Chapter 12: The Cell Cycle

Concept 12.1

Cell division requires the distribution of identical genetic material—DNA—to two daughter cells.

● A dividing cell duplicates its DNA, allocates the two copies to opposite ends of the cell, and then
splits into two daughter cells.
● A cell’s genetic information, packaged as DNA, is called its genome.
○ In prokaryotes, the genome is often a single long DNA molecule.
○ In eukaryotes, the genome consists of several DNA molecules.
● A human cell must duplicate about 2 m of DNA and separate the two copies such that each
daughter cell ends up with a complete genome.
● DNA molecules are packaged into chromosomes.
● Human somatic cells (body cells) have 46 chromosomes, made up of two sets of 23 (one from
each parent).
● Human gametes (sperm or eggs) have one set of 23 chromosomes, half the number in a somatic
cell.
● Eukaryotic chromosomes are made of chromatin, a complex of DNA and associated protein.
● The associated proteins maintain the structure of the chromosome and help control gene activity.
● Before cell division, chromatin condenses, coiling and folding to make a smaller package.
● Each duplicated chromosome consists of two sister chromatids, which contain identical copies of
the chromosome’s DNA.
● Later in cell division, the sister chromatids are pulled apart and repackaged into two new nuclei at
opposite ends of the parent cell.
● Mitosis, the formation of the two daughter nuclei, is usually followed by division of the cytoplasm,
cytokinesis.
● In contrast, gametes (eggs or sperm) are produced only in gonads (ovaries or testes) by a
variation of cell division called meiosis.
○ Meiosis yields four nonidentical daughter cells, each with half the chromosomes of the
parent.
○ In humans, meiosis reduces the number of chromosomes from 46 to 23.
○ Fertilization fuses two gametes together and doubles the number of chromosomes to 46
again.

Concept 12.2

The mitotic (M) phase of the cell cycle alternates with the much longer interphase.

○ The M phase includes mitosis and cytokinesis.

○ Interphase accounts for 90% of the cell cycle.
● During interphase, the cell grows by producing proteins and cytoplasmic organelles, copies its
chromosomes, and prepares for cell division.
● Interphase has three subphases: the G1 phase (“first gap”), the S phase (“synthesis”), and the G2
phase (“second gap”).
● For convenience, mitosis is usually broken into five subphases: prophase, prometaphase,
metaphase, anaphase, and telophase.
● In late interphase, the chromosomes have been duplicated but are not condensed.
○ A nuclear membrane bounds the nucleus, which contains one or more nucleoli.
○ The centrosome has replicated to form two centrosomes.
○ In animal cells, each centrosome features two centrioles.
● In prophase, the chromosomes are tightly coiled, with sister chromatids joined together.
○ The nucleoli disappear.
○ The mitotic spindle begins to form.
■ It is composed of centrosomes and the microtubules that extend from them.
○ The radial arrays of shorter microtubules that extend from the centrosomes are called
asters.
○ The centrosomes move away from each other, apparently propelled by lengthening
microtubules.
● During prometaphase, the nuclear envelope fragments, and microtubules from the spindle
interact with the condensed chromosomes.
○ Each of the two chromatids of a chromosome has a kinetochore, a specialized protein
structure located at the centromere.
○ Kinetochore microtubules from each pole attach to one of two kinetochores.
○ Nonkinetochore microtubules interact with those from opposite ends of the spindle.
● The spindle fibers push the sister chromatids until they are all arranged at the metaphase plate,
an imaginary plane equidistant from the poles, defining metaphase.
● At anaphase, the centromeres divide, separating the sister chromatids.
○ Each is now pulled toward the pole to which it is attached by spindle fibers.
○ By the end, the two poles have equivalent collections of chromosomes.
● At telophase, daughter nuclei begin to form at the two poles.
○ Nuclear envelopes arise from the fragments of the parent cell’s nuclear envelope and
other portions of the endomembrane system.
○ The chromosomes become less tightly coiled.
● Cytokinesis, the division of the cytoplasm, is usually well underway by late telophase.
● In animal cells, cytokinesis involves the formation of a cleavage furrow, which pinches the cell in
two.
● In plant cells, vesicles derived from the Golgi apparatus produce a cell plate at the middle of the
cell.
● The spindle fibers elongate by incorporating more subunits of the protein tubulin.
● Assembly of the spindle microtubules starts in the centrosome.
● During interphase, the single centrosome replicates to form two centrosomes.
● As mitosis starts, the two centrosomes are located near the nucleus.
○ As the spindle microtubules grow from them, the centrioles are pushed apart.
○ By the end of prometaphase, they are at opposite ends of the cell.
● An aster, a radial array of short microtubules, extends from each centrosome.
● The spindle includes the centrosomes, the spindle microtubules, and the asters.
● Each sister chromatid has a kinetochore of proteins and chromosomal DNA at the centromere.
○ The kinetochores of the joined sister chromatids face in opposite directions.
● During prometaphase, some spindle microtubules (called kinetochore microtubules) attach to the
kinetochores.
● When a chromosome’s kinetochore is “captured” by microtubules, the chromosome moves
toward the pole from which those microtubules come.
● When microtubules attach to the other pole, this movement stops and a tug-of-war ensues.
 ● Eventually, the chromosome settles midway between the two poles of the cell, on the metaphase
plate.
● Nonkinetochore microtubules from opposite poles overlap and interact with each other.
● By metaphase, the microtubules of the asters have grown and are in contact with the plasma
membrane.
● The spindle is now complete.
● Anaphase commences when the proteins holding the sister chromatids together are inactivated.
○ Once the chromosomes are separate, full-fledged chromosomes, they move toward
opposite poles of the cell.
● Mitosis in eukaryotes may have evolved from binary fission in bacteria.
● Prokaryotes reproduce by binary fission, not mitosis.
● Most bacterial genes are located on a single bacterial chromosome that consists of a circular
DNA molecule and associated proteins.
● While bacteria are smaller and simpler than eukaryotic cells, they still have large amounts of DNA
that must be copied and distributed equally to two daughter cells.
● The circular bacterial chromosome is highly folded and coiled in the cell.
● In binary fission, chromosome replication begins at one point in the circular chromosome, the
origin of replication site, producing two origins.
● How did mitosis evolve?
○ There is evidence that mitosis had its origins in bacterial binary fission.
○ Some of the proteins involved in binary fission are related to eukaryotic proteins.
○ Two of these are related to eukaryotic tubulin and actin proteins.

Concept 12.3

● A checkpoint in the cell cycle is a critical control point where stop and go-ahead signals regulate
the cycle.
○ The signals are transmitted within the cell by signal transduction pathways.
○ Animal cells generally have built-in stop signals that halt the cell cycle at checkpoints until
overridden by go-ahead signals.
○ Many signals registered at checkpoints come from cellular surveillance mechanisms.
○ These indicate whether key cellular processes have been completed correctly.
○ Checkpoints also register signals from outside the cell.
● Three major checkpoints are found in the G1, G2, and M phases.
● For many cells, the G1 checkpoint, the “restriction point” in mammalian cells, is the most
important.
○ If the cell receives a go-ahead signal at the G1 checkpoint, it usually completes the cell
cycle and divides.
○ If it does not receive a go-ahead signal, the cell exits the cycle and switches to a
nondividing state, the G0 phase.
■ Most cells in the human body are in this phase.
■ Liver cells can be “called back” to the cell cycle by external cues, such as growth
factors released during injury.
■ Highly specialized nerve and muscle cells never divide.
● Cancer cells have escaped from cell cycle controls.
 ● Cancer cells divide excessively and invade other tissues because they are free of the body’s
control mechanisms.
○ Cancer cells do not stop dividing when growth factors are depleted.
○ This is either because a cancer cell manufactures its own growth factors, has an
abnormality in the signaling pathway, or has an abnormal cell cycle control system.
● If and when cancer cells stop dividing, they do so at random points, not at the normal checkpoints
in the cell cycle.

Chapter 13: Meiosis and Sexual Life Cycles

Concept 13.1

● The transmission of hereditary traits has its molecular basis in the precise replication of DNA.
○ This produces copies of genes that can be passed from parents to offspring.
● In plants and animals, sperm and ova (unfertilized eggs) transmit genes from one generation to
the next.
● In asexual reproduction, a single individual is a sole parent to donate genes to its offspring.
○ Single-celled eukaryotes can reproduce asexually by mitotic cell division to produce two
genetically identical daughter cells.
○ Some multicellular eukaryotes, like Hydra, can reproduce by budding, producing a mass
of cells by mitosis.
● An individual that reproduces asexually gives rise to a clone, a group of genetically identical
individuals.
○ Members of a clone may be genetically different as a result of mutation.
● Unlike a clone, offspring produced by sexual reproduction vary genetically from their siblings and
their parents.

Concept 13.2

● Each chromosome can be distinguished by size, the position of the centromere, and pattern of
staining with certain dyes.
● The two chromosomes comprising a pair have the same length, centromere position, and staining
pattern.
● The pattern of inheritance of the sex chromosomes determines an individual’s sex.
○ Human females have a homologous pair of X chromosomes (XX).
○ Human males have an X and a Y chromosome (XY).
● Only small parts of the X and Y are homologous.
○ Most of the genes carried on the X chromosome do not have counterparts on the tiny Y.
○ The Y chromosome also has genes not present on the X.
● The occurrence of homologous pairs of chromosomes is a consequence of sexual reproduction.
● We inherit one chromosome of each homologous pair from each parent.
 ○ The 46 chromosomes in each somatic cell are two sets of 23, a maternal set (from your
mother) and a paternal set (from your father).
● Any cell with two sets of chromosomes is called a diploid cell and has a diploid number of
chromosomes, abbreviated as 2n.
● A gamete with a single chromosome set is haploid, abbreviated as n.
● Any sexually reproducing species has a characteristic haploid and a diploid number of
chromosomes.
○ For humans, the haploid number of chromosomes is 23 (n = 23), and the diploid number
is 46 (2n = 46).

Concept 13.3

Many steps of meiosis resemble steps in mitosis.

○ Both are preceded by the replication of chromosomes.

● However, in meiosis, there are two consecutive cell divisions, meiosis I and meiosis II, resulting in
four daughter cells.
○ The first division, meiosis I, separates homologous chromosomes.
○ The second, meiosis II, separates sister chromatids.
● The four daughter cells have only half as many chromosomes as the parent cell.
● Meiosis I is preceded by interphase, in which the chromosomes are replicated to form sister
chromatids.
○ These are genetically identical and joined at the centromere.
○ The single centrosome is replicated, forming two centrosomes.
● Division in meiosis I occurs in four phases: prophase I, metaphase I, anaphase I, and telophase I.
● Prophase I
● Prophase I typically occupies more than 90% of the time required for meiosis.
● During prophase I, the chromosomes begin to condense.
● Homologous chromosomes loosely pair up along their length, precisely aligned gene for gene.
○ In crossing over, DNA molecules in nonsister chromatids break at corresponding places
and then rejoin the other chromatid.
○ In synapsis, a protein structure called the synaptonemal complex forms between
homologues, holding them tightly together along their length.
○ As the synaptonemal complex disassembles in late prophase, each chromosome pair
becomes visible as a tetrad, or group of four chromatids.
○ Each tetrad has one or more chiasmata, sites where the chromatids of homologous
chromosomes have crossed and segments of the chromatids have been traded.
○ Spindle microtubules form from the centrosomes, which have moved to the poles.
○ The breakdown of the nuclear envelope and nucleoli take place.
○ Kinetochores of each homologue attach to microtubules from one of the poles.
● Metaphase I
● At metaphase I, the tetrads are all arranged at the metaphase plate, with one chromosome facing
each pole.
○ Microtubules from one pole are attached to the kinetochore of one chromosome of each
tetrad, while those from the other pole are attached to the other.
● Anaphase I
● In anaphase I, the homologous chromosomes separate. One chromosome moves toward each
pole, guided by the spindle apparatus.
 ○ Sister chromatids remain attached at the centromere and move as a single unit toward
the pole.
● Telophase I and cytokinesis
● In telophase I, movement of homologous chromosomes continues until there is a haploid set at
each pole.
○ Each chromosome consists of two sister chromatids.
● Cytokinesis usually occurs simultaneously, by the same mechanisms as mitosis.
○ In animal cells, a cleavage furrow forms. In plant cells, a cell plate forms.
● No chromosome replication occurs between the end of meiosis I and the beginning of meiosis II,
as the chromosomes are already replicated.
● Meiosis II
● Meiosis II is very similar to mitosis.
○ During prophase II, a spindle apparatus forms and attaches to kinetochores of each sister
chromatid.
■ Spindle fibers from one pole attach to the kinetochore of one sister chromatid,
and those of the other pole attach to kinetochore of the other sister chromatid.
● At metaphase II, the sister chromatids are arranged at the metaphase plate.
○ Because of crossing over in meiosis I, the two sister chromatids of each chromosome are
no longer genetically identical.
○ The kinetochores of sister chromatids attach to microtubules extending from opposite
poles.
● At anaphase II, the centromeres of sister chromatids separate and two newly individual
chromosomes travel toward opposite poles.
● In telophase II, the chromosomes arrive at opposite poles.
○ Nuclei from around the chromosomes, which begin expanding, and cytokinesis separates
the cytoplasm.
● At the end of meiosis, there are four haploid daughter cells.

Concept 13.4

● Sexual life cycles produce genetic variation among offspring.

● Three mechanisms contribute to genetic variation:
1. Independent assortment of chromosomes.
2. Crossing over.
3. Random fertilization.
● Crossing over produces recombinant chromosomes, which combine genes inherited from each
parent.
● Crossing over begins very early in prophase I as homologous chromosomes pair up gene by
gene.
● In crossing over, homologous portions of two nonsister chromatids trade places.
1. For humans, this occurs an average of one to three times per chromosome pair.
● The three sources of genetic variability in a sexually reproducing organism are:
1. Independent assortment of homologous chromosomes during meiosis I and of
nonidentical sister chromatids during meiosis II.
2. Crossing over between homologous chromosomes during prophase I.
3. Random fertilization of an ovum by a sperm.