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PubMed2006

The genetic analysis of family structured inbreeding depression studies.

An important question emerging from theoretical studies of mating system evolution is whether the fitness of a randomly extracted, fully inbred genotype will exceed the mean of outbred individuals. We introduce two statistics (I(1) and I(2)) related to the probability of extracting a high line. I(1) and I(2) can be estimated from the family structured experimental designs typically used to estimate inbreeding depression (ID). Maximum likelihood procedures are developed from an explicit genetic model. These yield parameter estimates and provide the likelihoods necessary to test hypotheses, for example, whether population-level ID is nonzero. Finally, we describe a new publicly available computer program titled 'IDG' (Inbreeding Depression Genetics) to execute these procedures.

Analysis of Variance↗

PubMed1999

Patterns of allozymic variation within calluna vulgaris populations at seed bank and adult stages

We investigated the spatial genetic structure within and between two plots of Calluna vulgaris and the extent to which the soil seed bank differed genetically from adults at seven allozyme loci. Averaged over the two plots, the seed bank and adult populations contained very similar levels of genetic diversity. Moreover, seeds contained in a single soil core (100 cm3) exhibited similar mean allozyme diversity to the surrounding adult population, indicating that the seed bank preserves genetic diversity at a very local scale. Few differences in allelic frequencies were found between the seed bank and its surrounding adult population in each plot. Mean GST indicated a lack of differentiation between the two plots at adult (GST = 0.008) and seed bank (GST = 0.002) stages. Low interplot differentiation is consistent with the outcrossing mating system of the population (tm = 0.91 in one plot) and its history of human disturbance. In contrast, spatial autocorrelation analysis of adults indicated a genetic structure at a very local scale, with positive autocorrelation for all alleles below 2 m in one plot and with a pattern of positive autocorrelation below 8 m in the two plots. Current limitation to seed dispersal rather than spatial extension of clones is thought to be responsible for local genetic structure.

Journal Article↗

PubMed1999

History can be more important than 'pollination syndrome' in determining the genetic structure of plant populations: the case of aconitum lycoctonum (Ranunculaceae)

The Aconitum lycoctonum complex is a widespread yellow-flowered group of species found in central and southern Europe. Because of extreme morphological variability, the systematics of this group is confusing, and hybridizations among taxa are often hypothesized. To determine whether hybridization, realized mating system within populations or colonization from different Pleistocene refugia might explain some of the morphological variation, the genetic structure of 19 populations from central and southern Europe was studied using starch gel electrophoresis and 10 enzyme loci, of which eight were polymorphic. A pattern typical of an outcrossing species was expected because A. lycoctonum flowers are adapted to pollination by long-tongued bumblebees. However, heterozygosity was very low (between 0.031 and 0.150), which is atypical of either widespread or outcrossing species. The inbreeding coefficient, FIS, also suggested inbreeding in more than half the populations. Analysis of molecular variance showed that 31% of the genetic variation is found among populations, again suggesting inbreeding. A neighbour-joining tree based on Reynolds's genetic distance showed a clear separation between the central/eastern European populations and the samples from the Iberian peninsula and Alpes Maritimes. These data are consistent with the hypothesized refugia and the phylogeographical histories of several European forest trees. The genetic identity of all populations was very high, suggesting that all investigated populations belong to the same species despite high morphological variability. Hybridization was not supported by the data.

Journal Article↗

PubMed1999

Inbreeding depression and partial selfing: evolutionary implications of mixed-mating in a coastal endemic, silene douglasii var. oraria (Caryophyllaceae)

Recent studies have found moderate to high levels of selfing in plants despite high inbreeding depression. Because both factors influence the evolution and persistence of rare plants, we conducted glasshouse and field studies of pollination and inbreeding in Silene douglasii var. oraria, a perennial tetraploid endemic to coastal prairies. We detected: (i) variation in reproduction or inbreeding depression among life stages, years and maternal families; (ii) partial selfing yet higher relative fitness in outcrossed than selfed progeny; (iii) differing values of selfing and inbreeding depression using population means vs. matched maternal families. Fruit and seed production varied significantly with pollination treatment and year in flowers manipulated in situ during three seasons of growth. Hand-pollinations providing pollen in excess of ovule production in 1996 yielded more seeds than marked, open-pollinated flowers, implying pollen limitation of seed production. However, among-year differences in reproductive success after open-pollination (i.e. values equivalent to autogamy, selfing or outcrossing) suggest that pollination levels also vary temporally. In pollinations matched by maternal family, selfing yielded significantly fewer seeds than outcrossing. Fitness differences between inbred and outbred progeny were significant (P < 0.05) for seed production, percentage germination, and biomass or fecundity, but not for survival. Maternal family data gave selfing rates intermediate between obligate outcrossers and predominant selfers (S = 0.34-0.51), but population-wide means gave unusually high values (S = 1.1-1.6). Cumulative inbreeding depression was 76% for maternal families, and 70-85% using population means; in all cases, inbreeding depression values were high in early and late life stages, and lowest for survival. Thus far, reproductive assurance offers the most cogent explanation for the coexistence of moderate selfing and high inbreeding depression in this strongly protandrous Silene once thought to be highly outcrossing. This possibility merits further study in other rare plants with mixed-mating systems, where inbreeding depression and pollinator scarcity may both compromise population persistence and raise the threshold below which selfing is favoured by evolution.

Journal Article↗

PubMed1999

The contribution of male-sterility mutations to inbreeding depression in Mimulus guttatus.

The magnitude of inbreeding depression can influence many aspects of a population's ecology and evolution, including the nature of selection acting on the mating system and the chances that the population will go extinct during periods of small population size. If inbreeding depression is caused primarily by recessive mutations of large effect on fitness, such as lethals or steriles, then it is expected to be purged rapidly by selection with moderate amounts of inbreeding. In contrast, inbreeding depression primarily caused by many genes with mild effects on fitness will not be rapidly purged with inbreeding, so it should be a more resilient barrier to the evolution of self-fertilization and a more significant threat to the survival of endangered species. Here I show that recessive male-sterility alleles at individual loci are common in a primarily outcrossing population of the plant Mimulus guttatus. Despite the high frequency of these major mutations, most of the inbreeding depression for male fertility and cumulative measures of lifetime fitness results from more mildly deleterious alleles. Male-sterility alleles contribute to 31% of the inbreeding depression for the fraction of viable pollen grains, and to 26% of the inbreeding depression for total fitness. These results suggest that most of the inbreeding depression for male fertility in this population would not be purged, in the short term, with moderate inbreeding.

Alleles↗

PubMed2001

Microsatellite analysis of female mating behaviour in lek-breeding sage grouse.

We used microsatellite DNA markers to genotype chicks in 10 broods of lek-breeding sage grouse, Centrocercus urophasianus, whose mothers' behaviour was studied by radio-tracking and observing leks. Previous behavioural studies suggested that almost all matings are performed by territorial males on leks and that multiple mating is rare. Two broods (20%) were sired by more than one male. Genetic analyses of the broods of eight females that visited an intensively studied lek were consistent with behavioural observations. Four females observed mating produced singly sired broods and males other than the individual observed copulating were excluded as sires for most or all of their chicks. Territorial males at the study lek were excluded as sires of broods of four other females that visited the lek but were not observed mating there. Radio-tracking suggested that two of these females mated at other leks. Our results confirm the reliability of mating observations at leks, but do not rule out a possible unseen component of the mating system.

Animals↗

PubMed2002

Male reproductive competition in spawning aggregations of cod (Gadus morhua, L.).

Reproductive competition may lead to a large skew in reproductive success among individuals. Very few studies have analysed the paternity contribution of individual males in spawning aggregations of fish species with huge census population sizes. We quantified the variance in male reproductive success in spawning aggregations of cod under experimental conditions over an entire spawning season. Male reproductive success was estimated by microsatellite-based parentage analysis of offspring produced in six separate groups of spawning cod. In total, 1340 offspring and 102 spawnings distributed across a spawning season were analysed. Our results show that multiple males contributed sperm to most spawnings but that paternity frequencies were highly skewed among males, with larger males on average siring higher proportions of offspring. It was further indicated that male reproductive success was dependent on the magnitude of the size difference between a female and a male. We discuss our results in relation to the cod mating system. Finally, we suggest that the highly skewed distribution of paternity success observed in cod may be a factor contributing to the low effective population size/census population size ratios observed in many marine organisms.

Animals↗

PubMed2001

Predictors of marker-informativeness for an outbred F2 design.

Generalization of the polymorphism information content (PIC) index to represent marker informativeness (MI) for a three-generation F2 design requires that two additional sources of non-informativeness be added to the PIC formula: the probability of matings between like-heterozygous F1 individuals, of which one is non-informative; and that of matings between like-heterozygous F1 individuals, which are both fully informative but where line of origin of the same alleles is reciprocal. Given the dense marker-maps currently available for some species, this F2 informativeness parameter constitutes the natural criterion for marker selection in F2 designs, and two computer programs to predict MI from grandparental marker-genotypes were developed for an F2 population originating from two divergent selection lines of outbred mice (F approximately 0.2). A total of 403 markers had been genotyped for the F0 grandparents (n=31), and 14 markers had also been genotyped in the complete pedigree including 559 F2 individuals. One program was based on assumptions of random-mating (RM), while the other (PED) accounted for the pedigreed mating structure. For the 403 markers, the correlation between MI from RM and from PED was 0.95, and the average deviation between the two predictions was 0.005 MI units (MI ranged from 0 to 1). Correlations between predicted and realized MI for the 14 fully genotyped markers were 0.97 for PED and 0.94 for RM, while the corresponding average of deviations between predicted and actual values were 0.01 and 0.04, respectively. Absolute deviations from realized MI never exceeded 0.09 and 0.16 for PED and RM, respectively. Simulated optimization of the mating system to maximize average MI of 28 markers on one chromosome led to improvements in the range of 15-20% average MI (0.07-0.09 MI units). The degree of relative advantage conferred by the F2 generalization of the PIC index over the traditional index was found to be of minor significance.

Animals↗

PubMed1999

RAPD variation within and between natural populations of the wild rice oryza rufipogon from china and brazil

Genetic variation within and between eight natural populations of Oryza rufipogon from China and Brazil was investigated at the DNA level by analysis of RAPD fragments. Out of 60 random primers, which were initially screened against DNA from four individuals, 20 generated highly reproducible RAPD fragments which were then used for further population analysis. With these primers, 95 discernible DNA fragments were produced and 78 (82.1%) were polymorphic, which indicated that high levels of genetic variation existed in these natural populations. In addition, the Chinese populations showed greater polymorphism than those from Brazil at both the population and regional levels. This is noteworthy considering that the Chinese populations are from a relatively restricted area of China. The factors responsible for these findings include the contrasting mating systems in the Brazilian and Chinese populations, and gene flow from annual cultivated rice to perennial natural populations in China. An Analysis of Molecular Variance (AMOVA) was used to apportion the variation between individuals within populations, between populations within regions, and between regions. Results showed that 61.8% of the total genetic diversity resided between the two continents, whereas only 14.9% and 23.3% was attributable to population differences within regions and to individual differences within a population, respectively. The great genetic differentiation between the Chinese and Brazilian populations is in agreement with recent treatment of the American form of O. rufipogon as a separate species, O. glumaepatula.

Journal Article↗

PubMed2000

Genetic structure and colonizing success of a clonal, weedy species, Pilosella officinarum (Asteraceae).

Introduced populations of weeds which are polyploid and reproduce primarily by apomixis are generally considered as having low levels of population genetic variation, highly differentiated populations and short evolutionary lifespans. Although polyploidy allows for habitat differentiation and colonization, lack of recombination because of apomixis means that long-term persistence is unlikely. However, variation can be introduced to a colonizing population by evolutionary changes in the mating system, or by somatic mutation and recombination. In this study hypersensitive genetic markers, inter-simple sequence repeats (ISSRs), were used to quantify genetic variation within Pilosella officinarum, a major weed of the New Zealand high country. Pilosella officinarum was introduced from Europe to New Zealand late in the 19th century and only polyploid, apomictic populations are thought to have survived. The combination of introduction history and breeding system has led to the assumption that New Zealand populations are necessarily genetically depauperate. However, our studies reveal variable levels of genetic variation and patterns of clonal distribution which indicate varying levels of sexual reproduction within New Zealand populations.

Asteraceae↗

PubMed2001

Population genetic structure of the cleistogamous plant species Viola pubescens Aiton (Violaceae), as indicated by allozyme and ISSR molecular markers.

Few studies of genetic variation have focused on species that reproduce through both showy, chasmogamous (CH) flowers and self-pollinated, cleistogamous (CL) flowers. Using two different techniques, genetic variation was measured in six populations of Viola pubescens Aiton, a yellow-flowered violet found in the temperate forests of eastern North America. Results from eight allozyme loci showed that there was considerable genetic variation in the species, and population structuring was indicated by the presence of unique alleles and a theta (F(ST)) value of 0.29. High genetic variation was also found using ISSR (inter-simple sequence repeat) markers, and population structuring was again evident with unique bands. Viola pubescens appears to have a true mixed-mating system in which selfing through CL and CH flowers contributes to population differentiation, and outcrossing through CH flowers increases genetic variation and gene flow among populations. Overall, allozyme and ISSR techniques yielded similar results, indicating that ISSR markers show potential for use in population genetic studies.

Alleles↗

PubMed2001

Genetic variation and population structure of the mixed-mating cactus, Melocactus curvispinus (Cactaceae).

Genetic diversity was measured in the mixed-mating cactus, Melocactus curvispinus, in Venezuela. Allozyme diversity was surveyed in 19 putative loci over 18 populations. Compared to other plant taxa, this cactus is rich in polymorphic loci (Ps=89.5%), with high numbers of alleles per polymorphic locus (APs=3.82), but moderate levels of heterozygosity (Hes=0.145). Substantial levels of inbreeding were detected across loci and populations at macrogeographic (FIS=0.348) and regional levels (FIS=0.194-0.402). Moderate levels of genetic differentiation among populations were detected at macrogeographical (FST=0.193) and regional (FST=0.084-0.187) scales, suggesting that gene flow is relatively restricted, but increases within regions without topographic barriers. The population genetic structure observed for this cactus was attributed to, at least, three factors: short-distance pollination and seed dispersal, the mixed-mating condition of the species, and genetic drift. High genetic identities between populations (I=0.942) supported the conspecific nature of all populations surveyed. The levels and patterns of genetic structure observed for M. curvispinus were consistent with its mating system and gene dispersal mechanisms.

Cactaceae↗

PubMed1999

Microsatellite and morphological analysis of population structure in the parthenogenetic freshwater snail Melanoides tuberculata: insights into the creation of clonal variability.

The distribution of variability was studied at various geographical scales in the tropical freshwater snail Melanoides tuberculata, in order to analyse the role of factors shaping this distribution, including the mating system and population dynamics. This parthenogenetic polyploid species reproduces mainly asexually, with males occurring at low frequency. About 800 individuals (38 sites) were sampled from Africa and the Middle East, where the species originated, and from recently colonized habitats in South and Central America, and especially the island of Martinique. We first described variation of general aspects and ornamentation of the shells. This analysis confirms the existence of discrete morphs. Second, individuals were studied at three microsatellite loci, showing that each morph is a genetic clone with some minor variation compatible with models of microsatellite evolution. The genetic analysis also showed much more variation within than between clones. However, two populations from Africa exhibited a large amount of variability, and a mixture of sexual and asexual reproduction might explain these genetic patterns. The worldwide distribution of variability is, therefore, compatible with the African origin of the species, and the introduction of a few clones in other parts of the world. These results also suggest that the distribution of variability in Martinique is influenced by flooding events, and that two morphs from Martinique can be interpreted as hybrids between two pre-existing morphs, based on morphological, genetic and geographical arguments.

Journal Article↗

PubMed1999

Amplified fragment length polymorphism (AFLP) markers reveal extra-pair parentage in a bird species: the bluethroat (Luscinia svecica).

We tested the use of amplified fragment length polymorphism (AFLP) to assess the frequency of extra-pair parentage in a bluethroat (Luscinia svecica namnetum) population. Thirty-six families totalling 162 nestlings were analysed. Using a combination of three primer pairs, we reached an exclusion probability of 93% for the population. This probability can reach 99% considering families independently. We revealed that extra-pair fertilizations are very common: 63.8% of all broods contain at least one extra-pair young, totalling 41.9% of all young analysed. However, with the technique and the three primer pairs used it was not possible to attribute the parentage exclusions to extra-pair paternity, maternity or both. As brood parasitism has never been reported in this species, it seems likely that the exclusions are due to extra-pair males. This study shows that dominant AFLP markers can be useful for studying the mating system of taxa for which no microsatellite primers are available. This technique allows the approximate estimation of parentage exclusions despite the fact that it is not possible to know which parent has to be excluded.

Animals↗

PubMed2000

Genetic differentiation of Euterpe edulis Mart. populations estimated by AFLP analysis.

Heart-of-palm (Euterpe edulis Mart.) is a wild palm with a wide distribution throughout the Atlantic Rainforest. Populations of E. edulis represent important renewable natural resources but are currently under threat from predatory exploitation. Furthermore, because the species is indigenous to the Atlantic Rainforest, which is located in the most economically developed and populated region of Brazil, social and economic pressures have devastated heart-of-palm forests. In order to estimate the partitioning of genetic variation of endangered E. edulis populations, 429 AFLP markers were used to analyse 150 plants representing 11 populations of the species distribution range. Analysis of the genetic structure of populations carried out using analysis of molecular variance (AMOVA) revealed moderate genetic variation within populations (57. 4%). Genetic differentiation between populations (FST = 0.426) was positively correlated with geographical distance. These results could be explained by the historical fragmentation of the Atlantic coastal region, together with the life cycle and mating system. The data obtained in this work should have important implications for conservation and future breeding programmes of E. edulis.

Base Sequence↗

PubMed2001

Population genetic structure of the lemon shark (Negaprion brevirostris) in the western Atlantic: DNA microsatellite variation.

DNA microsatellite markers were used to characterize the population genetic structure of the lemon shark, Negaprion brevirostris, in the western Atlantic. This study demonstrates for the first time the usefulness of microsatellites to study population genetic structure and mating systems in the Chondricthyes. Lemon sharks (mostly juveniles) were sampled non-destructively from four locations, Gullivan Bay and Marquesas Key in Florida, Bimini, Bahamas, and Atol das Rocas, Brazil. At least 545 individuals were genotyped at each of four dinucleotide loci. The number of alleles per locus ranged from 19 to 43, and expected heterozygosities ranged from 0.69 to 0.90. Relatively little genetic structure was found in the western Atlantic, with small but significant values for estimators of F(ST) and R(ST) among populations, theta (0.016) and rho (0.026), respectively. No sharp discontinuities were found between the Caribbean sites and Brazil, and most alleles were found at all four sites, indicating that gene flow occurs throughout the western Atlantic with no evidence for distinct stocks.

Animals↗

PubMed2001

Genetic variation and clonal diversity in four clonal sedges (Carex) along the Arctic coast of Eurasia.

We studied the structure of genetic variation (at both ramet- and genet-level) and clonal diversity within and among populations in the four closely related arctic clonal sedges Carex bigelowii, C. ensifolia, C. lugens and C. stans by use of allozyme markers. Compared to other sedges and arctic plants, the studied taxa all had high levels of genetic variation, both within populations and taxa. These taxa contained most of the total gene diversity (H(T)) within populations and a small part of the diversity among populations (G(ST) ranged 0.05--0.43). Carex bigelowii had genetic variation (H(S) = 0.173, mean for populations) at a comparable level to other outbreeding arctic plants and to other widespread, rhizomatous and mainly outbreeding Carex species. In contrast, C. ensifolia (H(S) = 0.335), C. lugens (H(S) = 0.339) and C. stans (H(S) = 0.294) had within-population variations that were higher than in most other studied Carex species and for arctic plants in general. Genetic variation was not related to any tested environmental variable, but it was lower in areas deglaciated only 10,000 years BP compared to areas deglaciated 60,000 years BP or not glaciated at all during the Weichselian. All the populations were multiclonal, except for two populations of C. stans that were monoclonal. In contrast to genetic variation, clonal diversity decreased with latitude and did not differ between areas with different times of deglaciation. In accordance with previous studies, C. bigelowii and C. lugens were found to be outbreeding, while C. ensifolia and C. stans had mixed mating systems.

Arctic Regions↗

PubMed2002

Historical biogeography and molecular systematics of the Indo-Pacific genus Dascyllus (Teleostei: Pomacentridae).

The phylogeographical and systematic relationships among species in the tropical marine fish genus Dascyllus were inferred using mitochondrial DNA (mtDNA) sequence data. Although our results were generally consistent with previously published phylogenies based on both morphological and mitochondrial data, our broad taxonomic and geographical sampling design revealed novel insights into the phylogenetic history of Dascyllus that had escaped previous notice. These results include: (a) the polyphyletic nature of D. reticulatus mtDNAs, representing two divergent and geographically separated lineages, one shared with D. flavicaudus and the second forming the sister lineage of D. carneus; (b) the paraphyly of D. trimaculatus relative to the closely related D. abisella; and (c) phylogeographical structure within the widespread taxa D. aruanus and D. trimaculatus. Application of a molecular clock permits us to posit a causative role for tectonic and oceanic changes regarding some Dascyllus speciation events. Finally, we mapped body size and the presence or absence of protogynous sex change on the mtDNA tree, and tested published hypotheses regarding determinants of the evolution of mating system and protogyny in the genus. Our data rejected a model based on body size but not one based on phylogenetic inertia. The ability to change sex arose once in the ancestor to the entire genus, and was lost once in the ancestor of the D. trimaculatus complex. For taxa that are as geographically widespread as many Indo-Pacific genera, this study highlights the importance of adequate geographical sampling when inferring patterns of species diversification and life history evolution.

Animals↗