THE RELATION BETWEEN MAXIMUM RATE OF PHOTOSYNTHESIS AND CONCENTRATION OF CHLOROPHYLL

Willstätter and Stoll have studied the rate of photosynthesis as a function of chlorophyll content. However, their experiments fail to reveal any regular relationship between these two quantities, probably because, in order to obtain material differing widely in chlorophyll content, they were obliged to use leaves which were not comparable in other respects. The writer describes a method for varying the chlorophyll content per unit volume of cells of Chlorella vulgaris, maintaining other factors constant. Experiments are described which show that the maximum rate of photosynthesis is a smooth function of the chlorophyll content.     

Relation between pigment content and photosynthetic characteristics in a blue-green algae

1. The blue-green alga Anacystis nidulans was cultured under steady state conditions at 25 and 39°C. and under several different light intensities to give five different types of cells. 2. Cells were submitted to pigment analysis based upon acetone extracts and aqueous extracts obtained by sonic disintegration. The different cell types show a threefold range of chlorophyll content and a fourfold range of phycocyanin content with only minor changes in the chlorophyll/phycocyanin ratio. Cells of highest pigment content were estimated to contain 2.8 per cent chlorophyll a and 24 per cent phycocyanin, the latter on a total chromoproteid basis. 3. Light intensity curves of photosynthesis were obtained for each of the cell types at 25 and at 39°C. The slopes of the light-limited regions of the curves are approximately linear functions of chlorophyll and phycocyanin contents. Maximum light-saturated rates of photosynthesis at 25 and 39° show no simple relation to pigment content.     

Chlorophyll a fluorescence and light-scattering kinetics displayed by leaves during induction of photosynthesis

Light-scattering, which can be taken as an indicator of the transthylakoid proton-gradient, and chlorophyll a fluorescence, have been followed simultaneously during re-illumination of spinach leaves at different energy fluence rates and carbon dioxide concentrations. The slow fluorescence transient (M peak), which has been associated with photosynthetic induction, was observed in air only at the lower fluence rates used. Data are presented that indicate that M peaks in chlorophyll fluorescence kinetics can only be observed if there is also a simultaneous transient in light-scattering and that these transients are observed when the dark period is relatively long, fluence rate relatively low, and CO₂ concentration relatively high.The results are discussed in relation to the varying demands on ATP by carbon assimilation during induction of photosynthesis at different carbon dioxide concentrations and the manner in which these variations influence the quenching of chlorophyll a fluorescence.Abbreviation Chl chlorophyll     

Effects of water stress on the chlorophyll content,nitrogen level and photosynthesis of leaves of two maize genotypes

The dynamics of leaf chlorophyll level, nitrogen content, photosynthesis and stomatal conductance were followed in detail in two cultivars of maize (Zea mays) during a short period of water stress, applied at tasseling, and during the subsequent recovery phase. Plants used in the experiment were grown in sand-nutrient solution culture under field weather conditions. Water stress reduced chlorophyll levels, stomatal conductance and photosynthesis, but the nitrogen content of the leaves was not affected. It is concluded that the stress-induced loss of chlorophyll is not mediated by a lack of nitrogen. Considerable differences were observed between genotypes in the rate of post-stress recovery of chlorophyll level. Recovery, upon rewatering, of stomatal conductance and photosynthesis preceded that of chlorophyll level. Losses of up to 40% of leaf chlorophyll content were insufficient to affect rates of photosynthesis measured at mid-day.     

Leaf photosynthetic light response: a mechanistic model for scaling photosynthesis to leaves and canopies

1. The response of photosynthesis to radiation is an often-studied but poorly understood process, represented empirically in most photosynthesis models. However, in scaling photosynthesis from leaf to canopy, predictions of canopy photosynthesis are very sensitive to parameters describing the response of leaves to Photosynthetic Photon Flux Density (PPFD).
2. In this study, a mechanistic, yet still simple, approach is presented that models the degree of light saturation in leaves explicitly, assuming a heterogeneous environment of PPFD and chlorophyll.
3. Possible mechanisms determining the ratio of chlorophyll to nitrogen are considered, including a direct dependence on PPFD, a mechanism involving the red/far-red ratio of light in the canopy, and an approach based upon maximizing photosynthesis.
4. Comparison of model predictions with two data sets of light, nitrogen and chlorophyll from canopies of Populus and Corylus suggests that the red/far-red mechanism is the most realistic. The data also show that the trees studied do not always optimize their nitrogen partitioning to maximize photosynthetic yield.
5. We then apply the model to the data sets, to predict the shape of light response curves of leaves within canopies and assess the applicability of simple scaling schemes, in which full acclimation of photosynthesis to PPFD justifies the use of big-leaf models. We conclude that, at least for the data used, basic assumptions of such schemes do not hold.     

PHYSIOLOGICAL EFFECTS OF MANGANESE DEFICIENCY RELATED TO AGE IN SOYBEANS (GLYCINE MAX)

Cooper , Eugene E., and Raymond E. Girton . (Purdue U., Lafayette, Ind.) Physiological effects of manganese deficiency related to age in soybeans (Glycine max). Amer. Jour. Bot. 50(2): 105–110. Illus. 1963.—Soybean plants when grown in manganese-deficient silica sand cultures developed typical manganese deficiency symptoms of interveinal chlorosis and necrosis. Physiological effects including depression of photosynthesis, respiration, growth, and relative chlorophyll contents were studied. The depression of photosynthesis was not always proportional to reduced chlorophyll content. This is taken to indicate the importance of manganese in reactions concerned in photosynthesis in addition to chlorophyll formation. Age of leaves related to position on the plant and actual aging of the plants with time sometimes produced different results when related to photosynthetic rates, which mainly decreased with age of plants. Chlorophyll content in young leaves increased with plant age, except for a consistent decrease after leaf maturity. Respiration rates generally decreased with age. For the most part, the effects of aging on photosynthesis, respiration, and chlorophyll contents were the same for soybeans as for other species reported in the literature.     

Identification of Genes Associated with Chlorophyll Accumulation in Flower Petals

Plants have an ability to prevent chlorophyll accumulation, which would mask the bright flower color, in their petals. In contrast, leaves contain substantial amounts of chlorophyll, as it is essential for photosynthesis. The mechanisms of organ-specific chlorophyll accumulation are unknown. To identify factors that determine the chlorophyll content in petals, we compared the expression of genes related to chlorophyll metabolism in different stages of non-green (red and white) petals (very low chlorophyll content), pale-green petals (low chlorophyll content), and leaves (high chlorophyll content) of carnation (Dianthus caryophyllus L.). The expression of many genes encoding chlorophyll biosynthesis enzymes, in particular Mg-chelatase, was lower in non-green petals than in leaves. Non-green petals also showed higher expression of genes involved in chlorophyll degradation, including STAY-GREEN gene and pheophytinase. These data suggest that the absence of chlorophylls in carnation petals may be caused by the low rate of chlorophyll biosynthesis and high rate of degradation. Similar results were obtained by the analysis of Arabidopsis microarray data. In carnation, most genes related to chlorophyll biosynthesis were expressed at similar levels in pale-green petals and leaves, whereas the expression of chlorophyll catabolic genes was higher in pale-green petals than in leaves. Therefore, we hypothesize that the difference in chlorophyll content between non-green and pale-green petals is due to different levels of chlorophyll biosynthesis. Our study provides a basis for future molecular and genetic studies on organ-specific chlorophyll accumulation.     

The effects of excess manganese on photosynthetic rate and concentration of chlorophyll in Triticum aestivum grown in solution culture

Manganese toxicity, which involves a broad array of physiological responses, has been identified as an important factor limiting plant growth on acid soils. In the experiments reported here, we examined the toxic effects of Mn on chlorophyll content, photosynthesis and respiration in two cultivars (Norquay and Columbus) of Triticum aestivum (wheat) which differ in tolerance of Mn. When grown over a range of concentrations of Mn (0–1 000 μ M ), the Mn-tolerant cultivar maintained higher rates of photosynthesis and respiration, and higher concentrations of chlorophyll a and chlorophyll b , than did the Mn-sensitive cultivar, despite greater accumulations of Mn in leaf tissues. After 5 days growth with 1 000 μ M Mn in solution, the photosynthetic rate fell to 25% of control in the sensitive cultivar and to only 75% of control in the tolerant cultivar. The concentration of chlorophyll a fell to 50% of control in the sensitive cultivar, but did not differ from control in the tolerant cultivar. Greater effects were seen on concentrations of chlorophyll b . which fell to 35% and 55% of control in the sensitive and tolerant cultivars, respectively. Rates of photosynthesis decreased in both cultivars as concentrations of chlorophyll decreased; however, the photosynthetic rate per unit chlorophyll remained constant or increased in the tolerant cultivar and decreased in the sensitive cultivar as concentrations of Mn in solution increased. Thus, in the sensitive cv. Columbus, Mn seemed to have a toxic effect on both chlorophyll content and photosynthesis per unit chlorophyll. In the tolerant cv. Norquay, the only clear effect of Mn was a reduction in chlorophyll content, although direct inhibition of photosynthesis could not be discounted.     

Chlorophyll synthesis in chlorella: regulation by degree of light limitation of growth

The degree of light limitation of growth is the primary controlling factor of chlorophyll synthesis during photoautotrophic growth of Chlorella. The chlorophyll content of the cells increases when light is limiting for growth as occurs in dense cultures, or in cultures under low incident light, or when the light is used less efficiently through partial inhibition of photosynthesis by 3-(p-chlorophenyl)-1, 1-dimethylurea. The chlorophyll content decreases when light is not limiting for growth, as occurs in cells in high light intensity and in dilute suspensions. The initial lag in rate of chlorophyll synthesis in a freshly inoculated culture can be attributed to light at first not being growth limiting, and then becoming growth limiting as the cell suspension becomes denser. Continuous culture experiments support the above conclusions by showing that under steady state conditions the chlorophyll content is inversely related to the relative available light.     

Effects of mechanical wounding, current application and heat treatment on chlorophyll fluorescence and pigment composition in tomato plants

Chlorophyll fluorescence quenching parameters and pigment content of wild-type tomato plants ( Lycopersicon esculentum Mill. cv. Moneymaker) were analyzed upon mechanical wounding, current application and heat treatment. Upon all stimuli the non-photochemical quenching increased significantly in local and systemic tissue. Vice versa, the photochemical quenching decreased. Total chlorophyll content remained almost constant, zeaxanthin content increased 5–6-fold, antheraxanthin content rose also, violaxanthin and neoxanthin on the other hand decreased significantly. These data indicate a similarity of processes on the level of photosynthesis and pigment composition that are induced by light-stress and those induced by mechanical wounding, current application and heat treatment, which all lead to an inhibition of photosynthesis.     

Summer algal communities and primary productivity in fish ponds

The paper presents data on primary productivity and phytoplankton communities in new experimental ponds which received the following treatments; ammonium nitrate and triplesuperphosphate, triplesuperphosphate, cracked corn (10% crude protein) and Auburn No. 3 fish feed (36% crude protein). Comparative data on algal communities were also obtained from production ponds which received feeds or fertilizers. Basic ecological data on macro-algae are also presented.

1. All nutrient additions to experimental ponds resulted in higher levels of gross photosynthesis and greater concentrations of chlorophyll a than were found in the control treatments. Fertilization with both nitrogen and phosphorus gave the highest values. Chlorophyll a and gross photosynthesis were higher in ponds receiving high protein content feed (Auburn No. 3) than in ponds to which low protein content feed (corn) was applied.
2. Persistent blooms of blue-green algae occurred in ponds receiving nitrogen and phosphorus fertilization. Phosphorus only fertilization produced blooms of blue-greens, but these blooms did not persist as in the ponds to which nitrogen was also added. Control ponds were dominated by green algae. Blue-green algae were seldom abundant in feed treatments.
3. Production ponds had high level of gross photosynthesis and large concentrations of chlorophyll a.
4. Many of the production ponds which received feed applications developed heavy blooms of blue-green algae.
5. The major species of blue-green algae observed in the present study were Oscillatoria sp., Raphidiopsis curvata, Anacystis nidulans, A. aeruginosa, Spirulina sp., and Anabaena circinalis. Heterocyst bearing forms, which can presumably fix nitrogen, were seldom noted in ponds that received continuous additions of nitrogen from fish feeds.
6. Macro-algae are abundant in many fish ponds. Data illustrating the competition of macro-algae with phytoplankton are presented.

     

磷胁迫对水曲柳幼苗叶绿素合成、光合作用和生物量分配格局的影响

采用沙培试验对水曲柳幼苗磷胁迫与叶绿素合成、光合作用和生物量分配的关系进行了研究．结果表明,在磷胁迫下,水曲柳幼苗叶片内5-氨基乙酰丙酸(ALA)合成速率和胆色素原(PBG)合成酶活性下降,但叶片中叶绿素a、叶绿素b、总叶绿素以及类胡萝卜素含量没有显著变化;水曲柳幼苗叶片和根中N浓度下降。P在各个部位的浓度也相应减少。表明N和P在苗木生理过程中具有协同作用．叶片中N浓度下降引起光系统Ⅱ(PSⅡ)最大光量子效率(Fv／Fm)、PSⅡ的光化学效率(ΦpsⅡ)、电子传递速率(ETR)以及最大光合速率(Amax)下降．磷胁迫对叶绿素合成的影响与Amax下降无关．在磷胁迫下,水曲柳幼苗总生物量下降,且生物量分配倾向于地下部分。而在P充足或过剩条件下,生物量主要分配于地上部分．     

Photosynthesis,photooxidation of chlorophyll and fluorescence of normal and manganese-deficient Chlorella with and without hydrogenase

Summary Growth, chlorophyll content, chloroplast structure, photosynthesis, photooxidation of chlorophyll and fluorescence were studied in normal and manganese-deficient, closely related Chlorella strains with hydrogenase (Chlorella vulgaris f. tertia 211-8k) and without hydrogenase (Chlorella vulgaris 211-8m).Under Mn-deficient conditions, algae with hydrogenase grow slowly for several weeks without becoming chlorotic and without any major change in thylakoid structure, whereas those without hydrogenase show a rapid loss of most of their chlorophyll, followed by a breakdown of thylakoid structure. In algae without hydrogenase, the inhibition of photosynthesis by Mn deficiency is much less pronounced when photosynthesis is determined on a chlorophyll rather than on a cell-volume or dry-weight basis. Thus the little remaining chlorophyll of these algae is photosynthetically much more active than the chlorophyll of the Mn-deficient, non-chlorotic algae with hydrogenase.Photooxidation of chlorophyll under pure O₂ in very strong light is always accelerated by Mn deficiency.Only in algae containing hydrogenase is there under aerobic conditions a pronounced peak of fluorescence at the beginning of illumination. This is indicative of a high degree of reduction of Q, the primary electron acceptor of System II of photosynthesis. The fluorescence peak can be eliminated by either 1 min preillumination with far red (=717 nm) or by treatment in the dark for 1 hr with pure O₂. In algae without hydrogenase, in contrast, fluorescence intensity at the onset of illumination is lower than the steady-state level, indicating an oxidized state of Q. In these algae a high start of fluorescence can only be produced by prolonged anaerobic incubation. These results indicate that even under aerobic conditions algal hydrogenase, or a biochemical system very closely associated with it, has some residual activity which enables it to feed electrons into System II of photosynthesis, thereby keeping Q in a reduced state with correspondingly high fluorescence at the beginning of illumination. In organisms without hydrogenase, in contrast, this can only be achieved by prolonged anaerobiosis. In addition, the presence of hydrogenase seems to protect the chlorophyll against photooxidative destruction when the cells are under Mn deficiency.This paper is dedicated to Professor A. Pirson on the occasion of his 60th birthday.     

Photosynthetic Capacity and Quantum Requirement of Three Secondary Mutants of Scenedesmus obliquus with Deletions in Carotenoid Biosynthesis*

Photosynthetic capacity and quantum requirement of photosynthesis were evaluated in three secondary mutants of Scenedesmus obliquus possessing specific blocks in carotenoid biosynthesis. These were derived from the mutant, C-2 A', a strain which develops chlorophyll only in the light. All strains were capable of a normal and rapid production of chlorophyll and the development of a functional photosynthetic apparatus when exposed to light. LHC-II levels in the mutants lacking the β,ε-carotenoids (strains C-2 A'-34,1 and C-2 A'-67, 3b) were considerably reduced in fully greened cells but normal in the β,β-epoxycarotenoid deficient-only strain (C-2 A'-67,1) (Bishop, 1996). The maximum photosynthetic capacity and the quantum efficiency (at λ=682nm) of photosynthesis of cells of C-2A'-34,1 and C-2 A'-67,1 showed no significant changes when compared to the original parent strain, C-2 A'. However, the quantum requirement of the strain lacking both types of carotenoids, C-2 A'-67,3b, was slightly increased. It is of particular interest that both strains lacking the β,ε-carotenoids also show a preferential loss of the oligomeric form of the LHC and its associated chlorophylls. The potential requirement of the different carotenoids in maintenance of photosynthetic efficiency in Scenedesmus is discussed.     

古尔班通古特沙漠白梭梭枝干光合及其影响因素

植物枝干光合（P_g）固定其自身呼吸所释放的CO₂,有效减少植物向大气的CO₂排放量。以古尔班通古特沙漠优势木本植物白梭梭（Haloxylon persicum）为研究对象,利用LI-COR 6400便携式光合仪与特制光合叶室（P-Chamber）相结合,观测白梭梭叶片、不同径级枝干的光响应及光合日变化特征;同时监测环境因子（大气温湿度、光合有效辐射、土壤温度及含水量等）与叶片/枝干性状指标（叶绿素含量、含水量、干物质含量、碳/氮含量等）,揭示叶片/枝干光合的主要影响因子;采用破坏性取样,量化个体水平上叶片与枝干的总表面积,阐明枝干光合对植株个体碳平衡的贡献。研究结果显示：（1）白梭梭叶片叶绿素含量是枝干叶绿素含量的12-16倍,各径级枝干叶绿素含量差异不显著;（2）枝干光饱和点低于叶片,枝干不同径级（由粗至细）,暗呼吸速率和枝干光合逐渐减小;（3）光合有效辐射、土壤含水量和空气温湿度是影响叶片光合的主要因子,对枝干光合无显著影响;（4）枝干光合可以固定其自身呼吸产生CO₂的73%,最高可达90%,枝干光合固定CO₂约占个体水平固碳量的15.4%。研究结果表明,忽视枝干光合的贡献来预测未来气候变化背景下荒漠生态系统碳过程,可能存在根本性缺陷,并且在估算枝干呼吸时,需要考虑枝干是否存在光合作用,以提高枝干呼吸的准确性。     

Causes of reduced leaf‐level photosynthesis during strong El Niño drought in a Central Amazon forest

Sustained drought and concomitant high temperature may reduce photosynthesis and cause tree mortality. Possible causes of reduced photosynthesis include stomatal closure and biochemical inhibition, but their relative roles are unknown in Amazon trees during strong drought events. We assessed the effects of the recent (2015) strong El Niño drought on leaf‐level photosynthesis of Central Amazon trees via these two mechanisms. Through four seasons of 2015, we measured leaf gas exchange, chlorophyll a fluorescence parameters, chlorophyll concentration, and nutrient content in leaves of 57 upper canopy and understory trees of a lowland terra firme forest on well‐drained infertile oxisol. Photosynthesis decreased 28% in the upper canopy and 17% in understory trees during the extreme dry season of 2015, relative to other 2015 seasons and was also lower than the climatically normal dry season of the following non‐El Niño year. Photosynthesis reduction under extreme drought and high temperature in the 2015 dry season was related only to stomatal closure in both upper canopy and understory trees, and not to chlorophyll a fluorescence parameters, chlorophyll, or leaf nutrient concentration. The distinction is important because stomatal closure is a transient regulatory response that can reverse when water becomes available, whereas the other responses reflect more permanent changes or damage to the photosynthetic apparatus. Photosynthesis decrease due to stomatal closure during the 2015 extreme dry season was followed 2 months later by an increase in photosynthesis as rains returned, indicating a margin of resilience to one‐off extreme climatic events in Amazonian forests.     

Photosynthesis responses to ozone in young trees of three species with different sensitivities, in a 2-year open-top chamber experiment (Curno, Italy)

This paper reports the findings of an open-top chamber experiment carried out in northern Italy (Forest nursery at Curno), during the 2004 and 2005 growth seasons, on Fagus sylvatica and Quercus robur seedlings and on Populus nigra cuttings, in order to test their photosynthesis response to ambient ozone. The experimental protocols were non-filtered air (NF), charcoal-filtered air (CF) and open air (OA). Tests performed included morphological features of leaves; development of foliar symptoms; chlorophyll content, determined by non-destructive means; chlorophyll fluorescence (direct fluorescence and JIP test) and gas exchanges and net photosynthesis (P_N). Main findings were as follows: (1) symptoms occurred early and were extensive in P. nigra , and they occurred later in F. sylvatica , whereas early degeneration of chlorophyll occurred in late summer in Q. robur ; (2) in conditions of ozone exposure, the three species all presented a decline in photosynthesis efficiency and a decrease in P_N, regardless of the symptomatology they displayed; (3) leaf traits are predictors of species-specific sensitivity to ozone—the high density of Q. robur foliar tissues prevents this species from developing visible symptoms and reduces the extent of physiological responses and (4) physiological responses varied from year to year in the same species—responses were lower in the second year of the experiment, when plants had become better acclimatized to plot conditions.     

The effect of Albugo Candida (white blister rust) on the photosynthetic and carbohydrate metabolism of leaves of Arabidopsis thaliana

Albugo candida (pers.) O. Kuntze (white blister rust) is a biotrophic fungus which infects cruciferous plants including Arabidopsis thaliana (L) Heynh. We report the effect of this pathogen on the photosynthetic and carbohydrate metabolism of A. thaliana. As infection progressed A. Candida caused a reduction in the rate of photosynthesis when measured at either ambient or saturating concentrations of CO₂. These data suggested that both chlorophyll and Rubisco were lost from regions of infected leaves, and measurements of chlorophyll, Rubisco content and activity supported these observations. The reduction in the rate of photosynthesis was not caused by closure of stomata as transpiration was unaffected by the disease. Infected leaves accumulated both soluble carbohydrates and starch. The activities of sucrose-phosphate synthase, sucrose synthase and ADP glucose pyrophosphorylase did not change in response to infection. However, the activities of both the wall-bound and soluble acid invertases were higher in infected leaves than in controls; a new soluble invertase isoform with a pl of 5-1 appeared in infected leaves. The possible origin of the increase in wall-bound and soluble invertase activities and its effect on the carbohydrate and photosynthetic metabolism of the leaf are discussed.     

Stomatal and Chlorophyll Distribution of Cox's Orange Pippin Apple Flowers Relative to Other Cluster Parts

Surface area, chlorophyll concentration and stomatal size anddensity were measured for all green components of floral clusterson Cox's Orange Pippin apple trees to assess their relativepotential for photosynthesis. Before flower opening, sepalsand receptacles had chlorophyll concentrations similar to thosefound in the laminae of rosette leaves. But, whereas the concentrationin rosette leaves continued to increased until fruit set, thosein the sepals and receptacles decreased. Sepals had a greater mature stomatal density than leaf laminaeshortly after bud burst; although both densities increased,that on the laminae later exceeded that on sepals. Flowers contributed significantly to the green surface are (40%),chlorophyll content (30%) and mature stomatal number (25%) offloral clusters at the green cluster stage, but these contributionsthen decreased. Cox apple flowers appear to have the potentialfor contributing significantly to their carbohydrate requirementat a time when the rosette leaves are not thought to be exportersof photosynthate.Copyright 1993, 1999 Academic Press Malus pumila, apples flowers, chlorophyll, stomata, surface area, flower photosynthesis