Variation of Loci Encoding Homologous Enzymes in the Evolutionary Series of Vertebrates

A study of variability of 11 allozyme loci (sAat, G3pdh, Gpi, sIdh, Ldh-A, Ldh-B, sMdh, sMe, sSod, Pgdh, and Sdh) in the evolutionary series of vertebrates from Cyclostomat to Mammalia revealed that (1) in vertebrates, these loci encoding multimeric enzymes are characterized by different heterozygosity levels, the extremes of which (represented by loci Ldh-A and Pgdh) differ from each other more than by a factor of 4; (2) classes of vertebrates markedly differed from one another in genetic variation; lower Tetrapoda are characterized by the highest level of genetic polymorphism, the classes representing the margins of the phyletic line—primitive (Cyclostomata and Chondrchthyes) and advanced (Aves and Mammalia)—have minimum heterozygosity levels, whereas Osteichthyes are characterized by intermediate heterozygosity level; (3) in the evolutionary series of vertebrates, heterozygosity varies rather independently in the groups of loci characterized by low, medium, and high variability. These patterns are explained in the context of intraorganismic factors: integration of mono- and polygenic traits (primarily, body size and ontogeny rate) and evolutionary specialization.     

A comparative analysis of allozyme variability in vertebrate animals]

Comparative analysis of levels of the allozymic variation in the vertebrates is conducted using evaluation of a) within population heterozygosity of ferment coding homologous loci, b) average (per each species) allele frequencies of homologous loci, and c) overall sample of the loci being analyzed. It is established that amphibians and reptiles are characterized by the highest, birds and mammals--by the lowest, and fishes--by the middle level of genetic diversity. The diversity of genetic systems, if the constant heterozygosity of duplicated loci is taken into consideration, decreases from fishes to mammals and from cold-blooded to warm-blooded vertebrates. This tendency could be considered as an extrapolation of the "progressive specialization" rule on molecular level. Three basic factors determining certain level of genetic diversity of a species are acknowledged position in the phylogenetic system, population structure, and level of introgressive hybridization. From methodological viewpoint, evaluation of the genetic diversity by homologous loci seems to be most valid in comparative studies.     

Characterization of long transcribed microsatellites in Helianthus annuus (Asteraceae)

? Premise of the study: Research on the evolutionary role of exonic microsatellites currently lacks an understanding of the evolutionary pressures that promote or limit their expansion. Contrasting levels of variability and genetic structures at anonymous and transcribed microsatellite loci of varying lengths are likely to provide useful insights regarding the relative strength of selection acting on different classes of microsatellites. We have developed primers for long transcribed microsatellites in Helianthus annuus to make these comparisons. ? Methods and Results: Eight relatively long microsatellites from sequences in the expressed sequence tag database of H. annuus were characterized. A total of 63 individuals from three populations in Kansas were genotyped. The number of alleles per locus ranged from four to 11 with an average observed heterozygosity of 0.723. ? Conclusions: Our study has generated suitable tools for studying the population genetics of long transcribed microsatellites that are potentially influenced by selection.     

Coarse-grained population structure in Central European sculpin (Cottus gobio L.): secondary contact or ongoing genetic drift?

Population genetic affinities of 261 European sculpins Cottus gobio L. across the Rhenanian-Danubian and the Rhenanian-Rhónian watersheds were assessed by horizontal agarose-gel electrophoresis of up to 20 allozyme systems (encoded by 29 genetic loci). Polymorphism P_mean= 0.0689 (range: 0.000-0.1379), and heterozygosity H_e(mean)= 0.0167 (range: 0.000-0.0507) indicated low genetic variability within local stocks from single streams. Significant genetic distances D_mean= 0.1917 ± 0.0336 (D_max= 0.2407), based on differential fixation at 3–6 loci (Acp-1**, Pgdh**, Fh**, Est-1**, Gpi-2**, and Pgm-1**) distinguished populations from the Neckar catchment basin from those of the Hochhrein-Oberrhein and Danube basins. Differential fixation of alleles and pronounced genetic distances also separated sculpins of the Rh6nian tributary Doubs from Neckarian populations (D_men= 0.2131 ± 0.0033; Ah**, Acp-1**, Fh**), of the Doubs from the Danube (D_mean= 0.2177 ± 0.0028; Gpi-2**, Pgm-1**, Pgdh**, Ah**), and of the Doubs from the Hochrhein-Oberrhein (D_mean= 0.1780; Pgm-1**, Pgdh**, Ah**). Genetic distances between streams within these drainages proved low (Neckar: D_men= 0.0047 ± 0.0014, Danube: D_mean= 0.026 ± 0.0179, Rhine: D_mean= 0.0308). Screening of another 16 small-size samples consisting of 55 sculpins for nine diagnostic loci (Aat-2**, Acp-1**, Acp-2**, Ah**, Est-1**, Fh**, Gpi-2**, Pgdh** and Pgm-1**) confirmed the genetic homogeneity of sculpins within the Danubian and Neckarian drainage systems, but Neckarian sculpins were similar to those from the river Main. Populations from Hochrhein-Oberrhein resembled the Danubian stock but contained a decreasing frequency of ‘Neckarian’ markers when approaching the Danubian region. The genetic divergence between Neckarian, Danubian and Rhdnian sculpins suggests the existence of hitherto neglected taxa of anteglacial divergence.     

The heterozygosity for the biochemical and immunological markers of genes and the variability of morphophysiological traits in man

The relationship between heterozygosity of 9 biochemical and 5 blood group loci and variability of body length and age of menarche were studied in 467 women and 336 men from Moscow population. High and low levels of individual heterozygosity were shown in men to be associated with the maximal values of the coefficient of variation (CV) of body length, while in women strong positive correlation between the CV of body length and individual heterozygosity was demonstrated. The highest level of heterozygosity was revealed in medium-height men and low-height women. Highly heterozygous women were characterized either by early or by late age of menarche; early onset of menarche causes growth retardation. Positive correlation between the CV of body length and heterozygosity in women is due to the accumulation of low-height individuals having early age of menarche. The results are discussed in terms of Lerner's concept of genetic homeostasis. It is concluded that an average level of heterozygosity is optimal for a population.     

The Influence of Translocation Strategy and Management Practices on the Genetic Variability of a Reestablished Elk (Cervus elaphus) Population

Reintroduction of terrestrial vertebrates with the goal of ecosystem restoration typically establishes small and isolated populations that may experience reduced genetic variability due to founder effects and genetic drift. Understanding the genetic structure of these populations and maintaining adequate genetic diversity is important for long‐term restoration success. We quantified genetic variability at six microsatellite loci for a reintroduced population of Cervus elaphus (elk) restored to the tallgrass prairie ecosystem of northeastern Kansas. Allelic richness, observed and expected heterozygosity were intermediate to levels reported in other North American elk populations. Current levels of genetic variability in restored North American elk populations were not well explained by founding population size, number of founding populations, or number of years since the last translocation. Simulation results suggest that the retention of genetic variability in isolated populations is strongly influenced by mating system while also being impacted by temporal variability in population size and population growth rate. Our results have implications for understanding how translocation strategies and post‐reintroduction management may influence genetic variability in restored populations.     

The distribution of individual heterozygosity in natural populations

Estimation of the distribution of the level of individual heterozygosity within natural populations is explored with both Monte-Carlo simulation studies and data from natural populations. Simulations indicate that heterozygosities estimated from as few as a dozen randomly chosen loci may, to some degree, reflect (r = 0.35) heterozygosity determined by 100 independent loci. The shape of the expected distribution of heterozygosity is heavily dependent upon levels of heterozygosity at the loci. Complete genetic data for 12 loci from 997 Fundulus heteroclitus are used to describe the distributions of heterozygosity for different localities, for age classes and for sexes. The distributions deviate from normality. Distributions from different localities are not different, but the distributions are heterogeneous among age classes at one of two localities and are heterogeneous between the sexes.     

Evaluation of genetic variability of the Kamchatka crab Paralithodes camtschaticus (Tilesius) using allozyme markers

Interspecific genetic variation in populations of red king crab Paralithodes camtschaticus Tilesius (Litholidae, Decapoda: Crustacea) was examined using allozyme markers. The activity of 57 enzymes and the general protein presumably encoded by 92 loci was detected. The level of allozyme variability was low: the expected heterozygosity and the proportion of polymorphic loci were respectively 0.027 +/- 0.008 and 6.5%. This level of heterozygosity is three times lower than the average value for 122 crustacean species (0.082 +/- 0.007). Although genetic variants were found at 22 loci, their frequencies were generally low: only in loci 6-Pgd, Alp-1, and Pep-1 did the frequencies of the most common alleles not exceed 0.9. All polymorphic loci except one had two alleles; the exception was 6-Pgd, which had three alleles. The possible reasons for the low level of allozyme variability in red king crab are discussed.     

Patterns of genetic variability at individual minisatellite loci in minke whale Balaenoptera acutorostrata populations from three different oceans

The genetic variability at six cloned minisatellite loci was analyzed in minke whale populations from the North Atlantic, North Pacific, and Antarctic Oceans. Three loci displayed only a few different alleles in each of the three populations, with heterozygosity ranging from 0.00 to 0.47, and three loci revealed many different alleles in at least two fo the three populations, with heterozygosity ranging up to 0.98. Using small sample sizes, samples from two adjacent Antarctic Management Areas were not found to differ significantly in allele frequencies at any of the six loci. The use of principal coordinate analysis to detect multilocus disequilibria was explored. No significant evidence was found of intrapopulation heterogeneity within the pooled Antarctic sample. Pronounced interoceanic differences were observed at every locus, confirming the existence of genetic isolation found earlier using more conventional marker systems. The populations from the three oceans appear to have diverged to such a degree that the hypervariable loci have had time to evolve independently and arrive at different evolutionary stages in different populations. The frequency of undetected "null" alleles is remarkably high in minke whale populations compared to human populations and is probably a result of the cloning protocol used. Minisatellite loci are shown to provide a powerful population genetic tool, supplying levels of resolution appropriate to different degrees of evolutionary divergence.      

Allozyme variation in 16 natural populations of Faidherbia albida (Del.) A. Chev

The objectives of this study were (1) to investigate the genetic variability; and (2) to determine the phylogenetic relationship in the leguminous tree Faidherbia albida through out its range of distribution in East, West and Southern Africa. A total of 16 populations were subjected to enzyme electrophoresis and 6 enzyme systems (Adh, Mdh, G6pdh, Idh, 6Pgdh, and alpha-Est) encoded by 12 loci were scored. A total of fifty one different alleles were detected, with an average of 2.5 alleles per locus. Forty three percent of the loci were polymorphic at a 95% criterion. The average expected heterozygosity (gene diversity index H(e)) was 0.141. All provenances showed significant deviation from Hardy-Weinberg expectation. The UPGMA cluster analysis, based on Modified Rogers distance revealed close similarities between Eastern and Southern African provenances, except Debre zeit from Ethiopia, which was closest to West African populations than to the East African populations. Also, Bignona from West Africa was peculiarly closer to East African provenances. Differentiation of West African populations from Eastern and Southern African provenances was quite evident, emphasising the clinical pattern of variation in these two major geographical races of F. albida in Africa. Wright's F-statistics showed an overall significant deficit of heterozygotes, a common feature in mixed mating, entomophilous, widespread species such as F. albida. The dendrogram analysis showed wide separation among the three Ethiopian provenances indicating a high level of genetic differentiation and diversity among them.     

Genetic structure of the Atlantic Rainforest tree species Luehea divaricata (Malvaceae)

The Atlantic Rain Forest is one of the most important Brazilian biomes and a hotspot for biodiversity that is characterized by its high level of endemism, where new species are still being described. Luehea divaricata (Malvaceae) is commonly found in riparian forests areas of the Atlantic forest. Because of the importance of this species in reforestation programs, we used nine pairs of microsatellite loci to study the genetic variability of this species along its distribution area and verify if fragmentation is compromising the survival of these populations. A total of 50 alleles were obtained with an average observed and expected heterozygosity of 0.53 and 0.67, respectively. Seven of the nine populations studied showed a heterozygosity deficit. Most of the genetic diversity was found within populations; while the level of genetic differentiation was moderated (6.84) between populations. Different levels of gene flow between the populations were detected. Positive and significant values of Fis were found for seven populations. The signal test for excess of heterozygosity indicated that a recent genetic bottleneck occurred in the fragmented populations. The dendrogram constructed by the UPGMA method revealed the formation of seven clusters, which was confirmed by the Bayesian analysis for number of K clusters. The presence of several pairs of loci in linkage disequilibrium confirms that these populations experienced a loss of genetic diversity caused by genetic drift. The results showed that it is necessary to develop management strategies for the conservation of these populations of L. divaricata as the viability of the next generations are severely compromised.     

栗属中国特有种居群的遗传多样性及地域差异

Genetic variability in 30 populations of three endemic Castanea (Tourn.) L. species (C. moUissima BI., C. seguinii Dode. and C. henryi (Skan) Rehd. et Wils. ) in China was investigated using isoelectric focusing in thin-layer polyacrylamide slab gels at 20 loci coding for 12 enzyme systems. C. mollissima was found to possess a significant higher value of genetic variability than that of the other two species. The percentage of polymorphic loci (P) and the expected heterozygosity (He) were 90.0% and 0. 311 at the species level, respectively; while P = 84.7 %, He = 0.295 at the population level. Comparison of the population genetic variability of C. mollissima in three regions of China, the populations from the Yangtze River valley showed a markedly higher mean expected gene heterozygosity. The results provided additional evidence that the Yangtze River valley, particularly in Shennongjia area, was the center of the genetic diversity of C. moUissima. Genetic relationships among populations and species were assessed by Nei genetic identity (I) and standard genetic distance (D), suggesting that C. mollissima and C. seguinii have a closer relationship, and genetic distances were correlated with geographical distances among populations. Genetic differentiation between populations in C. mollissima, C. seguinii and C. henryi was 7.5%, 10.9% and 22.1%, respectively, and the gene flow rate (Nm) was 3.20, 2.05 and 0.88 respectively. The study provides useful imfonnation for understanding of the origin and evolutionary events in genus Castanea and planning an effective conservation strategy.     

Genetic divergence between populations of two closely related troglobitic beetle species (Speonomus: Bathysciinae,Coleoptera)

Genetic variability and divergence at 13 enzyme loci were studied in two species of troglobitic beetles (Speonomus zophosinus andhydrophilus 12 populations) collected from the Pyrenees mountains. These allopatric species exhibit a high degree of specialization to underground life and a very small geographical range. Four diagnostic loci arePgi, Est-2, Pac-1, Phi. Within each species polymorphic loci exhibited marked spatial variation of allele frequencies, sometimes over short distances (5–10 km). The levels of genetic variability are comparable to those observed in non-cave invertebrates (frequency of polymorphic loci: 0.31; average expected heterozygosity: 0.112±0.015). Moreover, a single species showed variability in the average level of heterozygosity (0.06 to 0.172). Patterns of genetic differentiation among species were also studied; mean genetic distance (D: 0.263±0.010) between the twoSpeonomus species was of the same order of magnitude as most data reported in the literature between species. A significant heterozygote deficiency occurs in local populations and for all loci. This deficit does not vary from locus to locus. Several hypotheses were examined in an attempt to account for these observations in connection with the species biology. This deficit probably is due to the breeding structure of the population (inbreeding and assortative mating) coupled with limited dispersal ability.     

Variation in life history and genetic traits of Hawaiian mosquitofish populations

Mosquitofish (Gambusia affinis) were collected from 17 reservoirs on three islands in Hawaii, USA. Genetic and life history traits for adult females from these populations were used to evaluate hypotheses concerning short-term evolutionary divergence of populations recently established from a common ancestral source. The effects of founder events and drift on genetic variability and population differentiation were also examined. Significant differences in life history characteristics, allele frequencies, and multi-locus heterozygosities (H) were found among fish populations collected from different reservoirs and between reservoirs classified as stable or fluctuating on the basis of temporal fluctuation in water level. Females from stable reservoirs exhibited greater standard length (35.1 vs 32.8 mm), lower fecundity (11.9 vs 15.2 embryos), lower reproductive allocation (18.2% vs 22.8%), but larger mean embryo size (1.95 vs 1.67 mg) than females from fluctuating reservoirs. Consistency in means among replicates of each reservoir class and concordance in direction and magnitude of differences reported here and results of sampling conducted from these same locations 10 years previously (Stearns, 1983a) suggest that ecological factors intrinsic to these two environments are important in determing population life history traits. Females from stable reservoirs exhibited lower heterozygosity than females from fluctuating reservoirs (0.134 vs 0.158, respectively). Levels and direction od differences in heterozygosity, the high proportion of polymorphic loci and lack of fixation of alternative alleles argue against a purely stochastic explanation for genetic and life history variation among reservoir populations. Levels of genetic variability and interpopulation differentiation were similar to those observed in mainland populations of this species. A high proportion of the genetic diversity was apportioned between populations and within populations due to differences between juveniles and adults. Significant genotypic differences between adult and juvenile age classes suggest that the genetic divergence of local populations may occur over short periods of time.     

PERMANENT GENETIC RESOURCES: Isolation, characterization and cross-species amplifications of microsatellite loci from Conradina (Lamiaceae)

We report the isolation of microsatellite loci from three species in the genus Conradina (Lamiaceae). To ensure their utility for multiple species, loci were screened for amplification and variability in all six Conradina species; 11 loci demonstrated high levels of amplification and polymorphism in most species. These 11 loci were characterized in 20 individuals from one population of Conradina brevifolia; alleles per locus ranged from five to 15, and observed heterozygosity ranged from 0.30 to 0.90. These microsatellites will be used to clarify species limits, detect interspecific hybridization, and understand the partitioning of genetic variation in each species of Conradina.     

Patterns of genetic variation in rare and widespread plant congeners

Rare species are typically considered to maintain low levels of genetic variation, and this view has been supported by several reviews of large numbers of isozyme studies. Although these reviews have provided valuable data on levels of variability in plant species in general, and rare species in particular, these broad overviews involve comparisons that may confound the effects of rarity with a multitude of other factors that affect genetic variability. Additionally, the statistical analyses employed assume the data to be independent, which is not the case for organisms that share a common phylogenetic history. As the role of evolutionary history and historical constraints has become better understood, more researchers have studied widespread congeners when investigating the genetic diversity of rare species in an effort to control for these effects. We summarize the available data from such studies, comparing for rare and widespread congeners (1) the levels of genetic variability at the population and species levels and (2) measures of population substructuring. At the population level, we summarized data for percentage polymorphic loci (%P(pop)), mean number of alleles per locus (A(pop)), and observed heterozygosity (H(o)). Species-level measures used were percentage polymorphic loci (%P(spp)), mean number of alleles per locus (A(spp)), and total genetic diversity (H(T)). Indices of population subdivision (either F(ST) or G(ST)) were also examined. Using Wilcoxon signed rank tests, we found significant, but small, differences between rare and widespread species for all diversity measures except H(T). However, there does not appear to be a difference between rare and widespread congeners in terms of how genetic variation is partitioned within and among populations. Levels of diversity, for all measures examined, between rare and widespread congeners are highly correlated.     

Levels of genetic variability in sculpins (Gottidae: Teleostei) of the North American Pacific coast and an assessment of potential correlates

We examined levels of allozymic variation in intertidal sculpins of the genera Oligocottus, Clinocottus and Artedius. Variability in these species, as assessed by the proportion of loci polymorphic and average heterozygosity per locus, was generally low but within the range of values observed for other fish groups. In reviewing the ecological and life-history attributes of these sculpins, we found only one outstanding correlate of allozyme variability–larger-bodied species tended to have lower levels of variation. Also, the level of heterozygosity was not related in any obvious way to historical/ evolutionary aspects such as position in phylogenetic trees or the inferred relative ages of species. Our study of a relatively well-defined lineage of fishes that co-occur in the same geographical region and in the same, or broadly similar, habitats, underscores the difficulty of detecting correlations between genetic variability and seemingly relevant ecological features of species.     

Isolation and characterization of microsatellite markers in pangolins (Mammalia,Pholidota, Manis spp.)

Thirty‐four polymorphic dinucleotide microsatellite loci were developed in the Malayan pangolin Manis javanica. Of the 34 markers, 32 and 18 were also amplified, respectively, in the Chinese pangolin (Manis pentadactyla) and the African tree pangolin (Manis tricuspis). Analysis of 24 Malayan, 12 Chinese and 2 African tree pangolins showed high levels of variability (heterozygosity ranging from 0.321 to 0.708). These are the first available microsatellite markers in Pholidota and will be an invaluable tool for evolutionary and conservation genetic studies in pangolins.     

Variability of nuclear microsatellite loci in the populations of Siberian dwarf pine (<Emphasis Type="Italic">Pinus pumila</Emphasis> (Pallas) Regel) from the Russian part of the range

Variability of nuclear microsatellite loci was examined in Siberian dwarf pine. Six microsatellite loci (RPS2, RPS6, RPS12, RPS124, RPS127, Pc18) demonstrated different polymorphism levels in ten populations of Siberian dwarf pine. The average number of alleles per locus was 4.88, the average observed heterozygosity was 0.465, and the average expected heterozygosity was 0.510. About 13% of total genetic variability was explained by the genetic differences between the populations (F _ST = 0.129). Genetic distances between the examined populations of Pinus pumila inferred from the data on the SSR marker frequencies statistically significantly correlated with the geographical distances between the population samples. The level of genetic variability of the populations from Kamchatka Peninsula was lower than that demonstrated by continental and island populations. The genetic differentiation of the Kamchatka–Magadan and other populations of Siberian dwarf pine observed in our study can be explained in terms of their formation from different Pleistocene refugial centers.     

Isolation and characterization of microsatellite loci from the yellow-eyed penguin (Megadyptes antipodes)

Twelve microsatellite loci were isolated and characterized in the endangered yellow-eyed penguin (Megadyptes antipodes) using enriched genomic libraries. Polymorphic loci revealed two to eight alleles per locus and observed heterozygosity ranged from 0.21 to 0.77. These loci will be suitable for assessing current and historical patterns of genetic variability in yellow-eyed penguins.