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 Since nutrients would have to be withdrawn from the leaves via the phloem, the pods do not induce yellowing by pulling nutrients out of the leaf and must be able to exert their influence via the xylem.     

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.     

Regulation of monocarpic senescence of Brassica campestris by the developing pods

Senescence of Brassica campestris L. cv. B-9 was studied with regard to seed maturation and source-sink relationships. In normal control plants leaf senescence (as determined by the change in chlorophyll level) started and proceeded in a progressive manner from base to apex during the period of early pod setting. Complete yellowing of the leaves occurred well before the seed maturation and pod wall senescence. The pod wall always senesced before the attainment of final seed weight. In two different sets of acrocarpous plants containing 65 pods and 10 pods, respectively, leaf senescence was delayed during the pod filling period. It started non-sequentially after complete yellowing and senescence of the pod wall. The degree of leaf senescence at the post-pod filling stage was almost proportional to the number of pods present. When peduncles of the acrocarpous 10-podded plants were removed after the pod filling stage of the plant, leaf senescence was delayed compared to plants whose pedicels were removed, although the senescence pattern of the upper three leaves was nonsequential in both cases. Defruiting at an early stage of development delayed leaf senescence, although the pattern of such senescence remained unaltered (i.e. nonsequential). Defoliation hastened the seed-filling process and pod wall senescence. Plants containing fewer pods had higher average seed weight, although yield per plant was reduced.
These results suggest that the pod wall serves as a temporary as well as intermediary storage organ and that foliar senescence is not directly related to seed maturation. The possible cause of uncoupling between foliar senescence and seed development is discussed.     

Sequence of morphological and physiological events during natural ageing and senescence of a castor bean leaf: sieve tube occlusion and carbohydrate back-up precede chlorophyll degradation

The development of castor bean ( Ricinus communis L. var. sanguineus) leaves from bud break to abscission was studied to determine whether senescence of phloem precedes or follows chlorophyll degradation in the course of natural ageing of leaves. The castor bean leaf blade took 20 days for full expansion and its average life span was 60 days. From the day of full expansion on it suffered a substantial loss in N, a small loss in C, K and P and a gain in Ca, Mg and S. The content of soluble sugars increased with time, paralleled by a decrease of photosynthetic activity. Starch accumulated shortly before chlorophyll breakdown. The amino acid level in the leaves decreased steadily together with nitrate reductase and glutamine synthetase activity. Reactive oxygen species increased and oxidation-protecting compounds decreased during the life span of the leaves. Shortly after full leaf expansion an increasing number of sieve plates showed strong callose depositions when visualized by aniline blue method. At day 40 only half of the sieve tubes appeared functional. Chlorophyll breakdown followed these processes with a time lag of approximately 10 days. The sieve tube sap of ageing leaves had the same sucrose concentrations as young leaves, whereas amino acid concentrations decreased. High levels of reduced ascorbic acid and glutathione together with increasing levels of glutaredoxin indicated oxidative strain during senescence. We speculate that the gradual increase of reactive oxygen species during ageing together with the import of calcium ions lead to the stimulation of callose synthesis in plasmodesmata and sieve plates with the consequence of inhibition of phloem transport leading to carbohydrate back-up in the leaf blade. The latter may finally induce chlorophyll breakdown and, at the end, leaf abscission at the petiole base. Thus phloem blockage would precede and may be causal for chlorophyll degradation in leaf senescence.     

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

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

Phloem unloading in tobacco sink leaves: insensitivity to anoxia indicates a symplastic pathway

Phloem unloading in transition sink leaves of tobacco (Nicotiana tabacum L.) was analyzed by quantitative autoradiography. Detectable levels of labeled photoassimilates entered sink leaves approx. 1 h after source leaves were provided with ¹⁴CO₂. Samples of tissue were removed from sink leaves when label was first detected and further samples were taken at the end of an experimental phloem-unloading period. The amount of label in veins and in surrounding cells was determined by microdensitometry of autoradiographs using a microspectrophotometer. Photoassimilate unloaded from first-, second-and third-order veins but not from smaller veins. Import termination in individual veins was gradual. Import by the sink leaf was completely inhibited by exposing the sink leaf to anaerobic conditions, by placing the entire plant in the cold, or by steam-girdling the sink-leaf petiole. Phloem unloading was completely inhibited by cold; however, phloem unloading continued when the sink-leaf petiole was steam girdled or when the sink leaf was exposed to a N₂ atmosphere. Compartmental efflux-analysis indicated that only a small percentage of labeled nutrients was present in the free space after unloading from sink-leaf veins in a N₂ atmosphere. The results are consistent with passive symplastic transfer of photoassimilates from phloem to surrounding cells.Symbol VI radio of ¹⁴C in veins and interveinal tissue     

Mobilization of Minerals to Developing Seeds of Legumes

The mineral nutrition of fruiting plants of Pisum sativum L.,Lupinus albus L. and Lupinus angustifolius L. is examined insand cultures supplying adequate and balanced amounts of essentialnutrients. Changes in content of specific minerals in leaves,pods, seed coat, and embryo are described. P, N and Zn tendto increase precociously in an organ relative to dry matteraccumulation, other elements more or less parallel with (K,Mn, Cu, Mg and Fe) or significantly behind (Ca and Na) dry weightincrease. Some 60–90 per cent of the N, P and K is lostfrom the leaf, pod and seed coat during senescence, versus 20–60per cent of the Mg, Zn, Mn, Fe and Cu and less than 20 per centof the Na and Ca. Mobilization returns from pods are estimatedto provide 4–39 per cent of the seeds' accumulations ofspecific minerals, compared with 4–27 per cent for testatransfer to the embryo. Endosperm minerals are of only minorsignificance in embryo nutrition. Comparisons of the mineral balance of plant parts of Lupinusspp. with that of stem xylem sap and fruit tip phloem sap supportthe view that leaves and pod are principal recipients of xylem-borneminerals and that export from these organs via phloem is themajor source of minerals to the seeds. Endosperm and embryodiffer substantially in mineral compostition from phloem sap,suggesting that selective uptake occurs from the translocationstream during seed development. Considerable differences are observed between species in mineralcomposition of plant organs and in the effectiveness of transferof specific minerals to the seeds Differences between speciesrelate principally to Ca, Na and certain trace elements.     

MINERAL DISTRIBUTION IN RELATION TO FRUIT DEVELOPMENT AND MONOCARPIC SENESCENCE IN ANOKA SOYBEANS

The changes in distribution of several important mineral nutrients (N, K, Ca, Mg, Mn, and Fe) were studied in relation to monocarpic senescence (measured as leaf yellowing) and fruit development in hydroponically-grown (and to a lesser extent field-grown) Anoka soybeans with particular emphasis on the leaves and seeds. Only N shows a clear redistribution from the leaves to the seeds as the seeds grow, and this transfer starts before visible leaf yellowing. K, Ca, Mn and Fe do not seem to redistribute, but Mg may undergo limited redistribution. Depodding prevents the drop in the amounts of minerals in attached leaves by blocking leaf shedding and/or redistribution and also creates some quantitative changes in mineral distribution. On a g fresh weight basis, only the N content of leaf blades decreases during yellowing; the K, Mg, Ca, Mn and Fe contents do not decrease. Therefore, depletion of the latter minerals from the leaves cannot be responsible for their yellowing. Although N deficiency alone could cause foliar chlorosis, the monocarpic yellowing pattern is distinctly different from that induced by N deprivation.     

Role of microbial immigration in the colonization of apple leaves by Aureobasidium pullulans

The role of microbial immigration in the veinal colonization pattern of Aureobasidium pullulans on the adaxial surface of apple leaves was investigated in two experiments at two periods (early and late seasons) in 2004 by applying green fluorescent protein (GFP)-tagged blastospores to the foliage of orchard trees. Individual leaves were resampled by a semidestructive method immediately after inoculation (t(0)) and about 1 (t(1)), 2 (t(2)), and 3 (t(3)) weeks later. At t(0), there were no significant (P < or = 0.05) differences in densities (cells/mm(2)) on veinal (excluding midvein) sites and those on interveinal sites, but at all points thereafter, densities were significantly higher on veins. GFP-tagged A. pullulans cells remained primarily as singletons on interveinal regions (> or =90% at all points), while > or =20% of cells over veins at t(3) were in colonies of > or =4 cells. The colonies that developed from single cells placed on midveins and other veins were significantly larger than those that developed on interveinal regions of detached field and seedling leaves incubated under controlled conditions. Colonies primarily developed linearly along veins, reaching average colony sizes (72 h) of 24.4 +/- 12.7 (mean +/- standard deviation) cells. In contrast, colonies on interveinal regions tended to average only 2.9 +/- 1.3 cells, with less linearity. To examine the potential role of A. pullulans growth-inhibiting factors associated with interveinal features, single GFP-tagged A. pullulans cells in droplets previously incubated on interveinal sites were placed on midveins and compared to midvein colonies derived from cells in a water-only suspension. No differences in colony size resulted. Our results indicate that immigration limitation and growth-inhibiting factors are not the primary factors responsible for A. pullulans veinal colonization patterns in the field. Rather, indirect evidence suggests that growth-promoting substances occur locally in the veinal areas.     

Leaf Senescence and Abscisic Acid in Leaves of Field-grown Soybean

Leaf senescence in field-grown soybean (Merrill) as defined by the period after full expansion, was studied by measuring abscisic acid (ABA), total soluble protein, and chlorophyll in leaves through the later part of the growing season. ABA concentrations increased significantly at the end of the season when leaves had started to turn yellow, well after total soluble protein and chlorophyll had started to decline. The results indicate that events occurring before leaf yellowing are more significant in evaluating leaf senescence since the yellowing condition and rise in ABA are effects of changes in physiological activity beginning when leaves are still green.     

Short-term Effects of Heat-girdles on Source Leaves of Vicia faba: Analysis of Phloem Loading and Carbon Partitioning Parameters

Petiole heat-girdle treatments (followed by a 5 min ¹⁴CO₂ assimilation)were performed on mature leaves of Vicia faba, in order to assesstheir effect on the partitioning of photo-assimilates to theminor vein phloem. Whole leaf autoradiographic evidence indicateda high leaf-to-leaf variation in the image intensity over theminor veins (relative to the mesophyll/epidermal background)in both control and heat-girdled groups of leaves. The averagedegree of minor vein labelling in heat-girdled leaves, however,was found to be significantly lower than that in controls. Comparativeassessment of vein labelling was based on microscopic densityreadings of silver grains over veinal and interveinal regionsin autoradiographic images. Investigations into the cause ofthis alteration in vein labelling indicated no involvement ofan inhibition of apoplasmic phloem loading, as both heat-girdledand control leaves of Vicia were shown to have comparable minorvein uptake of exogenously supplied ¹⁴C-sucrose. Heat-girdlingwas shown, however, to increase significantly the partitioningof recently fixed carbon into the insoluble (mainly starch)fraction relative to the ethanol-soluble fraction, within 12min of the treatment. We suggest that this carbon partitioningchange can primarily account for the change in vein labelling,since an increase in the insoluble fraction would result in(1) more ¹⁴C-activity remaining in the leaf mesophyll and (2)less ¹⁴C-activity going into the mesophyll export pool, andthus, less ¹⁴C-sucrose being transferred to the minor vein region.Additionally, although leaf export was completely halted inheat-girdled leaves, ¹⁴C-activity was found within the majorveins as far as the point of petiole heat-girdling (followinga 5 min assimilation and 4 h chase). Apparently, continued (butlimited) solution flow within the sieve elements is maintainedby transport pathway unloading within the treated leaves. Key words: Phloem loading, carbon partitioning, heat-girdle, Vicia faba     

7-Hydroxymethyl chlorophyll a reductase functions in metabolic channeling of chlorophyll breakdown intermediates during leaf senescence

During natural or dark-induced senescence, chlorophyll degradation causes leaf yellowing. Recent evidence indicates that chlorophyll catabolic enzymes (CCEs) interact with the photosynthetic apparatus; for example, five CCEs (NYC1, NOL, PPH, PAO and RCCR) interact with LHCII. STAY-GREEN (SGR) and CCEs interact with one another in senescing chloroplasts; this interaction may allow metabolic channeling of potentially phototoxic chlorophyll breakdown intermediates. 7-Hydroxymethyl chlorophyll a reductase (HCAR) also acts as a CCE, but HCAR functions during leaf senescence remain unclear. Here we show that in Arabidopsis, HCAR-overexpressing plants exhibited accelerated leaf yellowing and, conversely, hcar mutants stayed green during dark-induced senescence. Moreover, HCAR interacted with LHCII in in vivo pull-down assays, and with SGR, NYC1, NOL and RCCR in yeast two-hybrid assays, indicating that HCAR is a component of the proposed SGR-CCE-LHCII complex, which acts in chlorophyll breakdown. Notably, HCAR and NOL are expressed throughout leaf development and are drastically down-regulated during dark-induced senescence, in contrast with SGR, NYC1, PPH and PAO, which are up-regulated during dark-induced senescence. Moreover, HCAR and NOL are highly up-regulated during greening of etiolated seedlings, strongly suggesting a major role for NOL and HCAR in the chlorophyll cycle during vegetative stages, possibly in chlorophyll turnover.     

A chloroplast envelope membrane protein containing a putative LrgB domain related to the control of bacterial death and lysis is required for chloroplast development in Arabidopsis thaliana

? A protein encoded by At1g32080 was consistently identified in proteomic studies of Arabidopsis chloroplast envelope membranes, but its function remained unclear. The protein, designated AtLrgB, may have evolved from a gene fusion of lrgA and lrgB. In bacteria, two homologous operons, lrgAB and cidAB, participate in an emerging mechanism to control cell death and lysis. ? We aim to characterize AtLrgB using reverse genetics and cell biological and biochemical analysis. ? AtLrgB is expressed in leaves, but not in roots. T-DNA insertion mutation of AtLrgB produced plants with interveinal chlorotic and premature necrotic leaves. Overexpression of full-length AtLrgB (or its LrgA and LrgB domains, separately), under the control of CaMV 35S promoter, produced plants exhibiting veinal chlorosis and delayed greening. At the end of light period, the T-DNA mutant had high starch and low sucrose contents in leaves, while the 35S:AtLrgB plants had low starch and high sucrose contents. Metabolite profiling revealed that AtLrgB appeared not to directly transport triose phosphate or hexose phosphates. In yeast cells, AtLrgB could augment nystatin-induced membrane permeability. ? Our work indicates that AtLrgB is a new player in chloroplast development, carbon partitioning and leaf senescence, although its molecular mechanism remains to be established.     

Effects of Bois noir on carbon assimilation, transpiration, stomatal conductance of leaves and yield of grapevine (Vitis vinifera) cv. Chardonnay

Bois noir (BN) is one of the main phytoplasma diseases of grapevine (Vitis vinifera). It is widespread, and can cause severe losses in European vineyards. The infective agent colonizes phloem elements and induces visible symptoms of leaf yellowing or reddening after a relatively long incubation period. As the most sensitive cultivars to BN, Chardonnay plants were grouped as healthy or symptomatic in spring, based on the records from the previous year. Leaf gas exchange and chlorophyll a fluorescence were measured weekly from July to September in healthy plants, and in symptomatic and asymptomatic leaves from symptomatic plants. The midday relative water content (mRWC) was measured once per month. The detection of phytoplasma DNA by nested-polymerase chain reaction revealed BN infection in symptomatic leaf samples at the end of September. A significant decrease in pigment content and maximum quantum efficiency of photosystem II (Fv/Fm) of these symptomatic leaves was detected from July to September, although in the asymptomatic leaves of the symptomatic plants the net photosynthesis (Pn) decrease was not significant. In the leaves from the healthy plants, Pn and transpiration were relatively stable. Of note, in July, an initially healthy plant showed a strong Pn reduction that was followed by visible leaf yellowing symptoms only in August. The phytoplasma infection also stimulated significant reductions in mRWC of the symptomatic leaves, with a final large decrease in yield.     

Characterization and kinetics of soybean maturation and monocarpic senescence

Daring monocarpic senescence in potted soybeans ( Glycine maxi (L.) Merrill cv. Anoka) grown in controlled-environment chambers, foliar chlorophyll, soluble protein nitrogen, total nitrogen, and starch decline (roughly in that order). All of these precede visible yellowing and, of course, abscission. The pattern of yellowing within a leaf is not uniform and is closely paralleled by starch loss. Unexpectedly, acid-soluble nitrogen rises slightly before the total foliar nitrogen declines. Foliar fresh weight and total dry matter/cm2 of leaf surface decline little if at all before shedding. Preceding and even during the foliar yellowing, the seeds rapidly accumulate dry matter and nitrogen. Yellowing appears first in the radicle tip, then in the rest of the axis and the leaves and finally in the carpels. Ability to germinate is acquired at about the time the radicle + hypocotyl turns yellow. The relationship between these changes and their role in senescence is discussed.     

Photosynthesis-Related Proteins Play Crucial Role During Senescence Stage in Flue-Cured Tobacco Plants: A Combine iTRAQ-PRM Study

Leaf aging is a significant process during herbaceous plant senescence, which is influenced by various internal and external factors. During leaf aging, chlorophyll catabolism is one of the most important metabolism pathways and results in leaf yellowing. Understanding the underlying mechanism is important for the regulation of senescence in tobacco leaf. However, there are few studies on explaining tobacco leaf senescence from the proteomics level. Here, photosynthesis experiments, cell ultrastructure, and proteomics were used to study tobacco leaves of different growth stages. We applied iTRAQ-based quantitative proteomics and parallel reaction monitoring (PRM) to determine the accumulation of proteins in aging tobacco leaves. Overall, we screened 4747 proteins. The result of KEGG pathways analysis showed that differently expressed proteins (DEPs) were involved in four pathways: metabolic pathways, biosynthesis of secondary metabolites, microbial metabolism in diverse environments, and starch and sucrose metabolism. This would be first report based on iTRAQ-PRM technique, in which we identified proteins related to photosynthesis showed a differently expressed during senescence stage in flue-cured tobacco plants.

     

Translocation of radioactive kinetin

Kinetin has generally been thought to be immobile in plants. This was confirmed in the case of laminar applications in this study, but not in regard to petiole, vein, or root applications. Radioactivity from kinetin-8-¹⁴C (Kn^*) moved freely in the vascular system of several types of leaves. This movement was usually distal to the point of application and seemed to occur with the transpiration stream. Basipetal as well as acropetal translocation of radioactive kinetin was achieved in tobacco leaves. The translocated material was extracted from veinal tissue, shown to be radioactive, and to be able to retard senescence. Similar but less decisive results were obtained from agar blocks inserted into the vascular system of leaves receiving Kn^* by petiole uptake.

A bioassay employing disks from primary bean leaves was developed for the qualitative determination of substances like kinetin which possess the ability to retard chlorophyll breakdown and plant senescence. The use of radioactive kinetin provided a refinement in this bioassay because treated non-senescent areas could be correlated with exposed areas on radioautographs made from dried leaf disks.

Root treatments showed that cotton seedlings did not take up Kn^* but that similarly treated tobacco seedlings both absorbed and translocated the isotope readily.

     

Leaf yellowing and anthocyanin accumulation are two genetically independent strategies in response to nitrogen limitation in Arabidopsis thaliana

For the first time in Arabidopsis thaliana, this work proposes the identification of quantitative trait loci (QTLs) associated with leaf senescence and stress response symptoms such as yellowing and anthocyanin-associated redness. When Arabidopsis plants were cultivated under low nitrogen conditions, we observed that both yellowing of the old leaves of the rosette and whole rosette redness were promoted. Leaf yellowing is a senescence symptom related to chlorophyll breakdown. Redness is a symptom of anthocyanin accumulation related to whole plant ageing and nutrient limitation. In this work, Arabidopsis is used as a model system to dissect the genetic variation of these parameters by QTL mapping in the 415 recombinant inbred lines of the Bay-0xShahdara population. Fifteen new QTLs and two epistatic interactions were described in this study. The yellowing of the rosette, estimated by visual notation and image processing, was controlled by four and five QTLs, respectively. The visual estimation of redness allowed us to detect six QTLs among which the major one explained 33% of the total variation. Two main QTLs were confirmed in near-isogenic lines (heterogenous inbred family; HIF), thus confirming the relevance of the visual notation of these traits. Co-localizations between QTLs for leaf yellowing, redness and nitrogen use efficiency described in a previous publication indicate complex interconnected pathways involved in both nitrogen management and senescence- and stress-related processes. No co-localization between QTLs for leaf yellowing and redness has been found, suggesting that the two characters are genetically independent.     

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