Effects of Water-Deficit Stress on Photosynthesis, Its Components and Component Limitations, and on Water Use Efficiency in Wheat (Triticum aestivum L.)

It is of theoretical as well as practical interest to identify the components of the photosynthetic machinery that govern variability in photosynthesis rate (A) and water-use efficiency (WUE), and to define the extent by which the component processes limit A and WUE during developing water-deficit stress. For that purpose, leaf exchange of CO₂ and H₂O was determined in two growth-chamber-grown wheat cultivars (Triticum aestivum L. cv TAM W-101 and cv Sturdy), and the capacity of A was determined and broken down into carboxylation efficiency (c.e.), light- and CO₂-saturated A, and stomatal conductance (g_s) components. The limitations on A measured at ambient CO₂ concentration (A₃₅₀) were estimated. No cultivar difference was observed when A₃₅₀ was plotted versus leaf water potential (Ψ_w). Light- and CO₂-saturated A, c.e., and g_s decreased with decreasing leaf Ψ_w, but of the corresponding photosynthesis limitations only those caused by insufficient c.e. and g_s increased. Thus, reduced stomatal aperture and Calvin cycle activity, but not electron transport/photophosphorylation, appeared to be major reasons for drought stress-induced inhibition of A₃₅₀. WUE measured as A₃₅₀/g_s first increased with stomatal closure down to a g_s of about 0.25 mol H₂O m⁻² s⁻¹ (Ψ_w = −1.6 MPa). However, it was predicted that A₃₅₀/g_s would decrease with more severe stress due to inhibition of c.e.     

Limitation factors for photosynthesis in ‘Bluecrop’ highbush blueberry (Vaccinium corymbosum) leaves in response to moderate water stress

The levels of stomatal, mesophyll and biochemical limitations in CO₂ assimilation of ‘Bluecrop’ highbush blueberry leaves were compared at two different levels of leaf water potential. The leaf water potentials were ?1.49 and ?1.94 MPa in daily-irrigated (DI) and non-irrigated (NI) shrubs, respectively. The NI shrubs represented plants under moderate water stress. Mesophyll conductance (g _m) and chloroplastic CO₂ concentration (C _c) were estimated by combined measurements of gas exchange and chlorophyll fluorescence under various intercellular CO₂ concentrations (C _i). Net CO₂ assimilation rates (A _n) as a function of C _c were used for calculating maximum carboxylation efficiency (α _cmax) at the real sites of CO₂ assimilation. Maximum A _n (A _nmax) from the light response curves at 400 μmol mol^?1 air of ambient CO₂ concentration (C _a) were lower in the leaves of NI shrubs than in those of DI ones. However, electron transport rates were higher in the leaves of NI shrubs than in those of DI ones. The decrease in CO₂ assimilation following water stress may be caused by a decrease in g _m rather than a decrease in stomatal conductance (g _s) according to limitation analysis. Limitation rates by g _s, calculated at 400 μmol mol^?1 air of C _a in A _n-C _i curves, were not significantly different between the leaves of DI and NI shrubs. However, limitation rates by g _m from A _n-C _c curves were significantly higher in the leaves of NI shrubs than in those of DI ones. Maximum carboxylation efficiency (α _cmax) values calculated from the A _n-C _c curve, contrary to those calculated from the A _n-C _i curve, were higher in the leaves of NI shrubs than in those of DI ones. Consequently, mesophyll limitation than stomatal and biochemical limitations mainly down-regulated the photosynthesis in the leaves of ‘Bluecrop’ blueberry shrubs during moderate water stress.     

Purification of the Trehalase GMTRE1 from Soybean Nodules and Cloning of Its cDNA. GMTRE1 Is Expressed at a Low Level in Multiple Tissues

The effects of ultraviolet-B (UV-B) radiation on stomatal conductance (g_s) in pea (Pisum sativum L.), commelina (Commelina communis L.), and oilseed rape (Brassica napus L.) plants were investigated. Plants were grown in a greenhouse either with three different high ratios of UV-B to photosynthetically active radiation or with no UV-B radiation. Pea plants grown in the highest UV-B radiation (0.63 W m⁻²) exhibited a substantial decrease of adaxial and abaxial g_s (approximately 80% and 40%, respectively). With growth in 0.30 W m⁻² of UV-B adaxial g_s was decreased by 23%, with no effect on abaxial g_s, and lower UV-B irradiance of 0.21 W m⁻² had no effect on either surface. Although abaxial g_s increased when leaves were turned over in control plants, it did not in plants grown with the highest UV-B. Adaxial g_s in commelina and oilseed rape also decreased on exposure to high UV-B (0.63 W m⁻²). For previously unexposed pea plants the time course of the effect of UV-B on g_s was slow, with a lag of approximately 4 h, and a time constant of approximately 3 h. We conclude that there is a direct effect of UV-B on stomata in addition to that caused by changes in mesophyll photosynthesis.     

Correlation of Stomatal Conductance with Photosynthetic Capacity of Cotton Only in a CO2-Enriched Atmosphere: Mediation by Abscisic Acid?

Some evidence indicates that photosynthetic rate (A) and stomatal conductance (g) of leaves are correlated across diverse environments. The correlation between A and g has led to the postulation of a “messenger” from the mesophyll that directs stomatal behavior. Because A is a function of intercellular CO₂ concentration (c_i), which is in turn a function of g, such a correlation may be partially mediated by c_i if g is to some degree an independent variable. Among individual sunlit leaves in a cotton (Gossypium hirsutum L.) canopy in the field, A was significantly correlated with g (r² = 0.41, n = 63). The relative photosynthetic capacity of each leaf was calculated as a measure of mesophyll properties independent of c_i. This approach revealed that, in the absence of c_i effects, mesophyll photosynthetic capacity was unrelated to g (r² = 0.06). When plants were grown in an atmosphere enriched to about 650 microliters per liter of CO₂, however, photosynthetic capacity remained strongly correlated with g even though the procedure discounted any effect of variable c_i. This “residual” correlation implies the existence of a messenger in CO₂-enriched plants. Enriched CO₂ also greatly increased stomatal response to abscisic acid (ABA) injected into intact leaves. The data provide no evidence for a messenger to coordinate g with A at ambient levels of CO₂. In a CO₂-enriched atmosphere, though, ABA may function as such a messenger because the sensitivity of the system to ABA is enhanced.     

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.     

Chloride as a macronutrient increases water‐use efficiency by anatomically driven reduced stomatal conductance and increased mesophyll diffusion to CO2

Chloride (Cl^?) has been recently described as a beneficial macronutrient, playing specific roles in promoting plant growth and water‐use efficiency (WUE). However, it is still unclear how Cl^? could be beneficial, especially in comparison with nitrate (NO₃^?), an essential source of nitrogen that shares with Cl^? similar physical and osmotic properties, as well as common transport mechanisms. In tobacco plants, macronutrient levels of Cl^? specifically reduce stomatal conductance (g_s) without a concomitant reduction in the net photosynthesis rate (A_N). As stomata‐mediated water loss through transpiration is inherent in the need of C₃ plants to capture CO₂, simultaneous increase in photosynthesis and WUE is of great relevance to achieve a sustainable increase in C₃ crop productivity. Our results showed that Cl^?‐mediated stimulation of larger leaf cells leads to a reduction in stomatal density, which in turn reduces g_s and water consumption. Conversely, Cl^? improves mesophyll diffusion conductance to CO₂ (g_m) and photosynthetic performance due to a higher surface area of chloroplasts exposed to the intercellular airspace of mesophyll cells, possibly as a consequence of the stimulation of chloroplast biogenesis. A key finding of this study is the simultaneous improvement of A_N and WUE due to macronutrient Cl^? nutrition. This work identifies relevant and specific functions in which Cl^? participates as a beneficial macronutrient for higher plants, uncovering a sustainable approach to improve crop yield.     

Effects of sulfur on the photosynthesis of intact leaves and isolated chloroplasts of sugar beets

Effects of sulfur on photosynthesis in sugar beets (Beta vulgaris L. cv. F58-554H1) were studied by inducing sulfur deficiency and determining changes in the photosynthesis of whole attached leaves and of isolated chloroplasts. The rates of photosynthetic CO₂ uptake by intact leaves, photoreduction of ferricyanide, cyclic and noncyclic photophosphorylation of isolated chloroplasts, and the rate of CO₂ assimilation by ribulose diphosphate carboxylase, decreased with decrease in total leaf sulfur from 2500 to about 500 μg g⁻¹ dry weight. Sulfur deficiency reduced photosynthesis through an effect on chlorophyll content, which decreased linearly with leaf sulfur, and by decreasing the rate of photosynthesis per unit chlorophyll. There was only a small effect of sulfur deficiency on stomatal diffusion resistance to CO₂ until leaf sulfur decreased below 1000 μg g⁻¹ when stomatal resistance became a more significant proportion of the total diffusion resistance to CO₂. Light respiration rates were positively correlated with photosynthesis rates and dark respiration was unchanged as leaf sulfur concentrations declined.     

Stomatal,mesophyll conductance,and biochemical limitations to photosynthesis during induction

The dynamics of leaf photosynthesis in fluctuating light affects carbon gain by plants. Mesophyll conductance (g_m) limits CO₂ assimilation rate (A) under the steady state, but the extent of this limitation under non-steady-state conditions is unknown. In the present study, we aimed to characterize the dynamics of g_m and the limitations to A imposed by gas diffusional and biochemical processes under fluctuating light. The induction responses of A, stomatal conductance (g_s), g_m, and the maximum rate of RuBP carboxylation (V_cmax) or electron transport (J) were investigated in Arabidopsis (Arabidopsis thaliana (L.)) and tobacco (Nicotiana tabacum L.). We first characterized g_m induction after a change from darkness to light. Each limitation to A imposed by g_m, g_s and V_cmax or J was significant during induction, indicating that gas diffusional and biochemical processes limit photosynthesis. Initially, g_s imposed the greatest limitation to A, showing the slowest response under high light after long and short periods of darkness, assuming RuBP-carboxylation limitation. However, if RuBP-regeneration limitation was assumed, then J imposed the greatest limitation. g_m did not vary much following short interruptions to light. The limitation to A imposed by g_m was the smallest of all the limitations for most of the induction phase. This suggests that altering induction kinetics of mesophyll conductance would have little impact on A following a change in light. To enhance the carbon gain by plants under naturally dynamic light environments, attention should therefore be focused on faster stomatal opening or activation of electron transport.

Gas diffusional and biochemical processes impose significant limitations to CO2 assimilation during photosynthetic induction.     

Photosynthetic Diffusional Constraints Affect Yield in Drought Stressed Rice Cultivars during Flowering

Global production of rice (Oryza sativa) grain is limited by water availability and the low ‘leaf-level’ photosynthetic capacity of many cultivars. Oryza sativa is extremely susceptible to water-deficits; therefore, predicted increases in the frequency and duration of drought events, combined with future rises in global temperatures and food demand, necessitate the development of more productive and drought tolerant cultivars. We investigated the underlying physiological, isotopic and morphological responses to water-deficit in seven common varieties of O. sativa, subjected to prolonged drought of varying intensities, for phenotyping purposes in open field conditions. Significant variation was observed in leaf-level photosynthesis rates (A) under both water treatments. Yield and A were influenced by the conductance of the mesophyll layer to CO₂ (g _m) and not by stomatal conductance (g _s). Mesophyll conductance declined during drought to differing extents among the cultivars; those varieties that maintained g _m during water-deficit sustained A and yield to a greater extent. However, the variety with the highest g _m and yield under well-watered conditions (IR55419-04) was distinct from the most effective cultivar under drought (Vandana). Mesophyll conductance most effectively characterises the photosynthetic capacity and yield of O. sativa cultivars under both well-watered and water-deficit conditions; however, the desired attributes of high g _m during optimal growth conditions and the capacity for g _m to remain constant during water-deficit may be mutually exclusive. Nonetheless, future genetic and physiological studies aimed at enhancing O. sativa yield and drought stress tolerance should investigate the biochemistry and morphology of the interface between the sub-stomatal pore and mesophyll layer.     

Leaf anatomical properties in relation to differences in mesophyll conductance to CO2 and photosynthesis in two related Mediterranean Abies species

Abies alba and Abies pinsapo are closely related species with the same ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) large subunit (rbcL) but contrasting hydraulic traits and mesophyll structure occurring in the Iberian Peninsula under contrasting conditions. As photosynthesis and hydraulic capacities often co‐scale, we hypothesize that these species differ in mesophyll conductance to CO₂ (g_m). g_m and key anatomical traits were measured in both species. Drought‐adapted population of A. pinsapo has higher photosynthesis than the more mesic population of A. alba, in agreement with its higher hydraulic capacity. However, A. alba exhibits the largest stomatal conductance (g_s), and so water use efficiency (WUE) is much higher in A. pinsapo. The differences in photosynthesis were explained by differences in g_m, indicating a correlation between hydraulic capacity and g_m. We report a case where g_m is the main factor limiting photosynthesis in one species (A. alba) when compared with the other one (A. pinsapo). The results also highlight the discrepancy between g_m estimates based on anatomical measurements and those based on gas exchange methods, probably due to the very large resistance exerted by cell walls and the stroma in both species. Thus, the cell wall and chloroplast properties in relation to CO₂ diffusion constitute a near‐future research priority.     

Evolution of protein transport to the chloroplast envelope membranes

Comparing with other angiosperms, most members within the family Orchidaceae have lower photosynthetic capacities. However, the underlying mechanisms remain unclear. Cypripedium and Paphiopedilum are closely related phylogenetically in Orchidaceae, but their photosynthetic performances are different. We explored the roles of internal anatomy and diffusional conductance in determining photosynthesis in three Cypripedium and three Paphiopedilum species, and quantitatively analyzed their diffusional and biochemical limitations to photosynthesis. Paphiopedilum species showed lower light-saturated photosynthetic rate (A _N), stomatal conductance (g _s), and mesophyll conductance (g _m) than Cypripedium species. A _N was positively correlated with g _s and g _m. And yet, in both species A _N was more strongly limited by g _m than by biochemical factors or g _s. The greater g _s of Cypripedium was mainly affected by larger stomatal apparatus area and smaller pore depth, while the less g _m of Paphiopedilum was determined by the reduced surface area of mesophyll cells and chloroplasts exposed to intercellular airspace per unit of leaf area, and much thicker cell wall thickness. These results suggest that leaf anatomical structure is the key factor affecting g _m, which is largely responsible for the difference in photosynthetic capacity between those two genera. Our findings provide new insight into the photosynthetic physiology and functional diversification of orchids.     

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.     

Abscisic Acid Induces Rapid Reductions in Mesophyll Conductance to Carbon Dioxide

The rate of photosynthesis (A) of plants exposed to water deficit is a function of stomatal (g_s) and mesophyll (g_m) conductance determining the availability of CO₂ at the site of carboxylation within the chloroplast. Mesophyll conductance often represents the greatest impediment to photosynthetic uptake of CO₂, and a crucial determinant of the photosynthetic effects of drought. Abscisic acid (ABA) plays a fundamental role in signalling and co-ordination of plant responses to drought; however, the effect of ABA on g_m is not well-defined. Rose, cherry, olive and poplar were exposed to exogenous ABA and their leaf gas exchange parameters recorded over a four hour period. Application with ABA induced reductions in values of A, g_s and g_m in all four species. Reduced g_m occurred within one hour of ABA treatment in three of the four analysed species; indicating that the effect of ABA on g_m occurs on a shorter timescale than previously considered. These declines in g_m values associated with ABA were not the result of physical changes in leaf properties due to altered turgor affecting movement of CO₂, or caused by a reduction in the sub-stomatal concentration of CO₂ (C_i). Increased [ABA] likely induces biochemical changes in the properties of the interface between the sub-stomatal air-space and mesophyll layer through the actions of cooporins to regulate the transport of CO₂. The results of this study provide further evidence that g_m is highly responsive to fluctuations in the external environment, and stress signals such as ABA induce co-ordinated modifications of both g_s and g_m in the regulation of photosynthesis.     

Seasonal relationships between leaf nitrogen content (photosynthetic capacity) and leaf canopy light exposure in peach (Prunus persica)

Abstract Field gas exchange measurements on intact peach (Prunus persica (L.) Batsch) leaves indicate that leaf nitrogen content (N_L) and leaf weight per unit leaf area (W_a) are highly correlated with CO₂ assimilation rate (A) and mesophyll conductance (g_m). Therefore, N_L and W_a were used to study seasonal relationships between leaf carboxylation capacity and natural light exposure in tree canopies. From mid-season onwards, N_L and W_a were linearly correlated with light exposure expressed as the amount of time during a clear day that a leaf was exposed to a photosynthetic photon flux density (Q) of ≥ 100 μmol m^?2 s^?1. The data support the hypothesis that whole-tree photosynthesis is optimized by partitioning of photosynthetic capacity among leaves in deciduous tree canopies with respect to natural light exposure.     

Rice varietal differences in responses of stomatal gas exchange to supplemental nitrogen application

In the field, supplemental application of N fertilizer to rice (Oryza sativa) shortly before the beginning of heading stage increases leaf N content and enhances photosynthesis during the grain-filling period. In search of varietal differences in leaf gas exchange in response to supplemental N application, we examined 13 rice varieties grown in the field during two successive years. The varieties included japonica and indica varieties, both of which are widely grown in Japan. The response to supplemental N application could not be separated clearly between variety groups; some of the japonica varieties, but none of the indica varieties, exhibited significant increase in stomatal conductance (g _s) after supplemental N application. Supplemental N was more effective to increase stomatal aperture in the varieties with inherently lower g _s. Varieties that showed greater response of g _s to supplemental N application might be able to adjust their stomatal aperture with appropriate N control. Although the internal-to-ambient CO₂ mole fraction ratio and the leaf carbon isotopic composition (δ¹³C) differed among varieties as a result of variations in stomatal aperture and the CO₂ requirement of mesophyll, supplemental N application barely influenced these parameters, because it only moderately affected stomatal aperture. Since δ¹³C tended to increase with increasing number of days from transplantation to heading stage in japonica varieties, δ¹³C values were more sensitive to differences in growth rate between years than to N application.     

The physiological basis for genetic variation in water use efficiency and carbon isotope composition in Arabidopsis thaliana

Ecologists and physiologists have documented extensive variation in water use efficiency (WUE) in Arabidopsis thaliana, as well as association of WUE with climatic variation. Here, we demonstrate correlations of whole-plant transpiration efficiency and carbon isotope composition (δ¹³C) among life history classes of A. thaliana. We also use a whole-plant cuvette to examine patterns of co-variation in component traits of WUE and δ¹³C. We find that stomatal conductance (g _s) explains more variation in WUE than does A. Overall, there was a strong genetic correlation between A and g _s, consistent with selection acting on the ratio of these traits. At a more detailed level, genetic variation in A was due to underlying variation in both maximal rate of carboxylation (V _cmax) and maximum electron transport rate (Jmax). We also found strong effects of leaf anatomy, where lines with lower WUE had higher leaf water content (LWC) and specific leaf area (SLA), suggesting a role for mesophyll conductance (g _m) in variation of WUE. We hypothesize that this is due to an effect through g _m, and test this hypothesis using the abi4 mutant. We show that mutants of ABI4 have higher SLA, LWC, and g _m than wild-type, consistent with variation in leaf anatomy causing variation in g _m and δ¹³C. These functional data also add further support to the central, integrative role of ABI4 in simultaneously altering ABA sensitivity, sugar signaling, and CO₂ assimilation. Together our results highlight the need for a more holistic approach in functional studies, both for more accurate annotation of gene function and to understand co-limitations to plant growth and productivity.     

Diffusional conductances to CO2 as a target for increasing photosynthesis and photosynthetic water-use efficiency

A key objective for sustainable agriculture and forestry is to breed plants with both high carbon gain and water-use efficiency (WUE). At the level of leaf physiology, this implies increasing net photosynthesis (A _N) relative to stomatal conductance (g _s). Here, we review evidence for CO₂ diffusional constraints on photosynthesis and WUE. Analyzing past observations for an extensive pool of crop and wild plant species that vary widely in mesophyll conductance to CO₂ (g _m), g _s, and foliage A _N, it was shown that both g _s and g _m limit A _N, although the relative importance of each of the two conductances depends on species and conditions. Based on Fick’s law of diffusion, intrinsic WUE (the ratio A _N/g _s) should correlate on the ratio g _m/g _s, and not g _m itself. Such a correlation is indeed often observed in the data. However, since besides diffusion A _N also depends on photosynthetic capacity (i.e., V _c,max), this relationship is not always sustained. It was shown that only in a very few cases, genotype selection has resulted in simultaneous increases of both A _N and WUE. In fact, such a response has never been observed in genetically modified plants specifically engineered for either reduced g _s or enhanced g _m. Although increasing g _m alone would result in increasing photosynthesis, and potentially increasing WUE, in practice, higher WUE seems to be only achieved when there are no parallel changes in g _s. We conclude that for simultaneous improvement of A _N and WUE, genetic manipulation of g _m should avoid parallel changes in g _s, and we suggest that the appropriate trait for selection for enhanced WUE is increased g _m/g _s.     

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.     

Stomatal and nonstomatal limitations of photosynthesis in 19 temperate tree species on contrasting sites during wet and dry years

A unique approach was used to evaluate stomatal and nonstomatal constraints to photosynthesis in 19 naturally occurring, deciduous tree species on xeric, mesic and wetmesic sites in central Pennsylvania, USA, during relatively wet (1990) and dry (1991) growing seasons. All species exhibited significantly decreased stomatal conductance to CO₂ (g_c) in 1991 compared to 1990. The mesic species had drought related decreases in photosynthesis (A) attributed primarily to increased absolute stomatal limitation to A (L_g), whereas in the wet-mesic species, the absolute mesophyll limitation (Lm) was at least as important as L_g in limiting A during drought. The xeric species maintained relatively high A during drought despite decreased g_c. In the xeric and mesic species, L_m decreased and L_g increased during drought due to stomatal closure. From xeric to mesic to wet-mesic, the relative stomatal limitation (I_g) generally decreased faster, and relative mesophyll limitations to A increased faster, with increasing g_c suggesting greater photosynthetic capacity (i.e. greater potential maximum A) with increasing drought tolerance rank of species. Few species exhibited a significant drought-related decrease in photosynthetic capacity. The results of this landscape-based study indicate that the interaction of stomatal and nonstomatal limitations of A vary in a manner consistent with species' drought tolerance and site conditions, and that nonstomatal constraints to A in field plants during a moderate, season-long drought were generally not as severe as reported in controlled studies.     

Isoprenoid emissions,photosynthesis and mesophyll diffusion conductance in response to blue light

The effects of blue light (BL) on leaf gas exchange of Populus × canadensis, a strong isoprene emitter, and Quercus ilex and Citrus reticulata, two monoterpene emitters with respectively small and large storage pools for monoterpenes, were studied. Leaves were initially exposed to a saturating photosynthetic photon flux density (PPFD) of white light (WL), which was then progressively reduced to perform WL-response curves. Leaves acclimated to saturating WL were then quickly exposed to equivalent BL levels to perform BL-response curves. Blue light did not significantly affect photosynthetic parameters in the light-limited portion of the PPFD-response curves in both P. × canadensis and Q. ilex. Whereas photosynthesis (A), stomatal conductance (g_s), and mesophyll conductance (g_m) were significantly decreased at high PPFDs of BL. A was similarly inhibited by BL in C. reticulata, but there was no significant effect of light quality on g_s. Overall these results show that the negative effect of BL on photosynthesis is widespread in tree species with different leaf characteristics, and that this involves coordinated reductions in g_s and g_m. BL negatively affected isoprene emission and, to a lesser extent monoterpene emissions, in concert with photosynthetic inhibition. Interesting, both isoprene and monoterpene emissions were shown to be inversely dependent upon intercellular [CO₂]. These results indicate that a change in light spectral quality, which can vary during the day, between days and within seasons, can alter photosynthesis and isoprenoid emissions, depending on the PPFD intensity. Such effects should be strongly considered in photosynthesis and volatile isoprenoid emission models.