Effect of nodulation on assimilate remobilization in soybean

The objectives of this work were to determine the effect of nodulation on dry matter, reduced-N, and phosphorus accumulation and partitioning in above-ground vegetative parts and pods of field-grown soybean (Glycine max [L.] Merr. cv Harosoy).

From comparison of nodulated and nonnodulated isolines, it was estimated that nodulated plants attained 81 and 71% of total-plant (above ground) N from uptake of soil N in 1981 and 1982, respectively. These data, along with visibly greener leaves of nodulated plants, led us to assume that nonnodulated plants were under a moderate N stress relative to nodulated plants. Nonnodulated plants accumulated less total-plant N and partitioned less dry matter and N to the pods, compared with nodulated plants. This occurred even though net photosynthesis, as estimated by rate and amount of dry matter accumulation, was the same for both nonnodulated and nodulated plants. Rate of dry matter and reduced-N accumulation in pods was less for nonnodulated than for nodulated plants while duration of podfill was similar for both isolines. From these data we concluded that moderate N stress affected partitioning of photosynthate rather than net photosynthesis, and that N played a role in translocation of photosynthate to the pods. Total plants (above-ground portion) and pods of both nodulated and nonnodulated plants accumulated similar amounts of phosphorus, which indicated that phosphorus and N accumulation were independent.

Remobilization of nitrogen and phosphorus from vegetation to pods preceded dry matter remobilization. It appeared that either more nitrogen accumulation prior to podfill, or continued nitrogen assimilation during podfill would increase nitrogen and dry matter partitioning to pods, but that increasing photosynthesis without concomitantly increasing nitrogen input may not necessarily result in enhanced seed production.

     

Effects of Pod Removal on Metabolism and Senescence of Nodulating and Nonnodulating Soybean Isolines: II. Enzymes and Chlorophyll

The objectives of this work were to determine the effect of sink strength (presence or absence of pods) and nitrogen source (nodulating versus nonnodulating plants) on enzymic activities, chlorophyll concentration, and senescence of soybean (Glycine max [L.] Merr. cv Harosoy) isolines. A 2-year (1981-1982) field study was conducted.

For both nodulated and nonnodulated plants, ribulose bisphosphate carboxylase (RuBPCase) activity of upper-canopy leaves was decreased by pod removal in both years, while chlorophyll concentration was decreased in 1981 only. Nonnodulated plants had lower RuBPCase activity in 1981 and lower chlorophyll concentration in both years compared with nodulated plants. In both years, and for all treatments, RuBPCase activity and chlorophyll began to decline at about the same time, but the rate of decline was less for depodded than for podded plants. Leaves in the middle and lower parts of the canopy had similar RuBPCase activity and chlorophyll concentration trends as upper-canopy leaves for all treatments.

Profiles of nitrate reductase activity (NRA) were similar for all treatments in both 1981 and 1982. Acetylene reduction profiles were similar for nodulated-podded and nodulated-depodded plants. The peak and decline in NRA profiles preceded the peak and decline in acetylene reduction profiles. The rate of decline in acetylene reduction activity was less for depodded plants, especially in 1982, but activities reached zero by the final sampling time. Thus, nodule senescence was not prevented by pod removal.

Based on seasonal profiles of RuBPCase activity, chlorophyll, NRA, and acetylene reduction activity, the initiation of senescence appeared to occur at the same approximate time for all treatments and, thus, did not depend on the presence or absence of pods or nodules. The hypothesis that nodules act as a nitrogen source and carbohydrate sink to delay senescence in the absence of pods was not correct.

     

Protein compositions of mesophyll and paraveinal mesophyll of soybean leaves at various developmental stages

Mesophyll and paraveinal mesophyll protoplasts (PVMP) were isolated from leaves of soybean (Glycine max) at various stages of physiological development, and protein compositions of the two protoplast types were analyzed by sodium dodecyl sulfate-polyacrylamide gel electrophoresis and immunoblotting. Polypeptides of 27, 29 (previously shown to be storage proteins), and 94 kilodaltons were found to be PVMP-specific proteins and were present in both nodulated and nonnodulated plants. The 27 and 94 kilodalton polypeptides were major PVMP constituents. All three polypeptides accumulate as early as one-quarter leaf expansion. Immunoblotting and immunocytochemical studies using antibodies against the 27/29 kilodalton proteins confirmed that they are specific to the paraveinal mesophyll (PVM) and that they are localized in the PVM vacuole. The 27 kilodalton polypeptide increased significantly by two weeks depodding, and this accumulation was restricted to the PVM vacuole. Radiolabeling experiments showed that the difference in relative amounts of the 27 and 29 kilodalton polypeptides was due to a greater rate of synthesis of the 27 kilodalton polypeptide. The 94 kilodalton polypeptide accumulated to a maximum at anthesis, but was absent at 2 weeks postanthesis in both depodded and podded nodulated plants, probably because they were nitrogen limited. In nonnodulated plants, it was present through 2 weeks postanthesis. The results confirm that the 27 and 29 kilodalton proteins of soybean leaf are stored in the PVM vacuole and show that they are accumulated early during leaf development while they are still strong sinks for nitrogen. The 94 kilodalton protein, previously found to accumulate in leaves after depodding, is also a PVM protein and is likely a third vegetative storage protein, although its accumulation appears to be more dependent on excess nitrogen availability. The results further support the hypothesis that the PVM is a specialized leaf tissue that functions in synthesis and compartmentation of storage proteins.     

Transmission of the Monocarpic Senescence Signal via the Xylem in Soybean

During monocarpic senescence in soybean (Glycine max [L.] Merrill cv. Anoka) there is a remobilization of nitrogen from the leaves to the seeds, and it has been hypothesized that this loss of nitrogen from the leaves induces foliar yellowing. The phloem in a small segment of the petiole between the pods and the target leaf can be inactivated with a jet of steam. When a plant is depodded except for a single pod cluster in the center of the plant, the pod cluster induces yellowing of the nearest leaf even if the petiole contains a zone of dead phloem, whereas most of the rest of the plant remains green. The nitrogen content of these leaves with a dead phloem zone in their petioles does not decrease greatly, even though the leaves turn yellow. A similar treatment of a single leaf on a fully depodded plant (leaves stay green) does not cause that leaf to turn yellow. Since nutrients would have to be withdrawn from the leaves via the phloem, the pods do not induce yellowing by pulling nutrients out of the leaf and must be able to exert their influence via the xylem.     

Soybean Lipoxygenase L-4, a Component of the 94-kilodalton Storage Protein in Vegetative Tissues: Expression and Accumulation in Leaves Induced by Pod Removal and by Methyl Jasmonate

A lipoxygenase L-4 gene was isolated from a soybean genomiclibrary. The amino acid sequence of lipoxygenase L-4 is highlyhomologous with the partial amino acid sequence of the 94-kDavegetative storage protein, vsp94, found in paraveinal mesophyllcells in the leaves of depodded soybean plants. No L-4 expressionwas observed in maturing seeds. The L-4 gene is highly expressedin the vegetative tissues of young seedlings, including cotyledons,hypocotyls, roots and primary leaves. L-4 expression followedthe same pattern as lipoxygenase activity in cotyledons peaking3 to 5 days after germination, and returning to a basal levelby 9 days after germination. L-4 gene expression was low inthe roots, stems and leaves of 10-week-old plants. Exposureof 4-week-old plants to atmospheric methyl jasmonate increasedL-4 mRNA in leaves. Continuous pod removal from 7-week-old plantsover a 2 week period resulted in dramatic accumulation of L-4mRNA in leaves. Accumulation of the L-4 protein and three otherlipoxygenase fractions in the leaves of depodded plants wasdemonstrated by ion exchange chromatography. These results indicatethat lipoxygenase L-4 is a component of vsp94. (Received May 31, 1993; Accepted August 9, 1993)     

Characterization of leaf senescence and pod development in soybean explants

Excised soybean (Glycine max [L.] Merrill) cv Anoka leaf discs tend to remain green even after the corresponding intact leaves have turned yello on fruiting plants. We have found that explants which include a leaf along with a stem segment (below the node) and one or more pods (maintained on distilled H₂O) show similar but accelerated leaf yellowing and abscission compared with intact plants. In podded explants excised at pre-podfill, the leaves begin to yellow after 16 days, whereas those excised at late podfill begin to yellow after only 6 days. Although stomatal resistances remain low during the first light period after excision, they subsequently increase to levels above those in leaves of intact plants. Explants taken at mid to late podfill with one or more pods per node behave like intact plants in that pod load does not affect the time lag to leaf yellowing. Explant leaf yellowing and abscission are delayed by removal of the pods or seeds or by incubation in complete mineral nutrient solution or in 4.6 micromolar zeatin. Like chorophyll breakdown, protein loss is accelerated in the explants, but minerals or especially zeatin can retard the loss. Pods on explants show rates and patterns of color change (green to yellow to brown) similar to those of pods on intact plants. These changes start earlier in explants on water than in intact plants, but they can be delayed by adding zeatin. Seed dry weight increased in explants, almost as much as in intact plants. Explants appear to be good analogs of the corresponding parts of the intact plant, and they should prove useful for analyzing pod development and mechanisms of foliar senescence. Moreover, our data suggest that the flux of minerals and cytokinin from the roots could influence foliar senescence in soybeans, but increased stomatal resistance does not seem to cause foliar senescence.     

Effect of pod removal on leaf photosynthesis and soluble protein composition of field-grown soybeans

Well nodulated, field-grown soybeans (Glycine max [L.] Merr. var Williams) were depodded just prior to seed development and near mid pod-fill. Both treatments caused a considerable increase in leaf dry weight, suggesting continued photosynthate production following pod removal. Moreover, depodding had a marked effect on leaf soluble protein without affecting total proteolytic activity. Early depodding caused a 50% increase in leaf protein, and both early and late depodding caused the retention of protein for several weeks following the decline in control leaves. But despite this retention of protein, leaves of depodded plants showed no difference in the onset of the irreversible decline in photosynthesis. Therefore, although depodding delayed the loss of leaf chlorophyll and protein, it did not delay the onset of functional leaf senescence and in fact, actually appeared to enhance the rate of decline in photosynthesis. There was a good correlation between the irreversible decline in ribulose bisphosphate carboxylase (activity and amount) and that of photosynthesis. In contrast, the correlation did not seem as good between stomatal closure and the onset of the irreversible decline in photosynthesis. The reason total soluble protein remained high following depodding while carboxylase, which normally comprised 40% of the soluble protein, declined was because several polypeptides increased in amounts sufficient to offset the loss of carboxylase. This change in leaf protein composition indicates a change in leaf function; this is discussed in terms of other recent findings.     

Differential Senescence of Maize Hybrids following Ear Removal : I. Whole Plant

Visual senescence symptoms and associated changes in constituent contents of three field-grown maize (Zea mays L.) hybrids (Pioneer brand 3382, B73 × Mo17, and Farm Service brand 854) were compared in response to ear removal. Whole plants were harvested at eight intervals during the grain-filling period, and analyzed for dry matter, total N and nitrate N, phosphorus, sugars, and starch.

Upper leaves of earless P3382 and B73 × Mo17 showed reddish discoloration by 25 days after anthesis (DAA) and all leaves had lost most of their chlorophyll by 40 DAA. In striking contrast, leaves of earless FS854 plants remained green and similar in appearance to eared controls throughout the grain-filling period.

For all hybrids, ear removal led to a decrease in dry weight, reduced N, total N, and phosphorus contents of the total plant, and an increase in carbohydrate content of the leaves and stalks, relative to respective controls. Although changes in carbohydrate and N contents, which previously had been associated with senescence, were observed for all earless hybrids, these changes were followed by accelerated senescence and early death only for P3382 and B73 × Mo17. By 30 DAA, earless P3382 and B73 × Mo17 plants ceased to accumulate dry weight, total N, and phosphorus, indicating a termination of major metabolic activities. In contrast, earless FS854 plants retained a portion of these metabolic activities until 58 DAA, indicating a role for roots in determining rate of senescence development. Thus, the course of senescence was more accurately reflected by measurements of metabolic activities than by measurements of metabolite contents at any given time. These results show that the ear per se does not dictate the rate or completion of the senescence process, and implicated an association between the continued accumulation of N and associated root activities with the delayed senescence pattern of the earless FS854 plants. It is evident that studies involving control of senescence among species must also consider genotypic influences within species.

     

DEVELOPMENTAL CHANGES IN ENERGY (ATP AND ADP) AND NUTRIENT LEVELS PRIOR TO SENESCENCE IN PODDED AND DEPODDED PEA PLANTS

Energy (ATP and ADP) levels in stem apices of depodded pea plants (Pisum sativum L. cv. Little Marvel) were significantly higher than those of podded plants during the pod-filling stage before whole plant senescence. This difference in energy content appeared before decreases in leaf chlorophyll and soluble proteins occurred in plants of both treatments. In contrast, the mineral nutrient levels (N, P, K, Ca, Mg, Fe, and Mn) in stem apices of plants from both treatments were similar. Energy levels in reproductive leaves from podded and depodded plants were similar. The mineral nutrients in leaves with fruits in their axils and similar leaves of depodded plants were comparable except for nitrogen. Plant growth measurements—dry weight, leaf area, leaf dry weight, root/shoot ratio—were significantly higher in depodded than podded plants. Whole plant senescence occurred significantly earlier in podded than in depodded pea plants.     

Biochemical Basis for Partitioning of Photosynthetically Fixed Carbon between Starch and Sucrose in Soybean (Glycine max Merr.) Leaves

The control of photosynthetic starch/sucrose formation in leaves of soybean (Glycine max L. Merr.) cultivars was studied in relation to stage of plant development, photosynthetic photoperiod, and nitrogen source. At each sampling, leaf tissue was analyzed for starch content, activities of sucrose-metabolizing enzymes, and labeling of starch and sucrose (by ¹⁴CO₂ assimilation) in isolated cells. In three of the four varieties tested, nodulated plants had lower leaf starch levels and higher activities of sucrose phosphate synthetase (SPS), and isolated mesophyll cells incorporated more carbon (percentage of total ¹⁴CO₂ fixed) into sucrose and less into starch as compared to nonnodulated (nitrate-dependent) plants. The variation among cultivars and nitrogen treatments observed in the activity of SPS in leaf extracts was positively correlated with labeling of sucrose in isolated cells (r = 0.81) and negatively correlated with whole leaf starch content (r = −0.66). The results suggested that increased demand for assimilates by nodulated roots may be accommodated by greater partitioning of carbon into sucrose in the mesophyll cells. We have also confirmed the earlier report (Chatterton, Silvius 1979 Plant Physiol 64: 749-753) that photoperiod affects partitioning of fixed carbon into starch. Within two days of transfer of nodulated soybean Ransom plants from a 14-hour to a 7-hour photoperiod, leaf starch accumulation rates doubled, and this effect was associated with increased labeling of starch and decreased labeling of sucrose in isolated cells. Concurrently, activities of SPS, sucrose synthase, and uridine diphosphatase in leaves were decreased.     

Growth, sucrose synthase, and invertase activities of developing Phaseolus vulgaris L. fruits

Activities of the sucrose-cleaving enzymes, acid and neutral invertase and sucrose synthase, were measured in pods and seeds of developing snap bean (Phaseolus vulgaris L.) fruits, and compared with ¹⁴C-import, elongation and dry weight accumulation. During the first 10 d post-anthesis, pods elongated rapidly with pod dry weight increase lagging behind by several days. The temporal patterns of acid invertase activity and import coincided closely during the first part of pod development, consonant with a central role for this enzyme in converting imported sucrose during pod elongation and early dry weight accumulation. Later, sucrose synthase became the predominant enzyme of dry weight accumulation and was possibly associated with the development of phloem in pod walls. Sucrose synthase activity in seeds showed two peaks, corresponding to two phases of rapid import and dry weight accumulation; hence, sucrose synthase was associated with seed sink growth. Acid invertase activities in seeds were low and did not show a noticeable relationship with import or growth. All neutral invertase activities, during pod and seed development, were too low for it to have a dominant role in sucrose cleavage. Changes in activities of certain sucrose-cleaving enzymes appear to be correlated with certain sink functions, including import, storage of reserves, and biosynthetic activities. The data supports the association of specific sucrose-cleaving enzymes with the specific processes that occur in the developing pods and seeds of snap bean fruits; for example, acid invertase with pod elongation and sucrose synthase with fruit dry matter accumulation.     

Studies on Genetic Male-Sterile Soybeans : IV. Effect of Male Sterility and Source of Nitrogen Nutrition on Accumulation, Partitioning, and Transport of Nitrogen

Soybean (Glycine max [L.] Merr.) germplasm, isogenic except for loci controlling male sterility (ms₁) and nodulation (rj₁), was used to investigate the effects of reproductive tissue development and source of nitrogen nutrition on accumulation, transport, and partitioning of nitrogen in a greenhouse experiment. Nodulated plants were supplied nitrogen-free nutrient solution, and nonnodulated plants were supplied nutrient solution containing 20 millimolar KNO₃. Plants were sampled from flowering until maturity (77 to 147 days after transplanting).

Accumulation rates of nitrogen in whole plants during reproductive growth were not significantly different among the four plant types. Nitrogen accumulation in the sterile, nonnodulated plants, however, ceased 2 weeks earlier than in fertile, nonnodulated or fertile and sterile, nodulated plants. This early cessation in nitrogen accumulation resulted in sterile, nonnodulated plants accumulating significantly less whole plant nitrogen by 133 days after transplanting (DAT) than fertile, nonnodulated plants. Thus, changing the site of nitrogen assimilation from nodules (N₂-fixing plants) to roots and leaves (NO₃-fed plants) resulted in similar whole-plant nitrogen accumulation rates in fertile and sterile plants, despite the absence of seed in the latter.

Leaflet and stem plus petiole tissues of both types of sterile plants had significantly higher nitrogen concentrations after 119 DAT than both types of fertile plants. Significantly higher concentrations and exudation rates of nonureide, reduced-nitrogen in xylem sap of sterile than of fertile plants after 105 DAT were observed. These latter results indicated possible cycling of nonureide, reduced-nitrogen from the downward phloem translocation stream to the upward xylem translocation stream in roots of sterile plants. Collectively, these results suggest a lack of sinks for nitrogen utilization in the shoots of sterile plants. Hence, comparison of nitrogen accumulation rates for sterile and fertile plants does not provide a definitive test of the hypothesis that reproductive tissue development limits photosynthate availability for support of N₂ fixation and nitrate assimilation in determinate soybeans.

Nitrogen assimilation during reproductive growth met a larger proportion of the reproductive-tissue nitrogen requirement of nitrate-dependent plants (73%) than of N₂-fixing plants (63%). Hence, vegetative-tissue nitrogen mobilization to reproductive tissue was a more prominent process in N₂-fixing than in nitrate-dependent plants. N₂-fixing plants partitioned nitrogen to reproductive tissue more efficiently than nitrate-dependent plants as the reproductive tissues of the former and latter contained 65 and 55%, respectively, of the whole-plant nitrogen at the time that nitrogen accumulation in reproductive parts had ceased (133 DAT).

     

Alteration of C-assimilate partitioning in leaves of soybeans having increased reproductive loads at one node

The objectives of this study were to determine if the partitioning of recently fixed carbon between starch and water-soluble compounds could be altered by increasing the pod load in the leaf axil, and if the presence of source leaves acropetal to such a node would influence the partitioning of carbon within the subtending leaf. Soybeans (Glycine max L. Merr. cv Hodgson 78) were grown to full-bloom in a controlled environment chamber, and then deflowered at all nodes except the eighth. This treatment resulted in an 83% increase in the number of pods at the eighth node. At 24 days after flowering, one-half of the treated plants were girdled above the untreated node. Forty-two hours later, the eighth trifoliolate was pulsed with ¹⁴CO₂ and sampled for radiolabeled starch and water-soluble compounds (WSC) at 0.5, 2, 4, 8, 12, and 24th after labeling.

When no girdling was applied above the increased pod load at the eighth node more label was accumulated by the pod walls (+6.9%) and seeds (+6.3%) when compared to the controls. Starch accumulation was not altered in the labeled leaf of the nongirdled plants. When the stem was girdled above the eighth node, significantly less starch was retained in the labeled leaf. Girdling also resulted in an increase in label accumulation by the pod walls (+5.4%) and seeds (+6.6%). These data suggest that the plant will change the distribution patterns of assimilate to supply added sink demand before altering the partitioning of recently fixed carbon in the subtending leaf.

     

Weather and nodule mediated variations in delta 13C and delta 15N values in field-grown soybean (Glycine max L.) with special interest in the analyses of xylem fluids

The nodulating soybean (Enrei) and its non-nodulating mutant (EN 1282) were grown in outdoor plots for 2 years (1994: extraordinary dry, high temperature, 1995: ordinary year). Carbon and nitrogen accumulation, delta 13C and delta 15N values in plant parts and xylem fluids and delta 15N values in the water-extractable soil N were analysed throughout the growing period. Plant growth in 1994 was rapid during the early growth stages, but no pods were produced. In 1995, plant growth was normal and pods were formed. The delta 13C values of the leaves were less negative in 1994 than in 1995 and the nodulated plants showed less negative delta 13C values than non-nodulated plants in both years. The delta 13C values of the leaves during the vegetative phase were positively correlated to the leaf N concentrations. Leaf N concentrations in their turn were influenced by nodulation and weather conditions and/or soil available N. The delta 15N values in the plants and xylem fluids were lower in the nodulated soybean than in non-nodulated soybean in both years, and estimates of the contribution of N2 fixation in nodulated plants based on plant top delta 15N values were 7-14% in 1994 and 37-63% in 1995. The delta 13C values of xylem fluids did not differ between nodulated and non-nodulated plants. Thus, the expected contribution by phosphopenolpyruvate carboxylase-mediated CO2 fixation in the root nodules to plant C-incorporation could not have been significant.     

Studies on Genetic Male-Sterile Soybeans: I. Distribution of Plant Carbohydrate and Nitrogen during Development

Soybeans (Glycine max [L.] Merr. cv. NC 69-2774) were used to study the nonstructural carbohydrate and nitrogen content of plant tissues, and nitrogenase activity throughout the development of male-sterile and male-fertile plants. Male-sterile plants set approximately 85% fewer pods plus seed than the male-fertile siblings and retained green leaves until a killing frost at 145 days after emergence. Reduced pod set caused increased carbohydrate accumulation in the leaf and root systems of male-sterile plants. Total carbohydrate in roots of male-sterile plants increased from 1.7 to 7.6 times that in the male-fertile roots. A high proportion (60 to 70%) of the male-sterile root carbohydrate was starch. Apparently, root starch was not metabolized by the male-sterile plants. Late in plant development per cent nitrogen was higher in the male-sterile soybean tissues. However, no difference was found in the ability of the nodulated root systems from either genotype to fix nitrogen.     

Role of aminolevulinic acid in improving biomass production in Vigna catjung, V. mungo, and V. radiata

A precursor in tetrapyrrole biosynthesis, 5-aminolevulinic acid (ALA), was applied via presowing soaking in Vigna catjung, V. mungo, and V. radiata. ALA increased plant growth and influenced dry matter accumulation in leaves, stems, and pods through increased chlorophyll content and photosynthetic CO₂ absorption. At harvest, ALA treated plants had increased number of pods per plant, seeds per pod, 100 seed dry matter, biological yield, and the harvest index. Therefore, pretreatment of seeds with optimal concentration of ALA is recommended for improving the growth and productivity of tropical legumes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Sink removal and leaf senescence in soybean : cultivar effects

Three cultivars of soybean (Glycine max [L.] Merr. cvs Harper, McCall, and Maple Amber) were grown in the field and kept continuously deflowered throughout the normal seedfill period. For all cultivars, deflowering led to delayed leaf abscission and a slower rate of chlorophyll loss. Compared to control plants, photosynthesis and ribulose 1,5-bis-phosphate carboxylase/oxygenase (Rubisco) level declined slightly faster for deflowered Harper, but for both McCall and Maple Amber, leaves of deflowered plants maintained approximately 20% of maximum photosynthesis and Rubisco level 1 month after control plants had senesced. Deflowering led to decreased leaf N remobilization and increased starch accumulation for all cultivars, but cultivars differed in that for McCall and Maple Amber, N and starch concentrations slowly but steadily declined over time whereas for Harper, N and starch concentrations remained essentially constant over time. SDS-PAGE of leaf proteins indicated that for all cultivars, deflowering led to accumulation of four polypeptides (80, 54, 29, and 27 kilodaltons). Western analysis using antisera prepared against the 29 and 27 kilodalton polypeptides verified that these polypeptides were the glycoproteins previously reported to accumulate in vacuoles of paraveinal mesophyll cells of depodded soybean plants. The results indicated that depending on the cultivar, sink removal can lead to either slightly faster or markedly slower loss of photosynthesis and Rubisco. This difference, however, was not associated with the ability to synthesize leaf storage proteins. For any particular cultivar, declines in chlorophyll, photosynthesis, and Rubisco were initiated at approximately the same time for control and deflowered plants. Thus, even though cultivars differed in rate of decay of photosynthetic rate and Rubisco level in response to sink removal, the initiation of leaf senescence was not influenced by presence or absence of developing fruits.     

Pathway and regulation of phosphate translocation to the pods of soybean explants

We investigated the degree to which developing fruit compete directly with leaves for mineral nutrients, e.g. phosphate coming up from the roots. When soybean ( Glycine max (L.) Merrill cv. Anoka) explants cut at mid-late podfill were given a 15-min pulse of ³²Pi via the cut stem and then transferred to distilled water, 75% of the ³²P accumulated in the leaves and 21% in stem and petiole during the first hour. The amount of ³²P entering the seeds was low (1%) initially, but thereafter increased to 30% in 48 h. An accumulation of ³²P in the seed coats preceded its entry into the embryos. Disruption (with hot steam) of the phloem between the leaf and the pods after pulse labelling indicated that more than 80% of the ³²Pi pulse moved to the leaf before redistribution to the pods. Increasing "sink" size by adjusting the pod load from 1 to 2–3 did not increase the ³²P accumulated by the pods proportionally. Conversely, excision of the seeds after pulse labelling did not prevent translocation of ³²P out of the leaves. These results suggest that the rate of transport of phosphate to the pods at mid-late podfill is controlled primarily by factors in the leaves. The results are consistent with the observation that the relative size of the sink (pod load) does not regulate leaf senescence.     

Light and Shade Effects on Abscission and C-Photoassimilate Partitioning among Reproductive Structures in Soybean

Field experiments were conducted in 1981 and 1982 to study the effects of low-irradiance supplemental light on soybean (Glycine max [L.] Merr. cv Evans) flower and pod abscission. Cool-white and red fluorescent lights illuminated the lower part of the soybean canopy during daylight hours for 3 weeks late in flowering. At the same time, flowers and young pods on half the plants were shaded with aluminum foil. Flowers were tagged at anthesis and monitored through abscission or pod maturity.

Responses to red and white lights were similar. Supplemental light tended to reduce abscission and increase seed weight per node compared to natural light. Shading flowers and pods increased abscission and reduced seed weight per node. Number of flowers produced per node, individual seed weight, and seeds per pod were not affected by light or shade treatments.

Further studies examined the effects of shading reproductive structures on their capacity to accumulate ¹⁴C-photoassimilates. Individual leaves were pulse labeled with ¹⁴CO₂ 1, 2, and 4 weeks post anthesis. Flowers and pods in the axil of the labeled leaf were covered with aluminum foil 0, 24, 72, and 120 hours before pulsing.

Shading flowers and pods resulted in a 30% reduction in the relative amount of radiolabel accumulated from the source leaf. The reduction in ¹⁴C accumulation due to shading was evident regardless of the length of the shading period and was most pronounced when the shades were applied early in reproductive development. We conclude that light perceived by soybean flowers and young pods has a role in regulating both their abscission and their capacity to accumulate photoassimilates.