The contribution of genetic adaptation and plasticity to intraspecific phenotypic variability remains insufficiently studied in long-lived plants, as well as the relevance of neutral versus adaptive processes determining such divergence. We examined the importance of phylogeographic structure and climate in modulating genetic and plastic changes and their interdependence in fitness-related traits of a widespread Mediterranean conifer (Pinus pinaster). Four marker-based, previously defined neutral classifications along with two ad hoc climate-based categorizations of 123 range-wide populations were analyzed for their capacity to summarize genetic and plastic effects of height growth and survival (age 20) in 15 common gardens. The plasticity of tree height and differential survival were interpreted through mixed modeling accounting for heteroscedasticity in the genotype-by-environment dataset. The analysis revealed a slight superiority of phylogeographic classifications over climate categorizations on the explanation of genetic and plastic effects, which suggests that neutral processes can be at least as important as isolation by climate as a driving factor of evolutionary divergence in a prevalent pine. The best phylogeographic classification involved eight geographically discrete genetic groups, which explained 92% (height) and 52% (survival) of phenotypic variability, including between-group mean differentiation and differential expression across trials. For height growth, there was high predictability of plastic group responses described by different reaction norm slopes, which were unrelated to between-group mean differentiation. The latter differences (amounting to ca. 40% among groups) dominated intraspecific performance across trials. Local adaptation was evident for genetic groups tested in their native environments in terms of tree height and, especially, survival. This finding was supported by Q_ST > F_ST estimates. Additionally, our range-wide evaluation did not support a general adaptive syndrome by which less reactive groups to ameliorated conditions would be associated with high survival and low growth. In fact, a lack of relationship between mean group differentiation, indicative of genetic adaptation, and predictable group plasticity for height growth suggests different evolutionary trajectories of these mechanisms of phenotypic divergence. Altogether, the existence of predictable adaptive-trait phenotypic variation for the species, involving both genetic differentiation and plastic effects, should facilitate integrating genomics and environment into decision-making tools to assist forests in coping with climate change.

中文翻译：

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系统发育地理学和气候塑造了常见针叶树的数量遗传景观和范围内的可塑性

在长寿植物中，遗传适应和可塑性对种内表型变异的贡献以及决定这种差异的中性过程与适应过程的相关性仍然没有得到充分研究。我们研究了系统发育地理结构和气候在调节遗传和可塑性变化方面的重要性，以及它们在广泛分布的地中海针叶树（Pinus pinaster）的适应性相关性状中的相互依赖性。对 123 个范围内种群的四个基于标记、先前定义的中性分类以及两个基于气候的临时分类进行了分析，以了解它们总结 15 个常见花园中身高生长和生存（20 岁）的遗传和塑料影响的能力。通过考虑基因型环境数据集中的异方差性的混合模型来解释树高的可塑性和差异存活率。分析显示，在解释遗传和可塑性效应方面，系统发育地理学分类比气候分类稍有优势，这表明中性过程至少与气候隔离一样重要，是流行松树进化分化的驱动因素。最好的系统地理学分类涉及八个地理上离散的遗传群体，这解释了 92%（高度）和 52%（生存）的表型变异，包括组间平均分化和试验中的差异表达。对于身高增长，通过不同反应范数斜率描述的塑料组反应具有高度可预测性，这与组间平均分化无关。后者的差异（组间差异约为 40%）主导了试验中的种内表现。对于在其原生环境中测试的基因组来说，在树高，尤其是生存方面，当地适应是显而易见的。这一发现得到了Q _ST  >  F _ST估计的支持。此外，我们的范围广泛的评估并不支持一般适应性综合症，即对改善条件反应较弱的群体与高存活率和低生长率相关。事实上，平均群体分化（表明遗传适应）与身高增长的可预测群体可塑性之间缺乏关系，这表明这些表型趋异机制的进化轨迹不同。总而言之，物种存在可预测的适应性特征表型变异，包括遗传分化和可塑性效应，应有助于将基因组学和环境整合到决策工具中，以帮助森林应对气候变化。