Carbon assimilation, 𝛅13C and water relations of Elaeagnus angustifolia grown at two groundwater depths in the Minqin desert,China

Abstract

We investigated the physiological responses of Elaeagnus angustifolia to variation in groundwater depth. Elaeagnus angustifolia seedlings were grown in the Minqin desert in lysimeters supplied with underground water at the soil depth of 1.40 m and 3.40 m. Results showed that constant access to groundwater allowed plants supplied with water at the lower soil layer to meet their water requirement and, consequently, they were not affected by water stress. There were no differences in A _max (the net CO₂ assimilation rate under conditions of photosynthetically photon flux density and CO₂ saturation), J _max (maximum rate of electron transport) and stomatal conductance between the two underground water treatments. However, plants with deeper groundwater had a significantly higher V _cmax (i.e. a higher carboxylation efficiency of Rubisco) and mesophyll conductance resulting in increased photosynthesis measured at the CO₂ growth condition (A) and, consequently higher intrinsic water-use efficiency (WUE). However, respiration was also increased in plants grown with deeper groundwater. This may have offset the increased A and led to a similar long-term WUE, as expressed by carbon isotope discrimination (δ¹³C), between the two ground water treatments. In the present study, we also found a high foliage nitrogen concentration in the E. angustifolia plants (3.75% on average), that may be very significant ecologically in improving soil properties. The physiological traits of E. angustifolia found in this study confirm that the use of perennial phreatophytic, nitrogen fixing species has significant potential to positively impact soil fertility and carbon sequestration under environmental conditions found in the Minqin desert.     

Effects of Elevated Temperature on Growth and Gas Exchange in Dominant Plant Species from Maowusu Sandland,China

We compared the effect of elevated temperature on morphological development, biomass accumulation and allocation, and gas exchange of three dominant plants (Caragana intermedia Kuanget H.C. Fu, Hedysarum mongolicum Turcz., and Artemisia ordosica Krasch.) growing in Chinese Maowusu sandland. Plants were grown in two temperature chambers (25/20, 28/23 °C, day/night) during 60 d. Tree height, number of leaves, and leaf area were increased in C. intermedia and H. mongolicum seedlings, while in A. ordosica temperature only affected tree height. Elevated temperature increased biomass and reduced the root : shoot ratio in C. intermedia and H. mongolicum seedlings, but not in A. ordosica seedlings. The net photosynthetic rate (P _N) and transpiration rate (E) were increased at days 40 and 60 in C. intermedia and H. mongolicum seedlings, while in A. ordosica seedlings no significant effects on E were observed, and P _N was increased only at day 60. Water use efficiency (WUE) was reduced at days 40 and 60 in H. mongolicum seedlings, and at day 60 in C. intermedia seedlings. No temperature effect on WUE was observed in A. ordosica seedlings. These different responses indicate that climate change could alter plant communities in Maowusu sandland.     

Photosynthesis,Transpiration, and Water Use Efficiency of Caragana microphylla,C. intermedia,and C. korshinskii

In the order C. microphylla — C. intermedia — C. korshinskii, compensation irradiance, saturation irradiance, and optimum temperature for photosynthesis increased, net photosynthetic rate (P _N) at low irradiance and low temperature decreased, optimum air humidity decreased, and P _N at low air humidity increased. Daily cumulative value of P _N increased while daily cumulative value of transpiration (E) decreased, and hence water use efficiency (WUE =P _N/E) increased. Diurnal course of P _N of C. microphylla was a double-peak curve, but the second peak in the curves of C. intermedia and C. korshinskii was not visible. These physiological characteristics are biological basis for the geographical distribution of these three Caragana species, and are in relation to water conditions of their habitats and distinctiveness in leaf hair of plant.     

Differences in gas exchange contribute to habitat differentiation in Iberian columbines from contrasting light and water environments

During photosynthesis, respiration and transpiration, gas exchange occurs via the stomata and so plants face a trade‐off between maximising photosynthesis while minimising transpiration (expressed as water use efficiency, WUE). The ability to cope with this trade‐off and regulate photosynthetic rate and stomatal conductance may be related to niche differentiation between closely related species. The present study explored this as a possible mechanism for habitat differentiation in Iberian columbines. The roles of irradiance and water stress were assessed to determine niche differentiation among Iberian columbines via distinct gas exchange processes. Photosynthesis–irradiance curves (P–I curves) were obtained for four taxa, and common garden experiments were conducted to examine plant responses to water and irradiance stress, by measuring instantaneous gas exchange and plant performance. Gas exchange was also measured in ten individuals using two to four field populations per taxon. The taxa had different P–I curves and gas exchange in the field. At the species level, water stress and irradiance explained habitat differentiation. Within each species, a combination of irradiance and water stress explained the between‐subspecies habitat differentiation. Despite differences in stomatal conductance and CO₂ assimilation, taxa did not have different WUE under field conditions, which suggests that the environment equally modifies photosynthesis and transpiration. The P–I curves, gas exchange in the field and plant responses to experimental water and irradiance stresses support the hypothesis that habitat differentiation is associated with differences among taxa in tolerance to abiotic stress mediated by distinct gas exchange responses.     

Gibberellic acid affects growth and flowering of Philodendron‘Black Cardinal’

Changes in net photosynthetic rate (P_N), stomatal conductance (g_s), intercellular CO₂ concentrations (C_i), transpiration rate (E) and water use efficiency (WUE) were measured in Plantago major L. plants grown under sufficient soil water supply or under soil water stress conditions. The plants had high P_N in a wide range of soil water potential and temperature regimes. Soil water had little effect on P_N under ambient CO₂ concentrations, which was explained by a high carboxylation rate, but increased the dark respiration rate. Carboxylation activity at low C_i depended on RuBP regeneration, whereas at high C_i it depended on the phosphate regeneration rate. The g_s and E values were low in plants under stress as compared to the controls that resulted in an increase of WUE. The results obtained show that Plantago major plants have different ways of adaptation to soil water deficit conditions.     

Ecophysiology of first and second generation hybrids in a natural plant hybrid zone

Hybrids between related species vary widely in relative fitness, and that fitness can depend upon the environment. We investigated aspects of physiology that might influence fitness patterns in a plant hybrid zone. Seeds of Ipomopsis aggregata, I. tenuituba, F1 hybrids, F2 hybrids, and offspring of crosses between natural hybrids were planted into the relatively mesic site of origin for I. aggregata and the drier site for natural hybrids. We measured rates of photosynthesis (A _max), transpiration (E), instantaneous (A/E) and long-term (δ¹³C) indices of water use efficiency (WUE), and leaf nitrogen and carbon. We also examined correlations of these traits with plant size. Photosynthetic rate and A/E were higher in vegetative than flowering plants. WUE varied between sites and years, but differences among genotypic classes were spatially and temporally consistent. Instantaneous WUE was higher for F1 hybrids than for the average of the parental species, thereby showing heterosis. There was no evidence of hybrid breakdown, as WUE was no different in the F2 than the average across the F1 and parental species. Nor did WUE depend on cross direction in producing F1 progeny. Carbon isotope discrimination revealed higher long-term water use efficiency in I. tenuituba than I. aggregata. Leaf nitrogen was higher in I. tenuituba than I. aggregata, and higher in offspring of natural hybrids than in the F2. Results indicate heterosis for water use efficiency, with no hybrid breakdown. Heterosis in WUE may help to explain the relatively high survival of both reciprocal F1 hybrids in dry sites within the natural hybrid zone.     

Relationships between gas-exchange characteristics and stomatal structural modifications in some desert grasses under high salinity

Two populations, one from lesser saline Derawar Fort (DF) and the other from highly saline Ladam Sir (LS) in the Cholistan desert, for each of the five grass species, Aeluropus lagopoides, Cymbopogon jwarancusa, Lasiurus scindicus, Ochthochloa compressa, and Sporobolus ioclados were examined to investigate the influence of salinity on structural and functional characteristics of stomata. Salinity tolerance in A. lagopoides mainly depended on controlled transpiration rate (E) and high water-use efficiency (WUE), which was found to be regulated by fewer and smaller stomata on both leaf surfaces as well as stomatal encryption by epidermal invaginations. C. jwarancusa had sunken stomata on the abaxial surface only, which largely reflected a reduced E, but less affected stomatal conductance (g _s) or WUE. L. scindicus had fewer but larger stomata along with hairs/trichomes which may function to avoid water loss through transpiration, and hence, to attain a high WUE. In O. compressa stomata were found only on the abaxial surface and these were completely encrypted by epidermal invaginations as well as a dense covering of microhairs, which was associated with a low E and high WUE under salinity stress. In S. ioclados, the traits of increased stomatal density and decreased stomatal area may be critical for stomatal regulation under salt-prone environments. High stomatal regulation depended largely on stomatal density, area, and degree of encryption under salinity, which is of great ecophysiological significance for plants growing under osmotic stresses.     

新疆策勒绿洲外围四种多年生植物的水分生理特征

对新疆策勒绿洲外围多年生植物胡杨、柽柳、沙拐枣和骆驼刺的水分生理特性进行了试验研究.结果表明：4种植物在生长季内没有受到严重的水分胁迫,灌溉对植物水分生理指标变化的影响不显著(P＞0.05).4种植物水分生理特性的季节变化各不相同.其中,骆驼刺的清晨水势和日均茎流量最高,但其平均水分利用效率最低;沙拐枣的平均水分利用效率最高,而且其清晨水势与日均茎流量的变化最为稳定,季节变幅不大;柽柳的清晨水势最低,具有较好的环境适应性;胡杨水分生理指标的季节变化相对平稳.在新疆策勒绿洲外围的极端干旱环境中,4种植物通过深根系和地下水相连接,并利用地下水来维持其生存与生长.     

The combined effect of constant water deficit and nitrogen supply on WUE, NUE and Δ13C in durum wheat potted plants

Water scarcity and nitrogen shortage are the main constraints on durum wheat productivity. This paper examines the combined effects of a constant water deficit and nitrogen supply (NS) on growth, photosynthesis, stomatal conductance (g_s) and transpiration, instantaneous and time‐integrated water use efficiency (WUE) and nitrogen use efficiency (NUE) and carbon isotope discrimination (Δ¹³C) in durum wheat genotypes grown in pots under greenhouse conditions. Three water levels (40%, 70% and 100% container capacity), two nitrogen doses (high and low N) and four genotypes were assayed in a total of 24 experimental treatments. Water and nitrogen treatments were imposed 2 weeks after plant emergence. The growth, nitrogen content and Δ¹³C of the shoot and the gas exchange in the flag leaf were determined about 2 weeks after anthesis. As expected, both water and NS had a strong positive effect on growth. However, a reduction in water supply had low effect decreasing photosynthesis and transpiration, Δ¹³C and NUE and increasing WUE. On the contrary, increasing the level of nitrogen supplied had a significant negative effect on g_s, which decreased significantly the ratio of intercellular to ambient CO₂ concentrations and Δ¹³C, and increased both instantaneous and time‐integrated WUE. In addition, a higher N level also negatively affected the instantaneous and time‐integrated NUE. The Δ¹³C of shoots correlated significantly and negatively with either instantaneous or time‐integrated measurements of WUE. Moreover, within each NS, Δ¹³C also correlated negatively with the integrated NUE. We concluded that under our experimental conditions, Δ¹³C gives information about the efficiency with which not just water but also nitrogen are used by the plant. In addition, this study illustrates that a steady water limitation may strongly affect biomass without consistent changes in WUE. The lack of effect of the different water regimes on gas exchange, WUE and Δ¹³C illustrate the importance of how stress is imposed during growth.     

四种荒漠植物保色浸渍标本实验研究

利用浸制标本能较长时间地保持原植物的形态、色泽、质地。以CuSO₄溶液浓度的不同梯度,选用L₉(3⁴)表进行4种荒漠植物保色浸渍标本正交试验研究,结果表明梭梭（Haloxylon ammodendron(C.A.Mey.)Bge.）在15%CuSO₄、沙拐枣（Calligonum mongolicum Turcz.）在10%CuSO₄溶液、中麻黄（Ephedra intermedia Schrenk）在5%CuSO₄溶液、多枝柽柳（Tamarix ramosissima Ledeb.）在5%CuSO₄溶液中浸渍效果最佳,推导出的最优配方与试验结果有很好的拟合性,有一定的实验价值。     

Morphological adaptations of emergent plants to water flow: a case study with Typha angustifolia, Zizania latifolia and Phragmites australis

1. Water velocity plays an important role in shaping plant community structure in flowing waters although few authors have yet attempted to explain the adaptation of plants to flow. 2. We aimed to test two hypotheses, that: (i) some emergent macrophytes reconfigure their shoot distribution in fast currents and form clumps, and (ii) the shape and morphology of such clumps minimises drag caused by the current. The study focuses on three emergent macrophytes that co‐occur along a gradient of water velocity. 3. The species showed a clear zonation in response to water depth and current velocity. Phragmites australis occupied shallower and more slowly flowing water than Typha angustifolia and Zizania latifolia, which had similar preferences. 4. Both T. angustifolia and Z. latifolia shoots were more clumped at high velocity, whereas they were more randomly distributed at low flow or in stagnant water. Because of the low shoot density, water flowed more easily through T. angustifolia clumps, whereas Z. latifolia clumps had a high shoot density and large amounts of trapped litter, causing stagnant water in the centre of the clump. The clumps of Z. latifolia with a high density of shoots were longer and narrower than T. angustifolia clumps. Phragmites australis was less tolerant of flow than the other two species and large amounts of litter trapped in the clumps impaired flow. 5. The shoot distribution of both T. angustifolia and Z. latifolia is reconfigured at high flow and this minimises drag on the clumps.     

Photosynthetic regulation of C4 desert plant Haloxylon ammodendron under drought stress

About 20-year-old desert plants of C₄ species, Haloxylon ammodendron, growing at the southern edge of the Badain Jaran Desert in China, were selected to study the photosynthetic characteristics and changes in chlorophyll fluorescence when plants were subject to a normal arid environment (AE), moist atmospheric conditions during post-rain (PR), and the artificial supplement of soil water (SW). Results showed that under high radiation, in the AE, the species down-regulated its net assimilation rate (A) and maximum photochemical efficiency of PS II (Fv/Fm), indicating photoinhibition. However, under the PR and SW environments, A was up-regulated, with a unimodal diurnal course of A and a small diurnal change in Fv/Fm, suggesting no photoinhibition. When the air humidity or SW content was increased, the light compensation points were reduced; light saturation points were enhanced; while light saturated rate of CO₂ assimilation (A _max) and apparent quantum yield of CO₂ assimilation (Φ_C) increased. Φ_C was higher while the A _max was reduced under PR relative to the SW treatment. It was concluded that under high-radiation conditions drought stress causes photoinhibition of H. ammodendron. Increasing air humidity or soil moisture content can reduce photoinhibition and increase the efficiency of solar energy use.     

Diurnal Gas Exchange and Superior Resources Use Efficiency of Typical C4 Species in Hunshandak Sandland,China

Net photosynthetic rate (P _N), transpiration rate (E), stomatal conductance (g _s), leaf water potential (ψ_leaf), leaf nitrogen content, and photosynthetic nitrogen use efficiency (PNUE) were compared between a typical C₄ plant, Agriophyllum squarrosum and a C₃ plant, Leymus chinensis, in Hunshandak Sandland, China. The plant species showed different diurnal gas exchange patterns on June 12–14 when photosynthetic photon flux density (PPFD), air temperature (T _air), and water potential were moderate. P _N, E, and g _s of A. squarrosum showed distinct single peak while those of L. chinensis were depressed at noon and had two peaks in their diurnal courses. Gas exchange traits of both species showed midday depression under higher photosynthetic photon flux density (PPFD) and T _air when Ψ_leaf was significantly low down on August 6–8. However, those of A. squarrosum were depressed less seriously. Moreover, A. squarrosum had higher P _N, Ψ_leaf, water use efficiency (WUE), and PNUE than L. chinensis. Thus A. squarrosum was much more tolerant to heat and high irradiance and could utilise the resources on sand area more efficiently than L. chinensis. Hence species like A. squarrosum may be introduced and protected to reconstruct the degraded sand dunes because of their higher tolerance to stress and higher resource use efficiency. This revised version was published online in August 2006 with corrections to the Cover Date.     

The effect of growth irradiance on leaf anatomy and photosynthesis in Acer species differing in light demand

Variation in light demand is a major factor in determining the growth and survival of trees in a forest. There is strong relation between the light‐demand and the effect of growth irradiance on leaf morphology and photosynthesis in three Acer species: A. rufinerve (light‐demanding), A. mono (intermediate) and A. palmatum (shade‐tolerant). The increase in mesophyll thickness and surface area of chloroplasts facing the intercellular airspaces (S_c) with growth irradiance was highest in A. rufinerve. Although the increase in light‐saturated photosynthesis (A_max) was similar among the species, the increase in water use efficiency (WUE) was much higher in A. rufinerve than that in the other species, indicating that the response to water limitation plays an important role in leaf photosynthetic acclimation to high light in A. rufinerve. The low CO₂ partial pressure at the carboxylation site (C_c) in A. rufinerve (130 µmol mol^?1) at high irradiance was caused by low stomatal and internal conductance to CO₂ diffusion, which minimized the increase in A_max in A. rufinerve despite its high Rubisco content. Under shade conditions, interspecific differences in leaf features were relatively small. Thus, difference in light demand related to leaf acclimation to high light rather than that to low light in the Acer species.     

The Effect of Salt Stress on the Production of Canthaxanthin and Astaxanthin by Chlorella zofingiensis Grown Under Limited Light Intensity

The fresh water green microalga Chlorella zofingiensisis known to accumulate ketocarotenoids – primarily astaxanthin but also canthaxanthin – when grown under stress conditions of high light irradiance and low nitrogen. We found that salt stress can replace light stress with respect to inducing carotenoid production: cells of C. zofingiensis grown under low light irradiance and subjected to salt and low nitrogen stress accumulated higher amounts of total secondary carotenoids than those growing under high light and low nitrogen stress. Furthermore, C. zofingiensis growing under conditions of salt stress and low light accumulated higher amounts of canthaxanthin than astaxanthin. It is suggested that for canthaxanthin accumulation under salt stress, light is not a limiting factor, but for astaxanthin accumulation high light irradiance is mandatory. These results may be applied in the future for the commercial production of canthaxanthin by C. zofingiensis in systems in which light availability is poor.     

露天煤矿煤粉沉降对矿区周边主要植物的生理影响

随着新疆准东煤田开采的日益加剧,露天开采及运输造成的煤粉尘不仅影响区域大气环境,同时在一定程度上影响周边植物的生理生长。为探讨煤粉尘对周边植物的生理影响,以准东露天煤矿周边人工绿化树种白榆、乡土植物梭梭和柽柳为研究对象,通过野外控制实验,探究覆尘、无尘条件下3种植物的光合生理特性的变化。结果表明:煤粉沉降对3种植物的光合生理均产生了明显影响,覆尘叶片在不同光照强度下净光合速率、气孔导度、蒸腾速率均降低,净光合速率柽柳(18.6%)梭梭(28.3%)白榆(30.7%),气孔导度梭梭(24.1%)柽柳(30.9%)白榆(33.2%),蒸腾速率梭梭(16.7%)白榆(18%)柽柳(39.1%);煤尘对白榆、梭梭的影响主要为非气孔因素,对柽柳的影响有待继续研究。因此露天煤矿开采造成的粉尘会在一定程度上影响周边植物的光合生理过程,抑制其生长,长期排放可能会导致植被覆盖度降低。     

ENVIRONMENTAL AND PHYSIOLOGICAL FACTORS INFLUENCING THE NATURAL DISTRIBUTION OF EVERGREEN AND DECIDUOUS ERICACEOUS SHRUBS ON NORTHEAST AND SOUTHWEST SLOPES OF THE SOUTHERN APPALACHIAN MOUNTAINS. I. IRRADIANCE TOLERANCE

Deciduous and evergreen species are segregated on northeast and southwest slopes of the southern Appalachian Mountains. The segregated distributions of three ericaceous shrubs (Rhododendron maximum valley positions; Rhododendron periclymenoides on northeast slopes; Kalmia latifolia on southwest slopes) were compared to the respective irradiance environments. Growth patterns of field plants, and photosynthetic acclimation of each species to three irradiance treatments in a phytotron were studied. Rhododendron maximum, an evergreen species, was found to be most sensitive to high radiation. In phytotron experiments, quantum yield, light saturated photosynthetic capacity, photosynthesis per chlorophyll, and water use efficiency decreased at high ambient irradiance for R. maximum. These characteristics limit the growth of R. maximum on high irradiance southwestern slopes. Both K. latifolia and R. periclymenoides were able to improve their photosynthetic performance at high ambient irradiance. Rhododendron periclymenoides, a deciduous species, was found to continue increasing leaf conductance at high irradiance without an increase in photosynthesis indicating a possible limitation by water in high light environments such as southwest slopes. Kalmia latifolia, an evergreen species, had reduced photosynthetic capacity and reduced water use efficiency when grown in low irradiance conditions which coincides with the higher K. latifolia abundance on high light, southwestern slopes.     

Stomatal and photosynthetic responses during sun/shade transitions in subalpine plants: influence on water use efficiency

Summary Different response patterns in net photosynthesis (A) leaf conductance (g) and water use efficiency (WUE= a/transpiration) in three subalpine plants occurred during experimental sun/shade transitions that simulated natural cloudcover. In Frasera speciosa Dougl., a large-leaved herb characteristic of open sites, g was relatively insensitive to transitions in irradiance and variations in A. However, large decreases in leaf temperature resulted in reduced transpiration during shade intervals and relatively constant WUE throughout the experimental sun/shade regime. In the understory herb, Arnica cordifolia Hook., patterns of A and g were similar during sun/shade transitions, but WUE was substantially reduced compared to steady-state levels. A third, somewhat intermediate pattern of A, g, and WUE was found in Artemisia tridentata L., an open site shrub. Higher intercellular CO₂ values in A. tridentata suggested that internal, cellular limitations to A were high relative to stomatal limitations in this shrub when compared to the herbaceous species.     

Mycorrhizal colonisation improves fruit yield and water use efficiency in watermelon (Citrullus lanatus Thunb.) grown under well-watered and water-stressed conditions

The effect of arbuscular mycorrhizal (AM) colonisation by Glomus clarum on fruit yield and water use efficiency (WUE) was evaluated in watermelon (Citrullus lanatus) cv. Crimson Sweet F1 under field conditions. Treatments were: (1) well-watered plants without mycorrhizae (WW-M), (2) well-watered plants with mycorrhizae (WW+M), (3) water- stressed plants without mycorrhizae (WS-M) and (4) water-stressed plants with mycorrhizae (WS+M). When soil water tension readings reached –20 and –50 kPa for well-watered (WW) and water-stressed (WS) treatments, respectively, irrigation was initiated to restore the top soil to near field capacity. Water stress reduced watermelon shoot and root dry matter, fruit yield, water use efficiency but not total soluble solids (TSS) in the fruit, compared with the non-stressed treatments. Mycorrhizal plants had significantly higher biomass and fruit yield compared to nonmycorrhizal plants, whether plants were water stressed or not. AM colonisation increased WUE in both WW and WS plants. Macro- (N, P, K, Ca and Mg) and micro- (Zn, Fe and Mn) nutrient concentrations in the leaves were significantly reduced by water stress. Mycorrhizal colonisation of WS plants restored leaf nutrient concentrations to levels in WW plants in most cases. This is the first report of the mitigation of the adverse effect of water stress on yield and quality of a fruit crop.     

Compensatory effects of elevated CO<Subscript>2</Subscript> concentration on the inhibitory effects of high temperature and irradiance on photosynthetic gas exchange in carrots

We determined the interactive effects of irradiance, elevated CO₂ concentration (EC), and temperature in carrot (Daucus carota var. sativus). Plants of the cv. Red Core Chantenay (RCC) were grown in a controlled environmental plant growth room and exposed to 3 levels of photosynthetically active radiation (PAR) (400, 800, 1 200 μmol m⁻² s⁻¹), 3 leaf chamber temperatures (15, 20, 30 °C), and 2 external CO₂ concentrations (C _a), AC and EC (350 and 750 μmol mol⁻¹, respectively). Rates of net photosynthesis (P _N) and transpiration (E) and stomatal conductance (g _s ) were measured, along with water use efficiency (WUE) and ratio of internal and external CO₂ concentrations (C _i/C _a). P _N revealed an interactive effect between PAR and C _a. As PAR increased so did P _N under both C _a regimes. The g _s showed no interactive effects between the three parameters but had singular effects of temperature and PAR. E was strongly influenced by the combination of PAR and temperature. WUE was interactively affected by all three parameters. Maximum WUE occurred at 15 °C and 1 200 μmol m⁻² s^{− 1} PAR under EC. The C _i /C _a was influenced independently by temperature and C _a. Hence photosynthetic responses are interactively affected by changes in irradiance, external CO₂ concentration, and temperature. EC significantly compensates the inhibitory effects of high temperature and irradiance on P _N and WUE.