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Urinary system (organa urinaria)   265



                  The  capillary  tuft of the glomerulus (rete capillare
       VetBooks.ir  glomerulare) develops from the division of the afferent
                  arteriole into four to eight branches. These branches form
                  up to 50 anastomosing capillary loops. The structural com-
                  ponents of the glomerulus (Figure 12.9) comprise:


                   ·  endothelial cells of the capillary wall,
                   ·  glomerular basement membrane,
                   ·  podocytes (= inner visceral layer of Bowman’s cap-
                      sule) and
                   ·  intraglomerular mesangial cells with loose connec-
                      tive tissue.

                  ENDOTHELIAL CELLS
                  The wall of the glomerular capillaries is composed of an
                  extremely thin endothelium with numerous round pores.
                  These openings are generally evenly spaced and lack a   12.14  Fine structure of a glomerular capillary, base-
                                                                 ment membrane and adjacent podocyte processes with
                  diaphragm. Substances that can pass through the endo-  slit diaphragms (‘blood–urine barrier’) (dog; x18,000).
                  thelium include water, sodium, urea and small proteins.
                  The endothelial surface contains negatively charged pro-  Bowman’s capsule
                  teoglycans that limit the filtration of large anionic proteins.
                  The endothelium also synthesises endothelin. Release of   VISCERAL LAYER (PODOCYTES, PODOCYTI)
                  endothelin results in vasoconstriction of the afferent and   The flattened cells of the visceral layer of Bowman’s
                  efferent arterioles of the glomerulus.         capsule (podocytes) cover the external surface of the glo-
                                                                 merular capillaries (Figures 12.15 to 12.17). Their nuclei
                  BASEMENT MEMBRANE (MEMBRANA BASALIS)           and cell bodies bulge into the urinary space. Extending
                  The basement membrane (thickness 0.1–0.2  μm) lies   from the cell bodies are long, actin-rich processes (primary
                  against the exterior of the capillary wall, where it acts as a   processes) that divide to form secondary and tertiary pro-
                  physical boundary and filter (Figure 12.14). The membrane   cesses (Figures 12.14 to 12.16). At the points of contact
                  consists of three layers:                      with the basement membrane, the tertiary processes
                                                                 expand to form foot processes (the origin of the term
                   ·  inner lamina rara interna (adjacent to endothelium),  ‘podocyte’). Their surfaces bear negatively charged mem-
                   ·  middle lamina densa and                    brane proteoglycans.
                   ·  outer lamina rara externa (adjacent to podocytes).  The finger-like processes are tightly interdigitated leav-
                                                                 ing narrow spaces (25 nm) between them. These spaces are
                  The basement membrane is composed of fine fibrillar   spanned by a thin slit diaphragm (6 nm thick) that is simi-
                  non-aggregated type IV collagen macromolecules as well   lar in structure to the diaphragm of fenestrated capillaries
                  as collagenous and non-collagenous glycoprotein subunits   (Figure 12.14). The slit diaphragm, containing P-cadherin
                  (laminin and fibronectin). These are embedded in a prote-  and nephrin, forms a further filtration barrier. Disruption
                  oglycan-rich matrix of sialic acids.           of this barrier leads to substantial proteinuria and oedema.
                     The basement  membrane is produced mainly by   Podocytes have several functions, including:
                  the capillary endothelium and partly by the podocytes.
                  Mesangial cells serve to remove and phagocytose sub-  ·  contribution to the glomerular filtration barrier,
                  stances filtered by the membrane.                ·  uptake of high-molecular weight molecules,
                     The lamina densa acts as a mechanical filter. The less   ·  regulation of pressure within the renal corpuscle
                  electron-dense lamina interna and externa are rich in   through contractile elements of the cytoskeleton
                  polyanionic molecules. Thus, the basement membrane   and
                  serves as a further barrier to proteins based on their   ·  production of fibrillar components of the glomeru-
                  negative charge. Neutral or positively charged molecules   lar basement membrane.
                  transit more readily than, for example, negatively charged
                  albumin.                                       PARIETAL LAYER AND URINARY SPACE
                     Renal disease, with associated loss of electric charge   (LUMEN CAPSULAE)
                  of the basement membrane, can result in significant   At the vascular pole (polus vascularis), the visceral layer
                  albuminuria.                                   of Bowman’s capsule reflects upon itself to continue as the









       Vet Histology.indb   265                                                                                  16/07/2019   15:03
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