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ceramide phosphorylethanolamine waxy lipid sphingosine fatty acid sphingomyelin cellular membrane sphingolipid cpe invertebrates biosynthetic mechanism rendering surfactant pathway substrate enzyme phosphatidylserine biosynthesis bilayer synthesis derivative choline glycerophospholipid contractor cholesterol bioprecursor lecithin cell cycle phosphocholine biogenesis precursor endogenous path phospholipid intermediate metabolism lysophosphatidylcholine phosphatidylethanolamine lipophilic