Photosynthate Partitioning into Starch in Soybean Leaves: I. Effects of Photoperiod versus Photosynthetic Period Duration

Photosynthesis, photosynthate partitioning into foliar starch, and translocation were investigated in soybean plants (Glycine max (L.) Merr. cv. Amsoy 71), grown under different photoperiods and photosynthetic periods to determine the controls of leaf starch accumulation. Starch accumulation rates in soybean leaves were inversely related to the length of the daily photosynthetic period under which the plants were grown. Photosynthetic period and not photoperiod per se appears to be the important factor. Plants grown in a 14-hour photosynthetic period partitioned approximately 60% of the daily foliar accumulation into starch whereas 7-hour plants partitioned about 90% of their daily foliar accumulation into starch. The difference in starch accumulation resulted from a change in photosynthate partitioning between starch and leaf residual dry weight. Residual dry weight is defined as leaf dry weight minus the weight of total nonstructural carbohydrates. Differences in photosynthate partitioning into starch were also associated with changes in photosynthetic and translocation rates, as well as with leaf and whole plant morphology. It is concluded that leaf starch accumulation is a programmed process and not simply the result of a limitation in translocation.     

Photosynthate Partitioning into Starch in Soybean Leaves: II. IRRADIANCE LEVEL AND DAILY PHOTOSYNTHETIC PERIOD DURATION EFFECTS

Two photosynthetic periods and photosynthetic photon flux densities (PPFD) were used to study the relationship between the rate of photosynthesis and starch accumulation in vegetative soybean leaves (Merr. cv Amsoy 71). Plants grown in short daily photosynthetic periods (7 hours) had higher rates of CO₂ fixation per unit leaf dry weight and of leaf starch accumulation than plants grown in long daily photosynthetic periods (14 hours) irrespective of PPFD. CO₂ fixation rates per unit leaf area were similar in 7-hour and 14-hour plants grown at low PPFD but were highest in 14-hour plants at the high PPFD. When single leaves of 14-hour plants were given 7-hour photosynthetic periods, their rates of starch accumulation remained unchanged. The programming of starch accumulation rate and possibly of photosynthetic rate by the length of the daily photosynthetic period is apparently a whole-plant, not an individual leaf, phenomenon. Programming of chloroplast starch accumulation rate by length of the daily photosynthetic and/or dark periods was independent of PPFD within the ranges used in this experiment.     

Effects of Nitrate Level on Nitrogen Metabolism in Winged Bean and Soya Bean

The effects of high (15 mM) and low (0.75 mM) solution nitratelevels on nitrogen metabolism in three genotypes (IL 7A, IL13 and IL 21) of winged beans [Psophocarpus tetragonolobus (L.)DC.] and one genotype (Williams) of soya bean [Glycine max (L.)Merrill] were investigated. Plants were grown for 42 days ina greenhouse in solution culture prior to sampling. The 15 mM nitrate treatment resulted in greater growth of allplant parts except roots. Growth of soya beans was more responsiveto nitrate level than was growth of winged beans. The high nitratelevel inhibited nodulation in all plants. The IL 13 and IL 21winged bean genotypes had similar nitrogenase activity (acetylenereduction per plant) as the soya bean and IL 7A winged beangenotype had lower activity. However, the IL 13 winged beangenotype had higher nitrogenase activity (acetylene reductionper unit nodule mass) than the other three genotypes which allhad similar activity. The 15 mM solution nitrate level stimulatedleaf and root nitrate reductase (NR) activity for all plants.All winged bean genotypes had higher leaf NR activity and higherpercentage reduced- and nitrate-nitrogen contents of leavesand stems compared with soya beans. However, total protein (reducednitrogen) was greater in soya beans when sampled indicatingthat more nitrate had been metabolized by soya beans than bywinged beans during the 42-day growth period. Psophocarpus tetragonolobus (L.) DC., winged bean, Glycine max (L.) Merrill, Soya bean, nitrate reductase, nitrogen fixation, nitrogenase activity, nodulation     

Sucrose and Starch Synthesis in Spinach Plants Grown under Long and Short Photosynthetic Periods

The flow of carbon into sucrose and starch was investigated in fully expanded primary leaves of spinach using the long to short day transition and partial defoliation as tools to manipulate sucrose/starch synthesis. Transfer from 12 hour to 7 hour photosynthetic periods resulted in a 4-fold increase in the initial rate of starch synthesis, a 50% increase in the initial rate of sucrose synthesis, a 30% increase in leaf sucrose, and a 40% decrease in fructose, 2,6-biphosphate. In addition, sucrose synthesis rates in cells isolated from shortened daylength plants are 80% higher than in cells isolated from control plants. These results show that, in spinach, an increase in the rates of both sucrose and starch synthesis can occur under short day conditions. In contrast, when short day plants are partially defoliated, starch levels remain high, fructose 2,6-biphosphate levels remain low, but the level of leaf sucrose drops by 50%. Thus, when demand exceeds supply, starch synthesis has priority over filling of leaf sucrose pools in the short day plant.     

Regulation of Photosynthetic Metabolism by Low-Temperature Treatment of Roots of Single-Rooted Soybean Plants

The single-rooted leaf of soybean (Glycine max L. Merr.) wasused to study source-sink relationships in photosynthesis. Theroots grown in a culture solution at 20 ? 1?C were treated withlow temperatures (minimum 6?C) for 4 days to change the source/sinkbalance. This treatment suppressed the increase of root dryweight, but by 2 days after the end of the treatment, the leafdry weight showed a 1.7-fold increase mainly due to the accumulationof starch and sucrose. In the roots, both contents also increasedsignificantly. The sucrose and starch contents in leaves andthe sucrose content in roots began decreasing immediately afterthe treatment was stopped, while the starch content in rootsremained at the high level. The leaf area did not change throughoutthe experimental period. The low-temperature treatment of theroots caused an abrupt decrease in the photosynthetic activity(41% of the initial value by day 4) with slow recovery afterthe stop treatment was stopped. A significant negative correlationwas found between the photosynthetic rate and the sucrose contentin the leaves given the low-temperature treatment, but no significantcorrelation was found between the rate and the starch content.This result indicates that the photosynthetic metabolism ismore directly influenced by the amount of sucrose in the cytoplasmthan by the starch content in the chloroplast when the source/sinkbalance is changed by low-temperature treatment to suppressroot growth. (Received May 6, 1986; Accepted December 6, 1986)     

Photosynthate partitioning into leaf starch as affected by daily photosynthetic period duration in six species

Starch accumulation rates increased up to five-fold in the leaves of five out of six species examined when plants were shifted from a long (12 or 14-h) to a short (7-h) photosynthetic period. The five species that responded were corn (Zea mays L.), pangola (Digitaria decumbens Stent.), soybean (Glycine max L.) Merr.), spinach (Spinacia oleracea L.), and sugarbeet (Beet vulgaris L.) However, the rate of starch accumulation in alfalfa (Medicago sativa L.) was unaltered following a shift from a long to a short photosynthetic period. We conclude that photosynthate partitioning into chloroplast starch, at least in a wide variety of species, is a programmable process that can be manipulated by altering the length of the daily photosynthetic period.     

Assumptions of Plastochron Index: Evaluation With Soya Bean Under Field Drought Conditions

Erickson and Michelini (1957) derived the plastochron index(PI) and a term sometimes referred to as the plastochron ratio(PR), as quantitative expressions of the vegetative developmentof plants. With the stable plant growth in environmental chambersand glasshouses, the assumptions used to derive these termshave been validated. However, more recently these expressionsare being used to characterize growth under the unstable conditionsresulting from the imposition of stress. This study examinesthe validity of the assumptions used to derive PI and PR forfield-grown soya beans [Glycine max (L.) Merrill] subjectedto drought stress. Under stress conditions, the assumptionswere not satisfied. In fact, observing change in PR appearedto be a good method for detecting drought stress in these plants.An alternate method for calculating PI based on a single, youngleaf was developed. This alternate method appeared to be a moresensitive indicator of changes in leaf emergence rate underunstable conditions. Plastochron index, plastochron ratio, Glycine max (L.), soya bean, drought, leaf growth     

Starch Metabolism in Developing and Germinating Soya Bean Seeds is Independent of {beta}-Amylase Activity

Developing seeds of soya bean cultivars Chestnut and Altonahave only trace amounts of ß-amylase activity. Comparedto a standard variety, Wells, ß-amylase activitieswere 200–300 times lower in Chestnut and Altona. Nevertheless,Chestnut and Altona accumulate starch as a transient reservematerial which is utilized later in development. Seeds of Chestnutand Altona also produce starch early in germination which subsequentlydeclines after the 4th day of germination. Throughout germinationß-amylase levels in these cultivars are about 300-foldlower than that observed in Wells, which has a similar patternof starch metabolism. Widely varying levels of ß-amylasein both developing and germinating seeds appear to be unrelatedto starch metabolism which is very similar in all cultivarsstudied. Consequently, ß-amylase activity seems irrelevantto starch metabolism in the soya bean seed. starch, ß-amylase, Glycine max. (L.), Merr, soya bean     

Effect of Flowering on the Photosynthetic Capacity of Ryegrass Leaves Grown With and Without Natural Shading

The photosynthetic capacity of newly expanded leaves of vernalizedor non-vernalized plants of S24 perennial ryegrass (Lolium perenneL.), grown in long or short photoperiods, was measured in twoexperiments. In the first, leaves were protected from shadingduring development, while in the second, the natural shade ofneighbouring tillers in a sward was allowed. In the first experiment there was little effect of vernalization,day length or flowering, and leaves in all treatments had photosyntheticrates at 250 W m^–2 of between 28 and 32 mg CO₂ dm^–2h_–1.In the second experiment the photosynthetic rate of successiveleaves fell as sward leaf area increased. This downward trendwas reversed, however, in flowering tillers in the vernalizedlong-day treatment, while in the other treatments, which didnot flower, photosynthetic capacity continued to fall. It isconcluded that the leaves of reproductive tillers have highphotosynthetic capacities because stem extension carries themto the top of the canopy where they are well illuminated duringexpansion. Lolium perenneL, ryegrass, photosynthetic capacity, flowering, shading, vernalization     

The Role of Cotyledons in Vegetative Growth and Nitrogen Assimilation by Soya Bean

The removal of both cotyledons from soya bean seedlings 10 daysafter sowing, when the primary leaves were unfolded, reducedtheir stem height, branching, leaf production and dry weightat flowering by a similar proportion whether they were nodulatedor nitrate-dependent. Nitrogen assimilation by the shoots ofnitrate-dependent plants was increased by the removal of onecotyledon and reduced by the removal of both cotyledons althoughthese effects were not significant. Both these treatments significantlyincreased the amount of nitrogen in the shoots of nodulatedplants at flowering, mainly by more than doubling the nitrogencontent of their leaves. In contrast, the proportion of thetotal plant nitrogen in the leaves of nitrate-dependent plantswas almost constant. These results suggest that the cotyledonsmarkedly inhibit nitrogen assimilation by nodulated plants butdo not appreciably affect nitrogen assimilation by plants dependentsolely on inorganic nitrogen for their nitrogen supply. Glycine mux (L) Merr., soya bean, cotyledons, nitrogen assimilation, growth     

Photosynthetic Determinants of Growth in Maize Plants: Effects of Nitrogen Nutrition on Growth, Carbon Fixation and Photochemical Features

Maize(Zea mays L.) plants were grown in a greenhouse with differentlevels of nitrate-N (2 to 20 millimolar). Nitrogen nutritionhad dramatic effects on plant growth and photosynthetic characteristicsof mature leaves. Increasing nitrogen resulted in greater biomassproduction, shoot/root ratios, and rates of leaf expansion duringthe day. The elongating zone of high-N plants had higher activities(per gram fresh weight) of sucrose synthase and neutral invertasethan low-N plants, suggesting that increased leaf growth wasrelated to a greater biochemical capacity for sucrose metabolism. Mature leaves of high-N plants had higher rates of photosynthesisand assimilate export (sucrose formation), and partitioned morecarbon into sucrose relative to starch. Increased photosyntheticrates (leaf area basis) were associated with higher levels ofribulose-l,5-bisphosphate carboxylase, phosphoenolpyruvate carboxylaseand pyruvate, phosphate dikinase (determined immunochemically).In addition, N-nutrition affected the functional organizationof chlorophyll in the leaves. Large increases in the numberof PS I reaction centers were observed which fully accountedfor increases in leaf chlorophyll content with increasing nitratesupply. Collectively, the results suggest that increased growth of maizeplants at high light and optimal nitrogen nutrition is relatedto greater capacity for photosynthesis and translocation inmature leaves, and possibly increased capacity for sucrose metabolismin expanding leaves. (Received May 22, 1989; Accepted August 28, 1989)     

Mineral Depletion and Leaf Senescence in Soya Bean as Influenced by Foliar Nutrient Application During Seed Filling

Senescence, as judged by the time courses of leaf lamina photosynthesis,soluble protein and chlorophyll contents, was studied in relationto mineral redistribution in field-grown soya beans [Glycinemax (L.) Merr] to investigate the hypothesis that the depletionof nutrients m the leaves by the developing seeds is the causeof soya bean senescence. A mineral nutrient solution was appliedto the canopy during the seed-filling period, and the effectson senescence and mineral depletion of the leaves were determinedin three cultivars, at two leaf positions, weekly from beginningof seed filling through physiological maturity. The onset of senescence occurred shortly after the beginningof rapid seed filling Photosynthetic rate declined about 60per cent within 3 weeks. Protein dropped by 52 per cent andchlorophyll by 48 per cent over the same period. Foliar nutrient application, at a rate previously shown to givesignificant yield increases in soya beans, increased the concentrationsof N, P and K in the leaf laminae, but tended only to delaytheir decline and failed to either delay the onset or alterthe course of senescence. The results of this experiment seem to indicate that, undernormal growth conditions, the events of senescence in the soyabean are not causally related to the N, P or K concentrationsof the leaf laminae Glycme max (L.) Merr., soya bean, nitrogen, phosphorus, potassium, leaf protein, chlorophyll, photosynthesis, foliar nutrient application, mineral depletion, leaf senescence     

Starch Accumulation and Photosynthetic Activity in Primary Leaves of Bean (Phaseolus vulgaris L.)

The effects of starch accumulation on photosynthesis and chloroplastultrastructure were studied in primary leaves of bean (Phaseolusvulgaris L. cv. Bulgarian). De-topping the shoot above the primaryleaf node, caused over an 8-day period, a considerable increasein the photosynthetic activity of the primary leaves, despitethe fact that a large quantity of starch had accumulated intheir chloroplasts. The accumulation of starch was greater inthe chloroplasts of spongy cells in comparison with that ofthe palisade cells. Initiation of starch grains was observedmainly in the peripheral part of the chloroplast, distant fromthe cell wall. As a result, most of the starch was accumulatedclose to the inner part of the cell, leaving a considerablemass of the chloroplast near the cell wall free of starch. Theaccumulation of starch was accompanied by the destruction, deformationand disorientation of grana and thylakoids. It is concludedthat the accumulation of starch is not inevitably a limitingfactor in photosynthesis and the results cast doubt on the hypothesisthat starch accumulation or dissipation is the main factor involvedin the regulation of photosynthesis. Phaseolus vulgaris L, bean, photosynthesis, starch accumulation, chloroplast ultrastructure     

Photosynthetic Acclimation in Pea and Soybean to High Atmospheric CO2 Partial Pressure

Nonnodulated pea (Pisum sativum L. cv Frosty) and soybean (Glycine max [L.] Merr. cv Wye) plants were grown under artificial lights from germination with ample nutrients, 600 [mu]mol photons m-2 s-1, and either 34 to 36 (control) or 64 to 68 Pa (enriched) CO2. For soybean, pod removal and whole-plant shading treatments were used to alter the source-sink balance and carbohydrate status of the plants. Growth of both species was substantially increased by CO2 enrichment despite some down-regulation of photosynthesis rate per unit leaf area ("acclimation"). Acclimation was observed in young pea leaves but not old and in old soybean leaves but not young. Acclimation was neither evident in quantum yield nor was it related to triose phosphate limitation of net photosynthesis. A correlation between levels of starch and sugars in the leaf and the amount of acclimation was apparent but was loose and only weakly related to the source-sink balance of the plant. A consistent feature of acclimation was reduced ribulose bisphosphate carboxylase (RuBPCase) content, although in vivo RuBPCase activity was not necessarily diminished by elevated growth CO2 owing to increased percentage of activation of the enzyme. A proposal is discussed that the complexity of photosynthetic acclimation responses to elevated CO2 is as an expression of re-optimization of deployment of within-plant resources at three levels of competition.     

The Effects of Shading and Cutting Treatments on the Photosynthetic Rate of Ryegrass Leaves

As the leaf area of miniature simulated swards of S24 perennialreygrass (Lolium perenne L.) grown in controlled environmentincreased, successive leaves which expanded on the main stemof a sample plant within the sward had progressively lower photosyntheticcapacities. When the plants surrounding the sample plant were cut, the photosyntheticcapacity of the next leaf to expand was increased, whether ornot other leaves on the sample plant had been removed. Whenall fully expanded leaves on the sample plant were removed butthe surrounding plants left uncut, the photosynthetic capacityof successive leaves on the sample plant continued to decreaseas in uncut swards. These results support the view that the changes in photosyntheticcapacity in grass swards are consequences of changes in thelight climate of the developing leaf rather than of changesin demand for assimilates or growth substance supply. The declinein photosynthetic capactiy was contributed to by both stomataland residual diffusion resistances.     

Starch Deposition and Carbohydrase Activities in Developing and Germinating Soya Bean Seeds

Immature soya bean seeds accumulate starch as a transient reservematerial which is utilized later in development. Germinatingseeds also accumulate starch reserves, probably as a resultof gluconeogenesis from storage lipid. Developing beans showa rapid increase in ß-amylase activity which continuesinto early germination before declining. Distribution of ß-amylaseactivity is not consistent with its supposed role in starchdegradation. Soya bean seeds also contain -amylase and -glucosidaseactivities which could be responsible for starch mobilization. Glycine max (L.) Merr., soya bean, starch, carbohydrase, amylase, -glucosidase     

Influence of Carbohydrates on Photosynthesis in Single, Rooted Soybean Leaves Used as a Source-Sink Model

The single rooted leaf of soybean (Glycine max L. Merr.) wasused to study source-sink relationships in photosynthesis. Whenthe leaves were kept under a regime of 10 h light (410–480µmol photons m^–2, 400–700 nm)–14 h dark,they did not expand, the increase in leaf dry weight almoststopped, and photosynthetic activity remained at a high andconstant level for 8 d while the dry weight of the roots increasedat a constant rate throughout the period. Thus, under this conditionthe leaf and the root system served as the only source and sinkorgans, respectively. When leaves grown for 7 d under this conditionwere placed under continuous light to alter the source/sinkbalance in photosynthate, the root dry weight increased at aconstant rate equal to that found under the 10 h light–14h dark condition. The leaf dry weight markedly increased andby day 5 of continuous light had increased 1.6-fold, mainlyas a consequence of accumulation of starch and sucrose, whichwere not translocated for root growth. The continuous lightcaused an abrupt decrease in the photosynthetic activity (40%of initial value by day 5). However, the activity recoveredalmost completely after a 32-h transfer to darkness. Significantnegative correlations existed between photosynthetic activityand the sucrose and starch contents in the rooted leaves placedunder continuous light. When the plants were treated with variouslight conditions, there was no significant difference (p<0.01)among the regression line slopes for photosynthetic activityon the sucrose content, but there was some deviation among thosefor the photosynthetic activity on the starch content. Thisresult suggests that sucrose accumulated in the leaf has a moredirect influence on photosynthetic activity when the source/sinkbalance was altered. (Received September 9, 1985; Accepted February 21, 1986)     

不同光质对桑树幼苗生长和光合特性的影响

光质可影响植物光合特性、形态以及生理过程。本试验研究了不同光质(白光W、红光R、红蓝混合光RB、蓝光B)对桑树植株生长、形态和光合作用的影响。结果表明:与白光对照相比,红光、蓝光和红蓝混合光处理下植株的生长、干物质积累受到抑制;红光处理下植株的株高、叶面积显著高于白光、红蓝混合光、蓝光处理;而白光、红蓝混合光、蓝光处理下植株的LMA、叶绿素a/b比值、可溶性蛋白含量、蔗糖、淀粉含量和叶片总N含量显著高于红光处理;红蓝混合光处理下植株的Pn、Gs、ΦPSⅡ与白光处理相近,红光、蓝光处理下植株的Pn、ΦPSⅡ低于白光、红蓝混合光处理,同时红光、红蓝混合光、蓝光处理下植株的抗氧化酶活性高于白光处理,而MDA含量低于白光处理;红光处理下植株的叶片厚度、栅栏组织和海绵组织厚度显著小于白光处理。因此,一定比例的红蓝混合光可以使桑树植株的生长、光合特性、生理特征和叶片解剖结构与白光下生长植株相近,并减少单质红光、单质蓝光对植株生长发育的不利影响。     

Effect of shortened photosynthetic period on C-assimilate translocation and partitioning in reproductive soyeans

Starch accumulation rate in leaves of vegetative soybeans is inversely related to the length of the daily photosynthetic period. However, it is not known whether a similar response would be observed during reproductive growth. Soybeans (Glycine max L. Merr. cv Amsoy 71) were grown to three stages of reproductive growth (beginning seed, mid seed-fill, and late seed-fill) under 12-hour daylengths, and then shifted to 6-hour photosynthetic periods (12-hour photoperiods) for 4 days. One and 4 days after treatment, a mid-canopy leaf was pulsed with ¹⁴CO₂, and sampled for radiolabeled starch and water-soluble compounds at 0.5, 1, 3, 9, and 21 hours after labeling.

Plants exposed to the 6-hour photosynthetic periods at the beginning seed stage retained and incorporated significantly more label as starch than did those given 12-hour photosynthetic periods. However, plants exposed to the shortened photosynthetic periods at the late seed-fill stage partitioned less label into starch. Plants exposed at mid seed-fill gave a variable response.

Shortened photosynthetic periods resulted in preferential partitioning of recently fixed carbon to the seed at the expense of the pod wall. The results of these experiments suggest that the increased sink demand present during late reproductive growth may be of greater importance in control of leaf starch accumulation than is the length of the daily photosynthetic period.

     

The Effect of Adenine and Mevalonic acid on the Endogenous Cytokinins of a Cytokinin-dependent Strain of Soya Bean Callus

The combined application of 10^–6 M adenine and 10^–6M mevalonic acid to soya bean callus accelerated its growth.Two biologically active compounds that co-chromatographed withzeatin and isopentenyl adenine were extracted from this callus.Studies with labelled adenine and mevalonic acid indicated thatthe cytokinin-dependent soya bean callus incorporated only avery small amount of the radioactive precursors into the biologically-activecompounds, making it extremely difficult to determine whetherthese compounds were synthesized de novo or whether they aroseas by-products of tRNA turnover. As cytokinins do not accumulatein rapidly-growing cytokinin-dependent soya bean callus culturedon kinetin as a source of cytokinin it seems as if biosynthesisde novo occurs when the callus is supplied with adenine andmevalonic acid. Glycine max (L.) Merrill, soya bean, callus culture, adenine, mevalonic acid, endogenous cytokinins