Ecophysiology of first and second generation hybrids in a natural plant hybrid zone

Abstract
Hybrids between related species vary widely in relative fitness, and that fitness can depend upon the environment. We investigated aspects of physiology that might influence fitness patterns in a plant hybrid zone. Seeds of Ipomopsis aggregata, I. tenuituba, F1 hybrids, F2 hybrids, and offspring of crosses between natural hybrids were planted into the relatively mesic site of origin for I. aggregata and the drier site for natural hybrids. We measured rates of photosynthesis (A max), transpiration (E), instantaneous (A/E) and long-term (δ13C) indices of water use efficiency (WUE), and leaf nitrogen and carbon. We also examined correlations of these traits with plant size. Photosynthetic rate and A/E were higher in vegetative than flowering plants. WUE varied between sites and years, but differences among genotypic classes were spatially and temporally consistent. Instantaneous WUE was higher for F1 hybrids than for the average of the parental species, thereby showing heterosis. There was no evidence of hybrid breakdown, as WUE was no different in the F2 than the average across the F1 and parental species. Nor did WUE depend on cross direction in producing F1 progeny. Carbon isotope discrimination revealed higher long-term water use efficiency in I. tenuituba than I. aggregata. Leaf nitrogen was higher in I. tenuituba than I. aggregata, and higher in offspring of natural hybrids than in the F2. Results indicate heterosis for water use efficiency, with no hybrid breakdown. Heterosis in WUE may help to explain the relatively high survival of both reciprocal F1 hybrids in dry sites within the natural hybrid zone.