Macro- and micro-geographic genetic variation in early fitness traits in populations of maritime pine (Pinus pinaster Ait.).

BACKGROUND AND AIMS: Assessing adaptive genetic variation and its spatial distribution is crucial to conserve forest genetic resources and manage species' adaptive potential. Macro-environmental gradients commonly exert divergent selective pressures that enhance adaptive genetic divergence among populations. Steep micro-environmental variation might also result in adaptive divergence at finer spatial scales, even under high gene flow, but it is unclear how often this is the case. Here, we assess genetic variation in early fitness traits among distant and nearby maritime pine (Pinus pinaster Ait.) populations, to investigate climatic factors associated with trait divergence, and to examine trait integration during seedling establishment. METHODS: Open pollinated seeds were collected from seven population pairs across the European species distribution, with paired populations spatially close (between <1 km up to 21 km) but environmentally divergent. Seeds were sown in semi-natural conditions at three environmentally contrasting sites, where we monitored seedling emergence, growth and survival. KEY RESULTS: At large spatial scales, we found significant genetic divergence among populations in all studied traits, with certain traits exhibiting association with temperature and precipitation gradients. Significant trait divergence was also detected between pairs of nearby populations. Besides, we found consistent trait correlations across experimental sites, notably heavier seeds and earlier seedling emergence were both associated with higher seedling survival and fitness over two years in all experimental conditions. CONCLUSIONS: We identified mean annual temperature and precipitation seasonality as potential drivers of P. pinaster population divergence in the studied early-life traits. Populations genetically diverge also at local spatial scales, potentially suggesting that divergent natural selection can override gene flow along local-scale ecological gradients. These results suggest the species exhibits substantial adaptive potential that has allowed it to survive and evolve under contrasting environmental conditions.

Polygenic adaptation and negative selection across traits, years and environments in a long-lived plant species (Pinus pinaster Ait., Pinaceae).

A decade of genetic association studies in multiple organisms suggests that most complex traits are polygenic; that is, they have a genetic architecture determined by numerous loci, each with small effect-size. Thus, determining the degree of polygenicity and its variation across traits, environments and time is crucial to understand the genetic basis of phenotypic variation. We applied multilocus approaches to estimate the degree of polygenicity of fitness-related traits in a long-lived plant (Pinus pinaster Ait., maritime pine) and to analyse this variation across environments and years. We evaluated five categories of fitness-related traits (survival, height, phenology, functional, and biotic-stress response) in a clonal common-garden network planted in contrasted environments (over 20,500 trees). Most of the analysed traits showed evidence of local adaptation based on Qst -Fst comparisons. We further observed a remarkably stable degree of polygenicity, averaging 6% (range of 0%-27%), across traits, environments and years. We detected evidence of negative selection, which could explain, at least partially, the high degree of polygenicity. Because polygenic adaptation can occur rapidly, our results suggest that current predictions on the capacity of natural forest tree populations to adapt to new environments should be revised, especially in the current context of climate change.

Genetic basis of growth, spring phenology, and susceptibility to biotic stressors in maritime pine.

Forest ecosystems are increasingly challenged by extreme events, for example, drought, storms, pest attacks, and fungal pathogen outbreaks, causing severe ecological and economic losses. Understanding the genetic basis of adaptive traits in tree species is of key importance to preserve forest ecosystems, as genetic variation in a trait (i.e., heritability) determines its potential for human-mediated or evolutionary change. Maritime pine (Pinus pinaster Aiton), a conifer widely distributed in southwestern Europe and northwestern Africa, grows under contrasted environmental conditions promoting local adaptation. Genetic variation at adaptive phenotypes, including height, spring phenology, and susceptibility to two fungal pathogens (Diplodia sapinea and Armillaria ostoyae) and an insect pest (Thaumetopoea pityocampa), was assessed in a range-wide clonal common garden of maritime pine. Broad-sense heritability was significant for height (0.219), spring phenology (0.165-0.310), and pathogen susceptibility (necrosis length caused by D. sapinea, 0.152; and by A. ostoyae, 0.021, measured on inoculated, excised branches under controlled conditions), but not for pine processionary moth incidence in the common garden. The correlations of trait variation among populations revealed contrasting trends for pathogen susceptibility to D. sapinea and A. ostoyae with respect to height. Taller trees showed longer necrosis length caused by D. sapinea while shorter trees were more affected by A. ostoyae. Moreover, maritime pine populations from areas with high summer temperatures and frequent droughts were less susceptible to D. sapinea but more susceptible to A. ostoyae. Finally, an association study using 4227 genome-wide SNPs revealed several loci significantly associated with each trait (range of 3-26), including a possibly disease-induced translation initiation factor, eIF-5, associated with needle discoloration caused by D. sapinea. This study provides important insights to develop genetic conservation and breeding strategies integrating species responses to biotic stressors.

Author Correction: The GenTree Dendroecological Collection, tree-ring and wood density data from seven tree species across Europe.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

The GenTree Dendroecological Collection, tree-ring and wood density data from seven tree species across Europe.

The dataset presented here was collected by the GenTree project (EU-Horizon 2020), which aims to improve the use of forest genetic resources across Europe by better understanding how trees adapt to their local environment. This dataset of individual tree-core characteristics including ring-width series and whole-core wood density was collected for seven ecologically and economically important European tree species: silver birch (Betula pendula), European beech (Fagus sylvatica), Norway spruce (Picea abies), European black poplar (Populus nigra), maritime pine (Pinus pinaster), Scots pine (Pinus sylvestris), and sessile oak (Quercus petraea). Tree-ring width measurements were obtained from 3600 trees in 142 populations and whole-core wood density was measured for 3098 trees in 125 populations. This dataset covers most of the geographical and climatic range occupied by the selected species. The potential use of it will be highly valuable for assessing ecological and evolutionary responses to environmental conditions as well as for model development and parameterization, to predict adaptability under climate change scenarios.

The heart of Blessed Anne-Madeleine Remuzat: a biomedical approach of "miraculous" heart conservation.

We present here the results of our inter-disciplinary examination of the mummified heart of Blessed Anne-Madeleine Remuzat (1696-1730). This organ has been examined in the context of a canonization process. This analysis is related to important aspects of the early history of anatomy in Europe, that of "Holy autopsies", and to the relationship between anatomical investigations, Catholic theology, and religious/medical customs. According to anatomical, genetic, toxicological, and palynological analyses, it has been shown that this organ has not been naturally ("miraculously") conserved but embalmed using myrtle, honey, and lime. Moreover, a right ventricle dilatation has been diagnosed, that may represent a post-tuberculosis condition and may have played a role in the cause of death of this religious figure.