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Principles of bioactive lipid signalling

Shown are model membranes that represent the plasma membrane, vesicular membranes and intracellular membranes, as well as the mechanism where sphingolipids could be transported between and across them. Sphingomyelin (SM) has two aliphatic chains plus a zwitterionic head group. Therefore, it just occasionally (whenever) flipflops across bilayers and has negligible aqueous solubility, but may perhaps have significant lateral movement. This movement could be hindered by selfaggregation and/or interaction with membrane sterols. Ceramide (Cer) contains two aliphatic chains but includes a neutral headgroup, so exhibits less aqueous solubility than SM, but is easily able to flipflop in model membranes. It's not known if interactions with other lipids or partitioning in microdomains would hinder this flipping. Both SM and Cer require vesicles or specific transfer proteins to maneuver between distinct membranes. CERT functions being a specific ceramide transfer protein that acts inside the endoplasmic reticulum (ER)toGolgi transport of Cer, either in the ER interface or as being a freely movable transfer protein. Sphingosine (Sph) has one aliphatic chain (usually 18 carbons in length) and appears to flipflop freely and transfer among membranes. Sphingosine1phosphate (S1P) has a zwitterionic headgroup (including a charged phosphate), and is also prone to move freely among membranes, but is not likely to flipflop spontaneously. Accordingly, two ABC transporters have already been suggested to possess roles in S1P transport: CFTR (cystic fibrosis transmembrane regulator) is implicated in S1P entry in to the cell, and ABCC1 is implicated in S1P efflux. CDase, ceramidase; SMase, sphingomyelinase; SK, sphingosine kinase; SMS, sphingomyelin synthase; SPPase, sphingosine phosphate phosphatase.

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