Mitosis and Meiosis
I.
These two processes function to pass chromosomes
from one cellular generation to the next in a very carefully controlled manner.
II.
Mitosis and Meiosis are both correctly described
as nuclear division; they are never correctly called cell
division, or any kind of reproduction. It is possible (and often quite normal)
for nuclei to divide when cells don't. And organisms reproduce; nuclei and
cells divide.
III.
Mitosis
A.
Mitosis is the division of a nucleus to produce
two genetically identical daughter nuclei.
B.
Mitosis is utilized for any function which
requires the production of more cells with identical genetic information. These
processes include the vase majority of cell production during the growth of an
organism, the cell division needed for healing and repair, and the division of
nuclei when an organism is in the process of asexual reproduction. Note:
Mitosis is not asexual reproduction, nor can it be called asexual cell (or even
nuclear) division.
C.
Because the vast majority of the cells in a
multicellular organism were produced by mitotic cell division, those cells all
have identical nuclei. They obviously don't all look or function alike. Cells
mature through a process called differentiation in which select sets of
genes are turned on and off, resulting in changes in the structure and function
of the cell.
IV.
Meiosis
A.
Meiosis is the division of one diploid (usually)
nucleus to produce four haploid (usually) nuclei, all genetically different.
Though the vast majority of the time the chromosome number reduction is from
diploid to haploid, in some cases it may be from, say, hexaploid (eg., wheat)
to triploid.
B.
Meiosis performs a key task necessary in a sexual
life cycle. Since fertilization (which is the actual sexual event in the life
cycle) automatically doubles chromosome number by combining the chromosomes of
an egg and a sperm, it is essential that some event occur somewhere in the life
cycle which reduced the chromosome number to compensate.
C.
Though Meiosis is part of a sexual life cycle, it is
not ever correctly described as, for instance, "sexual cell
division," and certainly not as "sexual reproduction." It is the
life cycle, and specifically the fertilization event, which constitute sexual
activity, not Meiosis.
D.
In animal life cycles, the meiotic cell division
in the life cycle immediately precedes the development of gametes (eggs and
sperm). However, this need not at all be the case. Plants have a somewhat
different sexual life cycle from animals which includes all of the same events,
including Meiosis, production of gametes, and fertilization, but also includes
an additional phase between Meiosis and gamete production.
V.
There are three differences between what Mitosis
accomplishes and what Meiosis accomplishes.
A.
Mitosis divides one nucleus into two; Meiosis
divides one nucleus into four.
B.
Mitosis conserves chromosome number; Meiosis
reduces it in half (usually from diploid to haploid).
C.
Mitosis produces genetically identical daughter
nuclei; Meiosis produces genetically different daughter nuclei.
VI.
Errors in nuclear division can produce chromosome
anomalies.
A.
Nuclei whose chromosomes do not form normal sets
(eg, which have extra or missing chromosomes in one or more of the chromosome
sets) are aneuploid.
1.
Aneuploidies typically result from nondysjunction
in either Meiosis I or Meiosis II.
2.
If a nucleus which should be diploid has three of
a chromosome instead of the normal two, that nucleus is trisomic. For
example, if the nucleus of a human cell has three of chromosome 18, that would
be trisomy 18. This term is also used for an organism, all of whose cells are
trisomic. Trisomy 21 can refer to a human, all of whose cells have three
twenty-first chromosomes instead of the expected two.
3.
If a nucleus which should be diploid has only one
of a chromosome in stead fo the normal two, that nucleus is monosomic.
An individual may be described as monosomic if all (or most) of the cells in
that individual are monosomic. For example, Turner's Syndrome female humans are
monosomic for the X chromosome.
4.
Other things being equal, it is significantly
worse to be missing chromosome material than to have extra chromosome
materials. Of course, the specific genes on the extra or missing chromosome
material also impact on how serious the effects of the anomaly will be.
B.
A nucleus which has extra sets of chromosomes
(like the wheat mentioned above) is polyploid.
1.
Polyploidy typically results from fertilization
inv0lving gametes with unreduced chromosome number. In other words, an error in
chromosome reduction during Meiosis produces a gamete which still has the
diploid number of chromosome sets, which then participates in fertilization
with a haploid gamete.
2.
An autopolyploid has more than two sets of
chromosomes from the same species. In other words, it has more than two
homologous sets of chromosomes. Commercially grown bananas are autotriploids;
they have three sets of banana chromosomes. Autopolyploids frequently have
difficulty performing Meiosis because the pairing mechanism in Prophase I
requires one-on-one pairing between two partners, and having more than two
homologues in the same nucleus creates confused pairing.
3.
An allopolyploid has chromosomes from more
than one species. Wheat is an allohexaploid. It has two each of the chromosome
sets from three different species of grasses, so each nucleus has six sets of
chromosomes, but each chromosome has only one homologous partner in the
nucleus. For this reason, allopolyploids are often perfectly fertile, as wheat
obviously is. Allopolyploids are generally formed by an accidental
fertilization between two different, closely related species (producing an
offspring which is technically diploid, but whose chromosomes don't match--a
mule is an example of this situation). In plants, it is frequently possible for
the plant to reproduce asexually, so the inability to perform Meiosis may not
be a problem which prevents survival and reproduction. (This is why commercial
bananas can reproduct successfully, despite being unable to do Meiosis.) Our
inter-species hybrid can thus generally reproduce and thrive. Eventually, an
accidental chromosome doubling event can double both chromosome sets, producing
a plant which is technically tetraploid (having four sets of chromosomes) but
functionally diploid (as each chromosome has only one homologous partner in the
nucleus). This restores the ability to reproduce sexually, and also creates a
brand new species. Clearly, this happens. It's happened twice in the natural
history of Triticum (wheat).
C.
Sometimes chromosomes lose segments, or acquire
extra copies of segments. These are called insertions and deletions,
or duplications and deficiencies. These generally arise due to
uneven crossovers during Prophase I of Meiosis. They can result from crossing
over in inversion heterozygotes.
D.
Occasionally, a segment of a chromosome will break
free and accidentally reattach with its ends switched around. This reversed
segment of the chromosome is an inversion.
1.
If the cell is an inversion homozygote,
which means that both chromosomes of a homologous pair carry the same
inversion, this causes no problems for the organism (though it might lead to
fertility problems with offspring, which could very easily be inversion
heterozygotes).
2.
If a cell has one chromosome of a pair which
carries an inversion, but the partner doesn't, the cell is an inversion
heterozygote. Inversion heterozygotes are, themselves, perfectly healthy,
but in Meiosis the inversion can create several problems.
a.
In Prophase I of Meiosis, chromosomes synapse on a
gene-by-gene basis. So in the region of the inversion, when the homologous
chromosomes pair they create an inversion loop as the frontward/backward
segments attempt to pair normally. This isn't, in itself, a problem, but if a
crossover occurs within the inversion loop, it will lead to duplication and
deficiency.
b.
If the inverted region does not include the
centromere of the chromosome, and a crossover occurs within the inverted
region, it will lead to one dicentric chromatid (a chromatid which has
two centromeres) and one acentric chromatid (a chromatid which has no
centromere). As the pull of spindle fibers on the centromeres is what moves
chromosomes around during Meiosis and Mitotis, this typically leads to
disaster.
E.
Sometimes, a segment of one chromosome gets
accidentally broken off and attached to a different chromosome. This is a translocation.
1.
If no genes are lost or damaged by the
translocation, and all genes are present in the correct quantity (generally
two), the result is a balanced translocation, which causes no problems
for the individual possessing the translocation, though it may cause problems
in gamete production and for the offspring.
2.
If the translocation involves trading pieces of
both chromosomes involved, it's called a reciprocal translocation.
3.
As with inversions, translocations may be homozygous
or heterozygous. A translocation homozygote has no problems; he
or she simply has an unusual arrangement of genes on his or her chromosomes,
but everything works perfectly.
4.
A translocation heterozygote has problems
in synapsis, just as the inversion heterozygote does, because the chromosomes
try to pair gene-to-gene, and the genes are located on different chromosomes in
the two homologous pairs. Again, crossing over in the wrong place can lead to
duplication and deficiencies, and to dicentric and acentric chromosomes. Note
that an individual can be a translocation heterozygote and still have a
perfectly balanced gene complement.
5.
Even if no unfortunate crossing over occurs, the
outcome of a fertilization between someone carrying a balanced translocation
and someone with the more usual chromosome complement can cause problems for
the offspring. The zygote will receive a normally arranged set of chromosomes
from one gamete, but may receive a translocated chromosome (but not its
reciprocal partner) from the other. A rare version of Down's Syndrome (which
generally results from trisomy of the twenty-first chromosome) results from
this sort of problem. The zygote receives two normal twenty-first chromosomes,
plus one normal fourteenth chromosome and one fourteenth chromosome which is
carrying a translocated copy of most of the twenty-first chromosome. The result
is a genome which has the appropriate number of chromosomes, but in which one
of the fourteenth chromosomes also carries a copy of the twenty-first
chromosome. Thus the child functionally has three copies of chromosome
twenty-one, and all of the normal characteristics of Down's Syndrome.
F.
Note that chromosome rearrangements like
inversions and translocations have important impact on speciation--the division
of one species into two. What distinguishes one species from another is the
inability to reproduce and produce fertile offspring, and if chromosome
rearrangements arise and become "fixed" within a section of a
species, that subgroup can easily become unable to reproduce with the original,
parent species. This is one of the things that happens when segments of a
species become isolated from other segments. Comparing the chromosomes of
closely related species shows us that this is a very significant way in which
they differ from each other. This is why, for example, a horse can breed with a
donkey (different, but closely related species) and produce a completely
healthy, even robust, hybrid: a mule. Mules are strong and healthy because the
chromosomes of the donkey and those of the horse carry very complementary kinds
of genes, and the two sets of genetic influences interact very well, with
little missing. However, the mule is sterile, because his horse chromosomes and
his donkey chromosomes have significant differences in arrangement (inversions
and translocations) and thus are unable to complete Meiosis.
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