Effects of N and water supply on water use-efficiency of a semiarid grassland in Inner Mongolia

Productivity of semiarid grasslands is primarily limited by seasonal rainfall amount and becomes increasingly limited by nutrient availability under wet conditions. Interactive effects of water and N availability on grassland productivity and parameters related to water use were studied on a grassland site in Inner Mongolia, China, in a 2-factorial experiment with two levels of water (rainfed: 158 mm; irrigated: 839 (N0) and 972 (N1) mm) and N supply (0 or 180 kg N ha^?1). RUE was calculated from ANPP and cumulative water supply. Bare soil evaporation (E) was calculated from climatic data and leaf area dynamics, and percolation (D) and transpiration (T) were estimated with HYDRUS-1D. Water-use efficiency (WUE, ANPP / (T + D)) and transpiration efficiency (TE, ANPP / T) were calculated. Resource availability had pronounced effects on the water-use efficiency of semiarid grassland. RUE, WUE, and TE all decreased under irrigated compared to rainfed conditions and were significantly increased with N fertilizer application at both levels of water supply. While the irrigation effect on parameters of water-use efficiency were accordingly reflected in stable carbon isotope signatures, N supply resulted in significantly less negative δ¹³C-values under rainfed but not irrigated conditions. It is concluded, that spatial or temporal gradients in resource availability have pronounced effects on the water-use efficiency of semiarid grassland. The decrease of water use-efficiency under high water supply was related to differences in TE and not to a relative increase of unproductive water loss. Carbon isotope discrimination was highly correlated with WUE and TE, but can be a poor predictor of RUE.     

Water-use efficiency and transpiration efficiency of wheat under rain-fed conditions and supplemental irrigation in a Mediterranean-type environment

Growth and water use were measured in wheat (Triticum aestivum L.) grown in northern Syria in a typical Mediterranean climate over five seasons 1991/92–1995/96. Water use was partitioned into transpiration (T) and soil evaporation (E_s) using Ritchie's model, and water-use efficiency (WUE) and transpiration efficiency (TE) were calculated. The aim of the study was to examine the influence of irrigation and nitrogen on water use, WUE and TE. By addition of 100 kg N ha^-1, E_s was reduced from 120 mm to 101 mm under rain-fed conditions and from 143 mm to 110 mm under irrigated conditions, and T was increased from 153 mm to 193 mm under rain-fed conditions and from 215 mm to 310 mm under irrigated conditions. Under rain-fed conditions, about 35% of evapotranspiration (ET) may be lost from the soil surface for the fertilized crops and 44% of ET for the unfertilized crops. Transpiration accounted for 65% of ET for the fertilized crops and 56% for the unfertilized crops under rain-fed. As a result of this, WUE was increased by 44% for dry matter and 29% for grain yield under rain-fed conditions, and by 60% for dry matter and 57% for grain yield under irrigated conditions. Transpiration efficiency for the fertilized crops was 43.8 kg ha^-1 mm^-1 for dry matter and 15 kg ha^-1 mm^-1 for grain yield, while TE for the unfertilized crops was 33.6 kg ha^-1 mm^-1 and 12.2 kg ha^-1 mm^-1 for dry matter and grain yield, respectively. Supplemental irrigation significantly increased post-anthesis water use, transpiration, dry matter and grain yield. Water-use efficiency for grain yield was increased from 9.7 to 11.0 kg ha^-1 mm^-1 by supplemental irrigation, although WUE for dry matter was not affected by it. Irrigation did not affect transpiration efficiency for grain yield, but decreased transpiration efficiency for dry matter by 16%. This was associated with higher harvest index as a result of good water supply in the post-anthesis period and increased transpiration under irrigated conditions.     

Identification of causal relationships among traits related to drought resistance in Stylosanthes scabra using QTL analysis

Previous studies have shown that a negative relationship exists between transpiration efficiency (TE) and carbon isotope discrimination (Delta) and between TE and specific leaf area (SLA) in Stylosanthes scabra. A glasshouse experiment was conducted to confirm these relationships in an F(2) population and to study the causal nature of these relationships through quantitative trait loci (QTL) analysis. One hundred and twenty F(2) genotypes from a cross between two genotypes within S. scabra were used. Three replications for each genotype were maintained through vegetative propagation. Water stress was imposed by maintaining plants at 40% of field capacity for about 45 d. To facilitate QTL analysis, a genetic linkage map consisting of 151 RAPD markers was developed. Results from this study show that Delta was significantly and negatively correlated with TE and biomass production. Similarly, SLA showed significant negative correlation with TE and biomass production. Most of the QTL for TE and Delta were present on linkage groups 5 and 11. Similarly, QTL for SLA, transpiration and biomass productivity traits were clustered on linkage groups 13 and 24. One unlinked marker was also associated with these traits. There were several markers coincident between different traits. At all the coincident QTL, the direction of QTL effects was consistent with phenotypic data. At the coincident markers between TE and Delta, high alleles of TE were associated with low alleles of Delta. Similarly, low alleles of SLA were associated with high alleles of biomass productivity traits and transpiration. At the coincident markers between trans-4-hydroxy-N:-methyl proline (MHP) and relative water content (RWC), low alleles of MHP were associated with high alleles of RWC. This study suggests the causal nature of the relationship between TE and Delta. Phenotypic data and QTL data show that SLA was more closely associated with biomass production than with TE. This study also shows that a cause-effect relationship may exist between SLA and biomass production.     

Efficiency of physiological trait-based and empirical selection approaches for drought tolerance in groundnut

Drought is the major abiotic constraint affecting groundnut productivity and quality worldwide. Most breeding programmes in groundnut follow an empirical approach to drought resistance breeding, largely based on kernel yield and traits of local adaptation, resulting in slow progress. Recent advances in the use of easily measurable surrogates for complex physiological traits associated with drought tolerance encouraged breeders to integrate these in their selection schemes. However, there has been no direct comparison of the relative efficiency of a physiological trait‐based selection approach (Tr) vis‐à‐vis an empirical approach (E) to ascertain the benefits of the former. The genetic material used in the present study originated from three common crosses and one institute‐specific cross from four collaborating institutes in India (total seven crosses). Each institute contributed six genotypes and each followed both the Tr and E selection approaches in each cross. The field trial of all selections, consisting of 192 genotypes (96 each Tr and E selections), was grown in 2000/2001 in a 4 × 48 alpha design in 12 season × location environments in India. The selection efficiency of Tr relative to E, RE_Tr, was estimated using the genetic concept of response to selection. Based on all the 12 environments, the two selection methods performed more or less similarly (RE_Tr= 1.045). When the 12 environments were grouped into rainy season and post‐rainy season, the relative response to selection in Tr method was higher in the rainy than in the post‐rainy season (RE_Tr= 1.220 vs 0.657) due to a higher genetic variance, lower G × E, and high h². When the 12 environments were classified into four clusters based on plant extractable soil‐water availability, the selection method Tr was superior to E in three of the four clusters (RE_Tr= 1.495, 0.612, 1.308, and 1.144) due to an increase in genetic variance and h² under Tr in clustered environments. Although the crosses exhibited significant differences for kernel yield, the two methods of selection did not interact significantly with crosses. Both methods contributed more or less equally to the 10 highest‐yielding selections (six for E and four for Tr). The six E selections had a higher kernel yield, higher transpiration (T), and nearly equal transpiration efficiency (TE) and harvest index (HI) relative to four Tr selections. The yield advantage in E selections came largely from greater T, which would likely not be an advantage in water‐deficient environments. From the results of these multi‐environment studies, it is evident that Tr method did not show a consistent superiority over E method of drought resistance breeding in producing a higher kernel yield in groundnut. Nonetheless, the integration of physiological traits (or their surrogates) in the selection scheme would be advantageous in selecting genotypes which are more efficient water utilisers or partitioners of photosynthates into economic yield. New biotechnological tools are being explored to increase efficiency of physiological trait‐based drought resistance breeding in groundnut.     

Transpiration efficiency and apparent cuticular transpiration in some c3 and c4 plants

Amphistomatous C₃ (Nicotiana tabacum L., Datura stramonium L.) and C₄ (Sorghum saccharatum Pers. and Zea mays L.) species were examined to find how (if at all) their inherent differences in water-use economy are reflected in apparent cuticular transpiration or vice versa. Transpiration efficiency (TE) was calculated from steady state photosynthesis (A) and transpiration (E) rates estimated for the upper side of the leaf after light induction of stomata opening. Apparent cuticular transpiration (’E_c) was measured as the part of transpiration which was not eliminated by convective counteraction of the air stream passing across the amphistomatous leaf: total pressure difference (AP) across the leaf was increased and the minimal value of E_ΔPτ0 was taken as the apparent cuticular transpiration rate (’E_c). ’E_c was treated relative to E at AP equal to zero (E_GDP=0), E’_cr. Measurements were carried out under two leaf-air vapour pressure differences (VPD). E_r (i.e. E_GDPτ0/E_GDP=0) versus GDP patterns differed qualitatively between the investigated C₃ and C₄ plants. TE increased and ’E_cr decreased from tobacco, stramony, maize to sorghum for both VPD of air. ’E_cr and TE were approximately linearly related, the slope being dependent on VPD. The increase in VPD resulted in larger E and slightly smaller epidermal conductance (g) at GDP equal to zero. Both E’_cr and E’_cr decreased markedly at the same time especially, for species with high TE. The results were considered as an indirect confirmation that E’_c values estimated by the technique used reflect species-specific differences in external peristomatal and cuticular vapour loss, at least in a relative sense.     

Drought tolerance in the alpine dandelion, Taraxacum ceratophorum (Asteraceae), its exotic congener T. officinale, and interspecific hybrids under natural and experimental conditions

We compared water relations and adaptations to drought stress in native and invasive exotic dandelions, Taraxacum ceratophorum and T. officinale. Photosynthesis (A), transpiration (E), and water use efficiency (WUE; carbon gained/water lost) were measured for the two species under extreme drought in the alpine tundra of Colorado, USA. We also subjected both species and F(1) hybrids to a dry-down experiment to determine how relative physiological performance varied with water availability. Photosynthesis and transpiration in the field were low and did not differ between Taraxacum congeners; however, native T. ceratophorum had higher WUE than T. officinale. After 6 days of greenhouse drought, photosynthesis and transpiration were reduced in T. officinale compared to T. ceratophorum. Taraxacum ceratophorum maintained high WUE under control and drought treatments. Conversely, WUE in T. officinale was highly plastic between watered (low WUE) and dry-down (high WUE) treatments. Hybrids did not exhibit heterosis; instead, they were similar to T. officinale in A and E and intermediate to the parental species in WUE. Overall, results suggest that native dandelions are more drought tolerant than invasive congeners or their hybrids, but have less plasticity in WUE. Arid habitats and occasional drought in mesic sites may provide native dandelions with refugia from negative interactions with invasives.     

The shifting influence of drought and heat stress for crops in northeast Australia

Characterization of drought environment types (ETs) has proven useful for breeding crops for drought‐prone regions. Here, we consider how changes in climate and atmospheric carbon dioxide (CO₂) concentrations will affect drought ET frequencies in sorghum and wheat systems of northeast Australia. We also modify APSIM (the Agricultural Production Systems Simulator) to incorporate extreme heat effects on grain number and weight, and then evaluate changes in the occurrence of heat‐induced yield losses of more than 10%, as well as the co‐occurrence of drought and heat. More than six million simulations spanning representative locations, soil types, management systems, and 33 climate projections led to three key findings. First, the projected frequency of drought decreased slightly for most climate projections for both sorghum and wheat, but for different reasons. In sorghum, warming exacerbated drought stresses by raising the atmospheric vapor pressure deficit and reducing transpiration efficiency (TE), but an increase in TE due to elevated CO₂ more than offset these effects. In wheat, warming reduced drought stress during spring by hastening development through winter and reducing exposure to terminal drought. Elevated CO₂ increased TE but also raised radiation‐use efficiency and overall growth rates and water use, thereby offsetting much of the drought reduction from warming. Second, adding explicit effects of heat on grain number and grain size often switched projected yield impacts from positive to negative. Finally, although average yield losses associated with drought will remain generally higher than that for heat stress for the next half century, the relative importance of heat is steadily growing. This trend, as well as the likely high degree of genetic variability in heat tolerance, suggests that more emphasis on heat tolerance is warranted in breeding programs. At the same time, work on drought tolerance should continue with an emphasis on drought that co‐occurs with extreme heat.     

CARD15 Gene Polymorphisms Are Associated with Tuberculosis Susceptibility in Chinese Holstein Cows

Bovine tuberculosis (BTB) is a significant veterinary and financial problem in many parts of the world. Associations between specific host genes and susceptibility to mycobacterial infections, such as tuberculosis, have been reported in several species. The objective of this study was to identify and evaluate the relationship of single-nucleotide polymorphisms (SNPs) in the CARD15 gene with susceptibility to BTB in Chinese Holstein cows. DNA samples from 201 Chinese Holstein cows (103 cases and 98 controls) were collected from Kunming City, Yuxi City, and Dali City in China. SNPs in the CARD15 gene were assessed using polymerase chain reaction (PCR) and restriction fragment length polymorphism-polymerase chain reaction (RFLP-PCR). Case-control association testing and statistical analysis identified six SNPs associated with susceptibility to BTB in Chinese Holstein cows. The frequency of genotypes C/T, A/G, A/G, A/G, C/T, and A/G in E4 (-37), 208, 1644, 1648, 1799, and E10 (+107), respectively, was significantly higher in cases than in controls, and also the alleles C, A, A, G, T, and A, respectively, were associated with a greater relative risk in cases than in controls. The distribution of two haplotypes, TGGACA and CAGACA, was significantly different between cases and controls. Overall, this case-control study suggested that E4 (-37)(C/T), 208(A/G), 1644(A/G), 1648(A/G), 1799(C/T), and E10 (+107)(A/G) in the CARD15 gene were significantly associated with susceptibility to BTB in Chinese Holstein cows and that haplotypes TGGACA and CAGACA could be used as genetic markers in marker-assisted breeding programs for breeding cows with high resistance to BTB.     

Physiological basis of genetic variation in leaf photosynthesis among rice (Oryza sativa L.) introgression lines under drought and well-watered conditions

To understand the physiological basis of genetic variation and resulting quantitative trait loci (QTLs) for photosynthesis in a rice (Oryza sativa L.) introgression line population, 13 lines were studied under drought and well-watered conditions, at flowering and grain filling. Simultaneous gas exchange and chlorophyll fluorescence measurements were conducted at various levels of incident irradiance and ambient CO(2) to estimate parameters of a model that dissects photosynthesis into stomatal conductance (g (s)), mesophyll conductance (g (m)), electron transport capacity (J (max)), and Rubisco carboxylation capacity (V (cmax)). Significant genetic variation in these parameters was found, although drought and leaf age accounted for larger proportions of the total variation. Genetic variation in light-saturated photosynthesis and transpiration efficiency (TE) were mainly associated with variation in g (s) and g (m). One previously mapped major QTL of photosynthesis was associated with variation in g (s) and g (m), but also in J (max) and V (cmax) at flowering. Thus, g (s) and g (m), which were demonstrated in the literature to be responsible for environmental variation in photosynthesis, were found also to be associated with genetic variation in photosynthesis. Furthermore, relationships between these parameters and leaf nitrogen or dry matter per unit area, which were previously found across environmental treatments, were shown to be valid for variation across genotypes. Finally, the extent to which photosynthesis rate and TE can be improved was evaluated. Virtual ideotypes were estimated to have 17.0% higher photosynthesis and 25.1% higher TE compared with the best genotype investigated. This analysis using introgression lines highlights possibilities of improving both photosynthesis and TE within the same genetic background.     

Differential gene expression of wheat progeny with contrasting levels of transpiration efficiency

High water use efficiency or transpiration efficiency (TE) in wheat is a desirable physiological trait for increasing grain yield under water-limited environments. The identification of genes associated with this trait would facilitate the selection for genotypes with higher TE using molecular markers. We performed an expression profiling (microarray) analysis of approximately 16,000 unique wheat ESTs to identify genes that were differentially expressed between wheat progeny lines with contrasting TE levels from a cross between Quarrion (high TE) and Genaro 81 (low TE). We also conducted a second microarray analysis to identify genes responsive to drought stress in wheat leaves. Ninety-three genes that were differentially expressed between high and low TE progeny lines were identified. One fifth of these genes were markedly responsive to drought stress. Several potential growth-related regulatory genes, which were down-regulated by drought, were expressed at a higher level in the high TE lines than the low TE lines and are potentially associated with a biomass production component of the Quarrion-derived high TE trait. Eighteen of the TE differentially expressed genes were further analysed using quantitative RT-PCR on a separate set of plant samples from those used for microarray analysis. The expression levels of 11 of the 18 genes were positively correlated with the high TE trait, measured as carbon isotope discrimination (Δ¹³C). These data indicate that some of these TE differentially expressed genes are candidates for investigating processes that underlie the high TE trait or for use as expression quantitative trait loci (eQTLs) for TE. Electronic Supplementary Material Supplementary material is available for this article at     

Genotypic and nutrition-dependent variation in water use efficiency and photosynthetic activity of leaves in winter wheat (Triticum aestivum L.)

Effects of high and reduced NPK nutrition on the genetic variation of components of water use efficiency at the leaf and whole-plant levels were examined in pot-grown old and modern cultivars of winter wheat (Triticum aestivum L.). At the subsequent growth stages, the photosynthetic rate (A), transpiration rate (E), leaf area (LA) and gas exchange efficiency (A/E) were measured on fully developed 4th, 5th, penultimate and flag leaves. At the plant canopy level, the total amount of water transpired was recorded during the whole life cycle to determine the efficiency of water use in the vegetative (WUEveg) and grain (WUEgen) matter formation. Considerable genotypic differences were found for the characters studied. The limited NPK supply caused a decrease in LA, A and A/E, but contributed to an increase in WUEgen. Examined cultivars (C) did not interact with nutrition levels (N) for these characteristics. However, the position of leaves (L) and the C x L and N x L interactions significantly affected the variance in leaf photosynthetic characteristics. A and A/E were negatively correlated with LA, and the flag leaves were photosynthetically less active and less efficient per unit area than the lower leaves. The whole-plant components of WUE were found to be more genetically stable than the photosynthetic leaf characteristics. Some modern cultivars tended to form leaves of higher A and A/E than the older ones, and this corresponds with a more efficient use of water in grain formation (WUEgen) of the former. Stay-green duration of flag leaves and harvest index showed positive correlations with WUEgen. However, no close associations were noticed between WUE components and stress tolerance, and the modern cultivars were usually less tolerant to NPK shortage.     

Validation of the effects of molecular marker polymorphisms in LcyE and CrtRB1 on provitamin A concentrations for 26 tropical maize populations

Vitamin A deficiency (VAD) compromises immune function and is the leading cause of preventable blindness in children in many developing countries. Biofortification, or breeding staple food crops that are rich in micronutrients, provides a sustainable way to fight VAD and other micronutrient malnutrition problems. Polymorphisms, with associated molecular markers, have recently been identified for two loci, LcyE (lycopene epsilon cyclase) and CrtRB1 (β-carotene hydroxylase 1) that govern critical steps in the carotenoid biosynthetic pathway in maize endosperm, thereby enabling the opportunity to integrate marker-assisted selection (MAS) into carotenoid breeding programs. We validated the effects of 3 polymorphisms (LcyE5′TE, LcyE3′Indel and CrtRB1-3′TE) in 26 diverse tropical genetic backgrounds. CrtRB1-3′TE had a two-ten fold effect on enhancing beta-carotene (BC) and total provitamin A (proA) content. Reduced-function, favorable polymorphisms within LcyE resulted in 0–30 % reduction in the ratio of alpha- to beta-branch carotenoids, and increase in proA content (sometimes statistically significant). CrtRB1-3′TE had large, significant effect on enhancing BC and total ProA content, irrespective of genetic constitution for LcyE5′TE. Genotypes with homozygous favorable CrtRB1-3′TE alleles had much less zeaxanthin and an average of 25 % less total carotenoid than other genotypes, suggesting that feedback inhibition may be reducing the total flux into the carotenoid pathway. Because this feedback inhibition was most pronounced in the homozygous favorable LcyE (reduced-function) genotypes, and because maximum total proA concentrations were achieved in genotypes with homozygous unfavorable or heterozygous LcyE, we recommend not selecting for both reduced-function genes in breeding programs. LcyE exhibited significant segregation distortion (SD) in all the eight, while CrtRB1 in five of eight digenic populations studied, with favorable alleles of both the genes frequently under-represented. MAS using markers reported herein can efficiently increase proA carotenoid concentration in maize.     

Carbon Isotope Discrimination is not Correlated with Transpiration Efficiency in Three Cool-Season Grain Legumes (Pulses)

The carbon Isotope discrimination (δ13C) of leaves has been shown to be correlated with the transpiration efficiency of leaves in a wide range of species. This has led to δ13C being used in breeding programs to select for improved transpiration efficiency. The correlation between δ13C and transpiration efficiency was determined under well-watered conditions during the vegetative phase In six genotypes of lentil (Lens culinaris Medikus), six genotypes of chickpea (Cicer arietinum L.) and 10 cultivars of narrow-leafed lupin (Lupinus angustifolius L.). Biomass (dry matter) accumulation and water use (transpiration)varied among the genotypes in all three species and transpiration efficiency was 40% to 75% higher In the most efficient compared with the least efficient genotypes. However, δ13C and transpiration efficiency were not significantly correlated in any of the species. This suggests that the δ13C technique cannot be used In selection for transpiration efficiency in the three grain legumes (pulses) studied.     

Large variation in whole-plant water-use efficiency among tropical tree species

It is well known that whole-plant water-use efficiency (transpiration efficiency of carbon gain, TE(C)) varies among plant species with different photosynthetic pathways. However, less is known of such variation among tree species within the C(3) group. Here we measured the TE(C) of seven C(3) tropical tree species. Isotopic analyses (delta(13)C, delta(18)O, and delta(15)N) and elemental analyses (carbon and nitrogen) were undertaken to provide insight into sources of variation in TE(C). Plants were grown over several months in approx. 80% full sunlight in individual 38-l containers in the Republic of Panama. Soil moisture content was nonlimiting. Significant variation was observed in TE(C) among the C(3) tree species. Values ranged from 1.6 mmol C mol(-1) H(2)O for teak (Tectona grandis) to 4.0 mmol C mol(-1) H(2)O for a legume, Platymiscium pinnatum. Variation in TE(C) was correlated with both leaf N concentration, a proxy for photosynthetic capacity, and oxygen-isotope enrichment, a proxy for stomatal conductance. The TE(C) varied with C-isotope discrimination within species, but the relationship broke down among species, reflecting the existence of species-specific offsets.     

Training does not protect against exhaustive exercise-induced lactate transport capacity alterations

The effects of endurance training on lactate transport capacity remain controversial. This study examined whether endurance training 1) alters lactate transport capacity, 2) can protect against exhaustive exercise-induced lactate transport alteration, and 3) can modify heart and oxidative muscle monocarboxylate transporter 1 (MCT1) content. Forty male Wistar rats were divided into control (C), trained (T), exhaustively exercised (E), and trained and exercised (TE) groups. Rats in the T and TE groups ran on a treadmill (1 h/day, 5 days/wk at 25 m/min, 10% incline) for 5 wk; C and E were familiarized with the exercise task for 5 min/day. Before being killed, E and TE rats underwent exhaustive exercise (25 m/min, 10% grade), which lasted 80 and 204 min, respectively (P < 0.05). Although lactate transport measurements (zero-trans) did not differ between groups C and T, both E and TE groups presented an apparent loss of protein saturation properties. In the trained groups, MCT1 content increased in soleus (+28% for T and +26% for TE; P < 0.05) and heart muscle (+36% for T and +33% for TE; P < 0.05). Moreover, despite the metabolic adaptations typically observed after endurance training, we also noted increased lipid peroxidation byproducts after exhaustive exercise. We concluded that 1) endurance training does not alter lactate transport capacity, 2) exhaustive exercise-induced lactate transport alteration is not prevented by training despite increased MCT1 content, and 3) exercise-induced oxidative stress may enhance the passive diffusion responsible for the apparent loss of saturation properties, possibly masking lactate transport regulation.     

Carbon Isotope Discrimination is not Correlated with Transpiration Efficiency in Three Cool-Season Grain Legumes （Pulses）

The carbon isotope discrimination （δ^13C） of leaves has been shown to be correlated with the transpiration efficiency of leaves in a wide range of species. This has led to δ^13C being used in breeding programs to select for improved transpiration efficiency. The correlation between δ^13C and transpiration efficiency was determined under well-watered conditions during the vegetative phase in six genotypes of lentil （Lens culinaris Medikus）, six genotypes of chickpea （Cicerarietinum L.） and 10 cultivars of narrow-leafed lupin （Lupinus angustifolius L.）. Biomass （dry matter） accumulation and water use （transpiration） varied among the genotypes in all three species and transpiration efficiency was 40% to 75% higher in the most efficient compared with the least efficient genotypes. However, δ^13C and transpiration efficiency were not significantly correlated in any of the species. This suggests that the δ^13C technique cannot be used in selection for transpiration efficiency in the three grain legumes （pulses） studied.     

Transpiration efficiency of a tropical pioneer tree (Ficus insipida) in relation to soil fertility

The response of whole-plant water-use efficiency, termed transpiration efficiency (TE), to variation in soil fertility was assessed in a tropical pioneer tree, Ficus insipida Willd. Measurements of stable isotope ratios (delta(13)C, delta(18)O, delta(15)N), elemental concentrations (C, N, P), plant growth, instantaneous leaf gas exchange, and whole-plant water use were used to analyse the mechanisms controlling TE. Plants were grown individually in 19 l pots with non-limiting soil moisture. Soil fertility was altered by mixing soil with varying proportions of rice husks, and applying a slow release fertilizer. A large variation was observed in leaf photosynthetic rate, mean relative growth rate (RGR), and TE in response to experimental treatments; these traits were well correlated with variation in leaf N concentration. Variation in TE showed a strong dependence on the ratio of intercellular to ambient CO(2) mole fractions (c(i)/c(a)); both for instantaneous measurements of c(i)/c(a) (R(2)=0.69, P <0.0001, n=30), and integrated estimates based on C isotope discrimination (R(2)=0.88, P <0.0001, n=30). On the other hand, variations in the leaf-to-air humidity gradient, unproductive water loss, and respiratory C use probably played only minor roles in modulating TE in the face of variable soil fertility. The pronounced variation in TE resulted from a combination of the strong response of c(i)/c(a) to leaf N, and inherently high values of c(i)/c(a) for this tropical tree species; these two factors conspired to cause a 4-fold variation among treatments in (1-c(i)/c(a)), the term that actually modifies TE. Results suggest that variation in plant N status could have important implications for the coupling between C and water exchange in tropical forest trees.