Periods of Shoot Chilling Sensitivity in Soybean Flower Development, and Compensation in Yield after Chilling

Soybeans [Glycine max (L.) Merr. cv. Ransom] grown at a constant25 °C were placed in a 12-h inductive photoperiod at twoweeks of age. Subgroups were shoot-chilled for one week at aconstant 10 °C during each of the first four weeks of floralinduction. Controls were photoinduced but not chilled. Chillingduring the first week of photoinduction inhibited productionof floral primordia, but did not increase the abscission rateof flowers and pods. Chilling during the second week did notaffect primordium production or abscission rate, but did causea significant increase in numbers of fused and malformed pods.Chilling during the third week caused loss of 77 per cent ofearly flowers and pods by abscission, while fourth week chillingcaused less severe losses by abscission. Inhibition of vegetativegrowth may have been responsible for primordium loss in first-weekplants, while disturbances in the development of flowers wereresponsible for the losses in the other chilling weeks. Althoughchilling during the first and third photoinduction weeks causeda significant reduction in early pod numbers, plants harvestedat 16 weeks of age showed no significant loss in seed yield.Low abscission rates late in pod filling and increased weightof individual seeds compensated for early losses of pods. Thesecompensatory responses to a chilling-induced loss of pods aresimilar to those reported for mechanically depodded soybeans. Glycine max (L.) Merr., soybean, temperature, chilling, floral initiation, anthesis, abscission, yield, compensation     

Density Studies on Lupins. II. Components of Seed Yield

Components of seed yield of cv. Ultra (Lupinus albus L.) andcv. Unicrop (L. angustifolius L.) were measured when grown atthree densities. The low density (10 plants m^–2) Unicropyield (34 g seed per plant) was 1.8 times that of Ultra as ithad more branches, pods and seeds per pod. Ultra seeds (310mg per seed) were heavier than Unicrop seeds (180 mg). The branchingpattern of Ultra was less dependent on plant density, henceat 93 plants m^–2 it gave a higher per plant yield (7.4vs 6.4 g) than Unicrop at lower densities (83 plants m^–2).Density had most influence on pod formation and only small effectson seeds per pod and seed weight. Yield components on the main-steminflorescence were influenced less by density than componentson branch inflorescences. Later formed, higher order generationsof inflorescences were most affected by increased inter- andintra-plant competition. Pod numbers on the main-stem were similarfor both species. Pods formed at higher flower nodes in Unicrop,but the lower flower nodes were less fertile than those in Ultra.Node position of flowers had no influence on seed set in main-stemUnicrop pods, but pods from higher nodes in Ultra formed fewerseeds. Seed weights in Unicrop were similar among main-stemnodes but in Ultra seed weights tended to increase at highernodes. Lupinus spp, lupins, seed yield, planting density     

A Flower and Pod Staging System for Soybean

Flower and pod abscission limit soybean yield. A system forquantifying flower and pod development based on the morphologicalappearance of the flower prior to and following anthesis hasbeen developed to aid in studies of pod abscission. Changesin the appearance of the corolla, primarily the banner petal,are used to distinguish the different stages of the system.External pistil dimensions have been correlated with internalfeatures for each stage of development. From anthesis to podset, pistil length and weight increase almost two- and fivefold,respectively, and ovule development progresses from unfertilizedegg cells to embryos surrounded by cellular endosperm. Pod determinedare correlated with ovule length and width and embryo cell number.Flower and pod stages can be determined in situ, thus permittingnon-destructive observation and experimental manipulation offlowers or pods without necessarily impeding their development.Stages have been identified that indicate precisely when podset occurs and when young pods cease growing and ultimatelyabscise. This system of flower and pod staging is useful instudies designed to assess effects of abiotic or biotic stressand genetic factors on pod set and abortion. Abscission, anthesis, Glycine max (L.) Merr, embryo development, pod set     

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     

Characters of Pod Anatomy Associated with Resistance to Pod-Shattering in Soybean

Seven characteristics of pod anatomy were studied for theirassociation with resistance to pod-shattering in 16 soybean[Glycine max (L.) Merrill] varieties and strains. The thicknessand length of the bundle cap on the dorsal side of the pod andpod-wall thickness were found to be significantly negativelycorrelated with the degree of pod-shattering. Further statisticalanalyses confirmed that these three anatomical characters werealmost equally important and could potentially serve as criteriafor the selection of resistance to pod-shattering. The identifiedtraits/sites in the pod represent sclerenchymous structuresand may provide the structural basis of resistance to pod-shatteringin soybean.Copyright 1995, 1999 Academic Press Pod, anatomy, shattering, soybean, Glycine max (L.) Merrill     

Defoliation and its Effects on Pod and Seed Development in Oil Seed Rape (Brassica napus L.)

Both in vivo and in vitro techniques have been used to followthe development of individual pods on the terminal inflorescenceof undefoliated and defoliated plants of oil-seed rape (Brassicanapus cv. Maris Haplona). For any pod, a rapid increase in podlength occurred between 2 d and 8 d after flower opening andthis preceded by approximately 2 d the increase in pod width,the rate of which was less than that for length. An increasein the diameter of individual seeds coincided with the increasein pod width. Regional increases in the length and width ofpods were associated with the presence of developing seeds inthese regions. Key words: Brassica napus L., Development, In vivo, Pod and seed, Stress     

Effects of pod removal on metabolism and senescence of nodulating and nonnodulating soybean isolines: I. Metabolic constituents

Field studies were conducted in 1981 and 1982 to ascertain the effects of pod removal on senescence of nodulating and nonnodulating isolines of soybean (Glycine max [L.] Merr. cv Harosoy) plants. Specifically, the test hypothesis was that nodules act as a nitrogen source and a carbohydrate sink which would in turn prevent or delay senescence in the absence of pods. Senescence was judged by changes in metabolite levels, in dry matter accumulation, and by visual observation.

For both nodulated and nonnodulated plants, pod removal had no effect on the magnitude or rate of dry matter and reduced-N accumulation by whole plants. Phosphorus accumulation was significantly less in both nodulated- and nonnodulated-depodded plants, compared with respective control plants with pods. These data suggested a role for pods in phosphorus uptake. Accumulation of dry matter, reduced N, and phosphorus ceased at approximately the same time for all treatments.

Pod removal did affect partitioning of plant constitments, with leaves and stems of depodded plants serving as a major alternate sink for accumulation of dry matter, reduced N, phosphorus, and nonstructural carbohydrates (primarily starch). While depodded plants eventually lost a significant amount of leaves, leaf drop was delayed relative to plants with pods; and depodded plants still retained some green leaves at 2 weeks past grain maturity of control (podded) plants.

The results indicated that senescence patterns of soybean plants were the same for nodulated and nonnodulated plants, and that pods did not control the initiation of senescence, but rather altered the partitioning of plant constituents and the visual manifestations of senescence.

     

Development of Oilseed Rape Buds, Flowers, and Pods In Vitro

A technique for growing buds, flowers, and pods of oilseed rape(Brassica napus L. cv. Haplona) on stem explants in vitro hasbeen developed. Open flowers and young pods underwent normaldevelopment on a basal medium of minerals, vitamins, and sucrosebut the development of buds was less successful. Young buds(3 mm long) did not develop and only limited development ofthe older buds (5 mm long) took place. Some 3 mm-long buds wereinduced to develop to open flowers by adding naphthyl aceticacid or gibberellic acid. Pod and seed set in open flowers werenot affected by adding plant growth substances to the medium,but pod elongation and pod dry weight were promoted by gibberellicacid, 105 M, and benzyl amino purine, 10⁷ M, respectively. Reducingthe supply of sucrose or minerals to open flowers reduced seedset, pod elongation and pod weight but did not affect pod set.The physiological significance of the results is discussed. Key words: In vitro cultures, oilseed rape, pod development, flower development     

Factors Affecting Shedding of Flowers in Soybean (Glycine max (L.) Merrill)

Flower shedding in soybean, Glycine max (L.) Merrill, was studiedusing cultivar ‘Clark’, isoline E₁t, which has relativelylong racemes for convenient identification and observation ofindividual flowers. On each raceme studied, pod set was greatestat the proximal (basal) positions, whereas shedding was greatestat the most distal positions. When proximal flowers were removedas they reached anthesis, pod set increased at the more distalpositions. Pod set was increased in some instances by applicationof water directly to the ovaries as a drop in the calyx cup.Peroxidase activity changed in parallel with ovary development,increasing rapidly in growing pods but not in shedding flowers.Increases in flower peroxidase was mainly in ovary walls. Flowerstaken at or near anthesis from positions with high percent podset could be grown in vitro with especially good ovary enlargement,whereas ovaries in flowers taken from positions of low pod setdid not enlarge in culture. Unidentified substances were extracted from young pods which,when incorporated into lanolin and tested in an in situ bioassay,could mimic the effect of proximal flowers in inducing sheddingof distal flowers. Indole-3-acetic acid resembled the extractedmaterials in inducing shedding, but differed by eliciting side-effectsthat extracts did not. The growth substances abscisic acid,gibberellic acid, and benzyladenine did not promote sheddingin the in situ test. The evidence was taken to indicate that soybean flower sheddingis induced in distal flowers by substances from the more proximal,fertilized ovaries, and that this is possibly due to interferencewith some of the intense metabolic changes that follow pollinationand fertilization.     

Carbon Exchange Rate of Intact Individual Soya Bean Pods. 1. Response to Step Changes in Light and Temperature

Carbon exchange rates (CER) of individual intact field-grownsoya bean [Glycine max (L.) Merr.] pods were measured continuouslywith a mobile gas analysis laboratory. Conditions in pod chamberssimulated those experienced normally by pods except for experimentalmodification of incident radiation or pod temperature. Undernormal conditions, CER (where positive CER represents CO₂ evolution)fluctuated diurnally with a morning rise followed by a slowafternoon and evening decline which was similar among pods whichwere measured simultaneously. The frequency of measurementspermitted detection of rapid CER responses to step changes inlight and pod temperature. CER rapidly decreased and increasedwhen the chamber was alternately exposed to full sunlight andcomplete darkness, respectively. CER responded similarly tosteps up [from ambient to elevated (+ 10°C) temperature]and steps down (from elevated to ambient temperature), respectively.Thus, a temperature sensitive process which regulated pod CERwas located within the pod. CER ranged from less than 0·1to more than 1·2 mg CO₂ h^–1 pod^–1 over theperiod of rapid dry-matter accumulation. Glycine max (L.) Merr., Soya bean, carbon exchange rate, light, temperature     

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.

     

The Influence of Genes ar and n on Senescence in Pisum sativum L.

The effect of genes ar (violet flowers, small hilum) and n (thick,fleshy pod wall) on whole plant in senescence peas was examinedby comparing Ar- with arar and N- with nn plants in segregatingprogenies. Homozygosity for ar or n significantly delayed the time whenthe plants were ready for harvest of their entire seed crop.These genes did not delay either the onset of reproduction orthe onset of apical arrest in the first instance. However, whereasAr- N- plants almost invariably senesced and died as the firstseed crop matured, the majority of arar and/or nn plants entereda period of secondary growth and a further fruiting cycle. Comparedwith Ar- plants, arar plants had over twice as many pods andseeds but individual seeds were 58 per cent lighter and totalseed yield (wt) was 19 per cent less. Pod length was unaffected.Compared with N-plants, nn plants had shorter pods (16 per cent),fewer seeds per pod (21 per cent), smaller seeds (20 per cent)and a lower total seed yield (wt 14 per cent less). It appearsthat ar and n impose a lower metabolic drain per reproductivenode as a consequence of their effects on hilum anatomy andpod morphology, respectively. These mutants disrupt the normalpattern of monocarpic senescence by breaking the coordinationbetween apical arrest and subsequent events. The developingseed crop delimited by the first arrest fails to cause plantdeath, possibly because sink size is less than in normal counterparts. Pisum sativum L, garden pea, senescence, hilum, pod, seed size, genetics     

In Vivo Fixation of CO2 by Attached Pods of Brassica campestris L.

In vivo net CO₂ exchange characteristics of attached Brassicapods were studied during the entire period of their growth anddevelopment after anthesis. ¹⁴CO₂ was fed both from the externalatmosphere and internally through the pod cavity, and the anatomyof the pod-wall was examined microscopically. Stomata were observedin the outer epidermal layer of the pod wall. Net in vivo CO₂fixation by the pods was observed throughout the period of theirdevelopment and was maximum on day 42 after anthesis (DAA).Compared to the internal feeding experiments, ¹⁴CO₂ fixationfrom the external environment was very high. Apparent translocationof fixed carbon from the pod wall to seeds was rapid. Pod photosynthesiscontributed substantially to seed growth. pods, Brassica campestris L, CO₂ fixation, stomata     

Rate of Increase in Size and Dry Weight of Individual Pods of Field Grown Soya Bean Plants

The dry weight of the whole fruit, the pod wall and an enclosedseed of randomly harvested soya beans is estimated from theexternal dimensions of the attached pod. The relations betweendimensions and dry weight are independent of cultivar and growthcondition and can be used on pods from 1 cm in length untilthe seeds reach their maximum fresh weight. Dimensions of tagged pods of three cultivars of field grownsoya beans differing in time to reach maturity were measuredevery 2–3 days from initial pod elongation until maturation.Dry weights for each pod were estimated from the dimensions,and the dry weight accumulation with time was fitted to thelogistic function to find the growth rate that best characterizedthe data for each pod. The final weight, the specific growthrate and the maximum growth rate of the whole fruit, the podwall and a single seed were subjected to analysis of variance. The most significant difference between pods of these cultivarswas the specific growth rate of individual seeds, which decreasedwith increasing maturity group. There were no differences ingrowth of the pod wall. However, most of the variation was betweenindividual pods within a cultivar, where the rate of dry weightaccumulation of the whole fruit, governed largely by the seedgrowth rate times the number of seeds, was highly correlatedwith the earlier growth of the pod wall. This suggests thatthe growth of individual whole fruit was determined early inpod development and was slightly influenced by factors appliedduring the period of rapid seed growth. Glycine max (L.) Merrill, Soya bean, seed growth analysis, specific growth rate     

In Vivo Fixation of CO2 by Attached Pods of Brassica campestris L.

In vivo net CO₂ exchange characteristics of attached Brassicapods were studied during the entire period of their growth anddevelopment after anthesis. ¹⁴CO₂ was fed both from the externalatmosphere and internally through the pod cavity, and the anatomyof the pod-wall was examined microscopically. Stomata were observedin the outer epidermal layer of the pod wall. Net in vivo CO₂fixation by the pods was observed throughout the period of theirdevelopment and was maximum on day 42 after anthesis (DAA).Compared to the internal feeding experiments, ¹⁴CO₂ fixationfrom the external environment was very high. Apparent translocationof fixed carbon from the pod wall to seeds was rapid. Pod photosynthesiscontributed substantially to seed growth. pods, Brassica campestris L., CO₂ fixation, stomata     

Soya Bean Seed Growth and Maturation by In Vitro Pod Culture

Immature Glycine max (L.) Merr. seeds initially at 50–70mg fresh weight were successfully grown and matured in vitroin detached pods. Surface sterilized pods were floated in aliquid medium containing 5 per cent sucrose, minerals, and glutaminein 125 ml Erlenmeyer flasks and incubated at 25 °C under350–400 µE m^–1 s^–1 white light. Seedswhich were matured in vitro increased tenfold in dry weight,were visually similar to commercial seeds of the same size,were tolerant to desiccation and germinated with normal seedlinggrowth. Excised pods transported dye from the pedicel to thegrowing seed within 120 min. Soya bean pod culture is a usefultechnique to study the influence of single or combinations ofchemical or environmental parameters on regulation of seed growth,seed maturation, and subsequent germination events without theconfounding interactions with the mother plant. Glycine max (L.) Merr., soya bean, pod culture, seed culture, seed growth, seed maturation, germination     

Comparisons of Earlier- and Later-formed Pods of Chickpeas (Cicer arietinum L.)

In chickpeas (Cicer arietinum L.) flowering and pod developmentproceed acropetally. In plants grown under normal field conditionsat Hyderabad, in peninsular India, and at Hissar in north India,at successively apical nodes of the branches there was a declinein pod number per node, weight per pod, seed number per podand/or weight per seed. The percentage of nitrogen in the seedswas the same in earlier and later-formed pods at Hyderabad;at Hissar the later-formed seeds contained a higher percentage.Earlier- and later-formed flowers contained similar numbersof ovules. The decline in seed number and/or weight per seedin the later-formed pods of 28 out of 29 cultivars indicatedthat pod-filling was limited by the supply of assimilates orother nutrients. By contrast, in one exceptionally small-seededcultivar there was no decline in the number or weight of seedsin later-formed pods, indicating that yield was limited by ‘sink’size. Cicer arietinum L., chickpea, flowering, pod development, seed number, seed weight, nitrogen content     

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.     

The Physiological Basis for Cytokinin Induced Increases in Pod Set in IX93-100 Soybeans

Previous investigations have shown the feasibility of increasing pod number on legumes by the application of 6-benzylaminopurine (BA) directly to the raceme. These investigations were designed to determine what reproductive parameter was affected by cytokinin application, and if these applications were overcoming a deficiency in root-produced cytokinins during late flowering. Five individual main stem racemes on greenhouse grown soybeans (Glycine max L. Merr.) were treated with 2 millimolar BA. A single application of BA when pods appeared at 25 to 50% of the proximal floral positions resulted in a 58% increase in pod set due primarily to a 33% reduction in floral abscission. Applications of BA at later intervals also resulted in significant reductions in total abscission. When three applications of BA were imposed on the upper five nodes of field grown soybeans, total pod number and seed weight were significantly increased in this section of the canopy by 27 and 18%, respectively. Throughout the flowering period, root pressure exudate was sampled for the subsequent separation and quantification of zeatin, dihydrozeatin, zeatin riboside, dihydrozeatin riboside, and isopentenyladenine. Total cytokinin flux peaked from 0 to 9 days after flowering began, and then dropped to one-half of this level by 15 days postanthesis. The probability that a flower would initiate a pod was directly related to the concentration of total cytokinins present in the exudate when the flower opened.     

Intersubgeneric hybridization of soybeans with a wild perennial species,Glycine clandestina Wendl

Summary The exploitation of wild perennial species of subgenus Glycine has been formidable in soybean breeding programs because of extremely poor crossability and an early pod abortion. The combination of gibberellic acid application to hybridized gynoecia and improved seed culture media formulations resulted in a new intersubgeneric hybrid between Glycine max (L.) Merr. (2n=40) and G. clandestina Wendt. (2n=40). Of the 31 immature seeds cultured, 1 regenerated 21 plants through organogenesis while the remaining 30 failed to germinate. All the regenerated plants were similar morphologically, carried expected 2n=40, possessed hybrid isozyme patterns and were completely sterile. Complete absence of chromosome pairing was observed in 40.9% sporocytes. The occurrence of 1 to 6 loosely paired rod bivalents suggests some possibilities of allosyndetic pairing. Hybrid plants set aborted pods after backcrossing to G. max.