Impact of elevated ozone on chlorophyll a fluorescence in field-grown oat (Avena sativa)

Oat (Avena sativa) plants were grown in the field near the urban area of Valencia, Eastern Spain. The data on air quality showed that ozone was the main phytotoxic pollutant present in ambient air reaching a 7-h mean of 46 nl l(-1) and a maximum hourly peak of 322 nl l(-1). The effect of ambient ozone on PSII activity was examined by measurements of chlorophyll (Chl) a fluorescence. In leaves with visible symptoms, the function of PSII was changed at high actinic irradiances. Nonphotochemical quenching (NPQ) was higher and quantum efficiency of PSII (Phi(PSII)), photochemical quenching (q(p)), quantum efficiency of excitation capture and PSII electron flow (F(v)'/F(m)') were lower. An enhanced susceptibility to photoinhibition was observed for symptom-exhibiting leaves compared to leaves that remain free of visible symptoms. Both the lowering of photosynthesis efficiency and the increased sensitivity to photoinhibition probably contribute to reduced crop yield in the field, to different extents, depending on growth conditions. To our knowledge, this is the first report that demonstrates that quantum efficiency of exciton trapping in PSII is associated with foliar injury in oat leaves in response to ambient concentration of ozone.     

Nonstomatal inhibition of photosynthesis by water stress. Reduction in photosynthesis at high transpiration rate without stomatal closure in field-grown tomato

Large underestimates of the limitation to photosynthesis imposed by stomata can occur because of an error in the standard method of calculating average substomatal pressures of carbon dioxide when heterogeneity of those pressures occurs across a leaf surface. Most gas exchange data supposedly indicating nonstomatal inhibition of photosynthesis by water stress could have this error. However, if no stomatal closure occurs, any reduction in photosynthesis must be due to nonstomatal inhibition of photosynthesis. Net carbon dioxide exchange rates and conductances to water vapor were measured under field conditions in upper canopy leaves of tomato plants during two summers in Beltsville, Maryland, USA. Comparisons were made near midday at high irradiance between leaflets in air with the ambient water vapor content and in air with a higher water content. The higher water content, which lowered the leaf to air water vapor pressure difference (VPD), was imposed either one half hour or several hours before measurements of gas exchange. In both seasons, and irrespective of the timing of the imposition of different VPDs, net photosynthesis increased 60% after decreasing the VPD from 3 to 1 kPa. There were no differences in leaf conductance between leaves at different VPDs, thus transpiration rates were threefold higher at 3 than at 1 kPa VPD. It is concluded that nonstomatal inhibition of photosynthesis did occur in these leaves at high transpiration rate.     

Modifications in photosystem II photochemistry in senescent leaves of maize plants

Analyses of CO2 exchange and chlorophyll fluorescence were carried out to assess photosynthetic performance during senescence of maize leaves. Senescent leaves displayed a significant decrease in CO2 assimilatory capacity accompanied by a decrease in stomatal conductance and an increase in intercellular CO2 concentration. The analyses of fluorescence quenching under steady-state photosynthesis showed that senescence resulted in an increase in non-photochemical quenching and a decrease in photo-chemical quenching. It also resulted in a decrease in the efficiency of excitation energy capture by open PSII reaction centres and the quantum yield of PSII electron transport, but had very little effect on the maximal efficiency of PSII photochemistry. The results determined from the fast fluorescence induction kinetics indicated an increase in the proportion of QB-non-reducing PSII reaction centres and a decrease in the rate of QA reduction in senescent leaves. Theoretical analyses of fluorescence parameters under steady-state photosynthesis suggest that the increase in the non-photochemical quenching was due to an increase in the rate constant to thermal dissipation of excitation energy by PSII and that the decrease in the quantum yield of PSII electron transport was associated with a decrease in the rate constant of PSII photochemistry. Based on these results, it is suggested that the decrease in the quantum yield of PSII electron transport in senescent leaves was down-regulated by an increase in the proportion of QB-non-reducing PSII reaction centres and in the non-photochemical quenching. The photosynthetic electron transport would thus match the decreased demand for ATP and NADPH in carbon assimilation which was inhibited significantly in senescent leaves.Key words: Chlorophyll fluorescence, gas exchange, maize (Zea mays L.), photochemical and non-photochemical quenching, photosystem II photochemistry.      

Influence of climatic and nutritional factors on yeast population dynamics in the phyllosphere of wheat

The role of saprophytic phyllosphere yeasts in removing aphid honeydew and other nutrients from wheat leaves was evaluated in growth cabinet experiments at different temperatures and relative humidities. Population densities of both pink and white yeasts (Sporobolomyces roseus and Cryptococcus laurentii, respectively) increased between 12 and 24°C, if nutrients were supplied. White yeast numbers increased rapidly at a constant vapor pressure deficit (VPD) of 0.10 kPa and alternating VPDs of 0.10 and 0.61 kPa (each 12 hours per day) but decreased at a constant VPD of 0.61 kPa. In growth cabinet experiments with aphids on wheat plants, the amount of aphid honeydew on the leaves was lower when yeast population densities were high. Addition of amino acids to leaves with honeydew had no effect on yeast population density or the rate of honeydew consumption. This indicated that low concentrations of amino acids in aphid honeydew are not a limiting factor for honeydew consumption by the yeasts. The naturally occurring saprophytes efficiently removed fructose, sucrose, and melezitose from the phyllosphere of field-grown wheat plants.     

Photosynthetic responses to vapour pressure deficit in temperate and tropical evergreen rainforest trees of Australia

Rainforests occur in high precipitation areas of eastern Australia, along a gradient in seasonality of precipitation, ranging from a summer dry season in the temperate south to a winter dry season in the tropical north. The response of net photosynthesis to increasing vapour pressure deficit (VPD) was measured in a range of Australian rainforest trees from different latitudes to investigate possible differences in their response to atmospheric drought. Plants were grown in glasshouses under ambient or low VPD to determine the effect of growth VPD on the photosynthetic response. Temperate species, which experience low summer precipitation, were found to maintain maximum net photosynthesis over the measurement range of VPD (0.5–1.9 kPa). In contrast, the tropical species from climates with high summer precipitation showed large reductions in net photosynthesis with increasing VPD. Temperate species showed higher intrinsic water-use efficiencies under low VPD than the tropical species, whereas their efficiencies were similar under high VPD. Growing plants under a low VPD had little effect on either the photosynthetic response to VPD or the intrinsic water-use efficiency of the species. These different responses of gas exchange to VPD shown by the tropical and temperate rainforest species may reflect different strategies to maximise productivity in their respective climates.     

The photosynthetic properties of rice leaves treated with low temperature and high irradiance

Photosynthetic characteristics in rice (Oryza sativa L.) leaves were examined after treatment with low temperature (15 degrees C) and high irradiance (1,500 micromol quanta m(-2) s(-1)). Decreases in quantum efficiencies in PSII (PhiPSII) and PSI (PhiPSI) and in the rate of CO2 assimilation were observed with a decrease in the maximal quantum efficiency of PSII (F(v)/F(m)) by simultaneous measurements of Chl fluorescence, P700+ absorbance and gas exchange. The decreases in PhiPSII were most highly correlated with those in CO2 assimilation. Although the initial (the activity immediately measured upon extraction) and total (the activity following pre-incubation with CO2 and Mg2+) activities of ribulose-1,5-bisphosphate (RuBP) carboxylase/oxygenase (Rubisco) decreased slightly, the maximal activity (the activity following treatment with SO4(2-)) of Rubisco remained almost constant. These results indicate that the decrease in CO2 assimilation rate with the decreasing F(v)/F(m) was not caused by a decrease in Rubisco activity but rather by a decrease in RuBP regeneration capacity which resulted from the decrease in the rate of the linear electron transport. On the other hand, the decrease in PhiPSI was very small and the ratio of PhiPSI to PhiPSII increased. The de-epoxidation state of xanthophyll cycle pigments also increased. Thus, the cyclic electron transport around PSI occurred in photoinhibited leaves.     

CaCl_2对UV-B辐射下菠菜叶片光合特性的影响

以"丹麦旺盛菠菜"为材料,通过UV-B和CaCl2复合处理,测定光合色素含量、Hill反应活力、叶绿素荧光、MDA含量和抗氧化酶活性等参数,探讨了CaCl2对UV-B辐射下菠菜叶片电子传递链和光合膜酶保护系统的影响。结果表明,UV-B处理下,光合色素含量、chl/car、类囊体膜上PSII潜在活性(Fv/Fo)、光化学淬灭系数(qP)、非光化学淬灭系数(qN)、PSII光量子产量(ΦPSⅡ)、原初光能转化效率(Fv/Fm),以及Hill反应活力等降低,chla/chlb和MDA含量升高;喷洒CaCl2可不同程度缓解UV-B的伤害。不同处理下,POD、SOD和CAT活性的变化呈现补偿效应。UV-B强度与菠菜叶片PSII功能受损程度呈正相关,CaCl2则主要通过提高chlb含量、类囊体膜上的光量子产量和POD活性,以缓解伤害。重度UV-B辐射下,CaCl2使chlb含量显著提高可能是导致PSII捕光效率提高的重要因素。     

The Effect of Drought and Vapour Pressure Deficit on Gas Exchange of Young Kiwifruit (Actinidia deliciosavar.deliciosa) Vines

To evaluate the effect of drought and vapour pressure deficit (VPD) on stomatal behaviour and gas exchange parameters, young kiwifruit vines (Actinidia deliciosavar.deliciosacv. Hayward) were exposed to alternating periods of drought and drought-relief over two growing seasons. Vines were grown either in the field or in containers. Stomatal conductance of fully-expanded leaves rapidly decreased as pre-dawn leaf water potential was reduced below a threshold value of -0.3MPa. Stomatal conductance reached minimum values of 10–20mmol m^-2s^-1. Transpiration rate was similarly sensitive to changes in leaf water status, whereas more severe drought levels were necessary to affect photosynthesis significantly. Net daily carbon gains were estimated at 4.7 and 2.7gm^-2for irrigated and droughted vines, respectively. Gas exchange parameters recovered to values of irrigated vines within a few hours after relief of stress. Rate of recovery depended on the level of stress reached during the previous drought period. There was a steady decline in stomatal conductance when VPD was increased from 0.8 to 2.5kPa in both irrigated and droughted vines. The VPD at which stomatal conductance reached 50% of maximum values was 2.1–2.2kPa for both treatments. We conclude that stomata were highly sensitive to changes in soil water status and that midday depression of photosynthesis measured in kiwifruit vines was related to water deficits arising in the leaf because of both transpirational losses and to the direct effect of increasing VPD.     

Combined effects of selenium and drought on photosynthesis and mitochondrial respiration in potato.

The possible effects of selenium (Se) foliar spraying and drought were studied for 3 months in potato (Solanum tuberosum L.) cultivar Desiree in Ljubljana, Slovenia. Four combinations of treatments were conducted: well-watered plants with and without Se foliar spraying, and drought exposed plants with and without Se foliar spraying. The following parameters were monitored 2 and 4 weeks after treatments: net photosynthesis, transpiration rate, quantum yield of photosystem II (PSII), and respiratory potential measured by electron transport system activity. After 3 months of treatments, leaf water potential and tuber yield were determined. The content of Se in tubers was measured after harvesting time. Several effects of drought and Se foliar spraying and their combinations were found. Net photosynthesis and respiratory potential were lower in drought exposed plants 4 weeks after treatments. Se induced higher respiratory potential in the leaves 4 weeks after treatments. Higher efficiency of energy conversion in PSII, expressed by a higher effective quantum yield, was observed in Se treated plants 2 weeks after treatments. Foliarly applied Se was efficiently absorbed by plant leaves and transported to the tubers.     

Photosynthesis, non-photochemical pathways and activities of antioxidant enzymes in a resilient evergreen oak under different climatic conditions from a valley-savanna in Southwest China

Plants in the savanna‐valleys in Southwest China are annually exposed to different combinations of multiple stresses from the hot‐rainy, to chill‐dry, and to warm‐dry seasons. This study monitored seasonal changes in photosynthesis and photoprotection in an evergreen oak (Cyclobalanopsis helferiana) from one of these valleys for four years during which usual and abnormal drought occurred. In general, during the study period with decreasing xylem water potential (Ψx), photosynthetic gas exchange, quantum yield of photosystem II (PSII) photochemistry and activities of most of the measured antioxidant enzymes decreased, while activities of the xanthophyll cycle and associated non‐photochemical energy dissipation and glutathione peroxidase (GP) (EC 1.11.1.9) increased. In a fairly severe chill period, high concentration of reactive oxygen species induced high activities of most of the antioxidant enzymes and relatively stronger decrease in gas exchange. In the most severe dry period, even when predawn Ψx decreased down to ?4 MPa, considerable Pn (maximum photosynthetic rate) (4 μmol m^–2 s^–1) was still maintained in midmorning. At this time, most of the antioxidant enzyme activities decreased to the lowest values, whereas the xanthophyll cycle and associated non‐photochemical energy dissipation and GP activities increased to their highest levels. High predawn antheraxanthin and zeaxanthin contents were observed in the severe and very severe drought periods. Superoxide dismutase maintained high and fairly constant activity (1500–1800 U mg^?1 protein) and predawn maximum photochemistry efficiency of PSII was always above 0.8 throughout the whole study period. These results indicated that the photosynthetic apparatus of the oak leaves was highly capable of maintaining its function under the multiple stresses in different seasons in the present valley‐savanna.     

NaCl胁迫增强杂交酸模(Rumex K-1)幼苗叶片光系统Ⅱ的耐热性

NaCl胁迫对杂交酸模幼苗光系统Ⅱ(PS Ⅱ)的最大光化学效率没有影响,但是增强了PS Ⅱ的耐热性.热胁迫条件下,与未经盐胁迫处理的叶片相比,经NaCl 200 mmol/L处理的杂交酸模幼苗叶片,其PS Ⅱ最大光化学效率下降较小,反映OEC受伤程度的指标Fk/Fj上升较小.此外,光化学猝灭系数(qP)、PS Ⅱ反应中心光能捕获效率(Fv1/Fm1)、PS Ⅱ光化学转换效率(ΦPS Ⅱ)的下降以及QB-非还原性反应PS Ⅱ反应中心的相对含量上升程度也较小.探讨了盐胁迫增强杂交酸模幼苗叶片PS Ⅱ耐热性的可能机理.     

Influence of atmospheric vapour pressure deficit on ozone responses of snap bean (Phaseolus vulgaris L.) genotypes

Environmental conditions influence plant responses to ozone (O(3)), but few studies have evaluated individual factors directly. In this study, the effect of O(3) at high and low atmospheric vapour pressure deficit (VPD) was evaluated in two genotypes of snap bean (Phaseolus vulgaris L.) (R123 and S156) used as O(3) bioindicator plants. Plants were grown in outdoor controlled-environment chambers in charcoal-filtered air containing 0 or 60 nl l(-1) O(3) (12 h average) at two VPDs (1.26 and 1.96 kPa) and sampled for biomass, leaf area, daily water loss, and seed yield. VPD clearly influenced O(3) effects. At low VPD, O(3) reduced biomass, leaf area, and seed yield substantially in both genotypes, while at high VPD, O(3) had no significant effect on these components. In clean air, high VPD reduced biomass and yield by similar fractions in both genotypes compared with low VPD. Data suggest that a stomatal response to VPD per se may be lacking in both genotypes and it is hypothesized that the high VPD resulted in unsustainable transpiration and water deficits that resulted in reduced growth and yield. High VPD- and water-stress-induced stomatal responses may have reduced the O(3) flux into the leaves, which contributed to a higher yield compared to the low VPD treatment in both genotypes. At low VPD, transpiration increased in the O(3) treatment relative to the clean air treatment, suggesting that whole-plant conductance was increased by O(3) exposure. Ozone-related biomass reductions at low VPD were proportionally higher in S156 than in R123, indicating that differential O(3) sensitivity of these bioindicator plants remained evident when environmental conditions were conducive for O(3) effects. Assessments of potential O(3) impacts on vegetation should incorporate interacting factors such as VPD.     

In vivo photosynthetic electron transport does not limit photosynthetic capacity in phosphate-deficient sunflower and maize leaves

The effects of extreme phosphate (Pi) deficiency during growth on the contents of adenylates and pyridine nucleotides and the in vivo photochemical activity of photosystem II (PSII) were determined in leaves of Helianthus annuus and Zea mays grown under controlled environmental conditions. Phosphate deficiency decreased the amounts of ATP and ADP per unit leaf area and the adenylate energy charge of leaves. The amounts of oxidized pyridine nucleotides per unit leaf area decreased with Pi deficiency, but not those of reduced pyridine nucleotides. This resulted in an increase in the ratio of reduced to oxidized pyridine nucleotides in Pi-deficient leaves. Analysis of chlorophyll a fluorescence at room temperature showed that Pi deficiency decreased the efficiency of excitation capture by open PSII reaction centres (φ_e), the in vivo quantum yield of PSII photochemistry (φ_PSII) and the photochemical quenching co-efficient (q_P), and increased the non-photochemical quenching co-efficient (q_N) indicating possible photoinhibitory damage to PSII. Supplying Pi to Pi-deficient sunflower leaves reversed the long-term effects of Pi-deficiency on PSII photochemistry. Feeding Pi-sufficient sunflower leaves with mannose or FCCP rapidly produced effects on chlorophyll a fluorescence similar to long-term Pi-deficiency. Our results suggest a direct role of Pi and photophosphorylation on PSII photochemistry in both long-and short-term responses of photosynthetic machinery to Pi deficiency. The relationship between φ_PSII and the apparent quantum yield of CO₂ assimilation determined at varying light intensity and 21 kPa O₂ and 35 Pa CO₂ partial pressures in the ambient air was linear in Pi-sufficient and Pi-deficient leaves of sunflower and maize. Calculations show that there was relatively more PSII activity per mole of CO₂ assimilated by the Pi-deficient leaves. This indicates that in these leaves a greater proportion of photosynthetic electrons transported across PSII was used for processes other than CO₂ reduction. Therefore, we conclude that in vivo photosynthetic electron transport through PSII did not limit photosynthesis in Pi-deficient leaves of sunflower and maize and that the decreased CO₂ assimilation was a consequence of a smaller ATP content and lower energy charge which restricted production of ribulose, 1-5, bisphosphate, the acceptor for CO₂.     

Interactive effects of elevated ozone and springtime frost on growth and physiology of birch (<Emphasis Type="Italic">Betula pendula</Emphasis>) in field conditions

We studied the impact of ozone enrichment and late frost, singly and interactively, on four birch (Betula pendula Roth) families selected from a naturally regenerated birch stand in southeastern Finland. Seedlings were exposed to 1.5× ambient ozone over one and a half growing seasons using free-air ozone enrichment system. Simulated springtime frost was implemented at the beginning of the second study year, 4 weeks after the bud burst. Plants were measured for timing of bud burst, visible ozone injuries, chlorophyll fluorescence, net photosynthesis and concentrations of photosynthetic pigments, as well as for growth and carbon allocation. Frost treatment caused a rapid 60% decline in net photosynthesis. The recovery of net photosynthesis from acute frost treatment was not complete during the subsequent 3 weeks, which led to significant growth reductions, decreased shoot/root ratio and accumulation of excess nitrogen in the leaves. Photosynthetic responses to ozone were very variable and family-specific. Concentrations of photosynthetic pigments were sensitive to both stress factors, while the maximum quantum yield of PSII was unaffected. Ozone exacerbated the effect of frost only on diameter increment. However, ozone and frost affected different seedling characters, e.g., ozone reduced pigments and frost collapsed net photosynthesis, and these effect combined appear to damage birch seedlings more than a single stress situation.     

High resolution imaging of photosynthetic activities of tissues, cells and chloroplasts in leaves

Through imaging of chlorophyll fluorescence, it is possible to produce parameterized fluorescence images that estimate the operating quantum efficiency of photosystem II (PSII) photochemistry and which can be used to reveal heterogeneous patterns of photosynthetic performance within leaves. The operating quantum efficiency of PSII photochemistry is dependent upon the effective absorption cross-section of the light-harvesting system of PSII and the photochemical capacity of PSII. The effective absorption cross-section is decreased by the process of down-regulation, which is widely thought to operate within the pigment matrices of PSII and which results in non-photochemical quenching of chlorophyll fluorescence. The photochemical capacity is non-linearly related to the proportion of PSII centres in the 'open' state and results in photochemical quenching of chlorophyll fluorescence. Examples of heterogeneity of the operating quantum efficiency of PSII photochemistry during the induction of photosynthesis in maize leaves and in the chloroplast populations of stomatal guard cells of a leaf of Tradescantia albifora are presented, together with analyses of the factors determining this heterogeneity. A comparison of the operating quantum efficiency of PSII photochemistry within guard cells and adjacent mesophyll cells of Commelina communis is also made, before and after stomatal closure through a change in ambient humidity.     

The influence of plant water stress on stomatal control of gas exchange at different levels of atmospheric humidity

Summary Leaves of well-watered and mildly water-stressed seedlings of Betula pendula Roth. and Gmelina aroborea L. were subjected to a range of vapour pressure deficits (VPD) between 10 and 24 kPa. The stomatal conductance of birch seedlings decreased as VPD was increased and at least in mildly-stressed seedlings this response seemed to be closely linked to the water status of the air rather than to the bulk water status of the plant. Mild water stressing enhanced the degree of the stomatal humidity-response and resulted in a significant increase in the efficiency of water use at high VPD. Stomata of Gmelina were apparently insensitive to variation in VPD, but were more sensitive to a decrease in bulk leaf water status than were stomata of birch. Water use efficiency of Gmelina seedlings was comparatively high, even when VPD was high and the stomata were fully open.     

Thermostability and photostability of photosystem II of the resurrection plant Haberlea rhodopensis studied by chlorophyll fluorescence

The stability of PSII in leaves of the resurrection plant Haberlea rhodopensis to high temperature and high light intensities was studied by means of chlorophyll fluorescence measurements. The photochemical efficiency of PSII in well-hydrated Haberlea leaves was not significantly influenced by temperatures up to 40 degrees C. Fo reached a maximum at 50 degrees C, which is connected with blocking of electron transport in reaction center II. The intrinsic efficiency of PSII photochemistry, monitored as Fv/Fm was less vulnerable to heat stress than the quantum yield of PSII electron transport under illumination (phiPSII). The reduction of phiPSII values was mainly due to a decrease in the proportion of open PSII centers (qP). Haberlea rhodopensis was very sensitive to photoinhibition. The light intensity of 120 micromol m(-2) s(-1) sharply decreased the quantum yield of PSII photochemistry and it was almost fully inhibited at 350 micromol m(-2) s(-1). As could be expected decreased photochemical efficiency of PSII was accompanied by increased proportion of thermal energy dissipation, which is considered as a protective effect regulating the light energy distribution in PSII. When differentiating between the three components of qN it was evident that the energy-dependent quenching, qE, was prevailing over photoinhibitory quenching, qI, and the quenching related to state 1-state 2 transitions, qT, at all light intensities at 25 degrees C. However, the qE values declined with increasing temperature and light intensities. The qI was higher than qE at 40 degrees C and it was the major part of qN at 45 degrees C, indicating a progressing photoinhibition of the photosynthetic apparatus.     

Modulation of photosynthetic energy conversion efficiency in nature: from seconds to seasons

Modulation of the efficiency with which leaves convert absorbed light to photochemical energy [intrinsic efficiency of open photosystem II (PSII) centers, as the ratio of variable to maximal chlorophyll fluorescence] as well as leaf xanthophyll composition (interconversions of the xanthophyll cycle pigments violaxanthin and zeaxanthin) were characterized throughout single days and nights to entire seasons in plants growing naturally in contrasting light and temperature environments. All pronounced decreases of intrinsic PSII efficiency took place in the presence of zeaxanthin. The reversibility of these PSII efficiency changes varied widely, ranging from reversible-within-seconds (in a vine experiencing multiple sunflecks under a eucalypt canopy) to apparently permanently locked-in for entire seasons (throughout the whole winter in a subalpine conifer forest at 3,000?m). While close association between low intrinsic PSII efficiency and zeaxanthin accumulation was ubiquitous, accompanying features (such as trans-thylakoid pH gradient, thylakoid protein composition, and phosphorylation) differed among contrasting conditions. The strongest and longest-lasting depressions in intrinsic PSII efficiency were seen in the most stress-tolerant species. Evergreens, in particular, showed the most pronounced modulation of PSII efficiency and thermal dissipation, and are therefore suggested as model species for the study of photoprotection. Implications of the responses of field-grown plants in nature for mechanistic models are discussed.     

Control of the Quantum Efficiencies of Photosystems I and II, Electron Flow, and Enzyme Activation following Dark-to-Light Transitions in Pea Leaves: Relationship between NADP/NADPH Ratios and NADP-Malate Dehydrogenase Activation State

The quantum efficiencies of photosystems I and II (PSI and PSII), [NADP]/[NADPH] ratios, and the activities of chloroplastic fructose-1,6-bisphosphatase and NADP-malate dehydrogenase were measured in intact pea (Pisum sativum L.) leaves in air following the transition from darkness to 750 microeinsteins per square meter per second irradiance. PSII efficiency declined from a low value to a minimum within the first 10 to 15 seconds of irradiance, after which it increased progressively to the steady-state value. The resistance of electron flow between the photosystems was high at this time, but it was not the principal factor limiting electron flow. Oxidation of P700 was restricted by acceptor side processes for approximately the first 60 seconds of illumination. Once the acceptor side limitation was relieved, the oxidation state of P700 was used to estimate the quantum efficiency of electron transport by PSI. This was observed to increase progressively with time. The quantum efficiencies of both photosystems increased in parallel, consistent with a predominant role for noncyclic electron transport. Fructose-1,6-bisphosphatase activity increased in an approximately sigmoidal fashion with time of irradiance, paralleling the changes in the quantum efficiencies of the photosystems. In contrast, the activation of NADP-malate dehydrogenase did not show a lag period but increased with time, reaching a maximum value at about 50 seconds of illumination, after which it declined. The NADP pool was not extensively reduced during the first 10 seconds of illumination, but became so subsequently. It remained in the reduced state until about 60 seconds of illumination and then became relatively oxidized. The empirical relationship between NADP-malate dehydrogenase activity and the reduction state of the NADP pool supports the suggestion that NADP-malate dehydrogenase activity is a useful estimate of the reduction state of the stroma.