Effect of Root Cooling on Photosynthesis of Artemisia tridentata Seedlings under Different Light Levels

Root chilling has been shown to inhibit shoot photosynthesis yet the mechanism for such an action is not clearly understood. A study was designed to elucidate the mechanism by which root cooling may affect net photosynthesis. Roots of Artemisia tridentata seedlings were cooled from 20°C to 5°C while their shoot temperature remained at 20°C. This was conducted at two light levels (700 and 1300 μmol m^?2 s^?1). The time course of shoot net photosynthesis (A), stomatal conductance to water vapor (g_s), intercellular CO₂ concentration (C_i) and root respiration (R_s) were determined on a whole-plant basis. Root cooling caused a 25% reduction in A at high PPFD, which was preceded by more than 50% reduction of g_s and about 10% reduction in C_i. A versus C_i curves for single branches showed no difference between cold and warm soil temperatures, although stomatal conductance was lower for the lower soil temperature. This suggests that a stomatal limitation may have been involved in the inhibition of A. Furthermore, a concomitant decrease of as much as 23% in leaf relative water content (RWC) indicated that root cooling affected stomatal closure due to decreased water supply to the foliage. At lower PPFD, root cooling did not cause a decrease in A of the whole plant despite a moderate drop in g_s, C_i and RWC. Cold soil also led to a substantial and rapid reduction in root respiration rate (R_s) regardless of the light level.     

The use of low [CO2] to estimate diffusional and non-diffusional limitations of photosynthetic capacity of salt-stressed olive saplings

In this study it has been shown that increased diffusional resistances caused by salt stress may be fully overcome by exposing attached leaves to very low [CO₂] (～ 50 µmol mol^?1), and, thus a non‐destructive‐in vivo method to correctly estimate photosynthetic capacity in stressed plants is reported. Diffusional (i.e. stomatal conductance, g_s, and mesophyll conductance to CO₂, g_m) and biochemical limitations to photosynthesis (A) were measured in two 1‐year‐old Greek olive cultivars (Chalkidikis and Kerkiras) subjected to salt stress by adding 200 mm NaCl to the irrigation water. Two sets of A–C_i curves were measured. A first set of standard A–C_i curves (i.e. without pre‐conditioning plants at low [CO₂]), were generated for salt‐stressed plants. A second set of A–C_i curves were measured, on both control and salt‐stressed plants, after pre‐conditioning leaves at [CO₂] of ～ 50 µmol mol^?1 for about 1.5 h to force stomatal opening. This forced stomata to be wide open, and g_s increased to similar values in control and salt‐stressed plants of both cultivars. After g_s had approached the maximum value, the A–C_i response was again measured. The analysis of the photosynthetic capacity of the salt‐stressed plants based on the standard A–C_i curves, showed low values of the J_max (maximum rate of electron transport) to V_cmax (RuBP‐saturated rate of Rubisco) ratio (1.06), that would implicate a reduced rate of RuBP regeneration, and, thus, a metabolic impairment. However, the analysis of the A–C_i curves made on pre‐conditioned leaves, showed that the estimates of the photosynthetic capacity parameters were much higher than in the standard A–C_i responses. Moreover, these values were similar in magnitude to the average values reported by Wullschleger (Journal of Experimental Botany 44, 907–920, 1993) in a survey of 109 C₃ species. These findings clearly indicates that: (1) salt stress did affect g_s and g_m but not the biochemical capacity to assimilate CO₂ and therefore, in these conditions, the sum of the diffusional resistances set the limit to photosynthesis rates; (2) there was a linear relationship (r² = 0.68) between g_m and g_s, and, thus, changes of g_m can be as fast as those of g_s; (3) the estimates of photosynthetic capacity based on A–C_i curves made without removing diffusional limitations are artificially low and lead to incorrect interpretations of the actual limitations of photosynthesis; and (4) the analysis of the photosynthetic properties in terms of stomatal and non‐stomatal limitations should be replaced by the analysis of diffusional and non‐diffusional limitations of photosynthesis. Finally, the C₃ photosynthesis model parameterization using in vitro‐measured and in vivo‐measured kinetics parameters was compared. Applying the in vivo‐measured Rubisco kinetics parameters resulted in a better parameterization of the photosynthesis model.     

Direct and indirect effects of elevated CO2 on transpiration from Quercus myrtifolia in a scrub-oak ecosystem

Elevated atmospheric carbon dioxide (C_a) usually reduces stomatal conductance, but the effects on plant transpiration in the field are not well understood. Using constant‐power sap flow gauges, we measured transpiration from Quercus myrtifolia Willd., the dominant species of the Florida scrub‐oak ecosystem, which had been exposed in situ to elevated C_a (350 µmol mol ^{? 1} above ambient) in open‐top chambers since May 1996. Elevated C_a reduced average transpiration per unit leaf area by 37%, 48% and 49% in March, May and October 2000, respectively. Temporarily reversing the C_a treatments showed that at least part of the reduction in transpiration was an immediate, reversible response to elevated C_a. However, there was also an apparent indirect effect of C_a on transpiration: when transpiration in all plants was measured under common C_a, transpiration in elevated C_a‐grown plants was lower than that in plants grown in normal ambient C_a. Results from measurements of stomatal conductance (g_s), leaf area index (LAI), canopy light interception and correlation between light and g_s indicated that the direct, reversible C_a effect on transpiration was due to changes in g_s caused by C_a, and the indirect effect was caused mainly by greater self‐shading resulting from enhanced LAI, not from stomatal acclimation. By reducing light penetration through the canopy, the enhanced self‐shading at elevated C_a decreased stomatal conductance and transpiration of leaves at the middle and bottom of canopy. This self‐shading mechanism is likely to be important in ecosystems where LAI increases in response to elevated C_a.     

Canopy gradients in leaf intercellular CO2 mole fractions revisited: interactions between leaf irradiance and water stress need consideration

Intercellular CO₂ mole fractions (C_i) are lower in the upper canopy relative to the lower canopy leaves. This canopy gradient in C_i has been associated with enhanced rates of carbon assimilation at high light, and concomitant greater draw‐downs in C_i. However, increases in irradiance in the canopy are generally also associated with decreases in leaf water availability. Thus, stress effects on photosynthesis rates (A) and stomatal conductance (G), may provide a further explanation for the observed C_i gradients. To test the hypotheses of the sources of canopy variation in C_i, and quantitatively assess the influence of within‐canopy differences in stomatal regulation on A, the seasonal and diurnal variation in G was studied in relation to seasonal average daily integrated quantum flux density (Q_int) in tall shade‐intolerant Populus tremula L. trees. Daily time‐courses of A were simulated using the photosynthesis model of Farquhar et al. (Planta 149, 78–90, 1980). Stable carbon isotope composition of a leaf carbon fraction with rapid turnover rate was used to estimate canopy gradient in C_i during the simulations. Daily maximum G (G_max) consistently increased with increasing Q_int. However, canopy differences in G_max decreased as soil water availability became limiting during the season. In water‐stressed leaves, there were strong mid‐day decreases in G that were poorly associated with vapour pressure deficits between the leaf and atmosphere, and the magnitude of the mid‐day decreases in G occasionally interacted with long‐term leaf light environment. Simulations indicated that the percentage of carbon lost due to mid‐day stomatal closure was of the order of 5–10%, and seasonal water stress increased this percentage up to 20%. The percentage of carbon lost due to stomatal closure increased with increasing Q_int. Canopy differences in light environment resulted in a gradient of daily average C_i of approximately 20 µmol mol^?1. The canopy variation in seasonal and diurnal reductions in G led to a C_i gradient of approximately 100 µmol mol^?1, and the actual canopy C_i gradient was of the same magnitude according to leaf carbon isotope composition. This study demonstrates that stress effects influence C_i more strongly than within‐canopy light gradients, and also that leaves acclimated to different irradiance and water stress conditions may regulate water use largely independent of foliar photosynthetic potentials.     

Steady-state stomatal responses of C3 and C4 species to blue light fraction: Interactions with CO2 concentration

Blue light induced stomatal opening has been studied by applying a short pulse (~5 to 60 s) of blue light to a background of saturating photosynthetic red photons, but little is known about steady-state stomatal responses. Here we report stomatal responses to blue light at high and low CO₂ concentrations. Steady-state stomatal conductance (g_s) of C₃ plants increased asymptotically with increasing blue light to a maximum at 20% blue (120 μmol m⁻² s⁻¹). This response was consistent from 200 to 800 μmol mol⁻¹ atmospheric CO₂ (C_a). In contrast, blue light induced only a transient stomatal opening (~5 min) in C₄ species above a C_a of 400 μmol mol⁻¹. Steady-state g_s of C₄ plants generally decreased with increasing blue intensity. The net photosynthetic rate of all species decreased above 20% blue because blue photons have lower quantum yield (moles carbon fixed per mole photons absorbed) than red photons. Our findings indicate that photosynthesis, rather than a blue light signal, plays a dominant role in stomatal regulation in C₄ species. Additionally, we found that blue light affected only stomata on the illuminated side of the leaf. Contrary to widely held belief, the blue light-induced stomatal opening minimally enhanced photosynthesis and consistently decreased water use efficiency.     

Water availability affects seasonal CO2‐induced photosynthetic enhancement in herbaceous species in a periodically dry woodland

Elevated atmospheric CO₂ (eCO₂) is expected to reduce the impacts of drought and increase photosynthetic rates via two key mechanisms: first, through decreased stomatal conductance (g_s) and increased soil water content (V_SWC) and second, through increased leaf internal CO₂ (C_i) and decreased stomatal limitations (S_lim). It is unclear if such findings from temperate grassland studies similarly pertain to warmer ecosystems with periodic water deficits. We tested these mechanisms in three important C₃ herbaceous species in a periodically dry Eucalyptus woodland and investigated how eCO₂‐induced photosynthetic enhancement varied with seasonal water availability, over a 3 year period. Leaf photosynthesis increased by 10%–50% with a 150 μmol mol^?1 increase in atmospheric CO₂ across seasons. This eCO₂‐induced increase in photosynthesis was a function of seasonal water availability, given by recent precipitation and mean daily V_SWC. The highest photosynthetic enhancement by eCO₂ (>30%) was observed during the most water‐limited period, for example, with V_SWC <0.07 in this sandy surface soil. Under eCO₂ there was neither a significant decrease in g_s in the three herbaceous species, nor increases in V_SWC, indicating no “water‐savings effect” of eCO₂. Periods of low V_SWC showed lower g_s (less than ≈ 0.12 mol m^?2 s^?1), higher relative S_lim (>30%) and decreased C_i under the ambient CO₂ concentration (aCO₂), with leaf photosynthesis strongly carboxylation‐limited. The alleviation of S_lim by eCO₂ was facilitated by increasing C_i, thus yielding a larger photosynthetic enhancement during dry periods. We demonstrated that water availability, but not eCO₂, controls g_s and hence the magnitude of photosynthetic enhancement in the understory herbaceous plants. Thus, eCO₂ has the potential to alter vegetation functioning in a periodically dry woodland understory through changes in stomatal limitation to photosynthesis, not by the “water‐savings effect” usually invoked in grasslands.     

Stomatal responses of C3, C3-C4 and C4Flaveria species to light and intercellular CO2 concentration: implications for the evolution of stomatal behaviour

Stomatal function mediates physiological trade‐offs associated with maintaining a favourable H₂O balance in leaf tissues while acquiring CO₂ as a photosynthetic substrate. The C₃ and C₄ species appear to have different patterns of stomatal response to changing light conditions, and variation in this behaviour may have played a role in the functional diversification of the different photosynthetic pathways. In the current study, we used gain analysis theory to characterize the stomatal conductance response to light intensity in nine different C₃, C₄ and C₃‐C₄ intermediate species Flaveria species. The response of stomatal conductance (g_s) to a change in light intensity represents both a direct (related to a change in incident light intensity, I) and indirect (related to a change in intercellular CO₂ concentration, C_i) response. The slope of the line relating the change in g_s to C_i was steeper in C₄ species, compared with C₃ species, with C₃‐C₄ species having an intermediate response. This response reflects the greater relative contribution of the indirect versus direct component of the g_s versus I response in the C₄ species. The C₃‐C₄ species, Flaveria floridana, exhibited a C₄‐like response whereas the C₃‐C₄ species, Flaveria sonorensis and Flaveria chloraefolia, exhibited C₃‐like responses, similar to their hypothesized position along the evolutionary trajectory of the development of C₄ photosynthesis. There was a positive correlation between the relative contribution of the indirect component of the g_s versus I response and water use efficiency when evaluated across all species. Assuming that the C₃‐C₄ intermediate species reflect an evolutionary progression from fully expressed C₃ ancestors, the results of the current study demonstrate an increase in the contribution of the indirect component of the g_s versus I response as taxa evolve toward the C₄ extreme. The greater relative contribution of the indirect component of the stomatal response occurs through both increases in the indirect stomatal components and through decreases in the direct. Increases in the magnitude of the indirect component may be related to the maintenance of higher water use efficiencies in the intermediate evolutionary stages, before the appearance of fully integrated C₄ photosynthesis.     

Stomatal response to blue light: water use efficiency in sugarcane and soybean*

Abstract. The significance of blue light-stimulated stomatal conductance for carbon assimilation (A), stomatal conductance (g), intercellular CO₂ (C_i), stomatal limitation of A (L), transpiration (E) and water use efficiency (W = A/E), was determined in a C₄ and a C₃ species. W and L were evaluated for steady-state gas exchange with constant, saturating red light (A_s, g_s, E_s), and for the integrated gas exchange above the steady state baseline induced by a single, brief pulse of blue light (A_p, g_p, E_p). Sugarcane (Saccharum spp. hybrid), a C₄ grass, and soybean (Glycine max) a C₃ dicot, were compared. Sugarcane exhibited typical C₄ behaviour, with A saturing at C_i of ca. 200 μmol mol^?1, compared to >500 μmol mol^?1 in soybean. Steady-state W was also considerably higher in sugarcane. The extent of stomatal opening in response to a blue light pulse, from baseline (g_s) to the maximum value of conductance during the opening response (g_m), was similar in the two species. More rapid opening and closing of stomata in sugarcane resulted in a smaller integrated magnitude of the conductance response (g_p) than in soybean. At the peak of the blue light response, both species exhibited similar levels of L. During the response to the pulse of blue light, A and C_i increased and L decreased to a greater extent in sugarcane than in soybean. As a result, the gas exchange attributed to the stomatal response to blue light exhibited a higher ratio of A_p to E_p (W_p) in sugarcane than in soybean. This W_p was lower in both species than was the W_s associated with the steady state gas exchange. The two species did not differ in the rate of induction of photosynthetic utilization of elevated C_i. The greater stimulation of A in sugarcane was attributed to its C₄ attributes of greater carboxylation efficiency (slope of the A versus C_i relationship), lower g_s and prevailing C_i,s, and greater L_s under steady-state red illumination. Despite saturation of A at low levels of C_i in C₄ species, the gas exchange attributed to the stomatal response to blue light decreased L and contributed considerably to carbon acquisition, while maintaining the high level of W associated with C₄ metabolism.     

The photosynthetic characteristics of papyrus in a tropical swamp

Summary Photosynthesis and transpiration was measured in the large emergent C₄ sedge Cyperus papyrus (papyrus) which occupies wide areas of wetland on the African continent. The maximum observed value of net assimilation was 35 mol CO₂ m^-2 s^-1 at full sunlight but light saturation of photosynthesis did not occur. The quantum yield of photosynthesis obtained from the initial slope of the light response curves (0.06 mol mol^-1 incident light) was relatively high and close to previously recorded values for some C₄ grasses. Measurements made over two days showed that stomatal conductance was sensitive to the ambient air vapour pressure deficit (VPD) and was consistently lower on the day when VPD's were higher. There was, however, no marked midday closure of the stomata. Photosynthesis was also reduced on the day when VPD's were higher. The relationship between net photosynthesis and stomatal conductance was close to linear over the range of measurement conditions, with the result that intercellular CO₂ concentrations (C _i ) did not vary markedly. There was some evidence that C _i decreased at high VPD's. The regulation of stomatal movement in papyrus appears to minimise excessive water loss while not severely limiting photosynthesis. The significance of this strategy for a wetland species with plentiful supplies of water is discussed.     

Critical responses of photosynthetic efficiency of goldspur apple tree to soil water variation in semiarid loess hilly area

Goldspur apple (Malus pumila cv. Goldspur) is one of the main fruit trees planted in semiarid loess hilly areas. The photosynthetic efficiency in leaves of eight-year-old trees were studied under different soil water conditions with a Li-6400 portable photosynthesis system and a Li-Cor1600 portable steady state porometer in order to explore the effects of soil water stress on photosynthesis and the suitable soil water content (SWC) for water-saving irrigation of apple orchards. The results showed that the leaf net photosynthetic rate (P _N), transpiration rate (E), water-use efficiency (WUE), stomatal conductance (g _s), intercellular CO₂ concentration (C _i), and stomatal limiting value (L _s) displayed different threshold responses to soil water variation. When SWC was within a range of about 60%–86% of field capacity (FC), P _N and E were maintained in a relative steady state. At an elevated level but below 60% of FC, both P _N and E decreased evidently with decreasing soil moisture. The SWC needed to support WUE in a relatively steady state and at a high level was in the range of about 50%–71% of FC. When SWC was less than 48% of FC, g _s and L _s declined with decreasing soil moisture, while C _i increased rapidly. Based on the analysis of the stomatal limitation of photosynthesis using two criteria (C _i and L _s) suggested by Farquhar and Sharkey, it was implied that the predominant cause of restricting P _N had changed from stomatal limitation to nonstomatal one under severe water stress. In terms of water-saving irrigation for enhancing water-use efficiency, it was concluded that in semiarid loess hilly areas, the suitable range of SWC for water-saving irrigation in goldspur apple orchards is in the range of about 50%–71% of FC, and the most severe degree of soil water stress tolerated for photosynthesis is about 48% of FC.     

In situ temperature relationships of biochemical and stomatal controls of photosynthesis in four lowland tropical tree species

Net photosynthetic carbon uptake of Panamanian lowland tropical forest species is typically optimal at 30–32 °C. The processes responsible for the decrease in photosynthesis at higher temperatures are not fully understood for tropical trees. We determined temperature responses of maximum rates of RuBP‐carboxylation (V_CMax) and RuBP‐regeneration (J_Max), stomatal conductance (G_s), and respiration in the light (R_Light) in situ for 4 lowland tropical tree species in Panama. G_s had the lowest temperature optimum (T_Opt), similar to that of net photosynthesis, and photosynthesis became increasingly limited by stomatal conductance as temperature increased. J_Max peaked at 34–37 °C and V_CMax ~2 °C above that, except in the late‐successional species Calophyllum longifolium, in which both peaked at ~33 °C. R_Light significantly increased with increasing temperature, but simulations with a photosynthesis model indicated that this had only a small effect on net photosynthesis. We found no evidence for Rubisco‐activase limitation of photosynthesis. T_Opt of V_CMax and J_Max fell within the observed in situ leaf temperature range, but our study nonetheless suggests that net photosynthesis of tropical trees is more strongly influenced by the indirect effects of high temperature—for example, through elevated vapour pressure deficit and resulting decreases in stomatal conductance—than by direct temperature effects on photosynthetic biochemistry and respiration.     

Shoots grafted into the upper crowns of tall Japanese cedar (Cryptomeria japonica D. Don) show foliar gas exchange characteristics similar to those of intact shoots

The lower foliar photosynthetic rates seen in shoots in the upper crowns of tall trees than those in seedlings could be caused by extrinsic factors, such as hydraulic conductance, for shoots or by irreversible intrinsic change in the meristems during tree development. To clarify which is most significant, we compared foliar gas exchange characteristics and water relations among scions of Japanese cedar (Cryptomeria japonica D. Don) grafted into the upper crowns of tall trees, rooted cuttings developed from scions of the same clones, and intact shoots in the upper crowns of the tall trees. Grafted shoots had the same water regime as intact shoots, as confirmed by their similar water potentials at the turgor loss point, which were more negative than those of the rooted cuttings. No significant difference was observed between the grafted and intact shoots in their light-saturated photosynthetic rate (P_max), stomatal conductance (g_s), photosynthetic capacity, carboxylation efficiency, ratio of intercellular to ambient CO₂ concentration (C_i/C_a), and carbon isotope composition (¹³C). Compared with the rooted cuttings, the grafted shoots showed significantly lower P_max, g_s, photosynthetic capacity, and carboxylation efficiency (to 49%, 31%, 68%, and 65%, respectively). The C_i/C_a and ¹³C indicated significantly stronger instantaneous and long-term stomatal limitation in the grafted shoots than in the rooted cuttings. These indicate that changes in extrinsic factors can reduce foliar photosynthetic rates in shoots in the upper crowns of tall trees as a result of stronger stomatal limitation and reduced photosynthetic activity, without irreversible intrinsic changes in the meristems.     

The analysis of photosynthetic performance in leaves under field conditions: A case study using Bruguiera mangroves

In this report, we analyze the photosynthetic capacity and performance of leaves under field conditions with a case study based on the mangroves Bruguiera parviflora and B. gymnorrhiza. Using a tower through a closed canopy at a field sight in North Queensland and portable infra-red gas analyzers, a large data set was collected over a period of 11 days early in the growing season. The set was used to analyze the relationship between net photosynthesis (P_net) and light, leaf temperature, stomatal conductance and intracellular CO₂ (C_i).There are three objectives of this report: (1) to determine photosynthetic potential as indicated by the in situ responses of P_net to light and stomatal conductance, (2) to determine the extent to which photosynthetic performance may be reduced from that potential, and (3) to explore the basis for and physiological significance of the reduction.The results indicate that even under harsh tropical conditions, the mangrove photosynthetic machinery is capable of operating efficiently at low light and with maximal rates of more than 15 mol CO₂ m^-2 s^-1. Though stomata were more often limiting than light, in any single measurement the average reduction of P_net from the maximum value predicted by light or conductance responses was 35%. Analysis of single leaf light and CO₂ responses indicated that photosynthetic performance was under direct photosynthetic, or non-stomatal, control at all light and conductance levels. Capacity was adjustable rapidly from a maximum value to essentially nil such that C_i varied inversely with P_net from ca. 150 L L^-1 at the highest rates of CO₂ exchange to ambient at the lowest.     

Why does photosynthesis decrease with needle age in Pinus pinaster?

Rates of photosynthesis vary with foliage age and typically decline from full-leaf expansion until senescence occurs. This age-related decline in photosynthesis is especially important in species that retain foliage for several years, yet it is not known whether the internal conductance to CO₂ movement (g _i) plays any role. More generally, g _i has been measured in only a few conifers and has never been measured in leaves or needles older than 1 year. The effect of ageing on g _i was investigated in Pinus pinaster, a species that retains needle for 4 or more years. Measurements were made in autumn when trees were not water limited and after leaf expansion was complete. Rates of net photosynthesis decreased with needle age, from 8 μmol m⁻² s⁻¹ in fully expanded current-year needles to 4.4 μmol m⁻² s⁻¹ in 3-year-old needles. The relative limitation due to internal conductance (0.24–0.35 out of 1) was in all cases larger than that due to stomatal conductance (0.13–0.19 out of 1). Internal conductance and stomatal conductance approximately scaled with rates of photosynthesis. Hence, there was no difference among year-classes in the relative limitations posed by internal and stomatal conductance or evidence that they cause the age-related decline in photosynthesis. There was little evidence that the age-related decline in photosynthesis was due to decreases in contents of N or Rubisco. The decrease in rates of photosynthesis from current-year to older needles was instead related to a twofold decrease in rates of photosynthesis per unit nitrogen and V _cmax/Rubisco (i.e., in vivo specific activity).     

Photosynthetic limitations in olive cultivars with different sensitivity to salt stress

Olive (Olea europea L) is one of the most valuable and widespread fruit trees in the Mediterranean area. To breed olive for resistance to salinity, an environmental constraint typical of the Mediterranean, is an important goal. The photosynthetic limitations associated with salt stress caused by irrigation with saline (200 mm ) water were assessed with simultaneous gas‐exchange and fluorescence field measurements in six olive cultivars. Cultivars were found to possess inherently different photosynthesis when non‐stressed. When exposed to salt stress, cultivars with inherently high photosynthesis showed the highest photosynthetic reductions. There was no relationship between salt accumulation and photosynthesis reduction in either young or old leaves. Thus photosynthetic sensitivity to salt did not depend on salt exclusion or compartmentalization in the old leaves of the olive cultivars investigated. Salt reduced the photochemical efficiency, but this reduction was also not associated with photosynthesis reduction. Salt caused a reduction of stomatal and mesophyll conductance, especially in cultivars with inherently high photosynthesis. Mesophyll conductance was generally strongly associated with photosynthesis, but not in salt‐stressed leaves with a mesophyll conductance higher than 50 mmol m^?2 s^?1. The combined reduction of stomatal and mesophyll conductances in salt‐stressed leaves increased the CO₂ draw‐down between ambient air and the chloroplasts. The CO₂ draw‐down was strongly associated with photosynthesis reduction of salt‐stressed leaves but also with the variable photosynthesis of controls. The relationship between photosynthesis and CO₂ draw‐down remained unchanged in most of the cultivars, suggesting no or small changes in Rubisco activity of salt‐stressed leaves. The present results indicate that the low chloroplast CO₂ concentration set by both low stomatal and mesophyll conductances were the main limitations of photosynthesis in salt‐stressed olive as well as in cultivars with inherently low photosynthesis. It is consequently suggested that, independently of the apparent sensitivity of photosynthesis to salt, this effect may be relieved if conductances to CO₂ diffusion are restored.     

Characterization of photosynthesis of Populus euphratica grown in the arid region

Net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (C _i), transpiration rate (E), water use efficiency (WUE), and stomatal limitation (L_s) of Populus euphratica grown at different groundwater depths in the arid region were measured. g _s of the trees with groundwater depth at 4.74 m (D₄) and 5.82 m (D₅) were lower and a little higher than that at 3.82 m (D₃), respectively. Compared with C _i and L_s of the D₃ trees, C _i decreased and L_s increased at 4.74 m, however, Ci increased and L_s decreased at D₅. Hence photosynthetic reduction of P. euphratica was attributed to either stomatal closure or non-stomatal factors depending on the groundwater depths in the plant locations. P _N of the D₃ trees was significantly higher than those at D₄ or D₅. The trees of D₄ and D₅ did not show a significant difference in their P _N, indicating that there are mechanisms of P. euphratica tolerance to mild and moderate drought stress.     

Environmental characteristics,field water relations,and photosynthetic responses of C4 Hawaiian Euphorbia species from contrasting habitats

Summary Four endemic Hawaiian Euphorbia species range in habitat from open arid coastal strand to shaded mesic forest and in growth-form from small prostrate shrubs to trees. As shown in the present study, these large differences in habitat and growth-form are paralleled by equally large differences in maximal photosynthetic rate (13.7 to 37.1 mol CO₂ m^-2s^-1), dark respiration rate (0.7 to 4.1 mol CO₂ m^-2s^-1), light level for saturation of photosynthesis (0.9 to over 2.0 mmol m^-2s^-1), light compensation point (0.01 to 0.11 mmol m^-2s^-1), leaf conductance to CO₂ (1.7 to 4.9 mm s^-1), and mesophyll conductance to CO₂ (3.7 to 8.5 mm s^-1). A principal consequence of this differentiation is that the capacity for photosynthesis at high light levels is higher in open site species, such as E. celastroides and E. degeneri, and at low light levels is higher in shade species, such as E. forbesii. E. hillebrandii, a species from intermediate semiopen habitats, exhibits an intermediate photosynthetic capacity at both high and low light levels. Despite this remarkable diversity, all four species exhibit the distinguishing physiological features of C₄ photosynthesis.     

The apparent feed-forward response to vapour pressure deficit of stomata in droughted, field-grown Eucalyptus globulus Labill

This study tested a multiplicative model of stomatal response to environment for drought‐affected trees of Eucalyptus globulus Labill. growing in southern Australia. The model incorporates a feed‐forward response to vapour pressure deficit of ambient air (δe_a) and performed well if evaluated using reduced major axis regression and log‐transformed data. There was strong evidence from gas‐exchange data, leaf water potentials and sapflow measurements of the feed‐forward response by stomata to leaf‐to‐air vapour pressure deficit (δe_l). The response of stomata to δe_l was irreversible. Stomatal conductance and the rate of net photosynthesis were highly correlated and declined, together with the rate of transpiration, throughout the afternoon as δe_a increased despite increasing leaf water potentials. The concentration of CO₂ inside leaves (c_i) increased as stomatal conductance declined indicating increasing non‐stomatal limitations to photosynthesis. The stomatal response to δe_l of E. globulus in the field is best described as an ‘apparent feed‐forward response’ that probably results from both slowly reversible depression of net photosynthesis and abscisic acid accumulation in guard cells. We suggest that the stomatal response to c_i may strengthen the link between photosynthetic capacity and stomatal conductance during leaf drying as a result of either drought or large δ e_l.     

Responses of Two Olive Tree (Olea Europaea L.) Cultivars to Elevated CO2 Concentration in the Field

Five-year-old plants of two olive cultivars (Frantoio and Moraiolo) grown in large pots were exposed for 7 to 8 months to ambient (AC) or elevated (EC) CO₂ concentration in a free-air CO₂ enrichment (FACE) facility. Exposure to EC enhanced net photosynthetic rate (P _N) and decreased stomatal conductance, leading to greater instantaneous transpiration efficiency. Stomata density also decreased under EC, while the ratio of intercellular (C _i) to atmospheric CO₂ concentration and chlorophyll content did not differ, except for the cv. Moraiolo after seven months of exposure to EC. Analysis of the relationship between photosynthesis and C _i indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase after five months of exposure to EC. Based on estimates derived from the P _N-C _i relationship, there were no apparent treatment differences in daytime respiration, CO₂ compensation concentration, CO₂-saturated photosynthetic rate, or photosynthetic rate at the mean C _i, but there was a reduction in stomata limitation to P _N at EC. Thus 5-year-old olive trees did not exhibit down regulation of leaf-level photosynthesis in their response to EC, though some indication of adjustment was evident for the cv. Frantoio with respect to the cv. Moraiolo.     

Diurnal Oscillation in the Intercellular CO2 Concentration of Spring Wheat Under the Semiarid Conditions

Yields of wheat in semiarid and arid zones are limited by drought, and water condition is very important at each stage of development. Studies carried out at Loess Plateau in the northwestern part of China indicated that yield of spring wheat (Triticum aestivum L.) cv. Dingxi 81-392 was reduced by 41% when subjected to water stress. The effects of two water regimens on net photosynthetic rate (P _N), stomatal conductance (g _s), and intercellular CO₂ concentration (C _i) were investigated at the jointing, booting, anthesis, and grain filling stages. Low soil moisture in comparison to adequate one had invariably reduced P _N during the diurnal variations at the four growth stages. P _N and g _s in both soil moisture regimes was maximally reduced at midday. C _i and the stomatal limitation fluctuated remarkably during photosynthesis midday depression processes, especially at the grain filling stage. Hence atmospheric drought at midday was one of the direct causes inducing stomata closure and the g _s depression, but it was beneficial for maintaining stable intrinsic water use efficiency. Fluctuation in C _i implicated that non-stomatal limitation also plays an important role during the period of photosynthesis midday depression. Consequently stomatal and/or non-stomatal limitation are the possible cause of the midday photosynthesis decline.