Photosynthetic photon flux density,carbon dioxide concentration,and vapor pressure deficit effects on photosynthesis in cacao seedlings

Independent short-term effects of photosynthetic photon flux density (PPFD) of 50–400 μmol m⁻² s⁻¹, external CO₂ concentration (C _a) of 85–850 cm³ m⁻³, and vapor pressure deficit (VPD) of 0.9–2.2 kPa on net photosynthetic rate (P _N), stomatal conductance (g _s), leaf internal CO₂ concentration (C _i), and transpiration rates (E) were investigated in three cacao genotypes. In all these genotypes, increasing PPFD from 50 to 400 μmol m⁻² s⁻¹ increased P _N by about 50 %, but further increases in PPFD up to 1 500 μmol m⁻² s⁻¹ had no effect on P _N. Increasing C _a significantly increased P _N and C _i while g _s and E decreased more strongly than in most trees that have been studied. In all genotypes, increasing VPD reduced P _N, but the slight decrease in g _s and the slight increase in C _i with increasing VPD were non-significant. Increasing VPD significantly increased E and this may have caused the reduction in P _N. The unusually small response of g _s to VPD could limit the ability of cacao to grow where VPD is high. There were no significant differences in gas exchange characteristics (g _s, C _i, E) among the three cacao genotypes under any measurement conditions.     

Factors determining the midday depression of photosynthesis in trees under monsoon climate

 Field studies of gas exchange of Populus deltoides, Prosopis juliflora and Acacia auriculiformis showed large diurnal changes in net photosynthesis (A) and stomatal conductance (g_s) during autumn. P. deltoides and P. juliflora undergo pronounced midday depression in A and g_s while A. auriculiformis showed a one-peak response. Several factors indicative of photosynthetic performance were found to be reversibly affected during afternoon decline. These include (i) decrease in initial slope of the CO₂ response curve (carboxylation efficiency), (ii) substantial increase in CO₂ compensation point and (iii) decrease in overall quantum yield of photosystem II. The phenomenon can be duplicated in potted plants by simulating a typical daily pattern of PPFD and VPD. It is found that high VPD induces significant decline in A and g_s at moderate temperature and saturating PPFD (800 μmol m^–2 s^–1) whereas these parameters are only marginally affected at high PPFD and low VPD. Fluorescence data show that the tree species under study have a high capacity for safe dissipation of excessive excitation energy. The activation of photorespiration, as evident from an increase in CO₂ compensation point, maintains constant internal CO₂ concentration (C_i) which may aid in minimizing photoinhibition during stomatal closure at midday. In case of P. deltoides and P. juliflora the stomata seem to be quite sensitive to the changes in humidity whereas this does not appear to be essential in case of A. auriculiformis because of its phyllode structure that endows it with mechanisms for conserving water without undergoing large-scale stomatal changes. Received: 16 October 1997 / Accepted: 5 March 1998     

Modelling photosynthetic photon flux density and maximum potential gross photosynthesis

Irradiance data software developed by the NREL Solar Radiation Laboratory (Simple Model of Atmospheric Radiative Transfer of Sunshine, SMARTS) has been used for modelling photosynthesis. Spectra and total irradiance were expressed in terms of quanta [mol m⁻² s⁻¹, photosynthetic photon flux density, PPFD (400–700 nm)]. Using the SMARTS software it is possible to (1) calculate the solar spectrum for a planar surface for any given solar elevation angle, allowing for the attenuating effects of the atmosphere on extraterrestrial irradiance at each wavelength in the 400–700 nm range and for the thickness of atmosphere the light must pass through during the course of a day, (2) calculate PPFD vs. solar time for any latitude and date and (3) estimate total daily irradiance for any latitude and date and hence calculate the total photon irradiance for a whole year or for a growing season. Models of photosynthetic activity vs. PPFD are discussed. Gross photosynthesis (P _g) vs. photosynthetic photon flux density (PPFD) (P _g vs. I) characteristics of single leaves compared to that of a canopy of leaves are different. It is shown that that the optimum irradiance for a leaf (I_opt) is the half-saturation irradiance for a battery of leaves in series. A C₃ plant, with leaves having an optimum photosynthetic rate at 700 μmol m⁻² s⁻¹ PPFD, was used as a realistic worked example. The model gives good estimates of gross photosynthesis (P _g) for a given date and latitude. Seasonal and annual estimates of P _g can be made. Taking cloudiness into account, the model predicts maximum P _g rates of about 10 g(C) m⁻² d⁻¹, which is close to the maximum reported P _g experimental measurements.     

Midday depression of photosynthesis in <Emphasis Type="Italic">Enkleia malaccensis</Emphasis>, a woody climber in a tropical rainforest

We measured the diurnal changes in net photosynthetic rate (P _N) and stomatal conductance (g _s) of the leaves of a liana, Enkleia malaccensis Griff. (Thymelaeaceae), at the canopy level in the lowland tropical rainforest at Pasoh, Peninsular Malaysia. The measurements were made from a canopy walkway system, 30 m from the ground for 3 d in March 2003. P _N increased with increasing photosynthetically active radiation (PAR) before noon, though P _N was not enhanced by the strong radiation hit in the afternoon. Plotting g _s at saturating PAR (>0.5 mmol m⁻² s⁻¹) against the vapour pressure deficit (VPD) failed to reveal a significant correlation between VPD and g _s, and g _s became very low at VPD >2.5 kPa. The relationship between P _N and g _s was fitted on the same regression line irrespective of measuring day, indicating that this relationship was not influenced by either VPD or leaf temperature (T _L). Therefore, in the liana E. malaccensis, an increase in VPD leads to partial stomatal closure and, subsequently, reductions in P _N and the midday depression of P _N of this plant.     

Non-uniform stomatal closure and the apparent convexity of the photosynthetic photon flux density response curve

The photosynthetic response of leaves to photosynthetic photon flux density (PPFD) may be described by parameters for the rate of dark respiration (R), the initial slope (Ф), the PPFD-saturated rate of net photosynthesis (A_mQ) and the apparent convexity (θ_a). We tested the hypothesis that non-uniformity in stomatal aperture across a leaf results in a clearly lower θ_a because PPFD saturation will occur at different irradiances in different regions of the leaf. A computer model was constructed to simulate the effects of bell-shaped and other distributions of stomatal conductance for CO₂ (g_s) across a model leaf. In the model, Ф and A_mQ decreased by up to 50% whereas θ_a decreased by at most 8%, essentially negating the hypothesis. The relationship between photosynthetic rate and g_s determined the size of the responses of θ_a and Ф to non-uniform g_s. In support of the model, experiments on sunflower leaves showed that Ф and A_mQ decreased by 32 and 52%, respectively, while no measurable change in θ_a occurred, when abscisic acid was used to induce patchiness in g_s. Although not all possible patterns of non-uniform stomatal conductance have been tested, it appears that if large variations in θ_a occur in nature they do not result from non-uniform g_s.     

Different photosynthetic responses of wild and cultivated plants to high irradiance

In addition to other factors, high altitude (HA) environment is characterized by high photosynthetic photon flux density (PPFD). Photosynthetic characteristics of wild and cultivated plants were studied at different irradiances at Losar, India (altitude 4 200 m). Wild plants were tolerant to high PPFDs. Slopes of curve between net photosynthetic rate (P _N) and intercellular CO₂ concentration (C _i) or stomatal conductance (g _s) increased with increase in irradiance suggesting insensitivity or tolerance of these plants to higher PPFD. Cultivated plants, however, were sensitive to higher PPFD, their slopes of curves between P _N and C _i or g _s decreased with increased PPFD. Tolerance or insensitivity to higher PPFD was an important parameter affecting plant performance at HA.     

Climate-ameliorating measures influence photosynthetic gas exchange of apple leaves

Sunburn has become one of the major threats to apple fruit production in South Africa and other countries with Mediterranean climate. Some climate‐ameliorating measures have been developed to control sunburn in apples. Effects of the climate‐ameliorating measures, viz. evaporative cooling, Surround^® WP and shade net, on leaf gas exchange of a 5‐year‐old orchard of ‘Cripps’ Pink’ apple were investigated during hot summer days in Stellenbosch, South Africa. Evaporative cooling increased net photosynthetic rate (A) and stomatal conductance (g_s) because of its lowering of leaf temperature and leaf‐to‐air vapour pressure difference (VPD). Shade net also reduced leaf temperature because of reduction in photosynthetic photon flux density (PPFD). Quantum efficiency of photosynthesis was increased under shade net to compensate for reduced PPFD. Shade net also reduced transpiration rate more than A, resulting in increased midday water‐use efficiency. The diurnal trends of A and g_s in the Surround WP and control treatments were similar, indicating limited ameliorative impact of Surround WP. Furthermore, Surround WP typically reduced maximum rate of carboxylation and the light‐saturated rate of electron transport. In all treatments, A decreased by 70% when leaf temperature increased from 35°C to 40°C. In conclusion, all treatments affected leaf photosynthetic gas exchange. Evaporative cooling enhanced leaf A and g_s because of distinct ameliorative effects on leaf temperature and VPD. Shade net reduced leaf temperature with no consistent effects on leaf gas exchange attributes. Surround WP had limited or no impact on leaf temperature and negatively affected leaf gas exchange attributes.     

Low temperature stress modifies the photochemical efficiency of a tropical tree species <Emphasis Type="Italic">Hevea brasiliensis</Emphasis>: effects of varying concentration of CO<Subscript>2</Subscript> and photon flux density

Two clones of Hevea brasiliensis (RRII 105 and PB 235) were grown for one year in two distinct agroclimatic locations (warmer and colder, W and C) in peninsular India. We simultaneously measured gas exchange and chlorophyll (Chl) fluorescence on fully mature intact leaves at different photosynthetic photon flux densities (PPFDs) and ambient CO₂ concentrations (C _a) and at constant ambient O₂ concentration (21 %). Net photosynthetic rate (P _N), apparent quantum yield for CO₂ assimilation (Φ_c), in vivo carboxylation efficiency (CE), and photosystem 2 quantum yield (Φ_PS2) were low in plants grown in C climate and these reductions were more predominant in RRII 105 than in PB 235 which was also reflected in their growth. We estimated in these clones the partitioning of photosynthetic electrons between CO₂ reduction (J_A) and processes other than CO₂ reduction (J^*) at low and high PPFDs and C _a. At high C _a (700 µmol mol⁻¹) most of the photosynthetic electrons were used for CO₂ assimilation and negligible amount went for other processes when PPFD was low (200–300 µmol m⁻² s⁻¹) both in the C and W climates. But at high PPFD (900-1 100 µmol m⁻² s⁻¹), J^* was appreciably high even at a high C _a. Hence at normal ambient C _a and high irradiance, electrons can be generated in the photosynthetic apparatus far in excess of what can be safely utilised for photosynthetic CO₂ reduction. However, at high C _a there was increased diversion of electrons to photosynthetic CO₂ reduction which resulted in improved photosynthetic parameters even in plants grown in C climate.     

Control of Photosynthesis and Stomatal Conductance in Ricinus communis L. (Castor Bean) by Leaf to Air Vapor Pressure Deficit

Castor bean (Ricinus communis L.) has a high photosynthetic capacity under high humidity and a pronounced sensitivity of photosynthesis to high water vapor pressure deficit (VPD). The sensitivity of photosynthesis to varying VPD was analyzed by measuring CO₂ assimilation, stomatal conductance (g_s), quantum yield of photosystem II (_II), and nonphotochemical quenching of chlorophyll fluorescence (q_N) under different VPD. Under both medium (1000) and high (1800 micromoles quanta per square meter per second) light intensities, CO₂ assimilation decreased as the VPD between the leaf and the air around the leaf increased. The g_s initially dropped rapidly with increasing VPD and then showed a slower decrease above a VPD of 10 to 20 millibars. Over a temperature range from 20 to 40°C, CO₂ assimilation and g_s were inhibited by high VPD (20 millibars). However, the rate of transpiration increased with increasing temperature at either low or high VPD due to an increase in g_s. The relative inhibition of photosynthesis under photorespiring (atmospheric levels of CO₂ and O₂) versus nonphotorespiring (700 microbars CO₂ and 2% O₂) conditions was greater under high VPD (30 millibars) than under low VPD (3 millibars). Also, with increasing light intensity the relative inhibition of photosynthesis by O₂ increased under high VPD, but decreased under low VPD. The effect of high VPD on photosynthesis under various conditions could not be totally accounted for by the decrease in the intercellular CO₂ in the leaf (C_i) where C_i was estimated from gas exchange measurements. However, estimates of C_i from measurements of _II and q_N suggest that the decrease in photosynthesis and increase in photorespiration under high VPD can be totally accounted for by stomatal closure and a decrease in C_i. The results also suggest that nonuniform closure of stomata may occur in well-watered plants under high VPD, causing overestimates in the calculation of C_i from gas exchange measurements. Under low VPD, 30°C, high light, and saturating CO₂, castor bean (C₃ tropical shrub) has a rate of photosynthesis (61 micromoles CO₂ per square meter per second) that is about 50% higher than that of tobacco (C₃) or maize (C₄) under the same conditions. The chlorophyll content, total soluble protein, and ribulose-1,5-bisphosphate carboxylase/oxygenase level on a leaf area basis were much higher in castor bean than in maize or tobacco, which accounts for its high rates of photosynthesis under low VPD.     

Stomatal and non-stomatal limitations to photosynthesis in field-grown grapevine cultivars

Diurnal changes of photosynthesis in the leaves of grapevine (Vitis vinifera × V. labrusca) cultivars Campbell Early and Kyoho grown in the field were compared with respect to gas exchanges and actual quantum yield of photosystem 2 (Φ_PS2) in late May. Net photosynthetic rate (P_N) of the two cultivars rapidly increased in the morning, saturated at photosynthetic photon flux density (PPFD) from 1200 to 1500 μmol m⁻² s⁻¹ between 10:00 and 12:00 and slowly decreased after midday. Maximum P_N was 13.7 and 12.5 μmol m⁻² s⁻¹ in Campbell Early and Kyoho, respectively. The stomatal conductance (g_s) and transpiration rate changed in parallel with P_N, indicating that P_N was greatly affected by g_s. However, the decrease in P_N after midday under saturating PPFD was also associated with the observed depression of Φ_PS2 at high PPFD. The substantial increase in the leaf to air vapour pressure deficit after midday might also contribute to decline of g_s and P_N.     

Photosynthetic characteristics of <Emphasis Type="Italic">Hordeum,Triticum, Rumex</Emphasis>, and <Emphasis Type="Italic">Trifolium</Emphasis> species at contrasting altitudes

Photosynthetic characteristics were compared between plants of low altitude (LA) grown at LA (Palampur; 1 300 m) and at high altitude, HA (Kibber; 4 200 m), and plants naturally occurring at different altitudes (Palampur, 1 300 m; Palchan, 2 250 m; and Marhi, 3 250 m). Net photosynthetic rate (P _N) was not significantly different between altitudes. However, the slopes of the curve relating P _N to intercellular CO₂ concentration (C _i) were higher in plants at Palchan, Marhi, and Kibber compared to those at Palampur, indicating that plants had higher efficiency of carbon uptake (the initial slope of P _N/C _i curve is an indication) at HA. They had also higher stomatal conductance (g _s), transpiration rate, and lower water use efficiency at HA. g _s was insensitive to photosynthetic photon flux density (PPFD) for plants naturally occurring at Palampur, Palchan, and Marhi, whereas plants from LA grown at Palampur and Kibber responded linearly to increasing PPFD. Insensitivity of g _s to PPFD could be one of the adaptive features allowing wider altitudinal distribution of the plants.This research is supported by the Department of Biotechnology (DBT), Government of India vide grant number BT/PR/502/AGR/08/39/966-VI.     

Photosynthesis of individual field-grown cotton leaves during ontogeny

Photosynthetic characteristics of field-grown cotton (Gossypium hirsutum L.) leaves were determined at several insertion levels within the canopy during the growing season. Single-leaf measurements of net photosynthesis (Pn), stomatal conductance to CO₂ (g_s·CO₂), substomatal CO₂, leaf area expansion, leaf nitrogen, and light intensity (PPFD) were recorded for undisturbed leaves within the crop canopy at 3–4 day intervals during the development of all leaves at main-stem nodes 8, 10, and 12. Patterns of Pn during leaf ontogeny exhibited three distinct phases; a rapid increase to maximum at 16–20 days after leaf unfolding, a relatively short plateau, and a period of linear decline to negligible Pn at 60–65 days. Analysis of the parameters which contributed to the rise and fall pattern of Pn with leaf age indicated the primary involvement of leaf area expansion, leaf nitrogen, PPFD, and g_s·CO₂ in this process. The response of Pn and g_s·CO₂ to incident PPFD conditions during canopy development was highly age dependent. For leaves less than 16 days old, the patterns of Pn and g_s·CO₂ were largely controlled by non-PPFD factors, while for older leaves Pn and g_s·CO₂ were more closely coupled to PPFD-mediated processes. Maximum values of Pn were not significantly different for any of the leaves monitored in this study, however, those leaves at main-stem node 8 did possess a significantly diminished photosynthetic capacity with age compared to upper canopy leaves. This accelerated decline in Pn could not be explained by age-related variations in g_s·CO₂ since all leaves showed similar changes in g_s·CO₂ with leaf age.Abbreviations g_s·CO₂ stomatal conductance to CO₂ - Pn net photosynthesis - PPFD photosynthetic photon flux density     

Photosynthetic light and carbon dioxide response of the invasive tree, Vochysia divergens Pohl, to experimental flooding and shading

Vochysia divergens Pohl is considered to be a flood-adapted, light-demanding pioneer species that has been invading grasslands of the Brazilian Pantanal. In these areas, a successful invasion requires an ability to tolerate physiologically wide fluctuations in surface hydrology and shading induced by a dense cover of grasses and other vegetation. We evaluated how flooding and shading affected the photosynthetic performance of V. divergens saplings by measuring light-saturated gas exchange (net photosynthetic rate, P _N; stomatal conductance, g _s), and intercellular CO₂ (P _N/C _i) and photosynthetic photon flux density (P _N/PPFD) response curves over a 61-d field experiment. Shading and flooding reduced significantly light-saturated P _N and g _s and affected multiple aspects of the leaf gas exchange response of V. divergens to variations in PPFD and CO₂. Flooding influenced the physiology of this species more than shading. Given the success of V. divergens at invading and expanding in seasonally flooded areas of the Pantanal, the results were surprising and highlighted the physiological ability of this species to tolerate suboptimal conditions. However, the consistently higher light-saturated P _N and g _s under nonflooded conditions suggested that the invasive success of V. divergens might not be related to its physiological potential during flooding, but to situations, when flooding recedes during the dry season and soil water availability is adequate.     

Influence of irradiance on photosynthesis and PSII photochemical efficiency in maize during short-term exposure at high CO<Subscript>2</Subscript> concentration

This work aimed to evaluate if gas exchange and PSII photochemical activity in maize are affected by different irradiance levels during short-term exposure to elevated CO₂. For this purpose gas exchange and chlorophyll a fluorescence were measured on maize plants grown at ambient CO₂ concentration (control CO₂) and exposed for 4 h to short-term treatments at 800 μmol(CO₂) mol⁻¹ (high CO₂) at a photosynthetic photon flux density (PPFD) of either 1,000 μmol m⁻² s⁻¹ (control light) or 1,900 μmol m⁻² s⁻¹ (high light). At control light, high-CO₂ leaves showed a significant decrease of net photosynthetic rate (P _N) and a rise in the ratio of intercellular to ambient CO₂ concentration (C _i/C _a) and water-use efficiency (WUE) compared to control CO₂ leaves. No difference between CO₂ concentrations for PSII effective photochemistry (Φ_PSII), photochemical quenching (q_p) and nonphotochemical quenching (NPQ) was detected. Under high light, high-CO₂ leaves did not differ in P _N, C _i/C _a, Φ_PSII and NPQ, but showed an increase of WUE. These results suggest that at control light photosynthetic apparatus is negatively affected by high CO₂ concentration in terms of carbon gain by limitations in photosynthetic dark reaction rather than in photochemistry. At high light, the elevated CO₂ concentration did not promote an increase of photosynthesis and photochemistry but only an improvement of water balance due to increased WUE.     

Analysis of gas exchange in seedlings of Acer saccharum: integration of field and laboratory studies

In the field, photosynthesis of Acer saccharum seedlings was rarely light saturated, even though light saturation occurs at about 100 mol quanta m^-2 s^-1 photosynthetic photon flux density (PPFD). PPFD during more than 75% of the daylight period was 50 mol m^-2 s^-1 or less. At these low PPFD's there is a marked interaction of PPFD with the initial slope (CE) of the CO₂ response. At PPFD-saturation CE was 0.018 mol m^-2 s^-1/(l/l). The apparent quantum efficiency (incident PPFD) at saturating CO₂ was 0.05–0.08 mol/mol. and PPFD-saturated CO₂ exchange was 6–8 mol m^-2 s^-1. The ratio of internal CO₂ concentration to external (C _i /C _a ) was 0.7 to 0.8 except during sunflecks when it decreased to 0.5. The decrease in C _i /C _a during sunflecks was the result of the slow response of stomates to increased PPFD compared to the response of net photosynthesis. An empirical model, which included the above parameters was used to simulate the measured CO₂ exchange rate for portions of two days. Parameter values for the model were determined in experiments separate from the daily time courses being sumulated. Analysis of the field data, partly through the use of simulations, indicate that the elimination of sunflecks would reduce net carbon gain by 5–10%.List of symbols A measured photosynthetic rate under any set of conditions (mol m^-2 s^-1) - A _m (atm) measured photosynthetic rate at saturating PPFD, 350 l/l CO₂ and 21% (v/v) O₂ (mol m^-2 s^-1) - C constant in equation of Smith (1937, 1938) - C _a CO₂ concentration in the air (l/l) - C _i CO₂ concentration in the intercellular air space (l/l) - C _i ^/* C _i corrected for CO₂ compensation point, i.e., C _i -I ^*, (l/l) - CE initial slope of the CO₂ response of photosynthesis (mol m^-2 s^-1/(l/l)) - CEM CE at PPFD saturation - E transpiration rate (mmol m^-2 s^-1) - F predicted photosynthetic rate (mol m^-2 s^-1) - G leaf conductance to H₂O (mol m^-2 s^-1) - I photosynthetic photon flux density (mol m^-2 s^-1) - N number of data points - P _m predicted photosynthetic rate at saturating CO₂ and given PPFD (mol m^-2 s^-1) - P _ml predicted photosynthetic rate at saturating CO₂ and PPFD (mol m^-2 s^-1) - R _d residual respiratory rate (mol m^-2 s^-1) - T _a air temperature (°C) - T _l leaf temperature (°C) - V reaction velocity in equation of Smith (1937, 1938) - V _max saturated reaction velocity in equation of Smith (1937, 1938) - VPA vapor pressure of water in the air (mbar/bar) - VPD vapor pressure difference between leaf and air (mbar/bar) - X substrate concentration in equation of Smith (1937, 1938) - initial slope of the PPFD response of photosynthesis at saturating CO₂ (mol CO₂/mol quanta) - (atm) initial slope of the PPFD response of photosynthesis at 340 l/l CO₂ and 21% (v/v) O₂ (mol CO₂/mol quanta) - I ^* CO₂ compensation point after correction for residual respiration (l/l) - PPFD compensation point (mol m^-2 s^-1)     

The importance of slope aspect and stand age on the photosynthetic carbon fixation capacity of forest: a case study with black locust (<Emphasis Type="Italic">Robinia pseudoacacia</Emphasis>) plantations on the Loess Plateau

The black locust (Robinia pseudoacacia L.) is an important tree species not only for the vegetation rehabilitation but also for the photosynthetic carbon dynamics on the Loess Plateau. Slope aspect and stand age play important roles in the photosynthesis of the black locusts. To investigate the photosynthetic carbon fixation capacity (PCFC) of the juvenile and mature black locusts located on the sunny and shady slopes, we have analyzed the capacity and daily dynamics of photosynthesis of the whole canopy of juvenile (6-year-old) and mature (18-year-old) black locusts located on the sunny (southeast facing) and shady (northwest facing) slopes. Mature plantations on the sunny slopes have lower average daily E, VPD, CE, A _n, LAI and PCFC than those on the shady slopes. Juvenile plantations have higher average daily g _s, E, C _i/C _a, CE, A _n and PCFC compared to the mature plantations. It is concluded that the lower average daily A _n and PCFC of the mature black locust plantations on the sunny slopes may be due to variations in the microclimatic conditions between sunny and shady slope aspects. The higher average daily A _n and PCFC of the juvenile black locust plantations are likely associated with stand age-related differences in tree sizes.     

Variable responses of mesophyll conductance to substomatal carbon dioxide concentration in common bean and soybean

Some reports indicate that mesophyll conductance (g _m) to carbon dioxide varies greatly with the substomatal carbon dioxide concentration (C _i) during the measurement, while other reports indicate little or no change in g _m with C _i. I used the oxygen sensitivity of photosynthesis to determine the response of g _m to C _i over the range of about 100 to 300 μmol mol⁻¹ C _i at constant temperature in common bean (Phaseolus vulgaris) and soybean (Glycine max) grown over a range of temperatures and photosynthetic photon flux densities (PPFD). In soybean grown and measured at high PPFD there was only a slight, approximately 15% decrease in g _m with C _i over the range of 100 to 300 μmol mol⁻¹. With lower PPFD during the measurement of g _m, and especially with low PPFD during plant growth, there was a larger decrease in g _m with C _i in soybean. In common bean, the same range in C _i resulted in about a 60% decrease in g _m for plants grown and measured at high PPFD, with an even larger decrease for plants at low growth or measurement PPFD. Growth temperatures of 20 to 30°C had little influence on the response of g _m to C _i or its absolute value in either species. It is concluded that these two species differed substantially in the sensitivity of g _m to C _i, and that PPFD but not temperature during leaf development strongly affected the response of g _m to C _i.     

The Influence of Previous Irradiance on Photosynthetic Induction in Three Species Grown in the Gap and Understory of a Fagus Crenata Forest

Photosynthetic induction responses to a sudden increase in photosynthetic photon flux density (PPFD) from lower background PPFD (0, 25, 50, and 100 mol m^–2 s^–1) to 1 000 mol m^–2 s^–1 were measured in leaves of Fagus crenata, Acer rufinerve Siebold & Zucc., and Viburnum furcatum growing in a gap and understory of a F. crenata forest in the Naeba mountains. In the gap, A. rufinerve exhibited more than 1.2-fold higher maximum net photosynthetic rate (P _Nmax) than F. crenata and V. furcatum. Meanwhile, in the understory F. crenata exhibited the highest P _Nmax among the three species. The photosynthetic induction period required to reach P _Nmax was 3–41 min. The photosynthetic responses to increase in PPFD depended on the background PPFD before increase in PPFD. The induction period required to reach P _Nmax was 2.5–6.5-fold longer when PPFD increased from darkness than when PPFD increased from 100 mol m^–2 s^–1. The induction period was correlated with initial P _N and stomatal conductance (g _s) relative to maximum values before increase in PPFD. The relationship was similar between the gap and the understory. As the background PPFD increased, the initial P _N and g _s increased, indicating that the degrees of biochemical and stomata limitations to dynamic photosynthetic performance decreased. Therefore, photosynthetic induction responses to increase in PPFD became faster with the increasing background PPFD. The differences in time required to reach induction between species, as well as between gap and understory, were mainly due to the varying of relative initial induction states in P _N and g _s at the same background PPFD.     

Compensatory effects of elevated CO<Subscript>2</Subscript> concentration on the inhibitory effects of high temperature and irradiance on photosynthetic gas exchange in carrots

We determined the interactive effects of irradiance, elevated CO₂ concentration (EC), and temperature in carrot (Daucus carota var. sativus). Plants of the cv. Red Core Chantenay (RCC) were grown in a controlled environmental plant growth room and exposed to 3 levels of photosynthetically active radiation (PAR) (400, 800, 1 200 μmol m⁻² s⁻¹), 3 leaf chamber temperatures (15, 20, 30 °C), and 2 external CO₂ concentrations (C _a), AC and EC (350 and 750 μmol mol⁻¹, respectively). Rates of net photosynthesis (P _N) and transpiration (E) and stomatal conductance (g _s ) were measured, along with water use efficiency (WUE) and ratio of internal and external CO₂ concentrations (C _i/C _a). P _N revealed an interactive effect between PAR and C _a. As PAR increased so did P _N under both C _a regimes. The g _s showed no interactive effects between the three parameters but had singular effects of temperature and PAR. E was strongly influenced by the combination of PAR and temperature. WUE was interactively affected by all three parameters. Maximum WUE occurred at 15 °C and 1 200 μmol m⁻² s^{− 1} PAR under EC. The C _i /C _a was influenced independently by temperature and C _a. Hence photosynthetic responses are interactively affected by changes in irradiance, external CO₂ concentration, and temperature. EC significantly compensates the inhibitory effects of high temperature and irradiance on P _N and WUE.     

Modelling Diurnal Courses of Photosynthesis and Transpiration of Leaves on the Basis of Stomatal and Non-Stomatal Responses,Including Photoinhibition

A mathematical model for photoinhibition of leaf photosynthesis was developed by formalising the assumptions that (1) the rate of photoinhibition is proportional to irradiance; and (2) the rate of recovery, derived from the formulae for a pseudo first-order process, is proportional to the extent of inhibition. The photoinhibition model to calculate initial photo yield is integrated into a photosynthesis-stomatal conductance (g _s) model that combines net photosynthetic rate (P _N), transpiration rate (E), and g _s, and also the leaf energy balance. The model was run to simulate the diurnal courses of P _N, E, g _s, photochemical efficiency, i.e., ratio of intercellular CO₂ concentration and CO₂ concentration over leaf surface (C _i/C _s), and leaf temperature (T ₁) under different irradiances, air temperature, and humidity separately with fixed time courses of others. When midday depression occurred under high temperature, g _s decreased the most and E the least. The duration of midday depression of g _s was the longest and that in E the shortest. E increased with increasing vapour pressure deficit (VPD) initially, but when VPD exceeded a certain value, it decreased with increasing VPD; this was caused by a rapid decrease in g _s. When air temperature exceeded a certain value, an increase in solar irradiance raised T ₁ and the degree of midday depression. High solar radiation caused large decrease in initial photon efficiency (). P _N, E, and g _s showed reasonable decreases under conditions causing photoinhibition compared with non-photoinhibition condition under high irradiance. The T ₁ under photoinhibition was higher than that under non-photoinhibition conditions, which was evident under high solar irradiance around noon. The decrease in C _i/C _s at midday implies that stomatal closure is a factor causing midday depression of photosynthesis.