Photosynthetic and growth responses of C3 and C4 grasses to short- duration sunflecks and resultant consequences for their performance in understory environments

Low C4 grass species abundance in understory environments is thought to be as a result of their high-light requirements, lack of photosynthetic advantage relative to C3 species in cooler environments, and an inability to adequately utilise sunflecks. This study sets out to investigate this theory, hypothesizing that C3 grass species outperform C4 grass species under the canopy, not as a result of quantum efficiency temperature effects, but as a result of C4 species inability to utilize short-duration sunflecks. Short sunflecks could result in a breakdown in assimilate movement between the mesophyll (MSC) and bundle sheath (BSC) cells. The role of BSC leakiness, stomata and PSII efficiency on the ability of C3 and C4 Alloteropsis semialata to utilize short-duration sunflecks was investigated using gaseous exchange and chlorophyll fluorescence techniques, while the growth of both subspecies under a simulated flecking-light environment tested whether these measured responses translated into effects on growth. As C3 grasses are known to possess higher levels of stomatal conductance in relation to C4 species, results showed that C3 A. semialata was able to utilize short-duration sunflecks as a result of increased stomatal conductance and an ability to induce photosynthesis under various light flecking conditions. In contrast, C4 A. semialata was unable to utilize sunflecks possibly as a result of energetic limitations of the carboxylation mechanism (PSII) and not because of increased bundle sheath leakiness. These photosynthetic responses translated into growth differences when both types were grown in an artificially flecking light environment. The photosynthetic differences noted for C3 and C4 A.semialata were also evident in shade adapted understory grasses; Erharta erecta (C3), Dactylotenium australe (C4) and Brachiaria chusqueoides (C4). As photosynthetic induction was marginally more rapid in all species relative to A. semialata it suggests some degree of adaptation in shade grasses, however the inability of the C4 species to utilise short sunflecks remained. As a result, it was hypothesized that C3 shade adapted E. erecta and C4 B. chusqueoides, co-inhabiting the same forest understory, are able to do so because of differences in light micro-environments associated with each species. However, canopy openness and light profiles determined for theses micro-climates showed no differences and that both species have persisted within a relatively stable understory environment. The coexistence was possible as the understory was supplied with a significant proportion of its daily light in flecks sufficiently long as to not compromise C4 productivity. A survey of 10 species of grasses under various canopy densities and in the open showed a strong negative linear relationship between canopy openness and the rate at which photosynthesis was induced by flecking light, which has not been shown before. This did not result from the phylogenetic relationship between species and could be shown for a single species (E. erecta) growing in a range of light environments. This demonstrates that C4 grasses, despite adaptation, would be limited from sunfleck environments if a significant proportion of the daily light available consists of flecks of short-duration. The generation of such environments may occur as a result of woody thickening and could help explain the observed decline in C4 grasses under these conditions