No abstract

Internal conductances to CO2 transfer from the stomatal cavity to sites of carboxylation (gi) in hypostomatous sun‐and shade‐grown leaves of citrus, peach and Macadamia trees (Lloyd et al. 1992) were related to anatomical characteristics of mesophyll tissues. There was a consistent relationship between absorptance of photosynthetically active radiation and chlorophyll concentration (mmol m−2) for all leaves, including sclerophyllous Macadamia, whose transmittance was high despite its relatively thick leaves. In thin peach leaves, which had high gi, the chloro‐plast volume and mesophyll surface area exposed to intercellular air spaces (ias) per unit leaf area were similar to those in the thicker leaves of the evergreen species. Peach leaves, however, had the lowest leaf dry weight per area (D/a), the lowest tissue density (Td) and the highest chloro‐plast surface area (Sc) exposed to ias. There were negative correlations between gi and leaf thickness or D/a, but positive correlations between gi and Sc or Sc/Td. We developed a one‐dimensional diffusion model which partitioned gi into a gaseous diffusion conductance through the ias (gias) plus a liquid‐phase conductance through mesophyll cell walls (gcw). The model accounted for a significant amount of variation (r2=0.80) in measured gi by incorporating both components. The gias component was related to the one‐dimensional path‐length for diffusion across the mesophyll and so was greater in thinner peach leaves than in leaves of evergreen species. The gcw component was related to tissue density and to the degree of chloroplast exposure to the ias. Thus the negative correlations between gi and leaf thickness or D/a related to gias whereas positive correlations between gi and Sc or Sc/Td, related to gcw. The gcw was consistently lower than gias, and thus represented a greater constraint on CO2 diffusion in the mesophylls of these hypostomatous species.

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.