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BACKGROUND AND AIMS: Reproductive systems enabling opportunities for self-fertilization influence population genetic structure and play a key role in colonization and genetic differentiation during range expansion. Because of their well-developed powers of dispersal, aquatic plants often have widespread disjunct geographical distributions, providing opportunities to investigate the role of reproductive systems in structuring genetic variation between parts of the range that differ in migration history and ecology. METHODS: We compared reproductive systems and spatial genetic structure of the freshwater aquatic macrophyte Sagittaria latifolia between disjunct western and eastern ranges of North America (NA). Populations of this species are most commonly either monoecious or dioecious. We examined chloroplast DNA haplotype variation to test the hypothesis that the western range of this species represents a secondary colonization from the east, and evaluated the roles of reproductive system differences and geography in structuring contemporary patterns of genetic variation at 11 polymorphic SSR (simple sequence repeat) loci. KEY RESULTS: Chloroplast haplotyping revealed a single haplotype in western NA compared to numerous haplotypes in eastern NA, consistent with a genetic bottleneck during westward migration. Estimates of genetic diversity in eastern NA populations differed significantly between reproductive systems, but this pattern was not evident in the western range. Eastern populations could be reliably assigned to genetic clusters based on their reproductive systems, whereas western populations clustered primarily by geographical location. CONCLUSIONS: The sparser distribution of aquatic habitats in the drier western range of S. latifolia, combined with secondary colonization of this region, probably cause the lower genetic diversity and increased differentiation among populations, possibly overriding the effects of reproductive system evident in the eastern portion of the range. Our findings demonstrate that the complex interplay between migratory history, reproductive systems and habitat availability plays an important role in structuring spatial patterns of genetic variation in disjunct plant populations.
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Variación Genética , Sagittaria , ADN de Cloroplastos/genética , Haplotipos/genética , Repeticiones de Microsatélite/genética , Filogenia , Reproducción/genética , Sagittaria/genéticaRESUMEN
Gene copy number variation (CNV) has been increasingly associated with organismal responses to environmental stress, but we know little about the quantitative relation between CNV and phenotypic variation. In this study we quantify the relation between variation in EPSPS (5-enolpyruvylshikimate-3-phosphate synthase) copy number using digital drop PCR and variation in phenotypic glyphosate resistance in 22 populations of Amaranthus palmeri (Palmer Amaranth), a range-expanding agricultural weed. Overall, we detected a significant positive relation between population mean copy number and resistance. The majority of populations exhibited high glyphosate resistance yet maintained low-resistance individuals, resulting in bimodality in many populations. We also investigated threshold models for the relation between copy number and resistance, and found evidence for a threshold of ~15 EPSPS copies: there was a steep increase in resistance below the threshold, followed by a much shallower increase. Across 924 individuals, as copy number increased the range of variation in resistance decreased, yielding an increasing frequency of high phenotypic resistance individuals. Among populations we detected a decline in variation (s.d.) as mean phenotypic resistance increased from moderate to high, consistent with the prediction that as phenotypic resistance increases in populations, stabilizing selection decreases variation in the trait. Our study demonstrates that populations of A. palmeri can harbour wide variation in EPSPS copy number and phenotypic glyphosate resistance, reflecting the history of, and template for future, resistance evolution.
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Amaranthus , Herbicidas , Amaranthus/genética , Variaciones en el Número de Copia de ADN , Dosificación de Gen , Glicina/análogos & derivados , Resistencia a los Herbicidas/genética , Herbicidas/farmacología , Humanos , GlifosatoRESUMEN
Plant mating systems play a key role in structuring genetic variation both within and between species. In hybrid zones, the outcomes and dynamics of hybridization are usually interpreted as the balance between gene flow and selection against hybrids. Yet, mating systems can introduce selective forces that alter these expectations; with diverse outcomes for the level and direction of gene flow depending on variation in outcrossing and whether the mating systems of the species pair are the same or divergent. We present a survey of hybridization in 133 species pairs from 41 plant families and examine how patterns of hybridization vary with mating system. We examine if hybrid zone mode, level of gene flow, asymmetries in gene flow and the frequency of reproductive isolating barriers vary in relation to mating system/s of the species pair. We combine these results with a simulation model and examples from the literature to address two general themes: (1) the two-way interaction between introgression and the evolution of reproductive systems, and (2) how mating system can facilitate or restrict interspecific gene flow. We conclude that examining mating system with hybridization provides unique opportunities to understand divergence and the processes underlying reproductive isolation.
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Flujo Génico , Hibridación Genética , Plantas/genética , Alelos , Simulación por Computador , Cruzamientos Genéticos , Modelos Biológicos , Reproducción/genética , Aislamiento Reproductivo , Autoincompatibilidad en las Plantas con Flores/fisiologíaRESUMEN
Clonality is often implicated in models of the evolution of dioecy, but few studies have explicitly compared clonal structure between plant sexual systems, or between the sexes in dioecious populations. Here, we exploit the occurrence of monoecy and dioecy in clonal Sagittaria latifola (Alismataceae) to evaluate two main hypotheses: (i) clone sizes are smaller in monoecious than dioecious populations, because of constraints imposed on clone size by costs associated with geitonogamy; (ii) in dioecious populations, male clones are larger and flower more often than female clones because of sex-differential reproductive costs. Differences in clone size and flowering could result in discordance between ramet- and genet-based sex ratios. We used spatially explicit sampling to address these hypotheses in 10 monoecious and 11 dioecious populations of S. latifolia at the northern range limit in Eastern North America. In contrast to our predictions, monoecious clones were significantly larger than dioecious clones, probably due to their higher rates of vegetative growth and corm production, and in dioecious populations, there was no difference in clone size between females and males; ramet- and genet-based sex ratios were therefore highly correlated. Genotypic diversity declined with latitude for both sexual systems, but monoecious populations exhibited lower genotypic richness. Differences in life history between the sexual systems of S. latifolia appear to be the most important determinants of clonal structure and diversity.
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Variación Genética , Genética de Población , Sagittaria/genética , Flores/fisiología , Ontario , Quebec , Reproducción , Sagittaria/fisiologíaRESUMEN
While homoploid hybridization was viewed as maladaptive by zoologists, the possibility that it might play a creative role in evolution was explored and debated by botanists during the evolutionary synthesis. Owing to his synthetic work on the ecological and genetic factors influencing the occurrence and effects of hybridization, G. Ledyard Stebbins' contributions to this debate were particularly influential. We revisit Stebbins' views on the frequency of hybridization, the evolution of hybrid sterility, and the evolutionary importance of transgressive segregation, introgression, and homoploid hybrid speciation in the context of contemporary evidence. Floristic surveys indicate that â¼10% of plant species hybridize, suggesting that natural hybridization is not as ubiquitous as Stebbins argued. There is stronger support for his contention that chromosomal sterility is of greater importance in plants than in animals and that selection drives the evolution of hybrid sterility. Stebbins' assertions concerning the frequent occurrence of transgressive segregation and introgressive hybridization have been confirmed by contemporary work, but few studies directly link these phenomena to adaptive evolution or speciation. Stebbins proposed a mechanism by which chromosomal rearrangements partially isolate hybrid lineages and parental species, which spurred the development of the recombinational model of homoploid speciation. While this model has been confirmed empirically, the establishment of reproductively independent hybrid lineages is typically associated with the development of both intrinsic and extrinsic reproductive barriers. We conclude by reflecting on outcomes of hybridization not considered by Stebbins and on possible future research that may extend our understanding of the evolutionary role of hybridization beyond Stebbins' legacy.
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Evolución Biológica , Cromosomas de las Plantas , Hibridación Genética , Plantas/genética , Ecología , Selección Genética , Especificidad de la EspecieRESUMEN
Our study evaluated whether exposure to naphthenic acid fraction compounds (NAFCs) extracted from oil sands process-affected waters (OSPW) has adverse effects on fish embryos that persist into later life. We exposed fathead minnow (Pimephales promelas) embryos to concentrations of NAFCs found in OSPW (2.5-54 mg/L) for 7 days (1 day postfertilization to hatch), then raised surviving larvae in outdoor mesocosms of uncontaminated lake water for 1 month. Embryos exposed to NAFCs were more likely to exhibit malformations (by up to 8-fold) and had slower heart rates (by up to 24%) compared to controls. Fish raised in uncontaminated lake water following exposure to NAFCs as embryos, were 2.5-fold less likely to survive during the larval stage than control fish. These fish also showed up to a 45% decrease in swim activity and a 36% increase in swim burst events during behavioral tests relative to controls. We conclude that exposure to NAFCs during the embryonic stage can have lasting effects on fish survival, physiology, and behavior that persist at least through the larval stage. These findings of delayed mortalities and persistent sublethal effects of embryonic NAFC exposure are relevant to informing the development of regulations on treated OSPW releases from mining operations. Environ Toxicol Chem 2022;41:1319-1332. © 2022 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.
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Cyprinidae , Contaminantes Químicos del Agua , Animales , Cyprinidae/fisiología , Larva , Yacimiento de Petróleo y Gas , Compuestos Orgánicos , Agua , Contaminantes Químicos del Agua/análisis , Contaminantes Químicos del Agua/toxicidadRESUMEN
BACKGROUND AND AIMS: Dimorphism among floral traits can evolve through variation in selection intensity between female and male performance, especially when sex functions are separated between flowers on a plant (monoecy), or between individuals (dioecy). In animal-pollinated species, male floral traits are predicted to be larger because competition for pollinators should favour larger displays. Floral dimorphism may be greater in dioecious than monoecious populations because of trade-offs between female and male function and opportunities for selfing in hermaphrodites. METHODS: These predictions were tested by surveying flower size, total flowers per inflorescence and daily display size in the insect-pollinated Sagittaria latifolia (Alismataceae). This species is useful for comparative analysis because populations are mostly either monoecious or dioecious. We examined floral dimorphism in 13 monoecious and 16 dioecious populations in eastern North America. KEY RESULTS: Male flowers were significantly larger than female flowers in monoecious and dioecious populations, but there was no evidence for greater flower size dimorphism in dioecious populations despite their larger flower sizes overall. Although inflorescences in both dioecious and monoecious populations produced more male flowers, daily floral displays were significantly larger for female than male function due to more synchronous female flower opening. Daily floral display dimorphism was significantly greater in dioecious populations, due to greater female daily floral displays. There was a positive relationship between mean flower size and total flowers per inflorescence for both sexes in dioecious populations, but no relationship for either sex function in monoecious populations. Flower size dimorphism was positively correlated with the frequencies of females in dioecious populations. CONCLUSIONS: The increased size and number of male flowers and protracted male floral displays in S. latifolia are probably shaped by sexual selection for more effective pollen dispersal.
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Óvulo Vegetal/fisiología , Polen/fisiología , Sagittaria/fisiología , Canadá , Ritmo Circadiano/fisiología , Inflorescencia/anatomía & histología , Inflorescencia/fisiología , Tamaño de los Órganos , Carácter Cuantitativo Heredable , Sagittaria/anatomía & histología , Razón de MasculinidadRESUMEN
Domesticated plants and animals often display dramatic responses to selection, but the origins of the genetic diversity underlying these responses remain poorly understood. Despite domestication and improvement bottlenecks, the cultivated sunflower remains highly variable genetically, possibly due to hybridization with wild relatives. To characterize genetic diversity in the sunflower and to quantify contributions from wild relatives, we sequenced 287 cultivated lines, 17 Native American landraces and 189 wild accessions representing 11 compatible wild species. Cultivar sequences failing to map to the sunflower reference were assembled de novo for each genotype to determine the gene repertoire, or 'pan-genome', of the cultivated sunflower. Assembled genes were then compared to the wild species to estimate origins. Results indicate that the cultivated sunflower pan-genome comprises 61,205 genes, of which 27% vary across genotypes. Approximately 10% of the cultivated sunflower pan-genome is derived through introgression from wild sunflower species, and 1.5% of genes originated solely through introgression. Gene ontology functional analyses further indicate that genes associated with biotic resistance are over-represented among introgressed regions, an observation consistent with breeding records. Analyses of allelic variation associated with downy mildew resistance provide an example in which such introgressions have contributed to resistance to a globally challenging disease.
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Helianthus/genética , Helianthus/microbiología , Hibridación Genética , Enfermedades de las Plantas/genética , Productos Agrícolas/genética , Productos Agrícolas/microbiología , Resistencia a la Enfermedad/genética , Ontología de Genes , Genes de Plantas , Variación Genética , Genoma de Planta , Enfermedades de las Plantas/microbiología , Recombinación Genética , Selección GenéticaRESUMEN
Geographically peripheral populations are expected to exhibit lower genetic diversity and higher differentiation than central populations because of their smaller size and greater spatial isolation. In plants, a shift from sexual to clonal asexual reproduction may further reduce diversity and increase differentiation. Here, these predictions were tested by assaying 36 inter-simple sequence repeat (ISSR) polymorphisms in 21 populations of the woody, clonal plant Vaccinium stamineum in eastern North America, from the range center to its northern limit where it has 'threatened' status. Populations decline in frequency, but not size or sexual reproductive output, across the range. Within-population diversity did not decline towards range margins. Modest genetic differentiation among populations increased slightly towards range margins and in small populations with high clonal propagation and low seed production, although none of these trends was significant. Low seed production and high clonal propagation were not associated with large-scale clonal spread. By combining demographic and genetic data, this study determined that increased population isolation, rather than reduced population size, can account for the weak increase in genetic differentiation at range margins.
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Variación Genética , Genética de Población , Vaccinium/genética , Análisis de Varianza , Geografía , Repeticiones de Minisatélite , Reproducción/genéticaRESUMEN
The stable coexistence within populations of females, males, and hermaphrodites (subdioecy) is enigmatic because theoretical models indicate that maintenance of this sexual system involves highly restricted conditions. Subdioecy is more commonly interpreted as a transitory stage along the gynodioecious pathway from hermaphroditism to dioecy. The widespread, North American, aquatic plant Sagittaria latifolia is largely composed of monoecious or dioecious populations; however, subdioecious populations with high frequencies of hermaphrodites (mean frequency = 0.50) characterize the northern range boundary of dioecy in eastern North America. We investigated two hypotheses for the origin of subdioecy in this region. Using polymorphic microsatellite loci, we evaluated whether subdioecy arises through selection on standing genetic variation for male sex inconstancy in dioecious populations, or results from hybridization between monoecious and dioecious populations. We found evidence for both pathways to subdioecy, although hybridization was the more common mechanism, with genetic evidence of admixture in nine of 14 subdioecious populations examined. Hybridization has also played a role in the origin of androdioecious populations in S. latifolia, a mechanism not often considered in the evolution of this rare sexual system. Our study demonstrates how hybridization has the potential to play a role in the diversification of plant sexual systems.
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Evolución Biológica , Hibridación Genética , Polimorfismo Genético , Sagittaria/fisiología , Canadá , Repeticiones de Microsatélite , Reproducción , Sagittaria/genética , Estados UnidosRESUMEN
In many angiosperm species, populations are reproductively subdivided into distinct sexual morphs including females, males and hermaphrodites. Sexual polymorphism is maintained by frequency-dependent selection, leading to predictable sex ratios at equilibrium. Charles Darwin devoted much of his book 'The Different Forms of Flowers on Plants of the Same Species' (1877) to investigating plant sexual polymorphisms and laid the foundation for many problems addressed today by integrating theory with empirical studies of the demography and genetics of populations. Here, we summarize our recent work on the ecological and genetic mechanisms influencing variation in sex ratios and their implications for evolutionary transitions among sexual systems. We present the results of a survey of sex ratios from 126 species from 47 angiosperm families and then address two general problems using examples from diverse angiosperm taxa: (i) the mechanisms governing biased sex ratios in dioecious species; (ii) the origins and maintenance of populations composed of females, males and hermaphrodites. Several themes are emphasized, including the importance of non-equilibrium conditions, the role of life history and demography in affecting sex ratios, the value of theory for modelling the dynamics of sex ratio variation, and the utility of genetic markers for investigating evolutionary processes in sexually polymorphic plant populations.
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Evolución Biológica , Genética de Población , Modelos Biológicos , Plantas/genética , Polimorfismo Genético , Razón de Masculinidad , Especificidad de la EspecieRESUMEN
We developed 11 microsatellite loci for Sagittaria latifolia, an aquatic plant common to wetlands of North America. From an (AG)-enriched library, we identified 66 unique microsatellite sequences for which primers could be designed. Twenty-two loci reliably amplified a clear single band of expected size, and 11 loci were scoreable and polymorphic. For these 11 loci, we genotyped a monoecious and a dioecious population, yielding four to 14 alleles per locus. Three loci exhibited significant linkage disequilibrium leaving eight independent variable loci. Eight loci also amplified in four other Sagittaria species. These microsatellite loci will be useful to compare genetic structure among monoecious and dioecious populations of S. latifolia.
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Geographically peripheral populations of widespread species are often the focus of conservation because they are locally rare within political jurisdictions. Yet the ecology and genetics of these populations are rarely evaluated in a broader geographic context. Most expectations concerning the ecology and evolution of peripheral populations derive from the abundant-center model, which predicts that peripheral populations should be less frequent, smaller, less dense, and have a lower reproductive rate than central populations. We tested these predictions and in doing so evaluated the conservation value of peripheral populations for the clonal shrub Vaccinium stamineum L. (Ericaceae, deerberry), which is listed as threatened in Canada. Based on 51 populations sampled from the center to the northern range limits over 2 years, population frequency and size declined toward the range limit, but ramet density increased. Sexual reproductive output varied widely among populations and between years, with many populations producing very few seeds, but did not decline toward range margins. In fact seed mass increased steadily toward range limit, and this was associated with faster germination and seedling growth, which may be adaptive in seasonal northern environments. Our results did not support the prediction that clonal reproduction is more prevalent in peripheral populations or that it contributed antagonistically to the wide variation in seed production. Peripheral populations of V. stamineum are as productive as central populations and may be locally adapted to northern environments. This emphasizes the importance of a broad geographical perspective for evaluating the ecology, evolution, and conservation of peripheral populations.