Allozyme differentiation of populations of the dogwhelk Nucella lapillus, (L.): the relative effects of geographic distance and variation in chromosome number

Geographic patterns of genie differentiation were compared with differentiation between karyotypes in the intertidal snail Nucella lapillus. Samples from 24 sites covering the species range in Europe and North America were analysed for allozyme variation at 16 soluble enzyme loci. Two homokaryotypes have been identified with diploid numbers 2n = 26 and 2n= 36 (variation is Robertsonian and hybrids have intermediate chromosome numbers) and samples were classified (on the basis of published data) according to karyotype. Group 1 consisted of samples from three English Channel populations of higher chromosome number (on average 2n > 32) and Group 2 consisted of the remaining 21 samples (presumed to be 2n= 26). Karyotype variation accounts for roughly the same amount of the absolute allozyme variance as geographic variation (46.3 °, and 53.7°, respectively). Yet the patterns of differentiation seen between karyotypes and with geographic separation are very different. In samples classified as 2n= 26 (Group 2), while there is a significant amount of heterogeneity (F_ST per locus averaged 0.128 for 10 polymorphic loci), allozyme variation occurs independently at different loci so mean genetic identity (Nei) is high: 0.972. There is only a slight decline in genetic identity with distance (genetic identity averaged 0.965 for amphi-atlantic comparisons) indicating that passive transport of juveniles or adults may contribute significantly to gene flow. Conversely, allozyme variation between karyotypes was concordant. High chromosome number populations possessed a suite of alleles at four allozyme loci (Esl-3, Lap-2, Mdh-1 and Pep-2) which were absent or rare in Group 2 samples resulting in high F_ST values for these loci (from 0.294 to 0.472) when karyotypic classes were combined. Consequently the mean genetic identity between these Robertsonian races is low, 0.856, and falls within the range more usually associated with congeneric comparisons than with con-specific comparisons. The mechanisms maintaining this genie difference are unclear. However the distribution of the karyotypes and physiological and morphological differences (in shell shape) between them strongly suggest that karyotypic variation in Nucella is adaptive.     

云南松居群遗传学研究的等位酶分析方法

针对１５个云南松Ｐｉｎｕｓｙｕｎｎａｎｅｎｓｉｓ居群,开展了１４种酶系统的水平切片淀粉凝胶电泳实验,在谱带遗传分析的基础上确定了３３个等位酶位点及其等位基因。其中有３２个等位酶位点是多态的（有２个以上的等位基因）,只有一个单态位点Ｄｉａ－４。有３个等位基因的位点有Ｌａｐ－１、Ｌａｐ－２、Ａａ－３、Ｓｋｄ－１、Ｓｋｄ－２、Ａｄｈ－１、Ａｄｈ－３、Ｇｄｈ、Ｐｇｄ－１、Ｐｇｍ－１、Ｐｇｍ－３、Ｐｇｉ－１、Ｐｇｉ－３、Ｍｄｈ－１、Ｍｅ、Ｇ６ｐｄ、Ｄｉａ－１、Ｔｐｉ－１、Ｔｐｉ－２、Ｔｐｉ－３和Ｔｐｉ－４,有４个等位基因的位点有Ｓｋｄ－３、Ａｄｈ－２、Ｐｇｄ－２、Ｍｄｈ－２、Ｍｄｈ－３、Ｍｄｈ－４和Ｄｉａ－２,有５个等位基因的位点有Ａａｔ－１和Ｄｉａ－３。云南松居群的等位基因平均数Ａ＝２１,在松属中居于中上水平。本研究揭示了云南松居群酶位点及其等位基因带谱的变异式样,为松属植物的遗传多样性研究提供了一批酶位点及其等位基因的参考图谱     

Polymorphism and inheritance of serum esterases and β-globulins in the paradise fish (Macropodus opercularis; Anabantidae)

Electrophoretic variants of serum esterases and β-globulins in two subspecies of paradise fish ( Macropodus opercularis ) were studied. Four esterase loci ( Est-1, Est-2, Est-3 and Est-4 ), a single transferin ( Tf ) and another major β-globulin locus ( Bg ) were identified by segregational analysis. Est-3 seems to be a monomorphic. locus. Three alleles of Est-1 , two of Est-2 , two of Est-4 , four of Tf and two alleles of Bg were found in the laboratory population. None of these loci were closely linked. Electrophoretic patterns of F₁ hybrids confirmed the monomeric structures of each of the studied proteins. Allelic segregation at the Tf and Bg loci was normal in F₂ and backcross populations. In crosses of the two Macropodus subspecies there were deviations from Mendelian ratios because of missing recombinant esterase phenotypes. Each of these would have been homozygous Est-2^f/f . We suppose that Est-2^f/f causes lethality in the early phase of development, except in the Est-1^c/c, Est-2^f/f combination characteristic of the parental subspecies M.o. concolor .     

Genetic differentiation in endemic Lobularia (Brassicaceae) in the Canary Islands

Patterns of genetic diversity were examined in five endemic subspecies of the Lobularia canariensis complex from six of the Canary Islands. The taxa are interfertile, insect-pollinated outbreeders with wind dispersal. Electrophoretic analysis revealed a high level of genetic polymorphism at ten loci coding soluble enzymes, with a mean of 2.38 alleles per locus, 73.7 % polymorphic loci, and a mean heterozygosity of 0.279. Excesses of homozygotes, indicating inbreeding, were observed in small populations. The average total diversity was high, F.,=0.518. Among-population diversity, F_ST=0.318, contributed more to the total diversity than within-population diversity, F_IS=0.222. Little geographic or taxonomic patterning of the allozyme variation was observed. The mean genetic identitity for pairwise comparisons of the 19 populations was 0.76, with a range of 0.51–0.96 and c. 17 % of the comparisons below 0.67 and c. 8 % above 0.90. The results contrast with the many cases of high genetic identities reported for populations of endemic plants on oceanic islands. High levels of allozyme divergence suggest a relatively old origin of the L. canariensis complex and a long period of isolation of some of the populations.     

Genetic differentiation between alpine and lowland populations of a butterfly is related to PGI enzyme genotype

Understanding the ecological process of population differentiation and identifying the molecular changes that contribute to adaptation is a central aspect of evolutionary biology. In this study we analyzed the geographic variation in allozyme allele frequencies (based on 15 enzyme systems representing 18 loci) across 18 populations of the butterfly Lycaena tityrus from different altitudes. Population genetic analyses showed that within population genetic diversity (e.g. mean number of alleles per loci: 1.8; expected heterozygosity: 12%) was within the typical value range for the Lepidoptera. The populations of L. tityrus investigated showed a remarkable genetic differentiation (F_ST: 0.065), being clearly divided into an alpine (high-altitude) and a non-alpine (low-altitude) cluster. This differentiation was almost entirely caused by variation at a single enzyme locus, PGI. Although the involvement of historical events cannot be ruled out, several lines of evidence suggest that the specific pattern of allozyme (and in this case particularly PGI) variation found is caused by thermal selection. For example, genetic variation was highly locus-specific rather than relatively uniform, as should be expected for effects of natural selection. Further, the PGI 2–2 genotype dominating in alpine (in contrast to low-altitude) populations is known to exhibit increased cold stress resistance and relatively long development times typical of alpine populations. Thus, PGI (or possibly a closely linked gene) is an obvious target for thermal selection in L. tityrus . This study exemplifies how allozyme analyses can be used to detect candidate loci likely to be under selection.     

The effect of natural selection on estimates of genetic divergence among populations of the Atlantic salmon

The effect of natural selection on the _mMEP-2 * locus on measures of genetic divergence among Atlantic salmon populations was investigated by examining the pattern of change in the level of genetic differentiation (F_ST) averaged over loci when data on the _mMEP-2 * locus were either included or excluded. The level of F_ST among populations at various geographic scales was estimated from allele frequencies at up to four loci (_s AAT-4 *, IDDH-1 *, IDHP-3 *, and _mMEP-2 *). At smaller geographic scales (within river systems or limited geographic regions) levels of variance in _mMEP-2 * allele frequencies were reduced relative to mean levels. At larger geographic scales (across continents or the species range) variation in _mMEP-2 * allele frequencies was greater than mean levels. These results suggest an a priori hypothesis for the effect of selection on the _mMEP-2 * locus which may be applied in future studies on variation in protein coding or other (e.g. mini- and microsatellite) loci in the Atlantic salmon. It is recommended that estimates of gene flow among populations of the Atlantic salmon based on mean F _ST estimates which include data on the _mMEP-2 * locus should be viewed with caution.     

Genetic control of allozymes from the juniper tall (<Emphasis Type="Italic">Juniperus excelsa</Emphasis> Bieb.) of Crimea

The genetic control of nine enzyme systems has been analyzed in the preserved juniper species (Juniperus excelsa Bieb.) from a natural population of the Mountain Crimea. Isozymes were extracted from endosperms of the haploid seeds to be studied by electrophoresis. As a result 16 loci have been identified, of which 14 loci were polymorphic (Gdh, Got-1, Mdh-1, Mdh-2, Mdh-3, Acp-1, Acp-2, Acp-3, Lap-1, Dia-1, Fdh, Sod-1, Sod-2, and Sod-3). Analysis of allele segregation in heterozygous trees confirmed monogenic inheritance of these alleles.     

Isoenzyme electrophoretic patterns in discus fish (Symphysodon aequifasciatus Pellegrin, 1904 and Symphysodon discus Heckel, 1840) from the Central Amazon

The discus is a very popular and expensive aquarium fish belonging to the family Cichlidae, genus Symphysodon, formed by three Amazon basin endemic species: Symphysodon aequifasciatus, S. discus and S. tarzoo. The taxonomic status of these fish is very controversial, with a paucity of molecular research on their population genetic structure and species identification. Information on molecular genetic markers, especially isoenzymes, in search of a better understanding of the population genetic structure and correct identification of fish species, has been receiving more attention when elaborating and implementing commercial fishery management programs. Aiming to contribute to a better understanding of the species taxonomic status, the present study describes the isoenzymatic patterns of 6 enzymes: esterase (Est - EC 3.1.1.1), lactate dehydrogenase (Ldh - EC 1.1.1.27), malate dehydrogenase (Mdh - EC 1.1.1.37), phosphoglucomutase (Pgm - EC 5.4.2.2), phosphoglucose isomerase (Pgi - EC 5.3.1.9), and super oxide dismutase (Sod - EC 1.15.1.1) extracted from skeletal muscle specimens and analyzed by starch gel electrophoresis. Monomorphic patterns, presumably controlled by 11 loci: Est-1, Est-2, Est-3, Ldh-1, Ldh-2, Mdh-1, Mdh-2, Pgi-1, Pgi-2, Pgm-1, and Sod-1 were fixed for the same alleles: Est-1(1), Est-2(1), Est-3(1), Ldh-1(1), Ldh-2(1), Mdh-1(1), Mdh-2(1), Pgi-1(1), Pgi-2(1), Pgm-1(1), and Sod-1(1), respectively, and detected in all 60 specimens examined (27 S. aequifasciatus from Manacapuru and 33 S. discus from Novo Air?o, Central Amazon). The failure in the present study to detect diagnostic loci, which could be very useful for differentiating S. aequifasciatus from S. discus species, and polymorphic loci, which could also be applied for possible identification and delimitation of their stocks, does not rule out the possibility of there existing in other isoenzyme gene loci to be analyzed in the future.     

Genic heterozygosity, chromosomal interchanges and fitness in rye: any relationship?

Relationship between heterozygosity at allozyme loci, chromosomal interchanges and fitness was analyzed in a rye cultivar showing a polymorphism for such rearrangements. Nine allozyme systems (ACO, ACPH, GOT, GPI, LAP, MDH, PER, PGD and PGM) and five components of fitness (number of fertile tillers, total offspring, egg cell fertility, flowers/ear and seeds/ear) were studied. The estimated selection coefficients against interchange heterozygotes ranged from s = 0.12 to s = 0.34. A significant effect of the genic heterozygosity on some fitness components was observed in interchange heterozygotes (tillering and total offspring), in their standard homozygous sibs (flowers/ear and seeds/ear) and in the descendants of the crosses between standard karyotypes (flowers/ear, seeds/ear and egg cell fertility). However, the main effect was linked to genetic background associated to different crosses. Significant differences for Acph-1, Gpi-1, Lap-1, Mdh-1, Mdh-4, Pgd-2 and Pgm-1 loci were also found in some of these crosses although these differences were inconsistent. This suggests that probably the allozyme loci analyzed were not directly contributing to the fitness and that they are linked, in some cases, to different deleterious alleles depending on both cross and locus. This fact could support the local effect hypothesis as explanation although we do not discard the existence of some inbreeding level (general effect hypothesis) since all crosses and loci studied show a overall consistent trend of increased fitness with increased heterozygosity.     

Evidence of a reproductive barrier between two forms of the marine periwinkle Littorina fabalis (Gastropoda)

Studies of allozyme variation may reveal unexpected patterns of genetic variation which challenge earlier conclusions of species delimitations based on morphological data. However, allozyme variation alone may not be sufficient to resolve this kind of problem. For example, populations of the marine intertidal snail Littorina fabalis (=Littorina mariae) from wave exposed parts end from protected parts of the same shores are distinguished by different alleles of arginine kinase (Ark) while indifferent, or very nearly so, in another 29 loci. Intermediate populations have large deficiencies of exposed/sheltered heterozygote classes of Ark and we have earlier suggested habitat-related selection in this locus as the explanation. In this study we estimated growth rate of individual snails of different Ark-genotypes in three different habitats (exposed, sheltered and intermediate). In all habitats the snails homozygous for alleles of ‘exposed’ type grew faster and matured at a larger size than did snails homozygous for alleles of ‘sheltered’ types. This relationship was indirectly confirmed in three additional sites of intermediate exposure where exposed AA-genotypes dominated among large (>8 mm) snails while the sheltered genotypes dominated among small (<5 mm) snails of truly sympatric samples. We furthermore found small differences in allele frequencies of two other loci (Pgi and Pgm-2) and in shell colour frequencies, comparing sympatric snails of exposed and sheltered Ark-homozygotes. Although we found no signs of habitat-related selection among snails of different Ark-genotype, or selection against heterozygotes, we cannot reject selection in Ark, as our experiments only covered one island, one season and grown-up snails. The coupling between allozyme and phenotypic characters in strictly sympatric samples of snails suggests the presence of two gene pools. Perhaps the large and small forms of L. fabalis represent very closely related cryptic taxa. However, introgression between them seems a possible explanation for the striking similarities in the vast majority of morphological and allozyme characters.     

Micro- and macrogeographic allozyme variation in Littorina fabalis; do sheltered and exposed forms hybridize?

Earlier studies of Swedish populations of the marine snail Littorina fabalis show that snails from different microhabitats (with a greater and lesser exposure to wave action) have almost diagnostic differences in one allozyme locus (arginine kinase, Ark ), and differ in adult size. Snails with 'sheltered' and 'exposed' Ark genotypes occur in sympatry in intermediary exposed sites and here adult sizes remain distinct. Approaching the microgeographic differentiation we studied the parts of two populations where the frequency of Ark changes dramatically over zones 50–120 m wide. The aim was to test if the transitional zones are best described as areas of mixing of two genetically separate populations, or if hybridization between the exposed and sheltered groups occurs. Heterozygotes were in deficiency along both clines but were still roughly twice as common as expected from a pure mixing of 'sheltered' and 'exposed' groups suggesting hybridization. Hybridization was also supported by the observation that snails homozygous for sheltered and exposed alleles mated at random with each other in both populations. On the macrogeographic scale, we found populations from exposed and sheltered sites in France and Wales being fixed for the same exposed and sheltered Ark alleles as found in Sweden. However, variation in three other highly polymorphic loci indicated geographic affinity rather than habitat similarity being the main factor of genetic coherence. These observations support a hypothesis of gene flow between exposed and sheltered populations of L. fabalis . Two Spanish populations were remarkably different with unique alleles at high frequencies in three of four strongly polymorphic loci.     

Hierarchical organization of genetic variation in the Costa Rican livebearing fish Brachyrhaphis rhabdophora (Poeciliidae)

I examined the geographic distribution of genetic variation in the livebearing freshwater fish Brachyrhaphis rhabdophora in northwestern Costa Rica as revealed by allozymes and mitochondrial DNA sequences. Allelic variability at 11 enzyme-coding loci surveyed across 12 localities revealed marked genetic differentiation among populations within drainages (0_P= 0.36) and among drainages within regions (0_D=0.17), but not between northern and southern geographic regions (0_R=– 0.02). Allozyme variation was hierarchically organized such that populations found within stream drainages were more similar to each other than to populations found in adjacent drainages, a result confirmed by cluster analysis. In contrast to the allozyme data, there was extremely little DNA sequence variation among populations in the mitochondrial control region (3 variable nucleotide positions out of 444 bp examined). The difference in genetic divergence between allozyme and mtDNA markers was unexpected and is discussed in terms of biogeographical colonization events and a molecular selective swéep on the mitochondrial genome, both processes that could explain the lack of mitochondrial variability in this highly subdivided species.     

Temporal and Microgeographic Variation in Allozyme Frequencies in a Natural Population of DROSOPHILA BUZZATII

Temporal variation in allozyme frequencies at six loci was studied by making monthly collections over 4 yr in one population of the cactophilic species Drosophila buzzatii. Ten sites were defined within the study locality, and for all temporal samples, separate collections were made at each of these sites. Population structure over microgeographic space and changes in population structure over time were analyzed using F-statistic estimators, and multivariate analyses of allele and genotype frequencies with environmental variables were carried out. Allele frequencies showed significant variation over time, although there were no clear cyclical or seasonal patterns. A biplot analysis of allele frequencies over seasons within years and over years showed clear discrimination among years by alleles at four loci. During the 4 yr, three alleles showed directional changes which were associated with directional changes in environmental variables. Significant associations with one or more environmental variables were found for allele frequencies at every locus and for both expected and observed heterozygosities (except those for Est-1 and Est-2). Thus, variation in allele frequencies over time cannot be attributed solely to drift. Significant linkage disequilibria were detected among three loci (Est-2, Hex and Aldox), but there was no evidence for spatial or temporal patterns. The F-statistic analyses showed significant differentiation among months within years for all loci, but the statistic used (coancestry) was heterogeneous among loci. Estimates of F (inbreeding) for all loci were significantly different from zero, with the loci in four groups, Adh-1 (negative), Pgm(small positive), Est-2 and Hex (intermediate) and Est-1 and Aldox (high positive). The correlation of genes within individuals within populations (f) for each locus in each month by site sample differed among loci, as did the (f) for each locus in each month by site sample differed among loci, as did the patterns of change in f over time (seasons). Heterogeneity in the F-statistic estimates indicates that natural selection is directly or indirectly affecting allele and genotype frequencies at some loci. However, the F-statistic analyses showed essentially no microgeographic structure (i.e., among sites), although there was significant heterogeneity in allele frequencies among flies emerging from individual rots. Thus, microspatial heterogeneity probably is most important at the level of individual rots, and coupled with habitat selection, it could be a major factor promoting diversifying selection and the maintenance of polymorphism.(ABSTRACT TRUNCATED AT 400 WORDS)     

Genetic evidence for population expansion in Hydrotaea irritans (Fallèn) (Diptera: Muscidae)

Allozyme variation in the sheep headfly Hydrotaea irritans was studied on two spatial scales. Geographic variation among seven Danish and one Dutch population revealed significant but rather low genetic differentiation with F _ST = 0.01 over all loci. The Dutch population was on average not more different from the Danish populations than the Danish populations from each other. Allele frequencies were very skewed with the most common allele always exceeding 0.85 and usually 0.9 in frequency, but with many rare alleles at some loci. Tests for neutrality of the variation at the nine polymorphic loci revealed highly significant deviations from expected homozygosity in this species, which was not found in a comparative analysis of allozyme variation at similar loci of seven other Hydrotaea species. To explain the peculiar observed pattern of allozyme variation in H. irritans , it is suggested that this species has successfully expanded its range and spread through northern and central Europe in the recent past. Alternatively, H. irritans may have recently invaded a new niche, resulting in increased abundance of the species and subsequent dispersal to former areas of the species distribution.     

Genetical control of the allozymes in juniper (Juniperus excelsa Bieb.) of the Crimea

Genetical control of nine enzyme systems has been studied in preserved juniper species (Juniperus excelsa Bieb.) of the natural population of the mountain Crimea. Isozymes were extracted from the haploid seed endosperms and separated elecrophoretically. As a result 16 loci have been identified. Fourteen of them were polymorphic (14--Gdh, Got-1, Mdh-1, Mdh-2, Mdh-3, Acp-1, Acp-2, Acp-3, Lap-1, Dia-1, Fdh, Sod-1, Sod-2, Sod-3). Analysis of the allele segragation of the heterozygous trees confirmed their monogenic inheritance.     

Molecular evolution of two linked genes, Est-6 and Sod, in Drosophila melanogaster.

We have obtained 15 sequences of Est-6 from a natural population of Drosophila melanogaster to test whether linkage disequilibrium exists between Est-6 and the closely linked Sod, and whether natural selection may be involved. An early experiment with allozymes had shown linkage disequilibrium between these two loci, while none was detected between other gene pairs. The Sod sequences for the same 15 haplotypes were obtained previously. The two genes exhibit similar levels of nucleotide polymorphism, but the patterns are different. In Est-6, there are nine amino acid replacement polymorphisms, one of which accounts for the S-F allozyme polymorphism. In Sod, there is only one replacement polymorphism, which corresponds to the S-F allozyme polymorphism. The transversion/transition ratio is more than five times larger in Sod than in Est-6. At the nucleotide level, the S and F alleles of Est-6 make up two allele families that are quite different from each other, while there is relatively little variation within each of them. There are also two families of alleles in Sod, one consisting of a subset of F alleles, and the other consisting of another subset of F alleles, designed F(A), plus all the S alleles. The Sod F(A) and S alleles are completely or nearly identical in nucleotide sequence, except for the replacement mutation that accounts for the allozyme difference. The two allele families have independent evolutionary histories in the two genes. There are traces of statistically significant linkage disequilibrium between the two genes that, we suggest, may have arisen as a consequence of selection favoring one particular sequence at each locus.     

Apparent Selection of Enzyme Alleles in Laboratory Populations of Drosophila

Twelve laboratory populations of recently collected Drosophila willistoni were begun with different frequencies of alleles at three enzyme loci, six populations at 25 degrees and six at 19 degrees . Periodic sampling of the populations allowed monitoring of the frequency changes in allozymes over time.-At Lap-5 (a locus coding for leucine amino peptidase), three alleles converged to the same frequencies in all populations at both temperatures. The apparent equilibrium frequency of the major allele was about.75; this is different from the frequency (.57) found in the natural population from which the experimental populations were begun. Allele frequency changes at the esterase-5 locus (Est-5) were slower but consistent in all cages. It is difficult to determine if an equilibrium has been reached. However, the frequency of the rare allele in all cages is about the same as in wild populations, 5%. Alleles at both Lap-5 and Est-5 are non-randomly associated with inversions in the chromosomes onto which they map. Because of these associations, it is impossible to unambiguously attribute the change in allele frequencies to selection at the loci being observed.-After one year, no significant gene frequency changes were detected at Est-7, the third locus studied.     

Quantitative trait and allozyme divergence in the Greenfinch (Carduelis chloris, Aves: Fringillidae)

Quantitative trait divergence and variability among 12 greenfinch populations across continental Europe was examined and compared to divergence in neutral genetic markers (allozymes). The added among locality variance component for 16 skeletal traits was large (mean 28%, range 4–48%) equalling a divergence of up to three SD units. The divergence in quantitative traits (Qst = 0.04-0.48) greatly exceeded that in alloymes (F_ST= 0.01-0.07), indicating the differentiation in quantitative traits to be larger than expected by mutation and drift alone. This conclusion was consistent also with results from the multivariate approach of Rogers & Harpending. However, genetic and morphometric distances between populations were positively correlated, even when controlling for the geographic distance separating pairs of populations. In concordance with Bergmann's rule, most traits were strongly and positively correlated with latitude, indicating latitudinally ordered genetic or/and environmental effects. However, the correlation between lower mandible width and latitude was strongly negative, demonstrating an inverse relationship between beak size and body size across the populations. These results are interpreted to reflect the re-colonization of history of northern Europe (genetic and geographic distances correlated) which has been paralleled by selection acting on quantitative traits (Q_ST>F_ST)- In particular, the counter-gradient variation in beak width, a functionally important trait, is suggestive of an adaptive basis for quantitative trait divergence.     

Genetic Variation and Clonal Diversity of the Two Divergent Types of Clonal Populations of Leymus chinensis Tzvel on the Song Liao Steppe in the West of Northeastern China

The genetic variation and clonal diversity of two divergent types (grey-green and yellow-green) of clonal populations of Leymus chinensis Tzvel at 14 loci were compared. Total gene diversity (HT) and the coefficient of genetic differentiation (GST) were all higher for the yellow-green type (HT = 0.270; GST =0.186) than for the grey-green type (HT = 0.250; GST = 0.157) of L. chinensis. Rare alleles usually occurred as heterozygotes rather than homozygotes and significant deviations from Hardy-Weinberg equilibrium were found only at a few loci. This indicated that these two types of populations were mainly out-crossing. Clonal diversity, evenness of clones, and mean clone size were not significantly different between the two types. We found that differences between the clone size and genetic variation of the yellow-green type of populations occurred with different climate and habitat population groups. However, for the grey-green type of populations, these genetic variations decreased under conditions of different climate and habitat population groups.     

Linkage relationships of seven enzyme and two pigmentation loci in the snail Biomphalaria glabrata

Crossing experiments with inbred stocks of the snail (Biomphalaria glabrata) demonstrated that variants at two loci determining pigmentation and seven enzyme-determining loci exhibited normal Mendelian segregation ratios in F2 progeny. Among 39 pairwise comparisons for joint segregation, there was evidence of genetic linkage between a locus controlling mantle pigmentation (S) and 6-phosphogluconate dehydrogenase (Pgd) and confirmation of a previously described linkage between esterase-2 (Est-2) and catalase (Cat). Recombination fractions were estimated to be 17 +/- 4 for S-Pgd and 33 +/- 5 for Est-2-Cat. The remaining five loci--Acon-1, Pgm-1, Lap-1, Lap-2, and Pgd--assorted independently. This brings to 17 the number of loci examined for segregation and assortment in this medically important species. As Biomphalaria has a chromosome number n = 18, markers should soon be available for most or all of the linkage groups.