exhibited a large growth advantage over the C 3 species at low [CO 2 ]. However, this advantage was reduced at low temperature, where the C 4 species produced 5× the total mass of the C 3 species versus 14× at the high temperature. This difference was due to a reduction in C 4 growth at low temperature, since the C 3 species exhibited similar growth between temperatures. Physiological differences between temperatures were not detected for either species, although photorespiration/net photosynthesis was reduced in the C 3 species grown at low temperature, suggesting evidence of improved carbon balance at this treatment. This system suggests that C 4 species had a growth advantage over C 3 species during low [ Studying plant responses to global changes of the past provides valuable insights for predicting future responses to a rapidly changing environment (Ward et al. 2005; Edwards et al. 2007; Jackson 2007). In addition, studies involving treatments that simulate past climates provide a baseline for understanding the physiological functioning of plants prior to anthropogenic influences (Polley et al. 1993a,b; Anderson et al. 2001 In addition to physiological models, empirical studies examining the growth and development of C 3 and C 4 species at reduced temperatures and low [CO 2 ] are also needed to better Table 1. The effects of modern and glacial temperatures on total mass (n = 13-14), leaf area (LA, n = 13-14), stomatal conductance (g s , n = 9-12), and specific leaf mass (SLM, n = 13-14) for Abutilon theophrasti (C 3 ) and Amaranthus retroflexus (C 4 ) grown at low CO 2 (200 μmol/mol) for 22 d Values are means ± 1 SE. Data from different chambers within the same treatment were combined because a chamber effect was not detected.Different superscript letters between temperatures and species indicate significant differences at the P < 0.05 level according to ANOVA.
ResultsOn an absolute basis, the C 3 (Abutilon) and C 4 species (Amaranthus) exhibited large differences in total mass when grown at low [CO 2 ] (200 μmol/mol) for 22 d. More specifically, the C 4 species had five times the total mass of the C 3 species when grown at the low temperature (22 light/16 dark • C), and almost 14 times the total mass of the C 3 species when grown at the high temperature (30/24 • C; Table 1). In this case, all plants were grown from seed for a total of 22 d, and therefore differences in final biomass mainly reflect differences in relative growth rate (change in biomass per unit biomass per unit time; hereafter RGR). Therefore, the RGR of the C 4 species greatly exceeded that of the C 3 species at low [CO 2 ]. In addition, Amaranthus has inherently small seeds compared with Abutilon, and therefore initial seed size would not have been a factor in producing a relative growth advantage of the C 4 species over the C 3 species when grown at low [CO 2 ]. Furthermore, the C 3 and C 4 species exhibited relative differences in their responses to temperature for total mass and LA (leaf area) (significant species X temperature inte...