perature response of Rubisco and A, we propose that Rubisco is a major limitation on C 4 photosynthesis in B. gracilis below 17°C. Based on these results and for theoretical reasons associated with how C 4 plants use Rubisco, we further suggest that Rubisco capacity may be a widespread limitation upon C 4 photosynthesis at low temperature.Key-words: Bouteloua; C4 photosynthesis; gas exchange; photosynthesis; Rubisco activity; sub-alpine plants; temperature responses.
INTRODUCTIONC 4 plants are common at low latitude and altitude, but are rare to absent at latitudes and altitudes having cool temperate or boreal climates (Long 1983;Collatz et al. 1998). Where growing season temperatures average less than 13°C, C 4 plants are rare; where they average above 22°C, C 4 plants are typically dominant members of herbaceous plant communities (Long 1983;Sage et al. 1999). In temperate grasslands where C 3 and C 4 grass species co-exist, the C 3 to C 4 biomass composition of the vegetation is correlated with seasonal temperature. C 3 plants have superior growth during cool springs, whereas C 4 grasses dominate during mid-summer (Kemp & Williams 1980; Monson et al. 1983). Despite this understanding of the geographic and seasonal distribution of C 4 plants, the underlying mechanisms controlling these trends are not clearly understood. In particular, is the lack of C 4 plants in cooler climates the result of an inherent limitation in the C 4 pathway, or is it a merely a reflection of recent C 4 origin in tropical climates? Numerous hypotheses have been proposed to explain the absence of C 4 species in low-temperature habitats (for reviews see Long 1983;Ehleringer & Monson 1993;Leegood & Edwards 1996). The leading explanations can be grouped into three general categories.(1) There is a loss of adaptive value of the C 4 syndrome at low temperature simply because it is less efficient than C 3 photosynthesis; no injury or enzyme dissociation explains C 4 failure.
ABSTRACTThe mechanisms controlling the photosynthetic performance of C 4 plants at low temperature were investigated using ecotypes of Bouteloua gracilis Lag. from high (3000 m) and low (1500 m) elevation sites in the Rocky Mountains of Colorado. Plants were grown in controlledenvironment cabinets at a photon flux density of 700 mmol m -2 s -1 and day/night temperatures of 26/16°C or 14/7°C. The thermal response of the net CO 2 assimilation rate (A) was evaluated using leaf gas-exchange analysis and activity assays of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), phosphoenolpyruvate carboxylase (PEPCase) and pyruvate,orthophosphate dikinase (PPDK). In both ecotypes, a reduction in measurement temperature caused the CO 2 -saturated rate of photosynthesis to decline to a greater degree than the initial slope of A versus the intercellular CO 2 response, thereby reducing the photosynthetic CO 2 saturation point. As a consequence, A in normal air was CO 2 -saturated at suboptimal temperatures. Ecotypic variation was low when grown at 26/16°C, with the major diffe...