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              Lipoprotein/Cholesterol Metabolism                                                          653

                       TABLE III Properties of the Apolipoproteins
                                    Molecular  Concentration
                       Apolipoprotein  weight (Da)  (mg/dl)    Tissue origin          Function
                       Apo-A1         28,300     90–130   Intestine, liver    LCAT activator
                       Apo-A2         17,000     30–50    Intestine, liver    Unknown
                       Apo-A4         46,000     10–30    Intestine, liver    Unknown
                       Apo-B100      516,000     80–100   Liver               Cholesterol, triglyceride transport,
                                                                               receptor recognition
                       Apo-B48       264,000      —       Intestine           Triglyceride, cholesterol transport
                       Apo-C1          6,500      4–7     Liver               LCAT activator
                       Apo-C2          8,800      3–8     Liver               LPL activator
                       Apo-C3          8,750      8–15    Liver               LPL inhibitor
                       Apo-D          33,000      10      Liver, intestine, pancreas,  Unknown
                                                            kidney, adrenals, brain
                       Apo-E          35,000      3–6     Liver, macrophages, brain,  Receptor recognition
                                                            Adrenal
                       CETP           74,000      —       Spleen, liver, small  Cholesterol ester transfer
                                                            intestine, adrenal
                       Lp(a)         400–700     0.4–80   Liver, testes, brain  Unknown



              X. OTHER LIPOPROTEIN RECEPTORS                    B. Cubulin
                                                                Cubulin is a 460-kDa protein localized to epithelial cells,
              A. LRP/α 2 -Macroglobulin Receptor
                                                                such as those of the kidney. It plays a role in albumin
              The lipoprotein receptor-related protein (LRP)/α 2 -  reabsorption; its absence leads to loss of albumin in the
              macroglobulin receptor (LRP) has a mass about five times  urine. Cubulin also mediates the uptake of the major HDL
              larger than the LDL receptor. It contains many of the  protein apo-A1 by the kidney.
              same structural domains as the LDL receptor, including a
              cysteine-rich repeat domain that binds to lipoproteins. In
                                                                C. Megalin (gp330)
              vitro studies have shown that the LRP does not bind to apo-
              B100, but binds to VLDL particles that have been enriched  This receptor is a member of the LDL receptor superfam-
              in apo-E. The binding of LRP to its ligands is dependent  ily. It is the antigen that elicits an autoimmune response
              upon cell surface proteoglycans. These are proteins con-  leading to a disease called Heymann’s glomerulonephritis.
              taining sulfated carbohydrate residues.           It interacts with cubulin and is thought to function together
                In addition to binding lipoproteins, the LRP also binds  with cubulin in the uptake of apo-A1 by the kidney.
              to α 2 -macroglobulin, a plasma protein that binds to pro-
              teases and, upon binding, is cleared by the liver. The LRP
              also binds to plasminogen activators and their inhibitors,  D. A VLDL Receptor (?)
              such as tissue-type plasminogen activator, urokinase-
                                                                A cDNA encoding another member of the LDL receptor
              type plasminogen activator, and their corresponding
                                                                family is expressed in muscle and adipose tissue, but not in
              inhibitors.
                                                                liver. It binds to apo-E-containing lipoproteins in vitro,but
                                                                not to LDL. Based on its tissue localization and binding
                                                                properties, this protein has been postulated to be a receptor
                  TABLE IV Apolipoproteins of the Plasma        for triglyceride-rich lipoproteins.
                  Lipoproteins
                  Chylomicrons  A-1, A-2, A-4, B-48, C-1, C-2, C-3, E
                                                                E. CD-36
                  VLDL       B-100, C-1, C-2, C-3, E, trace amounts
                               of A-1, A-2, and A-4
                                                                CD-36 is expressed in a variety of tissues, including
                  LDL        90–98% apo B-100
                                                                adipocytes and macrophages. It functions in fatty acid
                  HDL        A-1, A-2, sometimes, E
                                                                transport and, like scavenger receptors, binds a variety
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