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Reeves, P. A. and R. G. Olmstead. 1998

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Reeves, P. A. and R. G. Olmstead. 1998 American Journal of Botany 85(9): 1047±1056. 1998. EVOLUTION OF NOVEL MORPHOLOGICAL AND REPRODUCTIVE TRAITS IN A CLADE CONTAINING ANTIRRHINUM MAJUS (SCROPHULARIACEAE)1 PATRICK A. REEVES AND RICHARD G. OLMSTEAD2 Department of Botany, University of Washington, Seattle, Washington 98195 Phylogenetic analysis of DNA sequences of the chloroplast genes rbcL and ndhF revealed a highly supported clade composed of the families Plantaginaceae, Callitrichaceae, and Hippuridaceae in close association with the model organism Antirrhinum majus and other members of family Scrophulariaceae. Plantago has miniature actinomorphic wind-pollinated ¯owers that have evolved from zygomorphic animal-pollinated precursors. The aquatic Hippuridaceae have reduced wind- pollinated ¯owers with one reproductive organ per whorl, and three, rather than four, whorls. In monoecious aquatic Cal- litrichaceae, further reduction has occurred such that there is only one whorl per ¯ower containing a single stamen or carpel. Optimization of character states showed that these families descended from an ancestor similar to Antirrhinum majus. Recent studies of plant developmental genetics have focused on distantly related species. Differences in the molecular mechanisms controlling ¯oral development between model organisms are dif®cult to interpret due to phylogenetic distance. In order to understand evolutionary changes in ¯oral morphology in terms of their underlying genetic processes, closely related species exhibiting morphological variation should be examined. Studies of genes that regulate morphogenesis in the clade described here could aid in the elucidation of a general model for such fundamental issues as how changes in ¯oral symmetry, organ number, and whorl number are achieved, as well as providing insight on the evolution of dicliny and associated changes in pollination syndrome. Key words: Antirrhinum; development; evolution; ¯ower morphology; MADS box genes; ndhF; phylogeny; rbcL; Scrophulariaceae. One of the fundamental questions in evolutionary bi- angiosperm groups. In recent years, plant developmental ology is the origin of the novel characters that distinguish genetics has made great advances in our knowledge of species. Any genetic change that affects morphology mechanisms important in ¯oral ontogeny. Much of this must do so by affecting the interactive and contingent research has been focused on a few model systems, most processes of development. For a nascent morphological notably Arabidopsis thaliana (Brassicaceae) and Antir- character to become associated with a discrete evolution- rhinum majus (Scrophulariaceae). ary lineage (e.g., a species or higher taxon), the character Antirrhinum majus L., the snapdragon, is a common must pass through the sieve of selection and the process ornamental species characterized by showy zygomorphic of speciation, emerging as part of an integrated, success- ¯owers arranged in racemes. Its history as a research or- fully adapted organism. ganism for genetics began with studies on the inheritance Thus, in order to begin to understand how morpholog- of ¯ower color variation (Wheldale, 1907) and the char- ical character differences between taxa have arisen, it is acterization of numerous mutants by Baur (1930). Early necessary to have an understanding of the character from genetic analyses culminated with Stubbe's Genetik und both a phylogenetic and ontogenetic perspective. Phylo- Zytologie von Antirrhinum (1966). The current status of genetic trees are useful in determining homologous struc- Antirrhinum majus as a model organism for molecular tures and processes and in revealing the direction of evo- genetics was spurred by the discovery and characteriza- lution between character states. Through the comparison tion of several highly mobile transposable elements (Bon- of homologous developmental processes at the molecular as et al., 1984; Sommer et al., 1985; Upadhyaya et al., level, a mechanistic, as opposed to purely descriptive, 1985; Coen and Carpenter, 1986), which were useful for explanation for observed morphological differences may mutagenesis and gene-tagging experiments (Martin et al., be discovered. 1985; Sommer and Saedler, 1986; Martin et al., 1991). In plant evolutionary biology, molecular systematics Subsequent studies of ¯oral homeotic mutants led to the has increased our understanding of phylogeny in many identi®cation and characterization of a group of transcrip- tion factors, the MADS box gene family, which play a regulatory role during ¯oral morphogenesis, most notably 1 Manuscript received 10 June 1997; revision accepted 4 December 1997. in the speci®cation of organ identity (Carpenter and The authors thank Tom Philbrick for providing plant tissue of Hip- Coen, 1990; Schwarz-Sommer et al., 1990, 1992). puris and DNA of Callitriche, and Tom Philbrick, Jackie Nugent, Mi- Studies of ¯oral development in Arabidopsis thaliana chael Frohlich, and Peter Endress for many helpful suggestions on the (Bowman, Smyth, and Meyerowitz, 1989, 1991; Yanof- manuscript. This work was supported in part by a National Science sky et al., 1990) and later examinations in other angio- Foundation Undergraduate Research Experience (REU) supplement sperms (Pnueli et al., 1991; Angenent et al., 1992; Han- grant to Richard Olmstead (NSF grant number BSR-9107827) and a grant from the Undergraduate Research Opportunities Program at the sen et al., 1993; Schmidt et al., 1993) con®rmed the pres- University of Colorado to Patrick Reeves. ence and characteristic expression patterns of a few genes 2 Author for correspondence. in the MADS family, indicating that the fundamental ge- 1047 1048 AMERICAN JOURNAL OF BOTANY [Vol. 85 C class PLENA promotes termination in the fourth whorl (TroÈbner et al., 1992; Davies and Schwarz-Sommer, 1994). As a consequence of their role in meristem deter- minancy, MADS box genes may be thought of as in¯u- encing the number of whorls within a ¯ower. Moreover, genes from all three classes have been shown to in¯uence the number of organs that develop within a whorl (Bow- man, Smyth, and Meyerowitz, 1991; Coen and Meyero- witz, 1991). Several other genes, unrelated to the MADS box fam- ily, function in shaping the angiosperm ¯ower. Antirrhi- num majus has proven to be a valuable system for the study of genes involved in the development of zygomor- phic ¯owers. Mutations in the genes CYCLOIDEA and DICHOTOMA, which in wild type are expressed only in the dorsal region of the ¯ower, result in the development of actinomorphic Antirrhinum majus ¯owers (Luo et al., Fig. 1. Model illustrating the pattern of expression of key devel- 1996). Additional evidence from the gene DIVARICATA, opmental genes during early ¯ower development in wild-type Antirrhi- which in¯uences the development of features unique to num and Arabidopsis (adapted from Coen and Meyerowitz, 1991). Of the ventral region of the ¯ower, leads to the suggestion these, all but APETALA-2 (AP2) belong to the MADS box gene family. that asymmetry is achieved through the differential ex- There are three distinct gene functions, each given a class designation: A, B, or C. Each gene is expressed in two consecutive whorls. In wild- pression of such genes along the dorsoventral axis of the type developing ¯oral meristems, expression of the B and C class genes developing ¯ower (Almeida, Rocheta and Galego, 1997). as shown is necessary and suf®cient for determining the development In Arabidopsis, numerous genes affect the number of of the appropriate ¯oral organs in whorls 3 and 4 (Yanofsky, 1995). organs that develop within a whorl, but often do so in a The A function has not been clearly characterized in Antirrhinum (Da- stochastic manner (Bowman, Smyth, and Meyerowitz, vies and Schwarz-Sommer, 1994). Phylogenetic analysis suggests that 1989; Roe et al., 1993; Running and Meyerowitz, 1996; DEFICIENS (DEFA) and APETALA-3 (AP3), GLOBOSA (GLO) and PISTILLATA (PI), and PLENA and AGAMOUS (AG) are orthologous Huang and Ma, 1997). In the CLAVATA class of genes, gene pairs (Doyle, 1994). an increase in organ and whorl number is correlated with an increase in ¯oral meristem size (Clark, Running, and Meyerowitz, 1993). This suggests a possible biomechan- netic processes governing ¯oral organ differentiation ical in¯uence on the determination of organ and whorl have been conserved over evolutionary time (Fig. 1). number (Green, 1992). While it is unclear precisely how the whorl-speci®c There is considerable interest in possible roles for expression patterns of MADS box genes are de®ned, MADS box and other developmental genes in the evo- some mechanisms have been suggested. Antirrhinum ma- lution of ¯oral diversity (Coen and Meyerowitz, 1991; jus DEFICIENS protein can move from inner to outer Coen and Nugent, 1994; Irish and Yamamoto, 1995). layers in whorl 2 primordia, but not vice versa, and is Changes in in¯orescence architecture, modi®cation of the limited in the degree to which it can move within one basic four-whorled ¯oral ground plan, the appearance of cell layer. Thus control over the movement of MADS box differences between organs within a single whorl, and proteins through plasmodesmata may be important in changes in ¯oral sex expression and symmetry have all maintaining their spatial boundaries (Perbal et al., 1996). been effected via mutations in such genes in model sys- MADS box proteins can act alone or in combination as tems (Coen, 1991). These same morphological character- repressors or activators of transcription at other MADS istics often are used
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