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136  Veterinary Histology of Domestic Mammals and Birds



                   lymphoid (lymphocyte) and myeloid (granulocyte/mono-  (erythroblastus polychromatophilicus). Due to the high
       VetBooks.ir  cyte, erythrocyte/megakaryocyte) cell lines.  haemoglobin content, the cytoplasm of polychromat-
                      Progenitor cells are morphologically similar to one  ophilic erythroblasts gradually becomes acidophilic with
                   another, being relatively small and strongly basophilic with  little remaining basophilia (Figure 7.2). With continued
                   a round nucleus.                               maturation and increasing haemoglobin concentration,
                      In vitro stimulation of progenitor cells using cytokines  the proportion of basophilic organelles (ribosomes,
                   or other growth factors results in the formation of clus-  rER) declines, giving rise to the orthochromatophilic
                   ters or colonies of cells. These clonogenic cells are thus  (oxyphilic  or  acidophilic)  erythroblast  (normoblast,
                   referred to as a ‘colony-forming unit’.        erythroblastus acidophilicus) (Figure 7.2).
                      Progenitor cells differentiate in the bone marrow into
                   mature blood cells that are found in the circulation. These   Species variation
                   cells (and their differentiation) are termed:    Mammals: Following an increase in the density and com-
                                                                    paction of the nuclear chromatin (pyknosis), the nucleus
                     ·  erythrocytes (erythropoiesis),              is ejected (enucleation). Loss of the nucleus during
                     ·  granulocytes (granulopoiesis),              erythropoiesis  is  characteristic  of  mammals,  includ-
                     ·  lymphocytes (lymphopoiesis),                ing humans. The remaining organelles (Golgi vesicles,
                     ·  monocytes (monopoiesis) and                 polyribosomes, ER) become clumped together in the
                     ·  platelets (thrombocytes) (thrombopoiesis).
                                                                    cytoplasm. The enucleated cell is referred to as a reticu-
                                                                    locyte (proerythrocyte). Within 24 hours, reticulocytes
                   Red blood cells (erythrocytes)                   lose their internal organelles, becoming the haemoglo-
                   A major function of the blood is the transport of O and CO ,
                                                         2     2    bin-carrying cells known as erythrocytes (Figure 7.2).
                   bound to the pigment haemoglobin. This globular chro-
                   moprotein with a prosthetic haem group is concentrated   Birds, reptiles, amphibians and fish (except cyclostomes):
                   in red blood cells, constituting over 30% of the cytoplasm   Enucleation does not take place. In these species, erythro-
                   (95% of dry weight). The high, reversible binding capacity   cytes are nucleated and larger than those of mammals.
                   of haemoglobin enables erythrocytes to take up oxygen in
                   the lungs and deliver it to the peripheral tissues, and to trans-  The morphological changes occurring during erythro-
                   port carbon dioxide in the opposite direction. The bright  poiesis can be summarised as follows:
                   red colour of arterial blood is derived from its content of
                   oxyhaemoglobin. Venous blood contains haemoglobin in its   ·  reduction in cell volume,
                   reduced state and is thus dark red in colour.    ·  decrease in basophilia,
                                                                    ·  reduction and loss of nucleoli and cellular organelles,
                   Development of red blood cells (erythropoiesis)  ·  alteration of nuclear structure,
                   In the initial stage of erythropoiesis, haemopoietic stem   ·  increase in acidophilia and
                   cells differentiate into precursor cells referred to as pro-  ·  loss of the nucleus (mammals).
                   erythroblasts  (Figure 7.2).  This process is driven by
                   hormonal factors, particularly by erythropoietin, a sialo-  Erythrocytes
                   glycoprotein produced by the kidney in response to low  In most mammals, the red blood cell, or erythrocyte, is a
                   blood oxygen levels (anaemia or hypoxia). Steroid hor-  non-nucleated, biconcave, round disc (Figure 7.3). The size
                   mones, peptides and Vitamin B  also act as regulators of  of erythrocytes varies considerably with species, ranging in
                                            12
                   erythrocyte development.                       diameter from 4.1 μm in the goat to 7.3 μm in the dog (Table
                      Proerythroblasts are basophilic cells measuring 20–25  7.1). In camels and dromedaries, the erythrocyte is oval. The
                   μm in size. They undergo rapid replication, during which the  number of circulating erythrocytes is related to their species-
                   size of the nucleus decreases. Accumulation of haemoglobin  dependent variation in size. The smaller the cells, the greater
                   in the cytoplasm begins in this early stage of differentia-  their unit volume, as seen particularly in the goat.
                   tion. Further mitotic division gives rise to the basophilic   Circulating erythrocyte numbers are further influenced
                   erythroblast (erythroblastus basophilicus) (Figure 7.2). By  by muscular effort, training, condition (horses), housing
                   this stage the nucleus has become noticeably smaller and  (pigs), altitude (e.g. alpine cattle) and sex (greater in males
                   heterochromatic. The cytoplasm is basophilic. Through  than females). Lacking a nucleus, erythrocytes exhibit a
                   endocytosis, basophilic erythroblasts acquire increasing  high degree of plasticity and elasticity and undergo con-
                   amounts of ferritin from surrounding macrophages and  tinuous, reversible modifications of their shape. This
                   reticular cells to support increased haemoglobin synthesis.  facilitates their passage through even the smallest capil-
                      The basophilic erythroblast undergoes further differen-  laries. In hypotonic solution, erythrocytes absorb fluid,
                   tiation to become the polychromatophilic erythroblast  resulting in swelling and possible disintegration. Under









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