Interrelationships of Heterozygosity, Growth Rate and Heterozygote Deficiencies in the Coot Clam, Mulinia Lateralis

Allozyme surveys of marine invertebrates commonly report heterozygote deficiencies, a correlation between multiple locus heterozygosity and size, or both. Hypotheses advanced to account for these phenomena include inbreeding, null alleles, selection, spatial or temporal Wahlund effects, aneuploidy and molecular imprinting. Previous studies have been unable to clearly distinguish among these alternative hypotheses. This report analyzes a large data set (1906 individuals, 15 allozyme loci) from a single field collection of the coot clam Mulinia lateralis and demonstrates (1) significant heterozygote deficiencies at 13 of 15 loci, (2) a correlation between the magnitude of heterozygote deficiency at a locus and the effect of heterozygosity at that locus on shell length, and (3) a distribution of multilocus heterozygosity which deviates from that predicted by observed single-locus heterozygosities. A critical examination of the abovementioned hypotheses as sources of these findings rules out inbreeding, null alleles, aneuploidy, population mixing and imprinting as sole causes. The pooling of larval subpopulations subjected to varying degrees of selection, aneuploidy or imprinting could account for the patterns observed in this study.     

Characterization of microsatellite loci for the detection of temporal genetic shifts within a single cohort of the brown demoiselle,Neopomacentrus filamentosus

Neopomacentrus filamentosus, a common damselfish on the Indo–Australian archipelago, undergoes significant shifts in size and mitochondrial genetic structure upon larval settlement and metamorphosis to juvenile stages. We characterized five polymorphic microsatellite loci in order to study temporal genetic shifts within a single generation of N. filamentosus sampled first as larval settlers then again as demersal juvenile recruits. All loci were extremely polymorphic and exhibited high levels of heterozygosity. While all loci from the larval samples conformed to Hardy – Weinberg expectations, significant heterozygote deficiencies were seen in two loci in the juvenile samples, likely due to extreme size‐selective mortality imposed post‐settlement.     

Extraordinary multilocus genetic organization in mole crickets, Gryllotalpidae

Abstract.— Allozymic diversity at 21 loci was analyzed in 470 individuals of three species of mole cricket superspecies, Gryllotalpa gryllotalpa (two new chromosomal species, G. tali and G. marismortui ) and G. africana in Israel, which are distributed along a southward transect of increasing aridity. Two outstanding findings emerged in G. tali and G. marismortui : (1) genetic polymorphism was high but heterozygosity very low, indicating significant deviations from Hardy-Weinberg expectations; and (2) significant linkage disequilibria at an unprecedented level for outbreeders and remarkable intersite differences. The results may characterize subterranean gryllotalpids worldwide because a single sample of Neocurtilla hexadactyla from Tefé, Amazonia, shows the same features. Significant variation of heterozygote paucity among loci , combined with the biology of the species, rejects the simple explanation of inbreeding or any other single explanatory model. Likewise, direct selection against heterozygotes or specific multilocus associations can explain, but is not necessary nor likely to explain, the observed results in mole crickets. To explain these results, we developed a multiple-factor mathematical model combining niche viability selection, niche choice, and positive assortative mating. This model involves a special case of Wahlund effect and inbreeding. Simulations based on this model showed that a combination of these three mechanisms may produce the observed distribution of alleles, via selection on a few loci, to affect the entire genome organization.     

Temporal genetic variation in a population of the pocket gopher,Geomys bursarius

Genetic variation at eight polymorphic loci was examined in a population of a subterranean rodent, the plains pocket gopher (Geomys bursarius), sampled over a 10-yr period. Two loci exhibited relatively minor changes in gene frequencies, while the remaining loci demonstrated major shifts in predominant alleles, loss of minor alleles, and addition of alleles due to migration. Significant deviations from Hardy-Weinberg expectations, concomitant to heterozygote deficiencies, were observed for several loci. Temporal heterogeneity, as measured by F_ST, was high and comparable to that exhibited by local populations sampled over relatively short periods of time. The high degree of temporal genetic variation is consistent with observations that fossorial rodents occur in locally isolated populations with small effective population sizes that are subject to genetic drift, bottlenecking, and inbreeding.     

Isolation of microsatellite loci from the scleractinian corals,Montastraea cavernosa and Porites astreoides

The ability to assess genetic variation is critical for determining genetic diversity and population structure. In corals, slow evolutionary rates in mitochondrial genomes have left allozymes as the only markers presently available to investigate patterns of intraspecific genetic variation. Characteristics of microsatellites render them more informative than allozymes for such analyses; however, few coral microsatellites are available. This study describes polymorphic microsatellite loci isolated from two scleractinian coral species. Most loci exhibit significant heterozygote deficiencies, likely due to nonrandom mating or Wahlund effects. These markers are being used to investigate gene flow among populations, providing insight into reef connectivity.     

Multilocus structure in Pinus contorta Dougl.

We studied isozyme variation at 21 loci in 66 populations from three subspecies of Pinus contorta Dougl.; 35 in spp. latifolia, 20 in spp contorta and 11 in spp. murrayana. The objectives were to assess gametic disequilibria and multilocus structure. There was considerable differentiation of allele frequencies at 19 polymorphic loci across the 66 populations and within the subspecies. Allele frequencies at many loci correlated with geographic variables. Genetic variability varied considerably among populations within subspecies but the subspecies means were similar. The mean number of polymorphic loci and the mean heterozygosity over 19 polymorphic loci were, respectively, 13 and 0.194 in latifolia, 12 and 0.196 in murrayana, and 12 and 0.180 in contorta. The mean heterozygosity correlated with longitude and altitude across the 66 populations and with latitude in latifolia. Gametic disequilibria were evident in 40 populations; 29 in latifolia, eight in murrayana and three in contorta. Gametic disequilibria correlated with latitude across the 66 populations and with longitude in latifolia. The single-locus F _ST averaged 0.0339 in latifolia, 0.0567 in murrayana, and 0.0764 in contorta. The multilocus F _STM was 0.1227 in latifolia, 0.2926 in murrayana, and 0.3328 in contorta. Multilocus Wahlund and founder effects, migration patterns, and natural selection, probably played significant roles in generating and maintaining the multilocus genetic structure in P. contorta in general and the subspecies latifolia in particular.     

Temporal and Spatial Allozyme Variation in the South American Cactophilic Drosophila antonietae (Diptera; Drosophilidae)

Drosophila antonietae is an endemic South American cactophilic species that uses Cereus hildmaniannus rotting cladodes as breeding sites. We assessed temporal and spatial intrapopulational allozyme variation of two natural populations. Our results suggest that environmental variation (rain precipitation) is probably influencing allozyme temporal variation. Moreover, it seems that D. antonietae does not have intrapopulation structure and has N _ev (variance effective size) 83 and N _ec (number of adult flies that colonize each rotting cladode) = 21. The deficiency of heterozygotes found must be due to null alleles, a temporal Wahlund effect, or selection against heterozygotes. Assortative mating and inbreeding are discarded. This is the first report on allozyme variation in D. antonietae. It gives some insight on intrapopulational genetics through space and time for this species. This is important to understand its general genetic variability and will be essential to future works on the natural history and evolution of this species.     

Flowering Ecotypes of Capsella bursa-pastoris(L.) Medik. (Brassicaceae) Analysed by a Cosegregation of Phenotypic Characters (QTL) and Molecular Markers

Capsella bursa-pastoris(Brassicaceae) is an annual to biennialpredominantly autogamous species distributed worldwide. Usinga linkage map with RAPDs and isozymes we studied quantitativetrait loci (QTL) controlling phenotypic traits in this invasivespecies. To obtain a mapping population we crossed two plantsoccurring in different climatic regions in California, USA (CentralValley and Sierra Nevada) with the most diverse ecotypes (phenotypicparameters) and genotypes (isozyme multilocus genotypes). Ahundred and thirteen F₂individuals were raised and analysedfor segregation at 107 RAPDs, six isozyme loci, and one locusdetermining leaf type. The number, location and magnitude ofgenes underlying 13 traits were determined by using both intervaland composite interval mapping. Two to five QTL affecting onecharacter have been detected. Altogether the 13 quantitativetraits produced 48 QTL. The inheritance patterns of traits rangedfrom those controlled by one QTL with a major effect to thosecontrolled by several QTL with only minor effects. Closely linkedQTL, e.g. onset of flowering with rosette leaf number, wereinterpreted as pleiotropic. Three major QTL account for onsetof flowering. These loci were linked to at least three isozymeloci and several other QTL responsible for developmental traitslike rosette leaf number. Heritability of quantitative traits,segregation of the leaf type, and segregation of the allozymeswas tested in the F₃generation. We conclude that historicalevents alone are insufficient to explain the distribution patternof isozyme multilocus genotypes during the colonization of newregions and habitats. The present evidence indicates that ecotypicadaptation and genetic linkage of isozyme loci with adaptivecharacters are involved. Copyright 2001 Annals of Botany Company Capsella bursa-pastoris, Shepherd’s purse, Brassicaceae, plant invasions, RAPDs, allozymes, Mediterranean multilocus genotype, flowering ecotypes, leaf type, fruit dimensions, cosegregation analysis, QTL     

普通野生稻小种群的交配系统与遗传多样性

小种群的遗传动态是保育遗传学关注的核心问题之一,而种群遗传动态又与交配系统密切相关.普通野生稻(Oryza rufipogon Griff.)是具有重要经济价值的濒危物种,目前其种群规模都较小,研究其小种群交配系统与遗传变异性对普通野生稻的保护具有重要意义.运用7对SSR引物,对采自江西东乡普通野生稻小种群的36份种茎和其中20个家系共计601份子代进行了分析.结果显示:该种群的表观异交率为0.318,多位点法估计(MLTR)的多位点异交率为0.481;50%以上的子代共享亲本,非随机交配明显;东乡普通野生稻种群交配系统属于混合交配类型.比较亲本和子代种群的遗传变异性显示:子代种群比亲本种群遗传变异性更丰富;子代种群的杂合子不足与种群变小自交比例上升有关;而亲本种群杂合子过剩可能与杂合基因型的选择优势有关.这些结果说明创造条件扩大种群规模对普通野生稻的原生境保护显得尤为重要.     

Mating system and inbreeding depression in whitebark pine (Pinus albicaulis Engelm.)

Mating system and inbreeding depression in quantitative traits of whitebark pine (Pinus albicaulis Engelm.) was determined using isozymes and a seedling common garden experiment. Simultaneous isozyme analysis of embryo and haploid megagametophyes from progeny arrays of families in three distinct geographic regions (Oregon, Montana, and southern British Columbia) was used to estimate parental and progeny inbreeding coefficients, as well as regional and family mean multilocus outcrossing rates (t _m). Quantitative trait family means of seedlings from the same families growing in two temperature treatments in a common garden experiment were regressed on the estimated inbreeding coefficient to determine the presence and magnitude of inbreeding depression. Regional estimates of t _m ranged from 0.73 to 0.93, with a mean over all regions of 0.86. Family mean t _m values indicated predominant outcrossing; however, some individuals experienced substantial inbreeding. The Oregon region had a significant excess of heterozygotes in the parental generation relative to Hardy–Weinberg equilibrium, while both the Oregon and southern BC regions had a heterozygote deficiency in progeny, suggesting selection against inbred individuals. Biomass in the ambient temperature treatment for the southern BC region was the only trait significantly related to inbreeding coefficient. The mean inbreeding coefficient for this region was 0.25, and based on this relationship, mean predicted biomass would be reduced by 19.6% in this region if inbred individuals are not removed by selection. The estimated outcrossing rate of whitebark pine is slightly lower than most wind-pollinated conifers, and while most individuals are highly outcrossing, some experience substantial inbreeding.     

Genetic variation and population structuring in two brooding coral species (<Emphasis Type="Italic">Siderastrea stellata</Emphasis> and <Emphasis Type="Italic">Siderastrea radians</Emphasis>) from Brazil

Siderastrea stellata and S. radians are scleractinian coral species that present a remarkable overlap of diagnostic characteristics and sympatric distribution. Moreover, both are viviparous with similar reproductive strategies and with a gregarious larval behavior. Samples of both species from the Brazilian coast were analyzed using 18 isozymic loci to quantify their genetic variability and populational structure. Results confirmed species identity, high intrapopulational variability and revealed moderate genetic structuring among all samples (S. stellata: F (ST) = 0.070; S. radians: F (ST) = 0.092). Based on genotypic diversity analysis, there was evidence that local recruitment may have a minor role in the populations (mean, G ( o ):G ( e )= 1.00 +/- 0.0003 SD for S. stellata and 0.99 +/- 0.0023 SD for S. radians). Deviations towards heterozygote deficiencies found in both Siderastrea species could be explained by the Wahlund effect, since there was evidence that populations might be composed of colonies of different ages. In S. radians it is also likely that there is some inbreeding occurring in the studied populations. Despite the brooding pattern and the gregarious larval behavior, our data suggest the occurrence of gene flow along the Brazilian coast. This is the first study on population genetics of Brazilian reef corals.     

Nine polymorphic microsatellite loci from the psyllid Cacopsylla pruni (Scopoli), the vector of European stone fruit yellows

Cacopsylla pruni is the vector of European stone fruit yellows, a quarantine disease of Prunus trees. Nine polymorphic microsatellite markers were developed from enriched DNA libraries. Allelic variability was assessed in a collection of 149 females obtained from five localities covering a large geographical area in France. The number of detected alleles ranged from 8 to 37. Within the localities, observed and expected heterozygosities averaged across loci ranged from 0.39 to 0.55, and from 0.68 to 0.81, respectively. A heterozygote deficiency was detected for almost all loci, possibly due to a high null allele frequency. Other possible causes of the homozygote excess (mode of reproduction, inbreeding, assortative mating or Wahlund effect) are discussed. These variable microsatellite loci can provide tools to assess overall genetic variation in this important vector species. They will be used to search for population structure and migration patterns of C. pruni.     

Temporal genetic changes between cohorts in a natural population of a marine fish, Diplodus sargus

Temporal changes at 17 allozyme loci in the Diplodus sargus population of Banyuls sur Mer (Mediterranean Sea, France) were monitored within a single population among ten year‐classes (cohorts) sampled over a 6‐month period. The genetic survey was combined with evaluation of the demographic structure of the population by determining variation of abundance between cohorts. The population showed variation in abundance among cohorts ranging from 16 to 214 individuals. Significant divergences in genetic structure were observed between cohorts (P < 0.0001) despite very low values of F_ST (multilocus F_ST over all cohorts = 0.0018). The heterozygosity of each cohort, as well as the F_IS values, was significantly correlated with the abundance of each cohort, with abundant cohorts showing lower heterozygosity and a significant deficit of heterozygotes (positive F_IS values). Finally, multilocus temporal genetic variance (F_k) computed between successive cohorts was higher in low abundance cohorts. Change of heterozygosity between cohorts, distribution of year‐class genetic structure, and change in the genetic structure within a cohort appear to be affected mostly by the abundance of the cohort and are therefore driven by genetic drift. We propose that the Diplodus sargus cohorts are built up from the mixing of families during the pelagic stage or later during recruitment, and that the decrease in heterozygosity leading to a deficit of heterozygotes is characteristic of a Wahlund effect. Such a Wahlund effect would derive from the mixing of the progeny of families made up of few individuals, but exhibiting high fecundity and high variability of reproductive success. Therefore, although cohorts derived from poor recruitment would only group a few families and would exhibit limited deficit of heterozygotes (higher heterozygosity values), they would lead to high genetic drift and appear more divergent (higher mean temporal genetic variance) than cohorts with high abundance. While not demonstrating directly the family structure of marine populations, our survey provides evidence of highly structured populations. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 76 , 9–20.     

家蚕AFLP连锁框架图谱的构建

利用改进的AFLP分子标记方法,对家蚕Bombyx mori回交一代BC1群体进行连锁图谱的构建。经17组AFLP引物的选择性扩增,共得到430个多态位点,卡方检验后有253个为有效位点。利用Mapmaker/Exp (Version 3.0b)软件作连锁分析,其中163个标记分属28个连锁群,连锁群标记数变化范围是2～28个,平均每个连锁群标记数为5.8个,该图覆盖的基因组长度为2.998.9cM(图距单位),连锁群长度变化范围为4.5～652.8 cM,连锁群的平均长度为107.1 cM,平均图距为4.5～36.7 cM。     

突尼斯非洲跳鼠(Jaculus jaculus)和埃及跳鼠(J.orientalis)群体的等位酶多态及遗传分化(英文)

运用16种酶蛋白编码的23个遗传座位对突尼斯非洲跳鼠(Jaculus jaculus)和埃及跳鼠(J. orientalis)自然群体的遗传变异和分化进行了电泳分析。结果表明,与其他啮齿动物等哺乳动物的相关数据比较,发现这两个种群体的遗传变异水平较低。非洲跳鼠群体的观测杂合度 (H_obs) 为0.08—0.19,多态座位百分比(P)为26.2%—45.2%,每个座位的平均等位基因数(A)为1.1—1.4;埃及跳鼠的H_obs为0.10—0.15,P为29.3%—44.1%,A为1.1—1.7。两个种群体各自的遗传分化程度较低(非洲跳鼠和埃及跳鼠的F_st分别为0.0017和0.0019)。而两个种群体间的F_st为0.607（P<0.05）,表明两个种之间高度的遗传分化。本研究支持这两个种分类地位的合法性,并强调了地理因素（环境类型和生物气候阶段）对两个种遗传结构的影响。     

Effect of rate of inbreeding on inbreeding depression in Drosophila melanogaster

Summary This experiment was designed to study the relationship between rate of inbreeding and observed inbreeding depression of larval viability, adult fecundity and cold shock mortality in Drosophila melanogaster. Rates of inbreeding used were full-sib mating and closed lines of N=4 and N=20. Eight generations of mating in the N=20 lines, three generations in the N=4 lines and one generation of full-sib mating were synchronised to simultaneously produce individuals with an expected level of inbreeding coefficient (F) of approximately 0.25. Inbreeding depression for the three traits was significant at F=0.25. N=20 lines showed significantly less inbreeding depression than full-sib mated lines for larval viability at approximately the same level of F. A similar trend was observed for fecundity. No effect of rate of inbreeding depression was found for cold shock mortality, but this trait was measured with less precision than the other two. Natural selection acting on loci influencing larval viability and fecundity during the process of inbreeding could explain these results. Selection is expected to be more effective with slow rates of inbreeding because there are more generations and greater opportunity for selection to act before F=0.25 is reached. Selection intensities seem to have been different in the three traits measured. Selection was most intense for larval viability, less intense for fecundity and, perhaps, negligible at loci influencing cold shock mortality.     

ALLOZYME VARIATION AMONG POPULATIONS OF BIOMPHALARIA PFEIFFERI (KRAUSS, 1848) (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis was used to study genetic diversityamong the freshwater, hermaphroditic snail Biomphalaria pfeifferiin Cameroon. Four of 19 loci studied were polymorphic. The allozymesfor two loci, aspartate aminotransferase-1 (AAT-1) and isocitratedehydrogenase (IDH), showed distribution patterns related toclimatic zones. AAT-1¹⁰⁰ and IDH¹⁰⁰ occurred predominantly inthe Tropical Climatic Zone, located in the northern half ofthe country. AAT-1¹⁴⁰ and IDH⁹⁰ were common in the EquatorialZone. Intrapopulation variations occurred in 5 of 19 populationsampled. Only one population, polymorphic for AAT-1 locus, waspanmictic. The other 4 populations were not in Hardy-Weinbergequilibrium. The deficiency of heterozygotes in those populationsis unusual for a member of the genus Biomphalaria and may bedue to a predominance of self-fertilization. Other explanationsinclude ecological factors or the presence of segregated sub-populationsthat do not outcross. This latter hypothesis is supported bythe total absence of heterozygotes for PGM alleles. (Received 24 August 1989; accepted 14 September 1989)     

Adult survival and microsatellite diversity in possums: effects of major histocompatibility complex-linked microsatellite diversity but not multilocus inbreeding estimators

Adult survival is perhaps the fitness parameter most important to population growth in long-lived species. Intrinsic and extrinsic covariates of survival are therefore likely to be important drivers of population dynamics. We used long-term mark-recapture data to identify genetic, individual and environmental covariates of local survival in a natural population of mountain brushtail possums (Trichosurus cunninghami). Rainfall and intra-individual diversity at microsatellite DNA markers were associated with increased local survival of adults and juveniles. We contrasted the performance of several microsatellite heterozygosity measures, including internal relatedness (IR), homozygosity by loci (HL) and the mean multilocus estimate of the squared difference in microsatellite allele sizes within an individual (mean d ²). However, the strongest effect on survival was not associated with multilocus microsatellite diversity (which would indicate a genome-wide inbreeding effect), but a subset of two loci. This included a major histocompatibility complex (MHC)-linked marker and a putatively neutral microsatellite locus. For both loci, diversity measures incorporating allele size information had stronger associations with survival than measures based on heterozygosity, whether or not allele frequency information was included (such as IR). Increased survival was apparent among heterozygotes at the MHC-linked locus, but the benefits of heterozygosity to survival were reduced in heterozygotes with larger differences in allele size. The effect of heterozygosity on fitness-related traits was supported by data on endoparasites in a subset of the individuals studied in this population. There was no apparent density dependence in survival, nor an effect of sex, age or immigrant status. Our findings suggest that in the apparent absence of inbreeding, variation at specific loci can generate strong associations between fitness and diversity at linked markers.     

Field and experimental evidence of preferential selfing in the freshwater mollusc Lymnaea truncatula (Gastropoda, Pulmonata)

We have conducted a thorough study of the mating system of Lymnaea truncatula, the intermediate host of the liver fluke, using three approaches: (i) a population genetics study, (ii) controlled pairings in the laboratory and (iii) a progeny-array analysis. The population genetics study revealed high levels of inbreeding in the studied populations, with strong clues that the extensive heterozygote deficiencies observed are due to selfing. However, Wahlund effects may also arise due to recolonisations from different source populations after bottleneck events. A breeding experiment helped to disentangle the mating system and the Wahlund effects, and showed that high levels of selfing occurred in isolation and in controlled pairings. However, the progeny-array analysis performed after a high-density culturing of the snails suggests that substantial outcrossing may also occur.     

Adaptation at Specific Loci. V. Metabolically Adjacent Enzyme Loci May Have Very Distinct Experiences of Selective Pressures

The polymorphic phosphoglucomutase (PGM) and glucose-6-phosphate dehydrogenase (G6PD) loci have been studied in parallel to experimental work on the phosphoglucose isomerase (PGI) polymorphism in Colias butterflies. PGI, PGM and G6PD are also autosomal in Colias. PGM and G6PD are loosely linked (and represent the first identified autosomal linkage group in Colias); they assort independently from PGI. Recombination occurs in both sexes. Neither PGM nor G6PD shows large, consistent differences in flight capacity through the day among its genotypes, as PGI does. PGM shows some change of allele frequencies, and match to Hardy-Weinberg expectation, with air temperature in middle and latter parts of the season, but not early in the season. G6PD may show some heterozygote excess over Hardy-Weinberg expectation early in the day, but more testing is needed. No evidence for differential survivorship was seen at PGM or G6PD, in contrast to PGI. At the PGM and G6PD loci, male heterozygotes are advantaged in mating with females, but without the evidence of female choice which occurs for PGI. These effects are not correlated among the three loci. There is no assortative mating at G6PD (nor at PGI). There is minor positive assortative mating of PGM heterozygotes, but it is too weak to account for the PGM-genotype-specific male mating advantage. No trends of multilocus genotype frequencies involving PGI are seen. Certain PGM-G6PD two-locus genotypes are over-represented, and others under-represented, in wild adult samples, particularly among males and uniformly among successfully mating males. Our results emphasize that enzyme loci sharing a substrate need not have common experience of the existence or strength of natural selection, and suggest initial food-resource processing and allocation as a possible context for fitness-related effects of the PGM and G6PD polymorphisms.