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.     

Probing Monocarpic Senescence and Pod Development Through Manipulation of Cytokinin and Mineral Supplies in Soybean Explants

Defined solutions containing cytokinin and/or mineral nutrientswere supplied in lieu of the roots through the cut stem baseof soybean explants (a leaf with associated pod and subtendingstem segment) in order to analyze the roles of cytokinin andmineral nutrients from the roots in pod development and foliarmaintenance. In explants cut at early-mid podfill, supplyingonly H₂O accelerated leaf senescence and pod maturation anddecreased seed d. wt relative to comparable parts of intactplants. Zeatin (Z) and/or minerals not only delayed leaf yellowingand the decline in foliar chlorophyll levels and photosyntheticrates but also inhibited leaflet and petiole abscission relativeto H₂O controls. Even large declines in foliar assimilatoryprocesses did not necessarily lead to abscission. Z and/or mineralsalso increased stomatal conductivity throughout podfill. Z showedsome positive synergistic effects with minerals on leaf maintenance.Pod wall, cotyledon and radicle yellowing were delayed by Zand/or minerals but not as much as leaf senescence. Mineralsonly or Z +minerals prolonged seed d. wt accumulation and increasedfinal dry seed wt to a level similar to that for intact plants.Seed growth showed a complex interrelation with pod wall andleaf f. wt and d. wt changes. A decline in cytokinin and mineralflux from the roots appears to be important for pod-inducedleaf senescence; however, pod development, foliar senescenceand their component processes may be affected differently. Thus,even though the explant is a physiological/nutritional moduleof the whole plant, it is influenced by cytokinin and mineralsfrom the roots and therefore only semiautonomous. Glycine max L. Merr. cv. Anoka, soybean, abscission, cytokinin, chlorophyll, mineral nutrients, seed development, semiautonomous physiological modules, senescence, stomatal resistance     

Retardation of soybean leaf senescence and associated effects on seed composition

Soybean leaf senescence, leaf abscission, and pod yellowing were markedly delayed by sprays of 10^–4 M 6-benzylamino-9-(tetrahydropyran-2-yl)purine plus 5×10^–5 M -naphthalene acetic acid. The pods on the sprayed plants turned yellow 5–7 days later than those on the control plants, and the treated leaves remained dark green even when the pods had already desiccated. The antisenescence spray did not change pod numbers, seed numbers, seed size, or the yield. By retarding senescence, seed nitrogen content was increased in both TCA-soluble and TCA-insoluble fractions. Seed total protein, buffer-extractable total protein, and globulin were increased by 26, 28, and 33 mg/g of seed flour, respectively, and albumin was decreased by 6 mg/g. The overall increase in seed protein caused by spray treatment is confined to the globulin portion.     

Postflowering photoperiod regulates vegetative growth and reproductive development of soybean

Soybean development is controlled by environmental factors, primarily photoperiod and temperature. To date, photoperiod effects on flowering have been well studied but the performances and mechanism of postflowering photoperiod responses have not been fully understood, especially for the photoperiod effects on vegetative growth after flowering. In the present study, the responses of vegetative growth and reproductive development in soybean to different postflowering photoperiod regimes were investigated in four separate experiments. Three varieties of different maturity groups (MG) including the early (Dongnong 36, MG 000), medium (Dandou 5, MG IV), and late (Zigongdongdou, MG IX) were exposed to two photoperiods, short (10, 12 h) and long (15, 16 or 18 h). The results showed that postflowering photoperiod not only regulated reproductive development but also affected vegetative growth. Even when flowers and pods were removed, short-day (SD) treatment promoted leaf senescence. The onset of leaf senescence among varieties tested appeared to be dependent on photoperiod sensitivity. Leaf senescence of the late-maturing variety of Zigongdongdou (sensitive to photoperiod) was delayed more significantly than that of the medium and early-maturing varieties (less sensitive to photoperiod). Long-day (LD) treatments delayed leaf senescence and seed maturation in the late-maturing variety of Zigongdongdou plants with only the SD-induced leaves produced before flowering. LD treatments imposed from the beginning bloom, beginning pod setting or beginning seed filling delayed leaf senescence and seed maturation of late-maturing soybean variety (Zigongdongdou). Results of night-break with red (R) and far-red (FR) light demonstrated that postflowering photoperiod responses of soybean were R/FR reversible reactions and the phytochromes seemed to be functional as receptors of photoperiod signals even after flowering. It was proposed that the regulation of photoperiod on development of soybean was effective from emergence through maturation, and the postflowering photoperiod signals were also mediated by phytochromes similar to those before flowering. The flowering reversion in late-MG soybean varieties under LD was a direct result of LD and was not due to secondary effect of abscission of pods and flowers. Soybean leaves not only received SD signals but also LD signals; furthermore, the LD effects reversed the SD effects and vice versa.     

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.     

EFFECT OF PETIOLE PHLOEM DISRUPTION ON STARCH AND MINERAL DISTRIBUTION IN SENESCING SOYBEAN LEAVES

Normally, starch (sugars) and minerals are redistributed from the leaves to the pods during monocarpic senescence in maturing soybean plants. Petiole phloem destruction (steam girdling), which blocked this redistribution by interrupting export through the petiole, altered the foliar senescence pattern producing a distinctive interveinal yellowing with green areas along the veins on pod-bearing plants. This suggests that blockage of the petiole phloem may cause nutrients to accumulate in the green zones along the leaf veins instead of being redistributed to the pods. In the leaves of untreated plants, starch showed the same distribution pattern as chlorophyll; however, starch was preserved in yellow areas as well as the green zones of the steam-girdled leaves. Mineral analyses of the veinal and interveinal zones of treated leaves and controls showed that the veinal green zones and interveinal yellowing in treated plants were not respectively enriched and depleted in minerals corresponding to a redistribution of minerals within the leaves. Depodding also blocked leaf yellowing, net mineral redistribution and starch breakdown. Thus, the pods are able to induce chlorophyll breakdown without net mineral redistribution or starch loss in leaves with petiole phloem destruction. This shows that chlorophyll breakdown is not obligatorily coupled with mineral redistribution or starch breakdown.     

Seed Development in Phaseolus vulgaris L. cv Seminole: I. Developmental Independence of Seed Maturation

Phaseolus vulgaris L. cv Seminole pods removed from the plant continued their development when incubated in suitable conditions. Seeds continued to grow and develop and pods and seeds passed through an apparently normal developmental sequence to dryness. Seed growth was at the expense of pod dry weight (DW) reserves. Losses of pod DW paralleled DW gains by seeds in detached pods and in pod cylinders containing a seed. The transfer activity was apparent only within the period 10 to 30 days after anthesis (DAA) with maximal activity between 15 to 20 DAA. This period corresponds to maximum pod growth and the attainment of maximal DW. Seeds are in only the early phase of seed growth at this time. No DW transfer was observed at developmental stages beyond 30 to 35 DAA when normal senescence DW losses in pods became evident and seeds were in the later phase of seed fill. Pods or pod cylinders remained green and succulent over the transfer period, later passing through yellowing and drying phases characteristic of normal development. DW transfer was dependent on funicle integrity and was readily detectable in pod cylinders after 7 days incubation. The DW transfer activity may contribute to continuing nutrition of seeds under conditions where the normal assimilate supply to seeds becomes limiting. Defoliation and water stress treatments applied to Phaseolus plants reduced seed yields but allowed persistence of seed maturation processes such that all seeds developing to dryness were capable of germination.     

Modification of Seed Growth in Soybean by Physical Restraint: Effect on Leaf Senescence

Crafts-Brandner, S. J. and Egli, D. B. 1987. Modification ofseed growth in soybean by physical restraint. Effect on leafsenescence.—J. exp. Bot. 38: 2043–2049. The effect of total plant sink size on leaf senescence in soybean[Glycine max (L.) Merrill] was investigated by using a simple,non-destructive method to decrease seed growth rate and totalplant fruit sink size without altering fruit or seed number.The treatment consisted of placing plastic pod restriction devices(PPRD), which were made from plastic drinking straws (6·35mm diameter), over the fruits so that all of the seeds werecontained within the PPRD's. The treatment did not alter thetime of initiation of leaf senescence for two cultivars (McCalland Maple Amber), but decreased the rate of leaf senescencebased on declines in chlorophyll, ribulose-l,5-hi'sphosphatecarboxylase/oxygenase level and carbon dioxide exchange rate.The treatment also delayed seed maturation. At the time of seedmaturation, the plants still retained green leaves. In a separate experiment, one seed in each fruit (40% of theseeds on the plant) was not restrained by the PPRD's. This treatmentled to an intermediate rate of leaf senescence compared to controland complete seed restriction treatments. The results indicatedthat, for the cultivars examined (1) leaf senescence was initiatedat the same time regardless of sink size (2) the rate of leafsenescence could be modified by altering sink size and (3) seedmaturation could occur without complete leaf yellowing and leafabscission. The effect of the PPRD treatments on leaf senescencewere similar to results obtained when fruits were physicallyremoved, which indicated that physical removal of fruits doesnot lead to artefacts due to wounding of the plants. Key words: Glycine max L, senescence, source-sink     

Effects of decreasing source/sink ratio in soybeans on photosynthesis, photorespiration, transpiration and yield

Abstract. Shading of all side leaflets of a determinate soybean cultivar during pod filling significantly increased rates of photosynthesis in the unshaded centre leaflets, compared to centre leaflets of controls. Higher rates were associated with both higher stomatal and mesophyll conductances, and were reversible within 2 days when shades were removed. These higher rates of photosynthesis were not associated with decreased percentage enhancement by low oxygen, indicating that treatment effects were probably not associated with changes in photorespiration relative to photosynthesis. Percentage enhancement did, however increase as the plants approached physiological maturity, chiefly because of a decrease in photosynthesis.
In spite of these increases in rates of photosynthesis seed weight per plant was decreased by 37% in plants with side leaflets shaded for the entire pod-filling period and by 28% in plants shaded for only the second half of the period. In plants where shades were removed during the second half of pod filling seed yield was reduced by only 19% because shade removal delayed leaf senescence. The four treatments reduced yield by different mechanisms. Plants shaded continuously during pod filling produced fewer seeds than controls, but the weight per seed was similar. When shading was applied during the second half of pod fillings seed number was unchanged but weight per seed was significantly reduced. In contrast when shades were removed for the second half of pod filling, seed number remained similar to that of continuously shaded plants, but seed weight increased.
Although all shading treatments reduced yield, the reduction was not proportional to the 63% reduction in leaf area available for photosynthesis. This was because (1) photosynthetic rates in the centre leaflet of shaded plants were higher than rates in controls, (2) stem and lower surface photosynthesis in shaded leaf-lets contributed to whole leaflet photosynthesis.     

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     

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.

     

Effects of morphactin and other auxin transport inhibitors on soybean senescence and pod development

Because triiodobenzoic acid increases pod number, albeit variably, in soybean (Glycine max), we tested other auxin-transport inhibitors. Morphactins, especially methylchlorflurenol (MCF), were found to be very active (optimal concentration 10 micromolar) when sprayed onto the foliage. Applications at 1 week after the start of flowering were most effective, producing a 40% increase in pod number with little inhibition (12%) of stem elongation. MCF increased the number of pods initiated (reaching 1 cm length) at least partially by prolonging the initiation period, while pod abortion (failure of pods > 1 cm long) remained low. Generally, MCF did not increase seed yield (dry weight/plant); more, but smaller seeds, were formed by the treated plants. The promotive effect of MCF on pod initiation seems to be independent of its inhibition of stem elongation, which is insignificant at 10 micromolar. MCF delayed pod maturation by 3 to 4 days, while foliar yellowing, blade abscission, and petiole abscission were retarded by 2, 4, and 2 days, respectively. MCF has only a small effect on senescence and that could be indirect, due to a delay in pod development. Other auxin-transport inhibitors tested, including N-1-naphthylphthalamic acid, produced little or no increase in pod number; however, 0.1 millimolar 5-[2′-carboxyphenyl]-3-phenylpyrazole caused a 27% increase. These results implicate auxin as a potential regulator of pod development, and they show that soybean seed yield is not simply sink limited.     

Effect of spikelet removal on the whole plant senescence of rice

The rice (Oryza sativa L.) cultivar IET 1444 showed a nonsequential mode of senescence as evident from the decline in chlorophyll and protein of the flag and second leaves at the senescent stage. Removal of 50,75 and 100 % spikelets from the panicle of rice plant or emasculation of the panicle by hot water treatment induced the development of secondary branch from the axil of second leaf but 25 % removal had no effect. Similarly, removal of 75 and 100 per cent spikelets from the panicle of secondary branch induced tertiary branch development, while 25 and 50 per cent removal had no such effect. Similar treatments on tertiary branch had no effect on further branch production. The pattern of leaf senescence of the untreated (control) main tiller, secondary and tertiary branches was identical, i. e. nonsequential, which could be changed into the sequential type only by the development of additional sinks (i. e. side branch). The leaf area and the seed number of secondary and tertiary branches were gradually reduced and reached a critical value in the tertiary branch. The removal of spikelets or emasculation delayed leaf senescence of the main tiller and the secondary and tertiary branches. Also the longevity of the whole plant could be increased by 40 d i. e., up to the senescence of the tertiary branch. Both leaves and reproductive parts control side branch production, which, in turn, controls the longevity of the whole rice plant.     

Gas exchange by pods and subtending leaves and internal recycling of CO(2) by pods of chickpea (Cicer arietinum L.) subjected to water deficits

Terminal drought markedly reduces leaf photosynthesis of chickpea (Cicer arietinum L.) during seed filling. A study was initiated to determine whether photosynthesis and internal recycling of CO(2) by the pods can compensate for the low rate of photosynthesis in leaves under water deficits. The influence of water deficits on the rates of photosynthesis and transpiration of pods and subtending leaves in chickpea (cv. Sona) was investigated in two naturally-lit, temperature-controlled glasshouses. At values of photosynthetically active radiation (PAR) of 900 micromol m(-2) s(-1) and higher, the rate of net photosynthesis of subtending leaves of 10-d-old pods was 24 and 6 micromol m(-2) s(-1) in the well-watered (WW) and water-stressed (WS) plants when the covered-leaf water potential (Psi) was -0.6 and -1.4 MPa, respectively. Leaf photosynthesis further decreased to 4.5 and 0.5 micromol m(-2) s(-1) as Psi decreased to -2.3 and -3.3 MPa, respectively. At 900--1500 micromol m(-2) s(-1) PAR, the net photosynthetic rate of 10-d-old pods was 0.9-1.0 micromol m(-2) s(-1) in the WW plants and was -0.1 to -0.8 micromol m(-2) s(-1) in the WS plants. The photosynthetic rates of both pods and subtending leaves decreased with age, but the rate of transpiration of the pods increased with age. The rates of respiration and net photosynthesis inside the pods were estimated by measuring the changes in the internal concentration of CO(2) of covered and uncovered pods during the day. Both the WW and WS pods had similar values of internal net photosynthesis, but the WS pods showed significantly higher rates of respiration suggesting that the WS pods had higher gross photosynthetic rates than the WW pods, particularly in the late afternoon. When (13)CO(2) was injected into the gas space inside the pod, nearly 80% of the labelled carbon 24 h after injection was observed in the pod wall in both the WW and WS plants. After 144 h the proportion of (13)C in the seed had increased from 19% to 32% in both treatments. The results suggest that internal recycling of CO(2) inside the pod may assist in maintaining seed filling in water-stressed chickpea.     

The effects of flower removal on the seed yield of pigeonpeas (Cajanus cajan)

In field experiments carried out at Hyderabad, India with early and mediumduration cultivars of Cajanus cajan sown at the normal time, in July, removal of all flowers and young pods for up to 5 wk had little or no effect on final yield. The flowering period of the deflowered plants was extended and their senescence delayed. The plants compensated for the loss of earlier-formed flowers by setting pods from later-formed flowers; there was relatively little effect of the deflowering treatments on the number of seeds per pod or weight per seed. The plants were also able to compensate for the repeated removal of all flowers and young pods from alternate nodes by setting more pods at the other nodes.
The removal of flowers from pigeonpeas grown as a winter crop resulted in yield reductions roughly proportional to the length of the deflowering period, probably because maturation of these plants was delayed and occurred under increasingly unfavourable conditions as the weather became hotter.     

Varietal differences in mungbean (Vigna radiata) for growth, yield, toxicity symptoms and cadmium accumulation

Increased cadmium (Cd) contamination of soils resulting from industrial activities is critical to crop production. The objective of this study was to find varietal differences for foliar chlorosis and necrosis, growth and Cd accumulation in mungbean (Vigna radiata). Despite substantial varietal differences, increased Cd levels reduced the shoot and root dry weight and the number and area of leaves at different growth stages. Applied Cd stress produced the foliar symptoms such as marginal and intervein chlorosis and scattered necrotic spots on younger leaves while accelerating the senescence of older leaves. Slope of regression equation and correlations of shoot Cd content with foliar Cd toxicity revealed that leaf chlorosis was more damaging than necrosis. At maturity, number of pods per plant and seeds per pod were maximally reduced to 37% and 26%, while 100‐seed weight, seed yield and harvest index showed 61%, 79% and 54% reduction, respectively, as a result of Cd toxicity. Results suggested that although varietal difference exists, the accumulated Cd is mainly toxic to the mesophyll tissue, most probably by interfering with the uptake of essential nutrients, thereby reducing growth and yield at various stages. Therefore, selection programmes based on foliar toxicity criteria may be beneficial for better utilisation of Cd‐polluted soils.     

Physiological characterization of 'stay green' mutants in durum wheat

Four mutants with delayed leaf senescence were selected from seed of durum wheat mutagenized with ethylmethane sulphonate. Changes in net photosynthetic rate, efficiency of photosystem II and chlorophyll concentration during the maturation and senescence of the flag leaves of both mutant and parental plants were determined under glasshouse conditions. The four mutant lines maintained photosynthetic competence for longer than the parental line and are therefore functionally 'stay green'. The mutant lines also had higher seed weights and grain yields per plant than the parental line.     

Influence of the triazole growth retardant BAS 111..W on phytohormone levels in senescing intact pods of oilseed rape

Foliar treatment of oilseed rape plants (Brassica napus L.ssp. napus cv. Linetta) with the growth retardant BAS 111..W at the 5th leaf stage delayed pod senescence during early maturation. Changes of immunoreactive cytokinin- and abscisic acid (ABA)- like substances and of the ethylene precursor 1-aminocyclo-propane-1-carboxylic acid (ACC) and its malonyl-conjugate (MACC) were determined in intact whole pods. When compared with control plants, higher levels of total chlorophyll correlated with four-fold and three-fold increases of trans-zeatin riboside- and dihydrozeatin riboside-type cytokinins, respectively, in the pods of plants treated with 0.25 mg BAS 111..W per plant. Isopentenyladenosine-type cytokinins and ACC and MACC contents remained virtually unchanged, whereas ABA levels dropped considerably below those of controls (60% reduction). However, when analysed at late pod maturity, BAS 111..W treatment no longer affected the total chlorophyll content, or the levels of cytokinins, ABA, ACC and MACC. We hypothesize that the retardant-induced changes in the hormonal status of the pods, favouring the senescence-delaying cytokinins as opposed to abscisic acid, could contribute to the developmental delay.     

花生衰老进程的研究

通过对鲁花11号和辐8707 2个高产花生品种的衰老进程研究表明：花生衰老具有地上部（叶片）渐进衰老和整株衰老的双重特点。花生从始花至花后60d左右为地上部（叶片）渐进衰老期：此期主茎高、侧枝长、分枝数、主茎、侧枝绿叶数、叶面积、茎、叶干重迅速增加,并接近或达到最大值,主茎及侧枝基部叶片逐渐由下向上开始衰老死亡,饱果开始出现,根系活力、固氮酶活性逐渐升高至接近最大值,始花后60－90d为整株缓衰期,此期地上部茎叶生长基因停止,逐渐开始衰老死亡,主茎、侧枝绿叶数开始减少,生殖体（荚果）干重迅速增长,根系活力、固氮酶活性缓慢降低;始花后90d以后称为整株速衰期,此期主茎、侧枝绿叶迅速减少,地上部迅速衰老死亡,生殖体（荚果）干重缓慢增长,根系活力、固氮酶活性迅速降低。地上部（叶片）渐进衰老期与开花及大量荚果形成相对应,整株缓衰期伴随着荚果迅速增重,整株速衰期与荚果缓慢增重一致。     

Gibberellic Acid Treatment Reduces the Tolerance of Field-Grown Common Bean to Leaf Removal

I studied the influence of gibberellic acid (GA₃) treatment in a field population of common bean on plant tolerance to leaf removal. Individual bean seedlings were treated with a foliar application of 10 μM GA₃ on day 7 and day 14 after emergence, which led to a significant increase in height in GA₃-treated plants. Twenty-eight days after emergence, either zero, one, two, or three leaflets from each trifoliate leaf were removed from each of 20 GA₃-treated and 20 control plants. All pods were harvested from each plant after plants became senescent 6 weeks later. Multivariate analyses revealed that leaf removal produced significant reductions in several yield components in both GA₃-treated and control plants, although the effects were not pronounced until at least two leaflets from each trifoliate leaf (67% of the total leaf area) were removed. However, GA₃-treated plants suffered greater reductions in total pod wall mass and total seed number than control plants after 33 and 67% leaf area removal. These results indicate that GA₃ treatment may have altered the assimilatory capacity or resource allocation pattern of treated plants in such a way as to decrease their ability to tolerate leaf removal, a negative consequence of the hormonal alteration of traits important to plant compensation for biotic stressors. Received December 6, 1996; accepted March 5, 1997