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An independent non-linear latitudinal cline for the sn-glycerol-3-phosphate (alpha- Gpdh ) polymorphism of Drosophila melanogaster from eastern Australia.

Latitudinal variation of the polymorphic sn-glycerol-3-phosphate (alpha-Gpdh) locus in Drosophila melanogaster has been characterized on several continents; however, apparent clinal patterns are potentially confounded by linkage with an inversion, close associations with other genetic markers that vary clinally, and a tandem alpha-Gpdh pseudogene. Here we compare clinal patterns in alpha-Gpdh with those of other linked markers by testing field flies from eastern Australian locations collected in two separate years. The alpha-Gpdh variation exhibited a consistent non-linear cline reflecting an increase in the alpha-GpdhF allele at extreme latitudes. This pattern was not influenced by the In(2L)t inversion wherein this locus is located, nor was it influenced by the presence of the alpha-Gpdh pseudogene, whose presence was ubiquitous and highly variable among populations. The alpha-Gpdh pattern was also independent of a cline in allozyme frequencies at the alcohol dehydrogenase (Adh) locus, and two length polymorphisms in the Adh gene. These results suggest clinal selection at the alpha-Gpdh locus that is partially or wholly unrelated to linear climatic gradients along the eastern coast of Australia.

Alleles↗

Latitudinal clines for morphometrical traits in Drosophila kikkawai: a study of natural populations from the Indian subcontinent.

Natural populations of Drosophila kikkawai were collected in India and Sri Lanka, along a latitudinal transect ranging from 6.8 degrees to 31.8 degrees N latitude. Six morphometrical traits were analysed: wing and thorax length, body weight, ovariole number, and abdominal and sternopleural bristle numbers. Significant clines were observed for the three-size related traits and for ovariole number, corresponding to a regular increase in the mean value with latitude, but not for bristle numbers. Due to the utilization of two types of laboratory food, data were distributed into two separate data sets. A low-nutrient food produced smaller flies on average because of more intense crowding. The two rearing conditions produced significant clines but with significantly different slopes. The wing/thorax ratio, which is inversely related to wing loading, also increased with latitude. The analysis of Indian climatic conditions suggested that winter temperature, decreasing from south to north, could be more efficient than summer temperature, which varies in an opposite way, as a selective factor for inducing the clinal variations. The sibling species D. leontia, which is known only from the humid tropics, was found to be much smaller than D. kikkawai and did not fit the clinal regressions. Such morphological differences should help to identify the two species when found in sympatry.

Animals↗

Clines in the colour polymorphism of Philaenus spumarius in Eastern Central Europe.

Populations of Philaemus spumarius (Homoptera) have been investigated in the Baltic States of the USSR, in Czechoslovakia and in Hungary. Colour morph frequencies in these highly polymorphic populations are compared with those reported earlier from northern Italy. Gradual changes in allele frequencies (clines) transect the study area, which lies in the most central part of the range of Philaenus in Europe. From north to south the allele pM increases, but pO decreases, in frequency. In their distribution, some of the other alleles also showed regional gradients. The possibility that visual selection varies regionally is dismissed on indirect grounds. Gradients in climatic severity, or in other environmental variables changing concomitantly with climate, are regarded as the primary cause of the observed clines and other differences between regions.

Alleles↗

Mating propensity of Indian Drosophila melanogaster populations with D. simulans: a nonadaptive latitudinal cline.

Crosses were investigated between a single strain of Drosophila simulans and 28 natural Indian populations of D. melanogaster. In each case, a mass culture and 10 isofemale lines of D. melanogaster were studied. Crosses were much easier between D. melanogaster females and D. simulans males than in the reciprocal case. Also, hybrid offspring were easier to obtain by using D. melanogaster flies (either males or females) from isofemale lines than from mass cultures. Finally, the crossability, estimated by progeny production, showed a clear-cut latitudinal cline: hybrids were easier to obtain with D. melanogaster populations from higher latitudes. As D. simulans does not occur in India, this cline does not reflect an evolutionary interaction between the two sibling species.

Adaptation, Physiological↗

Molecules versus morphology: the detection of selection acting on morphological characters along a cline in Drosophila melanogaster.

This work examines the nature of north-south clinal variation in morphological characters in Drosophila melanogaster. Isofemale lines were established from flies collected along a transect extending from Winnipeg, Manitoba (Canada) to Tampa Bay, Florida (U.S.A.). Offspring from different lines within each location were then cultured under standardized conditions and used to examine phenotypic variation in seven morphological characters along the cline. In addition, allozyme variation at seven polymorphic loci was examined for the same set of clinal populations. Scutellum length and wing length show the strongest clinal trends. Clinal variation is nonmonotonic, with larger flies in the middle latitudes and smaller flies in the north and south. This result contrasts with other studies which have shown monotonic clines. Patterns of population subdivision were different for the different characters. This implies that there are different selective forces acting on the different morphological characters. Based on a comparison of morphological and molecular population subdivision for adjacent populations it is inferred that natural selection is operating to maintain a high level of population subdivision for wing width and the first principal component between one of the sets of populations. A combined approach using molecules and morphology may provide an alternative to retrospective selection analysis for detecting selection in nature.

Animals↗

A morphological cline in Eucalyptus: a genetic perspective.

The putative hybrid zone between Eucalyptus populnea and E. brownii is examined using morphological and molecular techniques. This species complex displays continuous morphological variation across the study area, which has been previously interpreted as the product of hybridization between allopatric species. A microsatellite analysis indicates that there was little genetic structuring across the morphological cline and only low levels of population differentiation. The nested clade analysis of the JLA+ region of the chloroplast DNA (cpDNA) indicates that the geographical distribution of cpDNA haplotypes is unlikely to be the result of historical hybridization events, and that restricted seed-mediated gene flow with isolation by distance is responsible for the phylogeographical distribution. A more plausible explanation for the origin and persistence of the morphological cline is that the process of continuous morphological diversification has been promoted by a directional selection gradient. This study addresses species status within Eucalyptus and the belief that hybridization is widespread and is an important process in the group's evolution.

Analysis of Variance↗

Epistatic interaction between Arabidopsis FRI and FLC flowering time genes generates a latitudinal cline in a life history trait.

Epistatic gene interactions are believed to be a major factor in the genetic architecture of evolutionary diversification. In Arabidopsis thaliana, the FRI and FLC genes mechanistically interact to control flowering time, and here we show that this epistatic interaction also contributes to a latitudinal cline in this life history trait within the species. Two major FLC haplogroups (FLC(A) and FLC(B)) are associated with flowering time variation in A. thaliana in field conditions, but only in the presence of putatively functional FRI alleles. Significant differences in latitudinal distribution of FLC haplogroups in Eurasia and North Africa also depend on the FRI genotype. There is significant linkage disequilibrium between FRI and FLC despite their location on separate chromosomes. Although no nonsynonymous polymorphisms differentiate FLC(A) and FLC(B), vernalization induces the expression of an alternatively spliced FLC transcript that encodes a variant protein with a radical amino acid replacement associated with the two FLC haplogroups. Numerous polymorphisms differentiating the FLC haplogroups also occur in intronic regions implicated in the regulation of FLC expression. The features of the regulatory gene interaction between FRI and FLC in contributing to the latitudinal cline in A. thaliana flowering time are consistent with the maintenance of this interaction by epistatic selection. These results suggest that developmental genetic pathways and networks provide the molecular basis for epistasis, contributing to ecologically important phenotypic variation in natural populations and to the process of evolutionary diversification.

Alternative Splicing↗

Random genetic drift in a cline.

A diffusion model is derived for random genetic drift in a cline. The monoecious organism occupies an unbounded linear habitat with constant, uniform population density. Migration is homogeneous, symmetric, and independent of genotype. The analysis applies to a single diallelic locus without dominance or mutation. Assuming that Hardy-Weinberg proportions obtain locally, partial differential equations are deduced for the expected gene frequency at each location and the covariance between gene frequencies at any two points of the habitat. For most selection gradients, the population density, rho, selection intensity, s, and migration variance, sigma(2), appear only combined in the single dimensionless parameter, beta, the ratio of the characteristic length for migration and random drift to the natural distance for migration and selection. Random drift is highly significant if beta << 1; it causes only relatively small variations of order 1/beta around the deterministic cline if beta >> 1. With beta >> 1, the correlation between the gene frequencies at any two points is very nearly independent of beta, and hence is parameter-free for particular forms of the selection gradient. For a very steep selection gradient, beta = 2 [unk] 2srhosigma, i.e., essentially the product of the square root of the selection intensity and the neighborhood size.

Journal Article↗

Temperature-related genetic changes in laboratory populations of Drosophila subobscura: evidence against simple climatic-based explanations for latitudinal clines.

Parallel latitudinal clines to the long-standing ones in the original Palearctic populations have independently evolved at different rates for chromosomal polymorphism and body size in South and North American populations of Drosophila subobscura since colonization around 25 years ago. This strongly suggests that (micro) evolutionary changes are largely predictable, but the underlying mechanisms are unknown. The putative role of temperature per se was investigated by using three sets of populations at each of three temperatures (13 degrees , 18 degrees , and 22 degrees C) spanning much of the tolerable range for this species. We found a lower chromosomal diversity at the warmest temperature; a quick and consistent shift in gene arrangement frequencies in response to temperature; an evolutionary decrease in wing size, mediated by both cell area and cell number, at 18 degrees C; no relationship between wing size and those inversions involved in latitudinal clines; and a shortening of the basal length of longitudinal vein IV relative to its total length with increasing standard dose. The trends for chromosomal polymorphism and body size were generally inconsistent from simple climatic-based explanations of worldwide latitudinal patterns. The findings are discussed in the light of available information on D. subobscura and results from earlier thermal selection experiments with various Drosophila species.

Animals↗

Genetic and environmental responses to temperature of Drosophila melanogaster from a latitudinal cline.

Field-collected Drosophila melanogaster from 19 populations in Eastern Australia were measured for body size traits, and the measurements were compared with similar ones on flies from the same populations reared under standard laboratory conditions. Wild caught flies were smaller, and latitudinal trends in size were greater. Reduced size was caused by fewer cells in the wing, and the steeper cline by greater variation in cell area. The reduction in size in field-collected flies may therefore have been caused by reduced nutrition, and the steeper cline may have been caused by an environmental response to latitudinal variation in temperature. No evidence was found for evolution of size traits in response to laboratory culture. The magnitude of phenotypic plasticity in response to temperature of development time, body size, cell size and cell number was examined for six of the populations, to test for latitudinal variation in plasticity. All characters were plastic in response to temperature. Total development time showed no significant latitudinal variation in plasticity, although larval development time showed a marginally significant effect, with most latitudinal variation at intermediate rearing temperatures. Neither thorax length nor wing size and its cellular components showed significant latitudinal variation in plasticity.

Adaptation, Physiological↗

Testing for asymmetrical gene flow in a Drosophila melanogaster body-size cline.

Asymmetrical gene flow is an important, but rarely examined genetic parameter. Here, we develop a new method for detecting departures from symmetrical migration between two populations using microsatellite data that are based on the difference in the proportion of private alleles. Application of this approach to data collected from wild-caught Drosophila melanogaster along a latitudinal body-size cline in eastern Australia revealed that asymmetrical gene flow could be detected, but was uncommon, nonlocalized, and occurred in both directions. We also show that, in contrast to the findings of a previous study, there is good evidence to suggest that the cline experiences significant levels of gene flow between populations.

Animals↗

Genetic drift in a cline.

A model is developed of genetic drift in a cline maintained by spatially varying natural selection and local dispersal of individuals. The model is analyzed by an approximation scheme which is valid for weak selection and small migration rates. The results, which are based on numerical iterations of the approximate equations, are that the cline is less steep than predicted on the basis of the deterministic theory but that for weak selection the correlation between random fluctuations in neighboring colonies is approximately the same as in models of migration and drift in the absence of selection.

Biological Evolution↗

Further evidence for latitudinal inversion clines in natural populations of Drosophila melanogaster from India.

Chromosomal analysis of eight Indian natural populations (six from north India and two from south India) of Drosophila melanogaster revealed the presence of 13 paracentric inversions, including one in the X chromosome. All four types of inversions--common cosmopolitan (4), rare cosmopolitan (3), recurrent endemic (2), and unique endemic (4)--were detected. The frequency of commonly occurring inversions and the level of inversion heterozygosity were found to be higher in the two southern populations. The south Indian populations are genetically more differentiated than those from the north. Also, latitudinal clines in the frequencies of the four common cosmopolitan inversions were detected. These results provide further evidence for the existence of inversion clines in Indian populations of D. melanogaster.

Animals↗

Evolution of additive and nonadditive genetic variance in development time along a cline in Drosophila serrata.

Latitudinal clines provide natural systems that may allow the effect of natural selection on the genetic variance to be determined. Ten clinal populations of Drosophila serrata collected from the eastern coast of Australia were used to examine clinal patterns in the trait mean and genetic variance of the life-history trait egg-to-adult development time. Development time significantly lengthened from tropical areas to temperate areas. The additive genetic variance for development time in each population was not associated with latitude but was associated with the population mean development time. Additive genetic variance tended to be larger in populations with more extreme development times and appeared to be consistent with allele frequency change. In contrast, the nonadditive genetic variance was not associated with the population mean but was associated with latitude. Levels of nonadditive genetic variance were greatest in the region of the cline where the gradient in the change in mean was greatest, consistent with Barton's (1999) conjecture that the generation of linkage disequilibrium may become an important component of the genetic variance in systems with a spatially varying optimum.

Analysis of Variance↗

Contrasting patterns of floral and molecular variation across a cline in Mimulus aurantiacus.

Steep clines in ecologically important traits may be caused by divergent natural selection. However, processes that do not necessarily invoke ongoing selection, such as secondary contact or restricted gene flow, can also cause patterns of phenotypic differentiation over short spatial scales. Distinguishing among all possible scenarios is difficult, but an attainable goal is to establish whether scenarios that imply selection need to be invoked. We compared the extent of morphological and genetic differentiation between geographically structured red and yellow floral races of Mimulus aurantiacus (bush monkeyflower; Phrymaceae). Flower color was assessed in a common garden as well as in the field to determine whether variation was genetic and to quantify the extent of geographical differentiation. Population genetic differentiation at marker loci was measured for both chloroplast and nuclear genomes, and the degree of population structure within and among the floral races was evaluated. Flower color shows both a strong genetic basis and a sharp geographic transition, with pure red-flowered populations in western San Diego County and pure yellow-flowered populations to the east. In the zone of contact, both pure and intermediate phenotypes occur. Patterns of genetic differentiation at marker loci are far less pronounced, as little of the variation is partitioned according to the differences in flower color. Phenotypic differentiation (Q(ST)) between populations with different flower colors is much greater than neutral genetic differentiation (F(ST)). When comparisons are made between populations of the same flower color, the opposite trend is evident. Limited neutral genetic structure between the floral races, combined with sharp differentiation in flower color, is consistent with the hypothesis that current or recent natural selection maintains the cline in flower color.

Biological Evolution↗

Latitudinal clines in the properties of a circadian pacemaker.

The circadian rhythm of eclosion activity and its pacemaker were analyzed in a series of latitudinal races of Drosophila auraria ranging from 34.2 degrees to 42.9 degrees N in Japan. The phase of the rhythm (psi EL) to the daily photoperiod (PP) changes as daylength is increased, and the slope of psi EL (PP) changes with latitude. Is is sufficiently greater in the north to cause a phase reversal of northern and southern races on long versus short photoperiods. This reversal is found in assays of the pacemaker's phase (psi PL) as well as that of the rhythm (psi EL). Assay of the pacemaker shows that its period (tau) is longer in northern than in southern races, and that the amplitude of its phase response curve (PRC) is lower in the north. The period of the rhythm in all latitudinal races is longer than 24 hr in short photoperiods (LD 1:23), but is probably less than 24 hr (as an aftereffect of photoperiod) in longer days such as LD 14:10. The observed north-south differences in the phase relation of both pacemaker and rhythm to the light cycle are explained by the latitudinal clines in pacemaker properties and a postulated aftereffect of photoperiod on tau. It is suggested that the latitudinal cline in PRC amplitude has functional significance in conserving the amplitude of the pacemaker's signal to the rest of the system it times. Computer simulation shows that without such a reduction in the perceived light intensity, pacemaker amplitude will be lowered by the increase in duration of the daily light at higher latitudes.

Animals↗

Analysis of Clines with Variable Selection and Variable Migration.

We report a likelihood-based method that estimates both dispersal and natural selection using the rate of change of the shape of a cline when selection and migration are not constant through time. We have investigated the case of local adaptation of the mosquito Culex pipiens to organophosphate insecticides in the Montpellier area in France. We have analyzed the modification of the clinal patterns at two resistance loci during the period from breeding to overwintering. We show that mosquitoes migrate extensively from breeding to overwintering sites at a rate that is markedly different from previous estimates made during the breeding season only. This migration is also strongly asymmetrical, which can be explained by different geographical distributions of breeding and overwintering sites, by variation in mosquito density along the transect, or by behavioral biases. We found that the starting time of overwintering is likely to vary between northern and southern populations and that substantial fitness costs are associated with resistance alleles at the two loci during overwintering. These results illustrate how demography and adaptive microevolution can be studied using selected markers. The method provides a framework to use population genetics and statistical models to reveal ecological and evolutionary processes.

asymmetrical gene flow↗

An arbovirus cline across the northern hemisphere.

The mode and tempo of arbovirus evolution and dispersal can help to explain the dynamics of pandemics, viral outbreaks, and emerging viruses. By comparing nucleotide and deduced amino acid sequences of their envelope proteins, we describe the continuous distribution of the tick-borne encephalitis (TBE) complex viruses, the most important flaviviruses in Europe, across major geographical areas and the conditions under which mutations occur. The analyses reveal a correlation between the geographical and genetic distances of these viruses. The arthropod host appears to be a key factor for the formation and maintenance of this cline by constraining TBE dispersal and evolution. This is also illustrated by comparisons with mosquito-borne flaviviruses.

Animals↗