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          were isolated from bone marrow and
          maintained in cell culture. These studies
  VetBooks.ir  demonstrated that in the right conditions,

          certain individual cells would form in vitro
          colonies that contained immature and
          mature cells of a specific cell line. The most
          undifferentiated cells that could form the
          cells of a specific lineage were termed col­
          ony‐forming units (CFUs). More current
          literature uses the stem cell terminology;
          for example, a myeloid stem cell is equiva-
          lent to a colony‐forming unit–granulo-  Figure 15-2.  Red blood cells as seen by scanning
          cytic,  erythrocytic,  monocytic,  and  electron microscopy.  Source: Cohen and Wood,
          megakaryocytic (CFU‐GEMM).              2000. Reproduced with permission of Lippincott
            Obviously, the proliferation and differ-  Williams & Wilkins.
          entiation of bone marrow stem cells must
          be highly regulated. For example, the gen-  exchange across the cell membrane.
          eration of erythrocytes should increase in   Erythrocytes have no nuclei and few
          response to blood loss after hemorrhage,   organelles (see Fig. 1‐9). Total erythrocyte
          while the generation of leukocytes should   counts are expressed as number of cells per
          increase in response to an infection. A   microliter  of  whole  blood,  and  most
          large variety of circulating chemical mes-  domestic animals have about 7 million per
          sengers have been found to regulate the   microliter (Table  15‐2). (Total leukocyte
          proliferation and differentiation of bone   and platelet counts are also expressed per
          marrow  stem cells. The general  term for   microliter of blood.)
          such agents is hematopoietin. An individ-  The protein  hemoglobin is the major
          ual hematopoietin may stimulate commit-  intracellular constituent of erythrocytes.
          ted cells within development pathways to   Hemoglobin is a complex molecule con-
          give rise to specific blood cells or may have   taining four amino acid chains (globin
          a  more  general  effect  by  stimulating  less   portion) held together by noncovalent
          committed stem cells. For example, eryth­  interactions. Each amino acid chain con-
          ropoietin is the hematopoietin that stimu-  tains a heme group (red porphyrin pigment),
          lates a specific increase in erythrocyte   and each heme group contains an atom of
          production, and  interleukin‐2 stimulates   iron. Hemoglobin concentration is meas-
          increases in the production of several leu-  ured in grams per 100 mL of blood, and
          kocytes. More information on individual   typical values for normal hemoglobin
          hematopoietins is provided in later sec-  concentrations  range  from  about  11  to
          tions of this chapter and in Chapter  16.   13 g/100 mL in domestic mammals.
          Based on the CFU terminology, hemat-       Hemoglobin functions in the transport
          opoietins have also been described as col­  of both  oxygen and  carbon dioxide.
          ony‐stimulating factors.                Oxygen binds to the ferrous iron in the
                                                  heme group to form  oxyhemoglobin
                                                  (HbO ); this process is termed oxygenation
                                                       2
          Erythrocytes                            (not oxidation). The amount of oxygen that
                                                  can be bound is proportional to the amount
          Erythrocytes (from the Greek  erythro‐,   of iron present, with one molecule of
          red; ‐cyte, cell) range from about 5 to 7 μm   oxygen combining with each atom of iron.
          in diameter. They are biconcave disks with   Because of the binding to hemoglobin,
          a thick circular margin and a thin center   blood can contain about 60 times as much
          (Fig. 15‐2). The biconcave shape provides   oxygen as would be dissolved in a similar
          a  relatively large surface area for gas   quantity of water in the same conditions.
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