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PubMed2004

Swift laboratory thermal evolution of wing shape (but not size) in Drosophila subobscura and its relationship with chromosomal inversion polymorphism.

Latitudinal clinal variation in wing size and shape has evolved in North American populations of Drosophila subobscura within about 20 years since colonization. While the size cline is consistent to that found in original European populations (and globally in other Drosophila species), different parts of the wing have evolved on the two continents. This clearly suggests that 'chance and necessity' are simultaneously playing their roles in the process of adaptation. We report here rapid and consistent thermal evolution of wing shape (but not size) that apparently is at odds with that suggestion. Three replicated populations of D. subobscura derived from an outbred stock at Puerto Montt (Chile) were kept at each of three temperatures (13, 18 and 22 degrees C) for 1 year and have diverged for 27 generations at most. We used the methods of geometric morphometrics to study wing shape variation in both females and males from the thermal stocks, and rates of genetic divergence for wing shape were found to be as fast or even faster than those previously estimated for wing size on a continental scale. These shape changes did not follow a neat linear trend with temperature, and are associated with localized shifts of particular landmarks with some differences between sexes. Wing shape variables were found to differ in response to male genetic constitution for polymorphic chromosomal inversions, which strongly suggests that changes in gene arrangement frequencies as a response to temperature underlie the correlated changes in wing shape because of gene-inversion linkage disequilibria. In fact, we also suggest that the shape cline in North America likely predated the size cline and is consistent with the quite different evolutionary rates between inversion and size clines. These findings cast strong doubts on the supposed 'unpredictability' of the geographical cline for wing traits in D. subobscura North American colonizing populations.

Analysis of Variance↗
PubMed1999

Geographic variation of six allozyme loci in Drosophila melanogaster: an analysis of data from different continents.

Based on data from 70 literature sources, the frequencies of common alleles at six allozyme loci were examined in Drosophila melanogaster populations from five geographic regions: North America (including Central America), South America, Europe-Africa, Asia, and Australasia. The analyzed loci were Adh, Odh, Gpdh, G6pd, Pgd, and Est-6, which have been previously reported by other authors to show latitudinal variation in North America, Eurasia and Australasia. We found five parallel latitudinal clines for AdhF and three clines for GpdhS in five geographic regions as well as four clines for G6pdF, three clines for Est-6S, and two clines for OdhF and PgdF in four regions (data from South America for G6pd, Odh, Est-6, and Pgd were not available). Such pattern of variation confirmed the possibility that considered allozyme polymorphisms are maintained by climatic selection. Significant differentiation of mean allele frequencies among geographic regions was in agreement with current evidence on history of D. melanogaster worldwide dispersion.

Animals↗
PubMed2026

The MexMAGIC population reveals the genetic architecture of traits exhibiting clinal variation in Mexican native maize.

Defining the genetic basis of local adaptation is a key goal of evolutionary biology and crop improvement. Theory predicts that when selective pressures follow differences in the environment, a cline will be established. Clines can be exploited to uncover adaptive variation by association of alleles with the environment. However, monotonic phenotypic change over a cline is not necessarily mirrored in the behavior of genetic variants and population structure can further complicate analysis. To study genetic and phenotypic variation across the environment, we developed a multi-parent advanced generation inter-cross (MAGIC) population using eight Mexican native maize (Zea mays L. ssp. mays) varieties sourced from distinct agroecological zones. We evaluated the population in a common garden in Mexico and mapped tassel branching and flowering time, two traits that exhibit clinal variation. Variation in tassel branching was dominated by a single QTL with allele effects aligning to a negative elevational cline. By contrast, allele effects associated with 11 identified flowering time QTL were not consistently correlated with any one source environmental factor. Our observations support the prediction that genotype-environment association will be strongest under simple genetic architecture, although, even then, analysis in native populations may be confounded by population structure.

MAGIC↗
PubMed1994

Clinal variation and selection of MDH allozymes in honey bee populations.

Latitudinal clines of malate dehydrogenase-1 (MDH-1) allozymes occur within honey bee populations on three continents: Europe, North America and South America. The North and South American populations are introduced and demonstrate that Mdh allelic clines were established within the last 150 years. The frequency of the 'medium' electrophoretic allele increases in frequency with increasing latitude while the 'fast' allele decreases with latitude on all the three continents. The clines are best explained by the average daily high temperature for July on all continents. These parallel clines provide evidence for selection on Mdh alleles in honey bees.

Alleles↗
PubMedDate not supplied

The association between inversion In(3R)Payne and clinally varying traits in Drosophila melanogaster.

In Drosophila melanogaster, inversion In(3R)Payne increases in frequency towards low latitudes and has been putatively associated with variation in size and thermal resistance, traits that also vary clinally. To assess the association between size and inversion, we obtained isofemale lines of inverted and standard karyotype of In(3R)Payne from the ends of the Australian D. melanogaster east coast cline. In the northern population, there was a significant association between In(3R)Payne and body size, with standard lines from this population being relatively larger than inverted lines. In contrast, the inversion had no influence on development time or cold resistance. We strengthened our findings further in a separate study with flies from populations from the middle of the cline as well as from the cline ends. These flies were scored for wing size and the presence of In(3R)Payne using a molecular marker. In females, the inversion accounted for around 30% of the size difference between cline ends, while in males the equivalent figure was 60%. Adaptive shifts in size but not in the other traits are therefore likely to have involved genes closely associated with In(3R)Payne. Because the size difference between karyotypes was similar in different populations, there was no evidence for coadaptation within populations.

Acclimatization↗
PubMed2000

The effect of epistasis on the structure of hybrid zones.

Within hybrid zones that are maintained by a balance between selection and dispersal, linkage disequilibrium is generated by the mixing of divergent populations. This linkage disequilibrium causes selection on each locus to act on all other loci, thereby steepening clines, and generating a barrier to gene flow. Diffusion models predict simple relations between the strength of linkage disequilibrium and the dispersal rate, sigma, and between the barrier to gene flow, B, and the reduction in mean fitness, W. The aim of this paper is to test the accuracy of these predictions by comparison with an exact deterministic model of unlinked loci (r = 0.5). Disruptive selection acts on the proportion of alleles from the parental populations (p,q): W = exp[-S(4pq)beta], such that the least fit genotype has fitness e-s. Where beta << 1, fitness is reduced for a wide range of intermediate genotypes; where beta >> 1, fitness is only reduced for those genotypes close to p = 0.5. Even with strong epistasis, linkage disequilibria are close to sigma 2p'ip'j/rij, where p'i, p'j are the gradients in allele frequency at loci i, j. The barrier to gene flow, which is reflected in the steepening of neutral clines, is given by [formula: see text] where r, the harmonic mean recombination rate between the neural and selected loci, is here 0.5. This is a close approximation for weak selection, but underestimates B for strong selection. The barrier is stronger for small beta, because hybrid fitness is then reduced over a wider range of p. The widths of the selected clines are harder to predict: though simple approximations are accurate for beta = 1, they become inaccurate for extreme beta because, then, fitness changes sharply with p. Estimates of gene number, made from neutral clines on the assumption that selection acts against heterozygotes, are accurate for weak selection when beta = 1; however, for strong selection, gene number is overestimated. For beta > 1, gene number is systematically overestimated and, conversely, when beta < 1, it is underestimated.

Alleles↗
PubMed1981

Latitudinal relationships of esterase-6 and phosphoglucomutase gene frequencies in Drosophila melanogaster.

Geographic variation in Esterase-6 (Est-6) and Phosphoglucomutase (Pgm) gene frequencies in Australasian populations of Drosophila melanogaster are compared with analogous data collated from 16 previous reports for North America and Europe/Asia. A large-scale latitudinal cline is found on all three zoogeographic zones for Est-6 and overall, Est-61.00 frequency increases from about 20 per cent around 20 degrees latitude to about 80 per cent approaching 50 degrees latitude. In contrast, there is no consistent evidence for the latitudinal cline in Pgm gene frequencies in any of the three zones with Pgm1.00 frequency generally about 85 per cent and Pgm1.20 and Pgm0.70 frequencies each between 5 per cent and 10 per cent. The consistent Est-6 clines are attributed to latitudinal selection gradients but not consistent correlations are found between Est-6 gene frequencies and maximum or minimum temperature or rainfall which might be associated with these gradients. The directions of the Est-6 clines in fact run counter to expectations based on the in vitro thermostabilities of the respective allozymes.

Animals↗
PubMed1997

Californian hybrid zone between Culex pipiens pipiens and Cx. p. quinquefasciatus revisited (Diptera:Culicidae).

Phallosome morphology (DV/D ratio) and allozyme variation were used to reexamine the transition from Culex pipiens pipiens L. to Cx. p. quinquefasciatus Say, detected in California from the northern Central Valley to the Mexican border of the United States of America. Significant deficiency of heterozygotes was observed at the diagnostic locus Mdhp-2 in populations from the central part of the hybrid zone. Long tails of introgression were detected: populations from both north and south ends of the transect were not genetically pure Cx. p. pipiens or Cx. p. quinquefasciatus, respectively, as previously considered, but included approximately 5% introgressed individuals. A narrow reversed cline from the Delta area into the Sacramento Valley, characterized by increasing frequencies of Cx. p. quinquefasciatus alleles proceeding to the north, was confirmed. Both these cline appear to be related mainly to temperature gradients. Over the last 50 yr, an increase in the proportion of Cx. p. pipiens DV/D phenotypes was detected proceeding north to south along the main latitudinal cline, as well as in the narrow reversed cline. Accordingly, the center of the main latitudinal hybrid zone has apparently moved approximately 100 km to the south. This phenomenon is only partially paralleled by the differentiated locus Pgm of the 3 for which comparison was possible. Similarities to and differences from previous studies are discussed, also in relation with comparable data from another hybrid zone between Cx. p. pipiens and Cx. p. quinquefasciatus recently detected in Madagascar. Hybrid index scores based on differentiated allozymes and the diagnostic locus Mdhp-2 prove to be better descriptors than the DV/D ratio of hybridization and introgression occurring between Cx. p. pipiens and Cx. p. quinquefasciatus. This seems to be caused mainly by the influence of temperature on male genitalia development, and the weaker association found between genetic markers and DV/D phenotypes in hybrid populations.

Animals↗
PubMed2002

A fine-scale spatial analysis of the mosaic hybrid zone between Gryllus firmus and Gryllus pennsylvanicus.

The pattern of character variation within a hybrid zone, the hybrid zone structure, has been used to infer the processes that maintain hybrid zones. Unfortunately it is difficult to infer process from structure alone because many different processes can produce the same pattern of character variation. Mosaic hybrid zones may be maintained by exogenous selection in a heterogeneous environment and/or endogenous selection against hybrid individuals; habitat preference, premating isolating barriers and/or fertility selection can also contribute. The spatial scale at which a hybrid zone is sampled affects its apparent structure; a hybrid zone may appear clinal at one scale and mosaic at another. Here, we sample the mosaic hybrid zone between two field crickets, Gryllus firmus and G. pennsylvanicus, at a scale that spans the boundaries between individual soil-habitat patches. From our analysis, we find that at fine scales, the mosaic hybrid zone resolves into a set of steep clines across patch boundaries. Both morphological and molecular traits exhibit sharp and generally concordant clines. However, clines for mitochondrial DNA and one anonymous nuclear marker are clearly displaced as a result of current hybridization or past introgression (the "ghost of hybridization past"). Thus, scale is important for the structure of this and probably other hybrid zones. The extremely sharp, concordant clines across patch boundaries indicate that the cricket hybrid zone is undoubtedly structured by selection. However, the detailed mechanisms responsible for the maintenance of the hybrid zone--whether endogenous selection against hybrids, exogenous selection by the environment, and/or behavioral preferences for mates or habitats--remain to be elucidated. Determining these mechanisms will depend on closer inspection of the organisms themselves and their interactions, as is the case for all hybrid zones.

Animals↗
PubMedDate not supplied

How does arthropod segment number evolve?--Some clues from centipedes.

Studies of intraspecific variation in the number of trunk segments of geophilomorph centipedes provide clues as to how different species of arthropods, and whole clades in some cases, come to be characterized by different segment numbers. However, although previous work in this area has revealed an interesting geographical pattern-a latitudinal cline in which segment number decreases with increasing latitude-the causality of the cline remains obscure. Is it because of selection on genetically based variation, or is it a result of a form of phenotypic plasticity in which the segmentation process is directly affected by a latitude-correlated factor such as temperature? Here, we provide some indirect evidence for plasticity. If the cline is indeed a plastic one, a paradox arises, because the cline mirrors interspecific variation-geophilomorph species with more northern ranges typically have fewer segments than those from further south-but interspecific differences cannot arise from nonheritable variation. We propose a resolution of this apparent paradox via a model in which genetic and environmental factors interact through selection acting on developmental reaction norms.

Animals↗
PubMed1973

Gene flow and population differentiation.

There are many possible spatial patterns of selection and gene flow that can produce a given cline structure; the actual geography of natural selection and gene flow must be worked out before an attempt is made to explain a given natural cline in terms of a model. The results of experimental and theoretical models show that it is possible for local differentiation to evolve parapatrically in spite of considerable gene flow if the selection gradients are relatively uniform. Irregularities in environmental gradients increase the sensitivity of clines to the effects of gene flow in proportion to the increase in the differences in gene frequencies between the emigrants and the demes receiving the immigrants. It is not necessary for a sharp spatial environmental change to be present for distinct differentiation to occur. In some cases even a gentle environmental gradient can give rise to marked spatial differentiation along a genetically continuous series of demes; such environmental differences may be below the practical limits of resolution in field studies. Any asymmetry in gene flow does not lead to dedifferentiation if the environmental gradient is smooth; it merely shifts the position of the transition zone between the differentiated areas from that which would be expected if there were no asymmetry. Abrupt geographic differences in gene, genotype, or morph frequencies should not, therefore, be interpreted as evidence for environmental changes in the immediate vicinity of the steepest part of the cline; neither should they be interpreted as evidence for geographic barriers, sharp environmental differences, or sexual isolation among the differentiated groups of populations when there are no other sources of evidence for these phenomena. Gene flow may be unimportant in the differentiation of populations along environmental gradients.

Animals↗
PubMed2005

Diversification before the most recent glaciation in Balanus glandula.

A deep genetic cline between southern populations of the barnacle Balanus glandula (from about Monterey Bay southward) and northern populations (from northern California through Alaska) has recently been described. If this pattern is due to historical isolation and genetic drift, we expect it to have formed recently and represent a transient, nonequilibrium state. However, this cline appears to have formed well before the last glacial maximum. Our assays of sequence diversity at a region of mitochondrial cytochrome oxidase I, combined with coalescent estimators of the time of separation for these two regions, suggest that a late Pleistocene event more than 100 thousand years ago may be responsible for the initial separation. This suggests that either strong oceanographic mechanisms or natural selection have maintained the cline, because there has clearly been adequate time for this cline or polymorphism to resolve itself by genetic drift and migration. However, reliance on only a single mitochondrial marker for which the substitution rate has been estimated still limits the resolution of our analysis.

Animals↗
PubMed2006

Hybridization of Bombina bombina and B. variegata (Anura, Discoglossidae) at a sharp ecotone in western Ukraine: comparisons across transects and over time.

Bombina bombina and B. variegata are two anciently diverged toad taxa that have adapted to different breeding habitats yet hybridize freely in zones of overlap where their parapatric distributions meet. Here, we report on a joint genetic and ecological analysis of a hybrid zone in the vicinity of Stryi in western Ukraine. We used five unlinked allozyme loci, two nuclear single nucleotide polymorphisms and a mitochondrial DNA haplotype as genetic markers. Parallel allele frequency clines with a sharp central step occur across a sharp ecotone, where transitions in aquatic habitat, elevation, and terrestrial vegetation coincide. The width of the hybrid zone, estimated as the inverse of the maximum gradient in allele frequency, is 2.3 km. This is the smallest of four estimates derived from different clinal transects across Europe. We argue that the narrow cline near Stryi is mainly due to a combination of habitat distribution and habitat preference. Adult toads show a preference for either ponds (B. bombina) or puddles (B. variegata), which is known to affect the distribution of genotypes within the hybrid zones. At Stryi, it should cause a reduction of the dispersal rate across the ecotone and thus narrow the cline. A detailed comparison of all five intensively studied Bombina transects lends support to the hypothesis that habitat distribution plus habitat preference can jointly affect the structure of hybrid zones and, ultimately, the resulting barriers to gene flow between differentiated gene pools. This study also represents a resampling of an area that was last studied more than 70 years ago. Our allele-frequency clines largely coincide with those that were described then on the basis of morphological variation. However, we found asymmetrical introgression of B. variegata genes into B. bombina territory along the bank of a river.

Animals↗
PubMed1997

Pattern of gradient of apolipoprotein E allele *4 frequencies in western Europe.

The apolipoprotein E gene (APOE) is located on chromosome 19. The three most common APOE alleles account for most of the corresponding peptide chain variations in most human populations. APOE*3 is the most common allele, coding for the product E3; APOE*2 codes for an Arg-158-->Cys substitution (E2), and APOE*4 codes for a Cys-112-->Arg product (E4). We completed a meta-analysis of APOE allele frequencies from 30 geographically defined populations in Europe, including Iceland and Turkey. We performed a weighted multiple regression using normalized geographic coordinates and a fourth-degree polynomial. Next, we constructed maps showing isofrequencies of the *4 allele in Europe. We found a clear north to south decline in *4 allele frequency for continental Western Europe. No such clinal pattern was apparent for the *2 allele frequencies, but for *3 we found an inverse south to north decreasing gradient. Symmetry between the clines of the *4 and *3 alleles is due to a negative correlation coefficient (r = -0.89). We also plotted APOE allele frequencies against latitude; a decreasing cline was evident for *4 frequencies (y = -0.152 + 0.006x, r = 0.904) and an increasing cline was evident for *3 frequencies (y = 1.087 - 0.006x, r = 0.809). Clines for the APOE alleles could be the result of natural selection.

Alleles↗
PubMed1990

NMDA receptor agonist and antagonists alter retinal ganglion cell arbor structure in the developing frog retinotectal projection.

The development of sensory maps is thought to require an activity-dependent structural rearrangement of afferent terminal arbors within the CNS which recreates the topographic relations of sensory somata present in the periphery. In the frog retinotectal projection, activation of the NMDA receptor plays a role in this structural plasticity. Exposure of the optic tectum of tadpoles to NMDA receptor antagonists results in a rearrangement of retinal ganglion cell arbors so that their organization into a topographic projection and eye-specific stripes is disrupted (Cline et al., 1987; Cline and Constantine-Paton, 1989). Exposure of the optic tectum to the receptor agonist, NMDA, increases the eye-specific segregation of these arbors (Cline et al., 1987). We examined the projection of the supernumerary retina and the morphology of individual retinal afferent arbors of untreated, NMDA-treated, APV-treated, MK801-treated, and MK801/NMDA-treated 3-eyed tadpoles and young postmetamorphic frogs in an effort to understand how NMDA receptor activation is involved in the growth and ordering of retinal arbors. Treatments with MK801 in combination with NMDA resulted in a desegregation of eye-specific stripes, whereas treatments with MK801 or NMDA alone did not. As reported previously, APV treatment resulted in stripe desegregation without increasing the tangential area (measured from 2-dimensional drawings) of the terminal arbors. However, a detailed analysis revealed that the APV-treated tadpole arbors have 35% reduction in branch density (branch tips/area) compared to untreated 3-eyed tadpole arbors. We treated the optic tectum with a range of concentrations of NMDA prepared in the slow-release plastic Elvax. NMDA at 10(-4) M in Elvax was the optimal concentration to produce the sharpening of stripe borders. Exposure of the tectum to NMDA at 10(-6) M in Elvax produced no change in the stripe pattern, while 10(-2) M NMDA in Elvax resulted in beading of the arbors. At the optimal concentration NMDA treatment results in a 75% reduction in the number of axons crossing from a stripe to an interstripe zone. Drawings of individual HRP-labeled, NMDA-treated arbors demonstrate that they have fewer branch points and fewer branch tips. NMDA treatment reduced arbor density by approximately 50%. Arbors drawn from untreated postmetamorphic frogs have twice the branch density of arbors from untreated tadpoles. NMDA treatment in these animals reduced the branch density by 55%, comparable to the reduction seen in tadpole branch density. Our data support a specific hypothesis for NMDA receptor involvement in the activity-dependent structural refinement process within the developing retinotectal projection.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed2003

Genetic diversity within the R408W phenylketonuria mutation lineages in Europe.

The R408W phenylketonuria mutation in Europe has arisen by recurrent mutation in the human phenylalanine hydroxylase (PAH) locus and is associated with two major PAH haplotypes. R408W-2.3 exhibits a west-to-east cline of relative frequency reaching its maximum in the Balto-Slavic region, while R408W-1.8 exhibits an east-to-west cline peaking in Connacht, the most westerly province of Ireland. Spatial autocorrelation analysis has demonstrated that the R408W-2.3 cline, like that of R408W-1.8, is consistent with a pattern likely to have been established by human dispersal. Genetic diversity within wild-type and R408W chromosomes in Europe was assessed through variable number tandem repeat (VNTR) nucleotide sequence variation and tetranucleotide short tandem repeat (STR) allelic associations. Wild-type VNTR-8 chromosomes exhibited two major cassette sequence organizations: (a1)5-b3-b2-c1 and (a1)5-b5-b2-c1. R408W-1.8 was predominantly associated with (a1)5-B5-B2-C1. Both wild-type vntr-3 and r408w-2.3 chromosomes exhibited a single invariant cassette sequence organization, a2-b2-c1. STR allele distributions associated with the cassette variants were consistent with greater diversity in the wild-type VNTR-8 lineage and were suggestive of different levels of diversity between R408W-1.8 and R408W-2.3. The finding of greater genetic diversity within the wild-type VNTR-8 lineage compared to VNTR-3 suggests that VNTR-8 may be older within the European population. However, in the absence of a more extensive STR data-set, no such conclusions are possible for the respective R408W mutant lineages.

Amino Acid Substitution↗
PubMed2000

Food choice behaviour may promote habitat specificity in mixed populations of clonal and sexual Potamopyrgus antipodarum.

Genetic polymorphism along an environmental gradient may be maintained if disruptive selection on habitat-specific traits leads to a correlated response in traits that reduce gene flow between habitats. We studied a short-distance cline in a population of freshwater snails Potamopyrgus antipodarum in which sexual and clonal snails coexist. Sexuals and clones show a life history cline by depth: snails reproduce at a smaller size in shallower habitats. Clones are also structured genetically across habitats and seem not to mix, even though habitats are within the dispersal distance of the snails and the opportunity for gene flow via migration must be considerable. Because habitat preference may promote divergence in both clones and sexuals along the depth gradient, we investigated whether snails show habitat-specific food choice behaviour that could reduce migration. We tested the food choice behaviour of the snails by exposing them simultaneously to food from their home and adjacent habitats. Both juvenile and adult snails from the shallow shore bank and a mid-water macrophyte habitat preferentially grazed on the vegetation of their original habitats. We suggest that the observed genetic and life history cline may be maintained by food choice behaviour that may promote a partial barrier to gene flow between the habitats. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed2004

Clinal genetic variation and the 'rare allele phenomenon' in random mating populations of Urophora cardui (Diptera: Tephritidae).

In the present study we investigate a contact zone between two population groups of the tephritid fly Urophora cardui. We investigate scenarios that may have produced the genetic differentiation of the two groups, and we describe the 'rare allele phenomenon' from the contact zone. The rare allele phenomenon refers to alleles that are found at high frequency in contact zones but are rare or lacking outside the contact zone. The phenomenon is often observed in hybrid zones between subspecies of limited reproductive compatibility, but seldom in populations with random mating. Clinal genetic variation was observed at three loci in the contact zone. Three alleles at the locus Aat showed steep clines, between 20-70 km wide. A rare Aat-A allele occurred at high frequency in the centre of the contact zone. Two further loci, Hk and Pgd, showed less steep clinal genetic variation, the transition being in and slightly south of the centre of the Aat cline. Populations showed Hardy-Weinberg proportions and there was no evidence for linkage disequlibrium. These findings suggest random mating and gradual introgression between the population systems, which may originate from at least two range expansions. Aat's steep clines and rare allele may indicate selection on Aat alleles, although we presently can not quantify any agents. Because U. cardui experiences random mating in the contact zone with no apparent 'hybrid' incompatibility, mating experiments offer the possibility for future enquiries about the genetic basis of the rare allele phenomenon.

Alleles↗