ch18 - The Cell Division Cycle-2 (JP)

Chapter 18
The Cell-Division Cycle
Essential Cell Biology, Fifth Edition Copyright © 2019 W. W. Norton & Company
CHAPTER CONTENTS
OVERVIEW OF THE CELL CYCLE

THE CELL-CYCLE CONTROL SYSTEM
G1 PHASE
S PHASE
M PHASE
MITOSIS
CYTOKINESIS
CONTROL OF CELL NUMBERS AND CELL SIZE

G1 PHASE
• Cdks Are Stably Inactivated in G1

• Mitogens Promote the Production of the Cyclins That
Stimulate Cell Division
• DNA Damage Can Temporarily Halt Progression Through G1
• Cells Can Delay Division for Prolonged Periods by Entering
Specialized Nondividing States
The transition from G1 to S phase offers the cell a crossroad
: start in the cell cycle.
G1 PHASE

Cdks Are Stably Inactivated in G1
1. eliminating all of the existing cyclins,
2. blocking the synthesis of new ones
3. deploying Cdk inhibitor proteins to muffle the activity of any remaining cyclin–Cdk complexes
the cell can grow and monitor its environment before committing to a new round of division.
G1 PHASE

One way in which mitogens stimulate cell proliferation is by
inhibiting the Rb protein.
G1 PHASE

DNA damage can arrest the cell cycle at a checkpoint in G1
Essential Cell Biology, Fifth Edition
Copyright © 2019 W. W. Norton & Company
G1 PHASE

The transition from G1 to S phase offers the cell a crossroad
: start in the cell cycle.
Cell Cycle Differences
Early Embryonic Cycle

• no growth occurs
• each daughter cell is half the size of parent cell
• cycle time is very short
• S phases and M phases alternate without any intervening G1 or G2 phases
G0 Phase
• exits the cycle at G1 (cancer cells do not enter G0)
• cell can leave the cell cycle (temporarily or permanently)
• temporarily - quiescent
• permanently - terminally differentiated
• cell will never reenter the cell cycle
• carry out their function until they die
• not simply the absence of signals for mitosis, active repression of
the genes needed for mitosis
Stem Cells
• pluripotent self-renewal cell cycle of human embryonic stem cells (hESCs)
• short G1 period without reducing periods of time allocated to S phase, G2, and mitosis.
S PHASE
• S-Cdk Initiates DNA Replication and Blocks Re-Replication

• Incomplete Replication Can Arrest the Cell Cycle in G2
S PHASE

S phase
- S-Cdk initiates DNA replication and helps block re-replication
- Cohesins help hold the sister chromatids of each replicated

chromosomes together
- DNA damage checkpoints help prevent the replication of damaged

DNA
The initiation of DNA replication takes place in two steps
S-Cdk Initiates DNA Replication Once Per Cycle
S-Cdk Initiates DNA Replication Once Per Cycle
Origin Recognition Complex (ORC)
- ORC remains bound to origins of

replications throughout the cell cycle
- In early G1, Cdc6 concentration

increases transiently.
- When Cdc binds to ORCs in G1, it

promotes the binding of additional
proteins (cdt1,Mcm) to form a pre-
replicative complex (Pre-RC).
S-Cdk triggers DNA replication and

ensures that DNA replication is
initiated only once per cell cycle
Cohesins Hold Sister Chromatids Together
Cohesin is a protein complex with 4 subunits
At the end of S phase, each replicated chromosome consists of a pair

of identical sister chromatids glued together along their length
Cohesins Hold Sister Chromatids Together
The ATPase domains are required for

cohesion loading on the DNA
S PHASE

cdc25
Inhibitory phosphorylation and Cdk inhibitory proteins (CKI) can
suppress Cdk activity
M-Cdk
Summary 1
• Duplication of the chromosome in S phase involves the accurate

replication of the entire DNA molecules in each chromosome, as
well as the duplication of the chromatin proteins that associate
with the DNA and govern various aspects of chromosome function.
• Chromosome duplication is triggered by the activation of S-Cdk,
which activates proteins that unwind the DNA (helicase) and
initiate its replication at replication origins.
• Once a replication origin is activated, S-Cdk also inhibits proteins
that are required to allow that origin to initiate DNA replication
again.
• Thus, each origin is fired once and only once in each S phase and
cannot be reused until the next cell cycle.
M PHASE
• M-Cdk Drives Entry into Mitosis

• Cohesins and Condensins Help Configure Duplicated
Chromosomes for Separation
• Different Cytoskeletal Assemblies Carry Out Mitosis and
Cytokinesis
• M Phase Occurs in Stages
M PHASE

Cytokinesis
M-Cdk drives entry in M phase and mitosis
- triggers the condensation of the replicated chromosomes into compact,

rod-like structures
- induces mitotic spindle that will separate the condensed chromosomes

and segregated into two daughter cells
Cdk1-cyclin B
Activated M-Cdk indirectly activates more m-Cdk, creating a positive
feedback loop
Dephosphorylation Activates M-Cdk at the Onset of Mitosis
M PHASE

Cytokinesis
Cohesins and Condensins Help Configure Duplicated
Cohesins help hold the sister chromatids of each replicated
chromosomes together
Defects in sister-chromatid
cohesin lead to major errors
in chromosome segregation
Condensins help to coil the mitotic chromatids into smaller, more
compact structures that can be more easily segregated during mitosis
SMC: Structural Maintenance of Chromosomes
Cohesin contains SMC1 and SMC3 and is involved in sister chromatid cohesion.
The SMC5/6 complex contains SMC5 and SMC6 and is implicated in recombinational repair.
Condensins help to coil the mitotic chromatids into smaller, more
compact structures that can be more easily segregated during mitosis
M-Cdk phosphorylates condensins leads to stimulate the coiling activity
2 m of DNA into ~200 µm
Chromosome condensation
Sister chromatid resolution
Cohesins and Condensins Help Configure Duplicated
M PHASE

Cytokinesis
Two transient cytoskeletal structures mediate M phase in animal cells
M PHASE

Cytokinesis
PANEL 18–1 THE PRINCIPAL STAGES OF M PHASE IN
AN ANIMAL CELL
Introduction
Introduction
Introduction
Introduction
Introduction
Introduction

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Chapter 18

The Cell-Division Cycle

OVERVIEW OF THE CELL CYCLE

CONTROL OF CELL NUMBERS AND CELL SIZE

• Cdks Are Stably Inactivated in G1

• Cdks Are Stably Inactivated in G1

• Cdks Are Stably Inactivated in G1

• Cdks Are Stably Inactivated in G1

• Cdks Are Stably Inactivated in G1

Early Embryonic Cycle

• S-Cdk Initiates DNA Replication and Blocks Re-Replication

• S-Cdk Initiates DNA Replication and Blocks Re-Replication

- Cohesins help hold the sister chromatids of each replicated

- DNA damage checkpoints help prevent the replication of damaged

Origin Recognition Complex (ORC)

- ORC remains bound to origins of

- In early G1, Cdc6 concentration

- When Cdc binds to ORCs in G1, it

S-Cdk triggers DNA replication and

Cohesin is a protein complex with 4 subunits

At the end of S phase, each replicated chromosome consists of a pair

The ATPase domains are required for

• S-Cdk Initiates DNA Replication and Blocks Re-Replication

• Duplication of the chromosome in S phase involves the accurate

• M-Cdk Drives Entry into Mitosis

• M-Cdk Drives Entry into Mitosis

- triggers the condensation of the replicated chromosomes into compact,

- induces mitotic spindle that will separate the condensed chromosomes

• M-Cdk Drives Entry into Mitosis

SMC: Structural Maintenance of Chromosomes

M-Cdk phosphorylates condensins leads to stimulate the coiling activity

2 m of DNA into ~200 µm

• M-Cdk Drives Entry into Mitosis

• M-Cdk Drives Entry into Mitosis

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