Estimation of diffusive resistance of bundle sheath cells to CO2 from modeling of C 4 photosynthesis

Bundle sheath resistance to diffusion of CO2 (rc) is a critical component of C4 photosynthesis which allows accumulation of inorganic carbon in bundle sheath cells of C4 plants. Several analyses were made to evaluate the magnitude of rc in C4 plants. Experimental data on the O2 inhibition of photosynthesis (Dai et al. (1993) Plant Physiol 103: 83-90; (1995) Plant Physiol 107: 815-825) and rates of photorespiration (de Veau and Burris (1989) Plant Physiol 90: 500-511) in Z. mays at different stages of development were analyzed using mathematical models of C4 photosynthesis. In young and senescing leaves modeled values of rc and the CO2 partial pressure in bundle sheath cells (Cbs) were lower and fractional leakage of CO2 from bundle sheath cells (fL) was higher than in mature leaves. Diffusive resistance of bundle sheath cells of C4 plants was also evaluated by analyzing the response of photosynthetic rates to varying CO2 in Amaranthus edulis in which the C4 cycle was dysfunctional by chemical mutagenesis (Dever et al. (1995) J Exp Bot 46: 1363-1376) and in Sorghum bicolor, Panicum maximum and Panicum miliaceum in which the C4 cycle was chemically inhibited (Brown and Byrd (1993) Plant Physiol 103: 1183-1188). These analyses indicate that in mature leaves of C4 plants the values of rc are substantially lower (ca. 50-200 m(2) s mol(-1)) than previous suggested (ca. 500-1500 m(2) s mol(-1)) for C4 photosynthesis and that there is considerable leakage of CO2 from bundle sheath cells. Nevertheless, rc and Cbs values are sufficiently high in mature leaves to minimize photorespiration in C4 plants under normal levels of CO2.