The evolution of tolerance to damage in Gentianella campestris: natural selection and the quantitative genetics of tolerance

In the framework of phenotypic plasticity, tolerance to browsing can be operationally defined as a norm of reaction comparing plant performance in undamaged and damaged conditions. Genetic variation in tolerance is then indicated by heterogeneity in the slopes of norms of reaction from a population. We investigated field gentian (Gentianella campestris) tolerance to damage in the framework of phenotypic plasticity using a sample of maternal lines from natural populations grown under common garden conditions and randomly split into either a control or an artificial clipping treatment. We found a diversity of tolerance norms of reaction at both the population and family level: the impacts of clipping ranged from poor tolerance (negative slope) to overcompensation (positive slope). We detected heterogeneity in tolerance norms of reaction in four populations. Similarly, we found a variety of plastic architectural responses to clipping and genetic variation in these responses in several populations. Overall, we found that the most tolerant populations were late flowering and also exhibit the greatest plastic increases in node (meristem) production in response to damage. We studied damage-imposed natural selection on plasticity in plant architecture in 10 of the sampled populations. In general, there was strong positive direct selection on final number of nodes for both control and clipped plants. However, the total selection on nodes (direct + indirect selection) within each treatment category depended heavily on the frequency of damage and cross-treatment genetic correlations in node production. In some cases, strong correlated responses to selection across the damage treatment led to total selection against nodes in the more rare environment. This could ultimately lead to the evolution of maladaptive phenotypes in one or both of the treatment categories. These results suggest that tolerance and a variety of architectural responses to damage may evolve by both direct and indirect responses to natural selection. While the present study demonstrates the potential importance of cross-treatment genetic correlations in directing the evolution of tolerance traits, such as branch or node production, we did not find any strong evidence of genetic trade-offs in candidate tolerance traits between undamaged and damaged conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Phylogeography of sex ratio and multiple mating in stalk-eyed flies from southeast Asia

The factors maintaining sex chromosome meiotic drive, or sex ratio (SR), in natural populations remain uncertain. Coevolution between segregation distortion and modifiers should produce transient SR distortion while selection can result in a stable polymorphism. We hypothesize that if SR is maintained by selection, then phylogenetically related populations should exhibit similar SR frequency and intensity. Furthermore, when drive is present, females should mate with multiple males more often both to insure fertility and to increase the probability of producing male progeny. In this paper we report on variation in SR frequency and multiple mating among seven populations and three species of stalk-eyed flies, genus Cyrtodiopsis, from southeast Asia. Using a phylogenetic hypothesis based on 1100 bp of mtDNA sequence we find that while sex chromosome meiotic drive is present in all populations of C. whitei and C. dalmanni, the frequency and intensity of drive only differs between populations or species with greater than 4.8% sequence divergence. The frequency of females mating with multiple males is higher in populations with SR. In addition, SR males mate less often, possibly to compensate for sperm depletion. Our results suggest that sex chromosome drive is maintained by balancing selection in populations of C. whitei and C. dalmanni. Nevertheless, coevolution between drive and suppressors deserves further study.     

Sequential gel electrophoretic analysis of esterase-2 in two populations of Drosophila buzzatii

Sequential electrophoresis, using three different buffer systems on cellulose acetate gels, was used to characterize the allelic variation for esterase-2 in two populations of D. buzzatii in Australia that are separated by 550 km. Twenty-five alleles were detected, of which nine were unique to one population, eight unique to the other, and only eight were common to both populations. Allele frequencies within each population were significantly different between the two major chromosome sequences (standard and j inversion), and for each chromosome sequence allele frequencies were significantly different between populations. Observed allelic frequency distributions were not significantly different from those predicted for selective neutrality using the homozygosity test statistic. However, estimates of the effective sizes of the populations derived from their observed differentiation, together with the history of the species in Australia, provide support for some form of balancing selection affecting at least some of the alleles.     

Biosystematic studies onCapsella bursa-pastoris (Brassicaceae): Enzyme polymorphism in natural populations

The genetic variability of natural populations ofCapsella bursapastoris in North- and Middle-Europe has been estimated by means of enzyme assays. Zymograms of 81 populations have been developed. 17 loci could be identified, and 8 of them can be heterozygous. Genetic variability is greater between populations than within. No correlation between actual population sizes and genetic heterogeneity could be detected. Some electromorphs shift their frequencies proportionally to increasing adversity of climatic conditions, some appear to be constant over the whole area, and others are characterized by an apparently irregular variation pattern. Marginal populations comprise a significantly higher proportion of heterozygous plants than central ones. Apart from this clinal variation pattern, a mosaic pattern, strongly related to habitat conditions, was observed: genetic heterogeneity is greater in more intensively disturbed sites. The pattern of genetic variation in natural populations ofCapsella bursa-pastoris is rather highly influenced by the breeding system.     

An experimental analysis of the relationships between fitness components and malic enzyme genotypes inTribolium confusum

The hypothesis that natural selection is capable of maintaining allozyme variation in natural populations was tested using a species of flour beetles,Tribolium confusum. We selected a polymorphic locus (a locus encoding variation for malic enzyme) in an experimental population ofT. confusum and scored the genotypes at this locus for a series of fitness components on different flour types. Measurements included survival rate, development time, fecundity, and rate of egg cannibalism. Flour type had significant effects on most traits. Significant differences among genotypes for fecundity and rates of egg cannibalism and the presence of genotype × flour type interactions for development time were demonstrated. Thus, changes in allele frequencies at the malic enzyme locus could in part be under the influence of natural selection. The existence of genotype × flour type interactions suggests that environmental heterogeneity could maintain allozyme variation at the malic enzyme locus.     

Homogeneity of common cosmopolitan inversion frequencies in Southeast AsianDrosophila melanogaster

East AsianDrosophila melanogaster are known for great variation in morphological and physiological characters among populations, variation that is believed to be maintained by genetic drift. To understand the genetic properties of AsianD. melanogaster populations, we initiated a population genetic study of chromosome inversion polymorphisms in hitherto unanalysed population samples from Southeast (SE) Asia. We generally found a high frequency of each of the four common cosmopolitan inversions in comparison to populations from Africa, Asia, and Australia. In contrast to the great phenotypic variation among Asian populations, however, we could not detect differences in inversion frequencies among populations. Furthermore, we observed neither correlations of inversion frequencies with population latitude and longitude, nor evidence for linkage disequilibrium between different inversion loci. We propose two explanations for the observed genetic homogeneity among these SE AsianD. melanogaster populations: (i) the observed pattern simply reflects the retention of ancestral polymorphisms originating from a panmictic population that was once present on a large single landmass (Sundaland), and/or is a consequence of high recent gene flow between populations; and (ii) it is caused by selective forces (e.g. balancing selection).     

Genetic structure of natural populations of the red-spotted newt,Notophthalmus viridescens

Genetic variation is described at 15 loci in 2 neotenic and 12 nonneotenic populations of red-spotted newts. Though high levels of genetic similarity (I=0.990) were found among all populations, allele frequencies at six of the eight most polymorphic loci show significant heterogeneity across populations. Change in allele frequencies at two of these loci (Pep-2 and Ldh-1) is significantly correlated with latitude. Interspecific homologies are established for newt peptidases based on substrate specificities and lactate dehydrogenases based on tissue distribution, thermal stability, and kinetic properties. Nonneotenic populations are highly variable (H=0.157) and neotenic populations are only slightly, but significantly, less variable (H=0.120). The high levels of heterozygosity detected in nonneotenic populations may result from large effective population size and/or environmental heterogeneity. The unexpectedly high heterozygosity values obtained for the neotenic populations may indicate adult dispersal or the presence of some previously undetected red efts at these localities. In any case, a major change in life history has apparently had little effect on the genetic structure of these populations.This research was supported by grants from the Blakeslee Fund of Smith College.     

Evidence for ecological divergence across a mosaic of soil types in an Amazonian tropical tree: Protium subserratum (Burseraceae)

Soil heterogeneity is an important driver of divergent natural selection in plants. Neotropical forests have the highest tree diversity on earth, and frequently, soil specialist congeners are distributed parapatrically. While the role of edaphic heterogeneity in the origin and maintenance of tropical tree diversity is unknown, it has been posited that natural selection across the patchwork of soils in the Amazon rainforest is important in driving and maintaining tree diversity. We examined genetic and morphological differentiation among populations of the tropical tree Protium subserratum growing parapatrically on the mosaic of white‐sand, brown‐sand and clay soils found throughout western Amazonia. Nuclear microsatellites and leaf morphology were used to (i) quantify the extent of phenotypic and genetic divergence across habitat types, (ii) assess the importance of natural selection vs. drift in population divergence, (iii) determine the extent of hybridization and introgression across habitat types, (iv) estimate migration rates among populations. We found significant morphological variation correlated with soil type. Higher levels of genetic differentiation and lower migration rates were observed between adjacent populations found on different soil types than between geographically distant populations on the same soil type. P_ST‐F_ST comparisons indicate a role for natural selection in population divergence among soil types. A small number of hybrids were detected suggesting that gene flow among soil specialist populations may occur at low frequencies. Our results suggest that edaphic specialization has occurred multiple times in P. subserratum and that divergent natural selection across edaphic boundaries may be a general mechanism promoting and maintaining Amazonian tree diversity.     

Large mainland populations of South Island robins retain greater genetic diversity than offshore island refuges

For conservation purposes islands are considered safe refuges for many species, particularly in regions where introduced predators form a major threat to the native fauna, but island populations are also known to possess low levels of genetic diversity. The New Zealand archipelago provides an ideal system to compare genetic diversity of large mainland populations where introduced predators are common, to that of smaller offshore islands, which serve as predator-free refuges. We assessed microsatellite variation in South Island robins (Petroica australis australis), and compared large mainland, small mainland, natural island and translocated island populations. Large mainland populations exhibited more polymorphic loci and higher number of alleles than small mainland and natural island populations. Genetic variation did not differ between natural and translocated island populations, even though one of the translocated populations was established with five individuals. Hatching failure was recorded in a subset of the populations and found to be significantly higher in translocated populations than in a large mainland population. Significant population differentiation was largely based on heterogeneity in allele frequencies (including fixation of alleles), as few unique alleles were observed. This study shows that large mainland populations retain higher levels of genetic diversity than natural and translocated island populations. It highlights the importance of protecting these mainland populations and using them as a source for new translocations. In the future, these populations may become extremely valuable for species conservation if existing island populations become adversely affected by low levels of genetic variation and do not persist.     

Chromosome Differentiated Populations of Cruzii: Evidence for a Third Sibling Species

Anopheles cruziiis the most common species of mosquito in Southeast Brazil and a vector of human and monkey malaria. The banding pattern of the ovarian polytene chromosomes and the frequencies of paracentric inversions of individuals from two populations were studied. A new sequence of bands on the sex chromosome, defined as form C, was disclosed. In both populations where forms A (considered as standard) and C are sympatric no heterozygotes were detected. A sequence of events that could account for the observed changes in the banding sequences of the X chromosome forms was proposed. The frequencies of 22 paracentric inversions were used to assess panmixia and the results indicated the presence of two distinct genetic pools in each population. We consider these results as evidence of another sibling species in the taxon cruzii, characterized by a distinctive form of the X chromosome and provisionally designated Anopheles cruziispecies C.     

Allozyme genotype-environment relationships in natural populations of Drosophila buzzatii

Allozyme frequency data from natural populations of Drosophila buzzatii were analyzed for genotype-environment relationships. Allele frequency and heterozygosity at six loci polymorphic throughout eastern Australia and a number of environmental factors (both means and variabilities) were examined by a variety of multivariate techniques. Significant genotype-environment associations were found for five of the six loci, and after correcting for geographic location significant associations remained for Est-2 and Adh-1 gene frequencies and heterozygosities and for Pgm gene frequencies. The results are discussed in relation to selection and gene flow and provide the basis for laboratory studies to disentangle confounded effects of (1) environmental means and environmental variabilities and (2) allele frequency and heterozygosity, and thus to further test for and determine the nature of any natural selection at particular allozyme loci.This work was supported by a grant from the Australian Research Grants Committee to J. S. F. B.     

Latitudinal clines in<Emphasis Type="Italic">Drosophila melanogaster</Emphasis>: Body size,allozyme frequencies,inversion frequencies,and the insulin-signalling pathway

Many latitudinal clines exist inDrosophila melanogaster: in adult body size, in allele frequency at allozyme loci, and in frequencies of common cosmopolitan inversions. The question is raised whether these latitudinal clines are causally related. This review aims to connect data from two very different fields of study, evolutionary biology and cell biology, in explaining such natural genetic variation inD. melanogaster body size and development time. It is argued that adult body size clines, inversion frequency clines, and clines in allele frequency at loci involved in glycolysis and glycogen storage are part of the same adaptive strategy. Selection pressure is expected to differ at opposite ends of the clines. At high latitudes, selection onD. melanogaster would favour high larval growth rate at low temperatures, and resource storage in adults to survive winter. At low latitudes selection would favour lower larval critical size to survive crowding, and increased male activity leading to high male reproductive success. Studies of the insulin-signalling pathway inD. melanogaster point to the involvement of this pathway in metabolism and adult body size. The genes involved in the insulin-signalling pathway are associated with common cosmopolitan inversions that show latitudinal clines. Each chromosome region connected with a large common cosmopolitan inversion possesses a gene of the insulin transmembrane complex, a gene of the intermediate pathway and a gene of the TOR branch. The hypothesis is presented that temperateD. melanogaster populations have a higher frequency of a ’thrifty’ genotype corresponding to high insulin level or high signal level, while tropical populations possess a more ’spendthrift’ genotype corresponding to low insulin or low signal level.     

Genetic analysis of ecologically relevant morphological variability in Plantago lanceolata L.

Summary Morphological variability was studied in two populations of Plantago lanceolata using diallel analysis. In each population, reciprocal crosses between all possible pairs of ten plants were made. In the greenhouse, six members of each family were grown and many characters were measured. Using the model of Cockerham and Weir, the contributions of the different genetic variance components were calculated. From earlier papers it was postulated in advance to what extent and by which effect the characters in both populations were genetically determined. The populations had been differentiated for life history and morphological characters, and varied also in the relative contribution of genetic components to variability. In the Merrevliet (Me) population, where strong biotic selection was assumed, low levels of additive-genetic variability were present and the relative dominance appeared to be high. The contrasting population, Westduinen (Wd), which is abiotically controlled and shows strong environmental variability, possessed higher levels of additive-genetic variability and lower levels of relative dominance. It is possible that differential natural selection has diminished additive-genetic variability to different extents in both populations: plasticity and environmental heterogeneity prevented the loss of additive-genetic variability in Wd, whereas in the stable population, Me, natural selection had the opportunity of not only changing the means of the characters but also of diminishing additive-genetic variability to a great extent.Grassland Species Research Group Publication No. 146     

Geographical differentiation of allozymic variability in natural Indian populations ofDrosophila melanogaster

Fourteen Indian populations ofD. melanogaster collected along a 22° latitudinal range were analyzed electrophoretically to compare the geographical patterns of allozymic variation at 13 loci. The data show higher genetic differentiation on the basis of moderate to higherF _ST values and significant statistical correlation of allelic frequencies at six polymorphic loci with latitude. The results add support to the hypothesis that the occurrence of parallel or complementary latitudinal clines across different continental populations provides evidence of natural selection maintaining such clinal variation.     

Genetic variability in Muscari comosum L. (Liliaceae) IV Geographical distribution and adaptive role of the polymorphic variants of chromosome 2

Muscari comosum L. (Liliaceae) has a chromosomal polymorphism for a pericentric inversion and a supernumerary chromosome segment probably due to an unequal interchange or insertional translocation. Both arrangements are widely distributed throughout the species range and the mean genetic distance among populations is D=0.131±0.075. There are no correlations between genetic distance and geographic distance or latitude. Only appreciable decreases in the frequencies of the inversion are detected in populations with ecologically marginal characteristics. There is a permanent and extended association between chromosomal inversion and an enzymatic locus (ADH). An excess of individuals heterozygous for the inversion was found and female productivity of heterozygotes is higher than that of corresponding homozygotes. A low rate of inversion heterozygosity in populations with ecologically marginal characteristics could be explained by natural selection. With respect to the adaptive role of the segment, although no homozygotes are found and may be selected against, heterozygotes could have heterotic effects.     

Pollen pool heterogeneity in jack pine (Pinus banksiana Lamb.): a problem for estimating outcrossing rates?

Summary Pollen pool heterogeneity, which violates an assumption of the mixed-mating model, is a major potential problem in measuring plant mating systems. In this study, isozyme markers were used to examine pollen pool heterogeneity in two natural populations of jack pine, Pinus banksiana Lamb., in northwestern Ontario, Canada. Population multilocus estimates of outcrossing rate ranged from 0.83 to 0.95 and differed significantly between populations. Single-tree multilocus outcrossing rates were found to be homogeneous among trees in both populations. Computer simulation studies indicated that a consanguineous pollen pool (pollen gametes related to the mother tree) was capable of biasing population outcrossing estimates downward. Random pollen pool heterogeneity (uncorrelated with maternal genotypes) did not appear to affect population outcrossing estimates in the simulations. Heterogeneity G-tests and Spearman rank tests showed that pollen pool heterogeneity existed in the two natural populations examined; however, it did not have a major effect on population outcrossing estimates, since the consanguineous pollen pool detected was probably a relatively minor component of the outcross pollen pool in both populations. In addition, heterogeneity G-tests were found to be not sensitive in detecting pollen pool heterogeneity caused by consanguineous pollen pool.     

A simple approach for discovering common nonelectrophoretic enzyme variability: A heat denaturation study in Drosophila melanogaster

A simple procedure is described to detect genetic heterogeneity within electrophoretic classes at a locus in Drosophila, based on electrophoresis and heat denaturation studies. Temperature-resistant (tr) and temperature-sensitive (ts) isoelectrophoretic alleles at the phosphoglucomutase locus (Pgm) are present at polymorphic frequencies in natural and in laboratory populations of Drosophila melanogaster.This work has been supported by a grant from the Consiglio Nazionale delle Ricerche (C.N.R.).     

Non-random associations between Lap and Pept-1 loci and gene arrangements of chromosome O in D. subobscura — experiments in laboratory populations

In this paper the well-known non-random associations between Lap and Pept-1 loci and gene arrangements of chromosome O are studied in laboratory populations of D. subobscura. An increase of the frequency of the allele Lap ^1.00, towards an equilibrium point (0.70), was found to be associated with an increase of the gene arrangement O₃₊₄. This is an accordance to the associations found in natural populations. On the contrary no such an increase was observed in populations polymorphic for Lap and Pep-1 loci but homokaryotypic for gene arrangement O₃₊₄₊₈ differing in the initial allele frequencies at these loci. Although epistatic selection cannot be completely ruled out, our results are better explained under the assumption of neutrality.     

Pollinator visitation in populations of tristylous Eichhornia paniculata in northeastern Brazil

Summary The frequencies of floral morphs in populations of tristylous Eichhornia paniculata often deviate from the theoretical expectation of equality. This variation is associated with the breakdown of tristyly and the evolution of self-fertilization. Differences in morph frequencies could result from selection pressures due to variable levels of insect visitation to populations and contrasting foraging behavior among the floral morphs. We estimated pollinator densities in 16 populations and quantified visitation sequences to morphs in five populations of E. paniculata in northeastern Brazil. Foraging behavior among floral morphs was measured as the frequency of visits to morphs relative to their frequency in the population (preference) and number of flights between inflorescences of the same versus different morphs (constancy). Pollinator density (number/m²/minute) was not correlated with population size, plant density or morph diversity. Pollinator densities varied most among populations of less than 200 plants. Whether pollinators discriminated among the morphs, depended on whether they primarily collected nectar or pollen. In four populations, nectar-feeding bees (Ancyloscelis and Florilegus spp.) and butterflies showed no consistent preference or constancy among the morphs. In contrast, pollen-collecting bees (Trigona sp.) visited a lower proportion of longstyled inflorescences than expected and tended to visit more mid-and short-styled inflorescences in succession, once they were encountered. Pollinator constancy for morphs did not result from differences in inflorescence production or spatial patchiness among the morphs. Although non-random pollinator visitation to morphs in heterostylous populations could potentially affect mating and hence morph frequencies, the observed visitation patterns in this study do not provide evidence that pollinators play a major role in influencing floral morph frequencies.     

S-element insertions are associated with the evolution of the Hsp70 genes in Drosophila melanogaster

The "selfish DNA" theory postulates that transposable elements (TEs) are intragenomic parasites, and that natural selection against deleterious effects associated with their presence is the main force preventing their genomic spread in natural populations. In agreement with this model, TEs in Drosophila melanogaster populations are usually found at low frequencies in most genomic locations. Only a few cases of fixation of TE insertions have been reported, usually in regions of low recombination, where selection is expected to be less effective. Here, we report a population genetics study on the apparent fixation of an S-element in a highly recombining region in two natural populations of D. melanogaster. Three similar fragments of an S-element are inserted into the 5' regions of three members of a heat shock gene family, Hsp70 (Hsp70Aa and Hsp70Ab in polytene chromosome band 87A, and Hsp70Bb in 87C). A PCR-based analysis suggests that the insertions are fixed or at high frequencies in the entire species. A population survey of the levels of nucleotide sequence variation at the insertion site in 87C in two natural populations of D. melanogaster provided evidence for reduced levels of variation in the region, normal levels of recombination, and selection, reflected in a significant departure from neutrality of the variant frequency spectrum. This was particularly strong for the S-element inverted repeats (IRs) and suggests that these are of functional significance for the host.