A critical appraisal of a combined stomatal-photosynthesis model for C3 plants

Gas-exchange measurements on Eucalyptus grandis leaves and data extracted from the literature were used to test a semi-empirical model of stomatal conductance for CO₂ g_Sc=g_o+a₁A/(c_s-I) (1+D_s/D_o)] where A is the assimilation rate; D_s and c_s are the humidity deficit and the CO₂ concentration at the leaf surface, respectively; g₀ is the conductance as A → 0 when leaf irradiance → 0; and D₀ and a₁ are empirical coefficients. This model is a modified version of g_sc=a₁A h_s/c_s first proposed by Ball, Woodrow & Berry (1987, in Progress in Photosynthesis Research, Martinus Mijhoff, Publ., pp. 221–224), in which h_s is relative humidity. Inclusion of the CO₂ compensation point, τ, improved the behaviour of the model at low values of c_s, while a hyperbolic function of D_s for humidity response correctly accounted for the observed hyperbolic and linear variation of g_sc and c_i/c_s as a function of D_s, where C_i is the intercellular CO₂ concentration. In contrast, use of relative humidity as the humidity variable led to predictions of a linear decrease in g_sc and a hyperbolic variation in c_i/c_s as a function of D_s, contrary to data from E. grandis leaves. The revised model also successfully described the response of stomata to variations in A, D_s and c_s for published responses of the leaves of several other species. Coupling of the revised stomatal model with a biochemical model for photosynthesis of C₃ plants synthesizes many of the observed responses of leaves to light, humidity deficit, leaf temperature and CO₂ concentration. Best results are obtained for well-watered plants.     

Drought-inhibition of photosynthesis in C3 plants: stomatal and non-stomatal limitations revisited

There is a long-standing controversy as to whether drought limits photosynthetic CO2 assimilation through stomatal closure or by metabolic impairment in C3 plants. Comparing results from different studies is difficult due to interspecific differences in the response of photosynthesis to leaf water potential and/or relative water content (RWC), the most commonly used parameters to assess the severity of drought. Therefore, we have used stomatal conductance (g) as a basis for comparison of metabolic processes in different studies. The logic is that, as there is a strong link between g and photosynthesis (perhaps co-regulation between them), so different relationships between RWC or water potential and photosynthetic rate and changes in metabolism in different species and studies may be 'normalized' by relating them to g. Re-analysing data from the literature using light-saturated g as a parameter indicative of water deficits in plants shows that there is good correspondence between the onset of drought-induced inhibition of different photosynthetic sub-processes and g. Contents of ribulose bisphosphate (RuBP) and adenosine triphosphate (ATP) decrease early in drought development, at still relatively high g (higher than 150 mmol H20 m(-2) s(-1)). This suggests that RuBP regeneration and ATP synthesis are impaired. Decreased photochemistry and Rubisco activity typically occur at lower g (<100 mmol H20 m(-2) s(-1)), whereas permanent photoinhibition is only occasional, occurring at very low g (<50 mmol H20 m(-2) s(-1)). Sub-stomatal CO2 concentration decreases as g becomes smaller, but increases again at small g. The analysis suggests that stomatal closure is the earliest response to drought and the dominant limitation to photosynthesis at mild to moderate drought. However, in parallel, progressive down-regulation or inhibition of metabolic processes leads to decreased RuBP content, which becomes the dominant limitation at severe drought, and thereby inhibits photosynthetic CO2 assimilation.     

Drought limitation of photosynthesis differs between C3 and C4 grass species in a comparative experiment

Phylogenetic analyses show that C₄ grasses typically occupy drier habitats than their C₃ relatives, but recent experiments comparing the physiology of closely related C₃ and C₄ species have shown that advantages of C₄ photosynthesis can be lost under drought. We tested the generality of these paradoxical findings in grass species representing the known evolutionary diversity of C₄ NADP‐me and C₃ photosynthetic types. Our experiment investigated the effects of drought on leaf photosynthesis, water potential, nitrogen, chlorophyll content and mortality. C₄ grasses in control treatments were characterized by higher CO₂ assimilation rates and water potential, but lower stomatal conductance and nitrogen content. Under drought, stomatal conductance declined more dramatically in C₃ than C₄ species, and photosynthetic water‐use and nitrogen‐use efficiency advantages held by C₄ species under control conditions were each diminished by 40%. Leaf mortality was slightly higher in C₄ than C₃ grasses, but leaf condition under drought otherwise showed no dependence on photosynthetic‐type. This phylogenetically controlled experiment suggested that a drought‐induced reduction in the photosynthetic performance advantages of C₄ NADP‐me relative to C₃ grasses is a general phenomenon.     

Antitranspirant functions of stem periderms and their influence on corticular photosynthesis under drought stress

Transpiration and photosynthesis of current-year stems and adult leaves of different deciduous tree species were investigated to estimate their probable influence on carbon balance. Peridermal transpiration of young stems was found to be rather small as compared to the transpiration of leaves (stem/leaf like 1/5–1/20). A characteristic that was mainly attributable to the lower peridermal conductance to water and CO₂, which made up only 8–28% of stomatal conductance. Water vapour conductance was significantly lower in stems, but also non-responsive to PAR, which led to a comparatively higher water use efficiency (WUE, ratio assimilation/transpiration). Thus, although corticular photosynthesis reached only 11–37% of leaf photosynthesis, it may be a means of improving the carbon balance of stems under limited water availability. The influence of drought stress on primary photosynthetic reactions was also studied. Under simulated drought conditions the drying time needed to provoke a 50% reduction (t ₅₀) in dark- and light-adapted PSII efficiency (Fv/Fm, ΔF/Fm′) was up to ten times higher in stems than in leaves. Nevertheless, up to a relative water deficit (RWD) of around 40–50% dark-adapted PSII efficiency of leaves and stems was rather insensitive to dehydration, showing that the efficiency of open PS II reaction centres is not impaired. Thus, it may be concluded that in stems as well as in leaves the primary site of drought damage is at the level of dark enzyme reactions and not within PSII. However, enduring severe drought caused photoinhibitory damage to the photosynthetic apparatus of leaves and stems; thereby RWD₅₀ values (= RWD needed to provoke a 50% reduction in Fv/Fm ad ΔF/Fm′) were comparably lower in stems as compared to leaves, indicating a possibly higher drought sensitivity of the cortex chlorenchyma.     

Seasonal differences in photosynthesis between the C3 and C4 subspecies of Alloteropsis semialata are offset by frost and drought

The regional abundance of C₄ grasses is strongly controlled by temperature, however, the role of precipitation is less clear. Progress in elucidating the direct effects of photosynthetic pathway on these climate relationships is hindered by the significant genetic divergence between major C₃ and C₄ grass lineages. We addressed this problem by examining seasonal climate responses of photosynthesis in Alloteropsis semialata , a unique grass species with both C₃ and C₄ subspecies. Experimental manipulation of rainfall in a common garden in South Africa tested the hypotheses that: (1) photosynthesis is greater in the C₄ than C₃ subspecies under high summer temperatures, but this pattern is reversed at low winter temperatures; and (2) the photosynthetic advantage of C₄ plants is enhanced during drought events. Measurements of leaf gas exchange over 2 years showed a significant photosynthetic advantage for the C₄ subspecies under irrigated conditions from spring through autumn. However, the C₄ leaves were killed by winter frost, while photosynthesis continued in the C₃ plants. Unexpectedly, the C₄ subspecies also lost its photosynthetic advantage during natural drought events, despite greater water-use efficiency under irrigated conditions. This study highlights previously unrecognized roles for climatic extremes in determining the ecological success of C₃ and C₄ grasses.     

外源褪黑素对高温胁迫下菊花光合和生理特性的影响

以切花菊品种'神马'为试材,研究外源褪黑素(MT)对菊花抗高温胁迫的影响.将供试菊花叶面喷施200 μmol·L-1的MT后,进行40℃(昼)/35℃(夜)高温胁迫,观察菊花叶片叶绿体和内囊体超微结构,测定光合和生理指标.结果表明:与常温对照(CK)相比,高温胁迫下菊花叶片叶绿体和类囊体受损,叶绿素含量和最大荧光(Fm...     

Biochar amendment boosts photosynthesis and biomass in C3 but not C4 plants: A global synthesis

Biochar is a carbon (C)‐rich solid produced from the thermochemical pyrolysis of biomass. Its amendment to soils has been proposed as a promising mean to mitigate greenhouse gas emissions and simultaneously benefit agricultural crops. However, how biochar amendment affects plant photosynthesis and growth remains unclear, especially on a global scale. In this study, we conducted a global synthesis of 74 publications with 347 paired comparisons to acquire an overall tendency of plant photosynthesis and growth following biochar amendment. Overall, we found that biochar amendment significantly increased photosynthetic rate by 27.1%, and improved stomatal conductance, transpiration rate, water use efficiency, and chlorophyll concentration by 19.6%, 26.9%, 26.8%, and 16.1%, respectively. Meanwhile, plant total biomass, shoot biomass, and root biomass increased by 25.4%, 22.1%, and 34.4%, respectively. Interestingly, plant types (C₃ and C₄ plants) showed greater control over plant photosynthesis and biomass than a broad suite of soil and biochar factors. Biochar amendment largely boosted photosynthesis and biomass on C₃ plants, but had a limited effect on C₄ plants. Our results highlight the importance of the differential response of plant types to biochar amendment with respect to plant growth and photosynthesis, providing a scientific foundation for making reasonable strategies towards an extensive application of biochar for agricultural production management.     

外源钙离子对盐胁迫玉米气孔特征、光合作用和生物量的影响

探讨盐胁迫下玉米气孔特征、光合作用和生物量对外源钙离子的响应,有助于深入理解添加外源钙离子(Ca²⁺)缓解玉米盐胁迫的作用机理.以‘京科665’品种为试材,研究了NaCl胁迫下(100 mmol·L^-1)添加不同浓度外源Ca²⁺(0、5、10、20、40、80 mmol·L^-1)对玉米幼苗气孔特征、光合作用和生物量的影响.结果表明: 不同Ca²⁺浓度对盐胁迫下玉米的气孔密度影响不大,但显著减小了气孔形状指数、气孔面积、气孔长度、气孔宽度和气孔周长.同时,随着外源Ca²⁺浓度的逐渐提高,玉米叶片的净光合速率(P_n)呈先升高后降低的趋势,且气孔导度(g_s)和胞间CO₂浓度(C_i)均显著降低,表明不同浓度Ca²⁺通过改变玉米气孔结构特征进一步限制光合作用过程,最终导致P_n降低.另外,外源Ca²⁺促进盐胁迫下玉米幼苗生物量增加,但根冠比显著降低,表明盐胁迫下添加外源Ca²⁺对地上部分的缓解作用大于地下部分.     

Analysis of inhibition of photosynthesis due to water stress in the C3 species Hordeum vulgare and Vicia faba: Electron transport, CO2 fixation and carboxylation capacity

A C₃ monocot, Hordeum vulgare and C₃ dicot, Vicia faba, were studied to evaluate the mechanism of inhibition of photosynthesis due to water stress. The net rate of CO₂ fixation (A) and transpiration (E) were measured by gas exchange, while the true rate of O₂ evolution (J _O2) was calculated from chlorophyll fluorescence analysis through the stress cycle (10 to 11 days). With the development of water stress, the decrease in A was more pronounced than the decrease in J _O2 resulting in an increased ratio of Photosystem II activity per CO₂ fixed which is indicative of an increase in photorespiration due to a decrease in supply of CO₂ to Rubisco. Analyses of changes in the J _O2 A ratios versus that of CO₂ limited photosynthesis in well watered plants, and RuBP pool/RuBP binding sites on Rubisco and RuBP activity, indicate a decreased supply of CO₂ to Rubisco under both mild and severe stress is primarily responsible for the decrease in CO₂ fixation. In the early stages of stress, the decrease in C _i (intercellular CO₂) due to stomatal closure can account for the decrease in photosynthesis. Under more severe stress, CO₂ supply to Rubisco, calculated from analysis of electron flow and CO₂ exchange, continued to decrease. However, C _i, calculated from analysis of transpiration and CO₂ exchange, either remained constant or increased which may be due to either a decrease in mesophyll conductance or an overestimation of C _i by this method due to patchiness in conductance of CO₂ to the intercellular space. When plants were rewatered after photosynthesis had dropped to 10–30% of the original rate, both species showed near full recovery within two to four days.Abbreviations A- net CO₂ assimilation rate - A ^*- net CO₂ assimilation rate plus dark respiration - ATP- adenosine triphosphate - CABP- carboxyarabinitol 1,5-bisphosphate - C _a- ambient CO₂ concentration - C _c- CO₂ concentration in the chloroplast - C _i- intercellular CO₂ concentration - E- transpiration rate - g _m- mesophyll conductance - g _s- stomatal conductance - J _O2 true rate of O₂ evolution - LSD- least significant difference - PPFD- photosynthetic photon flux density - PS II- Photosystem II - R _n- dark respiration rate - Rubisco- ribulose 1,5-bisphosphate carboxylase/oxygenase - RuBP- ribulose 1,5-bisphosphate - RWC- relative water content - _c- rate of carboxylation - _o- rate of oxygenation - _PSII- quantum yield of Photosystem II - - CO₂ compensation point in the absence of R _n - - water potential     

Physiological significance of proline and glycinebetaine: Maintaining photosynthesis during NaCl stress in wheat

Experiments on the physiological significance of accumulation of proline and glycinebetaine (GB) in sustaining photosynthesis during salt stress in wheat in vivo showed that pre-treatment with GB, but not proline, alleviated NaCl-induced stomatal and non-stomatal inhibition of photosynthesis completely. A permeating and non-dissociating osmoticum, 3-orthomethyl-glucopyranose, also alleviated NaCl-induced perturbations of photosynthesis, suggesting that GB may work by maintaining chloroplast volume and not by specific effects on photosynthetic processes.     

Drought stress effects on three cultivars of Eragrostis curvula: photosynthesis and water relations

Water stress effects were studied on three cultivars ofEragrostis curvula. Leaf water potential, RWC, total plantleaf area, green dry weight mass percentage and CO₂ gas-exchangeweremeasured during the onset of stress and after recovery. After 3 days of waterstress, RWC of cv Tanganyika plants was around 30–40% of controls,while RWC of cvs Ermelo and Consol was around 50–60% of controls.However midday and predawn water potentials were lower in cvs Tanganyka andErmelo than in cv Consol. After re-watering, RWC and water potentials recoveredonly in Consol plants. A strong decrease of leaf area was recorded in cvsErmeloand Consol during water stress (about 91–94% less than the leafarea of controls). Photosynthesis decreased as a function of the degree ofwaterstress severity in all cultivars. Also, light saturated photosynthesis,CO₂ quantum yield and light at which saturated photosynthesisoccurred, were strongly reduced by water stress. Recovery of photosynthesis wasfound in cv Consol after five days re-watering. Cv Consol showed a betterconservation of water and higher resistance to water stress than the other twocvs.     

Structural, biochemical, and physiological characterization of C4 photosynthesis in species having two vastly different types of kranz anatomy in genus Suaeda (Chenopodiaceae)

C (4) species of family Chenopodiaceae, subfamily Suaedoideae have two types of Kranz anatomy in genus Suaeda, sections Salsina and Schoberia, both of which have an outer (palisade mesophyll) and an inner (Kranz) layer of chlorenchyma cells in usually semi-terete leaves. Features of Salsina (S. AEGYPTIACA, S. arcuata, S. taxifolia) and Schoberia type (S. acuminata, S. Eltonica, S. cochlearifoliA) were compared to C (3) type S. Heterophylla. In Salsina type, two layers of chlorenchyma at the leaf periphery surround water-storage tissue in which the vascular bundles are embedded. In leaves of the Schoberia type, enlarged water-storage hypodermal cells surround two layers of chlorenchyma tissue, with the latter surrounding the vascular bundles. The chloroplasts in Kranz cells are located in the centripetal position in Salsina type and in the centrifugal position in the Schoberia type. Western blots on C (4) acid decarboxylases show that both Kranz forms are NAD-malic enzyme (NAD-ME) type C (4) species. Transmission electron microscopy shows that mesophyll cells have chloroplasts with reduced grana, while Kranz cells have chloroplasts with well-developed grana and large, specialized mitochondria, characteristic of NAD-ME type C (4) chenopods. In both C (4) types, phosphoenolpyruvate carboxylase is localized in the palisade mesophyll, and Rubisco and mitochondrial NAD-ME are localized in Kranz cells, where starch is mainly stored. The C (3) species S. heterophylla has Brezia type isolateral leaf structure, with several layers of Rubisco-containing chlorenchyma. Photosynthetic response curves to varying CO (2) and light in the Schoberia Type and Salsina type species were similar, and typical of C (4) plants. The results indicate that two structural forms of Kranz anatomy evolved in parallel in species of subfamily Suaedoideae having NAD-ME type C (4) photosynthesis.     

The acclimation of photosynthesis and respiration to temperature in the C3–C4 intermediate Salsola divaricata: induction of high respiratory CO2 release under low temperature

Photosynthesis in C₃–C₄ intermediates reduces carbon loss by photorespiration through refixing photorespired CO₂ within bundle sheath cells. This is beneficial under warm temperatures where rates of photorespiration are high; however, it is unknown how photosynthesis in C₃–C₄ plants acclimates to growth under cold conditions. Therefore, the cold tolerance of the C₃–C₄ Salsola divaricata was tested to determine whether it reverts to C₃ photosynthesis when grown under low temperatures. Plants were grown under cold (15/10 °C), moderate (25/18 °C) or hot (35/25 °C) day/night temperatures and analysed to determine how photosynthesis, respiration and C₃–C₄ features acclimate to these growth conditions. The CO₂ compensation point and net rates of CO₂ assimilation in cold‐grown plants changed dramatically when measured in response to temperature. However, this was not due to the loss of C₃–C₄ intermediacy, but rather to a large increase in mitochondrial respiration supported primarily by the non‐phosphorylating alternative oxidative pathway (AOP) and, to a lesser degree, the cytochrome oxidative pathway (COP). The increase in respiration and AOP capacity in cold‐grown plants likely protects against reactive oxygen species (ROS) in mitochondria and photodamage in chloroplasts by consuming excess reductant via the alternative mitochondrial respiratory electron transport chain.     

Effects of silicon on photosynthesis,water relations and nutrient uptake of <Emphasis Type="Italic">Phaseolus vulgaris</Emphasis> under NaCl stress

A greenhouse experiment was conducted to investigate the effects of silicon application on Phaseolus vulgaris L. under two levels of salt stress (30 and 60 mM NaCl in the irrigation water). Salinity significantly reduced growth, stomatal conductance and net photosynthetic rate, and increased Na⁺ and Cl⁻ content mainly in roots. Silicon application enhanced growth of salt stressed plants, significantly reduced Na⁺ content especially in leaves and counterbalanced the effects of NaCl on gas exchange; the effect was more evident at 30 mM NaCl. Cl⁻ content in shoots and roots was not significantly modified by silicon application; the drop in K⁺ content caused by salinity was partially counterbalanced by silicon, especially in roots.     

Correlation between the inhibition of photosynthesis and the decrease in area of detached leaf discs or volume/absorbance of protoplasts under osmotic stress in pea (Pisum sativum)

Exposure to osmotic stress reduces leaf area and protoplast volume while decreasing photosynthesis. But the measurement of protoplast volume is tedious, while rapid determinations of leaf area in the field are difficult. We evaluated the quantitative relationship between the extent of decrease in area of detached leaf discs or the volume of protoplast of pea ( Pisum sativum ) and reduction in their photosynthetic capacity under osmotic stress. Osmotic stress was induced by increasing sorbitol concentration in the surrounding medium of the leaf discs from zero to 1.0 M (-3.1 MPa), and in case of protoplasts from 0.4 M (-1.3 MPa, isotonicity) to 1.0 M (-3.1 MPa, hypertonicity). There was a high degree of positive correlation between the extent of reduction in the area of detached leaf discs or the volume of protoplasts (indicated by diameter or absorption at 440 nm) and the decrease in photosynthesis. The correlation coefficients between inhibition of photosynthesis and the decrease in leaf disc area or protoplast volume were 0.96 and 0.99, respectively. We therefore suggest that the decrease in absorbance at 440 nm (corrected for turbidity at 750 nm) can be used as a simple measure to predict the inhibition due to osmotic stress of photosynthesis in mesophyll protoplasts. Similarly, the reduction in area of detached leaf discs could also be a very simple and useful criterion to assess osmotic tolerance of photosynthesis.     

No evidence for triose phosphate limitation of light‐saturated leaf photosynthesis under current atmospheric CO2 concentration

The triose phosphate utilization (TPU) rate has been identified as one of the processes that can limit terrestrial plant photosynthesis. However, we lack a robust quantitative assessment of TPU limitation of photosynthesis at the global scale. As a result, TPU, and its potential limitation of photosynthesis, is poorly represented in terrestrial biosphere models (TBMs). In this study, we utilized a global data set of photosynthetic CO₂ response curves representing 141 species from tropical rainforests to Arctic tundra. We quantified TPU by fitting the standard biochemical model of C₃ photosynthesis to measured photosynthetic CO₂ response curves and characterized its instantaneous temperature response. Our results demonstrate that TPU does not limit leaf photosynthesis at the current ambient atmospheric CO₂ concentration. Furthermore, our results showed that the light‐saturated photosynthetic rates of plants growing in cold environments are not more often limited by TPU than those of plants growing in warmer environments. In addition, our study showed that the instantaneous temperature response of TPU is distinct from temperature response of the maximum rate of Rubisco carboxylation. The new formulations of the temperature response of TPU derived in this study may prove useful in quantifying the biochemical limits to terrestrial plant photosynthesis and improve the representation of plant photosynthesis in TBMs.     

Heterogeneous inhibition of photosynthesis over the leaf surface of Rosa rubiginosa L. during water stress and abscisic acid treatment: induction of a metabolic component by limitation of CO2 diffusion

The contribution of changes in stomatal conductance and metabolism in determining heterogeneous photosynthesis inhibition during dehydration and abscisic acid (ABA) feeding was investigated using detached leaves of Rosa rubiginosa L. The steady-state and maximal rates of electron transport under a transient high CO₂ concentration were monitored using chlorophyll fluorescence imaging. The decrease in electron transport rate induced by dehydration and ABA treatment almost reverted to the control rate under transient high CO₂ availability. Therefore, inhibition of photosynthesis was mainly mediated through stomatal closure. However, since reversion was not complete, a metabolic inhibition was also identified as a decrease in the maximal electron transport rate driven by carboxylation. Under dehydration or ABA feeding, as under low ambient CO₂ treatment, in 21% or 0.4% O₂, the lower the steady-state electron transport was, the lower was the maximal electron transport rate during transient high CO₂ availability. We conclude that low CO₂ availability reduced the capacity of ribulose-1,5-bisphosphate carboxylase-oxygenase (Rubisco) to drive electron transport. The potential contribution of Rubisco deactivation mediated by stomatal closure is discussed. Received: 1 February 1999 / Accepted: 15 June 1999     

New application of Bacillus strains for optically pure l-lactic acid production: general overview and future prospects

Members of the genus Bacillus are considered to be both, among the best studied and most commonly used bacteria as well as the most still unexplored and the most wide-applicable potent bacteria because novel Bacillus strains are continuously being isolated and used in various areas. Production of optically pure l-lactic acid (l-LA), a feedstock for bioplastic synthesis, from renewable resources has recently attracted attention as a valuable application of Bacillus strains. l-LA fermentation by other producers, including lactic acid bacteria and Rhizopus strains (fungi) has already been addressed in several reviews. However, despite the advantages of l-LA fermentation by Bacillus strains, including its high growth rate, utilization of various carbon sources, tolerance to high temperature, and growth in simple nutritional conditions, it has not been reviewed. This review article discusses new findings on LA-producing Bacillus strains and compares them to other producers. The future prospects for LA-producing Bacillus strains are also discussed.     

Internal conductance does not scale with photosynthetic capacity: implications for carbon isotope discrimination and the economics of water and nitrogen use in photosynthesis

Central paradigms of ecophysiology are that there are recognizable and even explicit and predictable patterns among species, genera, and life forms in the economics of water and nitrogen use in photosynthesis and in carbon isotope discrimination (delta). However most previous examinations have implicitly assumed an infinite internal conductance (gi) and/or that internal conductance scales with the biochemical capacity for photosynthesis. Examination of published data for 54 species and a detailed examination for three well-characterized species--Eucalyptus globulus, Pseudotsuga menziesii and Phaseolus vulgaris--show these assumptions to be incorrect. The reduction in concentration of CO2 between the substomatal cavity (Ci) and the site of carbon fixation (Cc) varies greatly among species. Photosynthesis does not scale perfectly with gi and there is a general trend for plants with low gi to have a larger draw-down from Ci to Cc, further confounding efforts to scale photosynthesis and other attributes with gi. Variation in the gi-photosynthesis relationship contributes to variation in photosynthetic 'use' efficiency of N (PNUE) and water (WUE). Delta is an information-rich signal, but for many species only about two-thirds of this information relates to A/gs with the remaining one-third related to A/gi. Using data for three well-studied species we demonstrate that at common WUE, delta may vary by up to 3 per thousand. This is as large or larger than is commonly reported in many interspecific comparisons of delta, and adds to previous warnings about simplistic interpretations of WUE based on delta. A priority for future research should be elucidation of relationships between gi and gs and how these vary in response to environmental conditions (e.g. soil water, leaf-to-air vapour pressure deficit, temperature) and among species.