Season effects on leaf nitrogen partitioning and photosynthetic water use efficiency in mango

The key parameters of photosynthetic capacity (maximum carboxylation rate (V_cmax), electron transport capacity (J_max) and dark respiration rate (R_d)) and the slope (m) of the stomatal conductance model of Ball et al. [Progress in photosynthetic research, Martinus Nijhoff, Dordrecht, 1987] were measured for a whole growing season in fully expanded leaves of 12-year-old mango trees cv. Cogshall in La Réunion island. Leaf nitrogen partitioning into carboxylation (P_c) and bioenergetic (P_b) pools were computed according to the model of Niinemets and Tenhunen [Plant Cell Environ 1997;20: 845–66]. V_cmax, J_max, R_d, P_c and P_b remained relatively stable over the whole study period, with the exception of the period of linear fruit growth when J_max, R_d and P_b were slightly lower, and leaf non-structural carbohydrate content higher. During the pre-floral and floral periods, m decreased by more than 50%, indicating an increase in photosynthetic water use efficiency and m increased again during the period of linear fruit growth. Our results show that, in tropical orchard conditions characterized by mild seasonal climatic changes and non-limiting water supply, leaf nitrogen partitioning is rather stable. Our results also advocate for more studies on the effect of phenology on m and photosynthetic water use efficiency, which is of paramount importance for building coupled biochemical models of photosynthetic carbon assimilation.     

不同沟灌模式对沙漠绿洲区葡萄生长和水分利用的效应

在甘肃河西荒漠绿洲区研究了覆膜与不覆膜条件下隔沟交替灌溉、常规沟灌对葡萄生长和水分利用的影响.结果表明,隔沟交替灌溉可以保证作物一部分根区处于比较湿润状态,另一部分根区处于相对干燥状态,湿润与干燥区域的交替出现可诱导葡萄的补偿生长效应.隔沟交替灌溉条件下葡萄叶片气孔开度减小,光合速率略有降低或下降不显著,而蒸腾速率明显下降,水分利用效率增大.光合作用日变化也表现出类似规律.隔沟交替灌溉与地膜覆盖技术相结合能显著提高水分利用效率,为在田间实施气孔最优化调控提供了一种有效途径.     

Photosynthetic characteristics and water use efficiency of three coastal dune plants

To clarify the ecophysiological characteristics of plants growing on a coastal dune, net photosynthetic rate (Pn) and leaf conductance (g1) of three perennial species in Japanese coastal regions,Ischaemum anthephoroides (C₄),Carex kobomugi (C₃) andCalystegia soldanella (C₃), were compared under controlled environmental conditions and field conditions at the kado-ori coast at Ohno Village, Ibaraki.I. anthephoroides achieved photosynthetic CO₂ saturation at ca. 100 μll⁻¹ intercellular CO₂ concentration (C ₁), and itsPn was not light-saturated at a high photosynthetically active photon flux density (PPFD) of 1000 μmol m⁻²s⁻¹. This C₄ species showed a high optimal leaf temperature forPn (35°C) and a lowg1 (0.1 mol H₂O m⁻²s⁻¹), permitting maintenance of the highest water use efficiency (WUE, the ratio ofPn to transpiration rate (Tr)) in the field among the three species. At light saturation,C. soldanella had the lowestPn andg1 and a similar carboxylation efficiency related toC ₁ (Pn/C₁ ratio) asCarex kobomugi. Calystegia soldanella also had a high optimal temperature forPn (30°C), and achieved higherWUE thanCarex kobomugi, as a result of efficient stomatal regulation. In contrast, at the optimum temperatureC. kobomugi had a highPn comparable toI. anthephoroides due to a highg1 of 0.3 mol H₂O m⁻²s⁻¹, but with high temperature treatmentsPn andg1 were significantly decreased.C. kobomugi always had the lowestWUE among the three species in the field. It is therefore clear thatI. anthephoroides is a heat-resistant species,Calystegia soldanella is a heat-enduring species andCarex kobomugi is a heat-evading species due to its eraly phenology.     

Stomatal and photosynthetic responses to shade in sorghum, soybean and eastern gamagrass

We studied photosynthetic and stomatal responses of grain sorghum ( Sorghum bicolor [L.] Moench cv. Pioneer 8500), soybean ( Glycine max L. cv. Flyer) and eastern gamagrass ( Tripsacum dactyloides L.) during experimental sun and shade periods simulating summer cloud cover. Leaf gas exchange measurements of field plants showed that short-term (5 min) shading of leaves to 300–400 μmol m⁻² s⁻¹ photosynthetic photon flux density reduced photosynthesis, leaf temperature, stomatal conductance, transpiration and water use efficiency and increased intercellular CO₂ partial pressure. In all species, photosynthetic recovery was delayed when leaves were reilluminated, apparently by stomatal closure. The strongest stomatal response was in soybean. Photosynthetic recovery was studied further with soybeans grown indoors (maximum photosynthetic photon flux density 1 200 μmol m⁻² s⁻¹). Plants grown indoors had responses to shade similar to those of field plants, except for brief nonstomatal limitation immediately after reillumination. These responses indicated the importance of the light environment during leaf development on assimilation responses to variable light, and suggested different limitations on carbon assimilation in different parts of the soybean canopy. Photosynthetic oxygen evolution recovered immediately upon reillumination, indicating that the light reactions did not limit soybean photosynthetic recovery. While shade periods caused stomatal closure and reduced carbon gain and water loss in all species, the consequences for carbon gain/water loss were greatest in soybean. The occurrence of stomatal closure in all three species may arise from their shared phenologies and herbaceous growth forms.     

田间不同条件下玉米叶片的气孔阻力及与光合、蒸腾作用的关系

以玉米17个自交系和10个杂交种为试材,在田间条件下研究了不同光强、不同叶位、不同生育期、不同源库比例和株间差异状态下的气孔阻力及其与光合(PH)、蒸腾(TR)和叶片水分利用效率(WUE)的关系,结果表明,不同自交系之间RS具有显著的差异,相差最大可达2.3倍以上;光照条件变弱、穗叶位差增加、生育期推延和源/库比例改变等均可引起RS值的增加;在各种情况下导致RS变化的同时也引起TR和PH产生相应的变化,RS与TR、PH表现出显著或极显著的负相关,且相关系数r_RS-TR>r_RS-PH;RS与WUE的相关关系在不同的条件下表现出不稳定性.     

A relationship between carbon dioxide, photosynthetic efficiency and shade tolerance

Net photosynthesis and transpiration of seedlings from shade tolerant, moderately tolerant and intolerant tree species were measured in ambient carbon dioxide (CO₂) concentrations ranging from 312 to 734 ppm. The species used, Fagus grandifolia Ehrh. (tolerant), Quercus alba L., Q. rubra L., Liriodendron tulipifera L. (moderately tolerant), Liquidambar styraciflua L. and Pinus taeda L. (intolerant), are found co-occurring in the mixed pine-hardwood forests of the Piedmont region of the southeastern United States. When seedlings were grown in shaded conditions, photosynthetic CO₂ efficiency was significantly different in all species with the highest efficiency in the most shade tolerant species, Fagus grandifolia , and progressively lower efficiencies in moderately tolerant and intolerant species. Photosynthetic CO₂ efficiency was defined as the rate of increase in net photosynthesis with increase in ambient CO₂ concentration. When plants which had grown in a high light environment were tested, the moderately tolerant and intolerant deciduous species had the highest photosynthetic CO₂ efficiencies but this capacity was reduced when these species grew in low light. The lowest CO₂ efficiency and apparent quantum yield occurred in Pinus taeda in all cases. Water use efficiency was higher for all species in enriched CO₂ environments but transpiration rate and leaf conductance were not affected by CO₂ concentration. High photosynthetic CO₂ efficiency may be advantageous for maintaining a positive carbon balance in the low light environment under a forest canopy.     

Effects of salinity on photosynthesis,leaf anatomy,ion accumulation and photosynthetic nitrogen use efficiency in five Indian mangroves

Five species of mangroves (Bruguiera gymnorrhiza, Excoecaria agallocha, Heritiera fomes, Phoenix paludosa and Xylocarpus granatum) were investigated with respect to their photosynthesis rate, chlorophyll content, mesophyll conductance, specific leaf area, stomatal conductance and photosynthetic nitrogen use efficiency under saline (15–27 PPT) and non-saline (1.8–2 PPT) conditions. Some inorganic elements were estimated from the leaf samples to compare the concentrations with change in salinity. Elevated assimilation rate coupled with increased chlorophyll content, more mesophyll and stomatal conductance and higher specific leaf area in non-saline condition indicates that these mangroves can grow well even with minimal salinity in soil. In B. gymnorrhiza, E. agallocha and P. paludosa the optimum PAR acquisition for photosynthesis was higher under salt stress, while the maximal rate of assimilation was lower even with minimal salinity. H. fomes and X. granatum followed the opposite trend, where the peak photosynthesis rate was lower under non-saline conditions even at a higher irradiance than in the saline forest. This indicates less affinity of H. fomes and X. granatum to high substrate salinity. Accumulation of Na⁺ increased in plants in saline substrate, while in most of the species, salinity imposed reduction in Ca⁺ and Mg⁺ uptake. Increased K⁺ content can be attributed to high substrate level K⁺ in non-saline soil. Trace amount of salinity induced Cu⁺⁺ detected in leaves of H. fomes may impart some toxic effects. Photosynthetic nitrogen use efficiency increased in non-saline soil that can be attributed to higher photosynthetic peak in most of the species and/or lower nitrogen accumulation in plant samples.     

Responses of Photosynthetic Rate and Stomatal Conductance to Water Stress in Soybean Leaves

Responses of photosynthetic rate and stomatal conductance to water stress as weI1 as the relationship between photosynthetic rate and stomatal conductance were investigated with soybean cultivars “Ludou No. 4” and “7605”. The former was a high yield cultivars widely used in Shandong province, and the latter was a small grain soybean line bred by Shandong Academy of Agricultural science. Soil water stress decreased leaf apparent photosynthetic rate and stomatal conductance of two soybean cultivars, and “Ludou No. 4” decreased more than “7605”. At the same value of water potential, photosynthetic rate and stomatal conductance of “7605” were higher than those of “Ludou No,4”,but the rate of stomatal closure for “7605” was higher than “Ludou No. 4”. Decreasing of stomatal conductance caused rising of leaf temperature of two soybean cultivars, and the rising of “7605” was more rapid than “Ludou No. 4”, but at the same treatment of water stress, leaf temperature of “Ludou No. 4” was higher than “7605”. Leaf water use efficiecy (WUE) of two soybean cultivars were decreased under water stress, and the rate of decreasing in “Ludou No.4” was more rapid than in “7605”. These results showed that “7605” was more resistant to water:stress than “Ludou No. 4”.     

Water use efficiency and carbon isotope composition of plants in a cold desert environment

Summary The effects of the availabilities of water and nitrogen on water use efficiency (WUE) of plants were investigated in a sagebrush steppe. The four species studied wereArtemisia tridentata (shrub),Ceratoides lanata (suffrutescent shrub),Elymus lanceolatus (rhizomatous grass), andElymus elymoides (tussock grass). Water and nitrogen levels were manipulated in a two-by-two factorial design resulting in four treatments: control (no additions), added water, added nitrogen, and added water and nitrogen. One instantaneous and two long-term indicators of WUE were used to testa priori predictions of the ranking of WUE among treatments. The short-term indicator was the instantaneous ratio of assimilation to transpiration (A/E). The long-term measures were 1) the slope of the relationship between conductance to water vapor and maximum assimilation and 2) the carbon isotope composition (¹³C) of plant material. Additional water decreased WUE, whereas additional nitrogen increased WUE. For both A/E and ¹³C, the mean for added nitrogen alone was significantly greater than the mean for added water alone, and means for the control and added water and nitrogen fell in between. This ranking of WUE supported the hypothesis that both water and nitrogen limit plant gas exchange in this semiarid environment. The short- and long-term indicators were in agreement, providing evidence in support of theoretical models concerning the water cost of carbon assimilation.     

田间不同条件下玉米叶片的气孔阻力及与光合,蒸腾作用的关系

以玉米 1 7个自交系和 1 0个杂交种为试材 ,在田间条件下研究了不同光强、不同叶位、不同生育期、不同源库比例和株间差异状态下的气孔阻力及其与光合 (PH)、蒸腾(TR)和叶片水分利用效率 (WUE)的关系 ,结果表明 ,不同自交系之间RS具有显著的差异 ,相差最大可达 2 .3倍以上 ;光照条件变弱、穗叶位差增加、生育期推延和源 /库比例改变等均可引起RS值的增加 ;在各种情况下导致RS变化的同时也引起TR和PH产生相应的变化 ,RS与TR、PH表现出显著或极显著的负相关 ,且相关系数rRS TR>rRS PH;RS与WUE的相关关系在不同的条件下表现出不稳定性     

The effect of atmospheric humidity on photosynthesis,transpiration and water use efficiency of leaves of several plant species

The effect of humidity on the gas exchange of leaves of the dicotyledons soybean (Glycine max (L.) Merrill), sunflower (Helianthus annuus L.), jojoba (Simmondsia chinensis (L.) Schneider), and saltbush (Atriplex halimus L.) and the monocotyledons wheat (Triticum aestivum L.), barley (Hordeum vulgare L.) sorghum (Sorghum bicolor (L.) Moench) and barnyard grass (Echinochloa crus-galli (L.) Beauv.) was examined under conditions of adequate soil moisture in a controlled environment. Photosynthesis and stomatal and internal diffusion resistances of whole, attached, single leaves were not affected by changes in humidity as the vapour pressure deficit between the leaf and atmosphere ranged from 8 to 27 mb. Transpiration increased linearly with increasing vapour pressure deficit. Whole plants of barley exhibited a different response. As humidity was increased, photosynthesis increased, transpiration expressed per unit of vapour pressure difference increased, and diffusion resistances became smaller. Reasons for the different behaviour of single leaves and whole plants are suggested. An index for water use efficiency, expressed per millibar of vapour pressure deficit, was calculated for single leaves of each species used in the experiments. This showed that water use efficiency was highest in the C₄ xerophytes and lowest in the C₃ mesophytes. The effect of environment on water use efficiency is examined using data from the literature.     

Water relations and stomatal characteristics of Mediterranean plants with different growth forms and leaf habits: responses to water stress and recovery

The aim of this study was to extent the range of knowledge about water relations and stomatal responses to water stress to ten Mediterranean plants with different growth forms and leaf habits. Plants were subjected to different levels of water stress and a treatment of recovery. Stomatal attributes (stomatal density, StoD), stomatal conductance (g _s), stomatal responsiveness to water stress (SR), leaf water relations (pre-dawn and midday leaf water potential and relative water content), soil to leaf apparent hydraulic conductance (K _L) and bulk modulus of elasticity (ε) were determined. The observed wide range of water relations and stomatal characteristics was found to be partially depended on the growth form. Maximum g _s was related to StoD and the stomatal area index (SAI), while g _s evolution after water stress and recovery was highly correlated with K _L. Relationships between SR to water deficit and other morphological leaf traits, such as StoD, LMA or ε, provided no general correlations when including all species. It is concluded that a high variability is present among Mediterranean plants reflecting a continuum of leaf water relations and stomatal behaviour in response to water stress.     

小麦进化材料水分利用效率与氮利用效率间相互关系

利用田间微区试验,在整株水平上研究了10种小麦进化材料水分利用效率(WUE)和氮利用效率(NUE)间的关系．结果表明,小麦在从二倍体→六倍体的长期进化过程中,WUE和NUE均逐渐增加．除法国黑麦外,其余9种小麦进化材料WUE和NUE间呈显著正相关,表明法国黑麦高的NUE可能与WUE以外的其它生理机制有关．     

Ozone uptake in the sun and shade crown of spruce: quantifying the physiological effects of ozone exposure

Summary The uptake of air pollutants depends both on pollutant concentration and on stomatal conductance. This paper deals with the uptake of ozone (O₃) from the air into the needles of Norway spruce [Picea abies (L.) Karst.] under ambient climatic conditions. Regulation of O₃ uptake by the stomata is shown and also the difference between the physiologically active O₃ concentration and the O₃ concentration of the ambient air. Data from the sun and shade crown of spruce trees at 1000 m a.s.l. are presented. Analysis of data from three vegetation periods has shown that at low ambient O₃ concentrations the O₃ uptake is largely regulated by stomatal conductance. Water vapour pressure deficit (VPD) of the atmosphere is the climatic factor which showed the highest positive correlation with O₃ concentration. However, a high leaf-air VDP led to stomatal closure, thus reducing the O₃ uptake in the needles despite high O₃ concentrations in the ambient air. The potential O₃ stress caused by high O₃ concentrations can be strongly mitigated by this natural closing of the stomata and the simultaneous occurrence of moderate drought stress.     

Gas exchange and water‐use efficiency in plant canopies

In this review, I first address the basics of gas exchange, water‐use efficiency and carbon isotope discrimination in C₃ plant canopies. I then present a case study of water‐use efficiency in northern Australian tree species. In general, C₃ plants face a trade‐off whereby increasing stomatal conductance for a given set of conditions will result in a higher CO₂ assimilation rate, but a lower photosynthetic water‐use efficiency. A common garden experiment suggested that tree species which are able to establish and grow in drier parts of northern Australia have a capacity to use water rapidly when it is available through high stomatal conductance, but that they do so at the expense of low water‐use efficiency. This may explain why community‐level carbon isotope discrimination does not decrease as steeply with decreasing rainfall on the North Australian Tropical Transect as has been observed on some other precipitation gradients. Next, I discuss changes in water‐use efficiency that take place during leaf expansion in C₃ plant leaves. Leaf phenology has recently been recognised as a significant driver of canopy gas exchange in evergreen forest canopies, and leaf expansion involves changes in both photosynthetic capacity and water‐use efficiency. Following this, I discuss the role of woody tissue respiration in canopy gas exchange and how photosynthetic refixation of respired CO₂ can increase whole‐plant water‐use efficiency. Finally, I discuss the role of water‐use efficiency in driving terrestrial plant responses to global change, especially the rising concentration of atmospheric CO₂. In coming decades, increases in plant water‐use efficiency caused by rising CO₂ are likely to partially mitigate impacts on plants of drought stress caused by global warming.     

作物群体CO2通量和水分利用效率的快速测定

In this paper,Eddy Correlation (EC) method was employed to measure the latent heat and CO₂ flux density and to calculate Water Use Efficiency (WUE) of winter wheat community in Yucheng district,Shandong Province in 1997.The results showed that the CO₂ flux density had an obvious diurnal change,with a maximum about1.5 mg·s^-1·m^-2,which appeared at about 9:00～10:00 am in general.The WUE of wheat community presented a fall trend from morning to afternoon,and the CO₂ flux density and WUE also had an obvious seasonal change,being lower in the early and late growth stages,and higher in the middle growth stage.The ranges of daily mean CO₂ flux density and WUE were 0.2～0.9 mg·s^-1·m^-2 and 5～20 gCO₂·kg^-1 H₂O,respectively.     

Changes in Stomatal Conductance,Transpiration and Water Use Efficiency of Ten Species Experienced in High CO2 Concentrations in Biosphere 2

The stomotal conductance, transpiration and water use efficiency (WUE) were measured using a LI-6400 portable photosynthesis system for 5 tropical rain forest species and 5 desert species in Biosphere 2, USA. All the species have experienced in very high CO2 ( > 2 200 μmol• mol- 1 ) for more than 4.5 years. The results showed that the stomatal conductance and transpiration of rain forest species decreased from ( 127.4 ± 65.6) and (2.04 ± 0.61 ) mmol• m- 2•s- 1 to (61.3 + 30.5) and ( 1.54 ± 0.65 ) mmol• m-2• s -1 respectively, while WUE increased from (2.90 ± 0.55) to (8.45 ± 2.71) μmol CO2 •mmo1-1 H2O, with CO2 increasing from 350 – 400 to 700 – 820 μmol• mol-l. For the desert species, stomatal conductance and transpiration decreased from respectively (142.8±94.6) and (2.09±0.71) mmol•m-2•s-1 to (57.7±35.8) and (1.36±0.52) mmolm-2•s-l, but WUE increased from (4.69 ± 1.39) to (9.68 ± 1.61) μmol CO2•mmo1-1 H2O, with the CO2 increase from 320 - 400 to 820 – 850 μtmol• mol- 1. The stomatal conductance, transpiration and WUE were less influenced by light intensity under high CO2 than low CO2 concentrations. Most rain forest species reached their light saturation points at light intensity of 500 μmol• m-2•s-1, while desert species at 1 000 μmol•m-2•s-1. Among different species, the desert C3 tree, Nicotiana glauca Grah., had the highest decrease in stomatal conductance and transpiration and the highest increase in WUE, by 78%, 69% and 310% respectively. The enhancement of increasing CO2 to the stomatal, transpiration and WUE of species with different photosynthesis pathway and life forms in Biosphere 2 could be concluded as: C3 species > C4 species, and desert C3 species > rain forest C3 species.     

Seasonal patterns of gas exchange,water relations and growth of it Roupala montana,an evergreen savanna species

Roupala montana is an evergreen species widespread in the seasonal savannas of the central plains of Brazil. I examined the degree of coupling of photosynthetic gas-exchange characteristics, water relations and growth responses of R. montana with regard to seasonal changes in soil water availability. Despite a rainless period of over three months soil water potential at 60 cm depth reached values of only about -1.0 MPa, while pre-dawn leaf water potential (^l) reached about -0.4 MPa by the end of the three-month drought. Thus, R. montana had access to deep soil water in the dry period, but pre-dawn ^l did not reach the high wet season values of -0.2 MPa. Most of the shoot growth was concluded in the onset of the rainy season. Although some individual branches might have shown some extension thereafter, most of them remained inactive during the rest of the rainy season and the subsequent dry season. New leaf production was also restricted to the first part of the wet period. R. montana remained evergreen in the dry season, but there was a 27% decrease in the number of leaves and herbivory removed about 16% of the leaf area still present in the plant. CO₂-exchange rates of these leaves reached only ca. 55% of the maximum rainy season values of 14 µmol m^-2 s^-1. Thus, the estimated potential daily carbon gain was about 34% of the maximum by the end of the dry period. These values will be even lower, if we considered the decrease in photosynthetic rates that occurred around midday. These reductions in photosynthetic rates as a result of partial stomatal closure were measured both in the wet and dry season and they were related to increases in the evaporative demand of the atmosphere. In conclusion, the combined effect of herbivory, leaf loss and reductions in photosynthetic rates limited plant productivity in the dry season.     

作物光合、蒸腾与水分高效利用的试验研究

通过田间试验,对作物光合、蒸腾、气孔行为及其影响因素进行了研究。结果表明,光合与蒸腾的非线性关系可以用抛物线方程表述,其中光合速率最高时的蒸腾速率为临界值,超出该值即为奢侈蒸腾,干旱处理的临界值较低,通过合适的调控措施,抑制奢侈蒸腾并不影响光合生产,综合分析光合速率、蒸腾速率与气孔导度的关系,气孑L导度大于0．12mol·m^-2·s^-1,实施提高气孔阻力并抑制蒸腾的措施,既节约水分又促进光合作用,增加产量．光合速率基本上随光合有效辐射的增加而提高,并有光饱和点存在,水分条件影响叶片光合作用达到饱和的早晚,干旱处理的光饱和点远远低于湿润处理,强光需要水分充足相耦合,才能充分发挥光能利用率,蒸腾与辐射的线性关系十分显著。从光合有效辐射入手,在光合有效辐射大于1000μmol·m^-2·s^-1时实施措施,既可大大降低蒸腾,又可改善光合,节水增产效果不言而喻。     

Interrelationships between water use and growth traits in biomass-producing willows

Abstract Water use, drought response and growth were examined under controlled conditions in four interbreeding willow species from different geographical origins (two clones of Salix viminalis L., one clone of S. viminalis × S. schwerenii E. Wolf and one clone of S. purpurea L.). The levels of soil water depletion that plants could sustain without wilting varied markedly between the clones. The level of drought resistance expressed this way was positively related to resistance to xylem cavitation, negatively related to the maximum stomatal conductance, and positively related to early stomatal closure. The rate of stomatal closure, however, was negatively related to the resistance to xylem cavitation. Prior to drought, there were no significant differences between leaf-specific hydraulic conductances of the clones when whole plants were considered. However, there were differences if the roots and shoots were considered separately. Drought resistance was negatively related to maximum growth yields. This is probably because resources were diverted away from leaf production to the production of denser wood (wood density was positively related to cavitation resistance), and, for one clone, to the growth of a larger root system. In addition, because the level of drought resistance was negatively related to the maximum stomatal conductance, growth may have been adversely affected as a result of reduced photosynthesis. Given its high water extraction ability, one of the clones started to wilt sooner than expected, although only lateral shoots were affected. This appeared to indicate a strategy of sacrificing expendable shoots.