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Transpiration efficiency of dry matter production (W), carbon-isotope discrimination () and dry matter partitioning were measured on six sunflower (Helianthus annuus L.) genotypes grown for 32 days in a glasshouse. Two watering regimes, one well watered (HW) and the other delivering half the water used by the HW plants (LW), were imposed. Four major results emerged from this study. (1) There was significant genotypic variation in W in sunflower and this was closely reflected in Δ for both watering treatments. (2) The low watering regime caused a decrease in Δ but no change in W; nonetheless the genotypic ranking for either Δ or W was not significantly altered by water stress. (3) A positive correlation between W and biomass accumulation occurred among genotypes of HW plants. (4) Q, the ratio of total plant carbon content to leaf area, was positively correlated with W and negatively correlated with Δ. These results are discussed with reference to the connection between transpiration efficiency and plant growth. In short, Δ can be used to select for W among young vegetative sunflower plants. However, selection for W may be accompanied by changes in other important plant growth characteristics such as Q.

The relationship between carbon isotope discrimination, Δ, measured in plant dry matter and the ratio of intercellular to atmospheric partial pressures of CO2, pi/pa, in leaves was examined in two glasshouse experiments using 14 wheat genotypes selected on the basis of variation in Δ of dry matter. Genotypic variation in Δ was similar in both experiments, with an average range of 1.8 x 10-3. Variation in pi/pa was significant but the range in pi/pa was relatively small, averaging 0.075. In both experiments, Δ measured in dry matter and pi/pa measured in flag leaves were positively correlated. Variation among genotypes in pi/pa was attributed, approximately equally, to variation in leaf conductance and in photosynthetic capacity. The relationship between plant transpiration efficiency, W* (the amount of above-ground dry matter produced per unit water transpired) and was also examined. There was a negative correlation between W * and Δ; under well watered conditions and under gradually increasing terminal water stress. The relationship between W* of stressed plants and Δ measured in well watered plants was also negative. These results indicate that genotypic variation in Δ measured in dry matter should provide a reasonable measure of genotypic variation in long-term mean leaf pi/pa in wheat. Further, selection for improved plant transpiration efficiency in wheat under both well watered and terminally water- stressed conditions should be possible based on Δ measured in well watered plants. The extent to which such selection will be effective in improving transpiration efficiency at the field canopy level may depend on the influence of boundary layer resistance on transpirationsal water loss. Under well watered conditions and at full canopy closure, the influence of boundary layer resistance on canopy water loss may be relatively large and stomatal control of water loss may be limited. Under water stress, stomatal control of canopy water loss will be greater.

Variation in water-use efficiency (W, g of total dry matter produced/kg water used), and its correlation with cultivar isotope discrimination in leaves (Δ) was assessed in peanut plants grown in small canopies in the field. Plants were grown in separate minilysimeters that were both embedded in the ground and positioned above the crop. Differences among cultivars were found in W and and the relationship between W and Δ was compared for plants grown in open and closed canopies. Genetic variability in W in plants grown in the field under non-limiting water conditions was demonstrated, with Tifton-8, of Virginia habit, having the highest W (3.71 g/kg) and Rangkasbitung, an Indonesian cultivar of Spanish habit, the lowest (2.46 g/ kg). Variability in W was due to variation in total dry matter production more than that of water use. A strong negative correlation was found between Δ and W, and also between Δ and total dry matter. The relationship between whole plant W, including roots, and Δ was stronger than that between shoot W, without roots and Δ. The improvement occurred because of variation among cultivars in the root to shoot ratio. This highlights the importance of taking account of root dry matter in studies concerning W. There were significant differences in W and Δ between plants in pots above-ground compared to pots in the ground, with above-ground plants having significantly lower values of both W and Δ. The ranking of W and Δ among cultivars was not affected by the contrast in environment, which suggests these parameters are under strong genetic control. Total above-ground dry matter yield at maturity was negatively correlated with Δ, while pod yield was not. It appears a negative association between harvest index and Δ may exist; however not all cultivars used in this and other studies follow this response. Both water-use efficiency, Wand total dry matter production are negatively correlated with Δ in leaves of peanut plants grown in small canopies in the field. Measurement of Δ may prove a useful trait for selecting cultivars with improved W and total dry matter yield under field conditions.

The discrimination, Δ, against 13C in carbon accumulation by different genotypes of peanut was investigated by looking at changes in composition of isotopes of CO2 during CO2 assimilation and at the 13C/12C ratio in dry matter. The short term (minutes) discrimination, which was measured as CO2 passed over leaves in a gas exchange cuvette, was positively related to pi/pa, the ratio of the pressure of CO2 inside the leaves to the pressure of CO2 around the leaves, which is independently related to transpiration efficiency, W. Heritability of Δ was studied in plants grown in pots and in the field. Crosses were made of cultivars with contrasting Δ and W. For these genotypes, small Δ (and large W) seemed to be dominant in the F1 progeny. Severe drought stress slightly decreased Δ and W in F2 plants but, for parental genotypes in another experiment, Δ decreased but W increased when they were less severely stressed. A strong negative phenotypic correlation (r = -0.78) was found between W (g DM/ kg water used) and Δ in the leaves of well-watered potted plants of F2 progeny from a cross between two cultivars with contrasting W and Δ. Transpiration efficiency varied significantly among cultivars of peanut irrespective of whether plants were drought-stressed or well-watered. Transpiration efficiency in the potted F2 plants had a heritability of 34% while Δ had a heritability of 53%. The broad sense heritability of Δ in fieid-grown cultivars was 81% and there was no significant genotype × environment interaction for Δ. The high heritability of Δ and its strong genetic correlation with W indicates that Δ is a suitable selection criterion for W in peanut breeding programs.

Properties and display of foliage were studied in relation to gas exchange characteristics of tropical mangrove species. Rates of assimilation of CO2 were maximal at leaf temperatures of approximately 30°C. The species operated with higher water use efficiencies than do most C3 species, and water-use characteristics became increasingly conservative with increase in the salinity tolerance of the species. Changes in three properties of leaves, i.e. inclination, area, and succulence, contributed to maintenance of leaf temperatures near air temperatures with minimal evaporative cooling. Interspecific differences in water-use characteristics, and their relationship to the maintenance of favourable leaf temperatures, could affect the competitive abilities of mangroves and the structure of mangrove forests.

Stable carbon isotope discrimination (Δ) showed both intraspecific and interspecific variations in leaves collected from field plants representing a wide range of peanut germplasm grown in similar environments. It was predicted, on the basis of theoretical models relating Δ and water-use efficiency (W), that there could be as much as 60% variation in W. The Δ of leaf material was used as a guide to select peanut genotypes for testing, in the glasshouse, for a correlation between Δ and W. Pot plants of nine peanut genotypes were grown in conditions of unlimited water availability or very restricted water supply. Water-use efficiency and Δ of leaves was measured on all plants. A strong correlation (r = -0.81) was found between Δ and W. Water-use efficiency actually varied as much as twofold. Carbon content and Δ were measured in all parts of some plants. The Δ values of all parts were highly correlated with and therefore well-represented by the Δ of the leaves. Calculation of W on a molar basis removes variation associated with carbon content of dry matter. On this basis, W was correlated with the discrimination against 13C of the whole plant. The link between Δ and W is not direct but the strong correlation between Δ and W suggests that other factors, which are discussed, do not interfere greatly. The data provide more evidence that Δ might be used as a selection technique for W in C3 species.

Conventional gas-exchange techniques that measure the stomatal conductance and rate of CO2 assimilation of leaves were combined with measurements of the carbon isotope composition of CO2 in air passing over a leaf. Isotopic discrimination during uptake was determined from the difference in the carbon isotope composition of air entering and leaving the leaf chamber. Isotopic discrimination measured over the short term correlated strongly with that determined from combusted leaf material. Environmental conditions were manipulated to alter the relative influences of stomatal conductance and carboxylation on the discrimination of carbon isotopes by intact leaves. With C3 plants, discrimination increased as the gradient in partial pressure of CO2 across the stomata decreased. For C4 plants there was little change in discrimination despite substantial changes in the diffusion gradient across the sto- mata. These results are consistent with, and provide the first direct experimental support for, theoretical equations describing discrimination during photosynthesis. Despite uncertainties about various processes affecting carbon isotope composition, the resistance to the transfer of CO2 from the intercellular airspaces to the sites of carboxylation in the mesophyll chloroplasts was estimated using this technique. For wheat the estimated resistance was 1.2-2.4 m2 s bar mol -1.

Variation in carbon-isotope composition among and between wheat genotypes was correlated with variation in water-use efficiency in separate pot experiments conducted in spring-summer and in winter. In the main, winter experiment, the water-use efficiencies ranged from 2.0 to 3.7 mmolC/mol H2O (means of four replicates) while the corresponding isotope effects for leaf material ranged from 1.0225 to 1.0194. 13C was more abundant in grain than in leaves and stems. It is suggested that carbon-isotope analysis may be a useful tool in selection for improved water-use efficiency in breeding programmes for C3 species.

The temperature dependence of net photosynthetic assimilation of CO2 by snowgum (Eucalyptus pauciflora Sieb. ex Spreng.) was investigated. CO2 assimilation was divided into its component processes, stomatal and biochemical. The biochemical limitation was investigated with gas-exchange techniques and found to conform well to a recent model of C3 photosynthesis. In line with the model, net assimilation was further divided into ribulose 1,5-bisphosphate (RuP2) regeneration or electron-transport/photophosphorylation limitation, limitation by RuP2 carboxylase/oxygenase (Rubisco; EC 4.1.1.39) activity, together with loss of CO2 in non-photorespiratory respiration. The estimated temperature dependence of electron transport agreed well with one published for uncoupled electron transport, while the estimated temperature dependence of the catalytic activity of Rubisco was slightly less than that reported from biochemical determinations. The estimated rate of non-photorespiratory respiration was about 0.6 times the rate of respiration at night and appeared to have the same temperature dependence. With this information the temperature dependence of the biochemical limitation was modelled. Stomatal conductance was assumed to follow the theory of constant marginal water cost of carbon assimilation (δE/δA) and net assimilation rate at ambient concentration of CO2 was predicted. It was concluded that, for a given Rubisco activity and RuP2-regeneration capacity, both temperature optimum and net assimilation rate at the optimum temperature increased with increasing stomatal conductance.

An empirical model of stomatal conductance is presented. The predictions of conductance correlate with experimental determinations of the responses to several kinds of short-term and long-term perturbations, including irradiance, temperature, CO2 and O2 partial pressures, and leaf age and nitrogen nutrition. The model is not mechanistic and cannot account for the observed functioning of stomata independent of mesophyll. Nevertheless its success in predicting responses in intact leaves means that the mathematical expressions used must be analogous to those describing actual functioning.

Theory is developed to explain the carbon isotopic composition of plants with the dicarboxylic acid (C4) pathway of carbon fixation. It is shown how leakage of inorganic carbon from the bundle sheath cells to the mesophyll cells can affect carbon isotopic discrimination. It is suggested that different types of C4 pathways will show variation in the degree of leakiness, reflecting variations in anatomy, and in the requirements for oxygen egress from the bundle sheath cells. It is also suggested that quantum yields may reflect these variations. A simplified expression is developed relating discrimination to leakiness and the ratio of intercellular and atmospheric partial pressures of CO2.

Theory is developed to explain the carbon isotopic composition of plants. It is shown how diffusion of gaseous CO2 can significantly affect carbon isotopic discrimination. The effects on discrimination by diffusion and carboxylation are integrated, yielding a simple relationship between discrimination and the ratio of the intercellular and atmospheric partial pressures of CO2. The effects of dark respiration and photorespiration are also considered, and it is suggested that they have relatively little effect on discrimination other than via their effects on intercellular p(CO2). It is also suggested that various environmental factors such as light, temperature, salinity and drought will also have effects via changes in intercellular p(CO2). A simple method is suggested for assessing water use efficiencies in the field.

In order to understand the factors influencing the isotopic composition of air above and within plant canopies, equations quantifying the effects of photosynthesis, respiration and turbulent transport on the isotopic composition of the surrounding CO2 are developed. These equations are then extended to the regional scale, allowing the average isotopic composition of CO2 within the convective boundary layer to be related to the isotopic composition of tropospheric CO2, and to isotopic fractionations during ecosystem carbon exchange. Equations presented have the potential to be inverted, allowing direct estimation of isotopic fractionations by vegetation at the local and regional scales. Equations allowing the estimation of the extent of refixation of respired CO2 ('recycling') at the regional scale are also presented. Using measurements of CO2 carbon isotopic composition in conjunction with ecosystem flux measurements, the theory is applied to a tropical rain forest in Amazonia and a boreal forest in Siberia. When examined on a ground area basis and over the course of a day it is observed that, by virtue of greater fluxes but similar isotopic fractionations, the tropical rainforest exerts much more influence over the isotopic composition of the surrounding air than does the boreal forest. Due to higher rates of ecosystem respiration, recycling of respired CO2 is modelled to be much greater for tropical rainforest, but values presented here are considerably lower than previously published estimates, the latter being based solely on the relationship between the isotopic composition and concentrations of CO2 within forest canopies. The reasons for these differences are examined.

Distributions of common species of Eucalyptus in south-eastem Australia are related to gradients in temperature and rainfall. To determine whether intrinsic water-use efficiency (as indexed by carbon isotope discrimination, Δ) or other leaf attributes were related to climate of native habitats, we sampled 17 populations representing 14 species of Eucalyptus growing in a common garden in south- central New South Wales. Phreatophytes were clearly distinguished from populations that are totally dependent upon soil moisture derived from rainfall by having higher Δ at a particular level of rainfall. Among 12 non-phreatophytic populations (11 species), Δ was positively correlated with mean annual precipitation (r = 0.75, P = 0.005), December-March precipitation (r = 0.79, P = 0.002), an index of annual soil moisture (r = 0.81, P = 0.001) and seasonality of precipitation (r = 0.85, P < 0.001). There were similarly strong but negative correlations between Δ and potential evaporation during the summer months, but Δ was not correlated with annual potential evaporation of the source sites. Leaf mass per unit area (&sigmaf;e) was negatively correlated with indices of water availability, positively correlated with nitrogen per unit leaf area (r = 0.90, P < 0.001), and negatively correlated with Δ (r = -0.73, P = 0.007). A was negatively correlated with area-based leaf nitrogen (r = -0.79, P = 0.002). These complementary correlations among Δ, &sigmaf;e, and nitrogen per unit leaf area indicate that variation in Δ may stem largely from variation in photosynthetic capacity. The results provide strong evidence that variation in Δ and &sigmaf;e reflect genetic adaptations to native habitats.

Variation in cahn isotope discrimination (Δ) and ribulose-1,5-bisphosphate carboxylase-oxygenase (Rubisco) content per unit leaf area was examined in leaves from upper and lower positions in the canopy of six groundnut (Arachis hypogaea L.) genotypes, grown under irrigated and mild water-deficit conditions in the field. The leaf mass per unit leaf area (&sigmaf;L) and soluble proteins in leaves were determined at 80, 96, 111 and 127 days after sowing (DAS), while Δ and Rubisco were determined at 80 DAS only. The mean Δ ranged from 18.2 to 20.20 among genotypes, representing a significant (P < 0.01) variation. Rubisco content per unit leaf area also varied significantly (P < 0.01) with genotype and leaf position. There was a trend to an increase in Rubisco content under water deficit, but the effects were not significant. Leaves at the top of the canopy had a higher Rubisco content and lower Δ, than leaves at the bottom of the canopy. Genotype × leaf position interaction was significant for Δ and Rubisco, indicating the importance of leaf position in selecting for water-use efficiency (W), using leaf traits in groundnut. Rubisco content and Δ were negatively related (r2 = 0.65, P < 0.01). There was a significantly positive correlation between Rubisco content and &sigmaf;L in the upper leaves (r2 = 0.60, P < 0.01 ), but not in the lower leaves in the canopy. However, the overall relationship between Rubisco and &sigmaf;L (r2 = 0.40) was not as strong as it was between Rubisco and Δ. The results suggest that, in groundnut, the basis of genotypic variation in was mostly (> 60%) attributable to Rubisco content per unit leaf area. In view of the leaf positional effects on Δ and Rubisco, the upper leaves in the canopy should be used for selecting genotypes for W based on leaf traits like &sigmaf;L or Δ.

Various aspects of the water economy were investigated for a range of herbaceous species varying in relative growth rate. Plants were grown in a growth chamber with a non-limiting supply of water and nutrients, and the rate of transpiration, short-term intercellular CO2 concentration and long-term carbon isotope discrimination (Δ) in the leaves were determined. No correlation was found between the relative growth rate of these species, and the transpiration rate per unit leaf area, the intercellular CO2 concentration and the 13C-discrimination. There was a positive correlation, however, with the rate of water uptake per unit root weight. From these observations we infer that the previously observed differences in photosynthetic nitrogen-use efficiency, the rate of photosynthesis per unit leaf nitrogen, can not be explained by variation in intercellular CO2 concentration. These data were also used to analyse correlations between Δ and both growth parameters and chemical composition. Apart from parameters related to the water economy, Δ only correlated (positively) with the fractional biomass allocation to the roots (root weight ratio) and the specific root length (root length divided by root weight), and not with any other investigated growth parameters.

Photosynthetic gas exchange measurements were combined with measurements of the carbon and oxygen stable isotopic composition of CO2 after it passed over a leaf of Phaseolus vulgaris or Senecio spp. plants held in a controlled environment chamber. Calculations were then made of discrimination by the leaf against 13CO2 and C18O16O. Leaves were maintained at different vapour pressure gradients in order to generate a range of leaf water 18CO/16CO ratios. The 18CO content of leaf water increased when plants were exposed to higher vapour pressure deficits. The observed C18O16O discrimination values also increased with an increase in the leaf-air vapour pressure gradient and the associated change in leaf water 18/CO16CO values. In addition, the observed C18O16O discrimination values were strongly correlated with values predicted by a mechanistic model of isotopic fractionation.

The relationship between transpiration ratio (R, ratio of water transpired to carbon fixed) or cabon isotopic discrimination (Δ) and leaf dry-matter mineral content (m) was examined within several C3 species including monocot and dicotyledonous species (Triticum aestivum, Hordeum spontaneum, Helianthus annuus and Nicotiana tabacum) and the C4 species, Sorghum bicolor. In all species, whether grown in the glasshouse or in the field, m was positively correlated to R; consistently, m was positively correlated to Δ in the C3 species, and negatively in sorghum. These genetic correlations between transpiration ratio and mineral content contrast with the absence of such a relationship when variations of R were environmentally induced by changes of atmospheric humidity or carbon dioxide concentration. In those circumstances, and consistent with earlier reports, ash content or its major constituents were remarkably stable and insensitive to a doubling or more of R. The genetic relationships shown in the present study are to some extent brought about by variations in assimilation and transpiration rates per se (i.e. passive mineral uptake), but the data indicate that they involve other quantitatively more important mechanisms, some of which may be related to the control of leaf water content. With the exception of Hordeum spontaneum, potassium was the element contributing the most to the relationship between R, or Δ, and mineral content. Depending on the species other elements showed a significant correlation with R or Δ. Further experiments are needed to unravel the nature of the genetic association between R and m. Meanwhile plant mineral content may be a useful criterion in selection programs to improve transpiration efficiency since its correlation with R was not much lower than that of carbon isotope discrimination.

Carbon isotope discrimination (Δ) and leaf gas-exchange were measured simultaneously for a number of C4 species. Linear relationships were found between A and the ratio of intercellular to ambient partial pressures of CO2, pI/pa. These data were used to estimate the fraction of CO2 released by C4-acid decarboxylation in the bundle sheath, which subsequently leaks out to the mesophyll. We define this fraction as the leakiness of the system and it is also a measure of the extent to which phosphoenolpyruvate (PEP) carboxylations exceed ribulose 1,5-bisphosphate (RuBP) carboxylations. For Sorghum bicolor and Amaranthus edulis, leakiness was estimated at 0.2 and was constant over a wide range of irradiances (between 480 and 1600 μmol quanta m-2 s-1), intercellular CO2 pressures (between 30 and 350 μbar) and leaf temperatures (from 21C to 34C). At irradiances less than 240 μmol quanta m-2 s-1, leakiness appeared to increase. For a number of dicotyledonous and monocotyledonous species, of the various C4-decarboxylation types, leakiness was also estimated at 0.2. Contrary to expectation, amongst the 11 species examined, those with suberised lamellae did not show lower values of leakiness than those without suberised lamellae. For one NAD-ME and one PCK monocot, the estimates of leakiness were significantly higher at 0.30 and 0.25, respectively. Long-term discrimination (assessed from carbon isotope composition of leaf dry matter) did not correlate well with these short- term measures of discrimination. We suggest that this may be due to differences between species in fractionations occurring after photosynthesis.

Wheat seedlings were grown on soils with different resistances to root penetration and at various ambient partial pressures of CO2, pa. The partial pressure pa was varied either (a) as soon as seedlings emerged and until the end of the experiment (i.e. until c. 2 weeks after sowing, until the three leaf stage) or (b) only after the effects of high soil resistance were well established, and then for two days. In the first week after emergence, a higher soil resistance induced slower growth of both roots and shoot. This effect was independent of pa. We conclude that at this stage, with much of the plant's carbon being supplied from hydrolysis of seed reserves, growth at high soil resistance was not C-source limited. There was, however, a positive relationship between shoot growth rate and the carbohydrate concentration in that tissue, the degree of which varied with soil resistance. A given carbohydrate concentration was associated with a lower growth rate at high, compared with low, soil resistance. We deduced that this reduced sensitivity of growth to internal availability of substrate carbohydrate may be one manifestation of a sink limitation. Subsequent to the first week following emergence, roots grew faster on soil with higher resistance, while the shoot continued to grow more slowly. As seed reserves were becoming exhausted, growth became sensitive to pa, i.e. somewhat source-limited. This response to partial pressure of CO2 was mainly seen in the roots. The correlation between growth rate and carbohydrate levels was maintained. We conclude that increased soil resistance induces a factor which retards shoot growth, partly by decreasing its sensitivity to carbohydrate levels, making more carbon available for root growth. It is unclear whether or not this factor also directly affects the physiology of the roots. A higher soil resistance resulted in greater respiratory losses as a proportion, Φ, of carbon fixation. Reduced pa also induced greater Φ. Theory is developed relating Φ to carbon assimilation, allocation and maintenance. It leads to the prediction that respiratory losses, as a proportion, Φ, should be increased under most environmental conditions which reduce relative growth rate.

Intact leaves of N. glauca and C. avellana were exposed to a range of humidities and their gas exchange monitored. Rates of transpiration and assimilation of carbon dioxide, and their sensitivities to changes in total conductance (leaf and boundary layer) were determined. The ratio of these sensitivities, δE/δA, remained substantially constant over the range of humidities. The results represent the first experimental support for a recent hypothesis that stomata vary their apertures in such a manner as to keep δE/δA constant, which optimizes carbon gain with respect to water loss.

Models for the responses of stomatal conductance to changes in humidity difference between leaf and air are developed. Feedforward of the stomatal response to humidity difference is present when stomata respond to a portion of the rate of transpiration of the leaf that is unaffected by changes in stomatal aperture (cuticular transpiration). A simple linear model of the responses of conductance is sufficient to predict many phenomena reported in the literature. These include the presence, under specified conditions, of a peak rate of transpiration, and an increase in the conductance and humidity difference at which this peak occurs, with increased general levels of stomatal opening.