Evidence For Adaptive Differentiation Research Articles

On morphological and ecological evidence of adaptive differentiation among stony cliff littoral Baikal gastropods

Three morpho- or ecotypes (‘ribbed’, ‘middle’ or ‘ribless’ shell and ‘smooth’) of lithophilous Maackia herderiana (Lindholm, 1909) (Caenogastropoda: Baicaliidae) were investigated along the hydrodynamic zones of four locations on the southwestern littoral of Lake Baikal in southeast Siberia. We found a prevalence of the ribbed-shell morphotype inhabiting stones at a depth of 1.2–2.0 m, corresponding to the hydrodynamic zone of strong wave activity, and a predominance of the smooth-shell morphotype living on cliffy canyons at 21–31 m depth in the hydrodynamic zone where wave action is very weak or totally absent. Mixed groups of snails with ribbed, ribless and/or smooth shells were present on stony-boulder substrates at 3 to 15–19 m depth of the wave-cut and wave-weakening hydrodynamic zones. Ribbed-morphotype snails living in inshore and wave-cut hydrodynamic zones had a smaller shell size than did smooth-morphotype snails inhabiting zones where wave activity was weak or totally absent. The thickness of ribbed shells in the non-ribbed areas was equal to the thickness of smooth shells; ribs enhance the thickness of the shell by 3–4 times, thereby increasing the strength of ribbed shells against destructive wave action. On the basis of the different correlation coefficients of shell weight, soft body mass and total mass with increasing shell height, we assumed a growth differentiation between the ribbed and smooth morphotypes. We compared our results with data on ecotypes of some freshwater and marine gastropods.

Quantitative genetic variance and multivariate clines in the Ivyleaf morning glory, Ipomoea hederacea.

Clinal variation is commonly interpreted as evidence of adaptive differentiation, although clines can also be produced by stochastic forces. Understanding whether clines are adaptive therefore requires comparing clinal variation to background patterns of genetic differentiation at presumably neutral markers. Although this approach has frequently been applied to single traits at a time, we have comparatively fewer examples of how multiple correlated traits vary clinally. Here, we characterize multivariate clines in the Ivyleaf morning glory, examining how suites of traits vary with latitude, with the goal of testing for divergence in trait means that would indicate past evolutionary responses. We couple this with analysis of genetic variance in clinally varying traits in 20 populations to test whether past evolutionary responses have depleted genetic variance, or whether genetic variance declines approaching the range margin. We find evidence of clinal differentiation in five quantitative traits, with little evidence of isolation by distance at neutral loci that would suggest non-adaptive or stochastic mechanisms. Within and across populations, the traits that contribute most to population differentiation and clinal trends in the multivariate phenotype are genetically variable as well, suggesting that a lack of genetic variance will not cause absolute evolutionary constraints. Our data are broadly consistent theoretical predictions of polygenic clines in response to shallow environmental gradients. Ecologically, our results are consistent with past findings of natural selection on flowering phenology, presumably due to season-length variation across the range.

Evidence for adaptive phenotypic differentiation inBalticSea sticklebacks

The evidence for adaptive phenotypic differentiation in mobile marine species remains scarce, partly due to the difficulty of obtaining quantitative genetic data to demonstrate the genetic basis of the observed phenotypic differentiation. Using a combination of phenotypic and molecular genetic approaches, we elucidated the relative roles of natural selection and genetic drift in explaining lateral plate number differentiation in threespine sticklebacks (Gasterosteus aculeatus) across the entire Baltic Sea basin (approximately 392000km(2) ). We found that phenotypic differentiation (PST =0.213) in plate number exceeded that in neutral markers (FST =0.008), suggesting an adaptive basis for the observed differentiation. Because a close correspondence was found between plate phenotype and genotype at a quantitative trait loci (QTL; STN381) tightly linked to the gene (Ectodysplasin) underlying plate variation, the evidence for adaptive differentiation was confirmed by comparison of FST at the QTL (FSTQ =0.089) with FST at neutral marker loci. Hence, the results provide a comprehensive demonstration of adaptive phenotypic differentiation in a high-gene-flow marine environment with direct, rather than inferred, verification for the genetic basis of this differentiation. In general, the results illustrate the utility of PST -FST -FSTQ comparisons in uncovering footprints of natural selection and evolution and add to the growing evidence for adaptive genetic differentiation in high-gene-flow marine environments, including that of the relatively young Baltic Sea.

Sequence variation patterns along a latitudinal cline in Scots pine (Pinus sylvestris): signs of clinal adaptation?

Adaptive clinal variation is abundant in nature, and its genetic basis is of great interest. The polygenic nature of this variation poses a challenge for identifying the causative loci underlying these adaptations. Here, we have examined the patterns of sequence variation in ten candidate genes for timing of bud set and cold tolerance, traits that form strong latitudinal clines in Scots pine (Pinus sylvestris). A set of reference genes was used for comparison and to infer a simplified demographic background model with approximate Bayesian computation methods. Against the resulting bottleneck model, the neutrality tests show little signs for local adaptation, but species-wide selection is suggested in some of the studied genes. In line with the theoretical expectations, we see little evidence for adaptive differentiation with F ST methods. Instead, allele frequency clines were found in three genes (dhn1, ftl2, and prr1). Our results add to the discussion on which molecular signals best characterize a gene subject to clinally varying selection. This will be especially relevant when these kinds of observations can be examined in parallel with association study results.

Natural selection favors rapid reproductive phenology in Potentilla pulcherrima (Rosaceae) at opposite ends of a subalpine snowmelt gradient

In high altitude plants, flowering quickly ensures reproductive success within a short snow-free period, but limits maturation time and fecundity. Natural selection on prefloration intervals may therefore vary in contrasting snowmelt environments and could influence the outcome of phenological responses to climatic change. This study investigated adaptive differentiation and plasticity of prefloration intervals in the subalpine perennial Potentilla pulcherrima. Three years of in situ field observations were combined with phenotypic selection analyses and a common garden experiment. Plants from high, intermediate, and low altitudes expressed similar prefloration intervals and plasticity when grown at common altitude, indicating no evidence for adaptive differentiation. Selection on the prefloration interval was negative at both low and high altitudes before and after accounting for strong positive selection on size. Environmental differences between high and low altitudes indicated that long, dry seasons and short, wet seasons both favored rapid reproduction. Therefore, quicker reproduction was adaptive in response to late snowmelt, but slower reproduction in response to earlier snowmelt appeared to be maladaptive. Selection differed marginally between late snowmelt years and dry ones. Plastic responses to future precipitation patterns may therefore have positive or negative effects on fitness within a single species, depending upon altitude and year.

LOCAL ADAPTATION AND AGENTS OF SELECTION IN A MOBILE INSECT.

The deme-formation hypothesis states that selection can produce adaptive genetic variation within and among phytophagous insect populations. We conducted three field experiments and tested this prediction by transferring eggs and measuring performance of a mobile leafmining insect, Stilbosis quadricustatella. In Experiment 1, we compared the rate of mine initiation of leafminers transferred to natal and novel sites. In Experiment 2, we compared mine-initiation rate of leafminers transferred to natal and novel host-plant species. In Experiment 3, we compared the mine-initiation rate, mine-completion rate, and sources of mortality of miners transferred to neighboring natal and novel Quercus geminata trees. In the first, second, and third experiments, leafminer larvae initiated significantly more mines at the natal site, on the natal plant species, and on the natal Q. geminata tree, evidence for adaptive differentiation. Furthermore, plant-mediated mortality was significantly lower among miners transferred to natal Q. geminata trees. This result supports a key assumption of the deme-formation hypothesis: insects adapt to the defensive phenotypes of individual trees. However, natural-enemy mortality was significantly higher among miners transferred to natal trees, essentially reversing the plant effect. Therefore, rates of successful mine completion were similar on natal (19%) and novel (17%) trees. This experiment suggests that host plants and natural enemies may represent opposing forces of selection. Leafminers adapted to individual trees may realize a selective advantage only when natural-enemy densities are low.

Potential for the Evolution of Heavy Metal Tolerance in Bryum argenteum, a Moss. II. Generalized Tolerances among Diverse Populations

Tolerance of copper, zinc, lead, and nickel were measured in two individuals from each ofseven populations of Bryum argenteum. The populations represented a range of habitats including industrial sites subject to atmospheric metal deposition, metal-contaminated mine tailings, serpen- tine barrens, and urban areas. Nevertheless, there was no evidence of adaptive differentiation in tolerance to any of the metals. Populations did differ significantly in general growth vigor across all experimental treatments. These observations contrast with results from studies of angiosperms, in which the evolution of heavy-metal tolerance almost always involves ecotypic differentiation among populations, but fit an emerging picture of B. argenteum as a plastic, broadly tolerant species with surprisingly little genetic differentiation among populations.