The Cell Cycle and How Cells Divide

Phases of the Cell Cycle

The cell cycle consists of Interphase normal cell activity The mitotic phase cell divsion
INTERPHASE

Growth G1 (DNA synthesis) Growth G2

Functions of Cell Division

100 m

200 m

20 m

(a) Reproduction. An amoeba, a single-celled eukaryote, is dividing into two cells. Each new cell will be an individual organism (LM).

(b) Growth and development. (c) Tissue renewal. These dividing This micrograph shows a bone marrow cells (arrow) will sand dollar embryo shortly after give rise to new blood cells (LM). the fertilized egg divided, forming two cells (LM).

Cell Division

An integral part of the cell cycle Results in genetically identical daughter cells Cells duplicate their genetic material Before they divide, ensuring that each daughter cell receives an exact copy of the genetic material, DNA

DNA

Genetic information - genome Packaged into chromosomes

Figure 12.3
50 m

DNA And Chromosomes

An average eukaryotic cell has about 1,000 times more DNA then an average prokaryotic cell. The DNA in a eukaryotic cell is organized into several linear chromosomes, whose organization is much more complex than the single, circular DNA molecule in a prokaryotic cell

Chromosomes

All eukaryotic cells store genetic information in chromosomes. Most eukaryotes have between 10 and 50 chromosomes in their body cells. Human cells have 46 chromosomes. 23 nearly-identical pairs

Structure of Chromosomes

Chromosomes are composed of a complex of DNA and protein called chromatin that condenses during cell division DNA exists as a single, long, doublestranded fiber extending chromosomes entire length. Each unduplicated chromosome contains one DNA molecule, which may be several inches long
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Structure of Chromosomes
Every 200 nucleotide pairs, the DNA wraps twice around a group of 8 histone proteins to form a nucleosome. Higher order coiling and supercoiling also help condense and package the chromatin inside the nucleus:

Structure of Chromosomes
The degree of coiling can vary in different regions of the chromatin: Heterochromatin refers to highly coiled regions where genes arent expressed. Euchromatin refers to loosely coiled regions where genes can be expressed.

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Structure of Chromosomes

Prior to cell division each chromosome duplicates itself. During this time, only the heterochromatin is visible, as dense granules inside the nucleus. There is also a dense area of RNA production called the nucleolus:
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Karyotype

An ordered, visual representation of the chromosomes in a cell Chromosomes are photographed when they are highly condensed, then photos of the individual chromosomes are arranged in order of decreasing size: In humans each somatic cell has 46 chromosomes, made up of two sets, one set of chromosomes comes from each parent

Pair of homologous chromosomes

5 m

Centromere

Sister chromatids

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Chromosomes

Non-homologous chromosomes Look different Control different traits Sex chromosomes Are distinct from each other in their characteristics Are represented as X and Y Determine the sex of the individual, XX being female, XY being male In a diploid cell, the chromosomes occur in pairs. The 2 members of each pair are called homologous chromosomes or homologues.
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Chromosomes

A diploid cell has two sets of each of its chromosomes A human has 46 chromosomes (2n = 46) In a cell in which DNA synthesis has occurred all the chromosomes are duplicated and thus each consists of two identical sister chromatids
Maternal set of chromosomes (n = 3) 2n = 6 Paternal set of chromosomes (n = 3)

Two sister chromatids of one replicated chromosome Centromere

Two nonsister chromatids in a homologous pair

Pair of homologous chromosomes (one from each set)

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Homologues
Homologous chromosomes: Look the same Control the same traits May code for different forms of each trait Independent origin - each one was inherited from a different parent

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Chromosome Duplication

In preparation for cell division, DNA is replicated and the chromosomes condense Each duplicated chromosome has two sister chromatids, which separate during cell division
0.5 m A eukaryotic cell has multiple chromosomes, one of which is represented here. Before duplication, each chromosome has a single DNA molecule.

Chromosome duplication (including DNA synthesis) Centromere

Once duplicated, a chromosome consists of two sister chromatids connected at the centromere. Each chromatid contains a copy of the DNA molecule.

Mechanical processes separate the sister chromatids into two chromosomes and distribute them to two daughter cells.

Separation of sister chromatids

Sister chromatids

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Centrometers
Sister chromatids

Chromosome Duplication

Because of duplication, each condensed chromosome consists of 2 identical chromatids joined by a centromere. Each duplicated chromosome contains 2 identical DNA molecules (unless a mutation occurred), one in each chromatid:
Non-sister chromatids

Centromere

Duplication

Sister chromatids Two unduplicated chromosomes

Sister chromatids

Two duplicated chromosomes

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Structure of Chromosomes

The centromere is a constricted region of the chromosome containing a specific DNA sequence, to which is bound 2 discs of protein called kinetochores. Kinetochores serve as points of attachment for microtubules that move the chromosomes during cell division:
Metaphase chromosome

Centromere region of chromosome Kinetochore microtubules

Kinetochore

Sister Chromatids
18
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Structure of Chromosomes

Diploid - A cell possessing two copies of each chromosome (human body cells). Homologous chromosomes are made up of sister chromatids joined at the centromere. Haploid - A cell possessing a single copy of each chromosome (human sex cells).

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Phases of the Cell Cycle

Interphase G1 - primary growth S - genome replicated G2 - secondary growth M - mitosis C - cytokinesis

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Interphase

G1 - Cells undergo majority of growth S - Each chromosome replicates (Synthesizes) to

produce sister chromatids Attached at centromere Contains attachment site (kinetochore)

G2 - Chromosomes condense - Assemble

machinery for division such as centrioles

21

Mitosis

Some haploid & diploid cells divide by mitosis. Each new cell receives one copy of every chromosome that was present in the original cell. Produces 2 new cells that are both genetically identical to the original cell.
DNA duplication during interphase

Mitosis

Diploid Cell

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Mitotic Division of an Animal Cell

G2 OF INTERPHASE

PROPHASE

PROMETAPHASE

Centrosomes (with centriole pairs)

Chromatin (duplicated)

Early mitotic spindle

Aster Centromere

Fragments of nuclear envelope

Kinetochore Nonkinetochore microtubules

Nucleolus

Nuclear Plasma envelope membrane

Chromosome, consisting of two sister chromatids

Kinetochore microtubule

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Mitotic Division of an Animal Cell

METAPHASE Metaphase plate

ANAPHASE

TELOPHASE AND CYTOKINESIS Cleavage furrow

Nucleolus forming

Spindle

Centrosome at Daughter one spindle pole chromosomes

Nuclear envelope forming

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G2 of Interphase

A nuclear envelope bounds the nucleus. The nucleus contains one or more nucleoli (singular, nucleolus). Two centrosomes have formed by replication of a single centrosome. In animal cells, each centrosome features two centrioles. Chromosomes, duplicated during S phase, cannot be seen individually because they have not yet condensed.

G2 OF INTERPHASE

Centrosomes (with centriole pairs)

Chromatin (duplicated)

The light micrographs show dividing lung cells from a newt, which has 22 chromosomes in its somatic cells (chromosomes appear blue, microtubules green, intermediate filaments red). For simplicity, the drawings show only four chromosomes.
Nucleolus Nuclear Plasma envelope membrane

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Prophase
The chromatin fibers become more tightly coiled, condensing into discrete chromosomes observable with a light microscope. The nucleoli disappear. Each duplicated chromosome appears as two identical sister chromatids joined together. The mitotic spindle begins to form. It is composed of the centrosomes and the microtubules that extend from them. The radial arrays of shorter microtubules that extend from the centrosomes are called asters (stars). The centrosomes move away from each other, apparently propelled by the lengthening microtubules between them.

PROPHASE

Early mitotic spindle

Aster Centromere

Chromosome, consisting of two sister chromatids

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Metaphase
Metaphase is the longest stage of mitosis, lasting about 20 minutes. The centrosomes are now at opposite ends of the cell. The chromosomes convene on the metaphase plate, an imaginary plane that is equidistant between the spindles two poles. The chromosomes centromeres lie on the metaphase plate. For each chromosome, the kinetochores of the sister chromatids are attached to kinetochore microtubules coming from opposite poles. The entire apparatus of microtubules is called the spindle because of its shape.
Spindle

METAPHASE Metaphase plate

Centrosome at one spindle pole

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The Mitotic Spindle

The spindle includes the centrosomes, the spindle microtubules, and the asters The apparatus of microtubules controls chromosome movement during mitosis The centrosome replicates, forming two centrosomes that migrate to opposite ends of the cell Assembly of spindle microtubules begins in the centrosome, the microtubule organizing center An aster (a radial array of short microtubules) extends from each centrosome
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The Mitotic Spindle

Some spindle microtubules attach to the kinetochores of chromosomes and move the chromosomes to the metaphase plate In anaphase, sister chromatids separate and move along the kinetochore microtubules toward opposite ends of the cell
Aster Microtubules Chromosomes Sister chromatids Centrosome Metaphase plate Kinetochores

Centrosome

1 m

Overlapping nonkinetochore microtubules

Kinetochore microtubules 0.5 m

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Anaphase
Anaphase is the shortest stage of mitosis, lasting only a few minutes. Anaphase begins when the two sister chromatids of each pair suddenly part. Each chromatid thus becomes a fullfledged chromosome. The two liberated chromosomes begin moving toward opposite ends of the cell, as their kinetochore microtubules shorten. Because these microtubules are attached at the centromere region, the chromosomes move centromere first (at about 1 m/min). The cell elongates as the nonkinetochore microtubules lengthen. By the end of anaphase, the two ends of the cell have equivalentand completecollections of chromosomes.

ANAPHASE

Daughter chromosomes

30

Telophase Two daughter nuclei begin to form in the cell. Nuclear envelopes arise from the fragments of the parent cells nuclear envelope and other portions of the endomembrane system. The chromosomes become less condensed. Mitosis, the division of one nucleus into two genetically identical nuclei, is now complete.

TELOPHASE AND CYTOKINESIS Cleavage furrow

Nucleolus forming

Nuclear envelope forming

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Mitosis in a plant cell

Chromatine Nucleus Nucleolus condensing Chromosome

1 Prophase. The chromatin is condensing. The nucleolus is beginning to disappear. Although not yet visible in the micrograph, the mitotic spindle is staring to from.

Metaphase. The 2 Prometaphase. 3 4 spindle is complete, We now see discrete and the chromosomes, chromosomes; each attached to microtubules consists of two at their kinetochores, identical sister are all at the metaphase chromatids. Later plate. in prometaphase, the nuclear envelop will fragment.

5 Anaphase. The chromatids of each chromosome have separated, and the daughter chromosomes are moving to the ends of cell as their kinetochore microtubles shorten.

Telophase. Daughter nuclei are forming. Meanwhile, cytokinesis has started: The cell plate, which will divided the cytoplasm in two, is growing toward the perimeter of the parent cell.

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Cytokinesis

Cleavage of cell into two halves Animal cells Constriction belt of actin filaments Plant cells Cell plate Fungi and protists Mitosis occurs within the nucleus
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Cytokinesis In Animal And Plant Cells

100 m

Cleavage furrow

Vesicles forming cell plate

Wall of patent cell

1 m

Cell plate New cell wall

Contractile ring of microfilaments

Daughter cells
Daughter cells

(a) Cleavage of an animal cell (SEM)

(b) Cell plate formation in a plant cell (SEM)

34

35

Meiosis and Sexual Life Cycles

Living organisms are distinguished by their ability to reproduce their own kind Heredity Is the transmission of traits from one generation to the next Variation Shows that offspring differ somewhat in appearance from parents and siblings

36

Inheritance of Genes

Genes are segments of DNA, units of heredity Offspring acquire genes from parents by inheriting chromosomes Genetics is the scientific study of heredity and hereditary variation

37

Inheritance of Genes

Each gene in an organisms DNA has a specific locus on a certain chromosome We inherit one set of chromosomes from our mother and one set from our father Two parents give rise to offspring that have unique combinations of genes inherited from the two parents - sexual reproduction

38

Asexual Reproduction

In asexual reproduction, one parent produces genetically identical offspring by mitosis

Parent Bud

Figure 13.2

0.5 mm
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Sexual Reproduction

Fertilization and meiosis alternate in sexual life cycles A life cycle is the generation-to-generation sequence of stages in the reproductive history of an organism
Key Haploid Diploid n Gametes n MEIOSIS FERTILIZATION n

Zygote 2n Diploid multicellular organism

2n

Mitosis (a) Animals

40

Sex Cells - Gametes

Unlike somatic cells, sperm and egg cells are haploid cells, containing only one set of chromosomes At sexual maturity the ovaries and testes produce haploid gametes by meiosis

41

Sexual Reproduction - The Human Life Cycle

Haploid gametes (n = 23)

During fertilization, sperm and ovum fuse forming a diploid zygote The zygote develops into an adult organism

Haploid (n) Diploid (2n)

Ovum (n)

Sperm Cell (n) MEIOSIS FERTILIZATION

Ovary

Testis

Diploid zygote (2n = 46)

Mitosis and development Multicellular diploid adults (2n = 46)

42

Meiosis

Reduces the chromosome number such that each daughter Cell has a haploid set of chromosomes Ensures that the next generation will have: Diploid number of chromosome Exchange of genetic information (combination of traits that differs from that of either parent)
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Meiosis

Only diploid cells can divide by meiosis. Prior to meiosis I, DNA replication occurs. During meiosis, there will be two nuclear divisions, and the result will be four haploid nuclei. No replication of DNA occurs between meiosis I and meiosis II.

44

Meiosis
Interphase

Meiosis reduces the number of chromosome sets from diploid to haploid Meiosis takes place in two sets of divisions

Homologous pair of chromosomes in diploid parent cell

Chromosomes replicate Homologous pair of replicated chromosomes

Sister chromatids

Diploid cell with replicated chromosomes

Meiosis I reduces the number of chromosomes from diploid to haploid Meiosis II produces four haploid daughter cells
Figure 13.7

Meiosis I

1 Homologous
chromosomes separate

Haploid cells with replicated chromosomes Meiosis II 2 Sister chromatids

separate

Haploid cells with unreplicated chromosomes

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Meiosis Phases

Meiosis involves the same four phases seen in mitosis prophase metaphase anaphase telophase They are repeated during both meiosis I and meiosis II. The period of time between meiosis I and meiosis II is called interkinesis. No replication of DNA occurs during interkinesis because the DNA is already duplicated.
46

Prophase I

Prophase I occupies more than 90% of the time required for meiosis Chromosomes begin to condense In synapsis, the 2 members of each homologous pair of chromosomes line up side-by-side, aligned gene by gene, to form a tetrad consisting of 4 chromatids During synapsis, sometimes there is an exchange of homologous parts between non-sister chromatids. This exchange is called crossing over Each tetrad usually has one or more chiasmata, X-shaped regions where crossing over occurred
Prophase I of meiosis Nonsister chromatids

Tetrad

Chiasma, site of crossing over

47

Metaphase I

At metaphase I, tetrads line up 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 Microtubules from the other pole are attached to the kinetochore of the other chromosome
PROPHASE I
Sister chromatids Chiasmata

METAPHASE I
Centromere (with kinetochore) Spindle Metaphase plate

ANAPHASE I
Sister chromatids remain attached

Tetrad

Microtubule attached to kinetochore

Homologous chromosomes separate Pairs of homologous chromosomes split up

Homologous chromosomes (red and blue) pair and exchange segments; 2n = 6

Tetrads line up

48

Anaphase I

In anaphase I, pairs of homologous chromosomes separate One chromosome moves toward each pole, guided by the spindle apparatus Sister chromatids remain attached at the centromere and move as one unit toward the pole
PROPHASE I
Sister chromatids Chiasmata

METAPHASE I
Centromere (with kinetochore) Spindle Metaphase plate

ANAPHASE I
Sister chromatids remain attached

Tetrad

Microtubule attached to kinetochore

Homologous chromosomes separate Pairs of homologous chromosomes split up

Homologous chromosomes (red and blue) pair and exchange segments; 2n = 6

Tetrads line up

49

Telophase I and Cytokinesis

In the beginning of telophase I, each half of the cell has a haploid set of chromosomes; each chromosome still consists of two sister chromatids Cytokinesis usually occurs simultaneously, forming two haploid daughter cells 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 because the chromosomes are already replicated

50

Prophase II

Meiosis II is very similar to mitosis In prophase II, a spindle apparatus forms In late prophase II, chromosomes (each still composed of two chromatids) move toward the metaphase plate
TELOPHASE I AND CYTOKINESIS PROPHASE II METAPHASE II ANAPHASE II TELOPHASE II AND CYTOKINESIS

Cleavage furrow

Sister chromatids separate

Haploid daughter cells forming

51

Metaphase II

At metaphase II, the sister chromatids are 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
TELOPHASE I AND CYTOKINESIS PROPHASE II METAPHASE II ANAPHASE II TELOPHASE II AND CYTOKINESIS

Cleavage furrow

Sister chromatids separate

Haploid daughter cells forming

52

Anaphase II

At anaphase II, the sister chromatids separate The sister chromatids of each chromosome now move as two newly individual chromosomes toward opposite poles
TELOPHASE I AND CYTOKINESIS PROPHASE II METAPHASE II ANAPHASE II TELOPHASE II AND CYTOKINESIS

Cleavage furrow

Sister chromatids separate

Haploid daughter cells forming

53

Telophase II and Cytokinesis

In telophase II, the chromosomes arrive at opposite poles Nuclei form, and the chromosomes begin decondensing Cytokinesis separates the cytoplasm At the end of meiosis, there are four daughter cells, each with a haploid set of unreplicated chromosomes Each daughter cell is genetically distinct from the others and from the parent cell
TELOPHASE I AND CYTOKINESIS
PROPHASE II METAPHASE II ANAPHASE II TELOPHASE II AND CYTOKINESIS

Cleavage furrow

Sister chromatids separate

Haploid daughter cells forming

54

A Comparison of Mitosis and Meiosis

Mitosis conserves the number of chromosome sets, producing cells that are genetically identical to the parent cell Meiosis reduces the number of chromosomes sets from two (diploid) to one (haploid), producing cells that differ genetically from each other and from the parent cell The mechanism for separating sister chromatids is virtually identical in meiosis II and mitosis

55

A Comparison of Mitosis and Meiosis

Three events are unique to meiosis, and all three occur in meiosis l:

Synapsis and crossing over in prophase I: Homologous chromosomes physically connect and exchange genetic information At the metaphase plate, there are paired homologous chromosomes (tetrads), instead of individual replicated chromosomes At anaphase I of meiosis, homologous pairs move toward opposite poles of the cell. In anaphase II of meiosis, the sister chromatids separate

56

A Comparison Of Mitosis And Meiosis

MITOSIS
Parent cell (before chromosome replication)

MEIOSIS
Chiasma (site of crossing over) MEIOSIS I

Prophase

Prophase I Chromosome replication 2n = 6 Chromosome replication Tetrad formed by synapsis of homologous chromosomes

Duplicated chromosome (two sister chromatids)

Metaphase

Chromosomes positioned at the metaphase plate

Tetrads positioned at the metaphase plate

Metaphase I

Anaphase Telophase

Sister chromatids separate during anaphase

Homologues separate during anaphase I; sister chromatids remain together

Anaphase I Telophase I Haploid n=3

Daughter cells of meiosis I

2n Daughter cells of mitosis

2n

MEIOSIS II

Daughter cells of meiosis II Sister chromatids separate during anaphase II

57

Comparison

Meiosis
DNA duplication followed by 2 cell divisions Sysnapsis Crossing-over One diploid cell produces 4 haploid cells Each new cell has a unique combination of genes

Mitosis
Homologous chromosomes do not pair up No genetic exchange between homologous chromosomes One diploid cell produces 2 diploid cells or one haploid cell produces 2 haploid cells New cells are genetically identical to original cell (except for mutation)

58

Sexual Reproduction - The Human Life Cycle

Haploid gametes (n = 23)

During fertilization, sperm and ovum fuse forming a diploid zygote The zygote develops into an adult organism

Haploid (n) Diploid (2n)

Ovum (n)

Sperm Cell (n) MEIOSIS FERTILIZATION

Ovary

Testis

Diploid zygote (2n = 46)

Mitosis and development Multicellular diploid adults (2n = 46)

59

Spermatocytes to Spermatids

Primary spermatocytes undergo meiosis I, forming two haploid cells called secondary spermatocytes Secondary spermatocytes undergo meiosis II and their daughter cells are called spermatids Spermatids are small round cells seen close to the lumen of the tubule Late in spermatogenesis, spermatids are nonmotile Spermiogenesis spermatids lose excess cytoplasm and form a tail, becoming motile sperm

60

Spermatogenesis

Figure 27.8b, c

61

Oogenesis

Production of female sex cells by meiosis In the fetal period, oogonia (2n ovarian stem cells) multiply by mitosis and store nutrients Primordial follicles appear as oogonia are transformed into primary oocytes Primary oocytes begin meiosis but stall in prophase I From puberty, each month one activated primary oocyte completes meiosis one to produce two haploid cells The first polar body The secondary oocyte The secondary oocyte arrests in metaphase II and is ovulated If penetrated by sperm the second oocyte completes meiosis II, yielding: One large ovum (the functional gamete) A tiny second polar body
62

Oogenesis

63