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.     

Water deficit, leaf rolling and susceptibility to photoinhibition in field grown sorghum

Chlorophyll fluorescence and gas-exchange techniques were used to investigate changes in photosynthelic performance in response to high light and mild water deficit, in two cultivars of the C., plant sorghum ( Sorghurn bicofor [L.] Moench). grown under field conditions. For all leaves fully exposed to the sun, the efficiency of phcttosystem 11 (PSII) showed a mid-day decline, hut with substantial over-night recovery: the magnitude of the mid-day decline was enhanced by water deficit. There was no corresponding decline in leaves not exposed to full sunlight, either because they were shaded by other leaves or else because of leaf-roiling. Net assimilation rates appeared more sensitive to water-deficit than was PSI1 efficiency. Shade-adapted leaves had lower rates of photosynthesis in full sun (and lower stomatal conductances) than well-exposed leaves. When these shade-adapted leaves were suddenly exposed to full sunlight, fluorescence quenching was slow. especially when plants were well-watered. For the latter, photochemical quenching (q_p)was small even after several minutes. indicating a continuing imbalance between energy funnelled to PSI1 and subsequent electron transport. Shade-adapted leaves that were water stressed were better able to withstand a sudden increase in irradiance than those that were well watered. It is suggested that the shade-adapted eaves from unirrigated plants. having a lower s'tomatal conductance than the irrigated leaves, had been acclimated by receiving energy in excess of that required to fix CO₂, thus leading to the operation of dissipative mechanisms. A shortened protocol for quenching analysis is proposed that enables non-photochemical quenching to be partitioned into rapidly and slowly relaxing components (the latter including photoinhibition) by relating results to a theoretical maximum yield of variable fluorescence. This is particularly suitable for screening field material.     

Biomass accumulation and allocation in soybean associated with genotypic differences in tolerance of nitrogen fixation to water deficits

Nitrogen fixation in soybean (Glycine max [L.] Merr.) is more sensitive to water deficits than many physiological processes and may therefore limit yield under nonirrigated conditions. Tolerance of nitrogen fixation to water deficits has been observed in the cultivar Jackson, however, the physiological basis for this is unclear. It was hypothesized that genotypes that could continue biomass production on limited soil water would prolong nitrogen fixation by continued photosynthate allocation to nodules. An initial greenhouse experiment compared biomass and N accumulation in six genotypes over an 8 d water deficit. Low stress intensity minimized genotypic expression of water-deficit tolerance; nevertheless, Jackson was clearly one of the most tolerant genotypes. In a second experiment, Jackson was compared to SCE82-303 at more severe stress levels. Biomass and N accumulation continued during water deficits for Jackson but ceased in SCE82-303. Individual nodule mass tended to increase during water deficits in Jackson and tended to decrease in SCE82-303, indicating greater allocation of photosynthate to Jackson's nodules in response to water deficits. Biomass accumulation of Jackson was contrasted with the USDA plant introduction (PI) 416937, which also has demonstrated tolerance to water deficits. For water-deficit treatments, total biomass accumulation was negligible for PI416937, but biomass accumulation continued at approximately 64% of the control treatment for Jackson. Transpirational losses for Jackson and PI416937 were approximately the same for the water-deficit treatment, indicating that Jackson had higher water use efficiency (WUE). Isotopic discrimination of ¹³C relative to¹² C also indicated that Jackson had superior WUE during water deficits. Carbon-14 allocation in Jackson was compared to KS4895, a cultivar that was identified as sensitive to water deficits in an initial experiment. The comparison of water-deficit treatments of Jackson with KS4895 indicated that Jackson exported significantly greater amounts of¹⁴ C from labeled leaves and allocated approximately four times greater amounts of ¹⁴C per g of nodule. Results indicated that Jackson's sustained biomass production during water deficits resulted in the continued allocation of photosynthate to nodules and prolonged nitrogenase activity.     

Nitrogen reallocation and photosynthetic acclimation in response to partial shading in soybean plants

The first trifoliate of soybean was shaded when fully expanded, while the plant remained in high light; a situation representative for plants growing in a closed crop. Leaf mass and respiration rate per unit area declined sharply in the first few days upon shading and remained rather constant during the further 12 days of the shading treatment. Leaf nitrogen per unit area decreased gradually until the leaves were shed. Leaf senescence was enhanced by the shading treatment in contrast to control plants growing in low light. Shaded leaves on plants grown at low nutrient availability senesced earlier than shaded leaves on plants grown at high nutrient availability. The light saturated rate of photosynthesis decreased also gradually during the shading treatment, but somewhat faster than leaf N, whereas chlorophyll contents declined somewhat slower than leaf N.
Partitioning of N in the leaf over main photosynthetic functions was estimated from parameters derived from the response of photosynthesis to CO₂. It appeared that the N exported from the leaf was more at the expense of compounds that make up photosynthetic capacity than of those involved in photon absorption, resulting in a change in partitioning of N within the photosynthetic apparatus. Photosynthetic nitrogen use efficiency increased during the shading treatment, which was for the largest part due to the decrease in leaf N content, to some extent to the decrease in respiration rate and only for a small part to change in partitioning of N within the photosynthetic apparatus.     

An attempt to establish a synthetic model of photosynthesis-transpiration based on stomatal behavior for maize and soybean plants grown in field

A synthetic model of photosynthesis-transpiration was established based on a comprehensive consideration of models of CO₂ and H₂O fluxes controlled by stomata of plant leaves.The synthetic model was developed by introducing the internal conductance to CO₂ assimilation, g_ic, and the general equation of stomatal conductance model to H₂O diffusion, g_sw = g₀+a₁A_mf(D_s)/(C_s-Γ), into models of CO₂ and H₂O diffusion through the plant leaves stomata. In the above expression, g₀ and a₁ are coefficients, C_s ambient CO₂ concentration at leaf surface, Γ CO₂ compensation point, and f(D_s) the general function describing the response of stomatal conductance to humidity. Using the data observed in maize (Zea mays L.) and soybean (Glycine max Merr.) plants grown in the field, the parameters in the model were identified, and the applicability of the model was examined. The verification indicated that the developed model could be used to estimate net assimilation rate, transpiration rate, and water use efficiency with a high enough level of precision. The examination also showed that when f(D_s) = h_s or f(D_s) = (1+D_s/D₀)⁻¹ was employed, the estimation precision of the synthetic model was highest. In the study, the parameter g_ic was estimated by means of a linear function of Q_P because it was shown to be mostly correlated with photosynthetic photon flux, Q_P, among various environmental factors.     

Stomatal response to humidity in sugarcane and soybean: effect of vapour pressure difference on the kinetics of the blue light response

Abstract. The effect of atmospheric humidity on the kinetics of stomatal responses was quantified in gas exchange experiments using sugarcane ( Saccharum spp. hybrid) and soybean ( Glycine max ). Pulses of blue light were used to elicit pulses of stomatal conductance that were mediated by the specific blue light response of guard cells. Kinetic parameters of the conductance response were more closely related to leaf-air vapour pressure difference (VPD) than to relative humidity or transpiration. Increasing VPD significantly accelerated stomatal opening in both sugarcane and soybean, despite an approximately five-fold faster response in sugarcane. In contrast, the kinetics of stomatal recovery (closure) following the pulse were similar in the two species. Acceleration of opening by high VPD was observed even under conditions where soybean exhibited a feedforward response of decreasing transpiration (E) with increasing evaporative demand (VPD). This result suggests that epidermal, rather than bulk leaf, water status mediates the VPD effect on stomatal kinetics. The data are consistent with the hypothesis that increased cpidermal water loss at high VPD decreases the backpressure exerted by neighbouring cells on guard cells. allowing more rapid stomatal opening per unit of guard cell metabolic response to blue light.     

An improved dynamic model of photosynthesis for estimation of carbon gain in sunfleck light regimes

A dynamic model of leaf photosynthesis for C₃ plants has been developed for examination of the role of the dynamic properties of the photosynthetic apparatus in regulating CO₂ assimilation in variable light regimes. The model is modified from the Farquhar-von Caemmerer-Berry model by explicitly including metabolite pools and the effects of light activation and deactivation of Calvin cycle enzymes. It is coupled to a dynamic stomatal conductance model, with the assimilation rate at any time being determined by the joint effects of the dynamic biochemical model and the stomatal conductance model on the intercellular CO₂ pressure. When parametrized for each species, the model was shown to exhibit responses to step changes in photon flux density that agreed closely with the observed responses for both the understory plant Alocasia macrorrhiza and the crop plant Glycine max. Comparisons of measured and simulated photosynthesis under simulated light regimes having natural patterns of lightfleck frequencies and durations showed that the simulated total for Alocasia was within ±4% of the measured total assimilation, but that both were 12–50% less than the predictions from a steady–state solution of the model. Agreement was within ±10% for Glycine max, and only small differences were apparent between the dynamic and steady–state predictions. The model may therefore be parametrized for quite different species, and is shown to reflect more accurately the dynamics of photosynthesis than earlier dynamic models.     

The effects of nitrogen, light and water availability on tropic leaf movements in soybean (Glycine max)

Abstract. Rapid, tropic leaf movements and photo-synthetic responses of the heliotropic plant, soybean, Glycine max cv. Cumberland, grown under two different nitrogen, three different light and two different water treatments were examined. Measurements of leaf orientation during midday periods outdoors, and tropic reorientation of leaflets in response to vertical illumination indoors, revealed a positive, linear relationship between leaf water potential and the cosine of the angle of incidence between the leaf and the direct beam of the excitation light. This relationship was altered by nitrogen availability, such that a lower cosine of incidence (lower leaf irradiance) for a given leaf water potential was measured for plants grown under low nitrogen compared to those grown under high nitrogen. Additionally, plants grown under low nitrogen and low water availability showed more rapid rates of leaf movement compared to plants receiving high levels of these resources. Light regime during growth had no effect on the relationship between the cosine of incidence and leaf water potential. Reduced water and nitrogen availabilities during growth resulted in lower photosaturated rates of photosynthesis and stomatal conductance, as well as alterations in the relationship between these parameters. Thus, higher values for the ratio of intercellular CO₂/ambient CO₂ were measured for low-N grown plants (higher nitrogen use efficiencies) and lower values of this ratio for water stressed plants (higher water use efficiencies). The results show that environmental growth conditions other than water availability have the potential to modify leaf orientation responses to vectorial light in heliotropic legumes such as soybean. This has implications for the potential of heliotropic movements to minimize environmental stress-induced damage to the photosynthetic apparatus, and to modulate leaf-level resource use efficiencies.     

Growth and photosynthesis of soybean (Glycine max (L.) Merr.) in simulated vegetation shade: influence of the ratio of red to far-red radiation*

Abstract. Glycine max (L.) Merr. was grown under several light conditions to determine the role of red and far-red radiation in plant adaptation to vegetation shade. Neutral density,‘neutral’ density with elevated far-red radiation, and green shade treatments were used in a greenhouse, producing calculated phytochrome photostationary state (P_fr/P_r+P_fr) values of 0.68, 0.63 and 0.51, respectively. Cool-white fluorescent lamps either alone or in conjunction with far-red fluorescent lamps were used in a growth chamber, providing P_fr/P_r+P_fr of 0.79 and 0.61, respectively. Daily photo-synthetically active radiation was about 25% of daylight and was approximately equal for both greenhouse (2.15MJ m^?2) and growth chamber (2.57MJ m^?2). Developmental stage 4 weeks after sowing was similar for all treatments, but axillary growth and rates of leaf area and dry matter accretion differed between plants from greenhouse and growth chamber. Light conditions simulating vegetation shade (i.e. a low ratio of red to far-red radiation) significantly promoted petiole elongation and retarded the rate of stem elongation in both greenhouse and growth chamber experiments. Other aspects of growth either were not significantly altered by spectral quality or were not modified consistently in both greenhouse and growth chamber environments. Net photosynthetic rates measured under growth conditions for unifoliate and first trifoliolate (TF₁) leaves of growth chamber plants between 9 and 21 d after sowing were generally unaffected by spectral quality, but supplemental FR enhanced TF₁ leaf area expansion. The latter effect was not correlated with increased dry matter accumulation. The significance of spectral quality for adaptation of soybeans to canopy closure and intercropping is discussed.     

Dirunal leaf movement, chlorophyll fluorescence and carbon assimilation in soybean grown under different nitrogen and water availabilities

Diurnal heliotropic leaf movements, photosynthetic gas exchange, and the ratio of variable fluorescence to maximum fluorescence (Fv/Fm) of unrestrained and horizontally restrained leaves from soybean (Glycine max cv. Cumberland) plants grown in two different water and two different nitrogen treatments were measured. Leaves of plants grown in low water or low nitrogen availability treatments displayed more pronounced diaheliotropism (solar tracking) in the afternoon and a longer period of paraheliotropism (light avoiding) at midday relative to those of well-watered, high-nitrogen-grown plants. Photosaturated photosynthetic rates and the photon flux required to saturate photosynthesis were reduced by water stress and nitrogen deficiency. Compared to horizontal leaves, irradiance on orienting leaves was nearer to the breakpoint of the photosynthetic light response curve, where photosynthesis is co-limited by ribulose biphosphate regeneration and carboxylation. This would increase the carbon return on investments of nitrogen into photosynthesis. A positive linear relationship between Fv/Fm and quantum yield of photosynthesis was measured. Leaves of low-nitrogen-grown plants had earlier and more prolonged reductions in Fv/Fm at midday compared to leaves of high nitrogen grown plants of the same water treatment. Within the same water and nitrogen treatment, horizontally restrained leaves had lower midday Fv/Fm in relation to orienting leaves. Nitrogen deficiency and water stress enhanced this difference such that horizontally restrained leaves of low water and low nitrogen grown plants had earlier and longer midday depressions in Fv/Fm.     

Variability in mesophyll conductance between barley genotypes,and effects on transpiration efficiency and carbon isotope discrimination

Leaf internal, or mesophyll, conductance to CO₂ (g_m ) is a significant and variable limitation of photosynthesis that also affects leaf transpiration efficiency (TE). Genotypic variation in g_m and the effect of g_m on TE were assessed in six barley genotypes (four Hordeum vulgare and two H. bulbosum). Significant variation in g_m was found between genotypes, and was correlated with photosynthetic rate. The genotype with the highest g_m also had the highest TE and the lowest carbon isotope discrimination as recorded in leaf tissue (Δ_p). These results suggest g_m has unexplored potential to provide TE improvement within crop breeding programmes.     

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.     

Shoot and root physiological responses to localized zones of soil moisture in cultivated and wild lettuce (Lactuca spp.)

Cultivated crisphead lettuce (Lactuca sativa L.) has a shallower root system than its wild relative, Lactuca serriola L. The effects of localized soil water, at depth, on plant water relations, gas exchange and root distribution were examined in the two species using soil columns with the soil hydraulic-ally separated into two layers, at (0–20 cm and 20–81) cm, but permitting root growth between the layers. Three treatments were imposed on 7-week-old plants, and maintained for 4 weeks: (i) watering, both layers to field capacity; (ii) drying the upper layer while watering the lower layer to field capacity, and (iii) drying both layers. Drying only 0–20 cm of soil had no effect on leaf water status, net photosynthesis, stomatal conductance or biomass production in L. serriola compared to a well-watered control, but caused a short-term reduction (10 d) in leaf water status and photosynthesis in L. sativa that reduced final shoot production. The different responses may be explained by differences in root distribution. Just before the treatments commenced, L. serriola had 50% of total root length at 20–80 cm compared to 35% in L. sativa. Allocation of total biomass to roots in L. serriola was approximately double that in L. sativa. The wild species could provide germplasm for cultivated lettuces to extract more soil water from depth, which may improve irrigation efficiency.     

Lack of photosynthetic or stomatal regulation after 9 years of elevated [CO2] and 4 years of soil warming in two conifer species at the alpine treeline

Alpine treelines are temperature‐limited vegetation boundaries. Understanding the effects of elevated [CO₂] and warming on CO₂ and H₂O gas exchange may help predict responses of treelines to global change. We measured needle gas exchange of Larix decidua Mill. and Pinus mugo ssp. uncinata DC trees after 9 years of free air CO₂ enrichment (575 µmol mol^?1) and 4 years of soil warming (+4 °C) and analysed δ¹³C and δ¹⁸O values of needles and tree rings. Tree needles under elevated [CO₂] showed neither nitrogen limitation nor end‐product inhibition, and no down‐regulation of maximal photosynthetic rate (A_max) was found. Both tree species showed increased net photosynthetic rates (A_n) under elevated [CO₂] (L. decidua: +39%; P. mugo: +35%). Stomatal conductance (g_H2O) was insensitive to changes in [CO₂], thus transpiration rates remained unchanged and intrinsic water‐use efficiency (iWUE) increased due to higher A_n. Soil warming affected neither A_n nor g_H2O. Unresponsiveness of g_H2O to [CO₂] and warming was confirmed by δ¹⁸O needle and tree ring values. Consequently, under sufficient water supply, elevated [CO₂] induced sustained enhancement in A_n and lead to increased C inputs into this ecosystem, while soil warming hardly affected gas exchange of L. decidua and P. mugo at the alpine treeline.     

Effects of doubled atmospheric carbon dioxide concentration on the responses of assimilation and conductance to humidity

Experiments were performed to determine if growth at elevated partial pressure of CO₂ altered the sensitivity of leaf water vapour conductance and rate of CO₂ assimilation to the leaf-to-air difference in the partial pressure of water vapour (Δw). Comparisons were made between plants grown and measured at 350 and 700 μPa Pa^?1 partial pressures of CO₂ for amaranth, soybean and sunflower grown in controlled environment chambers, soybean grown outdoors in pots, and orchard grass grown in field plots. In amaranth, soybean and orchard grass, both the absolute and the relative sensitivity of conductance to Δw at the leaf surface were less in plants grown and measured at the elevated CO₂. In sunflower, there was no change in the sensitivity of conductance to Δw for the two CO₂ partial pressures. Tests in soybeans and amaranth showed that the change in sensitivity resulted from elevated CO₂ during the measurement of the Δw response. Assimilation rate of CO₂ was not altered by Δw in amaranth, which has C₄ metabolism. In sunflower, the assimilation rate of plants grown and measured at elevated CO₂ was insensitive to Δw, consistent with the response of assimilation rate to intercellular CO₂ partial pressure in the prevailing range. In soybean, the sensitivity of assimilation rate to Δw was not different between CO₂ treatments, in contrast to what would be expected from the response of assimilation rate to intercellular CO₂ partial pressure.     

The Glycine-Glomus-Bradyrhizobium symbiosis. IX. Nutritional, morphological and physiological responses of nodulated soybean to geographic isolates of the mycorrhizal fungus Glomus mosseae.

The objective of the work was to determine differences in plant response to geographic isolates of a vesicular-arbuscular mycorrhizal (VAM) fungus, and to demonstrate the need for such determinations in the selection of desirable host-endophyte combinations for practical applications. Soybean ( Glycine max (L.) Merr.) plants were inoculated with Bradyrhizobium japonicum and isolates of the VAM-fungal morphospecies Glomus mosseae (Nicol. & Gerd.) Gerd. and Trappe, collected from an arid (AR), semiarid (SA) or mesic (ME) area. Inoculum potentials of the VAM-fungal isolates were determined and the inocula equalized, achieving the same level of root colonization (41%, P >0.05) at harvest (50 days). Plants of the three VAM treatments (AR, SA and ME) were evaluated against von VAM controls. Significant differences in plant response to colonization were found in dry mass, leaf K, N and P concentrations, and in root/shoot, nodule/root, root length/leaf area and root length/root mass ratios. The differences were most pronounced and consistent between the AR and all other treatments. Photosynthesis and nodule activity were higher ( P <0.05) in all VAM treatments, but only the AR plants had higher ( P <0.05) photosynthetic water-use efficiency than the controls. Nodule activity, evaluated by H₂ evolution and C₂H₂ reduction, differed significantly between treatments. The results are discussed in terms of nutritional and non-nutritional effects of VAM colonization on the development and physiology of the tripartite soybean association in the light of intraspecific variability within the fungal endophyte.     

Mesophyll CO2 conductance and leakiness are not responsive to short‐ and long‐term soil water limitations in the C4 plant Sorghum bicolor

Breeding economically important C₄ crops for enhanced whole‐plant water‐use efficiency (WUE_plant) is needed for sustainable agriculture. WUE_plant is a complex trait and an efficient phenotyping method that reports on components of WUE_plant, such as intrinsic water‐use efficiency (WUE_i, the rate of leaf CO₂ assimilation relative to water loss via stomatal conductance), is needed. In C₄ plants, theoretical models suggest that leaf carbon isotope composition (δ¹³C), when the efficiency of the CO₂‐concentrating mechanism (leakiness, ?) remains constant, can be used to screen for WUE_i. The limited information about how ? responds to water limitations confines the application of δ¹³C for WUE_i screening of C₄ crops. The current research aimed to test the response of ? to short‐ or long‐term moderate water limitations, and the relationship of δ¹³C with WUE_i and WUE_plant, by addressing potential mesophyll CO₂ conductance (g_m) and biochemical limitations in the C₄ plant Sorghum bicolor. We demonstrate that g_m and ? are not responsive to short‐ or long‐term water limitations. Additionally, δ¹³C was not correlated with gas‐exchange estimates of WUE_i under short‐ and long‐term water limitations, but showed a significant negative relationship with WUE_plant. The observed association between the δ¹³C and WUE_plant suggests an intrinsic link of δ¹³C with WUE_i in this C₄ plant, and can potentially be used as a screening tool for WUE_plant in sorghum.     

Stomatal responses to light and humidity in sugarcane: prediction of daily time courses and identification of potential selection criteria

Abstract Stomatal sensitivities to light and VPD have potential as quantitative selection criteria in programs designed to enhance water-use efficiency of sugarcane and other crops. These responses were characterized using gas exchange techniques and then simulated by a mathematical relationship describing conductance as a function of photon fluence rates and VPD values. The same form of relationship simulated stomatal responses of well-watered greenhouse- and field-grown plants. A comparison between simulated and measured conductance values showed a close correlation, indicating that light and VPD responses of stomata are dominant input signals modulating stomatal conductance in sugarcane. Observed conductance of Hawaiian sugarcane in a commerical production area appeared larger than required to support prevailing rates of carbon assimilation, since predicted intercellular CO₂ was greater than required to saturate its C₄ photosynthesis. Manipulation of the relationship describing stomatal conductance allowed us to simulate the responses of plants with hypothetically altered stomatal sensitivities to VPD or to light, using micrometeorological data collected in the field. Further simulation indicated that selection for clones with altered stomatal sensitivity to either light or VPD could improve the water-use efficiency of sugarcane without inhibiting current high levels of productivity.     

Water relations, gas exchange, photochemical efficiency, and peroxidative stress of four plant species in the Heihe drainage basin of northern China

Haloxylon ammodendron, Calligonum mongolicum, Elaeagnus angustifolia, and Populus hosiensis had different adaptations to limited water availability, high temperature, and high irradiance. C. mongolicum used water more efficiently than did the other species. Because of low transpiration rate (E) and low water potential, H. ammodendron had low water loss suitable for desert conditions. Water use efficiency (WUE) was high in E. angustifolia, but high E and larger leaf area made this species more suitable for mesic habitats; consequently, this species is important in tree shelterbelts. P. hosiensis had low WUE, E, and photosynthesis rates, and therefore, does not prosper in arid areas without irrigation. High irradiances caused photoinhibition of the four plants. The decrease of photochemical efficiency was a possible non-stomata factor for the midday depression of C. mongolicum. However, the species exhibited different protective mechanisms against high irradiance under drought stress. H. ammodendron and C. mongolicum possessed a more effective antioxidant defence system than E. angustifolia. These three species showed different means of coping with oxidative stress. Hence an enzymatic balance is maintained in these plants under adverse stress conditions, and the concerted action of both enzymatic and non-enzymatic reactive oxygen species scavenging mechanisms is vital to survive adverse conditions.     

Foliar gas exchange responses of two deciduous hardwoods during 3 years of growth in elevated CO2: no loss of photosynthetic enhancement

Responses of photosynthesis and stomatal conductance were monitored throughout a 3-year field exposure of Liriodendron tulipifera (yellow-poplar) and Quercus alba (white oak) to elevated concentrations of atmospheric CO₂. Exposure to atmospheres enriched with +150 and +300 umol mol^-1 CO₂ increased net photosynthesis by 12–144% over the course of the study. Net photosynthesis was consistently higher at +300 than at +150 umol mol^-1 CO₂. The effect of CO₂ enrichment on stomatal conductance was limited, but instantaneous leaf-level water use efficiency increased significantly. No decrease in the responsiveness of photosynthesis to CO₂ enrichment over time was detected, and the responses were consistent throughout the canopy and across successive growth flushes and seasons. The relationships between internal CO₂ concentration and photosynthesis (e.g. photosynthetic capacity and carboxylation efficiency) were not altered by growth at elevated concentrations of CO₂. No alteration in the timing of leaf senescence or abscission was detected, suggesting that the seasonal duration of effective gas-exchange was unaffected by CO₂ treatment. These results are consistent with data previously reported for these species in controlled-environment studies, and suggest that leaf-level photosynthesis does not down-regulate in these species as a result of acclimation to CO₂ enrichment in the field. This sustained enhancement of photosynthesis provides the opportunity for increased growth and carbon storage by trees as the atmospheric concentration of CO₂ rises, but many additional factors interact in determining whole-plant and forest responses to global change.