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temporal distributions in two greenhouse experiments. Plants (‘Elgin 50 d in field and greenhouse experiments (Illipronti et
87’) were exposed to two levels of shade (60 and 90%) from growth al., 2000; unpublished data, 2002).
stage R1 to maturity. Other plants were removed from 90% shade
The temporal distribution of flower or pod production
or placed under 90% shade midway through flowering (transfer treat-
ments). Temporal distributions of pod production and pod survival
was sensitive to planting date (Constable and Ross,
were determined by marking all unmarked pods ⱖ 10 mm long on 1988) and varied among years (Saitoh et al., 1998). Vari-
plants every three days with different colored paint. The color of ation in plant productivity created by changes in plant
paint on the mature pods identified when they started development. density (Torigoe et al., 1982), CO2 enrichment (Naka-
Continuous shade reduced mature pods by 27 (60% shade) and 82% moto et al., 2001), or N nutrition (Torigoe et al., 1982)
(90% shade), but it shortened the pod-production period in only one had no effect on the temporal distribution of flower pro-
of four comparisons. Pod production responded quickly to transfer duction.
treatments, and the mature pod load was always greater (nearly three Pod and seed number in soybean respond to changes
fold) than the continuous 90% shade treatment and less (average of in photosynthesis that are maintained during the entire
53%) than the control. The mature pod load failed to recover from flowering and pod set period (Hardman and Brun, 1971;
early shade because the increase in radiation did not lengthen the
pod-production period and not enough pods were produced. Pod
Schou et al., 1978; Egli and Zhen-wen, 1991) or just a
production was often more important than pod abortion in determin- portion of the period (Jiang and Egli, 1993). The tempo-
ing mature pod number. Adding the temporal distribution of pod ral distribution of flower and pod production may play
production and survival to models predicting pod and seed number an important role in these adjustments (Bruening and
will improve their accuracy. 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
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;
(Torigoe et al., 1982; Nakamoto et al., 2001), but the
effects on the distribution of pod production (appear-
Frederick et al., 1998). However, the mechanisms by ance of small pods) or pod survival have not been deter-
which the plant regulates the number of pods and seeds mined. These relationships must be defined before we
it produces are not completely understood. Recent evi- can completely understand the role these distributions
dence (Bruening and Egli, 1999, 2000) suggests that play in determining pod and seed number and yield in
the temporal distribution of flower and pod production soybean. Consequently, our objective was to investigate
should be added to the traditional determinants of pod the effect of large changes in photosynthesis (created
and seed number—photosynthesis or assimilate avail- by shade treatments) on the temporal distribution of
ability and sink (seed) characteristics (Charles-Edwards pod production and survival in soybean. Small pods
et al., 1986; Egli, 1998). (ⱖ10 mm long) were marked at regular intervals to
The asynchronous flowering characteristic of soybean identify when pods were produced and when the pods
is well documented. Flowering periods (first to last that survived until maturity initiated growth.
flower on a plant) are frequently 30 d long or longer
(Hansen and Shibles, 1978; Yoshida et al., 1983; Gai et MATERIALS AND METHODS
al., 1984; Dybing, 1994). The length of the flowering
Soybean plants (Elgin 87, Maturity Group II) were grown
period was sensitive to daylength (Guiamet and Naka- in a greenhouse at the University of Kentucky using 3-L pots
yama, 1984) and planting date (Constable and Ross, (one plant per pot after overseeding and thinning) filled with
1988; Dybing, 1994), but CO2 enrichment (Nakamoto a 2:1 (v:v) mixture of a silt loam surface soil and vermiculite.
et al., 2001), plant density (Torigoe et al., 1982; Saitoh Experiment 1 was planted on 1 May and Experiment 2 on 14
August 2003. Air temperature in the greenhouse was main-
Dep. of Plant and Soil Science, Univ. of Kentucky, Lexington, KY tained between 20 and 30⬚C and the photoperiod was never
40546-0312. Published with the approval of the Director of the Ken- less than 14 h, but the natural photoperiod exceeded 14 h
tucky Agric. Exp. Stn. as paper 04-06-147. Received 21 Sept. 2004. during Exp. 1. Supplemental radiation (120 mol m⫺2 s⫺1
*Corresponding author (degli@uky.edu). photosynthetic photon flux density) was provided by high-
Published in Crop Sci. 45:1764–1769 (2005).
pressure sodium lamps (430 W). The plants were not inocu-
Crop Physiology & Metabolism lated with Bradyrhizobium japonicum and the roots were not
doi:10.2135/cropsci2004.0557 nodulated. A complete fertilizer (20–20–20, N–P–K) was ap-
© Crop Science Society of America plied approximately once every 2 wk.
677 S. Segoe Rd., Madison, WI 53711 USA At the beginning of flowering (approximately growth stage
1764
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.
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 1767
tems that respond quickly (within days) to changes in and level of stress will still allow a complete recovery?)
photosynthesis. In fact, the control/90% shade treatment remains to be determined.
increased abortion of pods produced before shade was Pod production and survival in these experiments re-
imposed (early pods, Table 2) which is not surprising given sponded dynamically to changes in photosynthesis after
evidence that pods are susceptible to abortion until rapid growth stage R1, to continuous changes that might dif-
seed development begins (Duthion and Pigeaire, 1991; ferentiate a high- from a low-yield environment, and to
Westgate and Peterson, 1993). Predictions of pod num- shorter fluctuations that could occur in many field envi-
ber from average measures of productivity will be accu- ronments. The length of the pod-production period was
rate only when environmental conditions are relatively almost completely insensitive to changes in photosyn-
Reproduced from Crop Science. Published by Crop Science Society of America. All copyrights reserved.
stable during the critical period, not a common occur- thesis, and most of the variation in mature pods was deter-
rence in the field. The magnitude of short-term fluctua- mined by small pod production. Pod abortion seemed to
tions in photosynthesis needed to reduce mature pod play a major role only under severe stress. Our results
number will probably depend on the relationship be- support previous contentions (Egli, 2005) that flower
tween photosynthesis, storage carbohydrates, and the and small pod production are usually more important
assimilate supply to reproductive structures as well as than abortion in determining the number of mature pods.
the length of time that a pod is sensitive to low levels The dynamic nature of pod production and survival means
of assimilate. that models predicting pod and seed number must include
The mature pod load never recovered to control levels the time component of flower and pod production and
in either experiment (mean pod load was 45% less than survival to accurately account for short term variations
the control) when the plants were removed from the in photosynthesis. Predictions based on average photo-
shade midway through the pod-production period (90% synthesis or crop growth rates during the critical period
shade/control treatment). Pod production continued for will probably accurately reflect large changes in environ-
approximately 20 d after the transfer, pod production mental conditions (e.g., high- vs. low-yield environments),
was well above the continuous 90% shade treatment, but they may not accommodate smaller changes resulting
and abortion was reduced, but these changes were not from short-term fluctuations in the environment and in
enough to recover the pods lost during the early shade. photosynthesis.
There were not enough small pods produced to replace
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