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272  Veterinary Histology of Domestic Mammals and Birds



                   into the peri-tubular tissue where it is taken up by medul-  epithelium of the collecting duct transforms into a double
       VetBooks.ir  lary vessels (refer to physiology textbooks for more detail).  layer of columnar epithelium in the papillary duct (diame-
                                                                  ter 200–300 μm). Fat droplets may be seen in the collecting
                     Species variation
                                                                  ducts of older animals and goblet cells may be present in
                     Birds: The loops of Henle and associated counter-cur-  horses. At the papilla, the epithelium consists of two layers
                     rent mechanism are less extensively developed in birds,   (three in the pig).
                     thus the capacity to concentrate urine is lower than   The final stages of urine concentration take place
                     in mammals.                                  in the collecting duct system. The collecting ducts are
                                                                  surrounded by the hypertonic environment of the renal
                   Distal straight tubule and distal convoluted   medulla. Water follows the osmotic gradient into the inter-
                   tubule                                         stitium. Only small quantities of urea and electrolytes pass
                   The ascending thin limb continues as the distal straight  into the tubular cells. Recent research has shown that the
                   tubule, also referred to as the thick ascending limb of the  collecting ducts are the main target of aldosterone in regu-
                                                                                    +
                   loop of Henle (Figures 12.8 and 12.18). The convoluted  lating transport of Na  and water.
                   portion is located near its associated glomerulus. Both the   Antidiuretic hormone (ADH), secreted by the poste-
                   straight and convoluted portions have a wide lumen (30–45  rior pituitary gland, increases permeability to water. Lack
                   μm). The epithelium is somewhat lower than that of the  of ADH or epithelial ADH receptors results in excretion of
                   proximal tubule. Instead of a brush border, the apical surface  large volumes of relatively dilute urine (diabetes insipidus).
                   bears a few irregularly spaced, small microvilli. The basal
                   labyrinth is particularly prominent, with numerous mito-  Juxtaglomerular apparatus (complexus
                   chondria occupying the infoldings of the plasmalemma.  juxtaglomerularis)
                      Na  ions (via basally located ion pumps) are transported  The juxtaglomerular apparatus (Figure 12.12) consists of:
                        +
                                                         +
                   into the interstitium in exchange for K , H  or NH . The dis-
                                                  +
                                               +
                                                        4
                   tal tubule thus plays an important role in maintaining water   ·  the macula densa,
                   and electrolyte balance. These mechanisms are hormonally   ·  juxtaglomerular cells and
                                                               +
                                                         +
                   regulated. Aldosterone influences epithelial Na  and K    ·  extraglomerular mesangial cells (Goormaghtigh
                   transport (see also below). Antidiuretic hormone increases   cells, lacis cells).
                   the permeability of the epithelium to water. Calcitonin
                                                               +
                   raises, and parathyroid hormone lowers, excretion of Ca2 .  Located at the vascular pole of the renal corpuscle, the
                                                                  juxtaglomerular apparatus regulates blood pressure and
                   Collecting duct system                         urinary sodium concentration by modifying glomerular
                   The collecting duct system (Figure 12.19) comprises the:  filtration rate and sodium reabsorption (renin–angioten-
                                                                  sin system).
                     ·  arched collecting tubule,
                     ·  straight collecting tubule (collecting duct) and  Macula densa
                     ·  papillary duct.                           The macula densa is a region of up to 40 modified epithe-
                                                                  lial cells located in the portion of the distal straight tubule
                   The collecting duct system begins with the short, arched  that lies adjacent to the vascular pole of the glomerulus,
                   collecting tubule, which opens into the straight collecting  between the afferent and efferent arterioles and the extra-
                   tubule or collecting duct (Figure 12.6). Up to ten nephrons  glomerular mesangial cells (Figure 12.10). Beyond this
                   drain into each collecting duct. In the cortex, the collect-  point, the distal tubule becomes convoluted.
                   ing ducts are grouped together in the medullary rays.   The epithelial cells are columnar with dense cytoplasm.
                   They continue to run parallel to one another within the  Short microvilli are present on the cell surface. Basally
                   medulla. In the inner zone of the medulla, the collecting  there is a relatively poorly developed basal labyrinth and
                   ducts merge to form the papillary ducts, which empty  a continuous basal lamina. The macula densa acts as a
                                                                                              +
                   by a series of openings at the tip of the renal papilla/crest  chemoreceptor, detecting the Na  ion concentration
                                                                                                  +
                   (area cribrosa) into the renal calyces/pelvis (Figure 12.3).  in the tubular fluid. A fall in tubular Na  triggers release
                      In contrast to the nephron, the collecting duct appara-  of renin by the juxtaglomerular cells. This activates the
                   tus constitutes a branched system of tubules in which the  angiotensin system which brings about increased Na  reab-
                                                                                                          +
                   epithelial lining changes gradually from the cortex to the  sorption, with water following by osmosis.
                   papilla. The epithelial cells are predominantly cuboidal in
                   the cortex and increase in height towards the papilla. The  Juxtaglomerular cells
                   cells have a centrally located nucleus, pale cytoplasm and  Juxtaglomerular cells are modified smooth muscle cells
                   clear cell boundaries. The simple cuboidal to columnar  found in the afferent arteriole, near its entry into the









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