Page 152 - Veterinary Histology of Domestic Mammals and Birds, 5th Edition
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                   Blood and haemopoiesis


                   (sanguis et haemocytopoesis)







                   Blood, the essential functional element of the circulatory  Formation of blood cells (haemopoiesis)
                   system, is distributed evenly throughout the body. It serves  The differentiation of blood cells begins during embry-
                   to maintain internal biological equilibrium, without which  onic development, commencing in the wall of the yolk
                   life cannot be sustained.                      sac (megaloblastic phase). The second phase takes place
                      The composition of blood is correspondingly complex.  in the liver and spleen (hepatosplenic phase). In the latter
                   It consists of fluid and cellular components, both of which  stages of foetal development, the red bone marrow (tex-
                   have important functions. Blood has three fundamental  tus haemopoeticus) takes over the role of haemopoiesis
                   roles: transport, homeostasis and immune defence.  (medullary phase). Blood cell formation in the liver and
                      Blood acts as a vehicle for the transport of oxygen and  spleen gradually declines but can be reactivated in the pres-
                   carbon dioxide, nutrients, intermediate and end products  ence of bone marrow disease. For the life of the animal,
                   of metabolism and waste materials. It provides a means  blood cells differentiate from haemopoietic stem cells
                   of distributing heat, and carries endogenous regulatory   (HPCs) and progenitor cells in the bone marrow (Figures
                   molecules (hormones), enzymes and vitamins.    7.1 and 7.2).
                      In conjunction with organs such as the liver, kidney and
                   lung, blood supports homeostasis by regulating tissue  Regulation of blood formation
                   fluid levels, osmotic pressure, ion concentration and pH.  Cytokines play a major role in regulating the differentia-
                      By way of its white blood cell population (granulocytes,  tion of cells during haemopoiesis. These highly potent
                   lymphocytes, monocytes), blood contributes to innate and  soluble signalling chemicals are produced by numerous
                   adaptive immunity. Substances produced by white cells  haemopoietic and non-haemopoietic cells. Cytokines act
                   (e.g. complement, leukotrienes, lymphokines, antibodies)  via surface receptors on target cells. They exhibit auto-
                   circulate in the blood, reaching all tissues of the body.  crine, paracrine and endocrine activity. As intercellular
                      The  ability  of  blood  to  perform  these  functions  is  mediators, they regulate the division, proliferation, dif-
                   dependent upon additional physiological processes that  ferentiation, mobility and activity of haemopoietic cells.
                   maintain a balance between the coagulation of blood, for  Cytokine expression, usually low, increases dramatically in
                   haemostasis, and lysis of blood clots. Platelets are the cellu-  response to tissue damage or infection. Physical strain also
                   lar component of the coagulation mechanism. In addition,  influences the homeostatic activity of cytokines.
                   plasma contains the soluble protein fibrinogen which,   The  various  cytokines  are  classified  into  partially
                   when clotting is activated by damage to a vessel wall, is  overlapping categories. Haemopoietic cytokines include
                   converted to fibrin. The platelets and fibrin, together with  chemokines, growth factors and transcription fac-
                   red blood cells, form a blood clot. The term serum refers  tors (e.g. c-Myb, E2A). Of major importance are the
                   to plasma from which clotting factors have been removed.  colony-stimulating factors  (CSF)  (e.g. granulocyte-
                      In domestic mammals, the  blood  cells  constitute  macrophage-colony-stimulating factor [GM-CSF]) and
                   between 32 and 45% of the blood volume. The blood cells  interleukins (IL). Among the interleukins, IL-3, IL-5 and
                   are comprised of:                              IL-15 have a particular influence on cell differentiation.
                                                                     The later stages of differentiation are stimulated in a
                     ·  red blood cells (erythrocytes),           targeted fashion by specific factors. Thrombopoietin pro-
                     ·  white blood cells (leucocytes):           duced by the liver and kidney regulates the development of
                          − granulocytes,                         platelets. A lack of oxygen or a fall in intravascular erythro-
                          − lymphocytes,                          cyte numbers triggers production of the sialoglycoprotein
                          − monocytes and                         erythropoietin  (EPO)  by interstitial connective tissue
                     ·  platelets (thrombocytes).                 cells located in the peri-tubular region of the nephrons,
                                                                  between the renal cortex and medulla. Erythropoietin









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