The fertilization process in Lithocarpus dealbatus (Fagaceae) and its implication on the sexual reproduction evolution of Fagales.

MAIN CONCLUSION: The pistillate flowers of Lithocarpus dealbatus show two pollen tube (PT) arresting sites (the style-joining and micropyle) within the pistil during the postpollination-prezygotic stage. The PT, arrested at the pre-ovule stage, enhanced PT competition allowing the most compatible PTs to enter the ovary to ensure the highest fertilization success. During the shift from animal pollination to wind pollination, plants require a series of changes in reproductive traits. The mode of pollination is striking labile in Fagaceae. Lithocarpus is insect pollinated and is closely related to wind-pollinated Quercus. Little is known about the sexual reproduction of Lithocarpus. This study aimed to reveal the sexual reproduction of Lithocarpus dealbatus and to explore the evolutionary pattern of the key sexual reproduction traits to better understand their possible role in labile pollination. We found that after pollination, L. dealbatus PTs grew slowly in the style reaching style-joining in mid-January of the second year; then PT growth was arrested at style-joining for four months. Only two to three PTs resumed growth in mid-May to reach the micropyle, where PT growth ceased for one month before one PT resumed growth and passed through the micropyle to the embryo sac. Fagaceae showed a generalized mating system. Vast pollen production, small-sized pollen grains, long stigmatic receptive time, and reduced perianth were compatible with beetle pollination syndrome, representing the plesiomorphic status in Fagaceae. A large stigmatic surface and dry pollen grains linked to wind pollination might be independently derived several times in fagaceous lineages. Beetle pollination syndrome can cope with the uncertainty of pollinators to ensure conspecific pollen capture, which represents pre-adaptation status and has a selective advantage when conditions change, favouring wind pollination. The arrest of the PT at style-joining is a unique mechanism in later derived fagaceous lineages to enhance PT competition and promote outcrossing.

Flower morphology as a predictor of pollination mode in a biotic to abiotic pollination continuum.

BACKGROUND AND AIMS: Wind pollination has evolved repeatedly in flowering plants, yet the identification of a wind pollination syndrome as a set of integrated floral traits can be elusive. Thalictrum (Ranunculaceae) comprises temperate perennial herbs that have transitioned repeatedly from insect to wind pollination while also exhibiting mixed pollination, providing an ideal system to test for evolutionary correlation between floral morphology and pollination mode in a biotic to abiotic continuum. Moreover, the lack of floral organ fusion across this genus allows testing for specialization to pollination vectors in the absence of this feature. METHODS: We expanded phylogenetic sampling in the genus from a previous study using six chloroplast loci, which allowed us to test whether species cluster into distinct pollination syndromes based on floral morphology. We then used multivariate analyses on floral traits followed by ancestral state reconstruction of the emerging flower morphotypes and determined whether these traits are evolutionarily correlated under a Bayesian framework with Brownian motion. KEY RESULTS: Floral traits fell into five distinct clusters, which were reduced to three after considering phylogenetic relatedness and were largely consistent with flower morphotypes and associated pollination vectors. Multivariate evolutionary analyses found a positive correlation between the lengths of floral reproductive structures (styles, stigmas, filaments and anthers). Shorter reproductive structures tracked insect-pollinated species and clades in the phylogeny, whereas longer structures tracked wind-pollinated ones, consistent with selective pressures exerted by biotic vs. abiotic pollination vectors, respectively. CONCLUSIONS: Although detectable suites of integrated floral traits across Thalictrum were correlated with wind or insect pollination at the extremes of the morphospace distribution, a presumed intermediate, mixed pollination mode morphospace was also detected. Thus, our data broadly support the existence of detectable flower morphotypes from convergent evolution underlying the evolution of pollination mode in Thalictrum, presumably via different paths from an ancestral mixed pollination state.

From pollen dispersal to plant diversification: genetic consequences of pollination mode.

Pollinators influence patterns of plant speciation, and one intuitive hypothesis is that pollinators affect rates of plant diversification through their effects on pollen dispersal. By specifying mating events and pollen flow across the landscape, distinct types of pollinators may cause different opportunities for allopatric speciation. This pollen dispersal-dependent speciation hypothesis predicts that pollination mode has effects on the spatial context of mating events that scale up to impact population structure and rates of species formation. Here I consider recent comparative studies, including genetic analyses of plant mating events, population structure and comparative phylogenetic analyses, to examine evidence for this model. These studies suggest that highly mobile pollinators conduct greater gene flow within and among populations, compared to less mobile pollinators. These differences influence patterns of population structure across the landscape. However, the effects of pollination mode on speciation rates is less predictable. In some contexts, the predicted effects of pollen dispersal are outweighed by other factors that govern speciation rates. A multiscale approach to examine effects of pollination mode on plant mating system, population structure and rates of diversification is key to determining the role of pollen dispersal on plant speciation for model clades.

Global patterns of population genetic differentiation in seed plants.

Evaluating the factors that drive patterns of population differentiation in plants is critical for understanding several biological processes such as local adaptation and incipient speciation. Previous studies have given conflicting results regarding the significance of pollination mode, seed dispersal mode, mating system, growth form and latitudinal region in shaping patterns of genetic structure, as estimated by FST values, and no study to date has tested their relative importance together across a broad scale. Here, we assembled a 337-species data set for seed plants from publications with data on FST from nuclear markers and species traits, including variables pertaining to the sampling scheme of each study. We used species traits, while accounting for sampling variables, to perform phylogenetic multiple regressions. Results demonstrated that FST values were higher for tropical, mixed-mating, non-woody species pollinated by small insects, indicating greater population differentiation, and lower for temperate, outcrossing trees pollinated by wind. Among the factors we tested, latitudinal region explained the largest portion of variance, followed by pollination mode, mating system and growth form, while seed dispersal mode did not significantly relate to FST . Our analyses provide the most robust and comprehensive evaluation to date of the main ecological factors predicted to drive population differentiation in seed plants, with important implications for understanding the basis of their genetic divergence. Our study supports previous findings showing greater population differentiation in tropical regions and is the first that we are aware of to robustly demonstrate greater population differentiation in species pollinated by small insects.

Flowering Phenology Shifts in Response to Functional Traits, Growth Form, and Phylogeny of Woody Species in a Desert Area.

Climatic factors are considered the major driving forces for variation of flowering phenology among species. Yet, whether flowering phenology of woody species varies with functional traits, growth form, and phylogeny in arid regions is unknown. In the present study, we evaluated the relationships of three characteristics of flowering phenology (i.e., first flowering date, end of flowering date, and flowering duration) against functional traits, growth form, and phylogeny across 59 woody plant species across 3 years in Ürümqi city of the Xinjiang Autonomous Region, in Northwest China. The results showed that, plant functional traits and growth form had significant influences on the variability of flowering phenology among species. The contributions of fruit type (34.7-43.5%) and flower color (30.1-30.7%) to the variability of flowering phenology were larger than those of pollination mode (4.6-14.4%), life form (8.4-14%) and maximum plant height (9.7-13.1%). Trees had the significant correlations in terms of flowering duration against first flowering date and end of flowering date, while shrubs showed the opposite pattern. The values of phylogenetic signal (Blomberg's K) of the three characteristics of flowering phenology ranged from 0.36 to 0.43, which were significantly lower than the expectation of the Brownian motion model. Our results suggested that functional traits, growth form and phylogeny all affected variability of flowering phenology among species. Our results provide a new perspective for correctly evaluating the relationship between global climate change and plant reproduction.