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MICROCOPY RESOLUTION TEST CHART
NATIONAL BUREAU OF STANDARDS - 1963
I
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PE
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MYMPTON *
ORNL - P 3082
Coxf-670503712
JUN 2 2 1965

MASTUCES Reichs
CFSTI RRICES
CARBON-14 CYCLING IN THE ROOT AND SOIL COMPONENTS H.C. $ 2.00 MN 65
OF A PRAIRIE ECOSYSTEM?
Roger C. Dahlman.
Radiation Ecology Section, Health Physics Division
Oak Ridge National Laboratory, Oak Ridge, Tennessee?
. : Clair L. Kucera. i
Professor of Botany, University of Missouri, Columbia, Missouri
TE
Ir.
ABSTRACT
*
Tall grasses of a central Missouri native prairie were labeled with
24co, in polyethylene enclosures during late summer in order to study carcon
transfer from foliage to the roots and from roots to soil organic matter.
81x-week-old regenerated foliage age imilated six to eight times more of the
mitocarbon thar follage which was mormodlNG Matulty. AirAN LMA 40.
and 85 percent of the assimilated label was translocated to the root system
in regenerated and mature plants respectively. Similar quantities of radio-
carbon were present in roots collected from both mature and regenerated
areas because of differential translocation processes. Over a two-year
period, -*c disappearance from the 0-10 inch depth of the root system
followed a linear trend as described by the regression Y - 27.1 - 0.528,
This investigation was supported in part by federal funds, NSF 702 ani
NIH predoctoral fellowship. USPH-5-71-CM29-091-02.
operated by Union Carbide Corporation for the U. 8. Atomic Energy

1
Commission.
LEGAL NOTICE
This report m. propered as an account of Government sponsored work. Neither the United
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A. Makos may warranty or repronontation, exprouand or implied, with respect to the accu-
racy, completeness, or unetulness of the information contained in this report, or that the wo
of any information, apparatus, method, or proces. dioolond to the report may not infringe
printoly owned righto; or
B. Assumes any liabilides with respect to the use of, or for denuo ruouting from the
un of any information, apparatus, method, or procede disclosed in this report,
As vand in the abovo, "person sottog og behalf of the Commission" Includes way om-
ploys or vontractor of the Commission, or employ of much contractor, to the extent that
buch employs or contractor of the Commission, or employee of such contractor preparo,
i dienominates, or provides accous to, any information pur minnt to do employmeat or contract
1, with the Commission, or his omployment with such contractor,
DESTROUTION OE THIS: QOCUMENT IJ ONLIMITED
--
----
---
1 Where Y 1s radioactivity and X is time in months. This relationship suurests
that the root system undergoes turnover approximately every four years. Roca
deeper in the profile exhibited a slower rate of -*c disappearance, and & I
near-equilibrium condition between loss to the soil and input from trans-
location was approached. Maximum cumulative transfer of -4c from the roots !
to the soil was observed 10 to 14 months after assimilation of the label
7 when approximately 45% of the cumulative 24c loss from roots was present in
the soil organic matter. Fourteen to 26 months later the -4c present in tre
soil decreased to 9% of the cumulative 1063 from roots, indicating that most
10 of the early decay products were rapidly decomposed by the soil microbes.
11 Both the rapid transfer from roots to soil and the large fraction of decay
products which disappeared as co, indicate a rather short residence time for
inost organic materials in these parts of the carbon cycle.'
-
-
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UCN•3507
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INTRODUCTION
Decaying fibrous crass roots supply much of the organic input 20%
carbon accwulation in grassland soils (Weaver et al. 1935, Goedewielen
and Schuurman 1950, Broadbent 1957, Konuro'r 1961). Carbon transfer processes
in situ, involving residues from functioning and dying roots, are au
integral part of humification. If decomposition increases in proportion to
the quantity of residue present in the soil (Jenny, et al. 1949, Bartholomew
and Kirkham 1960), new organic matter equilibria develop after decomposition
balances the input. Despite attempts to quantify carbon cycling (Olson 1963,
Russell 1964, Jenkinson 1966), additional information is needed about transfer
parameters and residence time of carbon in different compartments of the
root-soll system. The principal objectives of the study reported by this
paper were to apply . *Label to a fraseland econyutem and evaluate
|(2) carbon assimilation, transfer and residence time in the intact root and
shoot system, (2) transfer between the root and soil compartments, (3) the
1residence time of the radiocarbon 10 the soil organio mattor from these
18 results, the biological ball-life was estimated for carbon in the root apa
BOLl compartments of the ecosystem. In practice, this estimate would apply
20 to the rate of turnover of both labeled and wlabeled carbon.

STUDY AREA
The study was conducted at the University of Missouri Prairie
Research Station located in central Missouri. The native grassland 18
situated on the Putnam Mexico catena, a s11t loam sequence derived from 2008
XKrusekoph and Scrivoer 1962). Numerous studies have characterized the soil,
UCN.ssor
1

particularly, mineralogical and chemical properties (Whiteside and March£22
1944), organic matter (Jenay 1930), nátrocen reserve (Mullar 1947), effect of
grazing on soil structure (Kucera 1958). The soil retains the characteristics
of the grass developed profile because the sod has never been broken for
cultivation. Dominant grass species are big bluestem (Andropogon Cerardi
Vitman), little bluestem (Andropogon scoparius Micx), and Indian grabs
(Sorghastrum nutans (1) Nash) according to floristic analysis by Drew (1947).
Dropseed (Sporobolus hererologis A. Gray) persists in localized areas.
Administrative control by the University Insured limited access to the
experimental area and minimized potential health hazard from the persistent
14c in the ecosystem. . .
METHODS
Prairie grasses were labeled with -*c in situ under field conoitions.
1s Important considerations at the time of labeling were attainment of a suit-
able specific activity with a single exposure, minimal disturbance of the
plants and site, and retention of the c-photosyothate in structural or
storage compounds of the root system. . since storage of carbohydrates
(Aldous 1930, McCarthy 1938) and active root growth (stucky 1942, Browa 1942)
occur during the late growing season, phenologically mature plants were
21 selected in order to obtain the maximum quantity of -*c in the roots.
Application of 24c to Prairie Grass
Native gracı oí ten representative prairie areas was labeled with
radiocarbon 10 September, 1963. Five areas, each 6.632, had a cover of
rogenerated growth, and five were walipped arons on which maturing vegeta-
tion was present. Polyethylene enclosures were placed over the areas inside
which **co, was released for assimilation by the grasses. The apparitus ued i
labeling procedure are described elsewhere (Dahlman 1967, Kucera and Datina
1967). Grasses were exposed to the *co, atmosphere for a 61x-hour period
from 11:00a.m. to 5:00 p.m. Approximately 1 mCi was released inside the
enclosure, providing 0.151 m01/sna. Air temperatures inside the chamber were
approximately 10° higher than ambient conditions (85-93° F). Shortly after
release of the 14c0g, water vapor condensed on the inside walls of the flim,
Indicating a saturated atmosphere. There was no attempt to modify the humida
ity por to cool the inside atmosphere. .
Collection of Plant Biomass :
To insure maximum translocation of the “*c-products to the root system,
12 foliage was collected only after the first killing frost in November.
Foliage was clipped, dried and weighed to determine end-of-year standing crop
after the 1963, 1964 and 1965 growing seasons. A 200 8 sample was randomly
selected from each plot harvest for radioassay. For mid-year production
16 estimates and radioassay, foliage was collected from 1/4 m? of each labeled
17 area in June of 1964 and 1965.
Roots were collected at 4-month intervals from November 1963 to November
1965 according to procedures of Dahlman and Kucera (1965) and Dahlman (1967).
Soll cores, containing the roots, were sectioned into 0-25, 25-40 and 40-70
em segments .corresponding with the respective Amo A, and B, horizons of the
profile. The A, horizon sample was further subdivided into o-10 and 10-25 cm
increments to study in more detail the horizon which contained over 80% of
the roots. 8012 was washed from the roots with a forced spray apparatus,
modified after Fribourg (1953).
Radiocarbon Assay of Plant Biomass
Uniformly ground plant material was assayed with a window-type
3 (1104c/cm) proportional detector according to a modification of O'Brien azd
Warálow's (1961) procedure. Witi standardized manipulations, radiotecay of
plant particles at saturation thickness gave consistent results. Several
conversion factors were derived to express the results in terms of specific
activity per unit of constituent carbon (Dahlman 1967).
Analysis of Soil Organic Matter
Soil samples were collected from each area and the organic matter was
10 examined for the presence of c. The first samples were taken in Truly of
the year following labeling of the root system, and thereafter a soil core
was collected concurrently with the roots. Root material was removed and the
soil was washed by a series of decantations util no plant fragments were
observable with 30 power magnification. Carbon was collected from the
15 washed soil by wet combustion (Allison 1965), and the carbonate was radio-
16 assayed by Liquid scintillation (Nathan, et al. 1958).
RESULTS
Assimilation of 14c
Eight weeks after the ***c was released for photosynthetic Incorporation
32 and 22% of the activity was detected in the biomass of the regenerated and
mature areas respectively. The more actively photosynthesizing foliage
accounted for greater assimilation, over 50% of which was translocated to
the root system. With the nearly mature foliage, the majority (80%) of the
assimilated *c was found in the root system. With both treatments, the
balance of the released 240, that not detected in the biomass, was 20st to
MSAVU *****
.
.
the atmosphere. Distribution of radiocarbon in shoots and roots 8 wecks
after assimilation are shown in Table I. The results were expresscd per wit
of biomass and per unit of soil surface in order to examine the respective
transfer processes within the intact plant and between plant and s011
components of the ecosystem. The uC1/? value for roots represented an
average from four differeri root increments. Calculation of this expression
was first performed for different root increments, and the above values are
& summation for the whole root system.
Quantity of biomass and activity per gram of foliage were relatively
10 consistent among each group of similarly labeled areas. The physiological
condition of the foliage affected the initial assimilation of the "+co.
because regenerated shoots contained five times more activity than those from
13 mature areas. A statistical treatment of the dissimilar results gave an
| LSD o value of 8.22 while the actual diffurence was 14.9 u21/m.
15 Differential retention of 1*c is the shoots was probably the result of
16 utiliza coa of new photosynthetic products in the structural compounds of the
actively growing foliage. With mature grass, however, a relatively greater
18 quantity of the total assimilated. *c was translocated to the root system
19 during accumulation of organic reserves in the underground component of the
20 plants. Similar quantities of radiocarbon were present in roots collected
from both mature and regenerated areas. While less *c was assimilated Lo
the mature foliage, a greater proportion of the photosyathate was translocated.
to the root system.
Radiocarbon Changes in Root Biomass as a function of Time
Disappearance of radiocarbon from the root system over a 2-year period
18 presented as an average of the ten labeled areas (Fig. 1). The results
WONDO
CY!
R14... P
u
uilo
NILUX
-
-
-
-
-
-

are expressed for different depth increments of the root system, and the
graphic trends of the 25-40 and 40-70 cm segments are on an expanded scale
compared with the 0-25 cm depth. These differences in scale should be
considered when comparing Pluctuations in the activity of the root incre-
ments. Brackets show the range of variability about the mean of ten samples
at the 95% confidence level. For the 0-25 ora increment of roots, this
range included from 20 to 50% of the mean, and it was difficult to determine
significant differences between seasonal collections, especially when the
average root activity values differed from each other by 1-5 uC1/. .
Fluctuating root biomass, and "hot or cold" radioactive spots in the root
system probably contributed to the variable expressions of root radiocarbon
content.
13 In the 0-25 cm increment, there was a distinct decrease in **c content
14 as a function oť time (Fig. 1). This trend is described by the linear
15 regression equation, Y = 27.1 - 0.52x, where Y is radiocarbon content and X
is time in months. According to this relationship, the time required for
17 | the root syster radiocarbon to approach o 18 calculated as 52 months, or
slightly more than four years. This rate of 1088 implies a biological ballo
211e of two years, and it should characterize the general root turnover
pattern, assiming that metabolic discrimination between 12c and 240 16
21 negligible.
Radiocarbon disappearance occurred very slowly for roots 'deeper in the
profile. Over the two-year period, the net differences between initial and
24 final values were negligible. Abest fit of the trend for the 40-70 cm
lnorament nearly parallels tha abeciosa.
I
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3
Transfer of 4c from Rooto to Soil
Accumulation of "*c in soil and corresponding losses from roots are
presented in Table II. Root radiocarbon losses were calculated as the
differences between the average initial quantity (28.8 uCi/m?) and that
present at the time under consideration. Maximum transfer of **c to the soil
occurred between 10 and 24 months following fixation of the label in the root
system. About 45% of that which had disappeared from the 0-25 cm root
increments appeared as insoluble soil organic matter. Residence time of this
initial "pulse" is short because 4 months later only 9% of the cumulative
root loss was still present in the soil. The balance was estimated
as release of 24co, due to microbial respiration.
DISCUSSION
..
Respiration losses of *co, undoubtedly occurred between cessation of
labeling in September and the first measurement in November. From plausible
community respiration rates (Lundegardh p. 71, 1931; Odum 1959), probable

and mature follage areas respectively. Compared with the quantity of radio-
carbon present in the plant biomass, these losses were negligible and con-
tributed litti
total budget.
Distribution and compartmental transfer of 14c in the ecosystem were
sumarized after periods of eight weeks, one year and two years, (Figs, 2, 3
4 and 5 respectively). Turnover of roots in the prairie ecosystem, as
determined by *c disappearance, agrees remarkably well with other observa-
tions (Dahiman and Kucera 1965). It was concluded from both early and
current investigations that tine root system undergoes an average turnover
approximately once every four years. Weaver (1961), working with rebraskus
prairie, indicated that some roots may persist for 15-25 years, although
annual replacement of the root system was not studied. Short life of song
root parts evidently outweighs the long life of others.
Maximum transfer of radiocarbon from roots to the soil occurrcă duriry
late summer of the year following incorporation by the roots. This observa-
tion agrees with those of Stucky (1942) and Browa (1943) where active root
development occurred during the early growing season followed by disintegra-
tion and decomposition later in the summer.. During the second year oł the
10 present study, microorganisms apparently acted on the fresh soil organic
matter, and 80% of the soil radiocarbon disappeared, probably as “cog.
Both the rapid transfer from roots to soil and the large fraction of decay
products which disappeared as co, indicated a rather short residence time
for most organic materials in the humification part of the carbon cycle.
During the first year there was negligible change in the 24c content of roots
16 from the 25-70 cm depth. Still, -*c accumulated in the soil. It appears
17 that dowoward translocation of *c metabolites compensated for root losses
resulting from respiration and transfer to soil organic matter.
19 / A wide range in decay rates is expected for different parts of soil
organic matter (Jenkinson 1966). With organic matter at equilibrium, which
21 is the assumed case for native grassland: soils of central United States, the
results of the present study suggest anual decay rates of 0.020 and 0.004
for fresh amendments and more resistant humus respectively. These variable
decay rates agree with results of Jenkinson (1965) where both rapid and slow
decay processes were evident in soil cultures amended with **c ryegrass.
Radiocarbon dates of humus suggest an annual turnover rate of 0.003 for the
most stable parts of humus (Simonson 1959).
Table I
lhe
BIOMASS AND 4C ACTIVITY IN ROOTS AND FOLIAGE 8 WEEKS FOLLOWING
EXPOSURE TO 1400, IN SEPTEMBER, 1953
Area
Number
Biomass g/m
Foliage Roots* Total
Activity uči/g biomass
Foliage Roots Total
Activity 4C1/22
. Foliage Roots Total
Regenerated
· Foliage
-
1
0.155
.154
126
133
152
154
2039
1539
1965
1390
1772
2155
1672
2017
1544
1943
0.024
.019
.032
.011
.015
0.179
.173
.143
.11
19.6
20.5
16.8
20.6
19.5
19.4
.134
37.3 56.9
24.9 45.4
34.7 51.5
20.7. 41.3
33.0 52.5
30.2 49.5
.134
.011
.245
172
.114
.129
Average
Mature
Foliage
.018
.012
.025
.030
.013
.018
272
278
231
245
259
1372
2618
1402
3206
1292
2644
1896
. 1633
3451
1551
.020
017
4.9
5.0
4.7
3.5
4.2
4.5
16.3
34.6
23.2
38.5
25.2
27.6
21.3
39.6
27.8
42.0
29.11
32.0
.014
.016
..014
.015
.028
.031
20
Averace
Average for all arca
28.8
*hch free
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Table II
PRESENCE OF .240 IN THE SOIL ORGANIC MATIER COMPARED
WITH LOSS FROM ROOTS AS A FUNCTION OF TIME!
14C Present
72 Time Following
*C Incorporation
into Roots
(months)
in Soil Organic
- Matter
(uC1/m2)
Loss of 14c
from Roots
(uci/m2)
Percentage of
Root Loss
Present in
Soil
0-25 cm Increment
0.13 :
.
5
(2.6 .
5.0
3.02
6.4
59
16o4
6
0
.
..
9.9
...39
13.4
25-40 cm Increment
0.03
gain
0.8
3.8
n
.
2.2
.
18:
11.2
1.
.
6
.07
7.3
6
:
0.2
25.0
.08
40-70 cm Increment
0.05
gain
.23
gain
.05
.58
6
Do
.23
200.0.
vom..
van
.
.
7
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---
Aldous, A. E. 1930. Relation of organic food reserves to the growth of some
Kausas pasture plants. J. Amer. Soc. Acron. 22: 385-392.
3 Allison, L. E. 1965. Organic Carbon. In Methods of Soil Analysis: Cherical
and Microbiological Properties. Vol. II: pp. 1367-1378.
Bartholomew, W., V. and Don Kirkham. 1960. Mathematical descriptions and
. interpretations of culture induced soil nitrogen changes. 7th Int.
Congr. Şoil Sci., III (2): 471-477.
8 Broadbent, F. E. 1957. Organic matter. In Soil, Yearbook of Agriculture,
91 2957. pp. 151-157.
10 Brown, E. Marion, 1943. Seasonal variation in the growth and chemical
composition of Kentucky. bluegrass. Mo. Agric. Expt. Sta. Res. Bull.
No. 360. 56pp.
Dahlman, Roger c. 1967. Carbon turnover and transfer studies on *c labeled
roots in a prairie ecosystem. Unpublished Pa. D. thesis. Univ. of
Missouri. 208 pp.
16 Dahlman, Roger C, and Clair L. Kucera. 1965. Root productivity and turnover
in native prairie. Ecol. .46:. 84-89. .
18 Drew, W. B. 1947. Floristic composition of grazed and ungrazed prairie
---------
...
Fribourg, H. A. 1953. A rapid method for washing roots. Agron. J. 45:334-335.
Goedewaagen, M. A. J. and J. J. Schuurman. 1950. Root production by agricul- i
tural crops on arable land and on grassland as a source of organic
: matter in the soil. 4th Int. Congr. Soil Sci., Trans. pp. 28-31.
Jenkinson, D. S. 1965. Studies on the decomposition of plant material in
soil. I. LOOBCs of carbon from 24c labeled rye grass incubated in soil
in the field. J. Soil. Sci. 16 (1): 104-115.
24
...
Jenkinson, D. S. 1966. The turnover of organic matter in soil. In The use of
1sotopes in soil organic matter studies. Rept. FAO/IAEA. Tech. Meeting.
Brunswick, 1963. pp. 287-197.
Jenny, Hans. 1930. A study on the influence of climate upon the nitrogen and
organic matter content of the soil. Mo. Agric. Expt. Sta. Res. Bull.
No. 152. 66 pp.
Jenny, Hans, S. P. Gessel and F. T. Bingham. 1949. Comparative study of
decomposition rates of organic matter in temperate and tropical regions.
S011 Sci. 68: 429-432.
Chernozen under different agricultural uses. Soviet Soil sci. 1961 (5):
533-537.
Krusekoph, 8. H. and C. L. Scrivner. 1962. Soil survey of Boone County,
Missouri. U. 8. Govt. Printing off. Washington, D. C. Ser. 1959, No. 12.
Kucera, C. L. 1958. Some changes in the soil environment of a grazed prairie
community in central Missouri. Ecol. 39: 538-540.
Kucera, C. L. and R. C. Dahlman. 1967. Uses of -4c to evaluate organic
turnover 10 a prairie ecosystem. Submitted to Ecology, October, 1966.
Lundegardh, Henrik. 1931. Bavironment and Plant Development. Trans. Eric
Ashby. Edward Arnold Co.: London, 325 pp.
McCarthy, Edward C. 1938. The relation of growth to the varying carbohydrate
content in mountain brome. V. 8. Dept. Agric. Tech. Bul. No. 598. 24 pp.
Millar, M. F. 1947. studies in soll nitrogen and organic matter maintenance.
Mo. Agric. Expt.,sta. Res. Bull. No. 409. pp. 1-32. '.
UCN-2007
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::
1
Nathan, David G., Jack D. Davidson, Jeanne G. Waggoner and Nathaniel I. Berlin.
1998. The counting of barium carbonate in a liquid scintillation spec-
trometer. J. Lab. Clin. Med. 32: 915-917.
O'Brien, T. P. and I. J. Wardlow. 1961. The direct assay of **c in dried
plant materials. Aust. J. Mol. Sci. 14: 361-367.
Odum, Eugene P. 1939. Tumdamentals of Ecology. W. B. Saunders Co.: Philadel-
| phia. Shố Tp.
Olson, Jerry 8. 1963. Energy storage and the balance of producers and
decomposers in ecological systems. Ecol. 44 (2): -322-331.
Russell, J. 8. 1964. Mathematical expression of seasonal changes in soil
organic matter. Nature 204: 161-162.
Simonson, Roy W. 1959. Modern concepts of soil genesis, a symposium. 8012
sei. Soc. Am., Proc. 23: 152-156.
stucky, I. 8. 1941. Seasonal growth of grass roots. A. J. B. 28: 486491.
Weaver, J. 8. 1961. The living network in prairie soils. Bot. Car. 123: 16-
28.
Weaver, J. E., V. H. Haugen and M. O. Weldon. 1935. Relation of root distri-
bution to organic matter in prairie soll. Bot. Gaz. 96: 389-420.
Whiteside, E. P. and C. E. Marshall. 1944. Mineralogical and chemical studies
of the Putnam silt loam soil. Mo. Agrio. Bapt. Sta. Ros. Bull. No. 386.
48 pp.


Figure 1. Averare *c present in different root increments. Brackets
represent the range of variability about the mean of 10 samples. Note the
expanded scale for the lower curves.
Figure 2. Carbon-14 distribution in the regenerated foliage areas of the eco-
system in November, 1963, 8 weeks after assimilation. Activity is expressed
in uči/m, and percentage distribution is in parentheses. Loss of +4C as
co, is attributed to plant and/or microbial respiration.
24
Figure 3. Carbon-14 distribution in the mature foliage areas of the eco-
system in November, 1963, 8 weeks after assimilation.
10 Figlire 4. Carbon-14 distribution in the ecosystem in November, 1964, 1 year
after assimilation. *Represents retranslocated or recycled -4c. **Removed
as litter prior to initiation of current growth.
13 Figure 5. Carbon-14 distribution in the ecosystem in November, 1965, 2 years
after assimilation. *Represents retranslocated or recycled -4c. **Removed
as litter prior to initiation of current growth.
ORNL-DWG 67-4052

30
T
0-25
cm
ACTIVITY IN μCi/m?

25-40
------I-
NOV | MAR JUL
1964
I
- -
NOV | MAR
40-70
-
JUL NOV |
1965
ORNL-DWG 67-4051

ESTIMATED COMPARTMENT
LOSS AS CO2 LOSS
19.4
A
1-10
12-17%)
1-10
L
'intensive
(2%)
ht.av
manca
17
1
TRANSFER
U14
-
-
.' -
-
-
- -
-
ORNL-DWG 67-4050

ESTIMATED
LOSS AS üüa
COMPARTMENT
LOSS
7)
(12%
come
CV
B
ay of the RF
NOT MEASURED
NOT MEASURED
TA1
ORNL-DWG 67-4049

ESTIMATED
LOSS AS CO2
COMPARTMENT
LOSS
NEGLIGIBLE
19.4**
4.4**
2.6
(9%)
5.9
TRANSFER
TO SOIL
News
(14%).
NO LOSS
Lys
daa
+
-
-
-
7
.
-
.
2
.
WWW.
IN

-
0144
.
.
ORNL-DWG 67-4048
-
1-
ESTIMATED
LOSS AS CO2
COMPARTMENT
LOSS
.
VALI
VAN
NEGLIGIBLE
0.6**
WW10.4*KLA
1
(18%)
I
...
.
m
8.3
(29%)
.
i ns
-
.
--
ineither.
ir
.
-
-
.
G
:
14.3
TRANSFER
TO SOIL
hat
10.7
(2%)
3.4.
END



DATE FILMED
8 / 7 /67







.