Clarkia tembloriensis exhibits a wide range of variation among its natural populations in outcrossing rate and in separation ofmale and female function in space (anther-stigma separation or herkogamy) and in time (protandry). Here we show that outcrossing rate is highly correlated with protandry and anther-stigma separation. Both genetic and environmental variation contribute to inter-and intrapopulation variation in protandry and anther-stigma separation. Interpopulation differentiation for protandry and anther-stigma separation was found to be polygenic. Genetic variation for protandry and anther-stigma separation within populations was demonstrated by a significant among-family variance in two populations with contrasting breeding systems. Environmental effects on the expression of mating system traits were manifested in two ways. First, significant variation among lathhouse benches suggests that small-scale environmental heterogeneity may affect the development of floral traits. Second, protandry was shortened under hot summer conditions. Hence, hotter and drier habitats, typical of the more self-pollinating populations of C. tembloriensis, can promote self-pollination purely through environmental effects.
Abstract. -We explored the extent to which the soil seed bank differed genetically and spatially in comparison to two actively growingstages in a natural population of Plantago lanceolata. All seedbank seeds, seedlings,and adults of P. lanceolata within eight subunits in a larger population were mapped, subjected to starch gelelectrophoresis, and allozyme analysis in 1988. Gel electrophoresis was also used to estimate the mating system in two years, 1986 and 1988. The spatial distributions of seeds, seedlings, and adults were highly coincident. Allele frequencies of the dormant seeds differed significantly from those of the adults for four of the five polymorphic loci. In addition, a comparison ofthe genotype frequencies of the three life-history stages indicated that the seed bank had an excessofhomozygotes. Homozygosity, relative to Hardy-Weinberg expectations, decreased during the life cycle (for seed bank, seedlings, and adults respectively: F;t = 0.19, 0.09, 0.01; F;, = 0.14, 0.04, -0.12). Spatial genetic differentiation increased sixfold during the life cycle: (for seed bank, seedling and adults: F" = 0.02, 0.05, 0.12). The apparent selfingrate was 0.01 in 1986 and 0.09 in 1988. These selfingrates are not large enough to account for the elevated homozygosity of the seed bank. Inbreeding depression, overdominance for fitness, and a "temporal Wahlund's effect" are discussed as possible mechanisms that could generate high homozygosity in the seed bank, relative to later life-history stages. In Plantago lanceolata, the influence of the mating system and the "genetic memory" of the seed bank are obscured by the time plants reach the reproductive stage.
Ribosomal internal transcribed spacer (ITS) sequences were used to evaluate the phylogenetics of Zea and Tripsacum. Maximum likelihood and polymorphism parsimony were used for phylogenetic reconstructions. Zea ITS nucleotide diversity was high compared to other plant species, but approximately equivalent to other maize loci. Coalescence of ITS alleles was rapid relative to other nuclear loci; however, there was still much diversity within populations. Zea and Tripsacum form a clade clearly differentiated from all other Poaceae. Four Zea ITS pseudogenes were identified by phylogenetic position and nucleotide composition. The phylogenetic position of Z. mays ssp. huehuetenangensis was clearly established as basal to the other Z. mays. The ITS phylogeny disfavored a Z. luxurians and Z. diploperennis clade, which conflicted with some previous studies. The introgression of Z. mays alleles into Z. perennis and Z. diploperennis was also established. The ITS data indicated a near contemporary divergence of domesticated maize and its two closest wild relatives.
Inbreeding depression is commonly observed in natural populations. The deleterious effects of forced inbreeding are often thought to be less pronounced in populations with self‐pollinating mating systems than in primarily outcrossing populations. We tested this hypothesis by comparing the performance of plants produced by artificial self‐ and cross‐pollination from three populations whose outcrossing rate estimates were 0.03, 0.26, and 0.58. Outcrossing rates and inbreeding coefficients were estimated using isozyme polymorphisms as genetic markers. Analysis of F statistics suggests that biparental inbreeding as well as self‐fertilization contribute to the level of homozygosity in the seed crop. Biparental inbreeding will reduce the heterozygosity of progeny produced by outcrossing, relative to random outcrossing expectations, and hence will reduce the effects of outcrossing versus self‐fertilization. Heterotic selection may increase the average heterozygosity during the life history. Selfed and outcrossed seeds from all three populations were equally likely to germinate and survive to reproduce. However, inbreeding depression was observed in fecundity traits of plants surviving to reproduction in all three populations. Even in the population whose natural self‐fertilization rate was 97%, plants grown from seed produced by self‐pollination produced fewer fruits and less total seed weight than plants grown from outcrossed seed. There was no detectable inbreeding depression in estimated lifetime fitness. Inbreeding effects for all reproductive yield characters were most severe in the accession from the most outcrossing population and least severe in the accession from the most self‐fertilizing population.
Nicotiana section Alatae exhibits great diversity among species in floral morphology, mating system, and predominant pollinators. As a first step towards estimating nectar's role in floral evolution, we studied nectar traits to determine whether they vary in association with predominant pollinators and mating system. Daily phenology determines when nectar becomes available to pollinators and differed between hummingbird- and moth-pollinated species. Nectar volume and concentration varied significantly among most species and pollinator groups, but were inversely correlated, so that total energy was similar among most species. In general, nectar volume was positively correlated with corolla length. The autogamous species, N. plumbaginifolia, had a nectar volume that matched expectations based on corolla length, but with lower concentration and total energy than predicted by corolla length, while nectar volume was lower than predicted by corolla length in the autogamous population of N. longiflora. Sugar and amino acid components (determined through HPLC) were similar among species, although differences did exist. The nectar of most species was sucrose-dominant, but the autogamous N. plumbaginifolia had nectar that contained similar proportions of sucrose, glucose, and fructose. Many nectar traits varied in association with the predominant pollinators and, in some cases, with the mating system.
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