An Epiallele at cly1 Affects the Expression of Floret Closing (Cleistogamy) in Barley

The swelling of the lodicule is responsible for floret opening in many grass species, allowing for pollen dispersal and cross-pollination. In barley, the closed floret habit (cleistogamy) is under the control of cly1, a gene that operates by inhibiting the development of the lodicule. In non-cleistogamous cultivars, cly1 mRNA is degraded by miR172-directed cleavage, allowing the lodicules to swell; however, in cultivars carrying the recessive allele cly1.b, a single-nucleotide substitution destroys the miR172 target site preventing mRNA cleavage. Barley cv. SV235 is cleistogamous; its cly1 coding sequence is identical to that of cly1.b, but its lodicules do develop, although insufficiently to produce a non-cleistogamous flower. In this cultivar, the downregulation of cly1 is unrelated to miR172-directed mRNA degradation, but rather is caused by an epiallele that represses transcription. Allelic relationships between known cly1 alleles were explored by the quantification of lodicule vascularization and an assessment of the response of the spike to the supply of exogenous auxin. The SV235 phenotype can be manipulated by a pre-anthesis application of 2,4-D, a feature that could be of interest in the context of hybrid barley grain production based on cleistogamy.