RESUMO
Extant conifer species may be susceptible to rapid environmental change owing to their long generation times, but could also be resilient due to high levels of standing genetic diversity. Hybridisation between closely related species can increase genetic diversity and generate novel allelic combinations capable of fuelling adaptive evolution. Our study unravelled the genetic architecture of adaptive evolution in a conifer hybrid zone formed between Pinus strobiformis and P. flexilis. Using a multifaceted approach emphasising the spatial and environmental patterns of linkage disequilibrium and ancestry enrichment, we identified recently introgressed and background genetic variants to be driving adaptive evolution along different environmental gradients. Specifically, recently introgressed variants from P. flexilis were favoured along freeze-related environmental gradients, while background variants were favoured along water availability-related gradients. We posit that such mosaics of allelic variants within conifer hybrid zones will confer upon them greater resilience to ongoing and future environmental change and can be a key resource for conservation efforts.
Assuntos
Adaptação Biológica/genética , Introgressão Genética/fisiologia , Especiação Genética , Traqueófitas/genética , Alelos , Arizona , Evolução Biológica , Variação Genética/fisiologia , Geografia , Hibridização Genética/fisiologia , México , Mosaicismo , Pinus/genética , Polimorfismo de Nucleotídeo Único , Traqueófitas/classificaçãoRESUMO
The phenotype of trees is determined by the relationships and interactions among genetic and environmental influences. Understanding the patterns and processes that are responsible for phenotypic variation is facilitated by studying the relationships between phenotype and the environment among many individuals across broad ecological and climatic gradients. We used Pinus strobiformis, which has a wide latitudinal distribution, as a model species to: (a) estimate the relative importance of different environmental factors in predicting these morphological traits and (b) characterize the spatial patterns of standing phenotypic variation of cone and seed traits across the species' range. A large portion of the total variation in morphological characteristics was explained by ecological, climatic and geographical variables (54.7% collectively). The three climate, vegetation and geographical variable groups, each had similar total ability to explain morphological variation (43.4%, 43.8%, 51.5%, respectively), while the topographical variable group had somewhat lower total explanatory power (36.9%). The largest component of explained variance (33.6%) was the four-way interaction of all variable sets, suggesting that there is strong covariation in environmental, climate and geographical variables in their relationship to morphological traits of southwest white pine across its range. The regression results showed that populations in more humid and warmer climates expressed greater cone length and seed size. This may in part facilitate populations of P. strobiformis in warmer and wetter portions of its range growing in dense, shady forest stands, because larger seeds provide greater resources to germinants at the time of germination. Our models provide accurate predictions of morphological traits and important insights regarding the factors that contribute to their expression. Our results indicate that managers should be conservative during reforestation efforts to ensure match between ecotypic variation in seed source populations. However, we also note that given projected large range shift due to climate change, managers will have to balance the match between current ecotypic variation and expected range shift and changes in local adaptive optima under future climate conditions.
RESUMO
Elucidating phylogenetic relationships and species boundaries within complex taxonomic groups is challenging for intrinsic and extrinsic (i.e., technical) reasons. Mexican pinyon pines are a complex group whose phylogenetic relationships and species boundaries have been widely studied but poorly resolved, partly due to intrinsic ecological and evolutionary features such as low morphological and genetic differentiation caused by recent divergence, hybridization and introgression. Extrinsic factors such as limited sampling and difficulty in selecting informative molecular markers have also impeded progress. Some of the Mexican pinyon pines are of conservation concern but others may remain unprotected because the species boundaries have not been established. In this study we combined approaches to resolve the phylogenetic relationships in this complex group and to establish species boundaries in four recently diverged taxa: P. discolor, P. johannis, P. culminicola and P. cembroides. We performed phylogenetic analyses using the chloroplast markers matK and psbA-trnH as well as complete and partial chloroplast genomes of species of Subsection Cembroides. Additionally, we performed a phylogeographic analysis combining genetic data (18 chloroplast markers), morphological data and geographical data to define species boundaries in four recently diverged taxa. Ecological divergence was supported by differences in climate among localities for distinct genetic lineages. Whereas the phylogenetic analysis inferred with matK and psbA-trnH was unable to resolve the relationships in this complex group, we obtained a resolved phylogeny with the use of the chloroplast genomes. The resolved phylogeny was concordant with a haplotype network obtained using chloroplast markers. In species with potential for recent divergence, hybridization or introgression, nonhierarchical network-based approaches are probably more appropriate to protect against misclassification due to incomplete lineage sorting. The boundaries among genetic lineages were delimited by the inclusion of morphological, geographical and ecological data in the haplotype network. These multiple lines of evidence help to assign species boundaries in this complex group. P. johannis, P. discolor, P. culminicola and P. cembroides are different species based on their genetic, morphological and ecological niche differences. We suggest a reevaluation of the conservation status of these species considering the information generated in this study.
Assuntos
Evolução Molecular , Filogenia , Pinus/classificação , Teorema de Bayes , Conservação dos Recursos Naturais , DNA de Cloroplastos/genética , DNA de Plantas/genética , Genoma de Cloroplastos , Haplótipos , México , Repetições de Microssatélites , Modelos Genéticos , Filogeografia , Pinus/genética , Análise de Sequência de DNARESUMO
BACKGROUND AND AIMS: The sexual separation in dioecious species has interested biologists for decades; however, the cellular mechanism leading to unisexuality has been poorly understood. In this study, the cellular changes that lead to male sterility in the functionally dioecious cactus, Opuntia stenopetala, are described. METHODS: The spatial and temporal patterns of programmed cell death (PCD) were determined in the anthers of male and female flowers using scanning electron microscopy analysis and histological observations, focusing attention on the transition from bisexual to unisexual development. In addition, terminal deoxynucleotidyl transferase-mediated dUTP nick-end labelling assays were used as an indicator of DNA fragmentation to corroborate PCD. KEY RESULTS: PCD was detected in anthers of both female and male flowers, but their patterns differed in time and space. Functionally male individuals developed viable pollen, and normal development involved PCD on each layer of the anther wall, which occurred progressively from the inner (tapetum) to the outer layer (epidermis). Conversely, functional female individuals aborted anthers by premature and displaced PCD. In anthers of female flowers, the first signs of PCD, such as a nucleus with irregular shape, fragmented and condensed chromatin, high vacuolization and condensed cytoplasm, occurred at the microspore mother cell stage. Later these features were observed simultaneously in all anther wall layers, connective tissue and filament. Neither pollen formation nor anther dehiscence was detected in female flowers of O. stenopetala due to total anther disruption. CONCLUSIONS: Temporal and spatial changes in the patterns of PCD are responsible for male sterility of female flowers in O. stenopetala. Male fertility requires the co-ordination of different events, which, when altered, can lead to male sterility and to functionally unisexual individuals. PCD could be a widespread mechanism in the determination of functionally dioecious species.