ALBERT

Notes: A model has been derived for the enrichment of heavy isotopes of water in leaves, including progressive enrichment along the leaf. In the model, lighter water is preferentially transpired leaving heavier water to diffuse back into the xylem and be carried further along the leaf. For this pattern to be pronounced, the ratio of advection to diffusion (Péclet number) has to be large in the longitudinal direction, and small in the radial direction. The progressive enrichment along the xylem is less than that occurring at the sites of evaporation in the mesophyll, depending on the isolation afforded by the radial Péclet number. There is an upper bound on enrichment, and effects of ground tissue associated with major veins are included. When transpiration rate is spatially nonuniform, averaging of enrichment occurs more naturally with transpiration weighting than with area-based weighting. This gives zero average enrichment of transpired water, the modified Craig–Gordon equation for average enrichment at the sites of evaporation and the Farquhar and Lloyd (In Stable Isotopes and Plant Carbon-Water Relations, pp. 47–70. Academic Press, New York, USA, 1993) prediction for mesophyll water. Earlier results on the isotopic composition of evolved oxygen and of retro-diffused carbon dioxide are preserved if these processes vary in parallel with transpiration rate. Parallel variation should be indicated approximately by uniform carbon isotope discrimination across the leaf.

Notes: A decrease in steady-state leaf transpiration rate with increased vapour pressure difference between leaf and air, which is reversible and independent of leaf water status, is evidence for feedforward control of stomatal aperture (Cowan 1977). A recent survey of gas exchange data by Monteith (1995), covering 52 sets of measurements on 16 species, reported that evidence for feedforward control was rare and usually reliant on a single point. We conducted gas exchange experiments on an additional 13 species and observed an apparent feedforward response in only two. However, the response was not reversible and depended upon experimental procedure. In view of this we discuss the appropriate use of the term ‘feedforward’.

Notes: Concurrent measurements of leaf gas exchange and on-line 13C discrimination were used to evaluate the CO2 conductance to diffusion from the stomatal cavity to the sites of carboxylation within the chloroplast (internal conductance; gi). When photon irradiance was varied it appeared that gi and/or the discrimination accompanying carboxylation also varied. Despite this problem, gi, was estimated for leaves of peach (Prunus persica), grapefruit (Citrus paradisi), lemon (C. limon) and macadamia (Macadamia integrifolia) at saturating photon irradiance. Estimates for leaves of C. paradisi, C. limon and M. integrifolia were considerably lower than those previously reported for well-nourished herbaceous plants and ranged from 1.1 to2.2μmol CO2 m−2 s−1 Pa−1, whilst P. persica had a mean value of 3.5 μmol CO2 m−2 s−1 Pa−1. At an ambient CO2 partial pressure of 33Pa, estimates of chloroplastic partial pressure of CO2 (Cc) using measurements of CO2 assimilation rate (A) and calculated values of gi, and of partial pressure of CO2 in the stomatal cavity (Cst) were as low as 11.2 Pa for C. limon and as high as 17.8Pa for peach. In vivo maximum rubisco activities (Vmax) were also determined from estimates of Cc. This calculation showed that for a given leaf nitrogen concentration (area basis) C. paradisi and C. limon leaves had a lower Vmax than P. persica, with C. paradisi and C. limon estimated to have only 10% of leaf nitrogen present as rubisco. Therefore, low CO2 assimilation rates despite high leaf nitrogen concentrations in leaves of the evergreen species examined were explained not only by a low Cc but also by a relatively low proportion of leaf nitrogen being used for photosynthesis. We also show that simple one-dimensional equations describing the relationship between leaf internal conductance from stomatal cavities to the sites of carboxylation and carbon isotope discrimination (Δ) can lead to errors in the estimate of gi. Potential effects of heterogeneity in stomatal aperture on carbon isotope discrimination may be particularly important and may lead to a dependence of gi upon CO2 assimilation rate. It is shown that for any concurrent measurement of A and Δ, the estimate of Cc is an overestimate of the correct photosynthetic capacity-weighted value, but this error is probably less than 1.0 Pa.

Notes: Abstract A negative correlation between water-use efficiency (W), defined as the ratio of moles of carbon in the plant to moles of water transpired, and carbon isotope discrimination (Δ) was established for barley in pot experiments using 12 cultivars. The correlation was strong in two independent experiments in four different controlled environment where ambient temperature and vapour pressure deficit were varied and plants were either well-watered or given limited amounts of water. Variation among cultivars was found in both Δ and W and rankings of both parameters, according to cultivar, were similar in different environments. Limiting water usually increased water-use efficiency of plants. Total dry matter can be substituted for moles of carbon when calculating water-use efficiency but the correlation between W and Δ were calculated using the carbon content of dry matter. There were differences varied significantly among cultivars. Despite these differences, correlations were also large between whole plant W and Δ of any of the plant parts. The amount of dry matter partitioned into reproductive growth varied genetically, as did the effect of stress on the partitioning. Growth, W and Δ of barley were compared with theory derived from gas exchange properties and with other literature. The effect on W of variation in vapour pressure deficit in these experiments was removed by multiplying W by vapour pressure deficit to derive the parameter, k(Pa mol C/mol H2O). This allowed comparisons among experiments with different vapour pressure deficits. The mean k for these barley cultivars was similar to that calculated by others for grasses. However, variation was found, and, in contrast with previous work which treats k as a species constant, we conclude that there is promise in selecting for increased k.

Notes: A new analytical model for the response of whole-leaf potential electron transport rate (J) to light is presented. The model treats incident irradiance at the upper and lower leaf surfaces independently, describes transdermal profiles of light absorption and electron transport capacity explicitly, and calculates J by integrating the minimum of light- and capacity-limited rates among paradermal chlorophyll layers. The capacity profile is assumed to be a weighted average of two opposed exponential profiles, each of which corresponds to the profile of light-limited rate when only one surface is illuminated; the weights may take on any values, provided they sum to unity, so the model can describe leaves with a wide range of ‘preferred’ illumination regimes. By treating irradiance at either surface independently and assuming the capacity profile is fixed on short time scales, the model predicts observed effects of leaf inversion on light-response curves and their apparent convexity. By assuming the capacity profile can adapt on developmental time scales, the model can predict the observed dependence of inversion effects on the growth lighting regime. It is suggested that the new model, which is mathematically compact and formally similar to the standard non-rectangular hyperbola model for J, be used in place of the standard model in studies in which the effects of leaf angle or diffuse light fraction on gas exchange are of interest.

Notes: The effectiveness of several leaf water models (‘string-of-lakes’, ‘desert river’ and the Farquhar–Gan model) are evaluated in predicting the enrichment of leaf water along a maize leaf at different humidities. Progressive enrichment of both vein xylem water and leaf water was observed along the blade. At the tip, the maximum observed enrichment for the vein water was 17.6‰ at 50% relative humidity (RH) whereas that for the leaf water was 50‰ at 34% RH and 19‰ at 75% RH. The observed leaf water maximum was a fraction (0.5–0.6) of the theoretically possible maximum. The ‘string-of-lakes’ and ‘desert river’ models predict well the variation of leaf water enrichment pattern with humidity but overestimate the average enrichment of bulk leaf water. However, the Farquhar–Gan model gives good prediction for these two aspects of leaf water enrichment. Using the anatomical dimensions of vein xylem overestimates the effective longitudinal Péclet number (Pl). Possible explanations for this discrepancy between the effective and the xylem-based estimate of Pl are discussed. The need to characterize the heterogeneity of transpiration rate over the leaf surface in studies of leaf water enrichment is emphasized. The possibility that past atmospheric humidity can be predicted from the slope of the Δ18O spatial variation of leaf macrofossils found in middens is proposed.

Notes: A mathematical model of stomatal conductance is presented. It is based on whole-plant and epidermal hydromechanics, and on two hypotheses: (1) the osmotic gradient across guard cell membranes is proportional to the concentration of ATP in the guard cells; and (2) the osmotic gradient that can be sustained per unit of ATP is proportional to the turgor pressure of adjacent epidermal cells. In the present study, guard cell [ATP] is calculated using a previously published model that is based on a widely used biochemical model of C3 mesophyll photosynthesis. The conductance model for Vicia faba L. is parameterized and tested As with most other stomatal models, the present model correctly predicts the stomatal responses to variations in transpiration rate, irradiance and intercellular CO2. Unlike most other models, however, this model can predict the transient stomatal opening often observed before conductance declines in response to decreases in humidity, soil water potential, or xylem conductance. The model also explicitly accommodates the mechanical advantage of the epidermis and correctly predicts that stomata are relatively insensitive to the ambient partial pressure of oxygen, as a result of the assumed dependence on ATP concentration.

Notes: Gas exchange experiments were performed with 13 plant species that differ from each other in growth-form and natural habitat. These comprised three herbaceous species, two ferns, two temperate deciduous trees, five rainforest trees and one liana from wet tropical forest. The aims were to investigate whether plants of similar growth-form and from similar habitats tended to respond similarly to a change in leaf-to-air vapour pressure difference (VPD), and to compare their ratio of intercellular to ambient partial pressures of CO2 for given conditions. Leaves were subjected to a step change in VPD and the initial and final steady rates of transpiration were used to calculate an index of sensitivity, φ, which enabled comparison of species. The results suggest that species of similar growth-form and habitat respond similarly to increasing VPD, with the temperate deciduous trees undergoing a greater reduction in stomatal conductance than the herbaceous plants in well-watered soil. Also, for these experimental conditions, the ratio of leaf internal to ambient CO2 partial pressure (pi/pa) was positively correlated with both CO2 assimilation rate and stomatal insensitivity to VPD, across the 13 species. The results are discussed in terms of growth strategies and possible advantages and limitations of hydraulic systems in different plants.