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Essential Cell Biology 5th edition

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Meiosis and Fertilization

657

(A)

MITOSIS

P1

M2

M1

(B)

MEIOSIS I

M1 P1

P2 M2

Figure 19−7 During meiosis I, duplicated

homologous chromosomes pair before

lining up on the meiotic spindle. (A) In

mitosis, the individual duplicated maternal

(M) and paternal (P) chromosomes line up

independently at the metaphase plate; each

consists of a pair of sister chromatids, which

will separate just before the cell divides.

(B) By contrast, in meiosis, duplicated

maternal and paternal homologs pair before

lining up at the metaphase plate. The

maternal and paternal homologs separate

during the first meiotic division, and the

sister chromatids separate during meiosis II.

The mitotic and meiotic spindles are shown

in green.

P2

duplicated homologous chromosomes

line up independently at the

metaphase plate

duplicated homologous chromosomes

pair before lining up at the

metaphase plate

Duplicated Homologous Chromosomes Pair During

Meiotic Prophase

Before a eukaryotic cell divides—by either meiosis or mitosis—it first

duplicates all of its chromosomes. The twin copies of each duplicated

chromosome, called sister

ECB5

chromatids,

E19.08/19.07

are tightly linked along their

length. The way these duplicated chromosomes are handled subsequently,

however, differs between meiosis and mitosis. In mitosis, as we

discuss in Chapter 18, the duplicated chromosomes line up, single file, at

the metaphase plate (Figure 19−7A). They are then segregated into the

two daughter nuclei.

In meiosis, however, the need to halve the number of chromosomes introduces

an extra demand on the cell-division machinery. The germ-line

cell must keep track of the maternal and paternal homologs—to ensure

that each of the four haploid cells produced by meiosis will receive a

single sister chromatid from each chromosome set. Meiosis therefore

begins with a complex and time-consuming process called pairing, in

which duplicated homologs are brought together during a stage called

meiotic prophase (or prophase I). It is these pairs of duplicated homologs

that line up at the metaphase plate in meiosis I (Figure 19−7B). Each

pairing forms a structure called a bivalent, in which all four sister chromatids

stick together until the cell is ready to divide (Figure 19−8).

The maternal and paternal homologs will separate during meiosis I,

and the individual sister chromatids will separate during meiosis II.

How the homologs (and the two sex chromosomes) recognize each other

during pairing is still not fully understood. In many organisms, the initial

association depends on an interaction between matching maternal

and paternal DNA sequences at numerous sites that are widely dispersed

along the homologous chromosomes. Once formed, bivalents are very

stable: they remain associated throughout meiotic prophase, a stage that

in some organisms can last for years.

duplicated

paternal

chromosome

duplicated

maternal

chromosome

centromere

sister

chromatids

Crossing-Over Occurs Between the Duplicated Maternal

and Paternal Chromosomes in Each Bivalent

The picture of meiosis we have just painted is greatly simplified, in that

it leaves out a crucial feature. In sexually reproducing organisms, the

pairing of the maternal and paternal chromosomes is accompanied by

bivalent

Figure 19−8 Duplicated maternal and

paternal chromosomes pair during

meiosis I to form bivalents. Each bivalent

ECB5 e19.09/19.08

contains four sister chromatids.

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