The Cell Cycle

Cellular Organization of the Genetic

Overview: The Key Roles of Cell Division Material
• The ability of organisms to produce more of their • All the DNA in a cell constitutes the cell’s genome
own kind best distinguishes living things from • A genome can consist of a single DNA molecule
nonliving matter (common in prokaryotic cells) or a number of DNA
• The continuity of life is based on the reproduction molecules (common in eukaryotic cells)
of cells, or cell division • DNA molecules in a cell are packaged into
• In unicellular organisms, division of one cell Chromosomes
reproduces the entire organism
• Multicellular organisms depend on cell division for
– Development from a fertilized cell
– Growth
– Repair
• Cell division is an integral part of the cell cycle,
the life of a cell from formation to its own division

• Eukaryotic chromosomes consist of chromatin, a

complex of DNA and protein that condenses
during cell division
• Every eukaryotic species has a characteristic
number of chromosomes in each cell nucleus
• Somatic cells (nonreproductive cells) have two
sets of chromosomes
• Gametes (reproductive cells: sperm and eggs)
Most cell division results in genetically have half as many chromosomes as somatic cells
identical daughter cells
• Most cell division results in daughter cells with Distribution of Chromosomes During
identical genetic information, DNA Eukaryotic Cell Division
• The exception is meiosis, a special type of division • In preparation for cell division, DNA is replicated
that can produce sperm and egg cells and the chromosomes condense
• Each duplicated chromosome has two sisters • Eukaryotic cell division consists of
chromatids (joined copies of the original – Mitosis, the division of the genetic material in the
chromosome), which separate during cell division nucleus
• The centromere is the narrow “waist” of the – Cytokinesis, the division of the cytoplasm
duplicated chromosome, where the two • Gametes are produced by a variation of cell
chromatids are most closely attached division called meiosis
• Meiosis yields nonidentical daughter cells that
have only one set of chromosomes, half as many
as the parent cell

The mitotic phase alternates with

interphase in the cell cycle
• In 1882, the German anatomist Walther Flemming
developed dyes to observe chromosomes during
mitosis and cytokinesis
• During cell division, the two sister chromatids of
each duplicated chromosome separates and move Phases of the Cell Cycle
into two nuclei • The cell cycle consists of
• Once separate, the chromatids are called – Mitotic (M) phase (mitosis and cytokinesis)
Chromosomes – Interphase (cell growth and copying of
chromosomes in preparation for cell division)
• Interphase (about 90% of the cell cycle) can be
divided into subphases
– G1 phase (“first gap”)
– S phase (“synthesis”)
– G2 phase (“second gap”)
• The cell grows during all three phases, but
chromosomes are duplicated only during the S
phase
• Mitosis is conventionally divided into five phases
– Prophase • At metaphase, the chromosomes are all lined up
– Prometaphase at the metaphase plate, an imaginary structure at
– Metaphase the midway point between the spindle’s two poles
– Anaphase
– Telophase
• Cytokinesis overlaps the latter stages of mitosis

The Mitotic Spindle: A Closer Look

• The mitotic spindle is a structure made of
microtubules that controls chromosome movement • In anaphase, sister chromatids separate and move
during mitosis along the kinetochore microtubules toward
• In animal cells, assembly of spindle microtubules opposite ends of the cell
begins in the centrosome, the microtubule • The microtubules shorten by depolymerizing at
organizing center their kinetochore ends
• The centrosome replicates during interphase,
forming two centrosomes that migrate to opposite
ends of the cell during prophase and
prometaphase
• An aster (a radial array of short microtubules)
extends from each centrosome
• The spindle includes the centrosomes, the spindle
microtubules, and the asters
• During prometaphase, some spindle microtubules
attach to the kinetochores of chromosomes and
begin to move the chromosomes
• Kinetochores are protein complexes associated
with centromeres
• Nonkinetochore microtubules from opposite poles Binary Fission in Bacteria
overlap and push against each other, elongating • Prokaryotes (bacteria and archaea) reproduce by
the cell a type of cell division called binary fission
• In telophase, genetically identical daughter nuclei • In binary fission, the chromosome replicates
form at opposite ends of the cell (beginning at the origin of replication), and the
• Cytokinesis begins during anaphase or telophase two daughter chromosomes actively move apart
and the spindle eventually disassembles • The plasma membrane pinches inward, dividing
the cell into two
Cytokinesis: A Closer Look
• In animal cells, cytokinesis occurs by a process
known as cleavage, forming a cleavage furrow
• In plant cells, a cell plate forms during cytokinesis

The Evolution of Mitosis

• Since prokaryotes evolved before eukaryotes,
mitosis probably evolved from binary fission
• Certain protists exhibit types of cell division that
seem intermediate between binary fission and
mitosis
The eukaryotic cell cycle is regulated by a
molecular control system
The Cell Cycle Control System
• The frequency of cell division varies with the type
of cell • The sequential events of the cell cycle are directed
• These differences result from regulation at the by a distinct cell cycle control system, which is
molecular level similar to a clock
• Cancer cells manage to escape the usual controls • The cell cycle control system is regulated by both
on the cell cycle internal and external controls
• The clock has specific checkpoints where the cell
Evidence for Cytoplasmic Signals cycle stops until a go-ahead signal is received
• The cell cycle appears to be driven by specific
chemical signals present in the cytoplasm
• Some evidence for this hypothesis comes from
experiments in which cultured mammalian cells at
different phases of the cell cycle were fused to
form a single cell with two nuclei
• For many cells, the G1 checkpoint seems to be the
most important
• If a cell receives a go-ahead signal at the G1
checkpoint, it will usually complete the S, G2, and
M phases and divide
• If the cell does not receive the go-ahead signal, it
will exit the cycle, switching into a nondividing
state called the G0 phase

Stop and Go Signs: Internal and External

Signals at the Checkpoints
• An example of an internal signal is that
kinetochores not attached to spindle microtubules
send a molecular signal that delays anaphase
• Some external signals are growth factors,
proteins released by certain cells that stimulate
other cells to divide
• For example, platelet-derived growth factor
The Cell Cycle Clock: Cyclins and Cyclin- (PDGF) stimulates the division of human fibroblast
Dependent Kinases cells in culture
• Two types of regulatory proteins are involved in
cell cycle control: cyclins and cyclin-dependent
kinases (Cdks)
• Cdks activity fluctuates during the cell cycle
because it is controlled by cyclins, so named
because their concentrations vary with the cell
cycle
• MPF (maturation-promoting factor) is a cyclin-Cdk
complex that triggers a cell’s passage past the G2
checkpoint into the M phase
• A clear example of external signals is density process called transformation
dependent • Cancer cells that are not eliminated by the
inhibition, in which crowded cells stop immune system, form tumors, masses of
dividing abnormal cells within otherwise normal tissue
• Most animal cells also exhibit anchorage • If abnormal cells remain at the original site, the
dependence, in which they must be attached to a lump is called a benign tumor
substratum in order to divide • Malignant tumors invade surrounding tissues and
• Cancer cells exhibit neither density-dependent can metastasize, exporting cancer cells to other
inhibition nor anchorage dependence parts of the body, where they may form additional
tumors

Loss of Cell Cycle Controls in Cancer Cells

• Cancer cells do not respond normally to the body’s
control mechanisms
• Cancer cells may not need growth factors to grow
and divide
– They may make their own growth factor
– They may convey a growth factor’s signal without
the presence of the growth factor
– They may have an abnormal cell cycle control
System
• A normal cell is converted to a cancerous cell by a