Effect of shading individual soybean reproductive structures on their abscisic Acid content, metabolism, and partitioning

Pod set in soybean is related to carbon partitioning and may be, at least partially, regulated by abscisic acid (ABA) concentrations. The studies reported here examine the relationship between carbon and ABA partitioning, reproductive abscission and ABA metabolism. The partitioning of radiolabeled ABA and photoassimilates from leaves to flowers and endogenous ABA concentrations were determined in shaded and unshaded reproductive structures. Aluminum foil was gently placed over individual soybean reproductive structures for 48 hours at 0, 4, 12, 17, and 22 days after anthesis (DAA). Shading of flowers at 12, 17, and 22 DAA resulted in significantly reduced concentration of ABA. However, shading had no effect on the catabolism of exogenously supplied [³H] ABA. The shading treatment on the first four of the five dates reduced partitioning of photoassimilates and ABA from the subtending leaf to the flower. Shading of reproductive structures also caused a significant reduction in the amount of assimilate exported from the subtending leaf, at 17 DAA. We conclude that shade-induced premature reproductive abscission in soybean is not stimulated by high levels of ABA within reproductive structures, but that ABA may inhibit abscission of reproductive structures by playing a role in preferential assimilate partitioning.     

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     

Source/Sink relations of abscising and nonabscising soybean flowers

The partitioning of recently fixed ¹⁴C to setting and abscising flowers within the axillary raceme of `Clark' isoline E1t soybeans (Glycine max L. Merr) was examined as a function of time after anthesis of individual flowers. In such racemes, the first four flowers showed a 17% abscission while the next four flowers showed 47% abscission.

Source/sink relations of flowers I-IV (normally setting) were compared to those of flowers V-VIII (normally abscising) by pulse labeling source leaves with ¹⁴CO₂ and determining the radioactivity of individual flowers after a 4-hour chase period. The relative specific activity (RSA;% disintegrations per minute per% dry weight), sink strength (% disintegrations per minute), and its components, sink size (milligrams dry weight) and sink intensity (% disintegrations per milligram dry weight) were then calculated as a function of days after anthesis.

Sink intensity (i.e. the competitive ability to accumulate photoassimilate per unit mass) was very high prior to anthesis of both setting and abscising flowers. Sink intensity then became very low for the first 3 days following anthesis after which it recovered in normally setting flowers, but failed to recover in normally abscising flowers. It is concluded that soybean reproductive abscission is determined at or very near the day of anthesis.

     

The Effect of Temperature on the Production and Abscission of Flowers and Pods in Snap Bean (Phaseolus vulgaris L.)

The effect of temperature on production and abscission of flowerbuds, flowers and pods was studied in a determinate snap-beancultivar (cv. Tenderette). Under moderate temperature (e.g.27/17°C) the onset of pod development was associated withcessation of flower bud production and with enhanced abscissionof flower buds. Raising night temperature from 17°C to 27°Cstrongly reduced pod production, mature pod size and seeds perpod, while an increase in day temperature from 22°C to 32°Chad smaller and less consistent effects. Pod production underhigh night temperature was not constrained by flower productionsince 27°C at night promoted branching and flower bud appearance.Under 32/27°C day/night temperature the large reductionin pod set was due to enhanced abscission of flower buds, flowersand young pods ( 3 cm). Flowers had the highest relative abscissionfollowed by young pods and flower buds. Therefore, the onsetof anthesis and of pod development were the plant stages mostsensitive to night temperature. Pods larger than 3 cm did notabscise but usually aborted and shrivelled under high nighttemperature. The effects of 32/27°C were not due to transientwater stresses and were observed even under continuous irrigationand mist-spraying. High temperature, flower production, pod set, seed set, abscission, snap bean, Phaseolus vulgaris L, Tenderette     

The Effect of Temperature on the Production and Abscission of Flowers and Pods in Snap Bean (Phaseolus vulgaris L.)

The effect of temperature on production and abscission of flowerbuds, flowers and pods was studied in a determinate snap-beancultivar (cv. Tenderette). Under moderate temperature (e.g.27/17 °C) the onset of pod development was associated withcessation of flower bud production and with enhanced abscissionof flower buds. Raising night temperature from 17 °C to27 °C strongly reduced pod production, mature pod size andseeds per pod, while an increase in day temperature from 22°C to 32 °C had smaller and less consistent effects.Pod production under high night temperature was not constrainedby flower production since 27 °C at night promoted branchingand flower bud appearance. Under 32/27 °C day/night temperaturethe large reduction in pod set was due to enhanced abscissionof flower buds, flowers and young pods (< 3 cm). Flowershad the highest relative abscission followed by young pods andflower buds. Therefore, the onset of anthesis and of pod developmentwere the plant stages most sensitive to night temperature. Podslarger than 3 cm did not abscise but usually aborted and shrivelledunder high night temperature. The effects of 32/27 °C werenot due to transient water stresses and were observed even undercontinuous irrigation and mist-spraying. High temperature, flower production, pod set, seed set, abscission, snap bean, Phaseolus vulgaris L., cv. Tenderette     

An Analysis of the Influence of Plant Density on the Growth of Vicia faba: II. THE SIGNIFICANCE OF COMPETITION FOR LIGHT IN RELATION TO PLANT DEVELOPMENT AT DIFFERENT DENSITIES

In a further analysis of the changing pattern of developmentinduced in Vicia faba by varying the density, it has been foundthat a reduction from a high to a low density has little influenceon the subsequent development unless such thinning is delayeduntil the flowering phase. By this time, save for widely spacedplants, the level of self shading within the population hasbecome marked. In fact, at high densities (55–65 plants/metre²)during the early-ripening phase the light intensity at groundlevel may fall to 0.03 daylight while a considerable proportionof the plant—up to 38 per cent.—may receive lessthan o-1 daylight. At low densities (11-I2 plants/metre²) theminimum intensity at ground level is 0.14 daylight and lessthan 3 per cent, of the shoot is subjected to o-1 daylight.In pot experiments, using a range of screens, it was establishedthat the compensation point is about o-I daylight. Thus, asthe density is increased the light gradient between the apexand the base becomes progressively steeper and the proportionof the leaves not actively assimilating correspondingly greater. To assess the ways in which such a light gradient operates,experiments were carried out in which either the apex or theinflorescences or leaves were removed over different sectionsof the stem or various parts of the shoot shaded and detailedrecords made of development, particularly of flower- and pod-production.Removal of the upper leaves or shading the apex primarily increasesthe rate of pod abscission after the flowers have set. Partialremoval of the inflorescences, especially at the lower nodes,has an opposite effect, while decapitation, though it augmentsthe percentage of flowers which produce immature pods, subsequentlycauses fewer pods to reach maturity. Shading of the lower nodesreduces at these nodes the number of mature pods but may resultin more pods maturing at the upper nodes. It is concluded that when the light gradient is such as to restrictthe internal supplies of substrates the growth of those organswith the least competitive ability, e.g. the newly formed pods,is arrested. It is at this phase that the factors controllingabscission come into play and that abscission is dependent upona balance between the levels of auxins and the production ofan abscission factor.     

A model for studying pollination and pod development in Brassica napus: the culture of isolated flowers

Summary A technique for cultivating isolated flowers of Brassica napus has been developed. Flowers were harvested at anthesis, the surface of their peduncles was then sterilized and they were cultivated in a hormonefree medium. We used an MS medium supplemented with 3% sucrose as a source of organic carbon. From our experiments, it was concluded that no exogenous growth regulator is required to ensure normal growth and development in vitro. The flowers, and thereafter the pods, can be kept in culture until seed maturity. After 30 days, seed development resulted in three types of seeds: (1) normal, (2) milky and (3) aborted. The results show that the number of seeds per pod was not dependent on the order of flowers on the raceme (except the first 10 flowers and flowers above row 50). Our study supports the validity of this model as an easy tool for studying pollination and early seed development.     

Morphological development of normal and phyllody expressing Rosa hybrida cv. Motrea flowers

A large number of soybean (Glycine max L. Merr.)flowers and young pods abscise rather than develop into mature pods. Flower andpod drop or abortion accounts for the majority of total reproductive abscissionand influences potential soybean yield. The objectives of this study were todetermine the patterns of flower, pod and seed development under treatmentswiththe growth regulators, 2-(2,4-dichlorophenoxy) propanoic acid (2,4-DP) and6-benzylaminopurine (BAP), applied at the early reproductive stages, and toexamine the association of reproductive abscission with growth characteristicsand agronomic traits, including seed yield and seed weight. Small seeded [cvPungsan (11.1±0.4 g100-seed^–1)] and large seeded [cv Manlee(21.0±0.5 g 100-seed^–1)]genotypes were separately planted in the greenhouse and field, and treated witheither 2,4-DP or BAP. 2,4-DP (a synthetic auxin) and BAP (a syntheticcytokinin)were each applied at three concentrations (i.e. high, intermediate or low):0.12mM, 0.08 mM, 0.04 mM, and 1.5mM, 1 mM, 0.5 mM respectively. High andlow concentrations were employed for greenhouse experiments to examine thenumber of flowers per plant in pots. With the exception of low BAP (0.5mM) treatment in Pungsan, all treatments increased total podnumberwith various numbers of seeds per pod. Low 2,4-DP (0.04 mM) inbothgenotypes or BAP (0.5 mM) in Manlee significantly reduced flowerabortion and delayed abscission of pods in both genotypes, resulting inincreased pod setting rates. Under field conditions using intermediateconcentrations, 1 mM BAP significantly increased 100-seed weightto22.3 g at R1 in Manlee and 11.9 g at R3 in Pungsan.BAP (1 mM) at R3 in Pungsan significantly improved seed yield(40.1g plant^–1). Maturity was not significantlyaffected by either application in Manlee, but was significantly affected by BAPin Pungsan. In Pungsan, 2,4-DP increased pod number, plant height and nodenumber, but decreased 100-seed weight in Pungsan treated at R1, causing nosignificant change of seed yield. This study suggested that exogenousregulatorssignificantly influenced reproductive and growth characteristics, andconsequently seed yield, but increase of pod number was not always beneficialfor seed yield.     

Rapid Changes in Translocation Patterns in Soybeans following Source-Sink Alterations

The effects of source-sink alterations on the translocation patterns to soybean (“Fiskeby V”) pods were studied using a pod leakage technique. The distribution of assimilates from a source leaf using double pulse experiments was followed at the pods at the source node and the node immediately below. Alterations were made by shading, illuminating, or excising two-thirds of the leaf area of the leaf at the node below. In control experiments both pulses exhibited identical time-course patterns at both nodes. Shading the lower leaf during the first half of the experiment and illuminating during the second reduced the distribution of ¹⁴C-assimilate to the lower node's pods from the source leaf by approximately 30 to 50% while having no effect at the source node. Illuminating the lower leaf during the first half of the experiment followed by excision of two-thirds of that leaf's area and shading increased the import from the source leaf by 4- to 33-fold relative to the control while reducing the distribution to the source node by up to 40%. The change in distribution pattern took place in less than 30 minutes with no apparent change in the source leaf net photosynthesis or in the rate of movement to the pods. The results indicate that any alterations in the source-sink balance will quickly produce a change in the distribution patterns to the pods.     

The Effect of Fruit Shading on Yield in Pisum sativum L.

Fruits of Pisum sativum L. cv. Feltham First which initiatesonly one flower per flowering node, were selectively shadedunder varying levels of defoliation. The purpose of the experimentswas to ascertain whether the foliage could compensate for lossof the fruit's contribution to its own growth. There was evidenceof this, but fruit and seed weight per fruit and per plant werereduced by fruit shading at all levels of defoliation. The lossin yield due to shading suggested that the contribution fromthe fruit was at least 12 per cent. The number of seeds whichdeveloped to maturity was the yield component most affectedby treatment. There was no evidence to suggest that shadinghad a different quantitative effect on final weight at differentnodes, but it did increase flower abscission at the first foweringnode in an experiment done at low radiant exposure. In an experimentat higher radiant exposure, very few flowers abscised at theearlier nodes, but leaflet removal reduced final fruit yieldat the first flowering node to a greater degree than at thesecond. These differential responses could contribute to variabilityof seed size in a crop of vining peas. Pisum sativum, pea, fruit, pod, light, shading, photosynthesis, yield     

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     

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     

Carbon Transfer and Partitioning between Vegetative and Reproductive Organs in Pisum sativum L

Assimilate partitioning was studied in the common pea (Pisum sativum L.) by feeding ¹⁴CO₂ to whole plants and measuring radioactivity in different organs 48 hours after labeling. Two experimental protocols were used. For the first, one reproductive node was darkened with an aluminum foil, to prevent photosynthesis during labeling. The aim was to study assimilate translocation among nodes. The second was carried out to assess any priority among sinks. Whole plants were shaded, during labeling, to reduce carbon assimilation. Various developmental stages between the onset of flowering and the final stage in seed abortion of the last pod were chosen for labeling. When all photosynthetic structures at the first reproductive node were darkened at any stage of development after the formation of the first flower, the first pod was supplied with assimilates from other nodes. In contrast, later developed pods, when photosynthetic structures at their node were darkened, received assimilates from other nodes only when they were beyond their final stage in seed abortion. Reducing illumination to 30% did not change distribution of assimilated carbon between vegetative and reproductive structures, nor among pods. It appears that the relative proportion of ¹⁴C allocated to any one pod, compared to other pods, depends on the dry weight of that pod as a proportion of the total reproductive dry weight. When the plant was growing actively, following the start of the reproductive phase until a few days before the end of flowering, the top of the plant (i.e., all the organs above the last opened flower) had a higher sink strength and a higher relative specific activity than pods, suggesting that it was a more competitive sink for assimilates. The pattern of assimilate distribution described here provides an explanation for pod and seed abortion.     

大豆花荚败育期间的植物激素变化

大豆开花结荚期,不同发育阶段的幼蕾与花荚的脱落率不同,其中以花后5d内的幼荚脱落最严重。与败育花荚相比,正常花荚中的干物质积累量均较高。细胞分裂素(DHZRs,ZRs,iPA)含量也较高,花后3～5d的幼荚中表现更明显。脱落酸(ABA)则是以败育幼蕾及花后3～5d的幼荚中含量较高。不同发育阶段的大豆生殖器官中,正常开放花中的玉米赤霉烯酮(ZEN)含量最高。     

不同时期叶喷植物生长调节剂对大豆花荚脱落率及多聚半乳糖醛酸酶活性的影响

在大田条件下,以大豆品种‘合丰50＇为材料,比较研究了在V3（第3节龄期,三叶期）、R1（初花期）和R3（始荚期）期叶面喷施DTA-6、S_（3307）和TIBA三种植物生长调节剂对大豆花荚脱落率及多聚半乳糖醛酸酶活性的影响。结果表明：V3期叶喷TIBA、S_（3307）、DTA降低了大豆花荚脱落率,降低了大豆花荚和脱落花荚多聚半乳糖醛酸酶活性;R1期叶喷植物生长调节剂显著降低了大豆花荚及脱落花荚中多聚半乳糖醛酸酶活性,以DTA-6调控效果最佳,S_（3307）次之;R3期叶喷植物生长调节剂降低了大豆荚及落荚的多聚半乳糖醛酸酶活性,以DTA-6调控效果最佳,TIBA次之。综合分析表明,V3、R1和R3期叶面喷施植物生长调节剂能够降低大豆花荚脱落率及多聚半乳糖醛酸酶活性,对大豆花荚的脱落有一定的调控作用,有利于提高产量,其综合调控效果为,V3期：S_（3307）〉DTA-6〉TIBA〉CK;R1期：DTA-6〉TIBA〉S_（3307）〉CK;R3期：DTA-6〉TIBA〉S_（3307）〉CK。     

大豆花荚败育期间的植物激素变化

大豆开花结荚期,不同发育阶段的幼蕾与花荚的脱落率不同,其中以花后５ｄ内的幼荚脱落最严重。与败育花荚相比,正常花荚中的干物质积累量均较高。细胞分裂素（ＤＨＺＲｓ,ＺＲｓ,ｉＰＡ）含量也较高,花后３～５ｄ的幼荚中表现更明显。脱落酸（ＡＢＡ）则是以败育幼蕾及花后３～５ｄ的幼荚中含量较高。不同发育阶段的大豆生殖器官中,正常开放花中的玉米赤霉烯酮（ＺＥＮ）含量最高     

Auxins,ABA and gibberellin-like activity in abscising and non-abscising flowers and pods of Vicia faba L.

Abscission probability varies among floral positions within inflorescences of Vicia faba L. Flowers from proximal positions have a greater chance to develop into mature pods than flowers from more distal positions which normally abscise either as older flowers or as young pods. In three field experiments with the indeterminate single stem variety Herz-Freya, changes in the contents of extractable auxins, abscisic acid (ABA) and gibberellins in flowers and pods during their development, and their possible influence on abscission were investigated.Inflorescences at different positions along the stem were divided into the two proximal and the remaining fruits. The content of all three hormones was at a low level during flower development, increased greatly in parallel with dry matter accumulation in the young pods, and then decreased to maturity. The first hormone to increase in the fruits was auxin and this took place when abscission from the distal positions began. ABA and gibberellins at this time were still at a low level. This ontogenic course of hormone production was very similar in fruits of both positions within an inflorescence, but in flowers and young pods from proximal positions, auxin content in most inflorescences was greater than in those from the abscising distal positions. No such positional differences were observed with ABA and gibberellins. Decapitation of the plants reduced flower and pod drop from the remaining reproductive nodes. Although decapitation resulted in less abscission among distal flowers and young pods from these nodes, it did not affect the ontogenic course of auxin and ABA production in these fruits.     

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.     

Yield components and cultivar, sowing date and density in field beans (Viciafaba)

Delay in sowing the cultivars Ostlers (Viciafaba var. equina) and Herz Freya (V.faba var. minor) beyond late February depressed yield. Densities of at least sixty plants per m² were required to ensure that yield was not limited by sowing rate. Differences in yield in response to changes in density were reflected particularly in changes in numbers of pod-bearing nodes per plant, but also to a lesser extent in numbers of pods per pod-bearing node. The inherently larger seed weight of cv. Ostlers compared with cv. Herz Freya did not confer any yield advantage, differences in number of beans per pod and number of pods per pod-bearing node tending to equalize the yield of the two cultivars. The most stable component of yield throughout was number of beans per pod, and the least stable was number of pod-bearing nodes per plant.