Phosphate Deficiency in Maize. I. Leaf Phosphate Status, Growth, Photosynthesis and Carbon Partitioning

Maize plants (Zea mays L.) were cultured with nutrient solutioncontaining 0.001 or 0.5 mM orthophosphate (Pi). Effects of lowphosphate (low-P) nutrition on growth, leaf phosphate status,photosynthesis, and carbon partitioning were investigated. Withlow-P treatment, the fresh weight of aerial parts decreasedby about 40% by 24 days after planting. Detailed studies ofthe effects of low-P treatment on the other characteristicsof maize leaves-were done using the middle part of the thirdleaf, counting from the base. Low-P treatment had almost noeffect on specific leaf weight or soluble protein content measured13 to 21 days after planting. Low-P treatment decreased Chicontent slightly (by 15% 19 days after planting). Twenty onedays after planting, low-P treatment had greatly decreased thelevels of leaf acid extractable Pi (by 77%) and photosynthesisrates (by 68%). The detrimental effects of low-P treatment onthe rates of photosynthesis and the amounts of acid extractablePi became progressively greater with time. There was a strongcorrelation between levels of leaf acid extractable Pi and ratesof photosynthesis. The minimum level of Pi necessary to sustainthe maximum photosynthesis rate was 0.6 mmol m^–2. Belowthis minimum content of Pi, the rate of photosynthesis decreasedsharply with decreasing Pi. To investigate the direct effectof Pi depletion on photosynthate partitioning at equivalentrates of photosynthesis, the rates in controls were reducedto almost the same as those in 18 or 19 day old low-P plants(about 50% of those in controls) by lowering light intensityand/ or ambient CO₂ concentration. The data clearly indicatesthat low-P treatment had a direct effect in lowering photosynthatepartitioning into starch. Starch mobilization during the nightwas also inhibited under low-P conditions. (Received January 7, 1991; Accepted March 5, 1991)     

Effects of Benzyladenine on Photosynthesis, Growth and Senescence of the Bean Plant

The net photosynthetic rate of attached primary bean leaves treated with benzyladenine (BA) decreased from 35 nano-grams carbon dioxide per square centimetre and second (ng cm^?2 s^?1) at week 2 to 17 ng cm^?2 s^?1 at week 4, and thereafter increased to 24 at week 6. By contrast the net photo-synthetic rate of the water-treated leaves decreased to 4–5 ng cm^?2 s^?1 at weeks 5 and 6. The stomatal resistance was not markedly affected by BA treatment. The BA-treated primary leaves had higher dry and specific weights, and the chlorophyll and carotenoid contents were greater than for the water controls. The pigment content of the BA-treated leaves steadily increased from week 2 to 6, whereas in the water controls it remained constant till week 4 and thereafter decreased. It is concluded that retardation of leaf senescence by BA is assoclatcd with a mnintenance of photosynthetic activity.     

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)     

Growth Activity of the Gibberellins of Dwarf French Bean, Potato, and Lettuce

Zones with similar Rf to gibberellic acid (GA) on chromatogramsof ethylacetate extracts of dwarf French bean and potato leavesand stems and lettuce heads all promoted growth of disks ofetiolated dwarf French bean leaves in darkness. The presence of a gibberellin was confirmed by tests with stemsof dwarf pea, but that the substance was not GA was shown bythe lack of effect in the tests with lettuce hypocotyls. Thetests with dwarf maize, effects on d-3 and d-5, but not d-1,identified it as gibberelliin A₅ (bean factor II). Materialthat promoted the growth of lettuce hypocotyls occurred in zonesclose to the starting-line of chromatograms of extracts of beanand lettuce leaves. Calibration of the growth tests with GA gave an estimate ofthe gibberellin content of the tissues extracted. In the dwarf-peatest the apparent gibberellin contents were similar to the valuesin the leaf-disk test, but very low values occurred in the maizetests.     

光强和施氮量对催吐萝芙木叶片生长及光合作用的影响

通过不同光强（15%、40%和70%自然光强）和施氮量（15、30和60g/株）的盆栽试验,研究了不同光照强度和施氮量对催吐萝芙木（Rauvolfia vomitoria Afzel.）叶片生长和光合特性的影响。结果表明：光强和施氮量显著影响了催吐萝芙木叶片净光合速率（Pn）、气孔导度（Gs）、水分利用效率（WUE）、叶绿素含量（Chl）、比叶面积（SLA）和叶生物量比（LMR）（p〈0.01）,Pn和Gs均随光强、施氮量的增加而增大,70%自然光强下叶片Pn和Gs显著高于15%和40%自然光强处理。总体而言,低光条件下,更有利于其叶绿素的合成,且施氮量对叶绿素含量的影响不大。低光处理和重度施氮量均有利于催吐萝芙木叶片SLA增大和叶生物量的分配,但实验中光强和施氮量处理并未引起催吐萝芙木叶绿素荧光参数Fv/Fm发生显著变化。光强和施氮量对催吐萝芙木叶片净光合速率（P）、气孔导度（G）、水分利用效率（WUE）等光合生理指标具有显著的交互作用（p〈0.05）。     

Effects of Elevated CO2 and High Temperature on Single Leaf and Canopy Photosynthesis of Rice

The increase of atmospheric CO2 concentration is indisputable. In such condition, photosynthetic response of leaf is relatively well studied, while the comparison of that between single leaf and whole canopy is less emphasized. The stimulation of elevated CO2 on canopy photosynthesis may be different from that on single leaf level. In this study, leaf and canopy photosynthesis of rice ( Oryza sativa L. ) were studied throughout the growing season. High CO2 and temperature had a synergetic stimulation on single leaf photosynthetic rate until grain filling. Photosynthesis of leaf was stimulated by high CO2, although the stimulation was decreased by higher temperature at grain filling stage. On the other hand, the simulation of elevated CO2 on canopy photosynthesis leveled off with time. Stimulation at canopy level disappeared by grain filling stage in beth temperature treatments. Green leaf area index was not significantly affected by CO2 at maturity, but greater in plants grown at higher temperature. Leaf nitrogen content decreased with the increase of CO2 concentration although it was not statistically significant at maturity. Canopy respiration rate increased at flowering stage indicating higher carbon loss. Shading effect caused by leaf development reached maximum at flowering stage. The CO2 stimulation on photosynthesis was greater in single leaf than in canopy. Since enhanced CO2 significantly increased biomass of rice stems and panicles, increase in canopy respiration caused diminishment of CO2 stimulation in canopy net photosynthesis, keaf nitrogen in the canopy level decreased with CO2 concentration and may eventually hasten CO2 stimulation on canopy photosynthesis. Early senescence of canopy leaves in high CO2 is also a possible cause.     

外源硅对干旱胁迫下烟草幼苗生长、叶片光合及生理指标的影响

采用营养液水培法,研究了施用不同浓度外源硅(0、0.5、1.0 mmol/L)对干旱胁迫(10%PEG、20%PEG)下烟草幼苗生长、叶片光合特性和生理指标的影响。结果表明:干旱胁迫严重抑制了烟草幼苗生长和光合作用,膜质稳定性降低和引起氧化应激反应;施用不同浓度外源硅有效改善了干旱胁迫下烟草幼苗生长,均表现为株高、叶面积、根系体积、根系干重和地上部干重等生长指标增加,提高叶绿素a、叶绿素b、叶绿素a+b和类胡萝卜素含量,显著提高净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)并降低胞间CO2浓度(Ci),膜质过氧化产物MDA含量显著降低,提高叶片含水量、膜稳定性系数和渗透调节物质(脯氨酸、可溶性糖)含量,显著提高SOD、POD和CAT等抗氧化酶活性,而且1.0 mmol/L Si处理对干旱胁迫下烟草幼苗生长和生理特性的影响显著优于0.5 mmol/L Si处理。以上结果说明,施用外源硅能提高干旱胁迫下烟草幼苗光合作用、抗氧化和渗透调节能力,缓解干旱胁迫对烟草幼苗的伤害,促进其生长。     

The Effects of Kinetin, Gibberellic Acid, and Light on Expansion and Cell Division in Leaf Disks of Dwarf Bean (Phaseolus vulgaris)

The effects of kinetin (Kn), gibberellic acid (G), and light(L) on cell expansion and division in disks from 6-day-old etiolatedprimary leaves of dwarf bean are described. Cell number wasdetermined by direct counting after disks were digested in apectinase/EDTA mixture to separate the cells. Kn increased leafexpansion wholly by increasing cell size. G also increased cellsize; it increased cell division in the dark but not in thelight. Light also increased cell size and cell division; iteliminated the effect of G on cell division but enhanced theeffect of G on cell expansion.     

Effects of Shading the First Leaf on Growth of Barley Plants: II. Effects on Photosynthesis

The contributions made by photosynthesis in the first leaf toseedling growth have been examined by a variety of methods includinginfra-red gas analysis and the use of ¹⁴CO₂. The first leafis fully expanded by day 8 and maximal rates of photosynthesisare achieved about I day later. Up to day 8 growth of the seedlingsresults from the redistribution of seed reserves and once theseare exhausted growth is dependent upon the first leaf, beingreduced to very low levels if this is shaded. The second leafwhich begins to expand rapidly after day 10 is contributingto growth by day 14, and the contribution from the first leafbegins to decline after day 12. Apart from greatly reducing photosynthesis in treated leaves,shade also affects the development of photosynthetic capacity.When applied for 2 days or more from day 6, shade reduces thepeak level of carbon fixation achieved on days 9 and 10 by upto 35 per cent. It is shown that development of the first leafin terms of increased dry weight and photosynthetic capacity,is dependent on photosynthesis in the developing leaf itself.The mechanisms by which shading affects development are discussed.     

Growth Rate, Photosynthesis and Respiration in Relation to Leaf Area Index

This work examined three possible explanations of growth rateresponses to leaf area index (LAI) in which growth rate perunit of ground area (crop growth rate, CGR) increased to a plateaurather than decreasing above an optimum LAI at which all lightwas intercepted. Single leaf photosynthetic measurements, andwhole plant 24 h photosynthesis and respiration measurementswere made for isolated plants and plants in stands using Amaranlhushybridus, Chenopodium album, and two cultivars of Glycine maxgrown at 500 and 1000 µimol m^–2 S^–1 photosyntheticphoton flux density at 25 °C. CGR, relative growth rate(RGR), and LAI were determined from 24 h carbon dioxide exchangeand leaf area and biomass measurements. Respiration increasedrelative to photosynthesis with crowding in A. hybridus andthere was an optimum LAI for CGR. In contrast, the ratio ofrespiration to photosynthesis was constant across plant arrangementin the other species and they had a plateau response of CGRto LAI. Neither increased leaf photosynthetic capacity at highLAI nor a large change in biomass compared to the change inLAI could account for the plateau responses. It was calculatedthat maintenance respiration per unit of biomass decreased withdecreasing RGR in C. album and G. max, but not A. hybridus,and accounted for the plateau response of CGR to LAI. Sincesimilar decreases in maintenance respiration per biomass atlow RGR have been reported for several other species, a constantratio of respiration to photosynthesis may occur in more speciesthan constant maintenance respiration per unit of biomass. Amaranlhus hybridus L., Chenopodium album L., Glycine max L Merr, soybean, photosynthesis, respiration, growth, leaf area index     

叶片花色素苷对植物光合作用影响的研究进展

近年来,花色素苷在彩叶植物呈色机制的研究中备受关注,但目前有关其对彩叶植物光合特性影响的机制仍缺乏系统的评述。本文简单介绍了花色素苷的基本特性,并基于国内外相关研究进展,综述了叶片花色素苷对植物光合特性的影响机制及其对叶片光合机构的保护意义,对彩叶植物光合作用的研究方向提出了建磷。     

Effect of Plant Growth Regulators on Nitrate Utilization by Roots of Nitrogen-Depleted Dwarf Bean

We examined the effect of pretreatments (18 h at 5 µmoldm^–3) with abscisic acid, the ethylene-releasing substance‘Ethephon’, gibberellic acid, indoleacetic acid,kinetin and zeatin on nitrate uptake and in vivo nitrate reductaseactivity (NRA) in roots of nitrogen-depleted Phaseolus vulgarisL. Nitrate uptake showed an apparent induction pattern witha steady state after about 6 h, in all treatments. The nitrateuptake rate after 6 h was unaffected or at most 30% lower aftertreatments with the plant growth regulators. Gibberellic acid, kinetin and zeatin induced substantial NRAin roots in the absence of nitrate, whereas Ethephon enhancedNRA only during nitrate nutrition. Kinetin-induced NRA (Ki-NRA)was maximal after a pretreatment at 1 µmol dm^–3,and showed a lag phase of 6–8 h. Ki-NRA was additive tonitrate-induced NRA (NO^–₃-NRA) for at least 24 h, independentof the induction sequence. After full induction, Ki-NRA approximated20% of NO-₃-NRA. Abscisic acid counteracted the developmentof Ki-NRA, but not of NO^–₃-NRA. Cycloheximide and tungstatewere equally effective to suppress the development of nitratereductase activity after supply of kinetin or NO^–₃. Our data are consistent with the operation of two independentenzyme fractions (Ki-NRA and NO^–₃-NRA) with apparentlyidentical properties but with separate control mechanisms. Theabsence of major effects of plant growth regulators on the time-courseand rate of nitrate uptake suggests that exogenous regulators,and possibly endogenous phytohormones are of minor importancefor initial nitrate uptake. The differential effect of someregulators on nitrate uptake and root NRA furthermore indicatesthat the processes of uptake and reduction of NO^–₃ arenot obligatory or exclusively coupled to each other.     

Effects of Nitrate Application on Amaranthus powellii Wats. : I. Changes in Photosynthesis, Growth Rates, and Leaf Area

Physiological effects of different nitrate applications were studied using the C₄ plant, Amaranthus powellii Wats. Plants were grown in a controlled environment chamber and watered daily with nutrient solutions containing 45, 10, 5, or 1 millimolar nitrate. Chloride and sulfate were used to keep the cation and phosphate concentrations constant. Total leaf nitrogen concentration, chlorophyll concentration, specific leaf mass, leaf area, relative growth rate, relative leaf growth rate, unit leaf rate (increase of dry mass per unit leaf area per day), net photosynthetic rate, and incident quantum yield decreased with decreasing nitrate concentration. The per cent decrease of unit leaf rate was similar to the decrease of light-saturated net photosynthetic rate; however, the decrease in relative growth rate was less than that of unit leaf rate because leaf area ratio (leaf area per unit dry mass) increased with decreasing nitrate concentration. Essential mineral concentrations per unit leaf area were about equal among all treatments. Leaf expansion, determined by stomatal density, decreased except for the 1 millimolar treatment which showed relatively more cell expansion but less cell division. Decreased nitrate application was correlated with higher osmotic potentials and lower pressure potentials (determined by pressure-volume curves), whereas leaf water potentials were equal among treatments. Even though total leaf area and shoot mass decreased with decreasing applied nitrate, the increase of the leaf area ratio may be related to selection for the highest possible growth rate.     

Analysis of Growth, Photosynthesis and Light Interception for Single Plants and Stands

The two traditionally distinct treatments of growth analysis,using either relative growth rate (RGR) or crop growth rate(CGR), can be integrated in a single treatment in which RGRis a component of CGR. CGR can alternatively be analysed asthe product of incident light receipt, efficiency of light interception(as determined by leaf area index and extinction coefficient)and efficiency of use of intercepted light in dry-matter productionor in canopy net photosynthesis. Further, the net assimilationrate or net photosynthetic rate can be resolved into two componentswhich quantify the dependence of light interception on leafarea and of CO₂ assimilation on intercepted light. These relationsprovide increased flexibility in the analysis of assimilationand growth in terms of light interception and the structureof plants and stands. The usefulness of growth analysis in elucidatingphysiological mechanisms is discussed in relation to the useof more complex mechanistic models. Crop growth rate, light interception, growth analysis, leaf area index     

Photosynthesis and Plant Growth at Elevated Levels of CO2

In this review, we discuss the effects of elevated CO₂ levelson photosynthesis in relation to the whole plant growth in terrestrialhigher C₃ plants. Short-term CO₂ enrichment stimulates the rateof photosynthesis. Plant mass is also enhanced by CO₂ enrichment.However, the effects of long-term CO₂ enrichment on photosynthesisare variable. Generally, the prolonged exposure to CO₂ enrichmentreduces the initial stimulation of photosynthesis in many species,and frequently suppresses photosynthesis. These responses areattributed to secondary responses related to either excess carbohydrateaccumulation or decreased N content rather than direct responsesto CO₂. Accumulation of carbohydrates in leaves may lead tothe repression of photosynthetic gene expression and excessstarch seems to hinder CO₂ diffusion. Therefore, the specieswhich have the sink organs for carbohydrate accumulation donot show the suppression of photosynthesis. The suppressionof photosynthesis by CO₂ enrichment is always associated withdecreases in leaf N and Rubisco contents. These decreases arenot due to dilution of N caused by a relative increase in theplant mass but are the result of a decrease in N allocationto leaves at the level of the whole plant, and the decreasein Rubisco content is not selective. Leaf senescence and plantdevelopment are also accelerated by CO₂ enrichment. However,they are independent of each other in some species. Thus, variousresponses to CO₂ observed at the level of a single leaf resultfrom manifold responses at the level of the whole plant grownunder conditions of CO₂ enrichment. (Received July 8, 1999; Accepted August 12, 1999)     

The Effects of Grain Nitrogen and Applied Nitrate on Growth, Photosynthesis and Protein Content of the First Leaf of Barley Cultivars

Five cultivars of barley with widely differing grain nitrogencontents were compared. In the absence of exogenous nitratesupply plants grown from high nitrogen grain showed a more rapidleaf emergence, greater leaf size, especially of the first leaf,higher photosynthetic rate and greater total souble proteinand Fraction 1 protein content of the first leaf, than plantsgrown from low nitrogen grain. However, early supply of nitrateto plants grown from low nitrogen grain enabled these to performas well as those from grain with a high nitrogen content. Regressionanalysis showed that Fraction 1 content of the first leaf isclosely correlated with grain nitrogen which exerts a progressivelygreater effect on content of this protein as application ofexogenous nitrate is delayed. The more rapid photosyntheticrate of plants grown with high nitrogen, and the consequentgreater rate of dry matter accumulation, is attributable mainlyto effects of nitrogen availability on leaf area and much lessto effects on leaf protein.     

Leaf Phosphate Status,Photosynthesis, and Carbon Partitioning in Sugar Beet (IV. Changes with Time Following Increased Supply of Phosphate to Low-Phosphate Plants)

Changes in photosynthesis, carbon partitioning, and growth following resupply of orthophosphate (Pi) to moderately P-deficient plants (low-P) were determined for sugar beets (Beta vulgaris L. cv F58-554H1) cultured hydroponically in growth chambers. One set of plants was supplied with 1.0 mM Pi in half-strength Hoagland solution (control plants), and a second set (low-P plants) was supplied with 0.05 mM Pi. At the end of 2 weeks, the low-P plants were resupplied with 1.0 mM Pi. Low-P plants rapidly accumulated large amounts of Pi, and the photosynthesis rate increased to control values within 4 to 6 h. The rate of photosynthesis appeared to be controlled by ribulose-1,5-bisphosphate (RuBP); low P reduced photosynthesis and RuBP levels, and P resupply increased photosynthesis and RuBP in a manner parallel with time. Low-P treatment reduced adenylate levels substantially but not nicotinamide nucleotides; adenylate levels recovered to control values over 3 to 6 h. With low P, more photosynthate is allocated to non-P carbon compounds (e.g. starch, sucrose) than to sugar phosphates. When P is resupplied, sugar phosphates increase as starch and sucrose pools decrease; this increase in leaf (chloroplast) sugar phosphates was most likely responsible for the increases in RuBP and photosynthesis and may have increased adenylate levels (through enhanced levels of ribose-5-phosphate).     

Effects of Bean Leaf Age on Pathogenicity by Botrytis fabae

Leaves from field bean plants grown out of doors were inoculated with conidia of B. fabae immediately after detaching from stems. The oldest leaves developed more lesions than youngest ones, although they were not chlorotic. On intact plants at high humidities, established lesions on young leaves increased in size at only half the rate of those on old. but still green leaves. Seven days after inoculation a higher proportion of lesions on old leaves bore conidia than those on young leaves, but leaf age had no significant effect on numbers of conidia per mm² of lesion area. Young leaflets from bean plants grown in a controlled environment or in the field challenged with β. cinerea accumulated more phytoalexins than did old ones.