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30  Veterinary Histology of Domestic Mammals and Birds



                   Endomitosis                                    ation exists in the events that are ascribed to each stage, the
       VetBooks.ir  Endomitosis is an incomplete internal form of division that  stages are consistently referred to as:
                   bears only partial similarity with mitosis. After replicating
                                                                    ·  leptotene (= thin strand),
                   during the S phase, the chromatids appear and separate,
                   but subsequent nuclear and cell division does not occur.   ·  zygotene (= yoke-like strand),
                   In contrast to mitosis the nuclear  envelope does not    ·  pachytene (= thick strand),
                   disintegrate, and a mitotic spindle is not formed.  ·  diplotene (= double strand) and
                      As the chromatids are not divided among two nuclei,   ·  diakinesis.
                   the nucleus retains double the usual number of chromo-
                   somes. Repeated endomitoses may occur, resulting in a  In the leptotene stage, chromatin fibrils undergo con-
                   high degree of polyploidy. This is usually accompanied by  densation to form elongated,  thin chromatin strands
                   the formation of multiple nucleoli, an increase in nuclear  (containing DNA that was duplicated in the previous S
                   volume, elevated metabolic activity and growth in cell  phase). Homologous pairs of  sister chromatids draw
                   volume. The nuclei of highly differentiated cells, such as  near to one another (homologous pairing) and become
                   megakaryocytes of the bone marrow or particular liver  connected by point-like  synaptonemal  complexes
                   cells, often undergo maturation through endomitosis.  (zygotene). The chromatin strands become shorter
                                                                  and thicker. In the  pachytene  stage the homologous
                   Amitosis                                       chromatid pairs are very closely associated. There is
                   In contrast to all of the previously described processes of  exchange of genetic material between maternal and
                   cell division, amitosis is characterised by direct division of  paternal chromosomes (crossing over). In the final stage
                   the nucleus and cytoplasm without the formation of vis-  of prophase I (diplotene), homologous chromosomes
                   ible chromosomes or breakdown of the nuclear envelope.  begin to separate. Junctions between the chromosomes
                   Division of the nucleus is brought about by microtubules  (chiasmata) represent sites at which crossing over has
                   that form a ‘drawstring’ around the nucleus, separating  occurred.
                   it into two portions of equivalent volume. The original   Towards the end of prophase, the nuclear envelope
                   organelle population is retained.              disintegrates and chromosomes become recognisable
                      Amitosis frequently results only in nuclear division, giv-  as separate entities. During this stage (diakinesis), kine-
                   ing rise to multinucleate cells.               tochore and spindle microtubules are formed.
                                                                     The subsequent phases (metaphase, anaphase and telo-
                   Meiosis                                        phase) proceed in a similar manner to their counterparts in
                   Mammalian fertilisation (syngamy, karyogamy) occurs  mitosis. In contrast to mitosis, however, the centromeres
                   through union of the nuclei of male and female gametes  that connect sister chromatids do not split. Consequently,
                   (oocytes and spermatozoa). In order to prevent doubling of  it is homologous chromatid pairs (rather than sister chro-
                   the number of chromosomes with each generation, the dip-  matids) that become separated. This results in halving of
                   loid set of chromosomes in the germ cells must be reduced  the number of chromosomes (Figure 1.39).
                   by half to become haploid. This is achieved by meiosis.
                      Meiosis involves two nuclear divisions. In the diploid  Meiosis II (equatorial division)
                   germ cell (2n), homologous chromosomes (equivalent  The sequence of events during meiosis II is similar to that
                   chromosomes of maternal and paternal origin) each  occurring during mitosis, with the exception that only a
                   form genetically identical chromatid pairs (4d). These  haploid set of chromosomes undergoes separation (Figure
                   exchange genetic material (‘crossing over’) before becom-  1.39). In contrast to meiosis I, the centromeres of each
                   ing separated in an initial division (meiosis I, reductional  chromatid pair become separated, allowing sister chro-
                   division). This gives rise to a haploid gamete in which  matids to be detached from one another. Division of the
                   the number of chromosomes and the amount of DNA is  two nuclei thus results in four haploid gametes (1n, 1d).
                   halved (1n, 2d). In a subsequent division, meiosis II (equa-  When the male and female gametes fuse at fertilisation,
                   torial division), the amount of DNA (but not the number  the two haploid nuclei combine to form a diploid somatic
                   of chromosomes) is halved again. This is achieved, as in  cell with a full complement of DNA (2n, 2d).
                   mitosis, by the separation of sister chromatids. In contrast
                   to mitosis, however, this division is not preceded by (fur-  Cell death
                   ther) duplication of DNA (Figure 1.39).        A physiological equilibrium exists in multicellular organs
                                                                  and tissues  between  the number of proliferating and
                   Meiosis I (reductional division)               degenerating cells. Abnormalities can result in an increase

                   Meiosis I begins with a prolonged prophase (prophase I)  in cell number (e.g. hyperplasia, neoplasia, autoimmune
                   that is subdivided into five stages (Figure 1.39). While vari-  disease) or a reduction in the cell population (e.g. atrophy,









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