Crops under past diversification and ongoing climate change: more than just selection of nuclear genes for flowering

Diversification and breeding following domestication and under current climate change across the globe are the two most significant evolutionary events experienced by major crops. Diversification of crops from their wild ancestors has favored dramatic changes in the sensitivity of the plants to the environment, particularly significantly in transducing light inputs to the circadian clock, which has allowed the growth of major crops in the relatively short growing season experienced in the Northern Hemisphere. Historically, mutants and the mapping of quantitative trait loci (QTL) have facilitated the identification and the cloning of genes that underlie major changes of the clock and the regulation of flowering. Recent studies have suggested that the thermal plasticity of the circadian clock output, and not just the core genes that follow temperature compensation, has also been under selection during diversification and breeding. Wild alleles that accelerate output rhythmicity could be beneficial for crop resilience. Furthermore, wild alleles with beneficial and flowering-independent effects under stress indicate their possible role in maintaining a balanced source-sink relationship, thereby allowing productivity under climatic change. Because the chloroplast genome also regulates the plasticity of the clock output, mapping populations including cytonuclear interactions should be utilized within an integrated field and clock phenomics framework. In this review, we highlight the need to integrate physiological and developmental approaches (physio-devo) to gain a better understanding when re-domesticating wild gene alleles into modern cultivars to increase their robustness under abiotic heat and drought stresses.