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98     Chapter 4    The Chromosome Theory of Inheritance


               4.3   Mitosis: Cell Division That                     eventually ends up enclosed in a separate nucleus in a sepa-
                                                                   rate cell. The two cells, known as daughter cells, are thus
              Preserves Chromosome Number                          genetically identical.
                                                                       The repeating pattern of cell growth (an increase in
                                                                   size) followed by division (the splitting of one cell into
                learning objectives                                two) is called the cell cycle (Fig. 4.9). Only a small part of
                                                                   the cell cycle is spent in division (or M phase); the period
                1.  Describe the key chromosome behaviors during mitosis.  between divisions is called interphase.
                2.  Diagram the forces and structures that dictate
                   chromosomal movement during mitosis.
                                                                   During Interphase, Cells Grow and
                                                                   Replicate Their Chromosomes
              The fertilized human egg is a single diploid cell that pre-  Interphase consists of three parts: gap 1 (G 1 ), synthesis (S),
              serves its genetic identity unchanged through more than   and gap 2 (G 2 ) (Fig. 4.9). G 1  lasts from the birth of a new cell
              100 generations of cells as it divides again and again to   to the onset of chromosome replication; for the genetic mate-
              produce a full-term infant ready to be born. As the newborn   rial, it is a period when the chromosomes are neither dupli-
              infant develops into a toddler, a teenager, and an adult, yet   cating nor dividing. During this time, the cell achieves most
              more cell divisions fuel continued growth and maturation.   of its growth by using the information from its genes to make
              Mitosis, the nuclear division that apportions chromosomes   and assemble the materials it needs to function normally. G 1
              in equal fashion to two daughter cells, is the cellular mech-  varies in length more than any other phase of the cell cycle.
              anism that preserves genetic information through all these   In rapidly dividing cells of the human embryo, for example,
              generations of cells. In this section, we take a close look at   G 1  is as short as a few hours. In contrast, mature brain cells
              how the nuclear division of mitosis fits into the overall
              scheme of cell growth and division.
                  If you were to peer through a microscope and follow the   Figure 4.9  The cell cycle: An alternation between
              history of one cell through time, you would see that for much   interphase and mitosis. (a) Chromosomes replicate to form sister
              of your observation, the chromosomes resemble a mass   chromatids during synthesis (S phase); the sister chromatids segregate
              of  extremely fine tangled string—called  chromatin—   to daughter cells during mitosis (M phase). The gaps between the S
              surrounded by the  nuclear envelope. Each convoluted   and M phases, during which most cell growth takes place, are called
              thread of chromatin is composed mainly of DNA (which   the G 1  and G 2  phases. In multicellular organisms, some terminally
                                                                   differentiated cells stop dividing and arrest in a G 0  stage. (b) Interphase
              carries the genetic information) and protein (which serves as   consists of the G 1 , S, and G 2  phases together.
              a scaffold for packaging and managing that information, as   (a)  The cell cycle
              described in Chapter 12). You would also be able to distin-
              guish one or two darker areas of chromatin called nucleoli               M
              (singular, nucleolus, literally small nucleus); nucleoli play a           Mitosis,
              key role in the manufacture of ribosomes, organelles that               cytokinesis
              function in protein synthesis. During the period between cell    G 2              G 1
              divisions, the chromatin-laden nucleus houses a great deal of
              invisible activity necessary for the growth and survival of the                      G
              cell. One particularly important part of this activity is the             Interphase   0
              accurate duplication of all the chromosomal material.                     duplication
                                                                                    Chromosome
                  With continued vigilance, you would observe a dramatic          S
              change in the nuclear landscape during one very short period
              in the cell’s life history: The chromatin condenses into dis-  (b)  Chromosomes replicate during S phase
              crete threads, and then each chromosome compacts even fur-
              ther into the twin rods clamped together at their centromeres   G : interphase,    A   a
                                                                           1
                                                                          gap before
              that can be identified in karyotype analysis (review Fig. 4.3).   duplication              B
              Each rod in a duo is called a chromatid; as described earlier,                                b
              it is an exact duplicate of the other sister chromatid to which   S: DNA synthesis
              it is connected. Continued observation would reveal the dou-  and chromosome
              bled chromosomes beginning to jostle around inside the cell,   duplication
              eventually lining up at the cell’s midplane. At this point, the                   A  A    a  B  B
              sister chromatids of each chromosome separate to opposite   G : interphase,           a
                                                                           2
              poles of the now elongating cell, where they  become identi-  gap before                     b   b
                                                                          mitosis
              cal  sets  of  chromosomes.  Each  of  the  two  identical  sets
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