Mixed-Mating Model of Reproduction Revealed in European Phytophthora cactorum by ddRADseq and Effector Gene Sequence Data

A population study of Phytophthora cactorum was performed using ddRADseq sequence variation analysis completed by the analysis of effector genes—RXLR6, RXLR7 and SCR113. The population structure was described by F-statistics, heterozygosity, nucleotide diversity, number of private alleles, number of polymorphic sites, kinship coefficient and structure analysis. The population of P. cactorum in Europe seems to be structured into host-associated groups. The isolates from woody hosts are structured into four groups described previously, while isolates from strawberry form another group. The groups are diverse in effector gene composition and the frequency of outbreeding. When populations from strawberry were analysed, both asexual reproduction and occasional outbreeding confirmed by gene flow among distinct populations were detected. Therefore, distinct P. cactorum populations differ in the level of heterozygosity. The data support the theory of the mixed-mating model for P. cactorum, comprising frequent asexual behaviour and inbreeding alternating with occasional outbreeding. Because P. cactorum is not indigenous to Europe, such variability is probably caused by multiple introductions of different lineages from the area of its original distribution, and the different histories of sexual recombination and host adaptation of particular populations.

Microsatellite analysis of the mating system of Bruguiera gymnorrhiza and Kandelia obovata

Microsatellite loci were used for estimating mating system for three populations of B. gymnorrhiza and K. obovata (Rhizophoracea) in Okinawa, Japan. Mother trees and thirty offspring of individual samples representing the population of both species were genotyped at five microsatellites. The mating system was examined using two approaches: a mixed mating model of multilocus testing, implemented by MLTR program and outcrossing rate from the level of inbreeding. Mating system analysis showed multilocus outcrossing rates (tm) for both species was 0.850-1.000 and 0.780-0.938 respectively. By contrast, according to inbreeding level, tm was lower than MLRT: 0.495-1.028 and 0.480-0.612 of both species respectively. However, biparental inbreeding (tm- ts) was diverse from zero both species for all three populations, showing that cross-fertilization events may ensue between the relatives both species. This data as well means the genetic relatedness (r) for B. gymnorrhiza and K. obovata were 0.108±0.025 and 0.032±0.09 respectively. Average relatedness was below 0.25, the value for a half-sib relationship. These results suggest that postulation of a half-sib relationship among progeny of open-pollinated families is opposed for both mangrove tree species.

Outcrossing rate of Couratari multiflora (J.Smith) Eyma (Lecythidaceae), a low-density tropical tree species from a central Amazonian rainforest

A multilocus mixed-mating model was used to evaluate the mating system of a population of Couratari multiflora, an emergent tree species found in low densities (1 individual/10 ha) in lowland forests of central Amazonia. We surveyed and observed phenologically 41 trees in an area of 400 ha. From these, only four mother trees were analyzed here because few of them set fruits, which also suffered high predation. No difference was observed between the population multilocus outcrossing rate (t mp = 0.953 ± 0.040) and the average single locus rate (t sp = 0.968 ± 0.132). The four mother trees were highly outcrossed (t m ~ 1). Two out of five loci showed departures from the Hardy-Weinberg Equilibrium (HWE) expectations, and the same results occurred with the mixed-mating model. Besides the low number of trees analyzed, the proportion of loci in HWE suggests random mating in the population. However, the pollen pool was heterogeneous among families, probably due to both the small sample number and the flowering of trees at different times of the flowering season. Reproductive phenology of the population and the results presented here suggest, at least for part of the population, a long-distance pollen movement, around 1,000 m.

Estimation of mating system parameters in two Italian natural populations of Fagussylvatica

Mating system parameters were estimated in two Italian natural populations of European beech (Fagussylvatica L.) using a mixed mating model and considering seven allozyme loci (Idh-A, Lap-A, Mdh-B, Pgd-A, Pgd-B, Pgd-C, Skd-A). High values of multilocus estimates of the outcrossing rate were found in both populations, ranging from 94 to 98%. Comparison of single- and multi-locus estimates of outcrossing rates seems to indicate the presence of consanguineous matings, probably because the populations are substructured. This hypothesis seems to be confirmed by the presence of a heterogeneity of the pollen allele frequencies across female parent trees and by the significant coefficient of the regression of pollen allele frequencies on ovule genotype. Variation in the fixation indices in different life-cycle phases was observed, indicating possible presence of selective processes between seed set and sexual maturity. Possible explanations of these results are presented.

Outcrossing in two faba bean cultivars under dryland conditions in Spain

Field outcrossing rates were estimated for two faba bean (Vicia faba L.) cultivars in 3 yr in Spain using isozyme markers and the mixed mating model for the determination of single and multilocus rates. Out-crossing rates were stable during the years for the local cultivar and higher than previously thought for the Mediterranean region. Key words:Vicia faba, outcrossing rates, isozyme marker

The Mating System and Genetic Diversity of the Australian Arid Zone Mallee, Eucalyptus rameliana

Estimates of parameters of the mixed mating model were made for three populations of the bird-pollinated arid zone eucalypt, Eucalyptus rameliana F. Muell. Levels of outcrossing ((t) over cap) varied significantly between populations from mixed mating with substantial selfing ((t) over cap = 0.54) to almost completely outcrossed ((t) over cap = 0.95). Comparison of single-locus and multilocus estimates suggested that the drop in outcrossing was due to increased self-pollination. The lowest outcrossing rate was attributed to the reduced ability of a population with low numbers of buds to attract bird pollinators. Outcrossing rates in E. rameliana are proposed to be a more direct reflection of pollination than estimates made for mass flowering, i.e. small-fruited eucalypts. The distribution of allozyme diversity in E. rameliana also appeared to reflect the impact of bird pollinators in promoting gene flow as well as the species capacity for outcrossing and introgression. Levels of diversity were comparable with other eucalypts, but the proportion of diversity between populations (GST = 9.2%) was only about half the mean for other eucalypts. Genetic distances between populations were low, but there was same significant differentiation of populations which was attributed to non-random bird migrations. The importance of bird pollination in the mating system and the distribution of genetic diversity in E. rameliana emphasises that enough habitat to support nomadic birds should be preserved in order to conserve this eucalypt species.

Mating system of Racosperma auriculiforme in a seed production area in Zaire

Analysis of the mating system of Racosperma auriculiforme (formerly Acacia auriculiformis) in a seed production area in Zaire was conducted using a mixed mating model at five polymorphic loci (Est-2, Est-3, G6p-dh, Nadhdh-2, Pgm-1). Single-locus estimates of outcrossing rate varied widely between 0.615 and 1.204 and were significantly heterogenous among and within the stands. Weighted means of single-locus estimates of outcrossing rates (ts) ranged from 0.859 to 1.032 while multilocus estimates of outcrossing rates (tm) ranged from 0.863 to 1.059. Comparisons between the single-locus estimates and the multilocus estimates of outcrossing rates indicated that most of the inbreeding detected was due to consanguineous (sibling) matings rather than selfing. Comparisons of observed and expected inbreeding coefficients indicated a general deficit of heterozygotes over the frequencies expected from random mating in the single stands and the pooled stands. Locus to locus variation in outcrossing rates suggested that violation of the mixed mating model might have occurred. Implications for seed production, collection, and distribution for reforestation are discussed. Key words: Acacia, mating system, Racosperma, seed production area, Zaire.

Definition and Properties of Disequilibrium Statistics for Associations Between Nuclear and Cytoplasmic Genotypes

ABSTRACT We define and establish the interrelationships of four components of statistical association between a diploid nuclear gene and a uniparentally transmitted, haploid cytoplasmic gene: an allelic (gametic) disequilibrium (D), which measures associations between alleles at the two loci; and three genotypic disequilibria (D 1, D 2, D 3), which measure associations between two cytotypes and the three respective nuclear backgrounds. We also consider an alternative set of measures, including D and the residual disequilibrium (d). The dynamics of these disequilibria are then examined under three conventional models of the mating system: (1) random mating; (2a) assortative mating without dominance (the "mixed-mating model"); and (2b) assortative mating with dominance ("O'Donald's model"). The trajectories of gametic disequilibria are similar to those for pairs of unlinked nuclear loci. The dynamics of genotypic disequilibria exhibit a variety of behaviors depending on the model and the initial conditions. Procedures for statistical estimation of cytonuclear disequilibria are developed and applied to several real and hypothetical data sets. Special attention is paid to the biological interpretations of various categories of allelic and genotypic disequilibria in hybrid zones. Genetic systems for which these statistics might be appropriate include nuclear genotype frequencies in conjunction with those for mitochondrial DNA, chloroplast DNA, or cytoplasmically inherited microorganisms.