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Macronutrients         99


                  portal circulation for metabolism by the liver.
        VetBooks.ir  Hepatic Handling of Dietary Lipid

                  The liver determines the fate of dietary lipid under the direc-
                  tion of hormonal signals related to energy balance.The primary
                  fuel source for hepatocytes is provided via  β-oxidation of
                  NEFA whether dietary or endogenous in origin (Figure 5-8).
                  The fate of dietary lipid from chylomicron origin can be traced
                  as follows (Figure 5-19) (Brody, 1994b):
                    • Chylomicron remnants attach to receptors on liver cells.
                    • Triglycerides are hydrolyzed to glycerol and CoA esterified
                                                                      Figure 5-17. Triglycerides are the main storage forms of fatty
                     fatty acids.
                                                                      acids, each molecule of which is composed of a three-carbon glyc-
                    • CoA esterified fatty acids may be either shunted to β-oxi-
                                                                      erol nucleus and three fatty acids (R , R , R ).
                                                                                                   2
                                                                                                      3
                                                                                                1
                     dation in mitochondria or repackaged into triglycerides and
                     then VLDL for use by peripheral tissues (storage or fuel).
                     Their fate depends on the energy status of the animal.
                    • Glycerol is converted to 3-P-glycerol (only in liver) and
                     enters the carbohydrate metabolic pathway.
                    Although fat storage is easily accomplished by de novo syn-
                  thesis (production of fat from carbohydrate or protein), it is
                  more energy efficient for animals to deposit dietary fat than to
                  synthesize it. When fat is deposited from foods, the fatty acid
                  profile tends to reflect the type of fat consumed. When fat is
                  synthesized, stored fat composition reflects the fat synthetic
                  enzyme activity of the animal.

                  Lipoprotein Metabolism
                  Lipoproteins are relatively large conglomerates of protein and
                  lipid that are necessary to transport hydrophobic lipids effective-
                  ly through the aqueous medium of physiologic solutions. Table
                  5-20 gives a generic relative composition of the different
                  lipoprotein classes observed in mammals (Chapter 28). As the
                  protein component increases, the relative density of the lipopro-
                  tein increases reflecting dilution of the buoyant density of fat.
                  Lipoproteins are made only in the liver (VLDL, HDL) and
                  enterocytes (chylomicrons). The protein fraction, before it is
                  integrated with the lipid component, is termed apolipoprotein.
                    Lipoprotein metabolism is very complex with species varia-
                  tion. However, a general introduction is necessary to under-
                                                                      Figure 5-18. Diagram of metabolic pathways (elongation and
                  stand lipoprotein metabolic disorders discussed in later chap-
                                                                      desaturation) for essential fatty acids.
                    Table 5-19. Essentiality and biologic function of common fatty acids.
                    Structure  Common name     Essential    Biologic function
                    14:0      Myristic         No           Energy use and storage, acylation of proteins
                    16:0      Palmitic         No           Energy use and storage, acylation of proteins
                    16:1      Palmitoleic      No           Energy use and storage
                    18:0      Stearic          No           Energy use and storage, membrane fluidity
                    18:1n-9   Oleic            No           Energy use and storage, phospholipid structure
                    18:2n-6   Linoleic         Yes          Energy use and storage, arachidonic acid precursor (20:4n-6)
                    18:3n-3   Alpha-linolenic  Yes          Energy use and storage, eicosapentaenoic acid precursor (20:5n-3)
                    18:3n-6   Gamma-linolenic  Yes          Energy use and storage, arachidonic acid precursor
                    20:4n-6   Arachidonic      Yes (cat)    Energy use and storage, synthesis of cytokines and eicosanoids, synthesis of
                                               No (dog)     steroid hormones, membrane fluidity, competitor of eicosapentaenoic acid
                                                              (20:5n-3)
                    20:5n-3   Eicosapentaenoic  Probably    Energy use and storage, synthesis of cytokines and eicosanoids, retinal and
                                                              nervous tissue development, membrane fluidity, competitor of arachidonic
                                                              acid (20:4n-6)
                    22:6n-3   Docosahexaenoic  Probably     Energy use and storage, retinal and nervous tissue development, membrane
                                                              fluidity, competitor of omega-6 fatty acids
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