Published March, 1972
Leaf Area-Leaf Weight Relationships in the Soybean Canopy1
H. R. Roller2
ABSTRACT
Leaf area (A) and leaf dry weight (LW) were deter-
mined throughout the growing season for lower, middle,
and upper main stem segments and branches of a field
soybean [Glycint: max (L.) Merrill] community. The
A/LW ratios were regressed on number of days after
planting, using polynomial regression models. For the
middle main-stem segment, A was also regressed on LW.
Mean seasonal A/LW decreased from the base to the
top of the canopy. Mean A/LW of branches was higher
than that of any main-stem segment.
The A/LW ralio of each main-stem segment continual-
ly declined until just prior to leaf senescence. The trend
subsequently reversed and A/LW increased rapidly dur-
ing leaf abscission. The trend in A/LW of the branches
differed in that there was an initial increase in A/LW,
followed by a decline and then a second increase during
leaf abscission.
Due to time related changes in A/LW, models for the
prediction of soybean leaf area from leaf weight should
also include time as a variable if observations are made
at different growth stages.
Additional key words: Glycine max (L.) Merrill, Ab-
scission of leaves.
ARIOUS methods are employed to estimate leaf
V area of crop plants. One reasonably accurate (4,
5, 8) and frequently used technique is based on the
ratio of leaf area (A) to leaf weight (LW). This ratio,
established by subsampling, is then used in conjunc-
tion with whole-sample leaf weight to estimate whole-
sample leaf area (8). Leaf weight is most often ex-
pressed on a dry basis, although fresh weight also has
been used (8, 9).
It is obvious that the accuracy of leaf area estima-
tion is dependent upon the accuracy of estimating the
A/LW ratio. However, to minimize the amount of
labor involved in determining leaf area it is desirable
to make as few determinations of A/LW as possible.
Prior knowledge of the factors that affect A/LW ratio
is useful in planning experiments in which this method
of leaf area estimation is to be used.
This paper reports the results of a study of the re-
lationship between A and LW in various parts of a
soybean (Glycine max (L.) Merrill) canopy through-
out the growing season. Time trends in A/LW ratio
are analyzed in terms of morphological development.
EXPERIMENTAL PROCEDURE
Leaf samples were obtained from 'Amsoy' soybean plants
grown in 1968 on the Purdue University Agronomy Research
Farm, near Lafayette, Ind. Seeding was delayed until June 9
due to excessive soil moisture. Row spacing was 76.2 cm and
within-row population averaged 18.5 plants/m. Natural rain-
fall was supplemented by approximately 5 cm of water, applied
by overhead sprinklers, on July 24 (full bloom stage). Canopy
closure took place about August 2. Moderate lodging occurred
on August 17. There were no significant weed, insect, or disease
problems, plant development appeared to be normal, and seed
yield was moderately high.
Samples were randomly selected from within a uniform area
of approximately 0.4 ha. The sampling unit consisted of a
single row of sufficient length to provide, on the basis of land
area, a 0.465-m2 sample. Sampling was carried out three times
per week during most of the growing season, with less frequent
sampling toward the end of the season. Three or four replicates
were taken on each sampling day.
Leaf laminae were grouped according to position on the
plant. All leaves from the branches were combined into a single
group. Leaves from the main stem were subgrouped on the
basis of consecutive four-node segments (designating the cotyle-
donary node as the first). Due to the normal small amount of
leaf development at nodes 1 through 4 and node 17 and beyond,
only three main-stem segments will be considered further: nodes
5 through 8, 9 through 12, and 13 through 16. These will be
referred to as the lower, middle, and upper main-stem segments.
Observations began as soon as the first leaf within a designated
segment unrolled. Leaf area was measured and leaves were
then dried to constant weight in a forced-air dryer at approxi-
mately 70 C and weighed. Leaf area measurements were made
with an optical planimeter, similar in design to that described
by Davis et al. (3). The planimeter was calibrated with soybean
leaf sections of known area.
The individual A/LW ratios were regressed on number of
days after planting. An appropriate computer program, em-
ploying the least squares method, was used to fit polynomial
functions (all terms in the polynomial being retained). The
regression model (degree of the polynomial) was determined
empirically. The coefficient of determination (R2) was calcu-
lated for each regression. The t-test was employed to determine
whether or not each partial regression coefficient differed sig-
nificantly from 0. In addition to the A/LW vs time regressions,
A was regressed on LW.
RESULTS
Relationships between A/LW and time in lower,
middle, and upper main-stem segments and branches
are shown in Fig. 1 through 4. Table 1 presents co-
efficients for the fitted polynomials and also gives the
number of observations and the R2 for each regression.
Successive leaves on the lower main-stem segment
(Fig. 1) unrolled 25, 29, 33, and 37 days after planting,
respectively. Leaf area index (LAI) of this segment
increased to 1.52 on the 46th day, remained constant
through day 60, then steadily declined to about 50%
of maximum by the 96th day. Rapid abscission then
began, and by the 105th day, all leaves had fallen.
On the middle main-stem segment (Fig. 2), leaves
unrolled on days 41, 45, 49, and 53. Maximum LAI
(1.43) was reached on the 62nd day after planting and
leaf area remained constant until the beginning of
1
Contribution from the Purdue University Agricultural Ex-
periment Station, Lafayette, Ind. 47907. Journal Paper No.
4491. Research supported in part by a grant from the Crop
Improvement Council, National Soybean Processors Association.
Received Feb. 21, 1971.
-' Assistant Professor of Agronomy.
 KOLLER: LEAF AREA-WEIGHT RELATIONSHIPS IN SOYBEAN CANOPY 181

rapid abscission on the 96th day. Leaf senescence was DISCUSSION

visible on day 92. All leaves had fallen by the l l0th Differences among ,segments in mean seasonal A/
day. LWratio are probably due to differing light intensi-
Leaves on the upper main-stem segment (Fig. 3) ties at various points in the canopy. Light intensity
unrolled on days 56, 60, 63, and 66. Maximum LAI
(0.55) was attained by the 70th day after planting. 4oo,
Leaf area remained constant until rapid abscission be-
gan on day 96. Leaf senescence was evident on the ~5c
92nd day and all leaves had abscissed by the ll0th day
after planting.
The first leaf on the branches (Fig. 4) unrolled ~oc
the 31st day after planting. Branch LAI increased to
0.94 on day 70, then steadily declined to about 50% ~
of maximum on day 96. At this time rapid abscission ~
.. ~5o
began and by the ll0tb day after planting all leaves ~
had fallen. 200
The relationship between A and LWin the middle
main-stem segment is shown in Fig. 5. Each point in ,50
Fig. 5 is associated with a specific point in the corre-
sponding A/LW vs time graph (Fig. 2). Two subsets %’0 io & ;o ~’o ;o ,do ,,’o
Days Mter Planting
of data in Fig. 5 are distinguished by different plotting
characters. The triangular points represent those ob- Fig. 2. Relationship between time and leaf area:leaf weight
ratio (A/LW)of the middle main-stem segment the soy-
servations made from the 71st day through the 94th bean plant. Eachpoint represents an individual observation,
day after planting. The round points represent all See Table 1 for regression parameters.
other observations. This subdivision of the data will
be used to illustrate a point to be made in the Discus-
300
sion section.
Table 2 presents coefficients and R2 for linear and v~
quadratic regressions of the entire set of data shown ~o ~o
in Fig. 5. Leaf area was designated the dependent
variable. ~.
Mean seasonal A/LW of lower, middle, and upper <) ~oo
main stem and branches was 255, 239, 222, and 388
cm2 X g-~, respectively. Tukey’s w-procedure was
used to determine that each mean is sign[icantly dif- ~o
ferent (P < .01) from all other tneans.
Days After Planting

Table 1. Estitnates of parameters of polynomial functions Fig. 3. Relationship between time and leaf area:leaf weight
selected to describe leaf area:leaf weight ratio vs time rela- ratio (A/LW)of the upper main-stemsegment of the soybean
tionships in various soybeanplant segments,fl~ is intercept; plant. Each point represents an individual observation. See
ft, fl..,, and~ are coefficients for first-, second-, andthird- Table 1 for regression parameters.
order terms.* Numberof observations n and coefficient of
determination R~ are shown.
Main stem
Lower Middle Upper Branches
’~ 50O
~30 3.38 x 101 -[,02 x 102 1.30 x IC 7.01 x 1(~
o-2,69
~I x 10 2.29 X 101 -2.7I x 101 5.08 x 101
-~
~ 1.93 x 10 -4.14 × 10-~ -s 1.65 x 10-~ -7.48 x 10-~
.......... 2. t8 × 10 .......... -~
~3n3 3.53 x L0
1(12 75 45O
53 90
R~ .30 .58 .66 .46
All partial regression coefficients (/~l , ~32, ~a) differ significantly fromzero at

3501

%’0 ~’o 2o io io ~b ob ~’o ,& go

Days After Planting %’0 ~’o do ;o ~’o ~ ,do ,,;

Fig. l. Relationship between time and leaf area:leaf weight Fig. 4. Relationship between time aud leaf area:leaf weight
ratio (A/LW)of the lower main-stemsegment of the soybean ratio (A/LW)ol the branch segment of the soybean plant.
plant. Each point represents an individual observation. See Eachpoint represents an individual observation. See Table
Table 1 for regression parameters. 1 for regression parameters.
189 CROPSCIENCE,VOL. 12, MARCH-APRIL
1972

Table2. Estiinate:~of parameters frompolynomial regressions 22

of leaf area on leaf weightfor the middlemain-stem segment
of the soybean
plant,fl0 is intercept;fll andf12 arecoefficients
for first- andsecond-order terms.*Coefficientof determina.
tion R~ is shown. 2O
Regression model
Linear Quadratic
~ ~
00 i.52x 10 3.54 x i0 18

* All partial regression coefficients (~, ~2 ) differ significantly from

decreases rapidl~ with depth in the soybean canopy

(7). Loweringthe light intensity is knownto increase
A/LW(1, 2). Therefore, an increase in A/LWtoward
the base of the canopy was not nnexpected. The data
suggest that leaves arising from the branches developed
under lower light intensities than lower main-stem
leaves. This is probably due to the fact that main-
stem leaves, during their early development, were
near the top, unshaded part of the plant, while most
branch leaves were continually shaded. A related
phenomenon that may account for part of the in-
creased A/L~¥ of branches and lower main stem is
leaf senescence. During much of the sampling period,
e
a proportion of the leaves on these segments was sen- o

escent and therefore very high in A/LWratio (note in

Fig. 2, 3, and 4 the rapid increase in A/LWfollowing
senescence).
There was a continuous decline in A/LWof each
main-stem segment until just prior to extensive leaf
senescence (Fig. l, ~, and S). At this point the trend
reversed and A/LWincreased rapidly during the pe-
riod of leaf abscission. The continuing decline in A/
LW following attainment of maximum LAI must be
due to increasing weight per nnit leaf area. Since A/
LWsubsequently increased, at least part of the weight
increase was probably in the form of materials that
were later retrans]ocated from the leaf.
The concavity of the A/LWvs time relationship
becomes more pronounced toward the top of the plant
(cf. Fig. 1, 9~, and 3). Leaves lower on the plant failed
to increase in weil~ht to the extent of leaves in richer
light environments. Therefore, A/LWratio of lower
leaves was more uniform th~:oughout the sampling
period.
The initial rise in A/LWof the branch leaves (Fig.
4) is unique. Thi~ increase in A/LWcan be partially
attributed to the continual initiation of new leaves,
which have high A/LW ratios. Furthermore, these
new leaves were probably developing under light in-
tensities that were continually decreasing as a result
of increased shading as the canopy developed. The
subsequent decline in branch A/LW, followed by an
increase, can be explained iu the same manner as for
the main-stem seg~ents.
Robison and Massengale (6), working with alfalfa
(Medicago saliva L.) at various growth stages, re-
gressed A on LWusing simple linear regression. Cor-
relation coefficients ranged from .90 to .98. Regression
coefficients did not differ significantly due to growth
stage. It was concluded that leaf weight is a useful
predictor of leaf area in alfalfa.
The linear regression of A on LWfor the middle
main-stem segment of the soybean (Fig. 3) is statistical
ly significant (P (.01) and ~ ~.9 1. Ho wever, th e
2.;
go ~’o .o
I
,oo
I
*~o
LW,g
points are those observationsfromthe 71st throughthe 94tb
days after planting. Roundpoints representall other ob-
servations.
second-order term in the quadratic regression is also
statistically significant, although R2 increases only to
.92 (Table 2). The curvature, as well as a substantial
proportion of the residual sums of squares, can be at-
tributed to nonconstancy of A/LWin time (Fig. 2).
This is graphically illustrated by arbitrarily separating
the data of Fig. 5 into two subsets. The triangular
points are those observations from the 71st through
the 94th days after planting, a period of relatively low
A/LWvalues (Fig. 2). The round points are all other
observations. The triangular points are clustered to-
ward the right of the scattergram as a result of their
lower A/LWratios. This shows that their distribution
is not random, but rather is time dependent. Similar
results were obtained for other segments, but are not
presented.
It is apparent that prediction models for soybean
leaf area should include time as a variable if observa-
tions are made at different stages of growth. Alterna-
tively, if all observations are made at approximately
the same time, the simple linear regTession of A on LW
is probably adequate.
 CARLETON & COOPER: SEED SIZE AND SEEDLING VIGOR IN LEGUMES 183

ACKNOWLEDGMENT
The helpful advice of Dr. W. E. Nyquist on statistical aspects
of this study is gratefully acknowledged.