A CYCLOIDEA-like gene mutation in sunflower determines an unusual floret type able to produce filled achenes at the periphery of the pseudanthium

The pseudanthium of sunflower (Helianthus annuus L.) consists of two floret types: zygomorphic sterile ray florets and actinomorphic hermaphrodite disc florets. In the tubular ray flower (turf) mutant,the loss-of-function of a CYCLOIDEA (CYC) gene generates hermaphrodite tubular-like ray florets that replace the normal sterile ray florets. We evaluated whether tubular-like ray florets have a multifaceted set of floral traits and the presence of heteromorphic seeds in the turf inflorescence. During early stages of floral ontogeny, primordia of both tubular-like ray florets and typical ray florets displayed a comparable shape. In contrast, during later stages of development, the form of tubular-like ray floret primordia was most similar to disc floret primordia. In mature tubular-like ray florets, corolla and ovary had both ray and disc floret characteristics but also displayed distinct identity traits. In open-pollinated tubular-like ray florets, the seed set was low but a noteworthy increase of filled achenes was obtained by hand pollination. Wild type ray achenes were always empty. Embryos of tubular-like ray florets were shorter and lighter than the embryos of disc florets but able to produce fertile plants. In conclusion, the different identity characteristics combined in tubular-like ray florets of the mutant evolved a capitulum type not described in the genus Helianthus.