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Convergence to spatial-temporal clines in the Fisher equation with time-periodic fitnesses.

The asymptotic behavior as t----infinity of the solutions with values in the interval (0, 1) of a reaction-diffusion equation of the form (Formula: see text) is studied. Conditions on m which are satisfied when m is nonincreasing in mu and which imply that every solution converges to some periodic limit function are found. Except in some very special and well-defined circumstances, the limit is the same for all solutions, so that it is a global attractor. This global attractor may be one of the trivial solutions 0 or 1, or it may be a spatial-temporal cline. The linear stability properties of the trivial states serve to distinguish between these cases.

Mathematics↗

Isolation by distance and a chromosomal cline in the Cayapa cytospecies of Simulium exiguum, the vector of human onchocerciasis in Ecuador.

The population genetic structure of the Cayapa cytospecies of Simulium exiguum, the vector of onchocerciasis, was analysed using allozyme frequency and chromosomal inversion polymorphism data from 6 and 15 populations respectively, collected in Ecuador. Eight allozyme loci were scored. No unique allozyme markers were found enabling us to identify biting adults of the vector from the non-vector Bucay cytotype. Mannose-phosphate isomerase (Mpi) contributed largely to the significant heterogeneity in gene frequency among populations of the Cayapa cytospecies and also to the overall population structuring (F(ST) = 0.015 +/- 0.014) which fitted the isolation by distance model. However, heterozygote deficits were recorded for Mpi in four of the six populations, which could indicate that selection is acting at this locus but this hypothesis will require further convincing evidence. Furthermore the significant population structuring of allozymes was not evident when Mpi was omitted from the analysis. All inversion polymorphisms (IIS-B, IIS-F, IIL-A and IIL-B) were in Hardy-Weinberg equilibrium, showed significant heterogeneity between populations and revealed the occurrence of an altitudinal cline in inversion IIS-B frequency. The inversion polymorphisms revealed a significant degree of population structuring (F(ST) = 0.083 +/- 0.027), which can be explained by the isolation by distance model. A UPGMA cluster analysis revealed the relatively remote, high altitude Rio Mira populations to be the most genetically distinct.

Animals↗

A second look at some data on a cline.

The release-recapture data given in table 1 of Kettlewell et al.'s (1969) paper on a cline in the frequencies of the typica and edda morphs of the moth. Amathes glareosa have been re-analysed. Estimates of daily survival probabilities have been calculated and these have been "explained" in terms of differences between localities of release and the morph involved. The conclusions of Kettlewell et al. (1969) with regard to the survival of the moths have not been confirmed.

Animals↗

Morphological variation in a natural population of Drosophila mediopunctata: altitudinal cline, temporal changes and influence of chromosome inversions.

To characterize the morphological variation in a natural population of Drosophila mediopunctata, males were collected on three occasions at a single locality. From each wild-caught male 14 body measures were taken and the karyotype for inversions on chromosomes X and II was determined. Through a principal components analysis, two sources of variation, identified as size and shape, accounted for approximately 80 and 6 per cent of the total morphological variability, respectively. The shape component was determined primarily by variations in the position of the wing second longitudinal vein. Differences between collections were detected both for size and shape. An altitudinal cline was observed in respect of wing shape, although altitude explained only a small part of the shape variation. Size and shape were affected by chromosome II inversions. However, in respect of size, no direct differences were detected between karyotypes but a significant interaction between collecting date and karyotype was found. This suggests that karyotypes might differ in their norms of reaction in the field.

Altitude↗

Evidence for a geographical cline of casein haplotypes in Portuguese cattle breeds.

Genetic variants of bovine milk proteins have been intensively used to characterize breeds and as markers for population/QTL studies throughout the world. However, a large number of cattle breeds including those found in Portugal, remain unstudied. In this work, we have analysed the genetic variation of six milk protein loci in 10 Portuguese cattle breeds by isoelectric focusing. High genetic diversities were generally found across breeds, with the exception of Mirandesa that showed a trend to fixation of the most common alleles in five loci, as well as of the rarer CSN3B allele. The casein haplotype BA2A was often the most frequent, followed by haplotypes BA2B and BA1A. Remarkably, CA2A was found to be the second most frequent haplotype in Southern breeds, supporting a geographical cline between Central-Northern European breeds and Bos indicus populations. Our data suggest that high genetic similarity among neighbouring Portuguese breeds is mainly caused by gene flow, and that the geographical distribution of particular casein haplotypes may indicate an influence of African cattle.

Animals↗

Clines of nuclear DNA markers suggest a largely neolithic ancestry of the European gene pool.

Comparisons between archaeological findings and allele frequencies at protein loci suggest that most genes of current Europeans descend from populations that have been expanding in Europe in the last 10, 000 years, in the Neolithic period. Recent mitochondrial data have been interpreted as indicating a much older, Paleolithic ancestry. In a spatial autocorrelation study at seven hypervariable loci in Europe (four microsatellites, two larger, tandem-repeat loci, and a sequence polymorphism) broad clinal patterns of DNA variation were recognized. The observed clines closely match those described at the protein level, in agreement with a possible Near Eastern origin for the ancestral population. Separation times between populations were estimated on the basis of a stepwise mutation model. Even assuming low mutation rates and long generation times, we found no evidence for population splits older than 10,000 years, with the predictable exception of Saami (Lapps). The simplest interpretation of these results is that the current nuclear gene pool largely reflects the westward and northward expansion of a Neolithic group. This conclusion is now supported by purely genetic evidence on the levels and patterns of microsatellite diversity, rather than by correlations of biological and nonbiological data. We argue that many mitochondrial lineages whose origin has been traced back to the Paleolithic period probably reached Europe at a later time.

Cell Nucleus↗

Imbalance of predator and prey armament: geographic clines in phenotypic interface and natural selection.

The escalation of defensive/offensive arms is ubiquitous in prey-predator evolutionary interactions. However, there may be a geographically varying imbalance in the armaments of participating species that affects the outcome of local interactions. In a system involving the Japanese camellia (Camellia japonica) and its obligate seed predator, the camellia weevil (Curculio camelliae), we investigated the geographic variation in physical defensive/offensive traits and that in natural selection on the plant's defense among 17 populations over a 700-km-wide area in Japan. The sizes of the plant defensive apparatus (pericarp thickness) and the weevil offensive apparatus (rostrum length) clearly correlated with each other across populations. Nevertheless, the balance in armaments between the two species was geographically structured. In the populations for which the balance was relatively advantageous for the plant's defense, natural selection on the trait was stronger because in the other populations, most plant individuals were too vulnerable to resist the attacks of the weevil, and their seeds were infested independent of pericarp thickness. We also found that the imbalance between the defensive/offensive armaments and the intensity of natural selection showed clear latitudinal clines. Overall, our results suggest that the imbalance of armament between sympatric prey and predator could determine the strength of local selection and that climatic conditions could affect the local and overall trajectory of coevolutionary arms races.

Animals↗

Gene flow and selection in a cline.

A model of the effect of gene flow and natural selection in a continuously distributed, infinite population is developed. Different patterns of spatial variation in selective pressures are considered, including a step change in the environment, a "pocket" in the environment and a periodically varying environment. Also, the problem of the effect of a geographic barrier to dispersal is analyzed. The results are: (1) there is a characteristic length scale of variation of gene frequencies, (see PDF). The population cannot respond to changes in environmental conditions which occur over a distance less than the characteristic length. The result does not depend either on the pattern of variation in selective pressures or on the exact shape of the dispersal function. (2) The reduction in the fitness of the heterozygote causes a cline in gene frequencies to become steeper. (3) A geographic barrier to dispersal causes a drastic change in the gene frequencies at the barrier only when almost all of the individuals trying to cross the barrier are stopped.

Environment↗

Patterns of variation within and between Greek populations of Ceratitis capitata suggest extensive gene flow and latitudinal clines.

Four natural Greek populations of Mediterranean fruit fly, Ceratitis capitata (Wiedemann), was studied for genetic variability at 25 enzyme loci. The comparison of polymorphism within and between populations shows two loci with high between-population heterozygosity (HT) and very high fixation index (F(ST)) values, suggesting the presence of balancing selection. The gradual decline of common allele frequency of the polymorphic loci tested indicated that latitudinal clines are present in Greece. Indirect estimates of gene flow based both on Wright's method (Nm*) and on the Slatkin's method (Nm*), which depends on the frequencies of rare alleles found in only one population, revealed a substantial amount of gene flow (Nm = 3.493 and Nm* = 3.197). These estimates of gene flow may well explain why the "introduced" Greek populations of C. capitata, in spite of their low genetic variability, display the same polymorphic loci. Gene flow in combination with natural selection and genetic drift may have played an important role to genetic differentiation in this species in Greece.

Alleles↗

A cline in frequency of autosomal males is not associated with insecticide resistance in house fly (Diptera: Muscidae).

Geographic variation in the chromosomal location of the male sex determining factor (M) was studied in four house fly, Musca domestica L., populations from the eastern United States. We found a strong clinal trend (29 degrees 41' latitude in Florida to 44 degrees 2' in Maine) in which the percentage of standard XY(M) males increased with increasing latitude. In Florida, 100% of the males possessed the M factor on the third autosome (III(M)). North Carolina had 20% III(M) males and 2.35% with both Y(M) and III(M). Fewer III(M) males were found in New York (4.35%). Populations from Maine contained 100% XY(M) males. In two of three standard laboratory-susceptible strains, all males carried M on an autosome ("autosomal males" or A(M)): CS (III(M)) and SRS (V(M)). Insecticide bioassays of four field-collected strains led us to conclude that resistance is not correlated with sex determination over a broad range of insecticides. For example, high levels of resistance to permethrin (86-99% survival at a diagnostic concentration) were found in all four field-collected strains. The five other insecticides evaluated showed varying levels of resistance among field strains. We conclude that a cline is present in house fly populations from the eastern United States with 100% III(M) males in the south and entirely Y(M) males in the north and that insecticide resistance is not a key factor influencing the evolution or linkage of M.

Animals↗

The Drosophila heat shock hsr-omega gene: an allele frequency cline detected by quantitative PCR.

The hsr-omega gene of Drosophila melanogaster produces RNA products both constitutively and at elevated levels in response to heat stress. A single-nucleotide difference in this gene that has been detected using denaturing gradient gel electrophoresis (DGGE) is responsible for an hsr-omegaa/b polymorphism, and selection experiments have indicated an association between the hsr-omegaa allele and susceptibility to heat stress. Since allele frequency estimates for population surveys using PCR and DGGE for single flies would be relatively time-consuming and expensive, we here develop a quantitative competitive-PCR method using mass-grind genomic DNA preparations for this purpose. Geographical and temporal variation of allele frequency at the hsr-omega locus in Australian populations of D. melanogaster are examined. Regular samples from a southern population through a summer season suggested stability of hsr-omegaa frequency. Field populations sampled from a approximately 2,250 km north-south transect along eastern Australia revealed a strong positive association between the frequency of hsr-omegaa and latitude, and marked spatial autocorrelation. Using appropriate analyses, strong association between population differences in hsr-omegaa frequencies and differences in temperature and rainfall measures, after controlling for latitudinal differences, support the idea that the cline in hsr-omegaa frequency may be attributable to some form of climatic selection.

Alleles↗

Phenotypic clines, plasticity, and morphological trade-offs in an intertidal snail.

Understanding the genetic and environmental bases of phenotypic variation and how they covary on local and broad geographic scales is an important goal of evolutionary ecology. Such information can shed light on how organisms adapt to different and changing environments and how life-history trade-offs arise. Surveys of phenotypic variation in 25 Littorina obtusata populations across an approximately 400-km latitudinal gradient in the Gulf of Maine revealed pronounced clines. The shells of snails from northern habitats weighed less and were thinner and weaker in compression than those of conspecifics from southern habitats. In contrast, body size (as measured by soft tissue mass) followed an opposite pattern; northern snails weighed more than southern snails. A reciprocal transplant between a northern and southern habitat revealed substantial plasticity in shell form and body mass and their respective measures of growth. Southern snails transplanted to the northern habitat produced lighter, thinner shells and more body mass than controls raised in their native habitat. In contrast, northern snails transplanted to the southern site produced heavier, thicker shells and less body mass than controls raised in their native habitat. Patterns of final phenotypic variation for all traits were consistent with cogradient variation (i.e., a positive covariance between genetic and environmental influences). However, growth in shell traits followed a countergradient pattern (i.e., a negative covariance between genetic and environmental influences). Interestingly, body growth followed a cogradient pattern, which may reflect constraints imposed by cogradient variation in final shell size and thickness. This result suggests the existence of potential life-history trade-offs associated with increased shell production. Differences in L. obtusata shell form, body mass, and their respective measures of growth are likely induced by geographic differences in both water temperature and the abundance of an invading crab predator (Carcinus maenas). Water temperatures averaged 6.8 degrees C warmer during the transplant experiment and C. maenas abundance is greater in the southern Gulf of Maine. Because both increased water temperature and crab effluent affect shell form in the same way, future experiments are needed to determine the relative importance of each. Nevertheless, it is clear that phenotypic plasticity has an important role in producing geographic variation in L. obtusata shell form. Moreover, the evolution of phenotypic plasticity in L. obtusata and other marine gastropods may be driven by architectural constraints imposed by shell form on body mass and growth.

Animals↗

Staggered clines in a hybrid zone between two chromosome races of the harvestman Gagrellopsis nodulifera (Arachnida: Opiliones).

We analyzed a hybrid zone between two chromosome races (2n = 16 and 2n = 22) of a Japanese harvestman, Gagrellopsis nodulifera Sato and Suzuki (Arachnida: Opiliones: Phalangiidae). The hybrid zone is located in the eastern part of Tottori Prefecture, western Honshu. The width of the zone is approximately 5 to 15 km. Three independent tandem fusions/fissions seem to be the main cause of the karyotypic differences between the parental races. Ten karyotypic variants were found in the hybrid zone. They differed by numbers of diploid chromosomes and trivalents detected in meiosis. In most of the collecting sites, karyotypic heterozygotes were less common than expected. A positive correlation was found between number of trivalents in a karyotype and its deficiency rate. In some sites, the deficit of heterozygous individuals was accompanied by an excess of the intermediate homozygotes. One of the three transects across the zone was studied in detail. We found that three types of single heterozygotes (2n = 17, 2n = 19 and 2n = 21) formed a series of successive, spatially separated peaks along the transect. Two types of intermediate homozygotes (2n = 18 and 2n = 20) were also spatially separated. The most parsimonious explanation of such a structure is the staggering of clines of three tandem (or Robertsonian) fusion/fission variants that differentiate the parental races caused by selection against multiple heterozygotes. Analysis of nondisjunction in single heterozygotes demonstrated that there was a strong interindividual variation in nondisjunction rate. The mean frequency of aneuploid MII in single heterozygotes was 0.10 +/- 0.03. Crossover exchanges in some critical regions of trivalents result in abnormal chromosomal configurations: chromosomes with unequal chromatids and dicentric chromosomes. Frequency of crossover-induced chromosomal abnormalities was low in single heterozygotes (approximately equal to 4%), and was unexpectedly high in the double heterozygotes (approximately equal to 15%). Selection against karyotypic heterozygotes is considered as a main evolutionary force responsible for the structuring of the hybrid zone. A positive association between diploid chromosome number and altitude was found. The race 2n = 16 tended to occupy lower altitudes than the 2n = 22 parental race. Differences in ecological preferences may be a result of previous adaptations to different environments in allopatry. A hypothesis concerning the origin and evolution of the hybrid zone is proposed.

Animals↗

Genetic differentiation and inversion clines in Indian natural populations of Drosophila melanogaster.

To study the genetic differentiation and inversion clines in Indian natural populations of Drosophila melanogaster, 14 natural populations (6 from the north and 8 from the south) were screened for chromosome inversions. The chromosomal analysis revealed the presence of 23 paracentric inversions, which include 4 common cosmopolitan, 4 rare cosmopolitan, 2 recurrent endemic, and 13 unique endemic (new inversions detected for the first time) inversions. The difference in karyotype frequencies between populations from the north and south were highly significant and the level of inversion heterozygosity was higher in populations from the south. Statistically significant negative correlations were found between each of the four common cosmopolitan inversions and latitude. These findings are in accord with results from other worldwide geographic regions and show that Indian populations of D. melanogaster have undergone considerable genetic differentiation at the level of inversion polymorphism.

Animals↗

Allele frequencies of human complement factor I in a sample from Iwate, northern Japan, with the description of geographical cline.

Serum samples from 270 healthy blood donors of Iwate prefecture, northern Japan, were examined for polymorphism of factor I (IF) by using polyacrylamide gel isoelectric focusing followed by semidry horizontal electroblotting with enzyme immunoassay. In 270 individuals four different patterns were observed, and these were controlled by two common alleles, IF*A and IF*B, and one rare allele, IF*A2. Allele frequencies were estimated to be 0.1019, 0.8963 and 0.0018 for IF*A, IF*B and IF*A2, respectively. The data of IF allele frequencies thus far reported in Japan excluding Okinawa Island were compared, and a statistically significant (p less than 0.01) geographical cline was detected for IF*A and IF*B alleles.

Alleles↗

Alpha-2-HS-glycoprotein gene frequencies in Galicia: further evidence for a cline in European populations.

alpha 2-HS-glycoprotein (AHSG) phenotypes were determined in 506 unrelated autochthonous individuals from the Galician population (NW Spain) by means of isoelectric focusing followed by silver stain immunofixation. The phenotype frequency distribution fitted with the Hardy-Weinberg law. No evidence of intrapopulation heterogeneity was observed. The allele frequencies obtained were: AHSG*1 = 0.7559 +/- 0.013 and AHSG*2 = 0.2441 +/- 0.013. Our findings lend support to the hypothesis of an AHSG gene frequency cline in European populations. Other population genetics' considerations have also been discussed.

Alleles↗

Clines, clusters, and the effect of study design on the inference of human population structure.

Previously, we observed that without using prior information about individual sampling locations, a clustering algorithm applied to multilocus genotypes from worldwide human populations produced genetic clusters largely coincident with major geographic regions. It has been argued, however, that the degree of clustering is diminished by use of samples with greater uniformity in geographic distribution, and that the clusters we identified were a consequence of uneven sampling along genetic clines. Expanding our earlier dataset from 377 to 993 markers, we systematically examine the influence of several study design variables--sample size, number of loci, number of clusters, assumptions about correlations in allele frequencies across populations, and the geographic dispersion of the sample--on the "clusteredness" of individuals. With all other variables held constant, geographic dispersion is seen to have comparatively little effect on the degree of clustering. Examination of the relationship between genetic and geographic distance supports a view in which the clusters arise not as an artifact of the sampling scheme, but from small discontinuous jumps in genetic distance for most population pairs on opposite sides of geographic barriers, in comparison with genetic distance for pairs on the same side. Thus, analysis of the 993-locus dataset corroborates our earlier results: if enough markers are used with a sufficiently large worldwide sample, individuals can be partitioned into genetic clusters that match major geographic subdivisions of the globe, with some individuals from intermediate geographic locations having mixed membership in the clusters that correspond to neighboring regions.

Alleles↗

Mitochondrial cytochrome B DNA variation in the high-fecundity atlantic cod: trans-atlantic clines and shallow gene genealogy.

An analysis of sequence variation of 250 bp of the mitochondrial cytochrome b gene of 1278 Atlantic cod Gadus morhua ranging from Newfoundland to the Baltic shows four high-frequency (>8%) haplotypes and a number of rare and singleton haplotypes. Variation is primarily synonymous mutations. Natural selection acting directly on these variants is either absent or very weak. Common haplotypes show regular trans-Atlantic clines in frequencies and each of them reaches its highest frequency in a particular country. A shallow multifurcating constellation gene genealogy implies young age and recent turnover of polymorphism. Haplotypes characterizing populations at opposite ends of the geographic distribution in Newfoundland and the Baltic are mutationally closest together. The haplotypes are young and have risen rapidly in frequency. Observed differentiation among countries is due primarily to clinal variation. Hypotheses of historical isolation and polymorphisms balanced by local selection and gene flow are unlikely. Instead the results are explained by demic selection of mitochondria carried by highly fit females winning reproductive sweepstakes. By inference the Atlantic cod, a very high-fecundity vertebrate, is characterized by a high variance of offspring number and strong natural selection that leads to very low effective to actual population sizes.

Analysis of Variance↗