Analysis of ancestry heterozygosity suggests that hybrid incompatibilities in threespine stickleback are environment dependent.

Hybrid incompatibilities occur when interactions between opposite ancestry alleles at different loci reduce the fitness of hybrids. Most work on incompatibilities has focused on those that are "intrinsic," meaning they affect viability and sterility in the laboratory. Theory predicts that ecological selection can also underlie hybrid incompatibilities, but tests of this hypothesis using sequence data are scarce. In this article, we compiled genetic data for F2 hybrid crosses between divergent populations of threespine stickleback fish (Gasterosteus aculeatus L.) that were born and raised in either the field (seminatural experimental ponds) or the laboratory (aquaria). Because selection against incompatibilities results in elevated ancestry heterozygosity, we tested the prediction that ancestry heterozygosity will be higher in pond-raised fish compared to those raised in aquaria. We found that ancestry heterozygosity was elevated by approximately 3% in crosses raised in ponds compared to those raised in aquaria. Additional analyses support a phenotypic basis for incompatibility and suggest that environment-specific single-locus heterozygote advantage is not the cause of selection on ancestry heterozygosity. Our study provides evidence that, in stickleback, a coarse-albeit indirect-signal of environment-dependent hybrid incompatibility is reliably detectable and suggests that extrinsic incompatibilities can evolve before intrinsic incompatibilities.

Heterosis counteracts hybrid breakdown to forestall speciation by parallel natural selection.

In contrast to ecological speciation, where reproductive isolation evolves as a consequence of divergent natural selection, speciation by parallel natural selection has been less thoroughly studied. To test whether parallel evolution drives speciation, we leveraged the repeated evolution of benthic and limnetic ecotypes of threespine stickleback fish and estimated fitness for pure crosses and within-ecotype hybrids in semi-natural ponds and in laboratory aquaria. In ponds, we detected hybrid breakdown in both ecotypes but this was counterbalanced by heterosis and the strength of post-zygotic isolation was nil. In aquaria, we detected heterosis in limnetic crosses and breakdown in benthic crosses, which is suggestive of process- and ecotype-specific environment-dependence. In ponds, heterosis and breakdown were three times greater in limnetic crosses than in benthic crosses, contrasting the prediction that the fitness consequences of hybridization should be greater in crosses among more derived ecotypes. Consistent with a primary role for stochastic processes, patterns differed among crosses between populations from different lakes. Yet, the observation of qualitatively similar patterns of heterosis and hybrid breakdown for both ecotypes when averaging the lake pairs indicates that the outcome of hybridization is repeatable in a general sense.

Patterns, Predictors, and Consequences of Dominance in Hybrids.

AbstractCompared to those of their parents, are the traits of first-generation (F1) hybrids typically intermediate, biased toward one parent, or mismatched for alternative parental phenotypes? To address this empirical gap, we compiled data from 233 crosses in which traits were measured in a common environment for two parent taxa and their F1 hybrids. We find that individual traits in F1s are halfway between the parental midpoint and one parental value. Considering pairs of traits together, a hybrid's bivariate phenotype tends to resemble one parent (parent bias) about 50% more than the other, while also exhibiting a similar magnitude of mismatch due to different traits having dominance in conflicting directions. Using data from an experimental field planting of recombinant hybrid sunflowers, we illustrate that parent bias improves fitness, whereas mismatch reduces fitness. Our study has three major conclusions. First, hybrids are not phenotypically intermediate but rather exhibit substantial mismatch. Second, dominance is likely determined by the idiosyncratic evolutionary trajectories of individual traits and populations. Finally, selection against hybrids likely results from selection against both intermediate and mismatched phenotypes.

Experimental Hybridization Studies Suggest That Pleiotropic Alleles Commonly Underlie Adaptive Divergence between Natural Populations.

The alleles used for adaptation can pleiotropically affect traits under stabilizing selection. The fixation of alleles with deleterious pleiotropic side effects causes compensatory alleles to be favored by selection. Such compensatory alleles might segregate in interpopulation hybrids, resulting in segregation variance for traits where parents have indistinguishable phenotypes. If adaptation typically involves pleiotropy and compensation, then the segregation variance for traits under stabilizing selection is expected to increase with the magnitude of adaptive phenotypic divergence between parents. This prediction has not been tested empirically, and I gathered data from experimental hybridization studies to evaluate it. I found that pairs of parents that are more phenotypically divergent beget hybrids with more segregation variance in traits for which the parents are statistically indistinguishable. This result suggests that adaptive divergence between pairs of natural populations proceeds via pleiotropy and compensation and that deleterious transgressive segregation variance accumulates systematically as populations diverge.

The morphogenesis of fast growth in plants.

Growth rate represents a fundamental axis of life history variation. Faster growth associated with C4 photosynthesis and annual life history has evolved multiple times, and the resulting diversity in growth is typically explained via resource acquisition and allocation. However, the underlying changes in morphogenesis remain unknown. We conducted a phylogenetic comparative experiment with 74 grass species, conceptualising morphogenesis as the branching and growth of repeating modules. We aimed to establish whether faster growth in C4 and annual grasses, compared with C3 and perennial grasses, came from the faster growth of individual modules or higher rates of module initiation. Morphogenesis produces fast growth in different ways in grasses using C4 and C3 photosynthesis, and in annual compared with perennial species. C4 grasses grow faster than C3 species through a greater enlargement of shoot modules and quicker secondary branching of roots. However, leaf initiation is slower and there is no change in shoot branching. Conversely, faster growth in annuals than perennials is achieved through greater branching and enlargement of shoots, and possibly faster root branching. The morphogenesis of fast growth depends on ecological context, with C4 grasses tending to promote resource capture under competition, and annuals enhancing branching to increase reproductive potential.

Multivariate phenotypic divergence along an urbanization gradient.

Evidence suggests that natural populations can evolve to better tolerate the novel environmental conditions associated with urban areas. Studies of adaptive divergence in urban areas often examine one or a few traits at a time from populations residing only at the most extreme urban and nonurban habitats. Thus, whether urbanization drives divergence in many traits simultaneously in a manner that varies with the degree of urbanization remains unclear. To address this gap, we generated seed families of white clover (Trifolium repens) collected from 27 populations along an urbanization gradient in Toronto, Canada, grew them in a common garden, and measured 14 phenotypic traits. Families from urban sites had evolved later phenology and germination, larger flowers, thinner stolons, reduced cyanogenesis, greater biomass and greater seed set. Pollinator observations revealed near-complete turnover of pollinator morphological groups along the urbanization gradient, which may explain some of the observed divergences in floral traits and phenology. Our results suggest that adaptation to urban environments involves multiple traits.

Bellevue Smart:: Development and Integration of a Smart City.

Key Takeaways In Washington, the City of Bellevue wanted to break away from the inefficiency of siloed operations and create a unified, data-based, smart-city strategy across all city departments. The Bellevue Smart Team tested a water module in its new City Portal, starting with a dashboard that would organize and optimize data for a better customer, utility, and city experience. The platform used to set up the City Portal allows for flexibility in adding modules and adopting new technologies.

Herbivores and plant defences affect selection on plant reproductive traits more strongly than pollinators.

Pollinators and herbivores can both affect the evolutionary diversification of plant reproductive traits. However, plant defences frequently alter antagonistic and mutualistic interactions, and therefore, variation in plant defences may alter patterns of herbivore- and pollinator-mediated selection on plant traits. We tested this hypothesis by conducting a common garden field experiment using 50 clonal genotypes of white clover (Trifolium repens) that varied in a Mendelian-inherited chemical antiherbivore defence-the production of hydrogen cyanide (HCN). To evaluate whether plant defences alter herbivore- and/or pollinator-mediated selection, we factorially crossed chemical defence (25 cyanogenic and 25 acyanogenic genotypes), herbivore damage (herbivore suppression) and pollination (hand pollination). We found that herbivores weakened selection for increased inflorescence production, suggesting that large displays are costly in the presence of herbivores. In addition, herbivores weakened selection on flower size but only among acyanogenic plants, suggesting that plant defences reduce the strength of herbivore-mediated selection. Pollinators did not independently affect selection on any trait, although pollinators weakened selection for later flowering among cyanogenic plants. Overall, cyanogenic plant defences consistently increased the strength of positive directional selection on reproductive traits. Herbivores and pollinators both strengthened and weakened the strength of selection on reproductive traits, although herbivores imposed ~2.7× stronger selection than pollinators across all traits. Contrary to the view that pollinators are the most important agents of selection on reproductive traits, our data show that selection on reproductive traits is driven primarily by variation in herbivory and plant defences in this system.

Urbanization drives the evolution of parallel clines in plant populations.

Urban ecosystems are an increasingly dominant feature of terrestrial landscapes. While evidence that species can adapt to urban environments is accumulating, the mechanisms through which urbanization imposes natural selection on populations are poorly understood. The identification of adaptive phenotypic changes (i.e. clines) along urbanization gradients would facilitate our understanding of the selective factors driving adaptation in cities. Here, we test for phenotypic clines in urban ecosystems by sampling the frequency of a Mendelian-inherited trait-cyanogenesis-in white clover (Trifolium repens L.) populations along urbanization gradients in four cities. Cyanogenesis protects plants from herbivores, but reduces tolerance to freezing temperatures. We found that the frequency of cyanogenic plants within populations decreased towards the urban centre in three of four cities. A field experiment indicated that spatial variation in herbivory is unlikely to explain these clines. Rather, colder minimum winter ground temperatures in urban areas compared with non-urban areas, caused by reduced snow cover in cities, may select against cyanogenesis. In the city with no cline, high snow cover might protect plants from freezing damage in the city centre. Our study suggests that populations are adapting to urbanization gradients, but regional climatic patterns may ultimately determine whether adaptation occurs.

Climatic niche differences between diploid and tetraploid cytotypes of Chamerion angustifolium (Onagraceae).

PREMISE OF THE STUDY: Polyploidy-the possession of more than two copies of each chromosome in the nucleus-is common in flowering plants. Polyploid plants can occupy different geographic ranges than their diploid progenitors, but the factors responsible for maintaining these range differences are poorly understood. Polyploidy can have significant physiological consequences, and the present study aims to determine whether previously described physiological differences between cytotypes are correlated with climatic niches and geographic distributions. METHODS: Prior research indicates that tetraploid plants of Chamerion angustifolium (Onagraceae) are more tolerant of drought and less tolerant of freezing than diploids, which suggests that they should occupy a niche that is warmer and drier than that of diploids. We extracted climate data for 134 populations of C. angustifolium classified as pure diploid, pure tetraploid, or mixed-ploidy. We compared climatic conditions between these population categories and generated ecological niche models to compare their geographic distribution with prior qualitative estimates. KEY RESULTS: Pure tetraploid populations occupy habitats that are warmer and drier than those of pure diploid populations. Mixed-ploidy populations occur in habitats that are not strictly intermediate between pure diploid and pure tetraploid populations, but are as cold as pure diploid populations and have intermediate soil moisture deficits. Our niche models were similar to previous qualitative estimates of cytotype geographic distribution. CONCLUSIONS: The correspondence between the physiological tolerances of cytotypes, their climatic niches, and their geographic distributions suggests that physiological traits are at least partially responsible for differences in the realized climatic niches of diploid and tetraploid C. angustifolium.

Does seed mass drive the differences in relative growth rate between growth forms?

The idea that herbaceous plants have higher relative growth rates (RGRs) compared with woody plants is fundamental to many of the most influential theories in plant ecology. This difference in growth rate is thought to reflect systematic variation in physiology, allocation and leaf construction. Previous studies documenting this effect have, however, ignored differences in seed mass. As woody species often have larger seeds and RGR is negatively correlated with seed mass, it is entirely possible the lower RGRs observed in woody species is a consequence of having larger seeds rather than different growth strategies. Using a synthesis of the published literature, we explored the relationship between RGR and growth form, accounting for the effects of seed mass and study-specific effects (e.g. duration of study and pot volume), using a mixed-effects model. The model showed that herbaceous species do indeed have higher RGRs than woody species, and that the difference was independent of seed mass, thus at all seed masses, herbaceous species on average grow faster than woody ones.

Physical dormancy in seeds: a game of hide and seek?

Historically, 'physical dormancy', or 'hard seededness', where seeds are prevented from germinating by a water-impermeable seed coat, is viewed as a dormancy mechanism. However, upon water uptake, resumption of metabolism leads to the unavoidable release of volatile by-products, olfactory cues that are perceived by seed predators. Here, we examine the hypothesis that hard seeds are an anti-predator trait that evolved in response to powerful selection by small mammal seed predators. Seeds of two legume species with dimorphic seeds ('hard' and 'soft'), Robinia pseudoacacia and Vicia sativa, were offered to desert hamsters (Phodopus roborovskii) in a series of seed removal studies examining the differences in seed harvest between hard and soft seeds. Volatile compounds emitted by dry and imbibed soft seeds were identified by headspace gas chromatography-mass spectrometry (GC-MS). Fourteen main volatile compounds were identified, and hamsters readily detected both buried imbibed seeds and an artificial 'volatile cocktail' that mimicked the scent of imbibed seeds, but could not detect buried hard or dry soft seeds. We argue that physical dormancy has evolved to hide seeds from mammalian predators. This hypothesis also helps to explain some otherwise puzzling features of hard seeds and has implications for seed dispersal.

The Ecology of Hybrid Incompatibilities.

Ecologically mediated selection against hybrids, caused by hybrid phenotypes fitting poorly into available niches, is typically viewed as distinct from selection caused by epistatic Dobzhansky-Muller hybrid incompatibilities. Here, we show how selection against transgressive phenotypes in hybrids manifests as incompatibility. After outlining our logic, we summarize current approaches for studying ecology-based selection on hybrids. We then quantitatively review QTL-mapping studies and find traits differing between parent taxa are typically polygenic. Next, we describe how verbal models of selection on hybrids translate to phenotypic and genetic fitness landscapes, highlighting emerging approaches for detecting polygenic incompatibilities. Finally, in a synthesis of published data, we report that trait transgression-and thus possibly extrinsic hybrid incompatibility in hybrids-escalates with the phenotypic divergence between parents. We discuss conceptual implications and conclude that studying the ecological basis of hybrid incompatibility will facilitate new discoveries about mechanisms of speciation.

Adaptive divergence and the evolution of hybrid trait mismatch in threespine stickleback.

Selection against mismatched traits in hybrids is the phenotypic analogue of intrinsic hybrid incompatibilities. Mismatch occurs when hybrids resemble one parent population for some phenotypic traits and the other parent population for other traits, and is caused by dominance in opposing directions or from segregation of alleles in recombinant hybrids. In this study, we used threespine stickleback fish (Gasterosteus aculeatus L.) to test the theoretical prediction that trait mismatch in hybrids should increase with the magnitude of phenotypic divergence between parent populations. We measured morphological traits in parents and hybrids in crosses between a marine population representing the ancestral form and twelve freshwater populations that have diverged from this ancestral state to varying degrees according to their environments. We found that trait mismatch was greater in more divergent crosses for both F1 and F2 hybrids. In the F1, the divergence-mismatch relationship was caused by traits having dominance in different directions, whereas it was caused by increasing segregating phenotypic variation in the F2. Our results imply that extrinsic hybrid incompatibilities accumulate as phenotypic divergence proceeds.

The diverse effects of phenotypic dominance on hybrid fitness.

When divergent populations interbreed, their alleles are brought together in hybrids. In the initial F1 cross, most divergent loci are heterozygous. Therefore, F1 fitness can be influenced by dominance effects that could not have been selected to function well together. We present a systematic study of these F1 dominance effects by introducing variable phenotypic dominance into Fisher's geometric model. We show that dominance often reduces hybrid fitness, which can generate optimal outbreeding followed by a steady decline in F1 fitness, as is often observed. We also show that "lucky" beneficial effects sometimes arise by chance, which might be important when hybrids can access novel environments. We then show that dominance can lead to violations of Haldane's Rule (reduced fitness of the heterogametic F1) but strengthens Darwin's Corollary (F1 fitness differences between cross directions). Taken together, results show that the effects of dominance on hybrid fitness can be surprisingly difficult to isolate, because they often resemble the effects of uniparental inheritance or expression. Nevertheless, we identify a pattern of environment-dependent heterosis that only dominance can explain, and for which there is some suggestive evidence. Our results also show how existing data set upper bounds on the size of dominance effects. These bounds could explain why additive models often provide good predictions for later-generation recombinant hybrids, even when dominance qualitatively changes outcomes for the F1.

Long-term resistance to simulated climate change in an infertile grassland.

Climate shifts over this century are widely expected to alter the structure and functioning of temperate plant communities. However, long-term climate experiments in natural vegetation are rare and largely confined to systems with the capacity for rapid compositional change. In unproductive, grazed grassland at Buxton in northern England (U.K.), one of the longest running experimental manipulations of temperature and rainfall reveals vegetation highly resistant to climate shifts maintained over 13 yr. Here we document this resistance in the form of: (i) constancy in the relative abundance of growth forms and maintained dominance by long-lived, slow-growing grasses, sedges, and small forbs; (ii) immediate but minor shifts in the abundance of several species that have remained stable over the course of the experiment; (iii) no change in productivity in response to climate treatments with the exception of reduction from chronic summer drought; and (iv) only minor species losses in response to drought and winter heating. Overall, compositional changes induced by 13-yr exposure to climate regime change were less than short-term fluctuations in species abundances driven by interannual climate fluctuations. The lack of progressive compositional change, coupled with the long-term historical persistence of unproductive grasslands in northern England, suggests the community at Buxton possesses a stabilizing capacity that leads to long-term persistence of dominant species. Unproductive ecosystems provide a refuge for many threatened plants and animals and perform a diversity of ecosystem services. Our results support the view that changing land use and overexploitation rather than climate change per se constitute the primary threats to these fragile ecosystems.

Urban domestic gardens: the effects of human interventions on garden composition.

Private domestic gardens contribute substantially to the biodiversity of urban areas and benefit human health and well-being. We previously reported a study of 267 gardens across five cities in the United Kingdom in which variation in geographical and climatic factors had little bearing on the richness, diversity and composition of plant species. We therefore hypothesise that garden management is an important factor in determining garden characteristics. Here, from the same sample of gardens, we investigate potential associations between the uses to which people put their gardens, the types of management activities they undertake, and the characteristics of those gardens. Householders (n = 265) completed a questionnaire detailing various aspects of garden use and management activities. The majority of respondents used their gardens chiefly for relaxation, recreation, and eating. Fewer than one fifth included "gardening" amongst their garden uses even though all performed some garden management, suggesting that not all management activity resulted from an interest in gardening. Garden-watering and lawn-mowing were the most prevalent activities and were predictors of other types of management including weeding, vegetation-cutting, leaf-collection, and dead-heading flowers. A number of these activities were associated with one another, the richness and composition of plant species, and the number of land uses in gardens. However, relationships between management activities and the amount of tall vegetation were less consistent, and garden management appeared to be independent of garden area. More species of amphibians, birds, and mammals were observed in gardens with ponds and in which efforts were made to attract wildlife, particularly by providing drinking water. This study supports the hypothesis that garden use and management is associated with garden characteristics.