Source-sink Relationships, Seed Sucrose Levels and Seed Growth Rates in Soybean

Field experiments using two soybean (Glycine max L. Merrill)cultivars (‘Elgin 87’ and ‘Essex’) wereconducted for 2 years near Lexington, KY, USA to evaluate theeffect of source-sink alterations on seed carbohydrate statusand growth. Sucrose concentrations in developing cotyledonsof control plants were consistently low (<50 m M) early inseed development, but they increased to 100–150 m M byphysiological maturity. The concentrations increased in bothyears by 47 to 59% when 90% of the pods were removed from ‘Elgin87’, but the increase had no effect on individual seedgrowth rate (SGR). Shading (80%) reduced cotyledon sucrose levelsand SGR in both years. The critical cotyledon sucrose concentration(the concentration providing 80% of the maximum cotyledon growthrate) was estimated fromin vitro cotyledon growth at sucroseconcentrations of 0–200 m M. These critical concentrationsvaried from 72–124 m M;in planta control cotyledon sucroseconcentrations were below this critical level during the firsthalf of seed growth but exceeded it in the later stages of growthin all experiments. The estimated critical concentration wasconsistent with the failure of in planta SGR to respond to anincrease in assimilate supply and with the reduction in SGRassociated with a decrease in assimilate supply. The resultssuggest that soybean SGR is generally sink limited if photosynthesisincreases during seed filling, but source limited if photosynthesisis reduced. Copyright 2001 Annals of Botany Company Glycine max(L.) Merrill, soybean, source-sink ratios, sucrose, starch, depodding, shade, in vitro culture     

Control of Seed Growth in Soya Beans [Glycine max (L.) Merrill]

The seed is the primary sink for photosynthate during reproductivegrowth and an understanding of the mechanisms controlling therate of seed growth is necessary to understand completely theyield production process. The growth rate of individual seedsof seven soya bean [Glycine max (L.) Merrill] cultivars withgenetic differences in seed size varied from 10.8 to 3.9 mgseed^–1 day^–1. The growth rates were highly correlatedwith final seed size. The growth rate of cotyledons culturedin a complete nutrient medium was highly correlated with thegrowth rate of seeds developing on the plant and with finalseed size. The number of cells per seed in the cotyledons variedfrom 10.2 to 5.7 x 10⁶ across the seven cultivars. The numberof cells per seed in the cotyledons was significantly correlatedwith final seed size and the seed growth rate both on the plantand in the culture medium. The data suggest that genetic differencesin seed growth rates are controlled by the cotyledons and thenumber of cells in the cotyledons may be the mechanism of control. Glycine max L., soya bean, seed size, growth rate, cell number, sink activity     

Partitioning and Utilization of Photosynthate Produced at Different Growth Stages after Anthesis in Soybean (Glycine max L. Merr.): Analysis by Long-term 13C-Labelling Experiments

Yamagata, M., Kouchi, H. and Yoneyama, T. 1987. Partitioningand utilization of photosynthate produced at different growthstages after anthesis in soybean (Glycine max L. Merr.): Analysisby long term ¹³C-labelling experiments.—J. exp. Bot. 38:1247–1259. Soybean (Glycine max L. Merr. var. Akishirome) plants were allowedto assimilate ¹³CO₂ with a constant specific activity for 10h at different growth stages (a total of seven times at aboutone week intervals) after anthesis. The plants were harvestedperiodically until the time of full maturity and the partitioningof ¹³C into individual plant parts was investigated with anemphasis on the contribution of carbon assimilated at differentgrowth stages to the seed formation. Carbon assimilated at the middle to late seed-filling stagecontributed most to the seed production; one day contributionaccounted for 3–4% in total carbon of the seed at fullmaturity. Integrated contribution of carbon assimilated afteranthesis was estimated as 96% of the final seed carbon. An approximationbased on the temporal data of the incorporation of labelledcarbon into the seeds indicates that 77% of the final seed carboncame from direct transfer of current photosynthate from sourceleaves, which occurred within a few days after the photosyntheticfixation, while the rest originated from remobilization of carbonreserved mainly in leaves and stems plus petioles. In comparison with the total carbon accumulation in the seeds,protein carbon in the seeds was relatively more dependent onphotosynthate produced during the early period of reproductivegrowth stage, whereas lipid carbon was more dependent on photosynthateproduced during the later reproductive stage. Key words: Photosynthate partitioning, soybean (Glycine max L. Merr.), ¹³CO₂ assimilation, seed formation     

Seed growth rate in grain legumes I. Effect of photoassimilate availability on seed growth rate

The dry weight of harvested grain legume seeds is strongly related to their growth rate during the period of storage accumulation in the cotyledons, which begins approximately at the end of embryo cell division. Depodding, defoliation, shading or changes in air CO2 concentration were applied during seed filling (i.e. during the decrease in seed water concentration) to field and glasshouse-grown plants, in order to affect the source-sink ratio. The experiments involved three legume species, namely pea (Pisum sativum L.), soybean (Glycine max L. Merr.) and white lupin (Lupinus albus L.). Some treatments affected the number of abortions of less developed seeds from younger pods, but they did not significantly affect the number or the growth rate of filling seeds, demonstrating the priority of carbohydrate partitioning to filling seeds. The maximum growth rate of seeds was achieved regardless of the intra-plant competition level, and the duration of seed growth was shortened if the photosynthetic activity was not sufficient to fulfil the assimilate demand of filling seeds.     

Effect of Water Deficits on Seed Development in Soybean : II. Conservation of Seed Growth Rate

Water deficits during seed filling decrease seed size in soybean (Glycine max L.). This may result from a reduction in the supply of assimilates from the maternal plant and/or an inhibition of seed metabolism. To determine whether maternal or zygotic factors limited seed growth, we examined the effects of a plant water deficit on the supply of sucrose to and its utilization by developing embryos. Plants were grown in the greenhouse, and water deficits were imposed by withholding water for a period of 6 days during linear seed fill. When water was withheld, leaf water potential decreased rapidly, inhibiting canopy photosynthesis completely within 3 days. However, seed dry weight (nodes 7-11) continued to increase at or near the control rate. The level of total extractable carbohydrates in leaf, stem, and pericarp tissue decreased by 70, 50, and 45%, respectively, indicating that reserves were mobilized to support seed growth. Cotyledon sucrose content decreased from about 60 milligrams per gram dry weight to 30 milligrams per gram dry weight. Similarly, the concentration of sucrose in the interfacial apoplast of the cotyledons decreased from approximately 100 millimolar to 50 millimolar. However, the rate of sucrose accumulation by excised embryos, measured in a short-term in vitro assay, increased in response to the water deficit. These results indicate that both source and sink activity in soybean are altered by water deficits to maintain the flux of assimilates to the developing embryos. This may explain why seed growth is maintained, albeit for a shorter duration, when soybean is exposed to water deficits during the seed filling period.     

Soya Bean Seed Growth and Maturation In vitro without Pods

Immature Glycine max (L.) Merrill seeds, initially between 50and 450 mg f. wt, were grown and matured successfully in vitro.Excised seeds were floated in a liquid medium containing 5 percent sucrose, minerals and glutamine in flasks incubated at25 °C under 300 to 350 µE m^–2 s^–1 fluorescentlight. During 16 to 21 d in culture, seeds grew to a matured. wt of 100 to 600 mg per seed at an average rate of 5 to 25mg d. wt per seed d^–1 depending on initial size. Growthrates were maximal during the first 8 to 10 d in vitro but declinedwith loss of green colour in the cotyledons. Seed coats rupturedwith rapid cotyledon expansion during the first 2 d in culture.Embryos were tolerant to desiccation and 80 to 90 per cent germinatedif removed from culture before complete loss of green colour.The growth of excised seeds in vitro exceeded the growth ofseeds in detached pods, but when windows were cut in pods topermit direct exposure of seeds to the medium, seed growth wascomparable. Glycine max (L.) Merrill, soya bean, seed culture, seed growth, seed maturation, germination     

Protein Synthesis During Rehydration, Germination and Seedling Growth of Naturally and Precociously Matured Soybean Seeds (Glycine max)

Soybean seeds [Glycine max (L.) Merr.] synthesize de novo andaccumulate several non-storage, soluble polypeptides duringnatural and precocious seed maturation. These polypeptides havepreviously been coined ‘maturation polypeptides’.The objective of this study was to determine the fate of maturationpolypeptides in naturally and precociously matured soybean seedsduring rehydration, germination, and seedling growth. Developingsoybean seeds harvested 35 d after flowering (mid-development)were precociously matured through controlled dehydration, whereasnaturally matured soybean seeds were harvested directly fromthe plant. Seeds were rehydrated with water for various timesbetween 5 and 120 h. Total soluble proteins and proteins radio-labelledin vivo were extracted from the cotyledons and embryonic axesof precociously and naturally matured and rehydrated seed tissuesand analyzed by one-dimensional PAGE and fluorography. The resultsindicated that three of the maturation polypeptides (21, 31and 128 kDa) that had accumulated in the maturing seeds (maturationpolypeptides) continued to be synthesized during early stagesof seed rehydration and germination (5–30 h after imbibition).However, the progression from seed germination into seedlinggrowth (between 30 and 72 h after imbibition) was marked bythe cessation of synthesis of the maturation polypeptides followedby the hydrolysis of storage polypeptides that had been synthesizedand accumulated during seed development. This implied a drasticredirection in seed metabolism for the precociously maturedseeds as these seeds, if not matured early, would have continuedto synthesize storage protein reserves. Glycine max (L.) Merr, soybean, cotyledons, maturation, germination/seedling growth     

Growth Analysis of Soybean Seedlings During the Lifespan of the Cotyledons

Patterns of seedling growth of Glycine max in light and darknesswere compared during the period from germination to cotyledonabscission. Fitted growth curves and the derived functions,relative growth rate, unit leaf rate, leaf area ratio and specificleaf area, were used to assess the relative importance in seedlinggrowth of cotyledon storage reserves, cotyledon photosynthesisand leaf photosynthesis. The cotyledons are of an intermediatetype with a predominant storage and a minimal photosyntheticfunction. Cotyledon reserves support seedling growth until theprimary leaves expand, after which growth depends on leaf photosynthesis. Glycine max (L.) Merr., soybean, cotyledons, growth analysis, seedling development     

Osmotic Stress as a Pregrowth Procedure for Cryopreservation: 1. Growth and Ultrastructure of Sycamore and Soybean Cell Suspensions

Sycamore and soybean cell suspensions were subjected to osmoticstress by culturing for one passage in media supplemented with6 per cent mannitol or sorbitol. The effects on growth wereto reduce cell number and biomass (d. wt) production throughoutthe culture period by about 30 per cent. Ultrastructural studiesat the early exponential phase of culture growth indicated similarreductions in cell wall thickness in both species. Osmoticallystressed sycamore cells became less vacuolate, but no such changeoccurred in stressed soybean cells. Acer pseudoplatanus L., sycamore, Glycine max L. var. Biloxi, soybean, suspension culture cells, osmotic stress, growth, ultrastructure     

Enhanced Nitrogen Fixation Increases Net Photosynthetic Output and Seed Yield of Hydroponically Grown Soybean

Ten to 20% of the net photosynthetic output of a tropical grainlegume may be consumed by the nodulation-nitrogen-fixation process.If plant growth activities during the reproductive phase werelimited by photosynthetic output, enhanced nitrogen fixationwould seemingly lower total plant mass and seed yield. To testthis possibility, soybean [Glycine max (L.) Merr.] plants weregrown hydroponically on nutrient medium supplemented with minimalurea or with an excess of either nitrate or nitrate plus urea.Acetylene reduction activities (i.e. nitrogen fixation rates)and transpiration rates were measured twice weekly on theseplants through pod fill. Of the plants inoculated, those grownon minimal urea revealed significantly greater acetylene reductionactivities and transpiration rates. At maturity, plants thathad fixed nitrogen at a rapid rate during pod fill had a significantlygreater seed size, total plant mass (i.e. net photosyntheticrate) and nitrogen content than uninoculated or poorly nodulatedplants grown on an excess of nitrate. It is concluded, therefore,that a rapid rate of nitrogen fixation during pod fill enhancesboth transpiration and net photosynthetic output. The increasedavailability of usable nitrogen (i.e. ureides), coupled withenhanced transpiration and photosynthetic output, significantlyincreases total plant mass and seed yield. Thus, enhanced nitrogenfixation seems to be an inexpensive means of increasing seedyield of soybean and perhaps of other tropical grain legumes. Key words: Glycine max, nodulation, nitrate, urea     

Interrelationship Between Plant Developmental Stage, Plant Growth Rate, Nitrate Utilization and Nitrogen Fixation in Hydroponically Grown Soybean

The growth rate of the indeterminate soybean plant [Glycinemax (L.)Merr.] slows as it proceeds from vegetative phase intoreproductive growth. Yet, the well-nodulated plant acquiresmost of its nitrogen during reproductive growth. Thus, the interrelationshipbetween plant developmental stage and nitrogen fixation wasexamined. It is shown that, regardless of the age of the hydroponicallygrown soybean plant at the time of its inoculation with Bradyrhizobiumjaponicum, the highest rate of nitrogen fixation occurs duringthe pod-filling stage (R5). Nevertheless, maximum total nitrogenfixation is generally achieved when inoculation occurs at thefull-bloom stage (R2). It is shown, however, that flower budsand flowering are not responsible for the enhanced nodulationand nitrogen fixation. Rather, the data suggest that the onsetof rapid nodulation occurs soon after the initiation of thedevelopmentally programmed drop in foliar nitrate reductaseactivity. The ensuing increase in nitrogen fixation providesthe plant with much of its needed nitrogen and hence stimulatesplant mass accumulation during pod-fill. It is suggested thatnitrogen fixation enhances growth of the soybean plant by increasingits net photosynthetic efficiency during reproductive growthand by providing the needed nitrogen at the appropriate timefor maximum seed growth. Key words: Glycine max, nitrate, nitrogen fixation, nodulation     

Changes in Seed Constituents During Germination and Seedling Growth of Precociously Matured Soybean Seeds(Glycine max)

During 7 d of precocious maturation of soybean seed (Glycinemax), the starch content declined and soluble sugar levels increasedin patterns similar to natural seed dehydration and maturation.Total seed protein content and total seed dry weight increasedwhereas oil content remained relatively unchanged. Overall,the proportions of the constituents in precociously maturedseeds were comparable to naturally mature seeds. Precociouslymatured soybean seeds showed much the same germination and seedlinggrowth frequency patterns as naturally matured seeds. Duringgermination and seedling growth of precociously matured seeds,starch, soluble sugar, protein and oil levels followed patternssimilar to naturally mature, germinating seeds and seedlings.Therefore, precocious maturation may be used as a model systemto investigate the control of the physiological and biochemicalevents occurring during seed maturation which lead to germinationand subsequently, seedling growth. Glycine max (L.) Merr., soybean, cotyledons, maturation, germination/seedling growth     

Cytokinin Regulation of Flower and Pod Set in Soybeans (Glycine max(L.) Merr.)

Exogenous application of cytokinin to raceme tissues of soybean(Glycine max(L.) Merr.) has been shown to stimulate flower productionand to prevent flower abortion. The effects of these hormoneapplications have been ascertained for treated tissues, butthe effects of cytokinins on total seed yields in treated plantshave not been evaluated. Our objectives were to examine theeffects of systemic cytokinin applications on soybean yieldsusing an experimental line of soybeans, SD-87001, that has beenshown to be highly sensitive to exogenous cytokinin application.Soybeans were grown hydroponically or in pots in the greenhouse,and 6-benzylaminopurine (BA) was introduced into the xylem streamthrough a cotton wick for 2 weeks during anthesis. After theplants had matured, the number of pods, seeds per pod, and thetotal seed weight per plant were measured. In the greenhouse,application of 3.4 x 10^-7 moles of BA resulted in a 79% increasein seed yield compared with controls. Results of field trialsshowed much greater variability within treatments, with consistent,but non-significant increases in seed number and total yieldsof about 3%. Data suggest that cytokinin levels play a significantrole in determining total yield in soybeans, and that increasingcytokinin concentrations in certain environments may resultin increased total seed production. Copyright 2001 Annals ofBotany Company Glycine max, soybean, flower abortion, cytokinin, 6-benzylaminopurine, hydroponic, seed yield, wicking     

Seed Maturation in Soybeans (Glycine max L. Merr.) is Independent of Seed Mass and of the Parent Plant, Yet is Necessary for Production of Viable Seeds

Freshly harvested seeds of soybean (Glycine max L. Merr.) orseeds shelled, then dried, were non-viable. Seeds dried in intactpods, even when only 17% of normal size, matured into viableseeds and produced healthy plants. These seeds maintained activityof various enzymes but gained little soluble protein while air-dryingin intact pods. There was a qualitative change in seed proteinsassociated with maturation. Seeds matured in intact pods havea greater proportion of protein as slow-moving bands and havecompletely lost one fast-moving band compared with seeds shelledbefore drying. Seed maturation is a distinct phase of seed production,is independent of the parent plant, and can be imposed on theseed at many stages of development.     

Changes in Dry Matter, Carbohydrate and Seed Yield Resulting from Lodging in Three Temperate Grass Species

The adverse effect of lodging on grass seed yield may be attributed,in part, to assimilate limitation during the seed filling period.This investigation examined plant dry matter assimilate partitioningand seed yield as affected by lodging in three species thatare closely related but phenotypically different: tall fescue(Festuca arundinacea Schreber.), Italian ryegrass (Lolium multiflorumLam.), and perennial ryegrass (L. perenne L.). Studies wereperformed in field plots at Corvallis, Oregon, USA. Seed yieldcomponents (seed number per inflorescence, seed yield per inflorescence,and single seed mass) and leaf, stem (lower, middle, and peduncle)and seed inflorescence dry mass were measured just prior toanthesis to seed maturity. Dry mass and water soluble carbohydrates(WSC) were determined for shoot components. The reduction indry mass and WSC in leaves and stem components following anthesiswas often greater in lodged plants compared to upright plants.The relatively low seed yield depression in lodged tall fescuesuggested a higher compensation potential for partitioning reserveassimilate from leaves and stems to support seed growth anddevelopment. This potential does not appear to be present tothe same degree in Italian ryegrass and to an even lesser extentin perennial ryegrass. These findings suggest that the potentialto compensate for reduced assimilate supply during the periodof high assimilate demand by seeds may be attributed, in part,to the total assimilate reserve accumulated prior to photoassimilatereduction caused by the lodged condition. Copyright 2000 Annalsof Botany Company Tall fescue, Festuca arundinacea Schreber., Italian ryegrass, Lolium multiflorum Lam., perennial ryegrass, L. perenne L., assimilate partitioning, source–sink     

Effect of Removal of Flower Buds, Open Flowers, Young Pods and Shoot Apex on Growth and Pod Set in Soybean

The complete removal of the reproductive structures once andshoot apices of soybeans (Glycine max L. Merrill) during earlyanthesis but before the rapid seed development stage significantlyincreased flowering and pod set in greenhouse and field grownplants. The treated plants had darker green leaves, shorterstems and petioles and retained their chlorophyll content longerthan control plants. Pod maturation was also delayed. Althoughdecapitation and the removal of reproductive structures increasedthe number of 3- or more-seeded pods in all varieties tested,seed weight per plant was not consistently increased. The possibleinvolvement of endogenous hormones in pod set and multi-loculepod production in soybeans is discussed. Key words: Glycine max (L.) Merr, Reproductive structures, Shoot apex, Growth, Flowering, Pod set, Multi-locule pods     

Germination and Early Growth of Plants Studied Using Neutron Radiography

Seed swelling, germination, root extension, lateral root initiationand shoot growth were studied in soils of different water contents,using non-destructive, serial neutron radiography. Seeds fromthree varieties of soya beans (Glycine max L.) and one varietyeach of maize (Zea mays L.) and vetch (Vicia sativa L.) wereused. The seeds germinated when they had increased in size bya certain amount, if germination is taken as the time when theradicle first appears. The rate at which roots and shoots extendalso depend on soil water content. Glycine max L., Vicia sativa L., Zea mays L., Soya bean vetch, maize, seed germination, root extension, lateral root initiation, neutron radiography     

Effect of Assimilate Supply on Development and Growth Potential of Axillary Buds in Roses

The effect of assimilate supply on axillary bud developmentand subsequent shoot growth was investigated in roses. Differencesin assimilate supply were imposed by differential defoliation.Fresh and dry mass of axillary buds increased with increasedassimilate supply. The growth potential of buds was studiedeither by pruning the parent shoot above the bud, by graftingthe bud or by culturing the bud in vitro. Time until bud breakwas not clearly affected by assimilate supply during bud development,Increase in assimilate supply slightly increased the numberof leaves and leaf primordia in the bud; the number of leavespreceding the flower on the shoot grown from the axillary budsubstantially increased. No difference was found in the numberof leaves preceding the flower on shoots grown from buds attachedto the parent shoot and those from buds grafted on a cutting,indicating that at the moment of release from inhibition thebud meristem became determined to produce a specific numberof leaves and to develop into a flower. Assimilate supply duringaxillary bud development increased the number of pith cells,but the final size of the pith in the subsequent shoot was largelydetermined by cell enlargement, which was dependent on assimilatesupply during shoot growth. Shoot growth after release frominhibition was affected by assimilate supply during axillarybud development only when buds sprouted attached to the parentshoot, indicating that shoot growth is, to a major extent, dependenton the assimilate supply available while growth is taking place.Copyright1994, 1999 Academic Press Assimilate supply, axillary bud, cell number, cell size, defoliation, development, growth potential, meristem programming, pith, Rosa hybrida, rose, shoot growth     

Seed Water Relations and the Regulation of the Duration of Seed Growth in Soybean

Soybean [Glycine max (L.) Merrill] seeds and cotyledons weregrown in an in vitro culture system to investigate the relationshipsbetween cell expansion (net water uptake by the seed) and drymatter accumulation. Seeds or cotyledons grown in a completenutrient medium containing 200 mol m^–3 sucrose continueddry matter accumulation for up to 16 d after in planta seedsreached physiological maturity (maximum seed dry weight). Seedor cotyledon water content increased throughout the cultureperiod and the water concentration remained above 600 g kg^–1fresh weight. These data indicate that the cessation of seeddry matter accumulation is controlled by the physiological environmentof the seed and is not a pre-determined seed characteristic.Adding 600 mol m^–3 mannitol to the medium caused a decreasein seed water content and concentration. Seeds in this mediumstopped accumulating dry matter at a water concentration ofapproximately 550 g kg^–1. The data suggest that dry matteraccumulation by soybean seeds can continue only as long as thereis a net uptake of water to drive cell expansion. In the absenceof a net water uptake, continued dry matter accumulation causesdesiccation which triggers maturation. Key words: Glycine max (L.) Merrill, solution culture, duration of seed growth, water content, dry matter accumulation     

Remobilization patterns of C and N in soybeans with different sink-source ratios induced by various night temperatures

The effects of increased sink-source ratios, induced by elevating night temperatures, on remobilization of ¹⁴C-assimilates and N within field-grown soybeans (Glycine max [L.] Merr.) was investigated from preflowering to maturity. Raising the mean minimum night temperature for the entire growing season from 10 (check, uncontrolled) to 16°C increased seed growth without appreciable effect on final leaf area. Increasing this temperature to 24°C increased seed growth and reduced final leaf area. Leaves, stems, petioles, and pods acted as intermediate storage sites for ¹⁴C assimilates. Only plants with higher night temperatures remobilized some of the stored assimilates during the period of rapid seed growth. Even the seeds in the 24°C plants with the largest sink-source ratios did not utilize all the C-assimilates potentially available for remobilization. Nitrogen was readily remobilized from petioles, stems, and pods of all treatments as early as the beginning of seed development, but from the leaves only during late seed-filling. However, only plants with elevated night temperatures tended to remobilize all of the available N from vegetative tissues and pods. We concluded that a larger portion of stored assimilates may be remobilized to the seed if a strong seed sink can be sustained. It also appeared that with increasing sink-source ratios, N shortage might limit seed yield before a lack of C-assimilates would. A proposed model for soybean assimilate demand, distribution, partitioning, and remobilization is presented.