/~~~~~~~~~~~~~~~~~~~~- C..
b~~~~~~~~~~~~~~~~~~~~o~,
5.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
/~~~~~~~~~~~~~~~~~~~~:"~..
0~~~~~~~~~~~~~~..








IL
= k f/' ~~t A c   -                       dIts=_L..    _   -'  A -'S,,       _~~- /        (;
//,f!,~..xr
_ C
ii   /I4"








722. ~  ~/.
X'2 "'i  ",    /          2
a:~~~~~~~~~~~/.  /3.,,
i'    i3         ~ rr-'-:   -  
/     5.
2- "S /                                      /9
42          5S0";                      it
~~~a~~~i~~~3, ~ ~
C.->,i;;429i~'                     t.
20.'J                       i-~~~~~~~
/                           U~~~~~~~~tr








w4'A
9/~~~~~~~~~~~~~~~1
/    eeS 7                    ie








V/~~~~~~~/
J~~~
II
/',                          v
17.~~~~~~~~.~~~~~7/ ~ ~ ~    2
/8ll 22/
I,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~J
27_ 2                                                      2:
1'; 
32.
d6,~ ~ ~ ~ ~ ~~~~~~8
424
4.'... 
r-               t:~~~~~~~~~~~~I Z — ~46.~~~~~~is
Q                )        i:.::.
iI~~~~~~~C:,.~::':t,.,~/~~~~~~~~~: i'~l
"0'~ ~?              JJ'    i,',.   i.3~~~~~~~~                  )
"%I           -.x
ep ~ ~ ~ ~ ~~..                         ~.~,~~~~~~~::
~~5'         a~~:'...          jO        2 a~- 
l~~~~j;    ~~~~~~~~~~   ~~~ -—,~~~O'








~L;z~~~~~~~~~F,aC
6'.     /       7a,:,C,,9.
e,3                                    i C'        ti'.           9.~~~~~~~~~~,3
10.'%            - 2.
16.~~~~~~7
K~~~~~~~
00~ ~~~~~~~~~~~~~~~~~ 
t                                                  2;:~~2
21.
/'. "   /:.:~    ~ ~~25.   t: i:'
2 7
33.~~~~~~~~~~~~~~~~~~~~~~~~l'.29  29                      C?7            {///
3''
41~~'~~~~~i0
9~l~                                    > 1 //oiv-;";"J~7~~~'
t,~.......?)._,,:~   ":.~        _~::i