Due to its hereditary tractability and increasing prosperity of accessible data,

Due to its hereditary tractability and increasing prosperity of accessible data, the candida is a magic size program of preference for the scholarly research from the genetics, biochemistry, and cell biology of eukaryotic lipid rate of metabolism. of signaling substances (Strahl and Thorner 2007). Therefore, the breakthroughs in candida glycerolipid metabolism talked about with this review content also have tremendous potential to lead essential insights into these essential tasks of lipids and lipid-mediated signaling in eukaryotic cells. Pathways of glycerolipid rate of metabolism Main glycerolipids of are the phospholipids Personal computer, PE, PI, PS (Shape 1), phosphatidylglycerol (PG), and cardiolipin (CL) (Rattray 1975; Henry 1982; Henry and Carman 1989; Paltauf 1992; Wenk and Guan 2006; Ejsing 2009). Small phospholipids consist of intermediates such as for example PA, CDP-diacylglycerol (CDP-DAG), phosphatidylmonomethylethanolamine (PMME), phosphatidyldimethylethanolamine (PDME), the D-3, D-4, and D-5 polyphosphoinositides, and lysophospholipids (Rattray 1975; Oshiro 2003; Strahl and Thorner 2007). Label and diacylglycerol (DAG) will be the main natural glycerolipids. The essential fatty acids that are generally esterified towards the glycerophosphate backbone of candida SU-5402 glycerolipids consist of palmitic acidity (C16:0), palmitoleic acidity (C16:1), stearic acidity (C18:0), and oleic acidity (C18:1) (Rattray 1975; Henry 1982; Martin and Bossie 1989; McDonough 1992; Martin 2007). The pathways for the formation of TAG and phospholipids are shown in Figure 2. The enzymes and transporters of glycerolipid metabolism and the genes that encode them are listed in Tables ?Tables11C3. The geneCprotein relationships shown in the tables have been confirmed by the analysis of gene mutations and/or by the biochemical characterization of the enzymes and transporters (Carman and Henry 1989; Greenberg and Lopes 1996; Henry and Patton-Vogt 1998; Carman and Henry 1999; Black and Dirusso 2007; Tehlivets 2007; Kohlwein 2010b; Carman and Han 2011). Synthesis SU-5402 SU-5402 and turnover of phospholipids In the pathways (Figure 2, Table 1), all membrane phospholipids are synthesized from PA, which is derived from glycerol-3-P via lysoPA by two fatty acyl CoA-dependent reactions that are catalyzed in the endoplasmic reticulum (ER) by the 1999b; Zheng and Zou 2001; Benghezal 2007; Rabbit Polyclonal to SFRS7. Chen 2007b; Jain 2007; Riekhof 2007b). The glycerol-3-P acyltransferase enzymes also utilize dihydroxyacetone-P as a substrate, and the product acyl dihydroxyacetone-P is converted to lysoPA by the lipid droplet (LD) and ER-associated 1996) or to DAG, catalyzed by 2006) (Figure 1). CDP-DAG synthase activity has been detected in the ER and in mitochondria (Kuchler 1986), whereas PA phosphatase is a cytosolic enzyme that must associate with membranes to catalyze the dephosphorylation of PA to produce DAG (Han 2006; Carman and Han SU-5402 2009a). CDP-DAG and DAG are used to synthesize PE and PC by two alternative routes, namely, the CDP-DAG and Kennedy pathways (Figure 2). In the CDP-DAG pathway, CDP-DAG is converted to SU-5402 PS by the ER localized 1980; Letts 1983; Bae-Lee and Carman 1984; Kiyono 1987; Nikawa 1987b). Yeast has two PS decarboxylases encoded, by the and genes. Psd1, localized to the inner mitochondrial membrane, accounts for the majority of the enzymatic activity in yeast, while the minor activity, Psd2, associates with Golgi/vacuole (Clancey 1993; Trotter 1993, 1995; Voelker 2003). PE then undergoes three sequential methylation reactions in the ER (Gaynor and Carman 1990), the first of which is catalyzed by the 1988; Kodaki and Yamashita 1989; McGraw and Henry 1989). The CDP-DAG pathway is the major route for synthesis of PE and PC when wild-type cells are grown in the absence of ethanolamine and choline, and mutants defective in this pathway possess choline/ethanolamine auxotrophy phenotypes (Atkinson 1980a Summers 1988; Henry and McGraw 1989; Voelker and Trotter 1995; Trotter 1995). Personal computer and PE synthesis in mutants defective in the CDP-DAG pathway may also be.