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A comparison of nuclear and mitochondrial cline shapes in a hybrid zone in the Sceloporus grammicus complex (Squamata; Phrynosomatidae).

The F5 and FM2 chromosome races of the Sceloporus grammicus complex form a hybrid zone in the Mexican state of Hidalgo. Previous studies of this zone have assessed genetic structure by averaging estimates of shape and width across three diagnostic chromosome markers. This approach is likely to mask subtle differences in cline shape among loci (e.g. selected vs. neutral), and obscure any displacement of cline centres (if present). Here we use maximum likelihood methods to construct the best fitting individual clines for three chromosomal markers, and also add two new markers; the mitochondrial DNA (mtDNA) locus, and the nuclear ribosomal DNA (rDNA) repeat. For each locus, hybrid zone models were fitted by cline shape and width, and the position and number of segments describing the centre of the zone. Pairwise comparisons between all clines revealed concordance between chromosomes 2 and 6, but significant discordance in cline structure among all other paired combinations. The concordance of chromosomes 2 and 6 suggests that these clines are maintained by genome-wide forces. The discordance of the chromosome 1 cline suggests an influence of asymmetric introgression, while the mtDNA cline is probably influenced by selection and drift. The rDNA locus reveals a pattern best explained by either extreme asymmetric introgression or gene conversion. The structure of zone indicates that genome-wide processes and locus specific selective forces as well as drift, are operating to different degrees on different loci. The locus-by-locus approach used here permits a finer discrimination among possible mechanisms responsible for the maintenance of the individual clines.

Animals↗

A comparison of the genetic basis of wing size divergence in three parallel body size clines of Drosophila melanogaster.

Body size clines in Drosophila melanogaster have been documented in both Australia and South America, and may exist in Southern Africa. We crossed flies from the northern and southern ends of each of these clines to produce F(1), F(2), and first backcross generations. Our analysis of generation means for wing area and wing length produced estimates of the additive, dominance, epistatic, and maternal effects underlying divergence within each cline. For both females and males of all three clines, the generation means were adequately described by these parameters, indicating that linkage and higher order interactions did not contribute significantly to wing size divergence. Marked differences were apparent between the clines in the occurrence and magnitude of the significant genetic parameters. No cline was adequately described by a simple additive-dominance model, and significant epistatic and maternal effects occurred in most, but not all, of the clines. Generation variances were also analyzed. Only one cline was described sufficiently by a simple additive variance model, indicating significant epistatic, maternal, or linkage effects in the remaining two clines. The diversity in genetic architecture of the clines suggests that natural selection has produced similar phenotypic divergence by different combinations of gene action and interaction.

Animals↗

Estimates of selection and gene flow from measures of cline width and linkage disequilibrium in heliconius hybrid zones.

Hybrid zones can yield estimates of natural selection and gene flow. The width of a cline in gene frequency is approximately proportional to gene flow (sigma) divided by the square root of per-locus selection (square root of s). Gene flow also causes gametic correlations (linkage disequilibria) between genes that differ across hybrid zones. Correlations are stronger when the hybrid zone is narrow, and rise to a maximum roughly equal to s. Thus cline width and gametic correlations combine to give estimates of gene flow and selection. These indirect measures of sigma and s are especially useful because they can be made from collections, and require no field experiments. The method was applied to hybrid zones between color pattern races in a pair of Peruvian Heliconius butterfly species. The species are Müllerian mimics of one another, and both show the same changes in warning color pattern across their respective hybrid zones. The expectations of cline width and gametic correlation were generated using simulations of clines stabilized by strong frequency-dependent selection. In the hybrid zone in Heliconius erato, clines at three major color pattern loci were between 8.5 and 10.2 km wide, and the pairwise gametic correlations peaked at R approximately 0.35. These measures suggest that s approximately 0.23 per locus, and that sigma approximately 2.6 km. In erato, the shapes of the clines agreed with that expected on the basis of dominance. Heliconius melpomene has a nearly coincident hybrid zone. In this species, cline widths at four major color pattern loci varied between 11.7 and 13.4 km. Pairwise gametic correlations peaked near R approximately 1.00 for tightly linked genes, and at R approximately 0.40 for unlinked genes, giving s approximately 0.25 per locus and sigma approximately 3.7 km. In melpomene, cline shapes did not perfectly fit theoretical shapes based on dominance; this deviation might be explained by long-distance migration and/or strong epistasis. Compared with erato, sample sizes in melpomene are lower and the genetics of its color patterns are less well understood. In spite of these problems, selection and gene flow are clearly of the same order of magnitude in the two species. The relatively high per locus selection coefficients agree with "major gene" theories for the evolution of Müllerian mimicry, but the genetic architecture of the color patterns does not. These results show that the genetics and evolution of mimicry are still only sketchily understood.

Animals↗

Cellular basis of wing size variation in Drosophila melanogaster: a comparison of latitudinal clines on two continents.

We investigated the cellular basis of two extensive, continuous, latitudinal, genetic, body size clines of Drosophila melanogaster by measuring wing area and cell size in the wing blade of adult flies reared under standard, laboratory conditions. We report that the contribution of cell size to an Australian cline is much smaller than that to a South American cline. The data suggest that neither cell size nor cell number were the targets of selection, but rather wing area itself, or a trait closely related to it. We hypothesize that the differences between the continents were caused by differences in the initial pattern of genetic variation for the cell traits and/or by the direction of selection on the source populations of the clines. Despite large differences between continents in the cellular basis of the latitudinal variation, multiple regression analysis, using the individual variation within populations, showed that the relationship between cell size and cell number was changed with latitude in the same way in the two clines. The relative contribution of cell number to wing area variation increased with latitude, probably because of compensatory interactions with cell size as a consequence of the latitudinal increase in cell number. Our findings are discussed in relation to the cellular basis of evolutionary change in laboratory thermal selection lines and natural populations along latitudinal clines.

Analysis of Variance↗

Molecular analysis of an allozyme cline: alcohol dehydrogenase in Drosophila melanogaster on the east coast of North America.

Clines may either be selectively maintained or be the by-product of nonadaptive processes related to population structure and history. Drosophila melanogaster populations on the east coast of North America show a latitudinal cline in the frequencies of two common electrophoretically distinguishable alleles at the alcohol dehydrogenase locus (Adh), designated Adh-S and Adh-F. This cline may either be adaptive or an artifact of a possible recent dual founding of North American D. melanogaster populations in which frequencies of Adh alleles differed between founder populations. By means of a high resolution restriction-mapping technique, we studied the distribution of 113 haplotypes derived from 44 polymorphic DNA markers within the Adh region in 1533 individuals from 25 populations throughout the cline. We found significant clinal differentiation at the polymorphism determining the mobility-difference causing amino acid replacement between Adh-F and Adh-S alleles. Hitchhiking was limited, despite extensive linkage disequilibrium, and other sites did not vary clinally. Such a pattern of differentiation implies that selection is responsible for the cline. To investigate whether selection acts only on the Adh-F/S site, we performed a "selective equivalence" test under the assumption that all variability within the specified allelic class is selectively neutral. This revealed selective equivalence among Adh-S-bearing haplotypes, whose frequencies showed no differentiation throughout the cline, implying high levels of frequency-homogenizing gene flow. Geographical heterogeneity among Adh-F-bearing haplotypes implied the action of selection on one or more additional variants in linkage disequilibrium with Adh-F. In a further study of a subset of the data (n = 1076 from 18 populations), we found a combined insertion/deletion polymorphism, designated delta 1, located in the 5' adult intron and in linkage disequilibrium with Adh-F, to show more marked clinal variation than Adh-F/S. Although the unequivocal identification of the precise target(s) of selection requires further study, we suggest that clinal selection may be acting epistatically on the Adh-F/S and delta 1 polymorphisms.

Alcohol Dehydrogenase↗

Rapid evolution of a geographic cline in size in an introduced fly.

The introduction and rapid spread of Drosophila subobscura in the New World two decades ago provide an opportunity to determine the predictability and rate of evolution of a geographic cline. In ancestral Old World populations, wing length increases clinally with latitude. In North American populations, no wing length cline was detected one decade after the introduction. After two decades, however, a cline has evolved and largely converged on the ancestral cline. The rate of morphological evolution on a continental scale is very fast, relative even to rates measured within local populations. Nevertheless, different wing sections dominate the New versus Old World clines. Thus, the evolution of geographic variation in wing length has been predictable, but the means by which the cline is achieved is contingent.

Animals↗

Gene frequency clines produced by kin-structured founder effects.

Two mechanisms are commonly invoked to explain geographic gradients in gene frequencies (clines): gene flow and natural selection. Thus gene frequency clines in Europe have been attributed to gene flow because expanding Neolithic farmers from the Near East spread and absorbed resident foragers, a process Cavalli-Sforza and his colleagues labeled demic diffusion. Alternatively, gradients in natural selection can produce the same genetic pattern. A third mechanism to explain European clines has recently been proposed by Barbujani et al. (1995). They found that European allele frequencies were consistent with distributions generated by a simple model of total replacement of foragers by expanding farmers. Clines were produced by repeated founder effects in colonizing farmer groups. Here, I present a simulation model using different population parameters to test the generality of the Barbujani model. Results of the simulation show that steep clines (measured by spatial autocorrelation statistics) can be produced by kin-structured founder effects. The implication of these findings is that several genetic mechanisms are consistent with the allele distributions in Europe. To discriminate among these alternatives, other information, such as better archeological and demographic characterization of interactions between farmers and foragers, is required. In any case, the mere presence of genetic clines does not necessarily validate the demic diffusion model.

Demography↗

Worldwide latitudinal clines for the alcohol dehydrogenase polymorphism in Drosophila melanogaster: what is the unit of selection?

Geographical clines may reflect the action of natural selection on genetic polymorphisms. In Drosophila melanogaster several latitudinal clines occur for many characters like allozymes, inversions and quantitative traits. The identical nature of these clines on the various continents, both on the Northern and Southern Hemispheres strongly suggests adaptation to specific stress factors. The alcohol dehydrogenase (Adh) polymorphism shows high frequencies of the S allele in tropical regions and declines with latitude. The reasons for this cline are difficult to determine because of the entanglement with other polymorphisms varying with latitude. In this paper the tentative connections with other polymorphisms like alpha-Gpdh, In(2L)t, body size and development time are reviewed with respect to the possible environmental stress factors involved. It is concluded, also from recent experiments, that the (2L)t inversion plays a dominant role in resistance to high temperature and is partly responsible for the Adh cline. Further research is aimed at the specific selective forces acting on Adh, focussing on the physiological and life history aspects.

Alcohol Dehydrogenase↗

Observations on the extent and temporal stability of latitudinal clines for alcohol dehydrogenase allozymes and four chromosome inversions in Drosophila melanogaster.

Previously we have presented evidence of large-scale latitudinal clines in the frequencies of four chromosome inversions and alleles at six enzyme loci in populations of D. melanogaster in Australasia, Asia and North America. Subsequent sampling by others in Japan and western U.S.A. has failed to repeat this observation for the steepest of the clines (alcohol dehydrogenase and the four chromosome inversions). We argue that this failure reflects the few populations and small latitudinal range sampled in these later studies. From extensive sampling over a long latitudinal transect in Australasia we here document Adh and inversion clines which are virtually identical to those originally obtained in different Australian populations four years earlier. We also repeat our observation that the Adh cline is largely independent of the cline in the linked inversion In(2L)t. We therefore retain our original conclusion that these polymorphisms are subject to natural selection. However the new Australasian data do not indicate an association between Adh and maximum rainfall which had been evident in the earlier data for Australasia, Asia and North America. We therefore retract our claim that the selective agent on Adh is related to rainfall.

Alcohol Dehydrogenase↗

Evidence for a link between local and seasonal cycles in gene frequencies and latitudinal gene clines in a cyclic parthenogen.

In an earlier study (Rhombergh et al., Can. J. Genet. Cytol. 27: 224-232, 1985) of natural populations of the cyclic parthenogenetic Rose aphids, Macrosiphum rosae, 6 out of 31 loci were found to be polymorphic and one locus (Esterase-4) showed cyclic seasonal changes in gene and genotypic frequencies. Assuming that the Est-4 polymorphism was balanced and due to some climatic factor that varies seasonally, and realizing that most environmental factors that vary seasonally also vary latitudinally, we predicted existence of a latitudinal gene cline at this locus. In the present study we surveyed four polymorphic loci (chosen to be used as markers) in six geographic populations spanning over 1200 km between the United States and Canada and found all four loci to have latitudinal clines. We think that the gene clines are due to a latitudinal cline in the degree of advancement of local populations through the seasonal cycle, and have called such a pattern a 'seasonal phase cline'. The results are discussed in relation to the temporal instability of local patterns and persistence of genetic variability on the large scale in aphids. It is argued that population structure of aphids makes retention of selectively neutral or weakly selected polymorphisms difficult.

Animals↗

Assessment of the ratio of pollen to seed flow in a cline for genetic variation in a quantitative trait.

A dispersal-selection cline model is analysed to evaluate the role of the ratio of pollen to seed flow (r) in spatial genetic variation, with a focus on clines in additive and dominant variances of major genes affecting a quantitative trait, assuming one locus with two alleles, no genetic drift and no mutation. It is shown that under weak selection, steady-state departures from the value at Hardy-Weinberg equilibrium (HWE) for genotype frequency (D) and additive variance (DeltaVa) generally display a small value on one side of the selection boundary point, a high value around the point, and a moderate value on the other side of the boundary point. A large value of r can enhance formation of this pattern. However, this is not the case for clines in the average effect of a gene substitution (Deltaalpha) or in dominance variance (DeltaVd), where large values of r can eradicate the clines. Direct use of their values at HWE to approximate the real case is acceptable. There is an expected turning point that divides clines of either additive variance or dominance variance into two subclines, each with a shorter width than that of allele frequency. Integration of these properties can help to indicate the existence of major genes affecting a quantitative trait.

Gene Frequency↗

Inference from clines stabilized by frequency-dependent selection.

Frequency-dependent selection against rare forms can maintain clines. For weak selection, s, in simple linear models of frequency-dependence, single locus clines are stabilized with a maximum slope of between square root of s/square root of 8 sigma and square root of s/square root of 12 delta, where sigma is the dispersal distance. These clines are similar to those maintained by heterozygote disadvantage. Using computer simulations, the weak-selection analytical results are extended to higher selection pressures with up to three unlinked genes. Graphs are used to display the effect of selection, migration, dominance, and number of loci on cline widths, speeds of cline movements, two-way gametic correlations ("linkage disequilibria"), and heterozygote deficits. The effects of changing the order of reproduction, migration, and selection, are also briefly explored. Epistasis can also maintain tension zones. We show that epistatic selection is similar in its effects to frequency-dependent selection, except that the disequilibria produced in the zone will be higher for a given level of selection. If selection consists of a mixture of frequency-dependence and epistasis, as is likely in nature, the error made in estimating selection is usually less than twofold. From the graphs, selection and migration can be estimated using knowledge of the dominance and number of genes, of gene frequencies and of gametic correlations from a hybrid zone.

Computer Simulation↗

Conditions for the existence of clines.

A very general partial differential equation in space and time satisfied by the gene frequency in a monoecious population distributed continuously over an arbitrary habitat is derived. The treatment is restricted to a single diallelic locus in the absence of mutation and random drift, and it is supposed that time is continuous, births and deaths occur at random, and migration is independent of genotype. With the further assumptions that migration is isotropic and homogeneous, the population density is constant and uniform (as permitted by the population regulation mechanism included in the formulation), and Hardy-Weinberg proportions obtain locally, this partial differential equation reduces to the simplest multidimensional generalization of the classical Fisher-Haldane cline model. The efficacy of migration and selection in maintaining genetic variability at equilibrium in this model is investigated by deducing conditions for the existence of clines under various circumstances. The effects of the degree of dominance, a neutral belt between the regions where a particular allele is advantageous and deleterious, finiteness of the habitat, and habitat dimensionality are evaluated. Provided at least one of the alleles is favored only in a finite region, excluding the special case in which its total effective selective coefficient is zero, if conditions for supporting a cline are too unfavorable because migration is too strong, selection is too weak, or both, a cline cannot exist at all. Thus, unless there is overdominance, the population must be monomorphic. It is possible for a cline which can barely exist under the prevailing ecological circumstances to show a large amount of variation in gene frequency.

Alleles↗

A hybrid zone comprising staggered chromosomal clines in the house mouse (Mus musculus domesticus).

In the vicinity of John o'Groats (Caithness, Scotland) there is a small karyotypic race of the house mouse (Mus musculus domesticus) characterized by a diploid number of 32 chromosomes, including the metacentrics 4.10, 9.12, 6.13 and 11.14. This race forms a hybrid zone with the standard British race (fully acrocentric chromosomes, 2n = 40). Although hybrid zones normally consist of several (or many) narrow character clines at the same position, this zone is unusual in that the chromosomal clines do not coincide. The cline for arm combination 11.14 is staggered relative to the 6.13 cline and both are separate from the clines for 4.10 and 9.12 (which may or may not coincide). A variety of explanations for the structure of the hybrid zone are discussed. It is possible that this may be a case of 'zonal raciation'.

Animals↗

Starvation resistance and adult body composition in a latitudinal cline of Drosophila melanogaster.

Latitudinal geographic variation in Drosophila melanogaster is pervasive. Parallel clines in traits such as body size, egg size, ovariole number, and development time have been found on several continents throughout the world. However, a cline in starvation resistance and fat content in D. melanogaster has so far been found only in India. Here we investigate starvation resistance and fat content in 10 populations from South America, in which clines in body size, egg size, and development time have previously been found. We find no evidence for a cline in starvation resistance or fat content in South America. We therefore suggest that the cline in starvation resistance in India may have evolved in response to specific climatic variation found only in India.

Animals↗

The amplitude of circadian oscillations: temperature dependence, latitudinal clines, and the photoperiodic time measurement.

This paper develops several propositions concerning the lability of the amplitude of Drosophila circadian pacemakers. The first is that the amplitude of the pacemaker's motion, unlike its period, is markedly temperature-dependent. The second is that latitudinal variation in pacemaker amplitude (higher in the north) is responsible for two very different sets of observations on Drosophila circadian systems at successively higher latitudes. One of these is a cline in D. auraria's phase-shifting response to light, which steadily weakens in a succession of more northerly strains. The other, concerning D. littoralis in the very far north, is a cline in the rate at which eclosion activity becomes arrhythmic (the circadian rhythm damps out) in constant darkness; damping is faster in the north. The third proposition concerns a plausible selection pressure for the cline in pacemaker amplitude that we propose underlies the two directly observed clines. Two points are emphasized: (1) The amplitude of the pacemaker's daily oscillation declines as the duration of the entraining light pulse (photoperiod) is increased; and (2) the duration of the daily photoperiods throughout the breeding season is steadily increased as one moves toward the poles. Selection for conservation of pacemaker amplitude (during the breeding season) would produce the latitudinal cline we propose. The fourth, and final proposition is that since the amplitude of the pacemaker's daily motion responds systematically to change in photoperiod, amplitude is clearly one way--and a temperature-dependent way--in which insect circadian systems may sense seasonal change. These propositions concerning the temperature and latitude dependence of pacemaker amplitude may be relevant to a wider array of circadian pacemakers than Drosophila.

Animals↗

Evolutionary flux of P element regulation in a Drosophila melanogaster hybrid dysgenesis cline.

Clines of P-induced hybrid dysgenesis provide a means for monitoring the evolution of transposition repression over space and time. We have studied the molecular and phenotypic profiles of flies taken from a 2900 km cline along the eastern coast of Australia, which had previously been characterized over 10 years ago as having P populations in the north, Q populations at central sites and M' populations in the south. We have found that Q and M' populations of flies have increased their range within the cline at the expense of P lines. Q populations were found to be in the north of the cline and M' populations in the south. Some of the northern Q lines transmit repression through both sexes and type I deletion elements have been isolated from them. We suggest that these elements are responsible for Q type repression. The results support our model that populations made up of Q individuals with strong biparentally transmitted repression form an evolutionarily stable strategy for the repression of hybrid dysgenesis in Drosophila melanogaster.

Animals↗

Gene frequency clines at X-linked or haplodiploid loci.

The equations governing gene frequency clines at X-linked or haplodiploid loci are derived. Clines at X-linked loci will be identical to corresponding clines at autosomal loci only under three restrictive conditions (a) no dominance, (b) dispersal distance the same in each sex, (c) average effect of alleles same in each sex. If any of these conditions are not met then the males and females will have different gene frequencies. Clinal variation in colour morph frequency in the bumble bee Bombus melanopygusis analysed. It is concluded that a minimum selection intensity acting on the queens on the order of 1 per cent or less is sufficient to maintain the cline.

Animals↗