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Essential Cell Biology 5th edition

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660 CHAPTER 19 Sexual Reproduction and Genetics

Figure 19−13 Chiasmata help ensure

proper segregation of duplicated

homologs during the first meiotic

division. (A) In metaphase of meiosis I,

chiasmata created by crossing-over hold the

maternal and paternal homologs together.

At this stage, cohesin proteins keep the

sister chromatids glued together along

their entire length (see Figure 19–10). The

kinetochores of sister chromatids function as

a single unit in meiosis I, and microtubules

that attach to them point toward the same

spindle pole. (B) At anaphase of meiosis I,

the cohesins holding the arms of the

sister chromatids together are suddenly

degraded, allowing the homologs to be

separated. Cohesins at the centromere

continue to hold the sister chromatids

together as the homologs are pulled apart.

(A)

(B)

metaphase

of meiosis I

kinetochores

on the two

sister chromatids

function as

a single unit

anaphase

of meiosis I

chiasma

kinetochore microtubules

attached to sister

chromatids point in same

direction

ARMS OF SISTER CHROMATIDS

BECOME UNGLUED, ALLOWING THE

DUPLICATED HOMOLOGS TO SEPARATE

chiasmata resist this pulling (Figure 19–13A). In so doing, the chiasmata

help to position and stabilize bivalents at the metaphase plate.

In addition to the chiasmata, which hold the maternal and paternal

homologs together, cohesin proteins (described in Chapter 18) keep the

sister chromatids glued together along their entire length at meiosis I

(see Figure 19−10). At the start of anaphase I, the cohesin proteins that

ECB5 e19.13/19.13

hold the chromosome arms together are suddenly degraded. This release

allows the arms to separate and the recombined homologs to be pulled

apart (Figure 19−13B). If the arms were not released in this way, the

duplicated maternal and paternal homologs would remain tethered to

one another by the homologous DNA segments they had exchanged and

would not separate during anaphase I.

The Second Meiotic Division Produces Haploid

Daughter Nuclei

To separate the sister chromatids and produce cells with a haploid

amount of DNA, a second round of division, meiosis II, follows soon after

the first—without further DNA replication or any significant interphase

period. A meiotic spindle forms, and the kinetochores on each pair of

sister chromatids now attach to kinetochore microtubules that point in

opposite directions, as they would in an ordinary mitotic division. At anaphase

of meiosis II, the remaining, meiosis-specific cohesins—located at

the centromere—are degraded, and the sister chromatids are pulled apart

(Figure 19−14). The entire process is shown in Movie 19.1.

Haploid Gametes Contain Reassorted Genetic

Information

Even though they share the same parents, no two siblings are genetically

the same (unless they are identical twins). These genetic differences

are initiated long before sperm meets egg, when meiosis I produces two

kinds of randomizing genetic reassortment.

First, as we have seen, the maternal and paternal chromosomes are shuffled

and dealt out randomly during meiosis I. Although the chromosomes

are carefully distributed so that each gamete receives one and only one

copy of each chromosome, the choice between the maternal or paternal

homolog is made by chance, like the flip of a coin. Thus, each gamete

contains the maternal versions of some chromosomes and the paternal

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