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Mitosis
occurs
in
both
haploid
and
diploid
cells.
It
a
continuous
process,
but
for
purposes
of
discussion
it is usually divided into four cytologically distinguish table stages called prophase,
metaphase, anaphase, and telophase. In photographs show the
typical
chromosome
morphology
in
interphase
and
in
the four stages of mitosis in plant (onion root tip) and
animal
(newt
lung
epithelium)
cells.
Shows
the
four
stages
in
simplified
diagrams.
PROPHASE. At the beginning of prophase, the chromatids are very elongated
and cannot be seen under the Iight microscope. In preparation for mitosis they
begin to coil tightly so that they appear shorter and fatter under the microscope. By
late
prophase
each
chromosome,
which
was
duplicated
during the preceding S phase, is seen to consist of two
Sister chromatids.
During prophase, the mitotic spindle (spindle apparatus) assembles outside the
nucleus. Each of the spindle fibers in the bipolar mitotic spindle is approximately 25
mm diameter and consists of microtubules made of special proteins called tubulins.
In most animal cells, the foci for spindle assembly are the centrioles.
Prior to the S phase, the cell's pair of centrioles have replicated and each neu
centriole pair becomes focus for a radial array of microtubules called the aster.
Early
in
prophase
the
two
asters
are
adjacent
to
one
close
to
the
nuclear
membrane.
By
late
prophase
the
two
asters
are
far
apart
along
the
oufside
of
nucleus
and are spanned by the microtubular spindle fibers.
Near
the
end
of
prophase
the
nuclear
membrane
breaks down and the nucleolus or nucleoli disappear, allowing the spindle to area.
Specialized structures called kinetochores form on either face of the centromere of
each chromosome and become attached to special microtubules called
kinetochore microtubule. These microtubules radiate in opposite directions from
each side of each cromosome and interact with the spindle microtubules.
METAPHASE. Metaphase begins when the nuclear membrane has completely
disappeared.
During metaphase become arranged so that their centromeres become
aligned in one plane halfway between the two spindle poles and with the long axes
the chromosomes at 90 degrees to the spindle axis. The kinetochore microtubules
are
responsible
for
this
chromosome
alignment
event.
Thee plane where in chromosomes become aligned is called the metaphase plate.
Shows electron micrographs of human chromosomes at this stage of the cell cycle.
Note the highly condensed state of the Sister chromatids. The electron micrograph
(EM) in Figure 1.22c shows a human chromosome from which much of the protein
has been removed. In the center is a dense framework of protein called a scaffold,
which retains the form of the chromosome. The scaffold is surrounded by a halo of
DNA
filaments
that
have
uncoiled
and
spread
outward.
ANAPHASE. Anaphase is initiated when sister chromatids break apart ar the
centromere,
splitting
the
chromosome.
Once
the
paired
kinetochores
on
each
chromosome
separate,
the
Sister
chromatid
pairs
undergo
disjunction
(separation),
and
the
daughter
chromosomes
(as
each
sister
chromatid
is
now
called)
move
toward
the
poles.
In
anaphase
the
two
of
the
daughter
chromosomes
migrate
toward the opposite poles of the cell. Now the chromosomes assume the
characteristic shapes related to the location of the along the chromosome's
length.
For
example,
a
metacentric
chromosome
Will
appear as a V as the
two
roughly equal-length
chromosome
arms
trail
the
centromere
in
its
migration
toward
the
pole.
Similarly,
a
submetacentric
chromosome
Will
appear
as
a
J.
gives
an
interpretation
of
the
mitotic
apparatus
in
anaphase
in
an
animal
cell.
TELOPHASE.- During telophase the migration of daughter chromosomes to the
two poles is completed. The two sets of progeny chromosomes are assembled into
two groups ac, opposite ends of the cell. The chromosomes begin to uncoil and
assume the extended state characteristic of interphase.
A nuclear membrane forms around each chromosome group, the spindle
microtubules disappear, and the nucleolus or nucleoli re-form. At this point, the cell
has
nuclei.
CYTOKINESIS.- Cytokinesis refers to division of the cytoplasm. In most cases the
telophase stage of mitosis is accompanied by cytokinesis. Cytokinesis
compartmentalizes the two new nuclei into separare dauchter cells, completing the
mitotic cell division process.
GENETIC SIGNIFICANCE OF MITOSIS. Mitosis maintains the genetic cotent of a
cell from generation to generation. Mitosis occurs in haploid or diploid cells after
DNA and chromosome duplication has taken place. It is a highly ordered process in
which a duplicated chromosome set partitioned equally into the two daughter cells.
Thus, for haploid (N) cell, chromosome duplication produces a cell with two sers of
chromosomes. The ensuing mitosis result in two progeny haploid cell, each which
one sec of chromosomes.
For a diploid cell, which has two sets of chromosomes, chromosome duplication
produces a cell in which each chromosome set has doubled its content- The
ensuing mitosis results in two progeny diploid cells, each with two sets of
chromosomes.
more
extended
and
again
are
invisible
under
the
light
microscope.
progeny nuclei produced by the process is extremely large. Given the genetic
features of meiosis, this process is of critical importance for understanding the
behavior of genes, as Will be seen in the following chapters.
KYYNOTE
Meiosis occurs In all sexually reproducing, cukaryotes. It is a process by which a
diploid (ZN) cell or cell nucleus transformed, through one round of
chromosome
replication
and
two
rounds
of
nuclear
division, into four haploid (N) cells or nuclei. In the first of two divisions, pairing
synapsis of homologous chromosomes occurs. The meiotic process results in
conservation of the number of chromosomes are combined in the progeny nuclei
and by crossing- over( the physical exchange of genes between maternally or
paternally derived homologs)
ates
gci-to
which
tbrough
and
combincd
(the
in
paternal
ihc
physical
the
hromosomes
progeny
exchange
variotif
nuclei
of
gcnes
and
by
bctwecn
ways
arc
crossing-ovcr
matcrnally
spindle
Homoiogous
pairs
of
dupiicated
chromosomes
(tetrads)
line
up
1:-
on
VIRLSES.-
omparison
Mitosis
CELLS,
spindie
AND
CELLULAR
FKGRE
of
mitosis
REPRODUCTION
1.28
and
meiosis
in
diploid
ceil-
interphase
Meiosis
o
tnterphase
DNA
DNA
replication
(dupiication
not
yet
visible)
-f
Pairing
of
Prophase
Metaphase
Metaphase
Duplication
becomes
visible
homotogous
ch
romosomes
Middie
Prophase
Late
Prophase
FIGURE
Two
1.29
possible
chromosomes
division.
arrangements
on
Paternal
the
of
metaphase
chromosomes
two
pairs
piare
are
shown
of
the
in
homoiozous
first
veliow
me10tic
and
green;
maternal
wo
chromosomes
pairs.of
of
Possible
pairing
of
ME/osIS
are
diploid
animals
produce
ca(lv
to
gametes
produce
are
Gaieres
male
the
cess
calied
egg,
In
only
produced
male
tozoa)
primordial
sperm,
the
produced
germ
the
cells
goniae
Spermatogonia
dnnsion,
cells
testes.
of
in
mitotl-
The
garnete
called
pro-
is
the
and
28).
sperma-
contain
secondarv
which
the
(p.
spemzatogonia),
into
cvcle.
In
through
testes
the
life
cells.
1.30
the
Thus
the
(also
produce
each
fuse
Spermatogenesis
Figure
transform
(merocytes),
of
female
(primary
dary
sis
in
such
which
organism-
produced
The
In
divides
specialized
sperm
by
mitonc
then
oogenesis.
illustrated
through
matocyctes
the
animals
cycle.
nuclei
stages
in
Cycle
gametes,
diploid
haploid
by
ammals
are
new
formed
is
life
cheir
zygote
spermatogenesis.
are
their
haploid
The
only
gamete
oogenesis
of
Life
multicellular
when
the
the
are
Most
zygote
process-
chromosomes
the
produces
diploid
Lertilization
in
most
meiosis
zan-
arranaements
ANIMALS.
through
and
separation
Meiosis
IN
purple
homoiogous
Direction
-?ions
In
which,
spermato-
przmary
undergoes
spermeio-
spemzatocytes.
I
and
gives
nse
ro
rwo
the
secon
FEGURE
1.28
continucd
Anaphase
Singlechrornatid
ch
romosomes
EUKARYOiES
27
Celi
division
Tefophase
Celt
division
ll
Telophase
ji
Twin-chromatid
chromosomes
Cell
division
Anaphase
Metaphase
ll
thcsc
two
which
four
eventually
edifferentiat
gametes,
In
animals
germ
givc
Trise
into
Primary
oogcncsisincicsis
spermatids
into
-the
female
dial
haploid
to
T
and
ovary
cells
uncqual
contains
(primary
secondary
he
manire
male
spermacozoa
the
oocytcs,
the
arisc,
oogonia.
which
diploid,
primary
cytok-tnesls
by
These
grow
until
oocycc
to
thc
givc
primormitosis,
cells
thc
gocs
two
transform
end
of
through
cclls:
largc
onc
calicd
the
onc
calicd
thc
first
division
the
cciis.
One
polar
body;
tnto
thc
may
or
gamctc
polar
sccondary
is
the
is
cgg
ceil,
nor
CThus
in
(thc
body.
small
Othcr
may
In
the
the
sccond
CCII
Oniy.
produccd
Thc
animal
by
Ineiotic
sccond
rapidly
matures
firsr
ovum
polar
is
only
Sinall
haploid
callcd
that
the
very
two
is
ovum.
femate
is
and
large
or
and
produces
cell
divide.
ovum)
oocytc
oocyte
very
Inaturc
gatncte.
sccondary
viable
one
tueiosis
bodv
of
mature
a
diploid
sexually
repro-
cell.
hffEIOSiS
IN
ducillg
PLANTS.
The
typically
has
plants
life
two
Anaphase
phyte
haploid
or
are
structurc
Figure
1.31
both
mcns
pbascs,
aiid
thc
in
(p.
male
ptstils,
singlc
in
which
thc
diploid
ganactcs
stage
produccd
angiospenns,
thc
ing
stage
sporophytc
spores
In
of
gamcro
II
or
and"the
cycle
by
flowcring
which
in
Shows
and
female
reproductivc
Each
which
haploid
thc
Ilowcr
rcproduction
28)
respectiveiy.
produccd,
tmeiosis-
plants,
sexual
arc
gencratizcd
sta;ncn
stalk,
occurs.
flowcr
org:ujs,
containthc
cons:sts
thc
fiianlcnr,
the
01)
top
of
wh;ch
is
aa
sta01
anthcr.
Front
contains
of
the
the
pollen;
111c
female
stigma,
the
grows;
aniher
style,
and
are
gametophy.tes
sticky
surface
thin
at
sralk
thc
the
pollcn
grains,
and
typica*ly
specialized
to
rcccive
thc
poltcn
tubc
down
basc
chc
which
of
consists
thc
structure,
ovary,
within
which
femalc
gamctophyte
egg
cclf
Meiosis
male
the
rclcased
arc
is
ffower
ovules.
with
fcrtilizcd,
occurs
parts
the
of
in
the
occurs
thc
singlc
ovuic
spccific
flowcr.
as
Each
egg
Mciokis
in
cncloscs
Whcn
thc
CCII.
dcvclops
ways
follows.
ovule
thc
into
sccd.
fcnralc
and
in
rhc
fc:nale
Wirhin
thc
ovary
parr
of
of
the
1.32
Megasporogcncsis: Meiosis in thc fcmalc part o the flowcr and the production of
thc
crnbryo
sac.
Ovule
(includes
megaspore
mother
cett)
Embryo
O.
sac
Cytcpiasmic
division
Endosperm
mother
ceti
Megaspore
mother
ceil
(diploid)
Megagametophyte
Mitosis
ee
Meiosis
Meiosis
4
haploid
daughter
cells
Megaspore
Mitosis
3
disintegrate
Mitosis
loid
cclls
haploid,
and
diploid
and
the
cells:
The
sporopbytc
gametophyte
cclis
arc
cells
arc
diploide
SUMMARY
In
this
tures
of
duction
chapter
wc
viruses
and
in
eukaryotes,
nutosis
As
rcvicwcd
cells,
with
and
the
simplest
and
basic
organizational
discussed
particular
cellu(ar
cmphasis
fearcproon
meiosis.
cellular
organisms,
prokaryotes
lack
membrane-bound
fission.
The
genetic
material
less,
localized
within
Oid
region-
Prokarvotic
single,
circular
fcw
DNA
distributed
Ants
have
ernation
two
Of
gesh
Chc
sporcs
nts
the
tcly
bccome
Ediploid
spores,
e
f
of
reproductive
(Figure
and
also
alternation
meiosis
in
ovulc.
generation.
turn,
the
of
plants
the
Meiosis
these
gcneration
bcgins
In
flowering
that
Fertilization
ultiinitiates
diploid
sporocells
gametophytes.
alternate
relationship
generations
of
called
31).
produce
the
singlc-
haploid
the
are
bc
between
The
second
indrcates
'vvith
chrornosomcs.
cells
specialized
consists
p:
the
of
ntcmbranc-bound
meiosis.
are
in
is
1.35;
nucle-
is
phases,
gametophyte
produces
which,
linear
by
the
consists
transition
produced
SPOCOphytc
1.35
the
distinct
pollen
sporophyte
-Igure
sevcral
noncthc-
called
cukaryotcs
among
equivalents
gencration
of
binary
is,
tnay
havc
the
by
tbat
orgapisms
haploid
arc
arca
matcrial
matcrial
COnstitute
an
molecule
generations
fertilizati0n
in
and
gcnctic
divide
prokaryotcs
gcnct:c
multiccllular
Thc
and
of
cell
Eukaryotic
or
nucieus.
the
DNA
proteins-
ccllcd
nucleus
generation.
of
each
to
meiosis.
The
spores.
The
spores
vand
mosses,
thc
cclls
in
gymnosperms
(cone-
ants)
and
3-Ytcs,
and
fcmalc
us
angiospcrms
the
cells
in
gamctophytc
in
all
that
the
ovules
arc
grecn
produce
of
all
plants
thc
malc
flowers
produccd
alternation
that
by
of
gen-
gansctopayre
and
grccn
sporophvte
gencrar:ons
planrs.
Gametophyte
generation
Male
(N)
gametophyte
(microgametophyte
potlen)
Sperm
(N)
Femaie
gametophyte
(megagametophyte)
31
PROi'.i.F.M.S
h:
(hcsc
chrotneso;ucsvpicallv,
chc
DNA
is
cukarvot;c
co:nplcxcd
ce!ls
with
contain
protc:n.s.
c;thcr
one
or
haploid
cclls
division
in
mitosis-
Mitosis
foliowcd
by
tWO
sets
and
borli
the
haploid
invoive.s
onc
of
(attcr
and
011c
round
chrotnoso:ncs:
are
diploid
diploid
round
of
thc
cciis-
cclls
of
fornicr
DNA
nuclear
Nuctcar
occurs
by
celi
by
rephcatjon
division,
oftcn
accompanicd
arc
dvtston-
-r-hus,
rcsults
the
production
chromosomc
to
onc
of
daughtcr
numbcrs
anothcr
in
nuclci
and
and
that
to
the
that
arc
contain
idcntical
gcncrically
parcnt
nuclcns
identical
frotn
which
cheysarose.
In
all
occurs
sexuaMy
at
thc_proccss
or
bv
CCII
tron
haploid
uets
particular
which
nuclcus
and
two
ccits
of
ter0(tuCing
stage
a
or
meiosis
of
are
ccil
onc
In
the
lifc
round
Mciosts
haploid
01
division
many
mmosts
cyclc-
(ncvcr
nuclear
nuclci-
gencrates
thc
diploid
undcrgoes
rounds
in
organisnt.s,
1)INA
to
gamctcs-
genct:c
CCII)
replica-
producc
cukaryotcs
is
(he
four
prod-
Unllke-mitosis,
luciosis
variability
two
mant
(1)
wnys:
through
paternal
and
dcrivcd
various
chromosomes
(2)
and
recombtnant
some
the
are
through
Ovule
Anther
in
combined
crossing-ovcr
patcrnally
chromosomes
derived
which
in
:uatcrnal
progcny
bctwcen
htnnologs
with
to
and
genes-
nuclci;
producc
(N)
and
tuatcrnallv
tnatcrnal
patcrnat
Spores
Meiosis
ways
Egg
(N)
Fertilization
Zygote
(ZN)
Embryo
Mature
Sporophyte generation (2N)
plant