Leaf Factors Affecting Light-Saturated Photosynthesis in Ecotypes of Solidago virgaurea from Exposed and Shaded Habitats

Plants of Solidago virgaurea L. from exposed and shaded habitats differ with respect to the response of the photosynthetic apparatus to the level of irradiance during growth. An analysis was carried out on leaf characteristies which might be responsible for the differences established in the rates of Hght-saturated CO₂ uptake. The clones were grown in controlled environment chambers at high and low levels of irradiance. Light-saturated rates of photosynthesis and transpiration were measured at natural and lower ambient CO₂ concentrations. A low temperature dependence of light-saturated CO₂ uptake at natural CO₂ concentrations, and a strong response to changes in stomatal width, suggested that the rate of CO₂ transfer from ambient air towards reaetion sites in chloroplasts was mainly limiting the pholosynthetic rate. Resistances to transfer of CO₂ for different parts of the pathway were calculated. There was a weak but significant correlation between stomatal conductance and the product stomatal frequency ± pore length. Mesopbyll conductance and dry weight per unit area were highly correlated in leaves not damaged by high irradiance. This suggests that mesophyll conductance increases with increasing cross sectional area (per unit leaf area) of the pathways of CO₂ transfer in the mesophyll from cell surfaces to reaction sites. The higher light-saturated photosynthesis in clones from exposed habitats when grown at high irradiance than when grown at low irradiance was attributable mainly to a lower mesophyll resistance. In shade clones the effect upon CO₂ uptake of the increase in leaf thickness when grown at high irradiance was counteracted by the associated inactivation of the photosynthetic apparatus. The difference in CO₂ uptake present between clones from exposed and shaded habitats when preconditioned to high irradiance resulted from differences in both mesophyll and stomatal resistances. A few hybrid clones of an F₁-population from a cross between a clone from an exposed habitat and a clone from a shaded habitat reacted, on the whole, in the same way as the exposed habitat parent. When grown at high irradiance, the hybrid clones showed higher photosynthetic rates than either parent; this was largely attributable to the unusually low stomatal resistance of the hybrid leaves.     

Zinc mediated effects on leaf CO2 diffusion conductances and net photosynthesis in Phaseolus vulgaris L.

Supra-optimal levels of zinc in primary leaves of Phaseolus vulgaris increased the CO₂ compensation point and inhibited net photosynthesis. Leaf morphology was modified: mesophyll intercellular area, stomatal slit length and interstomatal distance were reduced, but stomatal density increased. Internal and stomatal conductances to CO₂ diffusion decreased. These changes are discussed in relation to the observed effects on leaf gas exchange and to the previously reported inhibition of different photosynthetic and photorespiratory enzymes.     

Photosynthetic parameters in seedlings of Eucalyptus grandis as affected by rate of nitrogen supply

Seedlings of Eucalyptus grandis were grown at five different rates of nitrogen supply. Once steady‐state growth rates were established, a detailed set of CO₂ and water vapour exchange measurements were made to investigate the effects of leaf nitrogen content (N), as determined by nitrogen supply rate, on leaf structural, photosynthetic, respiratory and stomatal properties. Gas exchange data were used to parametrize the Farquhar–von Caemmerer photosynthesis model. Leaf mass per area (LMA) was negatively correlated to N. A positive correlation was observed between both day (R_d) and night respiration (R_n) and N when they were expressed on a leaf mass basis, but no correlation was found on a leaf area basis. An R_d/R_n ratio of 0·59 indicated a significant inhibition of dark respiration by light. The maximum net CO₂ assimilation rate at ambient CO₂ concentration (A_max), the maximum rate of potential electron transport (J_max) and the maximum rate of carboxylation (V_cmax) significantly increased with N, particularly when expressed on a mass basis. Although the maximum stomatal conductance to CO₂ (g_scmax) was positively correlated with A_max, there was no relationship between g_scmax and N. Leaf N content influenced the allocation of nitrogen to photosynthetic processes, resulting in a decrease of the J_max/V_cmax ratio with increasing N. It was concluded that leaf nitrogen concentration is a major determinant of photosynthetic capacity in Eucalyptus grandis seedlings and, to a lesser extent, of leaf respiration and nitrogen partitioning among photosynthetic processes, but not of stomatal conductance.     

Photosynthesis and stomatal conductance of potato leaves—effects of leaf age,irradiance, and leaf water potential

Potatoes (Solanum tuberosum L., cv. Bintje) were grown in a naturally lit glasshouse. Laboratory measurements on leaves at three insertion levels showed a decline with leaf age in photosynthetic capacity and in stomatal conductance at near saturating irradiance. Conductance declined somewhat more with age than photosynthesis, resulting in a smaller internal CO₂ concentration in older relative to younger leaves. Leaves with different insertion number behaved similarly. The changes in photosynthesis rate and in nitrogen content with leaf age were closely correlated. When PAR exceeded circa 100 W m^–2 the rate of photosynthesis and stomatal conductance changed proportionally as indicated by a constant internal CO₂ concentration. The photosynthesis-irradiance data were fitted to an asymptotic exponential model. The parameters of the model are AMAX, the rate of photosynthesis at infinite irradiance, and EFF, the slope at low light levels. AMAX declined strongly with leaf age, as did EFF, but to a smaller extent. During drought stress photosynthetic capacity declined directly with decreasing water potential (range –0.6 to –1.1 MPa). Initially, stomatal conductance declined faster than photosynthetic capacity.Abbreviations LNx leaf number x, counted in acropetal direction - DAP days after planting - DALA days after leaf appearance - C_i CO₂ concentration in the leaf - C_a CO₂ concentration in ambient air - LWP leaf water potential - OP osmotic potential - PAR photosynthetically active radiation     

Alterations in Rubisco activity and in stomatal behavior induce a daily rhythm in photosynthesis of aerial leaves in the amphibious-plant Nuphar lutea

Nuphar lutea is an amphibious plant with submerged and aerial foliage, which raises the question how do both leaf types perform photosynthetically in two different environments. We found that the aerial leaves function like terrestrial sun-leaves in that their photosynthetic capability was high and saturated under high irradiance (ca. 1,500 μmol photons m⁻² s⁻¹). We show that stomatal opening and Rubisco activity in these leaves co-limited photosynthesis at saturating irradiance fluctuating in a daily rhythm. In the morning, sunlight stimulated stomatal opening, Rubisco synthesis, and the neutralization of a night-accumulated Rubisco inhibitor. Consequently, the light-saturated quantum efficiency and rate of photosynthesis increased 10-fold by midday. During the afternoon, gradual closure of the stomata and a decrease in Rubisco content reduced the light-saturated photosynthetic rate. However, at limited irradiance, stomatal behavior and Rubisco content had only a marginal effect on the photosynthetic rate, which did not change during the day. In contrast to the aerial leaves, the photosynthesis rate of the submerged leaves, adapted to a shaded environment, was saturated under lower irradiance. The light-saturated quantum efficiency of these leaves was much lower and did not change during the day. Due to their low photosynthetic affinity for CO₂ (35 μM) and inability to utilize other inorganic carbon species, their photosynthetic rate at air-equilibrated water was CO₂-limited. These results reveal differences in the photosynthetic performance of the two types of Nuphar leaves and unravel how photosynthetic daily rhythm in the aerial leaves is controlled.     

Photosynthesis in leaves of the juvenile and adult phase of ivy (Hedera helix)

Abstract Photosynthetic and anatomical parameters of leaves from the juvenile and adult part of an ivy plant (Hedera helix L.) have been determined and compared with each other. Light-saturated net photosynthesis (per unit leaf area) was about 1.5 times higher in adult leaves than in juvenile ones. The lower photosynthetic capacity of juvenile leaves was caused by a lower stomatal and especially a lower residual conductance to the CO₂-transfer. This corresponds with anatomical features of the leaves, i.e. lower stomatal frequency, fewer chloroplasts per cell, and – most important – thinner leaves, as well as with a less efficient photosynthetic apparatus measured as Hill reaction of isolated broken chloroplasts and activity of ribulose bisphosphate carboxylase. No differences in the respiration in light (relative to net photosynthesis) and in the CO₂-compensation concentration could be detected between the two leaf types. These observed anatomical and photosynthetic parameters of the juvenile and adult ivy leaves resemble those reported for shade and sun leaves, respectively, although the leaves investigated originated from the same light environment.     

Seasonal changes in photosynthetic characteristics of Pachysandra terminalis (Buxaceae), an evergreen woodland chamaephyte,in the cool temperate regions of Japan

Summary Seasonal changes in photosynthetic capacity, and photosynthetic responses to intercellular CO₂ concentration and irradiance were investigated under laboratory conditions on intact leaves of Pachysandra terminalis. Photosynthetic capacity and stomatal conductance under saturating light intensity and constant water vapor pressure deficit showed almost the same seasonal trend. They increased from early June just after the expansion of leaves, reached the maximum in late-Septemer, and then decreased to winter. In over-wintering leaves they recovered and increased immediately after snow-melting, reached a first maximum in late April, and then decreased to early July in response to the reduction of light intensity on the forest floor. There-after, they increased from mid August, reached a second maximum in late September, and then decreased to winter. The parallel changes of photosynthesis and stomatal conductane indicate a more or less constant intercellular CO₂ concentration throughout the year. The calculated values of relative stomatal limitation of photosynthesis were nearly constant throughout the year, irrespective of leaf age. The results indicate that the seasonal changes in light-saturated photosynthetic capacity are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Indeed, carboxylation efficiency assessed by the inital slope of the Ci-photosynthesis curve changed in proportion to seasonal changes of the photosynthetic capacity in both current-year and over-wintered leaves. High photosynthetic capacity in current-year leaves as compared with one-year-old leaves was also due to the high photosynthetic capacity of mesophyll. Nevertheless, stomatal conductance changed in proportion to photosynthetic capacity, indicating that stomatal conductance is regulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations are maintained constant. The quantum yield also changed seasonally parallel with that in the photosynthetic capacity.Contribution No. 2893 from the Institute of Low Temperature Science     

Photosynthetic and structural acclimation to light direction in vertical leaves of Silphium terebinthinaceum

The azimuth of vertical leaves of Silphium terebinthinaceum profoundly influenced total daily irradiance as well as the proportion of direct versus diffuse light incident on the adaxial and abaxial leaf surface. These differences caused structural and physiological adjustments in leaves that affected photosynthetic performance. Leaves with the adaxial surface facing East received equal daily integrated irradiance on each surface, and these leaves had similar photosynthetic rates when irradiated on either the adaxial or abaxial surface. The adaxial surface of East-facing leaves was also the only surface to receive more direct than diffuse irradiance and this was the only leaf side which had a clearly defined columnar palisade layer. A potential cost of constructing East-facing leaves with symmetrical photosynthetic capcity was a 25% higher specific leaf mass and increased leaf thickness in comparison to asymmetrical South-facing leaves. The adaxial surface of South-facing leaves received approximately three times more daily integrated irradiance than the abaxial surface. When measured at saturating CO₂ and irradiance, these leaves had 42% higher photosynthetic rates when irradiated on the adaxial surface than when irradiated on the abaxial surface. However, there was no difference in photosynthesis for these leaves when irradiated on either surface when measurements were made at ambient CO₂. Stomatal distribution (mean adaxial/abaxial stomatal density = 0.61) was unaffected by leaf orientation. Thus, the potential for high photosynthetic rates of adaxial palisade cells in South-facing leaves at ambient CO₂ concentrations may have been constrained by stomatal limitations to gas exchange. The distribution of soluble protein and chlorophyll within leaves suggests that palisade and spongy mesophyll cells acclimated to their local light environment. The protein/chlorophyll ratio was high in the palisade layers and decreased in the spongy mesophyll cells, presumably corresponding to the attentuation of light as it penetrates leaves. Unlike some species, the chlorophyll a/b ratio and the degree of thylakoid stacking was uniform throughout the thickness of the leaf. It appears that sun-shade acclimation among cell layers of Silphium terebinthinaceum leaves is accomplished without adjustment to the chlorophyll a/b ratio or to thylakoid membrane structure.     

Photosynthesis in the water-stressed C4 grass Setaria sphacelata is mainly limited by stomata with both rapidly and slowly imposed water deficits

A comparison of the effects of a rapid and a slowly imposed water deficit on photosynthesis was performed in Setaria sphacelata var. splendida (Stapf) Clayton, a C₄ NADP‐ME grass. Gas exchange was measured in rapidly and slowly dehydrated adult leaves either under atmospheric CO₂ partial pressure with an infrared gas analyser or under saturating CO₂ partial pressure with a leaf disc oxygen electrode. These measurements were used to calculate stomatal and non‐stomatal limitations to photosynthesis. These were further investigated using modulated chlorophyll a fluorescence measurements and photosynthetic pigment quantification. The decrease of net photosynthesis, leaf conductance and water use efficiency was more pronounced under rapid stress than in slow stress. However, photosynthesis is always mainly limited by stomata in both types of stress, albeit the contribution of non‐stomatal limitations increases at severe water deficits in slow stress experiments. The substomatal CO₂ partial pressure significantly increased in both types of stress, suggesting an increased resistance due to an internal barrier to CO₂ diffusion. Physical alterations in the structure of the intercellular spaces due to leaf shrinkage may account for these results. The maximal photochemical efficiency of photosystem II (PSII) was remarkably resistant to stress, as the F_v/F_m ratio decreased only at severe water deficit. On the contrary, the effective photochemical efficiency of PSII (ΔF/F′_m) measured under high actinic light decreased linearly in both types of stress, although in a more pronounced way under rapid stress. A similar variation in photochemical quenching suggests that the decrease of ΔF/F′_m is mainly due to the closure of PSII reaction centres. The non‐photochemical quenching did not change significantly except under severe dehydration indicating that the energization state of thylakoids remained stable under stress. The decrease observed in photosynthetic pigments may be an adaptation to stress rather than a limiting factor to photosynthesis. Results suggests that, although intrinsic mesophyll metabolic inhibitions occur, stomatal limitation to CO₂ diffusion is the main reason for the decrease in photosynthesis.     

Stomatal function,density and pattern,and CO2 assimilation in Arabidopsis thaliana tmm1 and sdd1‐1 mutants

• Stomata modulate the exchange of water and CO₂ between plant and atmosphere. Although stomatal density is known to affect CO₂ diffusion into the leaf and thus photosynthetic rate, the effect of stomatal density and patterning on CO₂ assimilation is not fully understood.
• We used wild types Col‐0 and C24 and stomatal mutants sdd1‐1 and tmm1 of Arabidopsis thaliana, differing in stomatal density and pattern, to study the effects of these variations on both stomatal and mesophyll conductance and CO₂ assimilation rate. Anatomical parameters of stomata, leaf temperature and carbon isotope discrimination were also assessed.
• Our results indicate that increased stomatal density enhanced stomatal conductance in sdd1‐1 plants, with no effect on photosynthesis, due to both unchanged photosynthetic capacity and decreased mesophyll conductance. Clustering (abnormal patterning formed by clusters of two or more stomata) and a highly unequal distribution of stomata between the adaxial and abaxial leaf sides in tmm1 mutants also had no effect on photosynthesis.
• Except at very high stomatal densities, stomatal conductance and water loss were proportional to stomatal density. Stomatal formation in clusters reduced stomatal dynamics and their operational range as well as the efficiency of CO₂ transport.

     

Photosynthetic Responses of a Temperate Liana to Xylella fastidiosa Infection and Water Stress

Xylella fastidiosa is a xylem‐limited bacterial plant pathogen that causes bacterial leaf scorch in its hosts. Our previous work showed that water stress enhances leaf scorch symptom severity and progression along the stem of a liana, Parthenocissus quinquefolia, infected by X. fastidiosa. This paper explores the photosynthetic gas exchange responses of P. quinquefolia, with the aim to elucidate mechanisms behind disease expression and its interaction with water stress. We used a 2 × 2‐complete factorial design, repeated over two growing seasons, with high and low soil moisture levels and infected and non‐infected plants. In both years, low soil moisture levels reduced leaf water potentials, net photosynthesis and stomatal conductance at all leaf positions, while X. fastidiosa‐infection reduced these parameters at basally located leaves only. Intercellular CO₂ concentrations were reduced in apical leaves, but increased at the most basal leaf location, implicating a non‐stomatal reduction of photosynthesis in leaves showing the greatest disease development. This result was supported by measured reductions in photosynthetic rates of basal leaves at high CO₂ concentrations, where stomatal limitation was eliminated. Repeated measurements over the summer of 2000 showed that the effects of water stress and infection were progressive over time, reaching their greatest extent in September. By reducing stomatal conductances at moderate levels of water stress, P. quinquefolia maintained relatively high leaf water potentials and delayed the onset of photosynthetic damage due to pathogen and drought‐induced water stress. In addition, chlorophyll fluorescence measurements showed that P. quinquefolia has an efficient means of dissipating excess light energy that protects the photosynthetic machinery of leaves from irreversible photoinhibitory damage that may occur during stress‐induced stomatal limitation of photosynthesis. However, severe stress induced by disease and drought eventually led to non‐stomatal decreases in photosynthesis associated with leaf senescence.     

Impairment of photosynthesis by chilling-temperatures in tomato

Chilling of attached tomato leaves (cv. Rutgers) in the dark for 16 hours at 1 C decreased both photosynthesis and transpiration. To separate the effects of chilling on stomatal CO₂ conductance from more direct effects of chilling on the chloroplasts' activities, measurements of photosynthesis and transpiration were made at atmospheric and saturating CO₂ levels. At atmospheric CO₂, the inhibition of photosynthesis was approximately 60%, of which about 35% was attributable to the impairment of chloroplast function and about 25% was attributable to decreased stomatal conductance. However, the affinity of the photosynthetic apparatus for CO₂ was not changed by chilling, since the dependence of the relative rate of photosynthesis on the intercellular CO₂ concentration was unaltered. The apparent quantum requirement for CO₂ reduction also was identical in chilled and unchilled plants. This observation contradicts the widely held notion that the chilling-induced inhibition of photosynthesis is caused by an impairment of the water oxidation mechanism. The impairment of chloroplast activity was not a consequence of an unfavorable water status within the leaf, since chilling caused only a small drop (1 bar) in water potential. A small loss of chlorophyll resulted as a secondary effect of chilling, but this loss of chlorophyll was eliminated as a cause of the inhibition of photosynthesis.     

Effects of atmospheric CO2 concentration, irradiance, and soil nitrogen availability on leaf photosynthetic traits of Polygonum sachalinense around natural CO2 springs in northern Japan

Long-term exposure to elevated CO₂ concentration will affect the traits of wild plants in association with other environmental factors. We investigated multiple effects of atmospheric CO₂ concentration, irradiance, and soil N availability on the leaf photosynthetic traits of a herbaceous species, Polygonum sachalinense, growing around natural CO₂ springs in northern Japan. Atmospheric CO₂ concentration and its interaction with irradiance and soil N availability affected several leaf traits. Leaf mass per unit area increased and N per mass decreased with increasing CO₂ and irradiance. Leaf N per area increased with increasing soil N availability at higher CO₂ concentrations. The photosynthetic rate under growth CO₂ conditions increased with increasing irradiance and CO₂, and with increasing soil N at higher CO₂ concentrations. The maximal velocity of ribulose 1,5-bisphosphate carboxylation (V _cmax) was affected by the interaction of CO₂ and soil N, suggesting that down-regulation of photosynthesis at elevated CO₂ was more evident at lower soil N availability. The ratio of the maximum rate of electron transport to V _cmax (J _max/V _cmax) increased with increasing CO₂, suggesting that the plants used N efficiently for photosynthesis at high CO₂ concentrations by changes in N partitioning. To what extent elevated CO₂ influenced plant traits depended on other environmental factors. As wild plants are subject to a wide range of light and nutrient availability, our results highlight the importance of these environmental factors when the effects of elevated CO₂ on plants are evaluated.     

Increased stomatal conductance induces rapid changes to photosynthetic rate in response to naturally fluctuating light conditions in rice

A close correlation between stomatal conductance and the steady-state photosynthetic rate has been observed for diverse plant species under various environmental conditions. However, it remains unclear whether stomatal conductance is a major limiting factor for the photosynthetic rate under naturally fluctuating light conditions. We analysed a SLAC1 knockout rice line to examine the role of stomatal conductance in photosynthetic responses to fluctuating light. SLAC1 encodes a stomatal anion channel that regulates stomatal closure. Long exposures to weak light before treatments with strong light increased the photosynthetic induction time required for plants to reach a steady-state photosynthetic rate and also induced stomatal limitation of photosynthesis by restricting the diffusion of CO₂ into leaves. The slac1 mutant exhibited a significantly higher rate of stomatal opening after an increase in irradiance than wild-type plants, leading to a higher rate of photosynthetic induction. Under natural conditions, in which irradiance levels are highly variable, the stomata of the slac1 mutant remained open to ensure efficient photosynthetic reaction. These observations reveal that stomatal conductance is important for regulating photosynthesis in rice plants in the natural environment with fluctuating light.     

Elevated CO<Subscript>2</Subscript> reduces stomatal and metabolic limitations on photosynthesis caused by salinity in <Emphasis Type="Italic">Hordeum vulgare</Emphasis>

The future environment may be altered by high concentrations of salt in the soil and elevated [CO₂] in the atmosphere. These have opposite effects on photosynthesis. Generally, salt stress inhibits photosynthesis by stomatal and non-stomatal mechanisms; in contrast, elevated [CO₂] stimulates photosynthesis by increasing CO₂ availability in the Rubisco carboxylating site and by reducing photorespiration. However, few studies have focused on the interactive effects of these factors on photosynthesis. To elucidate this knowledge gap, we grew the barley plant, Hordeum vulgare (cv. Iranis), with and without salt stress at either ambient or elevated atmospheric [CO₂] (350 or 700 μmol mol⁻¹ CO₂, respectively). We measured growth, several photosynthetic and fluorescence parameters, and carbohydrate content. Under saline conditions, the photosynthetic rate decreased, mostly because of stomatal limitations. Increasing salinity progressively increased metabolic (photochemical and biochemical) limitation; this included an increase in non-photochemical quenching and a reduction in the PSII quantum yield. When salinity was combined with elevated CO₂, the rate of CO₂ diffusion to the carboxylating site increased, despite lower stomatal and internal conductance. The greater CO₂ availability increased the electron sink capacity, which alleviated the salt-induced metabolic limitations on the photosynthetic rate. Consequently, elevated CO₂ partially mitigated the saline effects on photosynthesis by maintaining favorable biochemistry and photochemistry in barley leaves.     

Seasonal changes in photosynthetic characteristics of Anemone raddeana,a spring-active geophyte,in the temperate region of Japan

Summary Seasonal changes in the photosynthetic characteristics of intact involucral leaves of Anemone raddeana were investigated under laboratory conditions. Net photosynthesis and constant water vapor pressure deficit showed almost the same seasonal trend. They increased rapidly from mid-April immediately after unfolding of the leaves and reached the maximum in late-April, before the maximum expansion of the leaves. They retained the maximum values until early-May and then decreased toward late-May with a progress of leaf senescence. The calculated values of intercellular CO₂ concentration and relative stomatal limitation of photosynthesis showed no significant change throughout the season. The carboxylation efficiency as assessed by the initial slope of Ci-photosynthesis curve and the net photosynthesis under a high Ci regime varied seasonally in parallel with the change of the light-saturated photosynthesis. The results indicate that the seasonal changes in light-saturated net photosynthesis are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Nevertheless, leaf conductance changed concomitantly with photosynthetic capacity, indicating that the seasonal change in stomatal conductance is modulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations is maintained constant. The shape of light-photosynthesis curve was similar to that of sun-leaf type. The quantum yield also changed simultaneously with the photosynthetic capacity throughout the season.Contribution No. 2965 from the Institute of Low Temperature Science     

Heterogeneity of leaf CO2 assimilation during photosynthetic induction

Spatial mid temporal variations in the distribution of photosynthesis over the leaf area were investigated during induction upon illumination of Rosa rubiginosa L. leaves. Gas exchange and maps of relative photosynthetie electron transport activity computed from chlorophyll fluorescence images were simultaneously monitored. In air, after 15 h of dark adaptation, linear electron transport was heterogeneously distributed over the leaf area during the induction. This patchy induction was explained by asynchronous metabolism activation for the first 10 min of illumination, concomitant asynchronous limitation by intrinsic metabolism and stomatal apertures (10–30 min) and finally by only stomatal limitation beyond 30 min. A brief transition to non-photorespiratory conditions after 20 min of illumination under subsaturating irradiance revealed a marked heterogeneity of CO₂ assimilation, presumably as a result of heterogeneous stomatal apertures. The frequency distribution of CO₂ assimilation was unimodal. During the induction, heterogeneity gradually decreased and photosynthesis was uniform at steady-state. After 10 min of dark adaptation, heterogeneity of linear electron transport activity occurred during the first 15 min of a second induction and mainly resulted from metabolic limitation.     

Diffusion limitations and metabolic factors associated with inhibition and recovery of photosynthesis from drought stress in a C3 perennial grass species

Stomatal closure and metabolic impairment under drought stress limits photosynthesis. The objective of this study was to determine major stomatal and metabolic factors involved in photosynthetic responses to drought and recovery upon re‐watering in a C₃ perennial grass species, Kentucky bluegrass (Poa pratensis L.). Two genotypes differing in drought resistance, ‘Midnight’ (tolerant) and ‘Brilliant’ (sensitive), were subjected to drought stress for 15 days and then re‐watered for 10 days in growth chambers. Single‐leaf net photosynthetic rate (A), stomatal conductance (g_s) and transpiration rate (Tr) decreased during drought, with a less rapid decline in ‘Midnight’ than in ‘Brilliant’. Photochemical efficiency, Rubisco activity and activation state declined during drought, but were significantly higher in ‘Midnight’ than in ‘Brilliant’. The relationship between A and internal leaf CO₂ concentration (A/Ci curve) during drought and re‐watering was analyzed to estimate the relative influence of stomatal and non‐stomatal components on photosynthesis. Stomatal limitation (Ls %), non‐stomatal limitation (Lns %), CO₂ compensation point (CP) and dark respiration (Rd) increased with stress duration in both genotypes, but to a lesser extent in ‘Midnight’. Maximum CO₂ assimilation rate (A_max), carboxylation efficiency (CE) and mesophyll conductance (g_m) declined, but ‘Midnight’ had significantly higher levels of A_max, CE and g_m than ‘Brilliant’. Maximum carboxylation rate of Rubisco (V_cmax) and ribulose‐1,5‐bisphospate (RuBP) regeneration capacity mediated by maximum electron transport rate (J_max) decreased from moderate to severe drought stress in both genotypes, but to a greater extent in ‘Brilliant’ than in ‘Midnight’. After re‐watering, RWC restored to about 90% of the control levels in both genotypes, whereas A, g_s, Tr and Fv/Fm was only partially recovered, with a higher recovery level in ‘Midnight’ than in ‘Brilliant’. Rubisco activity and activation state restored to the control level after re‐watering, with more rapid increase in ‘Midnight’ than in ‘Brilliant’. The values of Ls, Lns, CP and Rd declined, and A_max, CE, V_cmax, J_max and g_m increased after re‐watering, with more rapid change in all parameters in ‘Midnight’ than in ‘Brilliant’. These results indicated that the maintenance of higher A and A_max under drought stress in drought‐tolerant Kentucky bluegrass could be attributed to higher Rubico activation state, higher CE and less stomatal limitation. The ability to resume metabolic activity (A_max, CE, Fv/Fm and Rubisco) was observed in the drought‐tolerant genotype and is the most likely cause for the increased recuperative ability of photosynthesis. Incomplete recovery of photosynthesis upon re‐watering could be attributable to lasting stomatal limitations caused by severe drought damage in both genotypes. Promoting rapid stomatal recovery from drought stress may be critical for plants to resume full photosynthetic capacity in C₃ perennial grass species.     

Adaptive temperature regulation in the little bird in winter: predictions from a stochastic dynamic programming model

Stomatal CO₂ responsiveness and photosynthetic capacity vary greatly among plant species, but the factors controlling these physiological leaf traits are often poorly understood. To explore if these traits are linked to taxonomic group identity and/or to other plant functional traits, we investigated the short-term stomatal CO₂ responses and the maximum rates of photosynthetic carboxylation (V _cmax) and electron transport (J _max) in an evolutionary broad range of tropical woody plant species. The study included 21 species representing four major seed plant taxa: gymnosperms, monocots, rosids and asterids. We found that stomatal closure responses to increased CO₂ were stronger in angiosperms than in gymnosperms, and in monocots compared to dicots. Stomatal CO₂ responsiveness was not significantly related to any of the other functional traits investigated, while a parameter describing the relationship between photosynthesis and stomatal conductance in combined leaf gas exchange models (g ₁) was related to leaf area-specific plant hydraulic conductance. For photosynthesis, we found that the interspecific variation in V _cmax and J _max was related to within leaf nitrogen (N) allocation rather than to area-based total leaf N content. Within-leaf N allocation and water use were strongly co-ordinated (r ² = 0.67), such that species with high fractional N investments into compounds maximizing photosynthetic capacity also had high stomatal conductance. We conclude that while stomatal CO₂ responsiveness of tropical woody species seems poorly related to other plant functional traits, photosynthetic capacity is linked to fractional within-leaf N allocation rather than total leaf N content and is closely co-ordinated with leaf water use.     

Photosynthesis and Transpiration as a Function of Gaseous Diffusive Resistances in Orange Leaves

The CO₂ and H₂O vapour exchange of single attached orange, Citrus sinensis (L.), leaves was measured under laboratory conditions using infrared gas analysis. Gaseous diffusive resistances were derived from measurements at a saturating irradiance and at a leaf temperature optimum for photosynthesis. Variation in leaf resistance (within the range 1.6 to 60 s cm^-1) induced by moisture status, or by cyclic oscillations in stomatal aperture, was associated with changes in both photosynthesis and transpiration. At low leaf resistance (ri less than 10 s cm^-1) the ratio of transpiration to photosynthesis declined with reduced stomatal aperture, indicating a tighter stomatal control over H₂O vapour loss than over CO₂ assimilation. At higher leaf resistance (ri greater than 10 s cm^-1) changes in transpiration and photosynthesis were linearly related, but leaf resistance and mesophyll resistance were also positively correlated, so that strictly stomatal control of photosynthesis became more apparent than real. This evidence, combined with direct measurements of CO₂ diffusive resistances (in a -O₂ gas stream) emphasised the presence of a significant mesophyll resistance; i.e., an additional and rate limiting resistance to CO₂ assimilation over and above that encountered by H₂O vapour escaping from the leaf.