The galactolipids, mono-and digalactosyldiacylglycerol (DGDG), are the most common nonphosphorous lipids in the biosphere and account for 80% of the membrane lipids found in green plant tissues. These lipids are major constituents of photosynthetic membranes (thylakoids), and a large body of evidence suggests that galactolipids are associated primarily with plastid membranes in seed plants. A null-mutant of Arabidopsis (dgd1), which lacks the DGDG synthase (DGD1) resulting in a 90% reduction in the amount of DGDG under normal growth conditions, accumulated DGDG after phosphate deprivation up to 60% of the amount present in the wild type. This observation suggests the existence of a DGD1-independent pathway of galactolipid biosynthesis. The fatty acid composition of the newly formed DGDG was distinct, showing an enrichment of 16-carbon fatty acids in the C-1 position of the glycerol backbone of DGDG. Roots with their rudimentary plastids accumulated large amounts of DGDG after phosphate deprivation, suggesting that this galactolipid may be located in extraplastidic membranes. Corroborating evidence for this hypothesis was obtained directly by fractionation of subcellular membranes from leaf tissue and indirectly by lipid analysis of the phosphate-deprived fad3 mutant primarily deficient in extraplastidic fatty acid desaturation. The discovery of extraplastidic DGDG biosynthesis induced by phosphate deprivation has revealed a biochemical mechanism for plants to conserve phosphate. Apparently, plants replace phospholipids with nonphosphorous galactolipids if environmental conditions such as phosphate deprivation require this for survival.O ne of the most powerful environmental stimuli affecting the overall glycerolipid composition of bacterial membranes is phosphate deprivation. The replacement of phospholipids by nonphosphorous lipids was discovered first in nonphotosynthetic bacteria (1) and later in those capable of photosynthesis (2, 3). Mutants of different photosynthetic bacteria lacking the acidic nonphosphorus sulfolipid sulfoquinovosyldiacylglycerol showed impaired growth only under phosphate-limited growth conditions (2, 4). This led to the hypothesis that sulfolipid can replace the acidic thylakoid phospholipid phosphatidylglycerol under these growth conditions (5). An inverse relationship between sulfolipid and phosphatidylglycerol as a function of phosphate availability was also observed for Arabidopsis (6). Experiments with the phosphate-deficient pho1 mutant of Arabidopsis (7) suggested that not only sulfolipid but also the amounts of digalactosyldiacylglycerol are increased after phosphate deprivation (8). Thus, it seemed reasonable to ask whether the substitution of phospholipids by nonphosphorous lipids is a more general phenomenon in plants.The nonphosphorous galactolipids mono-and digalactosyldiacylgycerol (MGDG and DGDG) constitute the bulk of membrane lipids in green tissues of seed plants where they are known to be located in the photosynthetic membranes (thylakoids) of the chloroplasts (9, 1...