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Thus, evolution of this biochemical system is expected to repeat itself qualitatively in different species. Simulated evolutionary trajectories indeed cluster narrowly around a ��mean path�� (p?this website some of these species were previously classified as either C3 or C4 based on other criteria ( McKown et?al., 2005). Each of the intermediate species constitutes a separate point on evolutionary trajectories that started at C3 biochemistry. We collected experimental estimates of the biochemical model parameters for each of the 20 species from the literature, and we extended this data set by experimentally determining Vpmax and �� for CB-5083 nmr several Flaveria species ( Experimental Procedures). With few exceptions, the experimentally determined parameter sets indeed lie very close to the predicted mean path through the fitness landscape ( Figure?5). The model predicts experimental parameter combinations much better than a null model assuming a random order of evolutionary changes ( Figure?6; p?FXR from diverse C3-C4 intermediates confirms the model��s ability to quantitatively predict biochemical evolution over a timescale of several million years ( Sage et?al., 2012). While the majority of the data describe the genus Flaveria, the model also correctly predicts data from two phylogenetically distant genera ( Figure?5). Comparisons to additional C3-C4 intermediates are currently limited by the availability of species-specific protocols for the separation of BS and M cells. A hypothesis for the evolutionary succession of biochemical and morphological changes in the evolution of the C4 syndrome was previously derived from phylogenetically informed analyses of C3-C4 intermediates (Sage et?al., 2012). This hypothesis assumes modular biochemical changes, starting with a shift of photorespiration to the BS, followed by the establishment of a C4 cycle in conjunction with a shift of RuBisCO to the BS, and finally an optimization stage in which parameters are fine-tuned.
