Rice CYP703A3, a cytochrome P450 hydroxylase, is essential for development of anther cuticle and pollen exine

Anther cuticle and pollen exine act as protective envelopes for the male gametophyte or pollen grain, but the mechanism underlying the synthesis of these lipidic polymers remains unclear. Previously, a tapetum‐expressed CYP703A3, a putative cytochrome P450 fatty acid hydroxylase, was shown to be essential for male fertility in rice（Oryza sativa L.）. However,the biochemical and biological roles of CYP703A3 has not been characterized. Here, we observed that cyp703a3‐2 caused by one base insertion in CYP703A3 displays defective pollen exine and anther epicuticular layer, which differs from Arabidopsis cyp703a2 in which only defective pollen exine occurs. Consistently, chemical composition assay showed that levels of cutin monomers and wax components were dramatically reduced in cyp703a3‐2 anthers. Unlike the wide range of substrates of Arabidopsis CYP703A2, CYP703A3 functions as an in‐chain hydroxylase only for a specific substrate, lauric acid, preferably generating 7‐hydroxylated lauric acid. Moreover, chromatin immunoprecipitation and expression analyses revealed that the expression of CYP703A3 is directly regulated by Tapetum Degeneration Retardation, a known regulator of tapetum PCD and pollen exine formation. Collectively, our results suggest that CYP703A3 represents a conserved and diversified biochemical pathway for in‐chain hydroxylation of lauric acid required for the development of male organ in higher plants.

Lipid Metabolism: Critical Roles in Male Fertility and Other Aspects of Reproductive Development in Plants

Fatty acids and their derivatives are essential building blocks for anther cuticle and pollen wall formation.Disruption of lipid metabolism during anther and pollen development often leads to genic male sterility(GMS).To date,many lipid metabolism-related GMS genes that are involved in the formation of anther cuticle,pollen wall,and subcellular organelle membranes in anther wall layers have been identified and characterized.In this review,we summarize recent progress on characterizing lipid metabolism-related genes and their roles in male fertility and other aspects of reproductive development in plants.On the basis of cloned GMS genes controlling biosynthesis and transport of anther cutin,wax,sporopollenin,and tryphine\r\Arabidopsis,rice,and maize as well as other plant species,updated lipid metabolic networks underlying anther cuticle development and pollen wall formation were proposed.Through bioinformatics analysis of anther RNA-sequencing datasets from three maize inbred lines(Oh43,W23,and B73),a total of 125 novel lipid metabolism-related genes putatively involved in male fertility in maize were deduced.More,we discuss the pathways regulating lipid metabolism-related GMS genes at the transcriptional and post-transcriptional levels.Finally,we highlight recent findings on lipid metabolism-related genes and their roles in other aspects of plant reproductive development.A comprehensive understanding of lipid metabolism,genes involved,and their roles in plant reproductive development will facilitate the application of lipid metabolism-related genes in gene editing,haploid and callus induction,molecular breeding and hybrid seed production in crops.