Water relations and photosynthetic capacity of two species of Calotropis in a tropical semi-arid ecosystem

Background and Aims

Calotropis procera and Calotropis gigantea, originally from warm parts of Africa and Asia, are now pan-tropical and in ecological terms considered an indicator of overgrazed, disturbed lands; they grow successfully in dry areas. Variations in water relations, morphology and photosynthesis of the two species growing in the same habitat were studied to assess possible mechanisms of tolerance to drought and how these relate to their ecophysiological success. Also the hypothesis that their photosynthetic rate (A) under drought would be affected by stomatal and non-stomatal limitations was tested.

Methods

Water relations, gas exchange, water use efficiency (WUE), fluorescence parameters, pubescence and specific leaf area (SLA) of Calotropis procera and C. gigantea plants growing in the field were evaluated during the wet (WS) and dry (DS) seasons.

Results

The xylem water potential (ψ) was similar in both species during the WS and DS; drought caused a 28 % decrease of ψ. In C. procera, A, stomatal conductance (g_s) and carboxylation efficiency (CE) were higher in the WS with half the values of those during the DS, this species being more affected by drought than C. gigantea. A high δ¹³C of C. gigantea (–26·2 ‰) in the WS indicated a higher integrated WUE, in agreement with its lower g_s. Leaves of C. gigantea were more pubescent than C. procera. Relative stomatal and non-stomatal limitation of A increased with drought in both species; no changes in maximum quantum yield of photosystem II (PSII; F_v/F_m) were observed. The decrease in the relative quantum yield of PSII (φ_PSII) and in the photochemical quenching coefficient (q_P) was more pronounced in C. procera than in C. gigantea.

Conclusions

The photosynthetic capacity of C. procera was higher than that of C. gigantea. During the DS, A was regulated by stomatal and non-stomatal factors in a coordinated manner and drought did not cause chronic photoinhibition. A higher density of trichomes and leaf angle in C. gigantea may contribute to the maintenance of A and confer more efficient protection of photochemical activity in the DS. Ecophysiological traits such as high photosynthetic rate throughout the year even during the DS, and high WUE, highly pubescent leaves and low SLA observed in both species contribute to the establishment and growth of Calotropis in dry conditions.     

Decreased CO2 availability and inactivation of Rubisco limit photosynthesis in cotton plants under heat and drought stress in the field

Heat and drought stresses are often coincident and constitute major factors limiting global crop yields. A better understanding of plant responses to the combination of these stresses under production environments will facilitate efforts to improve yield and water use efficiencies in a climatically changing world. To evaluate photosynthetic performance under dry-hot conditions, four cotton (Gossypium barbadense L.) cultivars, Monseratt Sea Island (MS), Pima 32 (P32), Pima S-6 (S6) and Pima S-7 (S7), were studied under well-watered (WW) and water-limited (WL) conditions at a field site in central Arizona. Differences in canopy temperature and leaf relative water content under WL conditions indicated that, of the four cultivars, MS was the most drought-sensitive and S6 the most drought-tolerant. Net CO₂ assimilation rates (A) and stomatal conductances (gs) decreased and leaf temperatures increased in WL compared to WW plants of all cultivars, but MS exhibited the greatest changes. The response of A to the intercellular CO₂ concentration (A–C_i) showed that, along with stomatal closure, non-stomatal factors associated with heat stress also limited A under WL conditions, especially in MS. The activation state of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) decreased in WL compared to WW plants, consistent with thermal inhibition of Rubisco activase activity. The extent of Rubisco deactivation could account for the metabolic limitation to photosynthesis in MS. Taken together, these data reveal the complex relationship between water availability and heat stress for field-grown cotton plants in a semi-arid environment. Both diffusive (drought-stress-induced) and biochemical (heat-stress-induced) limitations contributed to decreased photosynthetic performance under dry-hot conditions.     

羊草叶片气体交换参数对温度和土壤水分的响应

 采用生长箱控制的方法研究了羊草(Leymus chinensis)幼苗叶片光合参数对5个温度和5个水分梯度的响应和适应。结果表明：轻度、中度土壤干旱并没有限制羊草叶片的生长,对气体交换参数亦无显著影响,反映了羊草幼苗对土壤水分胁迫的较高耐性。叶片生物量以26 ℃时最大,其它依次为23 ℃、20 ℃、29 ℃和32 ℃。温度升高使气孔导度和蒸腾速率增加, 却使光合速率和水分利用效率降低。水分和温度对叶片生物量、光合速率、气孔导度和蒸腾速率存在显著的交互作用,表明高温加强了干旱对叶片生长和气体交换的影响, 降低了羊草对土壤干旱的适应能力。高温和干旱的交互作用将显著减少我国半干旱地区草原的羊草生产力。     

Effects of temperature on leaf hydraulic architecture of tobacco plants

Main Conclusion

Modifications in leaf anatomy of tobacco plants induced greater leaf water transport capacity, meeting greater transpirational demands and acclimating to warmer temperatures with a higher vapor pressure deficit. Temperature is one of the most important environmental factors affecting photosynthesis and growth of plants. However, it is not clear how it may alter leaf hydraulic architecture. We grew plants of tobacco (Nicotiana tabacum) ‘k326’ in separate glasshouse rooms set to different day/night temperature conditions: low (LT 24/18 °C), medium (MT 28/22 °C), or high (HT 32/26 °C). After 40 days of such treatment, their leaf anatomies, leaf hydraulics, photosynthetic rates, and instantaneous water-use efficiency (WUE_i) were measured. Compared with those under LT, plants exposed to HT or MT conditions had significantly higher values for minor vein density (MVD), stomatal density (SD), leaf area, leaf hydraulic conductance (K _leaf), and light-saturated photosynthetic rate (A _sat), but lower values for leaf water potential (ψ _l) and WUE_i. However, those parameters did not differ significantly between HT and MT conditions. Correlation analyses demonstrated that SD and K _leaf increased in parallel with MVD. Moreover, greater SD and K _leaf were partially associated with accelerated stomatal conductance. And then stomatal conductance was positively correlated with A _sat. Therefore, under well-watered, fertilized conditions, when relative humidity was optimal, changes in leaf anatomy seemed to facilitate the hydraulic acclimation to higher temperatures, meeting greater transpirational demands and contributing to the maintenance of great photosynthetic rates. Because transpiration rate increased more with temperature than photosynthetic rate, WUE_i reduced under warmer temperatures. Our results indicate that the modifications of leaf hydraulic architecture are important anatomical and physiological strategies for tobacco plants acclimating to warmer temperatures under a higher vapor pressure deficit.     

Midday depression in net photosynthesis and stomatal conductance in Yucca glauca

Summary Yucca glauca in the Colorado shortgrass prairie undergoes a pronounced midday depression in net photosynthesis and stomatal conductance under summer field conditions. This phenomenon can be duplicated in the laboratory using potted plants by simulating a typical summer daily pattern of leaf temperature and leaf-to-air water vapor concentration difference (w). The decrease in photosynthetic rate appears to be due primarily to high leaf temperatures, while the decrease in stomatal conductance can be attributed mainly to high w values. Stomatal conductance also decreases when leaf temperatures exceed a critical threshold value, even when w is artificially maintained at a constant level. The threshold temperature is commonly attained for leaves in situ, but only after substantial stomatal closure has already occurred as a result of high w values.The photosynthetic temperature optimum and threshold temperature which promotes stomatal closure increases substantially as the growing season progresses. As a result, the midday depression in photosynthesis occurs at higher temperatures in mid-summer than in late spring. Preliminary evidence suggests that the photosynthetic temperature optimum closely follows the naturally-occurring morning leaf temperatures, while the threshold temperature for stomatal closure matches afternoon leaf temperatures.     

Earlier summer drought affects leaf functioning of the Mediterranean species Cistus monspeliensis L.

The Mediterranean vegetation is characterized by a high diversity of growth forms, habits and phenology that enable it to endure under harsh environmental conditions. It is however unclear whether these adaptations may allow plant survival under more extreme conditions, as predicted by climatic models under the perspective of climate change. A manipulative experiment aiming at anticipating summer aridity has been run to analyse the effects of the experimental drought on spring-leaf functioning and characteristics of the leaf-dimorphic Mediterranean shrub Cistus monspeliensis L.Assimilation rates were reduced under anticipated summer aridity due to a decrease of stomatal conductance, but only before morphological adaptations to drought (an increase of leaf mass per area) occurred. These adaptations were anticipated under experimental dry conditions, and causes photosynthetic performances to recover compared to previous dates. When natural summer aridity occurred, the leaf mass per area also changed in the control. However, this causes no recovery of the photosynthetic performances, because of the decrease of stomatal conductance due to low soil water content and leaf water potential values. Moreover, under experimental drought, leaf shedding was anticipated to reduce water losses, causing an overall reduction of leaf lifespan.     

Changes in leaf epicuticular wax,gas exchange and biochemistry metabolism between <Emphasis Type="Italic">Jatropha mollissima</Emphasis> and <Emphasis Type="Italic">Jatropha curcas</Emphasis> under semi-arid conditions

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.     

Photosynthesis in Encelia farinosa Gray in Response to Decreasing Leaf Water Potential

Photosynthetic responses of intact leaves of the desert shrub Encelia farinosa were measured during a long term drought cycle in order to understand the responses of stomatal and nonstomatal components to water stress. Photosynthetic rate at high irradiance and leaf conductance to water vapor both decreased linearly with declining leaf water potential. The intercellular CO₂ concentration (c_i) remained fairly constant as a function of leaf water potential in plants subjected to a slow drought cycle of 25 days, but decreased in plants exposed to a 12-day drought cycle. With increasing water stress, the slope of the dependence of photosynthesis on c_i (carboxylation efficiency) decreased, the maximum photosynthetic rates at high c_i became saturated at lower values, and water use efficiency increased. Both the carboxylation efficiency and photosynthetic rates were positively correlated with leaf nitrogen content. Associated with lower leaf conductances, the calculated stomatal limitation to photosynthesis increased with water stress. However, because of simultaneous changes in the dependence of photosynthesis on c_i with water stress, increased leaf conductance alone in water-stressed leaves would not result in an increase in photosynthetic rates to prestressed levels. Both active osmotic adjustment and changes in specific leaf mass occurred during the drought cycle. In response to increased water stress, leaf specific mass increased. However, the increases in specific leaf mass were associated with the production of a reflective pubescence and there were no changes in specific mass of the photosynthetic tissues. The significance of these responses for carbon gain and water loss under arid conditions are discussed.     

Green roof <Emphasis Type="Italic">Petunia</Emphasis>, <Emphasis Type="Italic">Ageratum</Emphasis>, and <Emphasis Type="Italic">Mentha</Emphasis> responses to water stress,seaweeds, and trinexapac-ethyl treatments

The availability of sufficient irrigation water and the development of drought-tolerant species are challenging factors in the design and maintenance of green roofs in modern cities. Green roof plants of Petunia hybrid Headliner^® Red Star, Ageratum hybrid Artist^® blue, and Mentha spicata L. grown in a simulated green roof pot system under controlled greenhouse conditions. The plants were watered every 2 or 6 days (2DWI/6DWI) for 8 weeks accompanied by either a 6-day treatment of seaweed extracts of Ascophyllum nodosum as a soil drench or foliar spray, or two concentrations of trinexapac-ethyl (TE) biweekly sprays. Following treatments, leaf number, leaf area, dry weights, plant height, stomatal conductanse, photosynthetic and transpiration rates and leaf water potential and relative water content were determined in two seasons during 2016 and 2017. The prolonged irrigation intervals reduced plant growth as revealed by morphological and physiological parameters. The application of SWE as drench treatment improved Petunia and Ageratum plant vegetative growth, stomatal conductance, photosynthetic and transpiration rates and leaf water potential and relative water content during prolonged irrigation intervals. TE increased the vegetative growth as well as the physiological performance of Ageratum plants. However, neither SWE nor TE treatments improved the performance of Mentha plants under prolonged irrigation intervals. It was suggested that improved photosynthetic rates were stimulated by enhanced stomatal conductance leading to improved leaf water potential as well as increased relative water content during prolonged irrigation conditions.     

Photosynthesis and micro-environmental factors in a spring ephemeral,Erythronium japonicum, from native and open habitats

Environmental factors affecting photosynthetic activity of the typical vernal speciesErythronium japonicum Decne were examined on the floor of a deciduous broad-leavedQuercus mongolica forest (Q.m. stand) and on bare land left undisturbed for 4 years after forest clearing (bare stand). Daytime solar irradiation and air and leaf temperatures at the bare stand were significantly higher than those at theQ.m. stand. The relative air humidity was very low and did not differ much between the stands, although the leaf-air vapor pressure differences (VPD) at the bare stand were about twice as high as those at theQ.m. stand. The plants at both stands were supplied with sufficient soil water throughout their growing season by a large snowmelt. However, the aboveground parts of the plants at the bare stand were subjected to much more severe heat stress, caused by the strong radiations and high leaf temperatures, and water stress, caused by the highly transpiring conditions, than those at theQ.m. stand. When the radiation on leaves, leaf temperatures and VPD in the assimilation chamber were changed from those observed at theQ.m. stand to those at the bare stand, the photosynthetic rate and stomatal conductance fell significantly. However, the rate and conductance were immediately restored to the respective values near those measured under the conditions at theQ.m. stand when only VPD was dropped to the value similar to that observed at theQ.m. stand. These results indicate that the photosynthetic rate of the plants at the bare stand was lowered largely by a decrease in stomatal conductance. The internal CO₂ partial pressure was considerably greater in leaves placed under environmental conditions similar to those at the bare stand.     

Influence of Drought,Rain and Artificial Irrigation on Photosynthesis,Gas Exchange and Water Relations of the Fynbos Plant Protea acaulos (L.) Reich at the End of the Dry Season

Protea acaulos, a prostrate fynbos shrub, often experiences very low air humidity at leaf temperatures over 10°C higher than mean air temperature. We determined to what degree this particular microclimate influenced photosynthetic performance, leaf conductance and water relations of non-irrigated and trickle-irrigated plants. Measurements were made at the end of the dry summer season in the sand plain lowland fynbos on the west coast of South Africa. Independent of water supply, plants showed a pronounced midday depression of gas exchange. While in non-irrigated plants leaf water potential dropped to ? 2.0 MPa around noon, it never fell below ?1.0 MPa in irrigated plants. On the other hand minimum pressure potential was similar in irrigated and non-irrigated plants. The latter showed higher turgor after rain, due to osmotic acclimation, which resulted from a reduction in maximum water volume. The main osmoticum was 1,5-anhydro-D-glucitol. Leaf temperature, directly or via the vapour pressure deficit between leaf and air (Δw), rather than plant water status, was the determinant of the midday depression of gas exchange. High Δw caused stomatal closure during times of saturating light, thus limiting photosynthetic CO₂ uptake and availability and enhancing the susceptibility for photoinhibition. This, as well as high leaf temperature per se, decreased the efficiency of photochemistry of photosystem II. Initial fluorescence remained constant until temperatures exceeded 35 °C, above which changes in fluorescence indicated both photoinhibition and heat stress. Unlike other fynbos plants, Protea acaulos could not use the improved soil water supply to increase carbon gain under hot summer condition.     

干旱胁迫下杨树无性系苗期光合与气孔形态变异研究

为了解不同干旱胁迫处理下,杨树叶片光合特性和气孔形态的变化规律,本研究以4个杨树无性系为材料,对其干旱胁迫下光合指标与气孔形态指标进行测定分析,方差分析结果表明：除气孔器长在处理间差异不显著外,其他性状在无性系间、处理间和无性系与处理交互作用间均达极显著差异水平。随着土壤相对含水量的逐渐降低,无性系净光合速率、蒸腾速率和气孔导度都呈现出不同程度的下降;而胞间二氧化碳浓度呈现为先降后升趋势,表明当土壤相对含水量大于40%时,光合作用的主要限制因子是气孔因素,当土壤相对含水量低于40%时,光合作用的主要限制因子转换为非气孔因素。气孔形态研究结果表明,各指标均随土壤相对含水量的持续降低呈现出持续下降趋势。利用隶属函数法对不同干旱胁迫下4个杨树无性系的抗旱性进行综合评价,结果表明：白城小青黑杨的抗旱性优于其他三个无性系。     

Differential adaptation of two varieties of common bean to abiotic stress: II. Acclimation of photosynthesis

The photosynthetic characteristics of two contrasting varieties of common bean (Phaseolus vulgaris) have been determined. These varieties, Arroz and Orfeo, differ in their productivity under stress conditions, resistance to drought stress, and have distinctly different stomatal behaviour. When grown under conditions of high irradiance and high temperature, both varieties displayed evidence of photosynthetic acclimation at the chloroplast level-there was an increase in chlorophyll a/b ratio, a decreased content of Lhcb proteins, and an increased xanthophyll cycle pool size. Both varieties also showed reduced chlorophyll content on a leaf area basis and a decrease in leaf area. Both varieties showed an increase in leaf thickness but only Arroz showed the characteristic elongated palisade cells in the high light-grown plants; Orfeo instead had a larger number of smaller, rounded cells. Differences were found in stomatal development: whereas Arroz showed very little change in stomatal density, Orfeo exhibited a large increase, particularly on the upper leaf surface. It is suggested that these differences in leaf cell structure and stomatal density give rise to altered rates of photosynthesis and stomatal conductance. Whereas, Arroz had the same photosynthetic rate in plants grown at both low and high irradiance, Orfeo showed a higher photosynthetic capacity at high irradiance. It is suggested that the higher yield of Orfeo compared with Arroz under stress conditions can be explained, in part, by these cellular differences.     

Water relations and gas exchange in Coespeletia moritziana (Sch. Bip) Cuatrec., a giant rosette species of the high tropical Andes

Giant rosettes are ones of the most striking features of the vegetation in the high tropical Andes, with Coespeletia moritziana reaching the highest altitudes up to 4,600 m a.s.l. Different from other giant rosettes, this species grows on rock outcrops with poorly developed soils and where water availability may be limited. Two questions are addressed in this study: How does this species respond in terms of water relations to maintain favorable gas-exchange conditions? Considering that adult plants rely on a water-reserving central pith, how do early stages respond to this environment??s extreme conditions? Water relations and gas-exchange studies were carried out on juveniles, intermediate and adult C. moritziana plants during wet and dry seasons in Páramo de Piedras Blancas at 4,200 m a.s.l. Adult plants maintained higher leaf water potentials (??_L) during the wet season, however, no differences between stages were found for the dry season. Minimum dry season ??_L were never near the turgor loss point in any of the stages. Juveniles show a more strict stomatal control during the dry season to maintain a favorable water status. Net photosynthesis significantly decreased in intermediate and juvenile stages from wet to dry seasons. Our results suggest that C. moritziana resists more extreme conditions compared to other Andean giant rosettes     

Photosynthesis and water relations of well-watered orange plants as affected by winter and summer conditions

The aim of this study was to evaluate how the summer and winter conditions affect the photosynthesis and water relations of well-watered orange trees, considering the diurnal changes in leaf gas exchange, chlorophyll (Chl) fluorescence, and leaf water potential (Ψ) of potted-plants growing in a subtropical climate. The diurnal pattern of photosynthesis in young citrus trees was not significantly affected by the environmental changes when compared the summer and winter seasons. However, citrus plants showed higher photosynthetic performance in summer, when plants fixed 2.9 times more CO₂ during the diurnal period than in the winter season. Curiously, the winter conditions were more favorable to photosynthesis of citrus plants, when considering the air temperature (< 29 °C), leaf-to-air vapor pressure difference (< 2.4 kPa) and photon flux density (maximum values near light saturation) during the diurnal period. Therefore, low night temperature was the main environmental element changing the photosynthetic performance and water relations of well-watered plants during winter. Lower whole-plant hydraulic conductance, lower shoot hydration and lower stomatal conductance were noticed during winter when compared to the summer season. In winter, higher ratio between the apparent electron transport rate and leaf CO₂ assimilation was verified in afternoon, indicating reduction in electron use efficiency by photosynthesis. The high radiation loading in the summer season did not impair the citrus photochemistry, being photoprotective mechanisms active. Such mechanisms were related to increases in the heat dissipation of excessive light energy at the PSII level and to other metabolic processes consuming electrons, which impede the citrus photoinhibition under high light conditions.     

The influence of water stress on the photosynthetic performance and stomatal behaviour of tree seedlings subjected to variation in temperature and irradiance

Summary Seedlings of Betula pendula Roth. and Gmelina arborea L. were subjected to variation in temperature and irradiance. The influence of a mild water-stressing treatment on the photosynthetic performance and stomatal behaviour of these plants was assessed. For both species, the shape of the relationships between irradiance and photosynthesis and temperature and photosynthesis resembled those reported for other species. The effect of water stress was to reduce the rate of photosynthesis, particularly at high temperatures. This was largely a function of a reduction in mesophyll conductance under these conditions. The optimum temperature for stomatal opening was significantly lower than the optimum temperature for photosynthesis, which was in turn lowered by the water stress treatment. The stomata of birch seedlings showed maximum opening at an intermediate temperature while the stomata of Gmelina generally exhibited a closing movement when leaf temperatures increased from 15° C. Mesophyll conductances of both species increased with increasing temperature.The physiological basis for the variation in photosynthetic performance and stomatal behaviour and the ecological significance of this variation are discussed.     

Stomatal versus biochemical limitations to dynamic photosynthetic performance in four tropical rainforest shrub species

Photosynthetic performance under dynamic light regimes was assessed in four different species of tropical shrubs from the family Rubiaceae via field gas exchange measurements conducted on Barro Colorado Island, Panamá. Rates of photosynthetic induction and induction loss were assessed throughout the day in both the wet and dry seasons in order to determine the relative roles of stomata and biochemistry in limiting photosynthetic performance under transient light conditions. A high degree of coordination was observed between stomatal conductance and biochemical capacity for CO₂ assimilation during induction. Rates of biochemical and overall photosynthetic induction sharply decreased when initial stomatal conductance fell below a narrow range of critical values. Time of day or season did not affect rates of biochemical deactivation upon shading, but did influence stomatal closure, which often exerted a significant influence over induction loss in the darkness. In measurements of total assimilation due to a 60-s light pulse, both biochemical activity and stomatal conductance were linearly related to total CO₂ uptake. Only during the mornings of the wet season was stomatal conductance consistently high enough to be non-limiting to dynamic photosynthetic performance. At all other times, stomatal behavior exercised significant influence over induction times, photosynthetic induction loss, and total CO₂ uptake from 60-s light pulses. Received: 17 March 1999 / Accepted: 26 October 1999     

Seasonal and diurnal changes in photosynthetic limitation of young sweet orange trees

This study tests the hypothesis that potted sweet orange plants show a significant variation in photosynthesis over seasonal and diurnal cycles, even in well-hydrated conditions. This hypothesis was tested by measuring diurnal variations in leaf gas exchange, chlorophyll fluorescence, leaf water potential, and the responses of CO₂ assimilation to increasing air CO₂ concentrations in 1-year-old ‘Valência’ sweet orange scions grafted onto ‘Cleopatra’ mandarin rootstocks during the winter and summer seasons in a subtropical climate. In addition, diurnal leaf gas exchange was evaluated under controlled conditions, with constant environmental conditions during both winter and summer. In relation to our hypothesis, a greater rate of photosynthesis is found during the summer compared to the winter. Reduced photosynthesis during winter was induced by cool night conditions, as the diurnal fluctuation of environmental conditions was not limiting. Low air and soil temperatures caused decreases in the stomatal conductance and in the rates of the biochemical reactions underlying photosynthesis (ribulose-1,5-bisphosphate (RuBP) carboxylation and RuBP regeneration) during the winter compared to the values obtained for those markers in the summer. Citrus photosynthesis during the summer was not impaired by biochemical or photochemical reactions, as CO₂ assimilation was only limited by stomatal conductance due to high leaf-to-air vapor pressure difference (VPD) during the afternoon. During the winter, the reduction in photosynthesis during the afternoon was caused by decreases in RuBP regeneration and stomatal conductance, which are both precipitated by low night temperature.     

The Effects of N Nutrition on the Water Relations and Gas Exchange Characteristics of Wheat (Triticum aestivum L.)

The purpose of this study was to characterize leaf photosynthetic and stomatal responses of wheat (Triticum aestivum L.) plants grown under two N-nutritional regimes. High- and low-N regimes were imposed on growth-chamber-grown plants by fertilizing with nutrient solutions containing 12 or 1 millimolar nitrogen, respectively. Gas-exchange measurements indicated not only greater photosynthetic capacity of high-N plants under well-watered conditions, but also a greater sensitivity of CO₂ exchange rate and leaf conductance to CO₂ and leaf water potential compared to low-N plants. Increased sensitivity of high-N plants was associated with greater tissue elasticity, lower values of leaf osmotic pressure and greater aboveground biomass. These N-nutritional-related changes resulted in greater desiccation (lowered relative water content) of high-N plants as leaf water potential fell, and were implicated as being important in causing greater sensitivity of high-N leaf gas exchange to reductions in water potential. Water use efficiency of leaves, calculated as CO₂ exchange rate/transpiration, increased from 9.1 to 13 millimoles per mole and 7.9 to 9.1 millimoles per mole for high- and low-N plants as water became limiting. Stomatal oscillations were commonly observed in the low-N treatment at low leaf water potentials and ambient CO₂ concentrations, but disappeared as CO₂ was lowered and stomata opened.