Shade and Temporal Distribution of Pod Production and Pod Set in Soybean
Dennis B. Egli* and William P. Bruening
ABSTRACT
The temporal distribution of pod production and pod survival play
an important role in determining pod and seed number in soybean
(Glycine max L. Merrill). We investigated the effect of changing
photosynthesis at growth stage R1 (beginning flowering) on these
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
temporal distributions in two greenhouse experiments. Plants (‘Elgin
87’) were exposed to two levels of shade (60 and 90%) from growth
stage R1 to maturity. Other plants were removed from 90% shade
or placed under 90% shade midway through flowering (transfer treat-
ments). Temporal distributions of pod production and pod survival
were determined by marking all unmarked pods ⱖ 10 mm long on
plants every three days with different colored paint. The color of
paint on the mature pods identified when they started development.
Continuous shade reduced mature pods by 27 (60% shade) and 82%
(90% shade), but it shortened the pod-production period in only one
of four comparisons. Pod production responded quickly to transfer
treatments, and the mature pod load was always greater (nearly three
fold) than the continuous 90% shade treatment and less (average of
53%) than the control. The mature pod load failed to recover from
early shade because the increase in radiation did not lengthen the
pod-production period and not enough pods were produced. Pod
production was often more important than pod abortion in determin-
ing mature pod number. Adding the temporal distribution of pod
production and survival to models predicting pod and seed number
will improve their accuracy.
P ods and seeds per unit area are an important deter-
minant of yield in many crop plants including soy-
bean (Jong et al., 1982; Pandy et al., 1984; Egli, 1998;
Frederick et al., 1998). However, the mechanisms by
which the plant regulates the number of pods and seeds
it produces are not completely understood. Recent evi-
dence (Bruening and Egli, 1999, 2000) suggests that
the temporal distribution of flower and pod production
should be added to the traditional determinants of pod
and seed number—photosynthesis or assimilate avail-
ability and sink (seed) characteristics (Charles-Edwards
et al., 1986; Egli, 1998).
The asynchronous flowering characteristic of soybean
is well documented. Flowering periods (first to last
flower on a plant) are frequently 30 d long or longer
(Hansen and Shibles, 1978; Yoshida et al., 1983; Gai et
al., 1984; Dybing, 1994). The length of the flowering
period was sensitive to daylength (Guiamet and Naka-
yama, 1984) and planting date (Constable and Ross,
1988; Dybing, 1994), but CO2 enrichment (Nakamoto
et al., 2001), plant density (Torigoe et al., 1982; Saitoh
Dep. of Plant and Soil Science, Univ. of Kentucky, Lexington, KY
40546-0312. Published with the approval of the Director of the Ken-
tucky Agric. Exp. Stn. as paper 04-06-147. Received 21 Sept. 2004.
*Corresponding author (degli@uky.edu).
Published in Crop Sci. 45:1764–1769 (2005).
Crop Physiology & Metabolism
doi:10.2135/cropsci2004.0557
© Crop Science Society of America
677 S. Segoe Rd., Madison, WI 53711 USA
1764
et al., 1998), and N fertilizer (Torigoe et al., 1982) had
no effect. Less is known about the survival of flowers
to produce pods, but some results suggest that pod pro-
duction is equally asynchronous. Pods that survived to
maturity and contained seeds were produced for 30 to
50 d in field and greenhouse experiments (Illipronti et
al., 2000; unpublished data, 2002).
The temporal distribution of flower or pod production
was sensitive to planting date (Constable and Ross,
1988) and varied among years (Saitoh et al., 1998). Vari-
ation in plant productivity created by changes in plant
density (Torigoe et al., 1982), CO2 enrichment (Naka-
moto et al., 2001), or N nutrition (Torigoe et al., 1982)
had no effect on the temporal distribution of flower pro-
duction.
Pod and seed number in soybean respond to changes
in photosynthesis that are maintained during the entire
flowering and pod set period (Hardman and Brun, 1971;
Schou et al., 1978; Egli and Zhen-wen, 1991) or just a
portion of the period (Jiang and Egli, 1993). The tempo-
ral distribution of flower and pod production may play
an important role in these adjustments (Bruening and
Egli, 1999, 2000), but little is known of the relationship
between these distributions and photosynthesis. Some
research suggests that the temporal distribution of flow-
ers is not very sensitive to variation in photosynthesis
(Torigoe et al., 1982; Nakamoto et al., 2001), but the
effects on the distribution of pod production (appear-
ance of small pods) or pod survival have not been deter-
mined. These relationships must be defined before we
can completely understand the role these distributions
play in determining pod and seed number and yield in
soybean. Consequently, our objective was to investigate
the effect of large changes in photosynthesis (created
by shade treatments) on the temporal distribution of
pod production and survival in soybean. Small pods
(ⱖ10 mm long) were marked at regular intervals to
identify when pods were produced and when the pods
that survived until maturity initiated growth.
MATERIALS AND METHODS
Soybean plants (Elgin 87, Maturity Group II) were grown
in a greenhouse at the University of Kentucky using 3-L pots
(one plant per pot after overseeding and thinning) filled with
a 2:1 (v:v) mixture of a silt loam surface soil and vermiculite.
Experiment 1 was planted on 1 May and Experiment 2 on 14
August 2003. Air temperature in the greenhouse was main-
tained between 20 and 30⬚C and the photoperiod was never
less than 14 h, but the natural photoperiod exceeded 14 h
during Exp. 1. Supplemental radiation (120 ␮mol m⫺2 s⫺1
photosynthetic photon flux density) was provided by high-
pressure sodium lamps (430 W). The plants were not inocu-
lated with Bradyrhizobium japonicum and the roots were not
nodulated. A complete fertilizer (20–20–20, N–P–K) was ap-
plied approximately once every 2 wk.
At the beginning of flowering (approximately growth stage
 EGLI & BRUENING: TEMPORAL DISTRIBUTION OF SOYBEAN POD SET 1765

R1, Fehr and Caviness, 1977), plants were placed under black
commercial shade cloth (60 and 90%) to reduce photosynthe-
sis. Some plants remained under the shade until maturity, while
others were moved from the unshaded control to 90% shade
and vice-versa midway in the flowering and pod set period. Air
temperature (0.5 h means) was measured with two shielded
thermistors per treatment at the top of the plants and the data
were recorded with a Li-Cor 1000 data logger. The average
daily maximum and minimum temperatures under the shades
between growth stage R1 and R6 were within 1.0⬚C of the
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.

controls in both experiments. Average maximum temperature

in the control treatment in Exp. 2 was ≈2⬚C higher than in
Exp. 1. The differences between experiments were smaller
under the shade. The control, 60 and 90% shade treatments
were each assigned to a single greenhouse bench. Four pots
(four replications) of each treatment were randomly assigned
to the appropriate bench and a completely randomized design
was used for the statistical analysis.
The temporal distribution of pod development and pod
survival of each plant was characterized by marking all un-
marked pods ⱖ10 mm long with acrylic paint on the pedicel
and the base of the pod at 3-d intervals as described previously
(Egli and Bruening, 2002), and the color of the paint was
changed at each marking. The number of marked pods was
recorded when they were marked to provide a temporal distri-
bution of pod production. The color of the paint on mature
pods indicated when pods that survived until maturity (mature Fig. 1. The effect of continuous shade on pod production profiles
full size pods that contained at least one developed seed) (each data point represents the number of pods ⱖ 10 mm long
began growth, that is, the temporal distribution of surviving that were marked on that date). Bars represent, for each treatment,
pods or pod set. the average standard error of the mean after excluding means
approaching zero. Shade treatments were applied at approximately
All surviving pods were harvested at maturity (all pods growth stage R1 and maintained until maturity. Times of reproduc-
were brown), separated by paint color, and location (main tive growth stages R1, R3, R5, and R6 are shown on the x axis.
stem or branches) and counted. Generally, ⬍5% of the surviv-
ing pods did not have paint on them at maturity, and these
pods were included in the totals but not in the temporal distri- ber (Table 1). Roughly 30% of the pods were produced
butions. Seeds were removed from the pods in Exp. 1 and after growth stage R5 (beginning of seed filling, Fehr
counted. Pod abortion, calculated as the difference between and Caviness, 1977).
marked pods and surviving pods divided by marked pods, Continuous shade significantly (P ⫽ 0.05) reduced
does not include abortion of flowers or pods ⬍ 10 mm long.
pod production in both experiments (an average of 24%
for 60% shade and 71% for 90% shade, Table 1). Shade
RESULTS had a greater effect after peak pod production than
Pod production (appearance of pods ⱖ 10 mm long) before, but only the 90% shade treatment shortened
by control plants continued for 48 d in Exp. 1 vs. 30 d the period and then only in Exp. 1 (Fig. 1).
in Exp. 2 (Fig. 1). In both experiments, pod production The continuous-shade treatments also significantly
on the controls increased to a maximum and then de- (P ⫽ 0.05) reduced the surviving pods (mature pods
clined to zero. The plants in Exp. 1 started flowering in containing a developed seed) in both experiments
early June and produced more than twice as many pods (Table 1). Pods on branches made a much larger contri-
as the plants in Exp. 2, which started flowering in Octo- bution to the total pod load in Exp. 1 than Exp. 2

Table 1. The effect of shade on the distribution of marked and surviving pods on main stem and branches.
Surviving pods†
Marked pods Exp. 1 Exp. 2
Treatment Exp. 1 Exp. 2 Main stem Branches Total Main stem Branches Total
Pods plant⫺1
Control 183 70 52 134 186 38 30 67
Shade, %
60 141 53 42 98 141 25 19 44
90 44 30 16 14 30 10 5 15
Control/90‡ 119 53 28 47 75 14 8 22
90/control§ 90 56 36 48 84 40 16 56
CV (%) 13 12 16 24 14 20 32 10
LSD (0.05) 23 10 8 25 21 8 8 7
† Full sized pods containing a developed seed at maturity. Includes pods that were not marked (usually ⬍5% of total pods).
‡ Shade (90%) from approximately midway in the flowering and pod set period to maturity.
§ Shade (90%) from initial bloom (growth stage R1) to midway in the flowering and pod set period.
 1766 CROP SCIENCE, VOL. 45, SEPTEMBER–OCTOBER 2005
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.

Fig. 2. The effect of continuous-shade treatments on surviving pod Fig. 3. The effect of large changes in the radiation environment mid-
profiles (marked pods that were full size and contained at least way in the flowering and pod set period on pod production (each
one developed seed at maturity). For each treatment, bars represent data point represents the number of pods ⱖ 10 mm long that were
the average standard error of the mean after excluding means marked on that date) profiles. Bars represent, for each treatment,
approaching zero. Shade treatments were applied at approximately the average standard error of the mean after excluding means
growth stage R1 and maintained until maturity. Times of reproduc- approaching zero. The arrows indicate when plants in the transfer
tive growth stages R1, R3, R5, and R6 are shown on the x axis. treatments were moved into and out of shade. Times of reproduc-
tive growth stages R1, R3, R5, and R6 are shown on the x axis.
(Table 1). Seeds per pod of all treatments in Exp. 1
were within ⫾10% of the control (data not shown). creases in total pod abortion (significant at P ⫽ 0.05),
The temporal patterns of surviving pods (mature pods) and the increases were larger late in the pod-production
(Fig. 2) closely followed the pod production curves in period where roughly 80% of the marked pods did not
Fig. 1 in both experiments, and some surviving pods survive to maturity (Table 2).
initiated growth after growth stage R5 (29% in Exp. 1 Variation in environmental conditions in the field
and 22% in Exp. 2). Most pods on control plants sur- can cause large rapid changes in photosynthesis during
vived to maturity (i.e., total pod abortion was low, flowering and pod set that could influence final pod num-
Table 2). The 60% shade treatment did not greatly in- ber. We simulated such changes by exchanging control
crease total pod abortion relative to the levels of control and shade (90%) plants approximately midway through
plants (significant, P ⫽ 0.05, only in Exp. 2, Table 2). flowering and pod set. Pod production responded rap-
However, continuous 90% shade caused substantial in- idly to the drastic increase in solar radiation when plants
were removed from the shade (Fig. 3, 90%/control treat-
Table 2. The relationship between the time of pod development, ment). The increase in radiation did not extend the pod-
shade, and pod abortion. production period beyond that of the control, but it did
Pod abortion† extend it beyond the continuous 90% shade treatment
in Exp. 1. Pod production after the switch was, at one
Early† Late‡ Total
point, double control levels in Exp. 2, but it never ex-
Treatment Exp. 1 Exp. 2 Exp. 1 Exp. 2 Exp. 1 Exp. 2 ceeded the control in Exp. 1. The reduction in radiation
% when the plants were moved under the shade (control/
Control 4 0 3 29 4 7 90%) caused an almost immediate decrease in pod pro-
Shade, % duction (Fig. 3), but again, it had only minimal effects,
60 3 7 9 61 4 20
90 16 32 78 87 30 49 if any, on the length of the pod-production period.
Control/90§ 24 42 54 97 37 58 Pod production in the early-shade treatment (90%/
90/control¶
LSD (0.05)
0
13
0
9
13
22
20
19
8
14
9
9
control) was significantly (P ⫽ 0.05) less than the late-
LSD (0.10) 11 7 18 15 11 8 shade treatment (control/90%) in Exp. 1, but there was
† Abortion ⫽ (marked pods ⫺ surviving marked pods)/marked pods ⫻ 100.
no significant difference in Exp. 2 (Table 1). Pod produc-
‡ Abortion of pods marked before (early) or after (late) plants in the tion in both partial-shade treatments was significantly
transfer treatments were placed under or removed from the shade mid- (P ⫽ 0.05) less than the control (20 to 50%) but substan-
way in the flowering and pod set period.
§ Shade (90%) from midway in flowering and pod set to maturity. tially larger than the continuous 90% shade treatment.
¶ Shade (90%) from R1 to midway in flowering and pod set. The temporal distribution of pods that survived to
 EGLI & BRUENING: TEMPORAL DISTRIBUTION OF SOYBEAN POD SET
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
Fig. 4. The effect of large changes in the radiation environment mid-
way in the flowering and pod set period on surviving pods (marked
pods that were full size and contained at least one developed
seed at maturity) profiles. Bars represent, for each treatment, the
average standard error of the mean after excluding means ap-
proaching zero. The arrows indicate when plants in the transfer
treatments were moved into and out of the shade. Times of repro-
ductive growth stages R1, R3, R5, and R6 are shown on the x axis.
maturity in the transfer treatments (Fig. 4) generally
followed the patterns of marked pods (Fig. 3). Abortion
of early and late pods was significantly (P ⫽ 0.05) in-
creased above the control when plants were moved from
the high radiation environment to shade (control/90%
shade) midway through the pod-production period
(Table 2). The abortion of early or late pods on the
early shade (90% shade/control) treatment was not sig-
nificantly (P ⫽ 0.10) different from the control, but it
was significantly (P ⫽ 0.05) lower than continuous 90%
shade treatment.
DISCUSSION
The number of mature pods and seeds is directly
related to photosynthesis during flowering and pod set
in soybean (Schou et al., 1978; Egli, 1993; Jiang and
Egli, 1995) and this relationship was confirmed here
when both continuous-shade treatments significantly re-
duced the number of surviving pods (up to a 76% reduc-
tion). There was little change in seeds per pod in Exp. 1,
so the number of seeds was primarily determined by the
number of surviving pods. The temporal patterns of pod
production and survival may play a role in determining
pod number, but little is known about how reductions
in photosynthesis and assimilate supplies affect these
patterns.
Continuous shade affected the length of the pod-pro-
duction period in only one of four comparisons. The
length of the flowering period was also tolerant of treat-
1767
ments that influence photosynthesis (CO2 enrichment,
Nakamoto et al., 2001) or individual plant productivity
(plant density or N nutrition, Torigoe et al., 1982). Con-
tinuous shade reduced marked pod (small pods) produc-
tion and increased pod abortion. But, the primary cause
of reduced pod load under moderate shade stress seems
to have been the production of fewer small pods (lower
rate of pod production) with little change in the length
of the pod-production period or pod abortion. Abortion
made a significant contribution only under severe stress
where it was much higher (ⱖ80%) in late than early
developing pods, in agreement with previous work (Heit-
holt et al., 1986; Huff and Dybing, 1980). Shade may
have reduced flowers per plant (nodes per plant) or per
node, as reported previously (Jiang and Egli, 1993), or
it could have stimulated flower and small-pod abortion,
which can make a significant contribution to total abor-
tion (Hansen and Shibles, 1978; Huff and Dybing, 1980;
Heitholt et al., 1986). It’s possible that both flower pro-
duction and flower and small pod abortion were im-
portant.
The pod-production periods were shorter in Exp. 2,
which was planted on 14 August (vs. 1 May for Exp. 1),
and the first pods were marked 39 d after planting in
Exp. 2 vs. 44 d in Exp. 1. Earlier flowering (relative
to planting) and shorter flowering and pod-production
periods are common with late plantings in the field
(Constable and Ross, 1988; Egli and Bruening, 2000).
It is reasonable to assume that Exp. 2 probably experi-
enced lower radiation levels during pod production since
it occurred in late September and early October com-
pared with June and early July in Exp. 1. Air tempera-
tures were slightly higher in Exp. 2 (average of 26.9⬚C
in the control) than in Exp. 1 (25.8⬚C), and the natural
photoperiod in Exp. 1 was slightly longer (maximum of
nearly 2 h) than the 14-h photoperiod maintained in
Exp. 2. It is not clear whether the shorter photoperiod
(suggested by Kantolic and Slafer, 2001) or lower radia-
tion levels (both would occur normally in late field plant-
ings) were responsible for the shorter pod-production
period. If low radiation was responsible, it must have been
a cumulative effect from seedling emergence, since low-
ering radiation levels after growth stage R1 (continuous-
shade treatments) had almost no effect on the length
of the pod-production period. A shorter pod-production
period and a reduced rate of small pod (marked pods)
production and survival (a function of lower radiation
levels) were probably responsible for the lower mature
pod load in Exp. 2 (about half of Exp. 1).
The plants responded almost immediately to large
changes in the radiation regime midway in the flowering
and pod set period by modifying the production and
survival of small (marked) pods. The transfer treatments,
however, had essentially no effect on the length of the
pod production or pod survival periods. Previous rela-
tionships between photosynthesis and pod and seed
number were frequently based on static relationships
(e.g., average plant or crop growth rates during flow-
ering and pod set; Pandy et al., 1984; Egli and Zhen-wen,
1991; Vega et al., 2001). Our data, however, demonstrate
clearly that pod production and survival are dynamic sys-
 1768 CROP SCIENCE, VOL. 45, SEPTEMBER–OCTOBER 2005
tems that respond quickly (within days) to changes in
photosynthesis. In fact, the control/90% shade treatment
increased abortion of pods produced before shade was
imposed (early pods, Table 2) which is not surprising given
evidence that pods are susceptible to abortion until rapid
seed development begins (Duthion and Pigeaire, 1991;
Westgate and Peterson, 1993). Predictions of pod num-
ber from average measures of productivity will be accu-
rate only when environmental conditions are relatively
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
stable during the critical period, not a common occur-
rence in the field. The magnitude of short-term fluctua-
tions in photosynthesis needed to reduce mature pod
number will probably depend on the relationship be-
tween photosynthesis, storage carbohydrates, and the
assimilate supply to reproductive structures as well as
the length of time that a pod is sensitive to low levels
of assimilate.
The mature pod load never recovered to control levels
in either experiment (mean pod load was 45% less than
the control) when the plants were removed from the
shade midway through the pod-production period (90%
shade/control treatment). Pod production continued for
approximately 20 d after the transfer, pod production
was well above the continuous 90% shade treatment,
and abortion was reduced, but these changes were not
enough to recover the pods lost during the early shade.
There were not enough small pods produced to replace
the lost pods; this failure was partially due to a lack of
time as the higher radiation levels did not extend the
pod-production period beyond the control. Much higher
rates of pod production would be needed without an
extension of the pod-production period. The fact that
the plants were smaller coming out of the shade, proba-
bly with less leaf area (photosynthesis per plant is par-
tially determined by leaf area in spaced plants) and
fewer nodes (flowers per plant are related to nodes per
plant, Egli, 2005) also could have limited pod production
and survival. The exact reasons for the failure of the
pod load to recover to control levels when the shade was
removed are not known, but it seems that the inability of
the plant to extend the pod-production period made some
contribution.
It is often assumed that the long period of pod produc-
tion and pod set in soybean may help stabilize pod
number in fluctuating environments (Shibles et al., 1975;
Loomis and Conner, 1992). However, if the plant cannot
increase late pod production (higher rates or extend the
period) enough to overcome early losses, and they could
not in our greenhouse experiments, then soybean may
be no more tolerant of variable environments than spe-
cies such as corn (Zea mays L.) that have shorter flow-
ering and seed set periods (Tollenaar and Daynard,
1978; Grant et al., 1989). Although our results with spaced
plants cannot be extrapolated directly to the field, there
are reports that field communities could not recover
from early stress (Kokubun and Watanabe, 1983; Jiang
and Egli, 1995). Collectively, these results suggest that
maximum pod and seed number, and yield, may require
a relatively stress free environment throughout flow-
ering and pod set. Just how stress free (what length
and level of stress will still allow a complete recovery?)
remains to be determined.
Pod production and survival in these experiments re-
sponded dynamically to changes in photosynthesis after
growth stage R1, to continuous changes that might dif-
ferentiate a high- from a low-yield environment, and to
shorter fluctuations that could occur in many field envi-
ronments. The length of the pod-production period was
almost completely insensitive to changes in photosyn-
thesis, and most of the variation in mature pods was deter-
mined by small pod production. Pod abortion seemed to
play a major role only under severe stress. Our results
support previous contentions (Egli, 2005) that flower
and small pod production are usually more important
than abortion in determining the number of mature pods.
The dynamic nature of pod production and survival means
that models predicting pod and seed number must include
the time component of flower and pod production and
survival to accurately account for short term variations
in photosynthesis. Predictions based on average photo-
synthesis or crop growth rates during the critical period
will probably accurately reflect large changes in environ-
mental conditions (e.g., high- vs. low-yield environments),
but they may not accommodate smaller changes resulting
from short-term fluctuations in the environment and in
photosynthesis.
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