Photosynthesis and associated signalling are influenced by the dorso-ventral properties of leaves. The degree of adaxial/abaxial symmetry in stomatal numbers, photosynthetic regulation with respect to light orientation and the total section areas of the bundle sheath (BS) cells and the surrounding mesophyll (M) cells on the adaxial and abaxial sides of the vascular bundles were compared in two C4 [Zea mays (maize) and Paspalum dilatatum] and one C3 [Triticum turgidum (Durum wheat)] monocotyledonous species. The C3 leaves had a higher degree of dorso-ventral symmetry than the C4 leaves. Photosynthetic regulation was the same on each side of the wheat leaves, as were stomatal numbers and the section area of the BS relative to that of the M cells (BS/M section area ratio). In contrast, photosynthetic regulation in maize and P. dilatatum leaves showed a marked surface-specific response to light orientation. Compared to the adaxial sides of the C4 monocotyledonous leaves, the abaxial surfaces had more stomata and the BS/M section area ratio was significantly higher. Differences in dorso-ventral structure, particularly in Kranz anatomy, serve not only to maximize photosynthetic capacity with respect light orientation in C4 monocotyledonous leaves but also allow adaxial and abaxial-specific signalling from the respective M cells.

The effects of dark chilling on the leaf-side-specific regulation of photosynthesis were characterized in the C4 grass Paspalum dilatatum. CO2- and light-response curves for photosynthesis and associated parameters were measured on whole leaves and on each leaf side independently under adaxial and abaxial illumination before and after plants were exposed to dark chilling for one or two consecutive nights. The stomata closed on the adaxial sides of the leaves under abaxial illumination and no CO2 uptake could be detected on this surface. However, high rates of whole leaf photosynthesis were still observed because CO2 assimilation rates were increased on the abaxial sides of the leaves under abaxial illumination. Under adaxial illumination both leaf surfaces contributed to the inhibition of whole leaf photosynthesis observed after one night of chilling. After two nights of chilling photosynthesis remained inhibited on the abaxial side of the leaf but the adaxial side had recovered, an effect related to increased maximal ribulose-1,5-bisphosphate carboxylation rates (Vcmax) and enhanced maximal electron transport rates (Jmax). Under abaxial illumination, whole leaf photosynthesis was decreased only after the second night of chilling. The chilling-dependent inhibition of photosynthesis was located largely on the abaxial side of the leaf and was related to decreased Vcmax and Jmax, but not to the maximal phosphoenolpyruvate carboxylase carboxylation rate (Vpmax). Each side of the leaf therefore exhibits a unique sensitivity to stress and recovery. Side-specific responses to stress are related to differences in the control of enzyme and photosynthetic electron transport activities.

Water deficit, when rapidly imposed on three C 4 grasses of the different metabolic subtypes, Paspalum dilatatum Poiret (NADP-malic enzyme), Cynodon dactylon (L.) Pers (NAD-malic enzyme) and Zoysia japonica Steudel (phosphoenolpyruvate carboxykinase), caused decreases in photosynthetic rates, in the quantum yield of PS II and photochemical quenching, and in the activities of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPC). The results provide evidence for non-stomatal limitations of photosynthesis differing in nature between the three species.Additional key words: chlorophyll a fluorescence; drought stress; NAD-malic enzyme; NADP-malic enzyme; phosphoenolpyruvate carboxykinase; phosphoenolpyruvate carboxylase; photosynthetic capacity; Rubisco.

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