From genotype to phenotype: unraveling the complexities of cold adaptation in forest trees1

Adaptation to winter cold in temperate and boreal trees involves complex genetic, physiological, and developmental processes. Genecological studies demonstrate the existence of steep genetic clines for cold adaptation traits in relation to environmental (mostly temperature related) gradients. Population differentiation is generally stronger for cold adaptation traits than for other quantitative traits and allozymes. Therefore, these traits appear to be under strong natural selection. Nonetheless, high levels of genetic variation persist within populations. The genetic control of cold adaptation traits ranges from weak to strong, with phenological traits having the highest heritabilities. Within-population genetic correlations among traits range from negligible to moderate. Generally, bud phenology and cold hardiness in the fall are genetically uncorrelated with bud phenology and cold hardiness in the spring. Analyses of quantitative trait loci indicate that cold adaptation traits are mostly controlled by multiple genes with small effects and that quantitative trait loci × environment interactions are common. Given this inherent complexity, we suggest that future research should focus on identifying and developing markers for cold adaptation candidate genes, then using multilocus, multiallelic analytical techniques to uncover the relationships between genotype and phenotype at both the individual and population levels. Ultimately, these methods may be useful for predicting the performance of genotypes in breeding programs and for better understanding the evolutionary ecology of forest trees.