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220  Veterinary Histology of Domestic Mammals and Birds



                      Smooth muscle cells (myocyti villi) originating from  gut flora frequently prevents colonisation of the mucosa
       VetBooks.ir  the lamina muscularis mucosae pass through the lamina  by pathogens.
                   propria along the longitudinal axis of the villus. These cells
                                                                     In its entirety, the mucosal barrier protects against
                   undergo rhythmic contraction, resulting in shortening and  absorption of antigens, modifies antigens (antigen han-
                   lengthening of the villus. Through these contractions,  dling) and hinders trans-epithelial uptake of antigens via
                   the cells act as a pumping station that aids in transport  the formation of complexes with secretory immunoglob-
                   of blood and lymph into deeper vessels (Figure 10.50).  ulin A (IgA) (see below).
                   Myofibroblasts lying perpendicular to the smooth muscle   Adaptive mechanisms comprise cellular responses that
                   cells extend to the sides of the villus. These may also play  result in localised production of immunoglobulins. These
                   a role in contraction of the villus.           antibodies, predominantly IgA, are synthesised by plasma
                      The lamina propria contains dense aggregates of  cells in the lamina propria. IgA is taken up by epithelial
                   lymphatic tissue rich in T and B lymphocytes, plasma  cells, coupled with transport peptides and delivered by exo-
                   cells, monocytes, macrophages, mast cells, neutrophils  cytosis from the intestinal glands to the intestinal lumen.
                   and eosinophils. This is a manifestation of the immune   Secretory IgA  is resistant to proteolytic secretions
                   responses that occur continuously within the gastrointes-  and thus has a long duration of action. It prevents bac-
                   tinal mucosa. The lymphatic tissue of the intestinal mucosa  teria from binding to the mucosa, agglutinates bacteria
                   is collectively termed  GALT  (gut-associated lymphoid  and neutralises viruses and toxins. IgA forms the first line
                   tissue)  and  resembles  the  BALT  (bronchus-associated  of immune defence in protecting the surface epithelium
                   lymphoid tissue) of the respiratory tract (see Chapter 8,  against agents of disease. Other immunoglobulins (IgM,
                   ‘Immune system and lymphatic organs’).         occasionally IgG) are of lesser importance in the local
                      In carnivores, and to a limited extent in horses, a stra-  immune response. As the immune system of neonates is
                   tum compactum is present at the base of the lamina  not fully developed, early development of localised immu-
                   propria near the lamina muscularis. This is an extension  nity in the gastrointestinal tract is particularly important.
                   of the stratum compactum of the gastric mucosa, which
                   it resembles morphologically.                  Tela submucosa
                                                                  The structure of the collagen fibre-rich connective tissue
                     Species variation                            of the tela submucosa is typical for the tubular digestive
                     Birds: Particularly in the chicken, numerous mucosa-  tract. In addition to adipose tissue, vessels and neural
                     associated lymphatic follicles may be present near   networks (plexus nervorum submucosus), the submucosa
                     Meckel’s diverticulum, in the distal jejunum and at the   contains solitary lymphatic nodules (noduli lymphatici
                     entry to the caecum. These tonsil-like lymphatic organs   solitarii) and aggregated lymphatic nodules (noduli lym-
                     (Peyer’s patches) are part of the immune system of the   phatici aggregati, Peyer’s patches). These are described in
                     avian gastrointestinal tract, in which discrete lymph   more detail below (see ‘Ileum’). Mucous glands are pres-
                     nodes are absent in most species.            ent in the tela submucosa of the duodenum (glandulae
                                                                  submucosae) (see below).
                   LAMINA MUSCULARIS MUCOSAE
                   The layers of the lamina muscularis mucosae resemble  Tunica muscularis and tunica serosa
                   those of the gastric mucosa. Species variation in muscle  The structure of the tunica muscularis and tunica serosa
                   layer thickness is observed.                   corresponds to the earlier description applying generally to
                                                                  the gastrointestinal tract. Macroscopically, the thickness of
                   INNATE AND ADAPTIVE PROTECTIVE MECHANISMS      the layers varies with species, in some cases to a consider-
                   OF THE INTESTINAL MUCOSA                       able degree (see anatomy textbooks).
                   Throughout the life of the organism, the intestinal
                   mucosa is exposed to a multitude of foreign materials  Distinguishing features of segments
                   and microorganisms (antigens within foodstuffs, viruses,  of the small intestine
                   bacteria, fungi, parasites). To prevent these from enter-
                   ing the body, the tunica mucosa incorporates innate and   DUODENUM
                   adaptive mechanisms that form a mucosal barrier. The  The duodenal mucosa is extensively folded with marked
                   bactericidal effect of mucus and the acid pH of the gastric  plicae circulares. The intestinal villi are uniform, dense and
                   juice have been described above. Other innate phenomena  relatively broad. Abundant intestinal glands are present,
                   include inactivation of enteropathogenic viruses by bile  and the epithelium contains isolated goblet cells.
                   and the bacteriostatic properties of lactoferrin secreted by   A defining histological feature of the duodenum is the
                   the intestinal mucosa. Interferon also contributes to local  presence of glandulae submucosae (Brunner’s glands)
                   immune defence. In addition, competition from normal  (Figures 10.48 and 10.53). Brunner’s glands lie in the tela









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