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In many angiosperm species, populations are reproductively subdivided into distinct sexual morphs including females, males and hermaphrodites. Sexual polymorphism is maintained by frequency-dependent selection, leading to predictable sex ratios at equilibrium. Charles Darwin devoted much of his book ‘The Different Forms of Flowers on Plants of the Same Species’ (1877) to investigating plant sexual polymorphisms and laid the foundation for many problems addressed today by integrating theory with empirical studies of the demography and genetics of populations. Here, we summarize our recent work on the ecological and genetic mechanisms influencing variation in sex ratios and their implications for evolutionary transitions among sexual systems. We present the results of a survey of sex ratios from 126 species from 47 angiosperm families and then address two general problems using examples from diverse angiosperm taxa: (i) the mechanisms governing biased sex ratios in dioecious species; (ii) the origins and maintenance of populations composed of females, males and hermaphrodites. Several themes are emphasized, including the importance of non-equilibrium conditions, the role of life history and demography in affecting sex ratios, the value of theory for modelling the dynamics of sex ratio variation, and the utility of genetic markers for investigating evolutionary processes in sexually polymorphic plant populations. [ABSTRACT FROM AUTHOR]

The role of mutations of small versus large effect in adaptive evolution is of considerable interest to evolutionary biologists. The major evolutionary pathways for the origin of dioecy in plants (the gynodioecy and monoecy-paradioecy pathways) are often distinguished by the number of mutations involved and the magnitude of their effects. Here, we investigate the genetic and environmental determinants of sex in Sagittaria latifolia, a species with both monoecious and dioecious populations, and evaluate evidence for the evolution of dioecy via gynodioecy or monoecy-paradioecy. We crossed plants of the two sexual systems to generate F1, F2 and backcross progeny, and grew clones from dioecious populations in low- and high-fertilizer conditions to examine sex inconstancy in females and males. Several lines of evidence implicate two-locus control of the sex phenotypes. In dioecious populations sex is determined by Mendelian segregation of alleles, with males heterozygous at both the male- and female-sterility loci. In monoecious populations, plants are homozygous for alleles dominant to male sterility in females and recessive to female sterility in males. Experimental manipulation of resources revealed sex inconstancy in males but not females. These results are consistent with predictions for the evolution of dioecy via gynodioecy, rather than the expected monoecy-paradioecy pathway, given the ancestral monoecious condition. [ABSTRACT FROM AUTHOR]

Negative frequency-dependent selection is a major selective force maintaining sexual polymorphisms. However, empirical demonstrations of frequency-dependent reproductive success are rare, particularly in plants. We investigate this problem by manipulating the frequencies of style morphs in a natural population of Narcissus assoanus, a self-incompatible herb with style-length dimorphism and intra-morph compatibility. We predicted that the reproductive success of morphs would vary negatively with their frequency because of the effects of morph-specific differences in sex-organ position on patterns of pollen transfer. This prediction was generally supported. The fruit and seed set of the two morphs did not differ significantly in plots with 1 : 1 morph ratios. However, short-styled plants produced significantly fewer seeds than long-styled plants in monomorphic plots, and significantly more seeds than long-styled plants in plots with 'long-biased' morph ratios. These patterns indicate that in the absence of physiological barriers to intra-morph mating, negative frequency-dependent selection contributes to the maintenance of stylar polymorphism through inter-morph pollen transfer. Our experimental results also provide insights into the mechanisms governing the biased style-morph ratios in populations of Narcissus species. [ABSTRACT FROM AUTHOR]