MITOSIS

The four stages of mitosis are prophase, metaphase, anaphase, and telophase. In prophase, replicated
chromosomes composed of sister chromatids condense into recognizable bodies. A spindle forms
between the centrosomes as they separate to opposite poles of the cell. At the end of prophase the
nuclear envelope disintegrates, and the kinetochores of each chromosome become attached to the
centrosomes by microtubules (kinetochore fi bers). At metaphase the sister chromatids are moved to
the center of the cell, held there by the kinetochore fi bers. At anaphase the centromeres divide, and
the sister chromatids are pulled apart by the attached kinetochore fibers of the mitotic spindle. At
telophase the sister chromatids, now called chromosomes, gather in the position of the nucleus in each
cell and revert to a diffuse chromatin network. A nuclear membrane reappears, and cytokinesis occurs.
At the end of mitosis and cytokinesis, two cells genetically identical to the parent cell have been
produced.

All cells arise from the division of preexisting cells. All cells found in most multicellular organisms
originated from the divisionm of a single cell, a zygote, which is the product of union (fertilization) of an
egg and a sperm (gametes). Cell divisionnprovides the basis for one form of growth, for both sexual and
asexual reproduction, and for transmission of hereditary qualities from one cell generation to another
cell generation. In the formation of body cells (somatic cells) the process of nuclear division is mitosis.
By mitosis each “daughter cell” is ensured a complete set of genetic instructions. Mitosis is a delivery
system for distributing the chromosomes and the DNA they contain to continuing cell generations. Thus,
a single zygote divides by mitosis to produce a multicellular organism, and damaged cells are replaced
by mitosis during wound healing. As an animal grows, its somatic cells differentiate and assume different
functions and appearances because of differential gene action. Although most of the genes in
specialized cells remain silent and unexpressed throughout the lives of those cells, every cell possesses a
complete genetic complement. Mitosis ensures equality of genetic potential; later, other processes
direct the orderly expression of genes during embryonic development by selecting from the genetic
instructions that each cell contains. (These fundamental properties of cells of multicellular organisms are
discussed further in Chapter 8.) In animals that reproduce asexually (see Chapter 7), mitosis is the only
mechanism for the transfer of genetic information from parent to progeny, and thus the progeny are
genetically identical to the parents in this case. In animals that reproduce sexually (see Chapter 7), the
parents must produce sex cells (gametes or germ cells) that contain only half the usual number of
chromosomes, so that progeny formed by the union of gametes will not contain double the parental
number of chromosomes. This requires a special type of reductional division called meiosis, described in
Chapter 5 (p. 77).

MEIOSIS

Although animal species differ greatly in the characteristic numbers, sizes, and shapes of chromosomes
present in their body cells, a common feature is that chromosomes occur in pairs. The two members of a
chromosomal pair contain similar genes encoding the same set of characteristics and usually, but not
always, have the same size and shape. The members of such a pair are called homologous
chromosomes; each individual member of a pair is called a homolog. One homolog comes from the
mother and the other from the father. Meiosis is a special pair of cell divisions in which the genetic
material replicates once followed by two roundsof cell division ( Figure 5.2 ). The result is a set of four
daughter cells, each of which has only one member of each homologous chromosome pair. The
chromosomes present in a meiotic daughter cell or gamete are collectively called a single set of
chromosomes. The number of chromosomes in a single set, which varies among species, is called the
haploid ( n ) number of chromosomes. When a pair of gametes unites in fertilization, each gamete
contributes its set of chromosomes to the newly formed cell, called a zygote, which has two complete
sets of chromosomes. The number of chromosomes in two complete sets is called the diploid (2 n )
number. In humans the zygotes and all body cells normally have the diploid number (2 n ), or 46
chromosomes; the gametes have the haploid number ( n ), or 23, and meiosis reduces the number of
chromosomes per cell from diploid to haploid. Thus each cell normally has two copies of each gene
coding for a given trait, one on each of the homologous chromosomes. Alternative forms of genes for
the same trait are allelic forms, or alleles. Sometimes only one of the alleles has a visible effect on the
organism, although both are present in each cell, and either During an individual’s growth, all dividing
cells contain the double set of chromosomes (mitosis is described on p. 52). In the reproductive organs,
gametes (germ cells) are formed after meiosis, which separates the chromosomes of each homologous
pair. Without this reductional division, the union of ovum (egg) and sperm would produce an individual
with twice as many chromosomes as the parents. Continuation of this process in just a few generations
could yield astronomical numbers of chromosomes per cell. Most unique features of meiosis occur
during prophase of the fi rst meiotic division ( Figure 5.2 ). Prior to meiosis, each chromosome has
already replicated to form two chromatids joined at one point, the centromere. The two members of
each pair of homologous chromosomes make side-by-side contact (synapsis) to form a bivalent, which
permits genetic recombination between the paired homologous chromosomes (p. 89). Each bivalent is
composed of two pairs of chromatids (each pair is a dyad, sister chromatids held together at their
centromere), or four future chromosomes, and is thus called a tetrad. The position or location of any
gene on a chromosome is the gene locus (pl., loci ), and in synapsis all gene loci on a chromatid normally
lie exactly opposite the corresponding loci on the sister chromatid and both chromatids of the
homologous chromosome. Toward the end of prophase, the chromosomes shorten and thicken and
then enter the fi rst meiotic division. In contrast to mitosis, the centromeres holding the chromatids
together do not divide at anaphase. As a result, each of the dyads is pulled toward one of the opposite
poles of the cell by microtubules of the division spindle. At telophase of the fi rst meiotic division, each
pole of the cell has one dyad from each tetrad formed at prophase. Therefore at the end of the fi rst
meiotic division, the daughter cells contain one chromosome of each homologous pair from the parent
cell, so that the total chromosome number is reduced to haploid. However, because each chromosome
contains two chromatids joined at a centromere, each cell contains twice the amount of DNA present in
a gamete.The second meiotic division more closely resembles events in mitosis. The dyads are split at
the beginning of anaphase by division of their centromeres, and single-stranded chromosomes move
toward each pole. Thus by the end of the second meiotic division, the cells have the haploid number of
chromosomes, and each chromatid of the original tetrad exists in a separate nucleus. Four products are
formed, each containing one complete haploid set of chromosomes and only one copy of each gene.
Only one of the four products in female gametogenesisbecomes a functional gamete (p. 146).
Meiosis separates homologous chromosomes, so that each gamete has half the somatic chromosome
number (haploid). In the fi rst meiotic division, centromeres do not divide, and each daughter cell
receives one of each pair of replicated homologous chromosomes with sister chromatids still attached to
the centromere. At the beginning of the fi rst meiotic division, replicated homologous chromosomes
come to lie alongside each other (synapsis), forming a bivalent. The gene loci on one set of chromatids
lie opposite the corresponding loci on the homologous chromatids. Portions of adjacent chromatids can
exchange with the nonsister chromatids (crossing over) to produce new genetic combinations. At the
second meiotic division, the centromeres divide, completing the reduction in chromosome number and
amount of DNA. The diploid number is restored when male and female gametes fuse to form a zygote.