Regulation of Transpiration to Improve Crop Water Use

Decreasing fresh water supplies and increasing agricultural drought threaten sustainable worldwide crop production. Consequently, there is a global priority to develop crops with higher water use efficiency (WUE): biomass production or yield per unit of water used. Water use efficiency varies substantially among species and genotypes within a species, and a major effort is now underway to identify the genetic determinants of WUE. Today, it is known that genotypes in primary gene pools exhibit allelic variation for WUE through mechanisms that regulate transpiration, which is the conductance of water through stomata, the cuticle, and the boundary layer. Because of the differential diffusion properties of water and carbon dioxide (CO₂) through these pathways, it is feasible that WUE could be improved by decreasing transpiration without a concomitant reduction in CO₂ uptake. Since CO₂ uptake and transpirational water loss occur predominantly through stomatal pores, it is not surprising that genes involved in stomatal development and stomatal opening/closing impact WUE. Furthermore, loss- and gain-of-function genetic screens have identified genes that regulate transpiration and WUE by yet undetermined mechanisms. This review will discuss the genetic determinants that regulate transpiration and WUE in the context of the modern agricultural goal of improving WUE while sustaining biomass and yield.     

The Arabidopsis GTL1 transcription factor regulates water use efficiency and drought tolerance by modulating stomatal density via transrepression of SDD1

A goal of modern agriculture is to improve plant drought tolerance and production per amount of water used, referred to as water use efficiency (WUE). Although stomatal density has been linked to WUE, the causal molecular mechanisms have yet to be determined. Arabidopsis thaliana GT-2 LIKE 1 (GTL1) loss-of-function mutations result in increased water deficit tolerance and higher integrated WUE by reducing daytime transpiration without a demonstrable reduction in biomass accumulation. gtl1 plants had higher instantaneous WUE that was attributable to ~25% lower transpiration and stomatal conductance but equivalent CO(2) assimilation. Lower transpiration was associated with higher STOMATAL DENSITY AND DISTRIBUTION1 (SDD1) expression and an ~25% reduction in abaxial stomatal density. GTL1 expression occurred in abaxial epidermal cells where the protein was localized to the nucleus, and its expression was downregulated by water stress. Chromatin immunoprecipitation analysis indicated that GTL1 interacts with a region of the SDD1 promoter that contains a GT3 box. An electrophoretic mobility shift assay was used to determine that the GT3 box is necessary for the interaction between GTL1 and the SDD1 promoter. These results establish that GTL1 negatively regulates WUE by modulating stomatal density via transrepression of SDD1.     

Morphological and physiological responses of two coffee progenies to soil water availability

Drought is a major environmental constraint affecting growth and production of coffee. The effects of water supply on growth, biomass allocation, water relations, and gas exchange in two coffee progenies representing drought-tolerant (Siriema) and drought-sensitive (Catucaí) genotypes were compared. They were grown in 12-L pots until 4-months old, when they were submitted to two watering treatments for 60 d: plants receiving either 100% transpired water (control plants) or a fraction (about 40%) of the amount of water transpired by control plants (drought-stressed plants). Under control conditions, Siriema grew faster than Catucaí. Regardless of the watering regimes and progenies, relative growth rate (RGR) was positively correlated both with net assimilation rate (NAR) and long-term water-use efficiency (WUE), but not with differences in biomass allocation. Both progenies responded to drought stress through (i) similar decreases in both RGR and NAR with marginal, if any, changes in allocation; (ii) decreases in leaf water potential, which occurred to a greater extent in Catucaí than in Siriema, even though they have showed similar abilities to adjust osmotically and elastically; (iii) similar reductions in net photosynthesis due mainly to nonstomatal factors; and (iv) decreases in transpiration rate coupled with increased long-term WUE. However, the lower transpiration rate and the higher long-term WUE as found in Siriema relative to Catucaí under control conditions persisted under drought conditions. Overall, the major differences between these progenies were largely associated with differences in plant water use, which was likely related to the improved water status of Siriema. The possible implications of selecting coffee genotypes for high WUE are discussed.     

Phenomics allows identification of genomic regions affecting maize stomatal conductance with conditional effects of water deficit and evaporative demand

Stomatal conductance is central for the trades‐off between hydraulics and photosynthesis. We aimed at deciphering its genetic control and that of its responses to evaporative demand and water deficit, a nearly impossible task with gas exchanges measurements. Whole‐plant stomatal conductance was estimated via inversion of the Penman–Monteith equation from data of transpiration and plant architecture collected in a phenotyping platform. We have analysed jointly 4 experiments with contrasting environmental conditions imposed to a panel of 254 maize hybrids. Estimated whole‐plant stomatal conductance closely correlated with gas‐exchange measurements and biomass accumulation rate. Sixteen robust quantitative trait loci (QTLs) were identified by genome wide association studies and co‐located with QTLs of transpiration and biomass. Light, vapour pressure deficit, or soil water potential largely accounted for the differences in allelic effects between experiments, thereby providing strong hypotheses for mechanisms of stomatal control and a way to select relevant candidate genes among the 1–19 genes harboured by QTLs. The combination of allelic effects, as affected by environmental conditions, accounted for the variability of stomatal conductance across a range of hybrids and environmental conditions. This approach may therefore contribute to genetic analysis and prediction of stomatal control in diverse environments.     

平茬措施对天然沙冬青生理特性的影响

沙冬青(Ammopiptanthus mongolicus(Maxim.)Cheng f.)是我国二级珍稀濒危植物,由于自然条件的不断恶化使这一古老残遗物种种群受到威胁。本文以西鄂尔多斯国家级自然保护区内生长的天然沙冬青为材料,对平茬后经过3年恢复生长的萌蘖灌丛和未平茬的沙冬青灌丛的地上部生物量、叶片相对含水量(RWC)、叶水势(WPB)、净光合速率(Pn)、蒸腾速率(Tr)、水分利用效率(WUE)、气孔导度(Gs)和胞间CO2浓度(Ci)等指标进行了测定,并对平茬处理的各项生理指标变化进行了研究。结果显示:平茬后的沙冬青萌蘖灌丛经过3年的生长能恢复到平茬前的生物量;平茬与未平茬(对照)沙冬青叶片相对含水量差异不显著(P0.05),但平茬后的沙冬青叶水势日间(7∶00-19∶00)均显著高于对照(P0.05);平茬与未平茬沙冬青日间净光合速率、蒸腾速率均出现两个高峰值,且均有"午休"现象,平茬后的沙冬青萌蘖丛Pn日均值比对照升高了15.73%、Tr日均值比对照升高了15.57%、WUE日均值比对照升高了13.92%(P0.05);Gs日均值比对照也有所上升,且与Pn、Tr的变化趋势基本一致,但Ci的变化趋势与Gs、Pn和Tr相反。说明平茬处理对增加地上部生物量、提高其生理指标均有明显作用,能提高沙冬青潜在的生产力。     

Growth, gas exchange, water-use efficiency, and carbon isotope composition of ‘Gale Gala’ apple trees grafted onto 9 wild Chinese rootstocks in response to drought stress

To determine the effects of rootstock choice on the scion response to drought stress, we compared the vegetative growth, biomass accumulation, gas exchange, and water-use efficiency (WUE) of ??Gale Gala?? apple (Malus domestica Borkh.) trees grafted onto nine wild Chinese Malus rootstocks. Compared with the well-watered control, drought treatment limited growth, as manifested by smaller increments in plant height (PH), trunk diameter (TD), total fresh biomass (TB), total dry biomass (TDB), total leaf area (LA), and relative growth rate (RGR). The extent of this effect differed among rootstocks. Stress conditions led to increases in the root/shoot ratio (RSR), leaf thickness (LT), water-holding capacity (WHC), carbon isotope composition (??¹³C), and WUE. Decreases were noted in stomatal density (SD), leaf relative water content (RWC), chlorophyll content (Chl), net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s), again varying by rootstock. Those that are generally considered more drought-tolerant, e.g., M. sieversii, M. prunifolia, and M. toringoides, had smaller declines in PH, TD, TB, TDB, LA, RGR, SD, RWC, Chl, P _N, E, and g _s and proportionally greater increases in RSR, LT, WHC, ??¹³C, and WUE compared with the droughtsensitive M. hupehensis and M. sieboldii. These results suggest that moisture stress has a significant dwarfing effect in the latter two species. Based on WUE calculations, trees on drought-tolerant rootstocks showed higher tolerance when stressed, whereas those on drought-sensitive rootstocks were less tolerant, as indicated by their lower WUE values.     

氮对苹果幼树水分利用效率的影响

以２年生盆栽新红星／平邑甜茶苹果树为试材,初步探讨了土壤不同水分状况下氮肥对植株水分利用效率（ＷＵＥ）及有关参数的影响,结果表明,充足供水时,随施Ｎ量的增加,植株ＷＵＥ降低,施Ｎ导致气孔导率（Ｇｓ）增大,对蒸腾（Ｔｒ）的提高幅度大于光合（Ｐｎ）;供水不足时,施Ｎ植株的ＷＵＥ明显高于对照,不同施Ｎ水平的ＷＵＥ表现为：高Ｎ〉中Ｎ〉低Ｎ,ＷＵＥ改善是由于叶肉羧化能力提高,导致光合增强。     

Genetic Control of Water Use Efficiency and Leaf Carbon Isotope Discrimination in Sunflower (Helianthus annuus L.) Subjected to Two Drought Scenarios

High water use efficiency (WUE) can be achieved by coordination of biomass accumulation and water consumption. WUE is physiologically and genetically linked to carbon isotope discrimination (CID) in leaves of plants. A population of 148 recombinant inbred lines (RILs) of sunflower derived from a cross between XRQ and PSC8 lines was studied to identify quantitative trait loci (QTL) controlling WUE and CID, and to compare QTL associated with these traits in different drought scenarios. We conducted greenhouse experiments in 2011 and 2012 by using 100 balances which provided a daily measurement of water transpired, and we determined WUE, CID, biomass and cumulative water transpired by plants. Wide phenotypic variability, significant genotypic effects, and significant negative correlations between WUE and CID were observed in both experiments. A total of nine QTL controlling WUE and eight controlling CID were identified across the two experiments. A QTL for phenotypic response controlling WUE and CID was also significantly identified. The QTL for WUE were specific to the drought scenarios, whereas the QTL for CID were independent of the drought scenarios and could be found in all the experiments. Our results showed that the stable genomic regions controlling CID were located on the linkage groups 06 and 13 (LG06 and LG13). Three QTL for CID were co-localized with the QTL for WUE, biomass and cumulative water transpired. We found that CID and WUE are highly correlated and have common genetic control. Interestingly, the genetic control of these traits showed an interaction with the environment (between the two drought scenarios and control conditions). Our results open a way for breeding higher WUE by using CID and marker-assisted approaches and therefore help to maintain the stability of sunflower crop production.     

Water and Nutrient Use Efficiency in Diploid, Tetraploid and Hexaploid Wheats

Three diploid (Triticum boeoticum, AA; Aegilops speltoides, BB and Ae. tauschii, DD), two tetraplold (T. dlcoccoides,AABB and T. dicoccon, AABB) and one hexaploid (T. vulgare, AABBDD) varieties of wheat, which are very important in the evolution of wheat were chosen in this study. A pot experiment was carried out on the wheat under different water and nutrient conditions (i) to understand the differences in biomass, yield, water use efficiency (WUE), and nutrient (N, P and K) use efficiency (uptake and utilization efficiency) among ploldles in the evolution of wheat; (ii) to clarify the effect of water and nutrient conditions on water and nutrient use efficiency; and (iii) to assess the relationship of water and nutrient use efficiency in the evolution of wheat. Our results showed that from diploid to tetraploid then to hexaploid during the evolution of wheat, both root biomass and above-ground biomass increased initially and then decreased. Water consumption for transpiration decreased remarkably, correlating with the decline of the growth period, while grain yield, harvest index, WUE, N, P and K uptake efficiency, and N, P and K utilization efficiency increased significantly. Grain yield, harvest index and WUE decreased in the same order: T.vulgare ＞ T. dicoccon ＞ T. dicoccoides ＞ Ae. tauschii ＞ Ae. speltoides ＞ T. boeoticum. Water stress significantly decreased root biomass, above-ground biomass, yield, and water consumption for transpiration by 47-52%, butremarkably increased WUE. Increasing the nutrient supply increased wheat above-ground biomass, grain yield,harvest index, water consumption for transpiration and WUE under different water levels, but reduced root biomass under drought conditions. Generally, water stress and low nutrient supply resulted in the lower nutrientuptake efficiency of wheat. However, water and nutrient application had no significant effects on nutrient utilization efficiency, suggesting that wheat nutrient utilization efficiency is mainly controlled by genotypes. Compared to theother two diploid wheats, Ae. squarrosa (DD) had significant higher WUE and nutrient utilization efficiency, Indicating that the D genome may carry genes controlling high efficient utilization of water and nutrient. Significant relationships were found between WUE and N, P and K utilization efficiency.     

A dynamic, architectural plant model simulating resource-dependent growth

BACKGROUND AND AIMS: Physiological and architectural plant models have originally been developed for different purposes and therefore have little in common, thus making combined applications difficult. There is, however, an increasing demand for crop models that simulate the genetic and resource-dependent variability of plant geometry and architecture, because man is increasingly able to transform plant production systems through combined genetic and environmental engineering. MODEL: GREENLAB is presented, a mathematical plant model that simulates interactions between plant structure and function. Dual-scale automaton is used to simulate plant organogenesis from germination to maturity on the basis of organogenetic growth cycles that have constant thermal time. Plant fresh biomass production is computed from transpiration, assuming transpiration efficiency to be constant and atmospheric demand to be the driving force, under non-limiting water supply. The fresh biomass is then distributed among expanding organs according to their relative demand. Demand for organ growth is estimated from allometric relationships (e.g. leaf surface to weight ratios) and kinetics of potential growth rate for each organ type. These are obtained through parameter optimization against empirical, morphological data sets by running the model in inverted mode. Potential growth rates are then used as estimates of relative sink strength in the model. These and other 'hidden' plant parameters are calibrated using the non-linear, least-square method. KEY RESULTS AND CONCLUSIONS: The model reproduced accurately the dynamics of plant growth, architecture and geometry of various annual and woody plants, enabling 3D visualization. It was also able to simulate the variability of leaf size on the plant and compensatory growth following pruning, as a result of internal competition for resources. The potential of the model's underlying concepts to predict the plant's phenotypic plasticity is discussed.     

On the Characteristics of Transpiration and Its Responses to Shade in Ginkgo biloba

With a CI-301PS portable photosynthesis system as a measuring device, a field study on the characteristics of transpiration and the responses of transpiration (E), stomatal resistance (R) and water use efficiency (WUE) to shade in Ginkgo biloba L. grown in Mt. Lushan was conducted. The results showed that with sufficient water in soil, the highest transpiration rate in a sunny summer day appeared in the afternoon. The WUE was maximum at about 8 a.m. and then decreased at noon remarkably. Different responses of E, R and WUE to shade were noticed which indicated that G. biloba was very suitable to the present climate. Moreover, the temperature of air, photosynthetic active radiation and R were the dominant factors affecting transpiration.     

半干旱黄土丘陵区五种植物的生理生态特征比较

通过测定陕北黄土丘陵区5种植物在旱季的光合速率、蒸腾速率和叶水势日变化,将植物划分为不同的水分生态适应类型.结果表明:柳枝稷的光合生理特征属于高光合、低蒸腾和高水分利用效率类型,其抗旱适应性特征属于高水势延迟脱水类型;苜蓿属于高光合高蒸腾低水分利用效率类型,耐旱性为高水势延迟脱水型;达乌里胡枝子属于低光合、低蒸腾、低水分利用效率类型,耐旱性为高水势延迟脱水型;白羊草属于高光合、蒸腾较高的水分利用效率中等型,耐旱性属于能忍耐脱水造成的低水势的一类.沙打旺属于高光合中等蒸腾速率中等水分利用效率类型,耐旱性为低水势延迟脱水型.     

Population differences in water-use efficiency of Eucalyptus microtheca seedlings under different watering regimes

Gas exchange, water-use efficiency (WUE), carbon isotope composition ( Δ ¹³C) and growth traits were compared among 5 populations of Eucalyptus microtheca F. Muell. Seedlings grown from seed collected across the natural distribution of the species were maintained under water-stressed and well-watered conditions. Gas exchange was measured in terms of net photosynthesis (A) and transpiration (E); WUE was measured in terms of instantaneous water-use efficiency (WUE_i) and transpiration efficiency (WUE_T); growth traits were measured in terms of total biomass (TB), root/shoot ratio (RS), and specific leaf area density (DEN). Significant differences in all traits were detected among the populations. Overall population variation was 1.68–2.50 and 1.48–2.26 μmol CO₂ uptake per mmol H₂O transpired (WUE_i), 1.97–3.04 and 1.64–2.36 g dry matter accumulation per kg water transpired (WUE_T), and Δ ¹³C was −28.81 to −26.75‰ and −30.56 to −30.04‰ under the water-stressed and well-watered conditions, respectively. In addition, WUE_i, WUE_T and Δ ¹³C were significantly correlated with A, E, RS, DEN and TB. The study indicated that measurement of WUE may be a useful trait for selecting genotypes with improved drought adaptation and biomass productivity under different environmental conditions.     

铝胁迫对蓼科植物生长和光合、蒸腾特性的影响

采用水培试验,设置5种铝处理浓度,研究了铝对3种蓼科植物酸模叶蓼、杠板归和辣蓼叶片光合、蒸腾和叶绿素荧光参数的影响。结果表明,高铝处理(400μmol.L-1)显著抑制3种蓼科植物地上部和根系生长,并且导致3种蓼科植物叶片叶绿素含量、Chla/Chlb、净光合速率(Pn)、水分利用效率(WUE)、PSII光合电子传递量子效率(φPSII)和光化学猝灭系数(qP)显著下降。中低铝处理(25~100μmol.L-1)时,与对照相比,酸模叶蓼生物量显著增加,杠板归显著减少,辣蓼先增加后减少。其中,酸模叶蓼和辣蓼叶绿素含量、Chla/Chlb、Pn、蒸腾速率(Tr)、胞间CO2浓度(Ci)、PSII最大光化学效率(Fv/Fm)、qP均未发生显著变化,但辣蓼WUE、φPSII和非光化学猝灭系数(NPQ)显著下降,酸模叶蓼无显著变化;而杠板归除Ci、Fv/Fm外,其余叶片光合、蒸腾及叶绿素荧光参数均出现显著下降。上述结果表明,酸模叶蓼在中低铝处理条件下可通过保持较高的叶绿素含量、Chla/Chlb、WUE、Pn、PSII反应中心光化学反应效率以及提高非辐射能量耗散来增强其对铝的耐性。     

硅对生姜叶片水、二氧化碳交换特性的影响

为探讨硅对生姜植株的生理效应,以莱芜大姜为试材,研究了水培营养液中不同硅素水平对生姜植株生长、硅含量及叶片光合作用和蒸腾作用的影响.结果表明：植株各器官硅含量及生物量均随营养液硅素水平的升高而显著增加,1.0 (T₁)、1.5 (T₂)、2.0 (T₃) mmol·L^-1硅(Si)处理植株叶片硅 (SiO₂) 含量分别比CK增加604.4%、834.8%和1130.4%,单株生物量分别比CK增加9.4%、19.4%和22.8%.随着硅素水平的升高,叶片Mg²⁺ TPase、Ca²⁺ ATPase活性及光合速率(P_n)和水分利用效率(WUE)提高,蒸腾速率(T_r)降低.一天中在11:00时,T₁、T₂、T₃处理的生姜叶片P_n和WUE分别比CK提高11.2%、21.8%、28.2%和23.1%、55.9%、54.8%,T_r分别比CK降低6.3%、17.1%和19.2%.此外,硅素还显著提高了生姜叶片光合作用饱和光强、CO2羧化效率及类胡萝卜素含量,但对叶绿素含量无显著影响.本试验条件下,以15～20 mmol·L-1硅素(Si)处理效果最好.     

Variance for water-use efficiency among ecotypes and recombinant inbred lines of Arabidopsis thaliana (Brassicaceae)

Knowledge of natural variation among ecotypes and recombinant inbred lines of Arabidopsis thaliana L. Heynh for season-long water-use efficiency (WUE, moles of carbon accumulated per mole of water used) is useful in the design of experiments to understand the genetic control of this phenomenon. Water-use efficiency among 31 container-grown Arabidopsis ecotypes ranged from a high of 2.40 to a low of 1.86 mg/g (grams of dry aerial biomass per gram of water used). Genetic variance for WUE was observed among 80 F₅ recombinant inbred lines (RILs) derived from a cross between the highest (Lip-0) by the lowest (Edi-0) ecotype. The heritability of WUE was calculated as 0.18 ± 0.07. The mean WUE for the ten highest and ten lowest RILs was 2.42 and 1.05 mg/g, respectively. A significant difference was observed in the composition of stable isotopes of plant carbon (δ¹³C) between the mean of the ten highest (-31.23±) and the mean of the ten lowest (-31.96±) RILs based on WUE. Characterization of the 80 RILs provides a structured population for further genetic and physiologic study of WUE.     

Phenotypic selection on leaf water use efficiency and related ecophysiological traits for natural populations of desert sunflowers

Plant water-use efficiency (WUE) is expected to affect plant fitness and thus be under natural selection in arid habitats. Although many natural population studies have assessed plant WUE, only a few related WUE to fitness. The further determination of whether selection on WUE is direct or indirect through functionally related traits has yielded no consistent results. For natural populations of two desert annual sunflowers, Helianthus anomalus and H. deserticola, we used phenotypic selection analysis with vegetative biomass as the proxy for fitness to test (1) whether there was direct and indirect selection on WUE (carbon isotope ratio) and related traits (leaf N, area, succulence) and (2) whether direct selection was consistent with hypothesized drought/dehydration escape and avoidance strategies. There was direct selection for lower WUE in mesic and dry H. anomalus populations, consistent with dehydration escape, even though it is the longer lived of the two species. For mesic H. anomalus, direct selection favored lower WUE and higher N, suggesting that plants may be “wasting water” to increase N delivery via the transpiration stream. For the shorter lived H. deserticola in the direr habitat, there was indirect selection for lower WUE, inconsistent with drought escape. There was also direct selection for higher leaf N, succulence and leaf size. There was no direct selection for higher WUE consistent with dehydration avoidance in either species. Thus, in these natural populations of two desert dune species higher fitness was associated with some combination direct and indirect selection for lower WUE, higher leaf N and larger leaf size. Our understanding of the adaptive value of plant ecophysiological traits will benefit from further consideration of related traits such as leaf nitrogen and more tests in natural populations.     

巨桉与5种木本植物幼树的耗水特性及水分利用效率的比较

巨桉因生长迅速且经济效益高,在我国南方被广泛用于营造短周期工业原料林,但其蒸腾耗水状况与其他常见或乡土树种存在怎样的差异,大面积种植是否会改变栽培区原有的水分平衡,是一个尚未明确的问题。利用LI-6400光合作用仪测定了巨桉与其他5种木本植物在不同光强、温度、湿度下的净光合速率(Pn)、蒸腾速率(Tr)和水分利用效率(WUE),用称重法测定了参试植物载叶量、生物量和耗水量,并对这些树种的蒸腾耗水特性进行了评价。结果表明:(1)相同环境条件下,巨桉的Tr最大,WUE最低,单位质量耗水量最多,单株蒸腾耗水量远高于其他树种,其明显较大的叶面积可能是重要原因之一,虽然其Pn仅次于杨树,生物量积累最大,因此为高光合、高蒸腾、低水分利用效率植物。(2)阔叶树种的Pn和Tr明显高于针叶树种,而WUE低于针叶树种。(3)环境因子(光照强度、温度和湿度)对植物Pn、Tr和WUE的影响较大,其中Pn主要受光照强度影响,Tr对湿度最为敏感,一般情况下WUE随湿度的增大而升高。在试验设置的温度范围(24—32℃)内,光合作用变化幅度不大。光照强度800μmol.m-.2s-1和温度28℃最有利于参试植物的光合作用。(4)巨桉等速生树种较强的光合作用使其生长迅速,固碳潜力大,但其高蒸腾和低水分利用效率的特点意味着在栽培区替代原有植被进行大规模造林时,可能会消耗更多的水资源而对生态环境造成一定的不利影响,因此在发展巨桉人工林时应选择水分条件好的区域(尤其是年降雨量充沛且季节间分配相对均匀的地区),并进行科学的经营管理。     

Water use efficiency as a function of leaf age and position within the cotton canopy

The gas exchange properties of whole plant canopies are an integral part of crop productivity and have attracted much attention in recent years. However, insufficient information exists on the coordination of transpiration and CO₂ uptake for individual leaves during the growing season. Single-leaf determinations of net photosynthesis (Pn), transpiration (E) and water use efficiency (WUE) for field-grown cotton (Gossypium hirsutum L.) leaves were recorded during a 2-year field study. Measurements were made at 3 to 4 day intervals on the main-stem and first three sympodial leaves at main-stem node 10 from their unfolding through senescence. Results indicated that all gas exchange parameters changed with individual main-stem and sympodial leaf age. Values of Pn, E and WUE followed a rise and fall pattern with maximum rates achieved at a leaf age of 18 to 20 days. While no significant position effects were observed for Pn, main-stem and sympodial leaves did differ in E and WUE particularly as leaves aged beyond 40 days. For a given leaf age, the main-stem leaf had a significantly lower WUE than the three sympodial leaves. WUE's for the main-stem and three sympodial leaves between the ages of 41 to 50 days were 0.85, 1.30, 1.36 and 1.95 μmol CO₂ mmol⁻¹ H₂O, respectively. The mechanisms which mediated leaf positional differences for WUE were not strictly related to changes in stomatal conductance (g_s·H₂O) since decreases in g_s·H₂O with leaf age were similar for the four leaves. However, significantly different radiant environments with distance along the fruiting branch did indicate the possible involvement of mutual leaf shading in determining WUE. The significance of these findings are presented in relation to light competition within the plant canopy during development.