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Environmentally covarying local adaptation is a form of cryptic local adaptation in which the covariance of the genetic and environmental effects on a phenotype obscures the divergence between locally adapted genotypes. Here, we systematically document the magnitude and drivers of the genetic effect (V G ) for two forms of environmentally covarying local adaptation: counter- and cogradient variation. Using a hierarchical Bayesian meta-analysis, we calculated the overall effect size of V G as 1.05 and 2.13 for populations exhibiting countergradient or cogradient variation, respectively. These results indicate that the genetic contribution to phenotypic variation represents a 1.05 to 2.13 s.d. change in trait value between the most disparate populations depending on if populations are expressing counter- or cogradient variation. We also found that while there was substantial variance among abiotic and biotic covariates, the covariates with the largest mean effects were temperature (2.41) and gamete size (2.81). Our results demonstrate the pervasiveness and large genetic effects underlying environmentally covarying local adaptation in wild populations and highlight the importance of accounting for these effects in future studies.

Genetic and physiological mechanisms of freezing tolerance in locally adapted populations of a winter annual

https://doi.org/10.1002/ajb2.1385

Sanderson, Brian J. ; Park, Sunchung ; Jameel, M. Inam ; Kraft, Joshua C. ; Thomashow, Michael F. ; Schemske, Douglas W. ; Oakley, Christopher G. ( November 2019 , American Journal of Botany)

Premise

Despite myriad examples of local adaptation, the phenotypes and genetic variants underlying such adaptive differentiation are seldom known. Recent work on freezing tolerance and local adaptation in ecotypes ofArabidopsis thalianafrom Italy and Sweden provides an essential foundation for uncovering the genotype–phenotype–fitness map for an adaptive response to a key environmental stress.

Methods

We examined the consequences of a naturally occurring loss‐of‐function (LOF) mutation in an Italian allele of the gene that encodes the transcription factorCBF2,which underlies a major freezing‐tolerance locus. We used four lines with a Swedish genetic background, each containing aLOFCBF2allele. Two lines had introgression segments containing the ItalianCBF2allele, and two contained deletions created usingCRISPR‐Cas9. We used a growth chamber experiment to quantify freezing tolerance and gene expression before and after cold acclimation.

Results

Freezing tolerance was lower in the Italian (11%) compared to the Swedish (72%) ecotype, and all four experimentalCBF2LOFlines had reduced freezing tolerance compared to the Swedish ecotype. Differential expression analyses identified 10 genes for which allCBF2LOFlines, and theITecotype had similar patterns of reduced cold responsive expression compared to theSWecotype.

Conclusions

We identified 10 genes that are at least partially regulated byCBF2that may contribute to the differences in cold‐acclimated freezing tolerance between the Italian and Swedish ecotypes. These results provide novel insight into the molecular and physiological mechanisms connecting a naturally occurring sequence polymorphism to an adaptive response to freezing conditions.