Interactions of Elevated CO2 Concentration and Drought Stress on Photosynthesis in Eucalyptus Cladocalyx F. Muell

Response of net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (c _i), and photosynthetic efficiency (F_v/F_m) of photosystem 2 (PS2) was assessed in Eucalyptus cladocalyx grown for long duration at 800 (C₈₀₀) or 380 (C₃₈₀) µmol mol^-1 CO₂ concentration under sufficient water supply or under water stress. The well-watered plants at C₈₀₀ showed a 2.2 fold enhancement of P _N without any change in g _s. Under both C₈₀₀ and C₃₈₀, water stress decreased P _N and g _s significantly without any substantial reduction of c _i, suggesting that both stomatal and non-stomatal factors regulated P _N. However, the photosynthetic efficiency of PS2 was not altered.     

On the Use of Leaf Spectral Indices to Assess Water Status and Photosynthetic Limitations in Olea europaea L. during Water-Stress and Recovery

Diffusional limitations to photosynthesis, relative water content (RWC), pigment concentrations and their association with reflectance indices were studied in olive (Olea europaea) saplings subjected to water-stress and re-watering. RWC decreased sharply as drought progressed. Following rewatering, RWC gradually increased to pre-stress values. Photosynthesis (A), stomatal conductance (g_s), mesophyll conductance (g_m), total conductance (g_t), photochemical reflectance index (PRI), water index (WI) and relative depth index (RDI) closely followed RWC. In contrast, carotenoid concentration, the carotenoid to chlorophyll ratio, water content reflectance index (WCRI) and structural independent pigment index (SIPI) showed an opposite trend to that of RWC. Photosynthesis scaled linearly with leaf conductance to CO₂; however, A measured under non-photorespiratory conditions (A_1%O2) was approximately two times greater than A measured at 21% [O₂], indicating that photorespiration likely increased in response to drought. A_1%O2 also significantly correlated with leaf conductance parameters. These relationships were apparent in saturation type curves, indicating that under non-photorespiratory conditions, CO₂ conductance was not the major limitations to A. PRI was significant correlated with RWC. PRI was also very sensitive to pigment concentrations and photosynthesis, and significantly tracked all CO₂ conductance parameters. WI, RDI and WCRI were all significantly correlated with RWC, and most notably to leaf transpiration. Overall, PRI correlated more closely with carotenoid concentration than SIPI; whereas WI tracked leaf transpiration more effectively than RDI and WCRI. This study clearly demonstrates that PRI and WI can be used for the fast detection of physiological traits of olive trees subjected to water-stress.     

Effects of elevated CO2, drought and temperature on the water relations and gas exchange of groundnut (Arachis hypogaea) stands grown in controlled environment glasshouses

Stands of groundnut (Arachis hypogaea L. cv. Kadiri‐3) were grown in controlled environment glasshouses at mean atmospheric CO₂ concentrations of 375 or 700 μmol mol^?1 and daily mean air temperatures of 28 or 32°C on irrigated or drying soil profiles. Leaf water (Ψ_l) and solute potential (Ψ_s), relative water content (RWC), stomatal conductance (g_l) and net photosynthesis (P_n) were measured at midday for the youngest mature leaf throughout the growing season. Elevated CO₂ and temperature had no detectable effect on the water relations of irrigated plants, but higher values of RWC, Ψ_l and Ψ_s were maintained for longer under elevated CO₂ during progressive drought. Turgor potential (Ψ_p) reached zero when Ψ_l declined to ?1.6 to ?1.8 MPa in all treatments; turgor was lost sooner when droughted plants were grown under ambient CO₂. A 4°C increase in mean air temperature had no effect on Ψ_s in droughted plants, but elicited a small increase in Ψ_l; midday g_l values were lower under elevated than under ambient CO₂, and Ψ_l and g_l declined below ?1.5 MPa and 0.25 cm s^?1, respectively, as the soil dried. Despite the low g_l values recorded for droughted plants late in the season, P_n was maintained under elevated CO₂, but declined to zero 3 weeks before final harvest under ambient CO₂. Concurrent reductions in g_l and increases in water use efficiency under elevated CO₂ prolonged photosynthetic activity during drought and increased pod yields relative to plants grown under ambient CO₂. The implications of future increases in atmospheric CO₂ for the productivity of indeterminate C₃ crops grown in rainfed subsistence agricultural systems in the semi‐arid tropics are discussed.     

Photorespiration and photoprotection of grapevine (<Emphasis Type="Italic">Vitis vinifera</Emphasis> L. cv. Cabernet Sauvignon) under water stress

In order to investigate the photoprotective function of photorespiration in grapevine under water stress, potted grapevines (Vitis vinifera L. cv. Cabernet Sauvignon) were randomly divided into three uniform groups for well-watered [watered every morning to keep the relative water content (RWC) of soil over 70 %], water-stress adapted (drought-adapted at 30 % relative soil water content for 30 days), and water stress without adaptation treatment (water-stressed to 30 % relative soil water content for 3 days). Net assimilation rate (A _N), stomatal conductance (g _s), substomatal CO₂ concentration (C _i), transpiration rate (E), actual photochemical efficiency of PSII (Φ_PSII), and maximum photochemical efficiency of PSII (F_v/F_m) were recorded by combining measurements of gas exchange and chlorophyll fluorescence. Gross photorespiration (P_r), photosynthetic electron partitioning (J_C/J_T), photochemical quenching coefficient (qP), and non-photochemical quenching (NPQ) were also calculated. The ratio of net assimilation rate to transpiration rate (A _N/E) was used as an indicator of water use efficiency (WUE). A _N, apparent P_r, Φ_PSII, F_v/F_m, qp, and g _s decreased, NPQ increased, and gross P_r sustained at a high level under water stress. This suggests that both photorespiration and energy dissipation play important roles in protecting photosynthetic apparatus against photoinhibition. C _i in water-stressed plants without adaptation treatment increased, which indicates the leaves suffered a non-stomatal limitation, while the water-stress adaped plants only suffered a stomatal limitation indicated by low C _i.     

Gas Exchange of Spring Barley and Wheat Grown under Mild Water Shortage

During mild water stress (decrease of full water capacity from 60 to 35 %) net photosynthetic rate (P _N) of four spring barley and wheat genotypes was about twice lower than that for unstressed plants and was mainly limited by non-stomatal factors. Availability of CO₂ from intercellular spaces did not change significantly when stomatal conductance (g _s) decreased from 0.25-0.35 to 0.15-0.20 mol(H₂O) m⁻² s⁻¹. There may be two main processes leading to similar intercellular CO₂ concentration (c _i) in stressed and unstressed seedlings despite of twice lower P _N under mild water stress: (a) lower diffusion of CO₂ through stomata represented by lower g _s, (b) lower consumption of CO₂ by photosynthetic apparatus of stressed plants. Last factor is partially pronounced by lower response of P _N to c _i observed for stressed than for control plants. This revised version was published online in August 2006 with corrections to the Cover Date.     

Interactions of Elevated CO2 Concentration and Drought Stress on Photosynthesis in Eucalyptus Cladocalyx F. Muell

Response of net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (c _i), and photosynthetic efficiency (F_v/F_m) of photosystem 2 (PS2) was assessed in Eucalyptus cladocalyx grown for long duration at 800 (C₈₀₀) or 380 (C₃₈₀) μmol mol^-1 CO₂ concentration under sufficient water supply or under water stress. The well-watered plants at C₈₀₀ showed a 2.2 fold enhancement of P _N without any change in g _s. Under both C₈₀₀ and C₃₈₀, water stress decreased P _N and g _s significantly without any substantial reduction of c _i, suggesting that both stomatal and non-stomatal factors regulated P _N. However, the photosynthetic efficiency of PS2 was not altered. This revised version was published online in June 2006 with corrections to the Cover Date.     

Free-air CO2 enrichment in a wheat-weed assembly – effects on water relations

In a three-year free-air CO₂ enrichment study (Mini-FACE), spring wheat associated with typical arable weeds were grown under present and elevated atmospheric carbon dioxide concentrations [CO₂] (ambient air+150 μmol mol^?1). Analyses of plant stable carbon isotope ratios and in vivo measurements of leaf gas exchange were used to describe the CO₂ effects on water relations. For most species examined elevated [CO₂] significantly increased the intrinsic water-use efficiency (A/g_s) as derived from carbon isotope analyses. In some of the species, seasonal averages of the ratio between leaf internal to atmospheric CO₂ (c_i/c_a) were found to be significantly reduced by elevated [CO₂]. Periodic leaf gas exchange measurements confirmed the increased water-use efficiency, but significant CO₂ effects became evident only over the entire season by carbon isotope analysis. In both types of analysis conducted, spring wheat was found to react significantly different from all other species examined. The relation between A/g_s and biomass production was significantly influenced by elevated [CO₂] in all three years of the study. At the end of the drier growing seasons 2003 and 2004, the soil water content tended to be increased in the CO₂ enriched plots indicating a water saving effect. These observations demonstrate the impact of elevated [CO₂] on plant water relations with a likely positive feedback leading to higher soil water availability. Due to the differences in the CO₂ responses of spring wheat compared to the weeds we suggest that rising [CO₂] may cause shifts in the species composition of crop-weed communities.     

Stomatal acclimation to increased CO2 concentration in a Florida scrub oak species Quercus myrtifolia Willd

Native scrub‐oak communities in Florida were exposed for three seasons in open top chambers to present atmospheric [CO₂] (approx. 350 μmol mol^?1) and to high [CO₂] (increased by 350 μmol mol^?1). Stomatal and photosynthetic acclimation to high [CO₂] of the dominant species Quercus myrtifolia was examined by leaf gas exchange of excised shoots. Stomatal conductance (g_s) was approximately 40% lower in the high‐ compared to low‐[CO₂]‐grown plants when measured at their respective growth concentrations. Reciprocal measurements of g_s in both high‐ and low‐[CO₂]‐grown plants showed that there was negative acclimation in the high‐[CO₂]‐grown plants (9–16% reduction in g_s when measured at 700 μmol mol^?1), but these were small compared to those for net CO₂ assimilation rate (A, 21–36%). Stomatal acclimation was more clearly evident in the curve of stomatal response to intercellular [CO₂] (c_i) which showed a reduction in stomatal sensitivity at low c_i in the high‐[CO₂]‐grown plants. Stomatal density showed no change in response to growth in high growth [CO₂]. Long‐term stomatal and photosynthetic acclimation to growth in high [CO₂] did not markedly change the 2·5‐ to 3‐fold increase in gas‐exchange‐derived water use efficiency caused by high [CO₂].     

Direct and acclimatory responses of stomatal conductance to elevated carbon dioxide in four herbaceous crop species in the field

In order to separate the net effect of growth at elevated [CO₂] on stomatal conductance (g_s) into direct and acclimatory responses, mid‐day values of g_s were measured for plants grown in field plots in open‐topped chambers at the current ambient [CO₂], which averaged 350 μmol mol^?1 in the daytime, and at ambient + 350 μmol mol^?1[CO₂] for winter wheat, winter barley, potato and sorghum. The acclimatory response was determined by comparing g_s measured at 700 μmol mol^?1[CO₂] for plants grown at the two [CO₂]. The direct effect of increasing [CO₂] from 350 to 700 μmol mol^?1 was determined for plants grown at the lower concentration. Photosynthetic rates were measured concurrently with g_s. For all species, growth at the higher [CO₂] significantly reduced g_s measured at 700 μmol mol^?1[CO₂]. The reduction in g_s caused by growth at the higher [CO₂] was larger for all species on days with low leaf to air water vapour pressure difference for a given temperature, which coincided with highest conductances and also the smallest direct effects of increased [CO₂] on conductance. For barley, there was no other evidence for stomatal acclimation, despite consistent down‐regulation of photosynthetic rate in plants grown at the higher [CO₂]. In wheat and potato, in addition to the vapour pressure difference interaction, the magnitude of stomatal acclimation varied directly in proportion to the magnitude of down‐regulation of photosynthetic rate through the season. In sorghum, g_s consistently exhibited acclimation, but there was no down‐regulation of photosynthetic rate. In none of the species except barley was the direct effect the larger component of the net reduction in g_s when averaged over measurement dates. The net effect of growth at elevated [CO₂] on mid‐day g_s resulted from unique combinations of direct and acclimatory responses in the various species.     

Low concentrations of glycine inhibit photorespiration and enhance the net rate of photosynthesis in <Emphasis Type="Italic">Caragana korshinskii</Emphasis>

The inhibition of photorespiration can be used to improve plant carbon fixation. In order to compare the effects of three photorespiration inhibitors [glycine, NaHSO₃, and isonicotinyl hydrazide (INH)], photosynthetic parameters of leaves sprayed respectively with these chemicals were examined and their inhibiting efficiency was evaluated in Caragana korshinskii. Our results showed that 5 mM glycine could reduce the photorespiratory rate (P_R) effectively, while the net photosynthetic rate (P_N), stomatal conductance (g_s), and intercellular CO₂ concentration (Ci) significantly increased. The ratio of electron flow for ribulose-1,5-bisphosphate (RuBP) carboxylation to RuBP oxygenation was elevated markedly. NaHSO₃ and INH could also suppress the PR in some cases, whereas PN was not improved. The glyoxylate content increased considerably after application of low concentrations of glycine. These results suggested that low concentrations of glycine could suppress photorespiration by feed-back inhibition of glyoxylate and enhance photosynthesis by regulating g_s, C_i, and the distribution of electron flow in C. korshinskii.     

Photosynthesis and Growth Responses of Parthenocissus quinquefolia (L.) Planch to Soil Water Availability

Photosynthesis and growth characteristics of Parthenocissus quinquefolia were measured under differing soil water availability within a pot. Decreased soil moisture significantly reduced the leaf relative water content (RWC) and the above- and below-ground biomass. However, more biomass was allocated to the root than to the leaf. Net photosynthetic rate (P _N), stomatal conductance (g _s), and transpiration rate (E) were also significantly decreased but water use efficiency (WUE) was increased. Midday depressions in P _N and g _s were not evident for the well-irrigated plants. With the lower water availability, midday reductions in P _N and g _s were much more marked and the duration of the depression was longer. Additionally, the P _N-irradiance response curves also indicated that water supply affected photosynthesis capacity. The growth and photosynthetic response of P. quinquefolia to water supply indicated that this species could resilient to water availabilities and adapt to Hunshandak conditions very well.     

Experimental and simulated light responses of photosynthesis in leaves of three tree species under different soil water conditions

Water deficit is one of the major limiting factors in vegetation recovery and restoration in loess, hilly-gully regions of China. The light responses of photosynthesis in leaves of two-year old Prunus sibirica L., Hippophae rhamnoides L., and Pinus tabulaeformis Carr. under various soil water contents were studied using the CIRAS-2 portable photosynthesis system. Light-response curves and photosynthetic parameters were analyzed and fitted using the rectangular hyperbola model, the exponential model, the nonrectangular hyperbola model, and the modified rectangular hyperbola model. Under high light, photosynthetic rate (P _N) and stomatal conductance (g _s) were steady and photoinhibition was not significant, when the relative soil water content (RWC) varied from 56.3–80.9%, 47.9–82.9%, and 33.4–92.6% for P. sibirica, H. rhamnoides, and P. tabulaeformis, respectively. The light-response curves of P _N, the light compensation point (LCP), and the dark respiration rate (R _D) were well fitted using the above four models. The nonrectangular hyperbola was the best model in fitting the data; the modified rectangular hyperbola model was the second, and the rectangular hyperbola model was the poorest one. When RWC was higher or lower than the optimal range, the obvious photoinhibition and significant decrease in P _N with increasing photosynthetic photon flux density (PPFD) were observed in all three species under high light. The light saturation point (LSP) and apparent quantum yield also decreased significantly, when the upper limit of PPFD was 200 μmol m^?2 s^?1. Under these circumstances, only the modified rectangular hyperbola model was able to fit well the curves of the light response, LCP, LSP, R _D, and light-saturated P _N.     

On the complementary relationship between marginal nitrogen and water-use efficiencies among Pinus taeda leaves grown under ambient and CO2-enriched environments

Background and Aims

Water and nitrogen (N) are two limiting resources for biomass production of terrestrial vegetation. Water losses in transpiration (E) can be decreased by reducing leaf stomatal conductance (g_s) at the expense of lowering CO₂ uptake (A), resulting in increased water-use efficiency. However, with more N available, higher allocation of N to photosynthetic proteins improves A so that N-use efficiency is reduced when g_s declines. Hence, a trade-off is expected between these two resource-use efficiencies. In this study it is hypothesized that when foliar concentration (N) varies on time scales much longer than g_s, an explicit complementary relationship between the marginal water- and N-use efficiency emerges. Furthermore, a shift in this relationship is anticipated with increasing atmospheric CO₂ concentration (c_a).

Methods

Optimization theory is employed to quantify interactions between resource-use efficiencies under elevated c_a and soil N amendments. The analyses are based on marginal water- and N-use efficiencies, λ = (∂A/∂g_s)/(∂E/∂g_s) and η = ∂A/∂N, respectively. The relationship between the two efficiencies and related variation in intercellular CO₂ concentration (c_i) were examined using A/c_i curves and foliar N measured on Pinus taeda needles collected at various canopy locations at the Duke Forest Free Air CO₂ Enrichment experiment (North Carolina, USA).

Key Results

Optimality theory allowed the definition of a novel, explicit relationship between two intrinsic leaf-scale properties where η is complementary to the square-root of λ. The data support the model predictions that elevated c_a increased η and λ, and at given c_a and needle age-class, the two quantities varied among needles in an approximately complementary manner.

Conclusions

The derived analytical expressions can be employed in scaling-up carbon, water and N fluxes from leaf to ecosystem, but also to derive transpiration estimates from those of η, and assist in predicting how increasing c_a influences ecosystem water use.     

Causes of decreased photosynthetic rate and metabolic capacity in water-deficient leaf cells: a critical evaluation of mechanisms and integration of processes

Background

Water deficit (WD) decreases photosynthetic rate (A) via decreased stomatal conductance to CO₂ (g_s) and photosynthetic metabolic potential (A_pot). The relative importance of g_s and A_pot, and how they are affected by WD, are reviewed with respect to light intensity and to experimental approaches.

Scope and Conclusions

With progressive WD, A decreases as g_s falls. Under low light during growth and WD, A is stimulated by elevated CO₂, showing that metabolism (A_pot) is not impaired, but at high light A is not stimulated, showing inhibition. At a given intercellular CO₂ concentration (C_i) A decreases, showing impaired metabolism (A_pot). The C_i and probably chloroplast CO₂ concentration (C_c), decreases and then increases, together with the equilibrium CO₂ concentration, with greater WD. Estimation of C_c and internal (mesophyll) conductance (g_i) is considered uncertain. Photosystem activity is unaffected until very severe WD, maintaining electron (e⁻) transport (ET) and reductant content. Low A, together with photorespiration (PR), which is maintained or decreased, provides a smaller sink for e⁻_, causing over-energization of energy transduction. Despite increased non-photochemical quenching (NPQ), excess energy and e⁻ result in generation of reactive oxygen species (ROS). Evidence is considered that ROS damages ATP synthase so that ATP content decreases progressively with WD. Decreased ATP limits RuBP production by the Calvin cycle and thus A_pot. Rubisco activity is unlikely to determine A_pot. Sucrose synthesis is limited by lack of substrate and impaired enzyme regulation. With WD, PR decreases relative to light respiration (R_L), and mitochondria consume reductant and synthesise ATP. With progressing WD at low A, R_L increases C_i and C_c. This review emphasises the effects of light intensity, considers techniques, and develops a qualitative model of photosynthetic metabolism under WD that explains many observations: testable hypotheses are suggested.Key words: Water stress, photosynthesis, photorespiration, stomata, ATP synthase, ATP, photoinhibition, electron transport, Rubisco, fluorescence, sucrose, mesophyll conductance     

Effects of Drought on Photosynthetic Performance and Water Relations of Four Vigna Genotypes

The effect of drought on plant water relations and photosynthesis of Vigna glabrescens (Vg) and Vigna unguiculata (cvs. 1183, EPACE-1 and Lagoa), which differ in their drought resistance, was compared. With the increase of drought severity, Vg showed a more gradual stomatal closure and maintained significantly higher levels of stomatal conductance (g_s) and photosynthetic activity (P_N) than the other genotypes even when minimum relative water content (RWC) values were observed. Furthermore, Vg was the only genotype able to accumulate significant amounts of proline already under moderate water deficit, what could explain the lower osmotic potential (ψ_s) values observed in these plants. The three V. unguiculata cultivars presented a similar stomatal control under increasing water deficit. A mesophyllic impairment of photosynthetic capacity (P_max) was detected for cv. 1183 from the beginning of drought onset (85-75 % RWC) while in the Vg plants the values remained unaffected along the whole drought period, indicating that P_N decrease observed in this genotype is mainly a consequence of stomatal closure. Such P_max maintenance suggests the existence of a high mesophyllic ability to cope with increasing tissue dehydration in Vg. This revised version was published online in September 2006 with corrections to the Cover Date.     

Exogenous application of trehalose induced H2O2 production and stomatal closure in Vicia faba

Trehalose can reduce stomatal aperture by a hydrogen-peroxide-dependent pathway in Vicia faba L. (cv. Daqingpi) resulting in significantly lower values of net photosynthetic rate (P_N), stomatal conductance (g_s), and transpiration rate (E). At 8 and 24 h, the lower P_N in trehalose-treated plants was accompanied by significant decrease in intercellular CO₂ concentration (c_i) suggesting that the reduction of P_N was caused by stomatal limitation. At 48 and 72 h, trehalose decreased apparent carboxylation efficiency (P_N/c_i) and did not decrease c_i and g_s compared with controls; therefore the reduction in photosynthesis was caused by non-stomatal limitation. Trehalose treatment resulted in significantly higher effective photochemical efficiency of PS II (Φ_PSII) and did not affect maximum photochemical efficiency of PS II (F_v/F_m). At 24, 48, and 72 h, trehalose decreased non-photochemical quenching (NPQ) and increased photochemical quenching (qP). Our results suggest that trehalose did not damage photosynthetic reaction centers.     

Soil drying and its effect on leaf conductance and CO2 assimilation of Vigna unguiculata (L.) Walp

Summary Well watered plants of Vigna unguiculata (L.) Walp cv. California Blackeye No. 5 had maximum photosynthetic rates of 16 mol m^-2 s^-1 (at ambient CO₂ concentration and environmental parameters optimal for high CO₂ uptake). Leaf conductance declined with increasing water vapour concentration difference between leaf and air (w), but it increased with increasing leaf temperature at a constant small w. When light was varied, CO₂ assimilation and leaf conductance were correlated linearly. We tested the hypothesis that g was controlled by photosynthesis via intercellular CO₂ concentration (c _i). No unique relationship between (1) c _i, (2) the difference between ambient CO₂ concentration (c _a) and c _i, namely c _a-c _i, or (3) the c _i/c _a ratio and g was found. g and A appeared to respond to environmental factors fairly independently of each other. The effects of different rates of soil drying on leaf gas exchange were studied. At unchanged air humidity, different rates of soil drying were produced by using (a) different soils, (b) different irrigation schemes and (c) different soil volumes per plant. Although the soil dried to wilting point the relative leaf water content was little affected. Different soil drying rates always resulted in the same response of photosynthetic capacity (A _max) and corresponding leaf conductance (g(_Amax)) when plotted against percent relative plant-extractable soil water content (W _e %) but the relationship with relative soil water content (W _e ) was less clear. Above a range of W _e of 15%–25%, A _max and g(_Amax) were both high and responded little to decreasing W _e . As soon as W _e fell below this range, A _max and g(_Amax) declined. The data suggest root-to-leaf communication not mediated via relative leaf water content. However, g(_Amax) was initially more affected than A _max.List of abbreviations A CO₂ assimilation - A _max photosynthetic capacity at favourable ambient conditions - c _a CO₂ concentration of the air in the leaf chamber - c _i intercellular - CO₂ concentration - E transpiration - g leaf conductance - g(_Amax) leaf conductance corresponding to photosynthetic capacity - I photon flux rate - T _l leaf temperature - W _e relative plant-extractable soil water content - W _e absolute plant-extractable soil water content - W _l relative leaf water content - W _s relative soil water content - w difference in water vapour mole fraction between leaf and air - leaf water potential     

Altitudinal variation in photosynthetic capacity, diffusional conductance and δ13C of butterfly bush (Buddleja davidii) plants growing at high elevations

In this study, we have examined several physiological, biochemical and morphological features of Buddleja davidii plants growing at 1300 m above sea level (a.s.l.) and 3400 m a.s.l., respectively, to identify coordinated changes in leaf properties in response to reduced CO₂ partial pressure (P_a). Our results confirmed previous findings that foliar δ¹³C, photosynthetic capacity and foliar N concentration on a leaf area basis increased, whereas stomatal conductance (g_s) decreased with elevation. The net CO₂ assimilation rate (A_max), maximum rate of electron transport (J_max) and respiration increased significantly with elevation, although no differences were found in carboxylation efficiency of Rubisco (V_cmax). Consequently, also the J_max to V_cmax ratio was significantly increased by elevation, indicating that the functional balance between Ribulose‐1,5‐biphosphate (RuBP) consumption and RuBP regeneration changes as elevation increases. Our results also indicated a homeostatic response of CO₂ transfer conductance inside the leaf (mesophyll conductance, g_m) to increasing elevation. In fact, with elevation, g_m also increased compensating for the strong decrease in g_s and, thus, in the P_i (intercellular partial pressure of CO₂) to P_a ratio, leading to similar chloroplast partial pressure of CO₂ (P_c) to P_a ratio at different elevations. Because there were no differences in V_cmax, also A measured at similar PPFD and leaf temperature did not differ statistically with elevation. As a consequence, a clear relationship was found between A and g_m, and between A and the sum of g_s and g_m. These data suggest that the higher dry mass δ¹³C of leaves at the higher elevation, indicative of lower long‐term P_c/P_a ratio, cannot be attributed to changes either in diffusional resistances or in carboxylation efficiency. We speculate that because temperature significantly decreases as the elevation increases, it dramatically affects CO₂ diffusion and hence P_c/P_a and, consequently, is the primary factor influencing ¹³C discrimination at high elevation.     

Leaf anatomy and photosynthetic carbon metabolic characteristics in <Emphasis Type="Italic">Phragmites communis</Emphasis> in different soil water availability

To investigate the variations of anatomical and photosynthetic carbon metabolic characteristics within one species in response to increasing soil water stress, leaf anatomical characteristics, gas exchange and the activity of key enzymes in photosynthesis and photorespiration were compared in different ecotypes of Phragmites communis growing in an oasis-desert transitional zone (ODTZ) from swamp habitat (plot 1–3) via heavy salt meadow (plot 4–7) and light salt meadow habitat (plot 8–9) to dune habitat (plot 10–13) in Northwest China. The results showed that interveinal distance (ID) decreased with increasing water stress except that in plots of dune reed (DR). Vein mean diameter (VMD) in plot 10, 11 and 12 of the DR was significantly larger than that in other ecotypes. Leaf specific porosity (LSP) enhanced from plot 4 to plot 13 from heave salt meadow reed (HSMR) to light salt meadow reed (LSMR) and to DR. Chlorophyll fluorescence in bundle sheath cells were microscopically found in four ecotypes, especially significantly in the DR. Net CO₂ assimilation rate (A _n) dropped rapidly from the swamp reed (SR) to the HSMR and then increased progressively from the LSMR to the DR. Stomatal conductance (g _s) decreased and the water use efficiency (WUE) rose from the wet to the dry ecotypes. Sensitivity of g _s to intercellular CO₂ concentration (C _i) increased, but glycolate oxidase (GO) activity gradually reduced with increasing soil water deficiency. The RuBPCase activity did not reduce in four ecotypes even in DR, but, the PEPCase and NAD-ME activities as well as the ratio of PEPCase/RuBPCase were gradually enhanced with increasing soil water stress. We concluded that anatomical and photosynthetic carbon assimilating characteristics in P. communis were developing to the direction of C₄ metabolism in response to the increasing drought stress in desert areas. The DR enduring severe water stress had more C₄ like photosynthetic features than the HSMR and LSMR as well as SR, according to significantly increased VMD and LSP and higher g _s sensitivity to C _i as well as higher PEPCase activity and lower GO activity in the DR.