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102    Chapter 4    The Chromosome Theory of Inheritance


              Figure 4.12  If cytokinesis does not follow mitosis, one   Figure 4.13  Checkpoints help regulate the cell cycle.
              cell may contain many nuclei. In fertilized Drosophila eggs,   Cellular checkpoints (red wedges) ensure that important events in
              13 rounds of mitosis take place without cytokinesis. The result is a   the cell cycle occur in the proper sequence. At each checkpoint,
              single-celled syncytial embryo that contains several thousand nuclei.   the cell determines whether prior events have been completed
              The photograph shows part of an embryo with dividing nuclei;   before it can proceed to the next step of the cell cycle. (For
              chromosomes are in red, and spindle fibers are in green. Nuclei at   simplicity, we show only two chromosomes per cell.)
              the upper left are in metaphase, while nuclei toward the bottom
              right are progressively later in anaphase. Membranes eventually   •  Is cell of sucient
              grow around these nuclei, dividing the embryo into cells.  size?         Chromosome
              © Dr. Byron Williams/Cornell University              •  Have proper signals    and       • Have the
                                                                    been received?     centrosome       chromosomes
                                                                      THEN: Duplicate   duplication     been completely
                                                                    chromosomes and                     duplicated?
                                                                    centrosomes                          THEN: Enter
                                                                                                        mitosis








                                                                                                            Prophase
                                                                                                 Mitosis
                                                                                Interphase

                                                                   Ongoing
                                                                   protein
                                                                   synthesis
                bodies, and (in plants) chloroplasts, must be parceled out to   and                         Metaphase
              the emerging daughter cells. The mechanism accomplish-  cell growth
              ing this task does not appear to predetermine which orga-     Telophase                  • Have all
              nelle is destined for which daughter cell. Instead, because   and                         chromosomes
                                                                                                        arrived and
              most cells contain many copies of these cytoplasmic struc-    cytokinesis       Anaphase  aligned at the
              tures, each new cell is bound to receive at least a few repre-                            metaphase
              sentatives of each component. This original complement of                                 plate?
              structures is enough to sustain the cell until synthetic activ-                            THEN: Initiate
                                                                                                        anaphase
              ity can repopulate the cytoplasm with organelles.
                  Sometimes cytoplasmic division does not immediately
              follow nuclear division, and the result is a cell containing   which the cell evaluates the results of previous steps—allow
              more than one nucleus. An animal cell with two or more   the sequential coordination of cell-cycle events (Fig. 4.13).
              nuclei is known as a syncytium. The early embryos of fruit   For example, the enzymes operating in one type of check-
              flies are multinucleated syncytia (Fig. 4.12), as are the pre-  point monitor DNA replication to ensure that cells do not
              cursors of spermatozoa in humans and many other animals.   begin mitosis until all the chromosomes have been com-
              A multinucleate plant tissue is called a coenocyte; coconut   pletely copied. If this checkpoint did not exist, at least one of
              milk is a nutrient-rich food composed of coenocytes.  the daughter cells would lose DNA every cell cycle.
                                                                       In a second illustration of the molecular basis of check-
                                                                   points, even a single kinetochore that has not attached to
              Regulatory Checkpoints Ensure Correct                spindle fibers generates a molecular signal that prevents the
              Chromosome Separation                                sister chromatids of all chromosomes from separating at
                                                                   their centromeres. This signal makes the beginning of an-
              The cell cycle is a complex sequence of precisely coordi-  aphase dependent on the prior proper alignment of all the
              nated events. In higher organisms, a cell’s “decision” to divide   chromosomes at metaphase. As a result of this cell-cycle
              depends on both intrinsic factors, such as conditions within   checkpoint, each daughter cell reliably receives the right
              the cell that register a sufficient size for division, and signals   number of chromosomes.
              from the environment, such as hormonal cues or contacts   Breakdown of the mitotic machinery can produce divi-
              with neighboring cells that encourage or  restrain division.   sion mistakes that have crucial consequences for the cell.
              Once a cell has initiated events leading to division, usually   Improper chromosome segregation, for example, can cause
              during the G 1  period of interphase, everything else follows   serious malfunction or even the death of daughter cells.
              like clockwork. A number of  checkpoints—moments at   Gene mutations that disrupt mitotic structures, such as the
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