Xin-Sheng Hu

On estimation of the ratio of pollen to seed flow among plant populations

Gene flow occurs in two ways for hermaphrodite plants; seed flow and pollen flow. Dispersal of biparentally inherited (nuclear) and paternally inherited (conifer chloroplast) genes can be mediated by both seed and pollen, whereas for maternally inherited (angiosperm chloroplast and most mitochondrial) genes only seed flow contributes to dispersal. This produces asymmetrical migration for biparentally, paternally and maternally inherited genes and may lead to different levels of population differentiation among them. This paper explores the effects of contrasting patterns of gene flow for different plant genes on their population structure under isolation by distance, on Neis genetic distance measure, on divergence in nucleotide sequence between populations and on gene phylogenies. The possibilities are discussed of using data on population structure, genetic distance, sequence divergence and gene phylogenies as a basis for estimating the ratio of pollen to seed flow among subpopulations. One important general result from the isolation-by-distance model is that population differentiation for maternally inherited genes is greater than that for paternally inherited genes, which, in turn, is greater than that for biparentally inherited genes as long as the dispersal of seeds and pollen grains takes place. This is consistent with results obtained previously for the island and stepping-stone models in which populations are discretely distributed.

Species diversity in local neutral communities

We extend the neutral theory of macroecology by deriving biodiversity models (relative species abundance and species‐area relationships) in a local community‐metacommunity system in which the local community is embedded within the metacommunity. We first demonstrate that the local species diversity patterns converge to that of the metacommunity as the size (scale) of the embedded local community increases. This result shows that in continuous landscapes no sharp boundaries dividing the communities at the two scales exist; they are an artificial distinction made by the current spatially implicit neutral theory. Second, we remove the artificial restriction that speciation cannot occur in a local community, even if the effects of local speciation are small. Third, we introduce stochasticity into the immigration rate, previously treated as constant, and demonstrate that local species diversity is a function not only of the mean but also of the variance in immigration rate. High variance in immigration rates reduces species diversity in local communities. Finally, we show that a simple relationship exists between the fundamental diversity parameter of neutral theory and Simpson’s index for local communities. Derivation of this relationship extends recent work on diversity indices and provides a means of evaluating the effect of immigration on estimates of the fundamental diversity parameter derived from relative species abundance data on local communities.

Seed and pollen flow and cline discordance among genes with different modes of inheritance.

The relationships between seed and pollen flow and cline discordance/concordance between cytoplasmic and nuclear genes, with the incorporation of the effects of natural selection, are formulated for one locus with two alleles, under assumptions of random mating, no drift and no mutation. Results show that under certain conditions, the relative roles of seed and pollen flow in shaping cline discordance/concordance are very similar to their roles in influencing population differentiation for selectively neutral markers with different modes of inheritance. Where the disequilibria between cytoplasmic and nuclear genes are of the order similar to selection coefficient, cline discordance/concordance can be predicted from the relative values of the ratio of pollen to seed flow and the ratio of selection coefficients. Where the disequilibria attained by seed and pollen flow are significant, the integrated cytonuclear data are recommended for cline analysis. In both cases, the relative rates of selection coefficients between cytoplasmic and nuclear genes can be roughly estimated according to their characteristic length.

On migration load of seeds and pollen grains in a local population

We have extended Wrights model of migration load to hermaphrodite plants showing variation at a single locus with two alleles. The model incorporates independent migration of seeds and pollen grains, the selection at both the haploid gametophyte and the diploid sporophyte stages, and a mixed mating system. The analytical relations between migration load and migration rate of seeds and pollen grains are explicitly formulated. The results show that under certain conditions, seed flow can have a more effect on migration load than pollen flow. Pollen selection at the gametophyte stage cannot substantially affect the migration load at the sporophyte stage. Selection at the diploid sporophyte stage is critical in determining the migration load of pollen grains. The relative migration loads of pollen versus seeds can be approximately estimated in predominantly outcrossing populations by the ratio of pollen flow to twice the seed flow, when the selection coefficient (sT) is greater than, or approximately equal to, the migration rate ([mtilde ]).

Structural Genomics: Correlation Blocks, Population Structure, and Genome Architecture

An integration of the pattern of genome-wide inter-site associations with evolutionary forces is important for gaining insights into the genomic evolution in natural or artificial populations. Here, we assess the inter-site correlation blocks and their distributions along chromosomes. A correlation block is broadly termed as the DNA segment within which strong correlations exist between genetic diversities at any two sites. We bring together the population genetic structure and the genomic diversity structure that have been independently built on different scales and synthesize the existing theories and methods for characterizing genomic structure at the population level. We discuss how population structure could shape correlation blocks and their patterns within and between populations. Effects of evolutionary forces (selection, migration, genetic drift, and mutation) on the pattern of genome-wide correlation blocks are discussed. In eukaryote organisms, we briefly discuss the associations between the pattern of correlation blocks and genome assembly features in eukaryote organisms, including the impacts of multigene family, the perturbation of transposable elements, and the repetitive nongenic sequences and GC-rich isochores. Our reviews suggest that the observable pattern of correlation blocks can refine our understanding of the ecological and evolutionary processes underlying the genomic evolution at the population level.

Assessment of the ratio of pollen to seed flow in a cline for genetic variation in a quantitative trait

A dispersal–selection cline model is analysed to evaluate the role of the ratio of pollen to seed flow (r) in spatial genetic variation, with a focus on clines in additive and dominant variances of major genes affecting a quantitative trait, assuming one locus with two alleles, no genetic drift and no mutation. It is shown that under weak selection, steady-state departures from the value at Hardy–Weinberg equilibrium (HWE) for genotype frequency (D) and additive variance (ΔVa) generally display a small value on one side of the selection boundary point, a high value around the point, and a moderate value on the other side of the boundary point. A large value of r can enhance formation of this pattern. However, this is not the case for clines in the average effect of a gene substitution (Δα) or in dominance variance (ΔVd), where large values of r can eradicate the clines. Direct use of their values at HWE to approximate the real case is acceptable. There is an expected turning point that divides clines of either additive variance or dominance variance into two subclines, each with a shorter width than that of allele frequency. Integration of these properties can help to indicate the existence of major genes affecting a quantitative trait.

Estimating the correlation of pairwise relatedness along chromosomes.

The ‘spatial’ pattern of the correlation of pairwise relatedness among loci within a chromosome is an important aspect for an insight into genomic evolution in natural populations. In this article, a statistical genetic method is presented for estimating the correlation of pairwise relatedness among linked loci. The probabilities of identity-in-state (IIS) are related to the probabilities of identity-by-descent (IBS) for the two- and three-loci cases. By decomposing the joint probabilities of two- or three-loci IBD, the probability of pairwise relatedness at a single locus and its correlation among linked loci can be simultaneously estimated. To provide effective statistical methods for estimation, weighted least square (LS) and maximum likelihood (ML) methods are evaluated through extensive Monte Carlo simulations. Results show that the ML method gives a better performance than the weighted LS method with haploid genotypic data. However, there are no significant differences between the two methods when two- or three-loci diploid genotypic data are employed. Compared with the optimal size for haploid genotypic data, a smaller optimal sample size is predicted with diploid genotypic data.

FST in the cytonuclear system

Selection on nuclear (or organelle) sites inevitably affects the spatial distribution of a neutral organelle (or nuclear) allele via transient cytonuclear disequilibrium. Here I examine this effect in terms of F(st) for a neutral allele by bringing together cytonuclear genomes with contrasting modes of inheritance. The relationships between cytonuclear disequilibrium and increment in F(st) are explored and confirmed through Monte Carlo simulations. Results show that the transient increment in F(st) for a neutral allele is not only related to the vectors of seed and pollen dispersal but also to the mode of its inheritance. Such increments can be substantial under certain conditions. Seed dispersal is more effective than pollen dispersal in changing the transient increment. The cumulative effects from multiple selective nuclear sites can amplify the transient increment in F(st) for a neutral paternal or maternal organelle allele. Selection on selective organelle sites facilitates the transient increment in F(st) for a neutral nuclear allele. Partial selfing can significantly reinforce the transient increment in F(st). These theoretical insights highlight the roles of transient cytonuclear disequilibrium as a biological factor in evolving population differentiation and refine our practical interpretations of F(st) with cytonuclear markers.