Spontaneous homeotic transformations have been described in natural populations of both plants and animals, but little is known about the molecular-genetic mechanisms underlying these processes in plants. In the ABC model of floral organ identity in Arabidopsis thaliana, the B-and C-functions are necessary for stamen morphogenesis, and C alone is required for carpel identity. We provide ABC model-based molecular-genetic evidence that explains the unique inside-out homeotic floral organ arrangement of the monocotyledonous mycoheterotroph species Lacandonia schismatica (Triuridaceae) from Mexico. Whereas a quarter million flowering plant species bear central carpels surrounded by stamens, L. schismatica stamens occur in the center of the flower and are surrounded by carpels. The simplest explanation for this is that the B-function is displaced toward the flower center. Our analyses of the spatio-temporal pattern of B-and C-function gene expression are consistent with this hypothesis. The hypothesis is further supported by conservation between the B-function genes of L. schismatica and Arabidopsis, as the former are able to rescue stamens in Arabidopsis transgenic complementation lines, and Ls-AP3 and Ls-PI are able to interact with each other and with the corresponding Arabidopsis B-function proteins in yeast. Thus, relatively simple molecular modifications may underlie important morphological shifts in natural populations of extant plant taxa.
INTRODUCTION An ABC Model-Based Hypothesis of the Developmental Genetic Factors Underlying the Unusual Reproductive Morphology of Lacandonia schismaticaThe ABC model for the specification of floral organ identity Coen and Meyerowitz, 1991;Meyerowitz et al., 1991) has played a critical role in the modern explanation of the molecular-genetic determinants of the ontogenetic development of reproductive structures in angiosperms. This combinatorial genetic model has guided diverse plant evolutionary developmental biology studies, especially during the formative years of this young field (Cronk, 2001;Cronk et al., 2002;Pruitt et al., 2003). As part of the larger discipline of evo-devo, the articulation of explanatory tools like the ABC model is considered essential for understanding the developmental mechanisms that underlie morphological innovation throughout evolutionary time (Cronk et al., 2002;Carroll et al., 2004;Gilbert, 2006;Wolpert et al., 2006).The ABC model of flower development was based on the interpretation of floral homeotic phenotypes in Arabidopsis thaliana and Antirrhinum majus (Coen and Meyerowitz, 1991). However, the participation of homeotic transformations and other forms of heterotopy in the appearance of new organ arrangements during the evolution of angiosperms had already been considered in the botanical literature, long before the age of plant developmental genetics (e.g., Meyer, 1966;Sattler, 1984;Bowman et al., 1989;Weston, 2000). Outstanding instances of spontaneous floral homeotic phenotypes have continued to be recorded in well-characterized taxa (s...