Maize adaptation across temperate climates was obtained via expression of two florigen genes.

Expansion of the maize growing area was central for food security in temperate regions. In addition to the suppression of the short-day requirement for floral induction, it required breeding for a large range of flowering time that compensates the effect of South-North gradients of temperatures. Here we show the role of a novel florigen gene,ZCN12, in the latter adaptation in cooperation withZCN8. Strong eQTLs ofZCN8andZCN12, measured in 327 maize lines, accounted for most of the genetic variance of flowering time in platform and field experiments.ZCN12had a strong effect on flowering time of transgenicArabidopsis thalianaplants; a path analysis showed that it directly affected maize flowering time together withZCN8. The allelic composition atZCNQTLs showed clear signs of selection by breeders. This suggests that florigens played a central role in ensuring a large range of flowering time, necessary for adaptation to temperate areas. Author summary The cultivation of maize in temperate climates required the suppression of photoperiod sensitivity and the selection of a wide range of flowering time for the adaptation to local environmental constraints. Photoperiodic flowering requires the production of a systemic protein signal referred to as the florigen, which is transcriptionally activated in leaves. A key question is to what extent the expression of the florigen gene can explain flowering time variability of temperate maize, where photoperiod sensitivity has been eliminated by artificial selection. Our results reveal large variability in two related florigen genes which is strongly correlated with variability in flowering time. Using association genetics approaches we could detect genomic regions responsible for the expression of the two florigens that precisely co-localise with flowering time-related regions, thereby supporting the significance of quantitative changes in florigen levels in driving flowering time variability. Markers associated with florigen expression/flowering time display significant signatures of selection indicating that variable patterns of florigen accumulation underpin the adaptation of temperate maize flowering.