The fabl mutant of Arabidopsis tbaliana, which contains increased levels of saturated fatty acids, was indistinguishable from the wild type when it was grown at 22 or 12°C. During the first 7 to 10 d after transfer to 2"C, the growth and photosynthetic characteristics of the fabl plants remained indistinguishable from the wild type, with values for the potential quantum efficiency of photosystem II decreasing from 0.8 to 0.7 in plants of both lines. Whereas wild-type plants maintained quantum efficiency of photosystem II at approximately 0.7 for at least 35 d at 2"C, this parameter declined rapidly in the mutant after 7 d and reached a value of less than 0.1 after 28 d at 2°C. This decline in photosynthetic capacity was accompanied by reductions in chlorophyll content and the amount of chloroplast glycerolipids per gram of leaf. Electron microscopic examination of leaf samples revealed a rapid and extensive disruption of the thylakoid and chloroplast structure in the mutant, which is interpreted here as a form of selective autophagy. Despite the almost complete loss of photosynthetic function and the destruction of photosynthetic machinery, fabl plants retained a substantial capacity for recovery following transfer to 22°C. These results provide a further demonstration of the importance of chloroplast membrane unsaturation to the proper growth and development of plants at low temperature.A remarkable feature of the chloroplast membranes of higher plants is the high number of double bonds that are found in the lipid acyl chains. Typically, only about 10% of the fatty acids that compose the hydrophobic mid-portion of the thylakoid bilayer lack double bonds altogether, whereas more than 80% have two or more double bonds (Harwood, 1982). The relevance of thylakoid lipid composition to the correct functioning of these membranes in photosynthesis has been the subject of considerable speculation and experimentation, but it is still not well understood (Quinn et al., 1989;Murata and Wada, 1995).
347To investigate the functional significance of chloroplast lipid composition, we have isolated a series of Arabidopsis mutants with specific alterations in leaf lipid composition (Browse and Somerville, 1991;Somerville and Browse, 1991). Five genetic loci have been identified that encode the genes that are involved in lipid-linked fatty acid desaturation of chloroplast lipids (fad4, fad5, fad6, fad7, and fad8).(The first four of these were previously known as f a d A , fadB, fadC, and fadD, respectively.) The actl mutants are deficient in the chloroplast glycerol-3-phosphate acyltransferase. Each of these mutants exhibits substantial changes in chloroplast fatty acid composition. For example, thefad5 mutant lacks cis-unsaturated 16-carbon fatty acids because of a deficiency in the desaturation of 16:O on MGD (Kunst et al., 1989a), whereas the fad6 mutant has increased levels of monoenoic fatty acids as a result of a mutation in the chloroplast 16:1/ 18:l desaturase (Browse et al., 1989). However, the m u t a n t s do not exh...