The very high proportlons of trlenolc fatty acids found in chloroplast membranes of all higher plants suggest that these llpld otructurtis mlght be essentlal for photosynthasis. We report here on the productisn of Arabidopsis triple mutants that contaln negllglble levels of trlenolc fatty acids. Photosynthesls at 22OC was barely affected, and vegetative growth of the mutants was ldentlcal wlth that of tha wlld type, dm"StratIng that any requlrement for trienoic acyl groups in membrane structure and functlon 1s relatively subtle. Although vegetative growth and development were unaffected, the triple mutants are male sterlle and produce no seed under normal conditlons. Comparisons of psllen development In wlld-type and triple mutant flowersestablished that pallan gralns in the mutant develeped to the tslcellular stege. Exogenous applicatlens of a-llnolenata or jasmonate restered fertlllty. T a k n tsgether, the results demonstrate that the critical role of trlenolc acids ln tha llfe cycle of plants 1s as the precursor of sxylipln, a signallng compound that regulates final maturatlon processes and the release of pollen.
INTRODUCTIONThe biophysical reactions of light harvesting and electron transport during photosynthesis take place in a uniquely constructed bilayer membrane, the thylakoid. In all photosynthetic eukaryotes, the complement of atypical glyceralipid molecules that form the foundation of this membrane are characterized by sugar headgroups and avery high leve1 of unsaturation in the fatty acid chains, which compose the central portion of the thylakoid lamella bilayer. For example, menogalactosyldiacylglycerol, the major thylakoid lipid, typically contains >90% af a-linolenic acid (183) er a combination of 183 and hexadecatrienoic (163) acids, depending on the plant species (Jamieson and Reid, 1971). These very high levels of trienoic fatty asids are noteworthy because free radicals that are byproducts of the photosynthetic llght reactions stimulate oxidation of polyunsaturated fatty acids. Because this oxidation might be expected ta medlate against a high degree of unsaturation, it has been inferred that there is a strong selective advantage to having such high levels of trienoic fatty acids in the thylakoid. Therefore, it could be reasoned that these lipid structures must have some critical role in maintaining photosynthetic functlon.To test this line of reasoning directly, we set out to develop Arabldopsis llnes with reduced levels of 18:3 and 163 fatty acids. There are two dlstinct pathways in plant 6811s for the biosynthesis of glycerolipids Rnd the associated praduction To whom correspondence should be addressed. of polyunsaturated fatty acids (Roughan et al., 1980;Browse and Somerville, 1991). 60th pathways are initiated by the synthesis of 16:O-acyl carrier protein (ACP) and 18:l-ACP by the combined action of a type II fatty acid synthase (Shimakata and Stumpf, 1982) and a soluble stearoyl-ACP desaturase (McKeon and Stumpf, 1982;Shanklin and Somerville, 1991) located in the chloroplasts or other p...