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1.
Functionality of the photosynthetic system under water stress is of major importance in drought tolerance. Oat (Avena sativa L.) doubled haploid (DH) lines obtained by pollination of F 1 oat crosses with maize were used to assess the differences in plant genotypic response to soil drought. The investigations were based on the measurements of gas exchange and chlorophyll a fluorescence kinetics. Drought was applied to 17-day-old seedlings by withholding water for 14 days and subsequent plant recovery. Non-stressed optimally watered plants served as controls. Yield components were determined when plants reached full maturity. It was shown differences among the oat lines with respect to drought stress susceptibility (SI) and stress tolerance index mean productivity and drought susceptibility index. Sensitivity to drought of individual DH lines was significantly different, as demonstrated by the correlation between drought susceptibility index and yield components, such as dry weight (GW) or grain number (GN) of the harvested plants. GW and GN were lower in drought-sensitive genotypes exposed to drought stress compared to those resistant to drought. The principal component analysis allow to separate three groups of lines differing in their sensitivity to drought stress and indicated that tolerance to drought in oat has a common genetic background.  相似文献   

2.
Phosphites, marketed as foliar fertilizers and resistance activators, have been shown to be useful for the control of diseases in many profitable crops. Despite the importance of white mold, caused by Sclerotinia sclerotiorum, to reduce common bean yield, knowledge of the phosphites´ effect on disease control at the physiological level is still missing. In this study, the leaf gas exchange and chlorophyll a fluorescence parameters variable-to-maximum chlorophyll a fluorescence ratio (Fv/Fm), photochemical yield [Y(II)], yield for dissipation by down-regulation [Y(NPQ)], yield for non-regulated dissipation [Y(NO)], and electron transport rate (ETR) as well as the concentrations of photosynthetic pigments in common bean plants that were sprayed with zinc (Zn) or copper (Cu) phosphites and challenged or not with S. sclerotiorum were determined. Based on the in vitro assays, Zn and Cu phosphites inhibited fungal mycelial growth in a dose-dependent manner, but the Cu phosphite showed to be more fungitoxic. Lesion area and white mold severity were reduced by Zn and Cu phosphites, but the Zn phosphite was more effective. Fungal infection dramatically decreased the values of net carbon assimilation rate, stomatal conductance to water vapor and transpiration rate on non-sprayed plants. Increases in internal CO2 concentration indicated that fungal-induced photosynthetic impairments were chiefly governed by biochemical limitations, but these impairments were greatly abrogated in the Zn and Cu phosphite-sprayed plants. Similarly, the photochemical dysfunctions stemmed from S. sclerotiorum infection were limited in the Zn and Cu phosphite-sprayed plants. Concentrations of chlorophyll a?+?b and carotenoids decreased on inoculated plants, but lower reductions were recorded on Zn and Cu phosphites-sprayed plants. In conclusion, the potential of Zn and Cu phosphites in attenuate the S. sclerotiorum-induced physiological impairments in common bean leaflets was demonstrated and may be an effective mean for managing this disease under field conditions.  相似文献   

3.
Previous works suggested that Pleurostima purpurea (Velloziaceae—Barbacenioideae) shows a remarkable capacity to endure desiccation of its vegetative tissues. P. purpurea occurs in monocotyledons mats on soil islands in the Pão de Açucar (Sugar Loaf) one of the most recognizable rock outcrops of the world, in Rio de Janeiro, southeastern Brazil. Mats of P. purpurea occur in cliffs by the sea some meters above the tidal zone. Although living in rock outcrops almost devoid of any soil cover, P. purpurea seems to occur preferably on less exposed rock faces and slightly shady sites. Usually, less extreme adaptations to drought would be expected in plants with the habitat preference of P. purpurea. Relying on this observation, we argue if a combination of different strategies of dealing with low water availability can be found in P. purpurea as on other desiccation tolerant angiosperms. This study aims to examine the occurrence of desiccation tolerant behavior in P. purpurea together with the expression of drought avoidance mechanisms during dehydration progression. For this, it was analyzed the gas exchanges, leaf pigments and relative leaf water content during desiccation and rehydration of cultivated mature individuals. P. purpurea behaved like typical drought avoiders under moderated drought condition with stomatal closure occurring around a relative leaf water content up to 90%. During this process, it was observed a delay in the leaf relative water content (RWC leaf) decrease comparing to the plant-soil relative water content (RWC plant-soil). As soil dehydration worsened, gas exchanges restrictions progressed until a lack of activity which characterizes anabiosis. The loss of chlorophyll occurs before the end of total dehydration, characterizing the presence of poikilochlorophylly. The chlorophyll degradation follows the RWC leaf decrease, which achieved the minimum average value of 17% without incurring in leaf abscission. The chlorophyll re-synthesis seems to start well after the full rehydration of the leaf. During all of this process, carotenoid content remained stable. These results are coherent with a combination of drought avoidance and desiccation tolerance in P. purpurea which seems to be coherent with the amplitude of water availability in the rock outcrop habitat where it occurs, suggesting that the periods of water availability are sufficiently long for the success of the costly desiccation tolerant behavior but too short to make a typical drought avoider species win the competition for exploring the rock outcrop substrate where P. purpurea occurs.  相似文献   

4.
Abscisic acid (ABA) is an important signaling molecule for plants under drought tolerance. However, ABA itself has many limitations to be used in agriculture practically. Recently, AM1 (ABA-mimicking ligand) has been found to replace ABA. In this study, we have investigated AM1’s potential role for drought tolerance by growing two contrasting rapeseed (Brassica napus L.) genotypes: Qinyou 8 (drought sensitive) and Q2 (drought resistant) with exogenous ABA or AM1 application under well-watered and drought-stressed conditions. Results demonstrate that drought stress has hampered plant growth (relative height growth rate, plant biomass, leaf area), plant water status (leaf relative water content, root moisture content, leaf water potential), photosynthetic gas exchange attributes like net photosynthesis rate (Pn), stomatal conductance (Gs), intercellular CO2 concentration (Ci), transpiration rate (E); chlorophyll fluorescence parameters like photosynthetic efficiency (Fv/Fm), effective quantum yield of PSII (Φ PSII ), photochemical quenching coefficient (qL), electron transport rate (ETR) and chlorophyll content, especially for Qinyou 8 significantly compared to well-watered plants. Whereas increased root/shoot ratio (R/S), water use efficiency (WUE) and non-photochemical quenching (NPQ) was recorded in both genotypes under drought stress. On the other hand, exogenous ABA or AM1 treatment has regulated all the above parameters in a rational way to avoid drought stress. Chloroplast transmission electron microscope images, especially for Qinyou8, have revealed that oxidative stress induced by drought has blurred the grana thylakoids, increased the size or number of plastoglobules due to lipid peroxidation, and the presence of starch granules depict weak capacity to convert them into simple sugars for osmotic adjustment. However, intact grana thylakoid, few plastoglobules with no or very few starch granules were observed in the chloroplast from ABA- or AM1-treated plants under drought. More importantly, AM1-treated plants under drought stress have responded in an extremely similar way like ABA-treated ones. Finally, it is suggested that AM1 is a potential ABA substitute for plant drought tolerance.  相似文献   

5.
An experiment was performed to study gas exchange and chlorophyll fluorescence responses of rice (Oryza sativa L.) to various regimes, such as flooding–midseason drying–flooding (FDF), flooding–midseason drying–saturation (FDS), and flooding–rain-fed (FR) regimes. Compared to FDF, FR resulted in an obvious decrease in net photosynthetic rate (PN), due to the decrease in stomatal conductance and the increase in stomatal limitation. In contrast, FDS plants did not suffer stomatal limitation and had comparable PN with FDF plants. For diurnal light-saturated electron transport rate and saturation irradiance, FDF performed the best, which was followed by FDS and FR successively. FR and FDS plants tended to suffer from midday depression. FDS reduced irrigated water by 17.2% compared to FDF for comparable yields. The results suggested that FDS can be an effective irrigation regime to save water.  相似文献   

6.
The present study aimed to determine effects of drought stress on Lycium ruthenicum Murr. seedlings. Our results showed that mild drought stress was beneficial to growth of L. ruthenicum seedlings. Their height, basal diameter, crown, leaf number, stem dry mass, leaf and root dry mass increased gradually when the soil water content declined from 34.7 to 21.2%. However, with further decrease of the soil water content, the growth of L. ruthenicum seedlings was limited. After 28 d of treatment, the seedlings were apparently vulnerable to drought stress, which resulted in significant leaf shedding and slow growth. However, growth was restored after rehydration. Drought treatments led to a decrease in contents of chlorophyll (Chl) a, b, and Chl (a+b) and increase in the Chl a/b ratio. After rewatering, the Chl content recovered to the content of the control plants. Under drought stress, minimal fluorescence and nonphotochemical quenching coefficient increased, thereby indicating that L. ruthenicum seedlings could protect PSII reaction centres from damage. Maximum fluorescence, maximum quantum yield, actual quantum yield of PSII photochemistry, and photochemical quenching decreased, which suggested that drought stress impacted the openness of PSII reaction centres. A comparison of these responses might help identify the drought tolerance mechanisms of L. ruthenicum. This could be the reference for the planting location and irrigation arrangements during the growing period of L. ruthenicum.  相似文献   

7.
The interactive effects of shade and drought on the morphological and physiological traits of Catalpa bungei plantlets were assessed. Seedling growth, biomass, biomass allocation, leaf morphology, chlorophyll (Chl) content and gas-exchange parameters were measured in plants raised for 3 months under three light levels [80% (HI), 50% (MI), 30% (LI)] and two water levels [moisture (M) and drought (D)]. The results showed that shade greatly decreased growth, biomass, leaf area (LA) and Chl a/b; increased specific leaf area (SLA) and Chl content; and reduced photosynthetic rate (P n). Drought reduced the growth, biomass, LA, SLA, Chl a/b, P n, stomatal conductance (G s), transpiration rate (T r) and intercellular carbon dioxide concentration (C i) and increased the Chl content. Stomatal closure was an early physiological response to water stress. Light, water and their interaction significantly affected plant traits and their bivariate relationships. The phenotypic plasticity index of light (0.47) was much higher than that of water (0.21), indicating that light was the main driver of the variations observed. Under drought stress, growth, biomass, leaf and stem biomass allocation significantly decreased in the HI and MI environments, whereas no significant difference was observed in growth or biomass parameters under the LI condition. Furthermore, no significant difference was observed in P n, G s, or T r under the LI condition under water stress. Our results showed that shade did not alter the negative effects caused by drought stress in MI but did alleviate the negative effects of the LI condition. In summary, the effect of drought on C. bungei plantlets depends on the irradiance conditions.  相似文献   

8.
9.
Medicago sativa L. cv. Longzhong is a nutritious forage plant in dryland regions of the Loess Plateau with strong drought tolerance and broad adaptability. To understand the adaptation mechanism of alfalfa (M. sativa L. cv. Longzhong) to drought stress, growth, and physiological parameters including levels of chlorophyll content, osmotic adjustment, reactive oxygen species (ROS), and antioxidant enzymes and antioxidants were measured under simulated levels of drought (? 0.40, ? 0.80, ? 1.20, ? 1.60, and ? 2.00 MPa). The changes in M. sativa L. cv. Longzhong were compared with those of plants of M. sativa L. cv. Longdong control (Variety I) suited to moderate rainfall areas and M. sativa L. cv. Gannong No. 3 (Variety II) suited to irrigated areas. The results showed that root–shoot ratio, the chlorophyll (a + b) and osmolytes contents, the degree of lipid peroxidation and ROS production, and the levels of antioxidative enzymes and antioxidants increased significantly with increasing drought stress, whereas plant height, aboveground biomass, chlorophyll a/b ratio, leaf water potential (Ψ1), and relative water content (RWC) decreased in response to drought. The Longzhong variety responded early to beginning drought stress (between 0 and ? 0.4 MPa) compared with the controls. Under drought stress (between ? 0.4 and ? 2.0 MPa), the Longzhong variety had significantly higher belowground biomass, root–shoot ratio, Ψ1, RWC, catalase (CAT) activity and reduced glutathione content than those of Varieties I and II, but hydrogen peroxide and hydroxyl free radical (OH·) contents were significantly lower. Step regression analysis showed that OH·, CAT, malondialdehyde, superoxide anion-free radical (O 2 ·? ), and superoxide dismutase of Longzhong had the most marked response to drought stress. In conclusion, the stronger drought tolerance of the Longzhong variety might be due to its higher water-holding capacity, root–shoot ratio, and ability to coordinate enzymatic and non-enzymatic antioxidant systems, which coordinate the peroxidation and oxidative systems.  相似文献   

10.
Jatropha curcas and Jatropha mollissima plants were evaluated under conditions of high (HSM) and low (LSM) soil moisture in a semi-arid environment, as changes in the content and concentration of epicuticular wax and the leaf metabolism which could have a relationship with drought tolerance. Besides epicuticular wax, gas exchange, antioxidant system and biochemical parameters of the photosynthetic metabolism were measured. The epicuticular wax content increased only in J. mollissima leaves 95 % under LSM, when compared with HSM conditions. Therefore, J. curcas invested less in the production of long-chain n-alkanes than did J. mollissima under LSM conditions. J. mollissima plants showed the highest CO2 assimilation rate during the HSM period compared to J. curcas. Both species showed high stability in some leaf biochemistry products, highlighting the highest sugar content, free amino acids, total soluble protein, and photosynthetic pigments in the leaves of J. mollissima plants under both of the soil moisture conditions. Moreover, the stability and performance of the different parameters, such as morphologic variables, seem to allow J. mollissima plants to tolerate semi-arid conditions.  相似文献   

11.
Aquaporin proteins are part of the complex response of common bean (Phaseolus vulgaris L.) to drought which affects the quality and quantity of yield of this important crop. To better understand the role of aquaporins in common bean, drought-induced gene expression of several aquaporins was determined in two cultivars, the more drought tolerant Tiber and the less tolerant Starozagorski ?ern. The two bean cultivars were selected among 16 European genotypes based on the tolerance to drought determined by time needed for plants to wilt after withholding irrigation and yield at harvest. The expression patterns of two plasma membrane intrinsic proteins, PvPIP1;2 and PvPIP2;7, and two tonoplast intrinsic proteins, PvTIP1;1 and PvTIP4;1 in leaves of 21 day old plants were determined by RT-qPCR in both cultivars under three degrees of drought stress, and under rehydration and control conditions. Gene expression of all four examined aquaporins was down-regulated in drought stressed plants. After rehydration it returned to the level of control plants or was even higher. The responses of PvPIP2;7 and PvTIP1;1 during drought and rehydration were particularly pronounced. The gene expression of PvPIP2;7 and PvTIP4;1 during drought was cultivar specific, with greater down-regulation of these two aquaporins in drought tolerant Tiber. Under drought stress the relative water content and water potential of leaves were higher in Tiber than in Starozagorski plants. The differences in these physiological parameters indicate greater prevention of water loss in Tiber during drought, which may be associated with rapid and adequate down-regulation of aquaporins. These results suggest that the ability of plants to conserve water during drought stress involves timely and sufficient down-regulation of gene expression of specific aquaporins.  相似文献   

12.
Salvia sinaloensis Fern. (sage) is a medicinal plant containing plant secondary metabolites (PSMs) with antioxidant properties. The current study investigated the effects of drought stress on S. sinaloensis morphological and ecophysiological traits, and active constituent production. Sage plants were cultivated in controlled conditions for 34 days and exposed to full irrigation as control, half irrigation, or no irrigation. Changes in growth index (G.I.), dry biomass, leaf water potential (LWP), physiological parameters, active compounds, volatilome (BVOCs) and essential oils (EOs) were determined. Not irrigated plants showed a decrease in total chlorophyll content (~???14.7%) and growth (G.I., ~???59.4%) from day 18, and dry biomass at day 21 (??56%), when the complete leaf withering occurred (LWP, ??1.10 MPa). Moderate drought stressed plants showed similar trends for chlorophyll content and growth but kept a constant LWP (??0.35 MPa) and dry biomass throughout the experiment, as control plants. Carotenoids were not affected by water regimes. The photosynthetic apparatus tolerated mild to severe water deficits, without a complete stomatal closure. Plants under both stress conditions increased the percentage of phenols and flavonoids and showed altered BVOC and EO chemical profiles. Interestingly Camphor, the main EO oxygenated monoterpene, increased in moderate stressed plants while the sesquiterpene hydrocarbon Germacrene D decreased. The same trend was seen in the headspace under stress severity. The data evidenced a possible role of the active molecules in the response of S. sinaloensis plants to drought stress. Taking together, these findings point at S. sinaloensis as a potential drought adaptive species, which could be used in breeding strategies to obtain sages with high quality PSMs, saving irrigation water.  相似文献   

13.
14.
The influence of arbuscular mycorrhiza (AM) and drought stress on aquaporin (AQP) gene expression, water status, and photosynthesis was investigated in black locust (Robinia pseudoacacia L.). Seedlings were grown in potted soil inoculated without or with the AM fungus Rhizophagus irregularis, under well-watered and drought stress conditions. Six full-length AQP complementary DNAs (cDNAs) were isolated from Robinia pseudoacacia, named RpTIP1;1, RpTIP1;3, RpTIP2;1, RpPIP1;1, RpPIP1;3, and RpPIP2;1. A phylogenetic analysis of deduced amino acid sequences demonstrated that putative proteins coded by these RpAQP genes belong to the water channel protein family. Expression analysis revealed higher RpPIP expression in roots while RpTIP expression was higher in leaves, except for RpTIP1;3. AM symbiosis regulated host plant AQPs, and the expression of RpAQP genes in mycorrhizal plants depended on soil water condition and plant tissue. Positive effects were observed for plant physiological parameters in AM plants, which had higher dry mass and lower water saturation deficit and electrolyte leakage than non-AM plants. Rhizophagus irregularis inoculation also slightly increased leaf net photosynthetic rate and stomatal conductance under well-watered and drought stress conditions. These findings suggest that AM symbiosis can enhance the drought tolerance in Robinia pseudoacacia plants by regulating the expression of RpAQP genes, and by improving plant biomass, tissue water status, and leaf photosynthesis in host seedlings.  相似文献   

15.
Drought and salinity are severe environmental stresses and limit soybean growth. In this study, a comparative analysis of physiological and molecular responses of two soybean (Glycine max L.) genotypes to these stresses was carried out. Plants of drought-tolerant genotype RD (cv. FD92) and sensitive genotype SD (cv. Z1303) were exposed to 15 % (m/v) PEG 6000, which simulated drought stress, or 150 mM NaCl. The RD plants maintained larger leaf area and higher net photosynthetic rate, chlorophyll content, stomatal conductance, and relative water content compared with the SD plants. Leaf proline content increased under both stresses more in RD than in SD. The drought tolerance of RD plants was also correlated with greater antioxidant activity and lower content of hydrogen peroxide and malondialdehyde under stress conditions. Amounts of abscisic acid, jasmonic acid, and salicylic acid under stress increased to a greater extent in RD than in SD plants. At the molecular level, the effects of 20-d stress treatments were manifested by relatively higher expression of drought- or salt-related genes: GmP5CS, GmDREB1a, GmGOLS, GmBADH, and GmNCED1 in RD plants than in SD plants. These results form the basis for understanding the mechanisms of the drought- and salt-stress tolerance in soybean.  相似文献   

16.
Christolea crassifolia HARDY: gene (CcHRD) belongs to the AP2/ERF-like tanscritpion factor family, and overexpression of HRD gene has been proved to result in improved water use efficiency and enhanced drought resistance in multiple plant species. In the present study, we cloned the CcHRD gene from Christolea crassifolia, which shares 99.1% sequence similarity with the HRD gene from Arabidopsis thaliana. We generated transgenic tomato plants expressing CcHRD gene by agrobacterium-mediated genetic transformation. Our results revealed that the transgenic tomato plants showed a more developed root system and higher fruit yield than the wild-type plants. Furthermore, the leaf relative water content, chlorophyll content and Fv/Fm value in transgenic plants were significantly higher than the wild type, while the relative conductivity and MDA content of transgenic plant leaves were markedly lower than those of wild type under drought stress. We also observed that the major agronomic traits of transgenic tomato plants were improved under natural drought stress compared with those of the wild type. In summary, results in this transgenic study showed that the CcHRD gene could enhance the drought resistance in tomato, and also provided important information for the application of drought-responsive genes in improving crop plant resistance to abiotic stresses.  相似文献   

17.
Insufficient attention has been paid to the physiological responses of sesame to drought and it is unclear if exogenous plant growth regulators are beneficial to drought-stressed sesame. Thus, a field study was conducted on seven Sesamum indicum genotypes affected by two levels of irrigation (60 and 80% depletions in available soil water) and by foliar-applied salicylic acid (SA; 0 and 0.6 mM). Water deficit led to depressions in net photosynthetic rate, stomatal conductance, leaf area index, chlorophyll a, b, and total chlorophyll contents, maximum quantum efficiency of PSII, and plant dry matter and seed yield, despite increases in carotenoid concentration, superoxide dismutase, catalase, peroxidase, and ascorbate peroxidase activities. SA was found beneficial in ameliorating the depressions in all of the above characteristics, indicating that it could be applied for lessening the harmful effects of the drought stress.  相似文献   

18.
Two greenhouse experiments were conducted in order to investigate the effects of different levels of water stress on gas exchange, chlorophyll fluorescence, chlorophyll content, antioxidant enzyme activities, lipid peroxidation, and yield of tomato plants (Solanum lycopersicum cv. Jinfen 2). Four levels of soil water content were used: control (75 to 80% of field water capacity), mild water stress (55 to 60%), moderate water stress (45 to 50%), and severe water stress (35 to 40%). The controlled irrigation was initiated from the third leaf stage until maturity. The results of two-year trials indicated that the stomatal conductance, net photosynthetic rate, light-saturated photosynthetic rate, and saturation radiation decreased generally under all levels of water stress during all developmental stages, while compensation radiation and dark respiration rate increased generally. Water stress also declined maximum quantum yield of PSII photochemistry, electron transfer rate, and effective quantum yield of PSII photochemistry, while nonphotochemical quenching increased in all developmental stages. All levels of water stress also caused a marked reduction of chlorophyll a, chlorophyll b, and total chlorophyll content in all developmental stages, while activities of antioxidant enzymes, such as superoxide dismutase, peroxidase, and catalase, and lipid peroxidation increased.  相似文献   

19.
The variation of light intensity has obvious effects on leaf external morphology, internal anatomy, and physiological characteristics; it even induces changes in secondary metabolite production. The effects of different irradiance levels on biomass, gas exchange parameters, and photosynthetic pigment contents in Mahonia bodinieri (Gagnep.) Laferr. were analyzed here. Combined analyses of physiology, cytology, and HPLC were used to study the differences in leaf morphology, structure, physiological characters, and alkaloid content in response to different irradiances. The results indicated that the highest foliar biomass was observed under I 50 (50 % of full sunlight) followed by I 30 (30 % of full sunlight), the highest net photosynthetic rate, stomatal conductance, transpiration rate values were observed under I 30 followed by I 50, and lower values occurred in I 10 (10 % of full sunlight) and I 100 (full sunlight). With increased light intensity, total leaf area and the contents of chlorophyll a (Chl a), chlorophyll b (Chl b), and chlorophyll (Chl a+b) per unit leaf area were clearly reduced, whereas leaf mass per area, carotenoid content, leaf thickness, thickness of palisade and spongy parenchyma, and stomatal density were all significantly increased. Electron microscopic observation revealed that the number of grana, stroma lamellae and the number of starch grains in chloroplasts were decreased, the number of plastoglobuli was increased when irradiance levels increased. The estimated total yield of alkaloids in a single plant was higher under I 30 and I 50 than under I 10 or I 100 as a result of the higher biomass of the plants. Therefore, I 30 and I 50 were not only beneficial to increase biomass, but also suitable for the synthesis and accumulation of the major secondary metabolites (alkaloids). Our findings provide valuable data for the determination and regulation of irradiance levels during artificial cultivation of M. bodinieri.  相似文献   

20.
Optical characteristics, contents of photosynthetic pigments, total soluble sugars, and starch, rates of gas exchange, chlorophyll (Chl) a fluorescence, and leaf water relations were analysed in three Vitis vinifera L. cultivars, Tinto Cão (TC), Touriga Nacional (TN), and Tinta Roriz (TR), grown in Mediterranean climate. Chl content was significantly lower in TC than in TN and TR leaves, while the Chl a/b ratio was higher. TR had the lowest net photosynthetic rate, stomatal conductance, and contents of soluble sugars and starch than TN and TC. In spite of low Chl content, TC showed the lowest photon absorbance and the highest photochemical efficiency of photosystem 2. TC had the lowest predawn and midday leaf water potential. The capability for osmotic adjustment was similar among cultivars and the calculated modulus of elasticity was higher in TC leaves. The typical lighter green leaves of TC seemed to be an adaptive strategy to high irradiance and air temperature associated to water stress.  相似文献   

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