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Clinal Variation in Flowering Time and Vernalisation Requirement across a 3000-M Altitudinal Range in Perennial Arabidopsis kamchatica Ssp.Kamchatica and Annual Lowland Subspecies Kawasakiana

Tanaka Kenta, Ayumu Yamada and Yoshihiko Onda

Clines in ecologically important traits across environmental gradients provide evidence of historical natural selection. Among the many adaptive traits that are relevant in natural habitats, flowering timing is a primary determinant of a plant’s lifetime fitness. In a laboratory experiment, we studied the divergence in flowering time and vernalisation requirement among 38 populations of two Arabidopsis subspecies: perennial A. kamchatica ssp. kamchatica across an altitudinal gradient from 30 to nearly 3000 m in the Japanese Alps, and its annual lowland subspecies, kawasakiana. Flowering time with vernalisation was not different and flowering time without vernalisation was slightly different between the subspecies; however, the altitude of the source populations was the dominant determinant of the variation in these traits for the species as a whole. The flowering time increased linearly and the effect of vernalisation to shorten the flowering time increased non-linearly with increasing altitude of the source population. Low-altitude populations of ssp. kamchatica from laboratory-collected seeds showed a stronger response to vernalisation than field-collected seeds, suggesting an effect of the maternal environment. By replicating the altitudinal gradient and explicitly accounting for the maternal effect, our results clearly suggest the existence of genetically based clinal patterns that provide signs of adaptive evolution. Early flowering is probably advantageous to completing reproduction before the hot summer at low altitudes. In ssp. kamchatica, which is perennial and can reproduce even if it does not flower in its first year, the strong vernalisation requirement may delay the first reproduction at high altitudes, where the growing season is short.