EFFECTS OF POPULATION BOTTLENECKS ON GENETIC DIVERSITY AS MEASURED BY ALLOZYME ELECTROPHORESIS

Low levels of allozyme heterozygosity in populations are often attributed to previous population bottlenecks; however, few experiments have examined the relationship between heterozygosity and bottlenecks under natural conditions. The composition and number of founders of 55 experimental populations of the eastern mosquitofish (Gambusia holbrooki), maintained under simulated field conditions, were manipulated to examine the effects of bottlenecks on three components of allozyme diversity. Correlations between observed and expected values of allozyme heterozygosity, proportions of polymorphic loci, and numbers of alleles per locus were 0.423, 0.602, and 0.772, respectively. The numbers of polymorphic loci and of alleles per locus were more sensitive indicators of differences in genetic diversity between the pre-bottleneck and post-bottleneck populations than was multiple-locus heterozygosity. In many populations, single- and multiple-locus heterozygosity actually increased as a result of the founder event. The weak relationship between a population's heterozygosity and the number and composition of its founders resulted from an increase in the variance of heterozygosity due to drift of allele frequencies. There was little evidence that selection influenced the loss of allozyme variation. When it is not possible to estimate heterozygosity at a large number of polymorphic loci, allozyme surveys attempting to detect founder events and other types of bottlenecks should focus on levels of locus polymorphism and allelic diversity rather than on heterozygosity.     

Genetic variation of polygenic characters and the evolution of genetic degeneracy

The classical model of mutation-selection balance for quantitative characters sums the effects of individual sites to determine overall character value. I develop an alternative version of this classical model in which character value depends on the averaging of the effects of the individual sites. In this new averaging model, the equilibrium patterns of variance in allelic effects and character values change with the number of sites that affect a character in a different way from the classical model of summing effects. Besides changing the patterns of variance, the averaging model favours the addition of loci to the control of character values, perhaps explaining in part the recent observation of widespread genetic degeneracy.     

A Comprehensive Study of Genic Variation in Natural Populations of Drosophila melanogaster. II. Estimates of Heterozygosity and Patterns of Geographic Differentiation

A study of genic variation in natural population of D. melanogaster was undertaken (1) to obtain a better estimate of heterozygosity by sampling a relatively large number of gene loci and (2) to identify different groups of polymorphic loci whose variation patterns might suggest different kinds of selection forces. A total of 117 gene loci (coding for 79 enzymes and 38 abundant proteins) were studied in 15 geographically distant populations originating from different continents. The findings of this study are as follows: (1) of the 117 gene loci studied, 61 are polymorphic and 56 are uniformly monomorphic everywhere. (2) An average population is polymorphic for 43% of its gene loci and an average individual is heterozygous for 10% of its gene loci. These estimates are remarkably similar among populations. (3) The average within-locality heterozygosity (H(S)) for polymorphic loci is uniformly distributed over the range of heterozygosity observed; i.e. , given that a locus has any local variation, it is nearly as likely to have a lot as a little. (4) The distribution of F(ST) (fixation index) is strongly skewed, with a prominent mode at 8-10% and a long tail of high values reaching a maximum of 58%. Two-thirds of all loci fall within the bell-shaped distribution centered on an F(ST) of 8-10%, a result compatible with the notion that they are experiencing a common tendency toward small interlocality differences owing to extensive gene flow among populations. (5) The distribution of total heterozygosity (H(T)) has a prominent bimodal distribution. The lower mode consists of loci with single prominent allele and a few uncommon ones and the upper mode consists of clinally varying loci with a high F(ST ) (e.g., Adh and G6-pd), loci with many alleles in high frequency (e.g., Ao and Xdh) and loci with two alleles in high frequency in all populations but, with little interpopulational differentiation (e.g., Est-6 and alpha-Fuc). The loci in the lower mode are probably under purifying selection; a large proportion of those in the latter mode may be under balancing selection. (6) Comparison of genic variation for loci located inside vs. outside inversions, comparison of F(ST) for inversions and their associated genes, and comparison of F(ST) and map position for pairs of loci all suggest that, while linkage has some influence, it does not seem to constrain the pattern of variation that a locus may develop. (7) Eighteen polymorphic loci show latitudinal variation in allele frequencies which are consistent in populations from different continents. (8) Estimates of Nei genetic distance between population pairs are generally low between populations on the same continent and high between populations on different continents. There are two important exceptions: population pairs for which both localities are in the temperate zone show no relationship to distance, and in cases where both populations are tropical or subtropical, the genetic distance is higher than for the temperate-tropical comparisons and seem even higher than one would expect from the geographic distance separating them. The latter observation suggests that either geographic separation outweighs differences in environment in determining the genetic composition of a population or that all tropical populations are not experiencing the same environment.-The results are discussed in relation to the neutralist-selectionist controversy of genic variation and two important conclusions are drawn: First, there is a negative correlation between the number of loci sampled and the resulting heterozygosity. This means that available estimates of heterozygosity, 85% of which are based on 30 or fewer loci, are high and hence not appropriate for making between-taxa comparisons. Secondly, there is a group of loci, comprising one-third of polymorphic loci (or about 15% of all loci studied), that is distinguishable by different patterns of variation within and among populations. Most of these loci have clinal variation which is consistent with the hypothesis that their genetic variation is maintained by balancing selection.     

Anthropometric multivariate structure and dermatoglyphic peculiarities in biochemically and morphologically different heterozygous groups

Multivariate analysis of the relationship between degree of heterozygosity at four blood group loci and the morphological variability in a human population was carried out. Additionally, the possibility that dermatoglyphic patterns correlate with biochemical and anthropometric variables was also investigated. A strong and significant increase in the frequency of morphologically multimodal individuals was observed, which paralleled the heterozygosity level. Discriminant analysis, by quantitative characters, of the closest pair of biochemically different samples yielded a satisfactory discrimination. Multiple correlations of each variable with all the others (18 traits), the communality of characters, the index of integration, and the Mahalanobis distances of the factor scores for each individual (all extracted from principal component analysis) were all indicative of the different multivariate structures of homo- and heterozygous individuals and thus supported the hypothesis that heterozygotes tend to cluster near the center of the joint multivariate distribution. The dermatoglyphic patterns showed a certain relationship with the morphological makeup of individuals. Correlations between biochemical heterozygosity at blood group loci and patterns of digital dermatoglyphics were rather irregular.     

Characterization of nine polymorphic microsatellite markers in sprat (Sprattus sprattus L.)

Nine polymorphic microsatellites were isolated from sprat (Sprattus sprattus) using a microsatellite enrichment protocol and selective hybridization with a biotinylated (AC)(12) probe. The loci showed different variation patterns in a Baltic Sea population (44 individuals) with mean number of alleles at 12.7 and mean observed heterozygosity at 0.78. These microsatellite loci are expected to be used for taxonomic considerations in sprat, stock differentiation and population genetic analysis.     

ASSOCIATION BETWEEN QUANTITATIVE TRAITS AND ALLOZYME HETEROZYGOSITY IN A TETRASOMIC SPECIES: DACTYLIS GLOMERATA

Tetraploid individuals of orchardgrass (Dactylis glomerata L. subsp. hispanica Roth.) sampled from a natural population were used to evaluate the correlation between both single and multilocus heterozygosity at 7 enzyme loci, and several quantitative traits (plant size at time of collection, leaf weight, and panicle number in experimental trials). Four hundred and forty-eight plants were studied at the 7 loci and 288 of these individuals were scored for an additional eighth locus. Five genotype classes (monogenic, simplex, and duplex digenic, trigenic, and tetragenic) were distinguished according to their heterozygosity level. Multilocus heterozygosity showed a significant positive correlation with both leaf and panicle yield in experimental conditions, but not with original plant size, which was found to be markedly influenced by environmental microheterogeneity. Multilocus heterozygosity, estimated from both the number of heterozygous loci and the number of distinct alleles per locus, had a significant influence on plant performance. Individual locus effects were positive and significant at two loci (GOT1 and PX1). Panicle number increased regularly with heterozygosity level (from monogenics to tetragenics) at the GOT1 locus, as did leaf weight and panicle number at the PX1 locus. Such variation would be predicted by overdominance at these loci or at linked loci. Significant relationships between leaf yield and heterozygosity level at the GOT1 locus distinguished the homozygotes from the heterozygotes (of any class) and was thus more consistent with inbreeding effects. No significant differences were observed among the five genotype classes for any quantitative trait at the six remaining loci. At both the GOT1 and PX1 loci, heterozygosity had a significant independent effect on leaf weight and panicle number even when the correlation between these traits was removed by analysis of covariance.     

Is there a pattern of gene differentiation in the Indian populations

Summary Indian populations divided into a number of endogamous groups consisting of different castes, languages, religions, and tribes provide unique opportunities for examining the extent and nature of genetic differentiation at a microevolutionary stage. The genetic relationships between some of these Indian population groups have been examined using electrophoretic data from several biochemical loci in a gene diversity analysis. Does this type of analysis provide any insight into what causes such gene differentiation? What patterns of genetic variation emerge from these empirical findings? Answers are sought by relating the observed heterozygosity, genetic distance, and allied statistics to a mutation-drift hypothesis. The statistics used are: (1) interlocus mean and variance of heterozygosity, (2) mean and variance of genetic distance, and (3) correlation of heterozygosity and gene identity. The observed relationships between these sets of statistics agree well with the ones predicted by the hypothesis that different alleles at protein loci are selectively equivalent and gene frequency change occurs predominantly due to genetic drift     

Assessing the allozyme variation in cultivars and Swedish landraces of rye (Secale cereale L.)

Genetic interpretation and diversity of 9 isozyme loci have been estimated in 7 improved varieties and 19 landraces from Sweden by means of starch gel electrophoresis. The isozyme systems were ACO, DIA, GPI, MDH, PGD and PGM. For the statistic analysis we used the following measures: average number of alleles per locus, percentage of polymorphic loci, average heterozygosity direct count and average heterozygosity Hardy-Weinberg expected unbiased estimate. The measures were made on species and population levels. The distribution of the total genetic diversity among populations was also calculated. To illustrate the genetic relationships among populations, genetic distances were measured and principal component analysis performed. As expected in a cross-pollinated crop we found high genetic diversity and a larger variation within than among the populations. Somewhat unexpectedly, however, we found that the currently used varieties have the same high level of heterozygosity as the landraces but in the dendrogram the two groups are separated. The dendrogram showed three main clusters. The large cluster included 21 populations and the two small clusters were clearly distinguishable from the rest. The landrace spring-type could not be separated from the landraces winter-type, but we did detect a difference between different spring types. A few populations had unique alleles for certain loci.     

Development and characterization of microsatellite loci for lotus (Nelumbo nucifera)

This paper reports the development of microsatellite primers for Nelumbo nucifera Gaerten. By screening genomic libraries enriched with 10 kinds of probes, Seventeen polymorphic loci were isolated and primers were designed. Polymorphism of these 17 loci was assessed in 24 individuals. All the 17 loci are polymorphic and the number of alleles ranged from two to seven. Observed heterozygosity and expected heterozygosity ranged from 0.0000 to 0.9176 and from 0.2837 to 0.7917 respectively. These microsatellite loci should be useful for studying the genetic diversity of N. nucifera.     

The genetic basis of developmental stability in Apis mellifera II. Relationships between character size, asymmetry and single-locus heterozygosity

Haploid male and diploid female honey bees, Apis mellifera, from colonies headed by queens polymorphic at the malate dehydrogenase (MDH) locus were examined for the influence of MDH genotype and heterozygosity on the size and asymmetry of six morphological characters. Although there were significant differences among MDH genotypes for mean character size within colonies, these effects were inconsistent between colonies. There were no significant relationships between MDH genotype or heterozygosity and asymmetry, indicating that genetic variation at this locus has no impact on developmental stability in this species.     

Isolation and characterization of 12 novel microsatellite loci for the red panda (Ailurus fulgens)

The information on dispersal patterns and mating systems of red pandas is quite important for the understanding of the genetic diversity and divergence of this species. And microsatellite marker is an ideal tool to analyze dispersal patterns and mating systems. Thus, we describe in this paper the isolation and characterization of 12 microsatellite loci in the red panda from genomic DNA-enriched libraries. These loci were highly polymorphic with numbers of alleles per locus in 24 individuals ranging from 2 to 14, observed heterozygosity from 0.143 to 0.864 and expected heterozygosity from 0.297 to 0.872. All loci except for RP6 locus followed Hardy–Weinberg expectations. No significant linkage association was found among all these loci. The 12 novel polymorphic microsatellite loci will be of use in studying dispersal patterns and mating systems of red pandas.     

椭圆叶花锚简化基因组的SSR信息分析及SSR引物开发

基于RAD-seq简化基因组测序技术获得椭圆叶花锚（Halenia ellipitica D.Don）简化基因组水平上的序列信息并开发SSR分子标记。利用SR search软件检测而得到双端各有至少100 bp的五种类型的SSR（二、三、四、五、六核苷酸）位点共6 201个,并成功设计其中3 865个SSR引物。在能成功设计引物的SSR位点中,三核苷酸SSR位点最多;重复序列长度包括17种（12~36 bp）;重复序列的基序共达316种,其中五核苷酸基序种类最多（91种）。从中挑选65对可对应五种SSR类型的引物,经梯度PCR检验后,利用椭圆叶花锚的4个居群32个个对可扩增的引物进行PCR和聚丙烯酰氨凝胶电泳检测,其中14对引物能在绝大多数个体中扩增,且13对具多态性。13个多态性位点的等位基因数量均值为5.462,多态性较高且不连锁（P<0.05）;其中10个位点在多数居群中偏离哈迪温伯格平衡（P<0.01）且存在较高的纯合子数量（观测杂合度H_o均值0.226）;近交系数在-0.443~1,均值为0.656;基因流N_m为0.474。     

Isolation and characterization of six polymorphic microsatellite loci in the western flower thrips Frankliniella occidentalis (Insecta,Thysanoptera)

Highly polymorphic microsatellite markers can provide important demographic information on founder events and range expansion following initial introduction of invasive insect species. Six microsatellite loci were isolated from an enriched DNA library in order to study the invasion patterns of the western flower thrips, Frankliniella occidentalis. All loci tested were found to be polymorphic and successfully amplified in all individuals. The number of alleles per locus ranged from five to nine and heterozygosity ranged from 45 to 73%. Some of the loci were also successfully amplified in other thrips species.     

How do population genetic parameters affect germination of the heterocarpic species Atriplex tatarica (Amaranthaceae)?

Background and Aims

The heterocarpic species Atriplex tatarica produces two types of seeds. In this study, how basic population genetic parameters correlate with seed germinability under various experimental conditions was tested.

Methods

Population genetic diversity was ascertained in eight populations of A. tatarica by assessing patterns of variation at nine allozyme loci. Germinability of both seed types from all sampled populations was determined by a common laboratory experiment under different salinity levels. Basic population genetic parameters, i.e. percentage of polymorphic loci, average number of alleles per locus and observed heterozygosity were correlated with observed population germination characteristics.

Key Results

Atriplex tatarica possesses a remarkable heterocarpy, i.e. one type of seed is non-dormant and the other shows different dormancy levels in relation to experimental conditions. Significant negative correlations have been detected between germination of both seed types and the coefficient of inbreeding, and a significant negative correlation between germination of dormant seeds and other population genetic parameters, i.e. percentage of polymorphic loci and average number of alleles per polymorphic locus. Moreover, populations from the region characterized by a shorter growing season manifested higher germinability, i.e. had lower dormancy, than those from the lower-latitude one.

Conclusions

In general, germination of non-dormant seeds is probably not under strong genetic control. Hence, they germinate as soon as conditions are favourable, thus ensuring survival in the short term, but populations risk local extinction if conditions become adverse (i.e. a high-risk strategy). In contrast, germination of the dormant type of seeds is under stronger genetic control and is significantly correlated with basic population genetic parameters. These seeds ensure long-term reproduction and survival in the field by protracted germination, albeit in low quantities (i.e. A. tatarica also adopts a low-risk strategy).Key words: Amaranthaceae, Atriplex, inbreeding depression, population genetics, seed dimorphism     

Isolation and characterization of 15 microsatellite loci in four endangered Amentotaxus species (Taxaceae)

? Premise of the study: Fifteen microsatellite loci were developed in an endangered species, Amentotaxus formosana, and were tested in an additional three species, A. argotaenia, A. yunnanensis, and A. poilanei, to evaluate the population structure for conservation efforts and reconstruct the phylogeographic patterns of this ancient lineage. ? Methods and Results: Polymorphic primer sets were developed from A. formosana; the number of alleles ranged from two to 10, with an observed heterozygosity ranging from 0 to 0.60. All of the loci were found to be interspecifically amplifiable. ? Conclusions: These polymorphic and transferable loci will be potentially useful for future studies that will focus on identifying distinct genetic units within species and establishing the phylogeographic patterns and the process of speciation among closely related species.     

Population structure and quantitative characters

A migration matrix model is used to investigate the behavior of neutral polygenic characters in subdivided populations. It is shown that gametic disequilibrium has a large effect on the variance among groups but none at all on its expectation. The variance of among-group variance is substantial and does not depend on the number of loci contributing to variance in the character. It is just as large for polygenic characters as for single loci with the same additive variance. This implies that one polygenic character contains exactly as much information about population relationships as one single-locus marker. The theory is compared with observed differentiation of dermatoglyphic and anthropometric characters among Bougainville islanders.     

New polymorphic microsatellite loci for the Greek smooth newt, Lissotriton vulgaris graecus, and their utility in the nominotypical subspecies

Eight polymorphic microsatellites were isolated from the Greek smooth newt (Lissotriton vulgaris graecus) using a microsatellite enrichment protocol and selective hybridization with a biotinylated (AC)(11) probe. The loci showed different variation patterns in a single breeding population (32 individuals) with mean number of alleles at 5.0 and mean observed heterozygosity at 0.520. The amplification success also in the nominotypical subspecies favours the use of these microsatellite loci in population genetic analyses as well as in the study of contact zones between smooth newt subspecies.     

New polymorphic microsatellite loci for the Macedonian crested newt, Triturus macedonicus, and cross-priming testing in four other crested newt species

Eleven microsatellites have been characterized for Triturus macedonicus. Nine loci showed different variation patterns in a sample of 40 individuals from a single breeding pond in Zagori province (Greece), with an average number of 4.2 alleles per locus and an expected heterozygosity ranging from 0.1199 to 0.8079. Distinct cross-priming amplification rates were recovered on four additional crested newt species. Two monomorphic T. macedonicus loci were polymorphic in other Triturus species. The microsatellites developed herein could be a useful intraspecific genetic tool to undertake fine-scale population genetic analyses as well as in the study of contact zones between crested newt species.     

Development and characteristics of microsatellite markers in Pinus merkusii

Pinus merkusii is an important industrial species that is distributed only in Southeast Asia. We isolated 10 microsatellite markers from this species using a dual‐suppression‐polymerase chain reaction technique. Of these markers, five loci were codominant and polymorphic. The number of alleles per locus ranged from three to six and the expected heterozygosity ranged from 0.389 to 0.728. These microsatellite markers will be available for analysis on population genetics and mating patterns.     

微卫星DNA标记分析德国镜鲤的遗传潜力

结合体重、体长、体高等数量性状, 用30个微卫星分子标记, 评估了3个德国镜鲤群体的遗传潜力, 共检测到287个等位基因, 559种基因型, 片段长度109~400 bp, 有效等位基因数1.1014~6.4665, 观察杂合度0.0968~0.9892, 期望杂合度0.0926~0.8554, 位点多态信息含量0.08787~0.8559, 其中中度多态(0.25≤PIC≤0.5)13个, 高度多态(PIC≥0.5)13个。统计结果显示: 3个群体的遗传潜力处于中度水平, 双来养殖群体的遗传潜力比换新和松浦繁殖群体低。同时, 用30个基因座的不同等位基因和基因型与双来群体的体重、体长、体高进行了连锁分析, 得到2个与镜鲤体长相关的微卫星分子标记(HLJ319, HLJ693)和1个与体高相关的位点(HLJ677), 与体长相关的1个位点(HLJ693)还与体重连锁。将此遗传标记在鲤鱼重组自交系中验证, 结果显示: 主要经济性状相连锁的3个遗传标记中HLJ319与鲤鱼体长性状的QTL定位结果基本一致。