Abstract. In C4 grasses belonging to the NADP-malic enzyme-type subgroup, malate is considered to be the predominant C4 acid metabolized during C4 photosynthesis, and the bundle sheath cell chloroplasts contain very little photosystem-II (PSII) activity. The present studies showed that Flaveria bidentis (L.), an NADP-malic enzyme-type C4 dicotyledon, had substantial PSII activity in bundle sheath cells and that malate and aspartate apparently contributed about equally to the transfer of CO2 to bundle sheath cells. Preparations of bundle sheath cells and chloroplasts isolated from these cells evolved O2 at rates between 1.5 and 2 gmol'min -1.mg 1 chlorophyll (Chl) in the light in response to adding either 3-phosphoglycerate plus HCO~-or aspartate plus 2-oxoglutarate. Rates of more than 2 ~tmol 02' min -1" mgChl were recorded for cells provided with both sets of these substrates. With bundle sheath cell preparations the maximum rates of light-dependent CO2 fixation and malate decarboxylation to pyruvate recorded were about 1.7 ~tmol -min-1. mg-1 Chl. Compared with NADPmalic enzyme-type grass species, F. bidends bundle sheath cells contained much higher activities of NADP-malate dehydrogenase and of aspartate and alanine aminotransferases. Time-course and pulse-chase studies following the kinetics of radiolabelling of the C-4 carboxyl of C4 acids from t4CO2 indicated that the photosynthetically active pool of malate was about twice the size of the aspartate pool. However, there was strong evidence for a rapid flux of carbon through both these pools. Possible routes of aspartate metabolism and the relationship between this metabolism and PSII activity in bundle sheath cells are considered.Abbreviations: DHAP = dihydroxyacetone phosphate; NADP-ME(-type) = NADP-malic enzyme (type); NADP-MDH = NADPmalate dehydrogenase; OAA = oxaloacetic acid; 2-OG = 2-oxoglutarate; PEP = phosphoenolpyruvate; PGA = 3-phosphoglycerate; Pi = orthophosphate; Ru5P = ribulose 5-phosphate Correspondence to: M.D. Hatch; FAX:61 (6)2465000; Tel.: 61 (6) 2465264

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