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278       Tissues and organs



             Lipoproteins                                     cosa and reach the blood via the lymphatic
                                                              system (see p. 266). In the peripheral vessel-
             Most lipids are barely soluble in water, and     s—particularly in muscle and adipose tis-
             many have amphipathic properties. In the         sue—lipoprotein lipase [1] on the surface of
             blood, free triacylglycerols would coalesce      the vascular endothelia hydrolyzes most of
             into drops that could cause fat embolisms.       the triacylglycerols. Chylomicron breakdown
             By contrast, amphipathic lipids would be de-     is activated by the transfer of apoproteins E
             positedin the bloodcells’membranes and           and C from HDL. While the fatty acids released
             would dissolve them. Special precautions are     and the glycerol are taken up by the cells, the
             therefore needed for lipid transport in the      chylomicrons gradually become converted
             blood. While long-chain fatty acids are bound    into chylomicron remnants,which areulti-
             to albumin and short-chain ones are dissolved    mately removed from the blood by the liver.
             in the plasma (see p. 276), other lipids are
             transported in lipoprotein complexes,of             VLDLs, IDLs,and LDLs are closely related to
             which there several types in the blood           one another. VLDLs formed in the liver (see
             plasma, with different sizes and composition.    p. 312) transport triacylglycerols, cholesterol,
                                                              and phospholipids to other tissues. Like chy-
                                                              lomicrons, they are gradually converted into
             A. Composition of lipoprotein complexes
                                                              IDL and LDL under the influence of lipoprotein
             Lipoproteins are spherical or discoid aggre-     lipase [1]. This process is also stimulated by
             gates of lipids and apoproteins.They consist     HDL. Cells that have a demand for cholesterol
             of a nucleus of apolar lipids (triacylglycerols  bind LDL through an interaction between
             and cholesterol esters) surrounded by a sin-     their LDL receptor and ApoB-100, and then
             gle-layered shell approximately 2 nm thick of    take up the complete particle through recep-
             amphipathic lipids (phospholipids and cho-       tor-mediated endocytosis. This type of trans-
             lesterol; the example shown here is LDL).        port is mediated by depressions in the mem-
             The shell, in which the apoproteins are also     brane (“coated pits”), the interior of which is
             deposited, gives the surfaces of the particles   lined with the protein clathrin. After LDL
             polar properties and thereby prevents them       binding, clathrin promotes invagination of
             from aggregating into large particles. The       the pits and pinching off of vesicles (“coated
             larger the lipid nucleus of a lipoprotein        vesicles”). The clathrin then dissociates off and
             is—i. e., the larger the number of apolar lipids  is reused. After fusion of the vesicle with ly-
             it contains—the lower its density is.            sosomes, the LDL particles are broken down
                Lipoproteins are classified into five groups.  (see p. 234), and cholesterol and other lipids
             In order of decreasing size and increasing       are used by the cells.
             density,  these   are:  chylomicrons,   VLDLs
             (very-low-density lipoproteins), IDLs (inter-       The HDLs also originate in the liver. They
             mediate-density lipoproteins), LDLs (low-        return the excess cholesterol formed in the
             density lipoproteins), and HDLs (high-density    tissues to the liver. While it is being trans-
             lipoproteins). The proportions of apoproteins    ported, cholesterol is acylated by lecithin cho-
             range from 1% in chylomicrons to over 50% in     lesterol acyltransferase (LCAT). The cholesterol
             HDLs. These proteins serve less for solubility   esters formed are no longer amphipathic and
             purposes, but rather function as recognition     can betransported in thecoreof the lipopro-
             molecules for the membrane receptors and         teins. In addition, HDLs promote chylomicron
             enzymes that are involved in lipid exchange.     and VLDL turnover by exchanging lipids and
                                                              apoproteins with them (see above).

             B. Transport functions
             The classes of lipoproteins differ not only in
             their composition, but also in the ways in
             which they originate and function.
                The chylomicrons take care of the transport
             of triacylglycerols from the intestine to the
             tissues. They are formed in the intestinal mu-


           Koolman, Color Atlas of Biochemistry, 2nd edition © 2005 Thieme
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