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.
Daily variations in net gas exchange, chlorophyll a fluorescence and water relations of mature, sun-acclimated grapefruit (Citrus paradisi Macfady.) and orange (Citrus sinensis L. Osbeck) leaves were determined in tree canopies either shaded with 50% shade screens or left unshaded (sunlit). Mean daily maximum photosynthetic photon flux density (PPFD) under shade varied from 500 to 700 micromol m-2 s-1 and was sufficient to achieve maximum net CO2 assimilation rates (A CO2). Responses of grapefruit and orange leaves to shading were remarkably similar. At midday, on bright clear days, the temperatures of sunlit leaves were 2-6 degrees C above air temperature and 1-4 degrees C above the temperatures of shaded leaves. Although midday depressions of stomatal conductance (gs) and A CO2 were observed in both sunlit and shaded leaves, shaded leaves had lower leaf-to-air vapor pressure differences (D) along with higher gs, A CO2 and leaf water-use efficiency than sunlit leaves. Estimated stomatal limitation to A CO2 was generally less than 25% and did not differ between shaded and sunlit leaves. Leaf intercellular CO2 partial pressure was not altered by shade treatment and did not change substantially with increasing D. Radiation and high temperature stress-induced non-stomatal limitation to A CO2 in sunlit leaves was greater than 40%. Reversible photoinhibition of photosystem II efficiency was more pronounced in sunlit than in shaded leaves. Thus, non-stomatal factors play a major role in regulating A CO2 of citrus leaves during radiation and high temperature stress.
Leaf water relations, net gas exchange and leaf and root constituent responses to 9 days of drought stress (DS) or soil flooding were studied in 6‐month‐old seedlings of Carrizo citrange [Citrus sinensis (L.) Osb. ×Poncirus trifoliata L.; Carr] and Cleopatra mandarin (Citrus resnhi Hort. ex Tanaka; Cleo) growing in containers of native sand in the greenhouse. At the end of the drought period, both species had similar minimum stem water potentials but Cleo had higher leaf relative water content (RWC) and higher leaf osmotic potential at full turgor () than Carr. Flooding had no effect on RWC but osmotic adjustment (OA) and were higher in Cleo than in Carr. Net CO2 assimilation rate (ACO2) in leaves was decreased more by drought than by flooding in both species but especially in Carr. Leaf water‐use efficiency (ACO2/transpiration) was lower in Carr and was decreased more by DS and flooding stress than in Cleo. Higher values of intercellular CO2 concentration (Ci) in stressed plants than in control plants indicated that non‐stomatal factors including chlorophyll degradation and chlorophyll fluorescence [maximum quantum efficiency of PSII (Fv/Fm, where Fm is the maximum fluorescence and F0, minimum fluorescence in dark‐adapted leaves)] were more important limitations on ACO2 than stomatal conductance. In both genotypes, leaf proline was increased by drought but not by flooding, whereas both stresses increased proline in roots. Soluble sugars in leaves were increased by DS, and flooding decreased leaf sugars in Cleo. In general, DS tended to increase the concentrations of Ca, K, Mg, Na and Cl in both leaves and roots, whereas flooding tended to decrease these ions with the exception of leaf Ca in Cleo. Based on water relations and net gas exchange, Cleo was more tolerant to short‐term DS and flooding stress than Carr.
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