Triacylglycerol assembly

Bafor, M., Smith, M. A., Jonsson, L., Stobart, K., and Stymne, S. 1991. Ricinoleic acid biosynthesis and triacylglycerol assembly in microsomal preparation from developing castor bean (Ricinus communis) endosperm. Biochem. J., 280, 507-514. 

Stymne, S., Bafor, M., Jonsson, L., Wiberg, E. and Stobart, A.K. (1990) Triacylglycerol assembly, in Plant Lipid Biochemistry, Structure and Utilization, eds. P.J. Quinn and J.L. Harwood, Portland, London, pp. 191-197. 

The proportions of triacylglycerols in plant cell cultures are generally low. There are a few exceptions to this rule, as given in Table 1, indicating that induction of fatty acid biosynthesis and triacylglycerol assembly is possible in cultured plant cells. In some cases, accumulation of these lipids may be explained by induction of cell differentiation, e.g. embryogenesis (cf. Sect. 4) or by senescence-related lipid changes [33]. 

FIGURE 24.3 (a) A duct at the junction of the pancreas and duodenum secretes pancreatic juice into the duodenum, the first portion of the small intestine, (b) Hydrolysis of triacylglycerols by pancreatic and intestinal lipases. Pancreatic lipases cleave fatty acids at the C-1 and C-3 positions. Resulting monoacylglycerols with fatty acids at C-2 are hydrolyzed by intestinal lipases. Fatty acids and monoacylglycerols are absorbed through the intestinal wall and assembled into lipoprotein aggregates termed chylomicrons (discussed in Chapter 25). 

HDL and VLDL are assembled primarily in the endoplasmic reticulum of the liver (with smaller amounts produced in the intestine), whereas chylomicrons form in the intestine. LDL is not synthesized directly, but is made from VLDL. LDL appears to be the major circulatory complex for cholesterol and cholesterol esters. The primary task of chylomicrons is to transport triacylglycerols. Despite all this, it is extremely important to note that each of these lipoprotein classes contains some of each type of lipid. The relative amounts of HDL and LDL are important in the disposition of cholesterol in the body and in the development of arterial plaques (Figure 25.36). The structures of the various... 

We first describe the biosynthesis of fatty acids, the primary components of both triacylglycerols and phospholipids, then examine the assembly of fatty acids into triacylglycerols and the simpler membrane phospholipids. Finally, we consider the synthesis of cholesterol, a component of some membranes and the precursor of steroids such as the bile acids, sex hormones, and adrenocortical hormones. 

Correct answer = A. Pancreatic lipase hydrolyzes dietary triacylglycerol primarily to 2-monoacylglycerol plus two fatty acids. These products of hydrolysis can be absorbed by the intestinal mucosal cells. Bile salts do not inhibit release of fatty acids from triacylglycerol, but rather are necessary for the proper solubilization and hydrolysis of dietary triacylglycerol in the small intestine. Short- and medium-chain length fatty acids enter the portal circulation after absorption from the small intestine. Synthesis of apolipoproteins, especially apo B-48, is essential for the assembly and secretion of chylomicrons. 

Chylomicrons are assembled in intestinal mucosal cells and cany dietary triacylglycerol, cholesterol, fat-soluble vitamins, and cholesteryl esters (plus additional lipids made in these cells) to the peripheral tissues (Figure 18.16). 

Assembly of chylomicrons The enzymes involved in triacylglycerol, cholesterol, and phospholipid synthesis are located in Ihe smooth ER. Assembly of the apolipoproteins and lipid into chylomicrons requires microsomal triacylglycerol transfer protein (see p. 229), which loads apo B-48 with lipid. This occurs during transition from the ER to the Golgi, where the particles are packaged in secretory vesicles. These fuse with the plasma membrane releasing the lipoproteins, which then enter the lymphatic system and, ultimately, the blood. 

The plasma lipoproteins include chylomicrons, very-low-density lipoproteins (VLDL), low-density lipoproteins (LDL), and high-density lipoproteins (HDL). They function to keep lipids (primarily triacylglyc-erol and cholesteryl esters) soluble as they transport them between tissues. Lipoproteins are composed of a neutral lipid core (containing triacylglycerol, cholesteryl esters, or both) surrounded by a shell of amphipathic apolipoproteins, phospholipid, and nonesterified cholesterol. Chylomicrons are assembled in intestinal mucosal cells from dietary lipids (primarily, triacylglycerol) plus additional lipids synthesized in these cells. Each nascent chylomicron particle has one molecule of apolipoprotein B-48 (apo B-48). They are released from the cells into the lymphatic system and travel to the blood, where they receive apo C-ll and apo E from HDLs, thus making the chylomicrons functional. Apo C-ll activates lipoprotein lipase, which degrades the... 

Chylomicrons are assembled in intestinal mucosal cells from dietary lipids (primarily triacylglycerol), plus additional lipids synthesized in these cells. Each nascent chylomicron particle has one molecule of apolipoprotein B-48 (apo B-48). They are released from the cells into the lymphatic system and travel to the blood, where they receive apo C-ll and apo E from HDLs. This makes the chylomicrons functional. 

Postulated scheme for the synthesis, assembly, and secretion of VLDL by a hepatocyte (liver cell). (1) Synthesis The apoproteins, phospholipid, triacylglycerol, cholesterol, and cholesteryl esters are synthesized in the endoplasmic reticulum. (2) Assembly These components are assembled into a prelipoprotein particle in the lumen of the endoplasmic reticulum. (3) Processing The particle moves to the Golgi apparatus, where modification of the apoproteins occurs. 

This chapter examines the biosynthesis of three important components of biological membranes—phospholipids, sphingolipids, and cholesterol (Chapter 12). Triacylglycerols also are considered here because the pathway for their synthesis overlaps that of phospholipids. Cholesterol is of interest both as a membrane component and as a precursor of many signal molecules, including the steroid hormones progesterone, testosterone, estrogen, and cortisol. The biosynthesis of cholesterol exemplifies a fundamental mechanism for the assembly of extended carbon skeletons from five-carbon units. 

The fatty acids and monoacylglycerol are absorbed by the intestinal cells, converted to fatty acyl CoA and reassembled into triacylglycerols. The triacyl glycerols then assemble with phospholipids and lipoproteins to form chylomicrons for transport through the lymph and blood to the tissues. 

Saturated fatty acids containing up to 16 carbon atoms (palmitate) are assembled in cytoplasm from acetyl-CoA. Depending on cellular conditions, the product of this process (palmitoyl-CoA) can be used directly in the synthesis of several types of lipid (e.g., triacylglycerol or phospholipids), or it can enter the mito-... 

The source of these may be the neutral lipid bodies (NLBs) containing triacylglycerol (TAG) and DAG, which are associated with the FVs (Jack-son et al., 2004). The scenario proposed is the double-membrane vesicle produced during cytostomal feeding (see p. 171) has its outer membrane fused with the FV membrane while its inner membrane is degraded by phospholipase C and/or lysosomal acid lipase (and found in the falciparum genome) the breakdown products are assembled into TAG and its precursors serve as promoters of p-hematin formation. 

The principal functions of the lipoprotein classes are determined by their apolipoprotein (apo) and lipid components. The CM are synthesized in the intestines for the transport of dietary triacylglycerols to various tissues (Chapter 19). VLDL are synthesized in the liver for the export of endogenous triacylglycerols (Chapter 19), while LDL arise from the metabolic transformation of VLDL in circulation (Chapter 20). The function of LDL is to deliver CE to peripheral tissues and to the liver. HDL are synthesized and assembled in the liver and intestine or are formed from metabolic transformations of other lipoproteins in circulation, and from cellular lipids at the cell membranes (see Chapter 20). HDL remove excess cholesterol from cells and transport it to liver and steroidogenic tissues for metabolism and excretion. 

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