Everything about Meiosis totally explained
In
biology or life science,
meiosis (pronounced mi-o-sis or me-o-sis) is a process of reduction division in which the number of chromosomes per cell is cut in half. In animals, meiosis always results in the formation of
gametes. The word "meiosis" comes from the Greek
meioun, meaning "to make small," since it results in a reduction in chromosome number in the gamete cell.
Meiosis is essential for
sexual reproduction and therefore occurs in all
eukaryotes (including single-celled organisms) that reproduce sexually. A few eukaryotes, notably the
Bdelloid rotifers, have lost the ability to carry out meiosis and have acquired the ability to reproduce by
parthenogenesis. Meiosis doesn't occur in
archaea or
bacteria, which reproduce via asexual processes such as
mitosis or
binary fission. Each cell has half the number of chromosomes as the parent cell.
During meiosis, the
genome of a
diploid germ cell, which is composed of long segments of
DNA packaged into
chromosomes, undergoes DNA replication followed by two rounds of division, resulting in four
haploid cells. Each of these cells contain one complete set of
chromosomes, or half of the genetic content of the original cell. If meiosis produces gametes, these cells must fuse during
fertilization to create a new diploid cell, or
zygote before any new growth can occur. Thus, the division mechanism of meiosis is a reciprocal process to the joining of two genomes that occurs at fertilization. Because the chromosomes of each parent undergo
genetic recombination during meiosis, each gamete, and thus each zygote, will have a unique genetic
blueprint encoded in its DNA. Together, meiosis and fertilization constitute sexuality in the eukaryotes, and generate genetically distinct individuals in populations.
In all plants, and in many protists, meiosis results in the formation of haploid cells that can divide vegetatively without undergoing fertilization. In these groups, gametes are produced by mitosis.
Meiosis uses many of the same biochemical mechanisms employed during
mitosis to accomplish the redistribution of chromosomes. There are several features unique to meiosis, most importantly the pairing and genetic recombination between
homologous chromosomes.
History
Meiosis was discovered and described for the first time in
sea urchin eggs in
1876, by noted German biologist
Oscar Hertwig (1849-1922). It was described again in
1883, at the level of chromosomes, by
Belgian zoologist
Edouard Van Beneden (1846-1910), in
Ascaris worms' eggs. The significance of meiosis for reproduction and inheritance, however, was described only in
1890 by
German biologist
August Weismann (1834-1914), who noted that two cell divisions were necessary to transform one diploid cell into four haploid cells if the number of chromosomes had to be maintained. In
1911 the
American geneticist
Thomas Hunt Morgan (1866-1945) observed
crossover in
Drosophila melanogaster meiosis and provided the first true genetics.
Evolution
Meiosis is thought to have appeared 1.4 billion years ago. The only supergroup of
eukaryotes which doesn't have meiosis in all organisms is
excavata. The other five major supergroups,
opisthokonts,
amoebozoa,
rhizaria,
archaeplastida and
chromalveolates all seem to have genes for meiosis universally present, even if not always functional. Some
excavata species do have meiosis which is consistent with the hypothesis that
excavata is an ancient,
paraphyletic grade. An example of eukaryotic organism in which meiosis doesn't exist is
euglenoid.
Occurrence of meiosis in eukaryotic life cycles
sexual reproduction, comprising of the constant cyclical process of meiosis and fertilization. This takes place alongside normal
mitotic cell division. In multicellular organisms, there's an intermediary step between the diploid and haploid transition where the organism grows. The organism will then produce the
germ cells that continue in the life cycle. The rest of the cells, called
somatic cells, function within the organism and will
die with it.
Cycling meiosis and fertilisation events produces a series of transitions back and forth between alternating haploid and diploid states. The organism phase of the life cycle can occur either during the diploid state (
gametic life cycle), or during the haploid state (
zygotic life cycle), or both (
sporic life cycle, in which there two distinct organism phases, one during the haploid state and the other during the diploid state). In this sense, there are three types of life cycles that utilize sexual reproduction, differentiated by the location of the organisms phase(s). In the
gametic life cycle, the species is diploid, grown from a diploid cell called the
zygote. In the
zygotic life cycle the species is haploid instead, spawned by the proliferation and differentiation of a single haploid cell called the
gamete. Humans, for example, are diploid creatures. Human stem cells undergo meiosis to create haploid gametes, which are
spermatozoa for males or
ova for females. These gametes then fertilize in the
Fallopian tubes of the female, producing a diploid zygote. The zygote undergoes progressive stages of mitosis and
differentiation, turns into a
blastocyst and then gets implanted in the
uterus endometrium to create an
embryo.
In the
gametic life cycle, of which humans are a part, the living organism is diploid in nature. Here, we'll generalize the example of human reproduction stated previously. The organism's diploid germ-line stem cells undergo meiosis to create haploid gametes, which fertilize to form the zygote. The diploid zygote undergoes repeated cellular division by
mitosis to grow into the organism. Mitosis is a related process to meiosis that creates two cells that are genetically identical to the parent cell. The general principle is that mitosis creates somatic cells and meiosis creates germ cells.
In the
zygotic life cycle, the living organism is haploid. Two organisms of opposing gender contribute their haploid germ cells to form a diploid zygote. The zygote undergoes meiosis immediately, creating four haploid cells. These cells undergo
mitosis to create the organism. Many
fungi and many
protozoa are members of the zygotic life cycle.
Finally, in the
sporic life cycle, the living organism alternates between haploid and diploid states. Consequently, this cycle is also known as the
alternation of generations. The diploid organism's germ-line cells undergo meiosis to produce gametes. The gametes proliferate by mitosis, growing into a haploid organism. The haploid organism's germ cells then combine with another haploid organism's cells, creating the zygote. The zygote undergoes repeated mitosis and differentiation to become the diploid organism again. The sporic life cycle can be considered a fusion of the gametic and zygotic life cycles.
Process
Because meiosis is a "one-way" process, it can't be said to engage in a
cell cycle as mitosis does. However, the preparatory steps that lead up to meiosis are identical in pattern and name to the interphase of the mitotic cell cycle.
Interphase is divided into three phases:
- Growth 1 (G1) phase: Immediately follows cytokinesis. This is a very active period, where the cell synthesizes its vast array of proteins, including the enzymes and structural proteins it'll need for growth. In G1 stage each of the 46 human chromosomes consists of a single (very long) molecule of DNA. At this point cells are 46,2N, identical to somatic cells.
- Synthesis (S) phase: The genetic material is replicated: each of its chromosomes duplicates (46,2N). The cell is still diploid, however, because it still contains the same number of centromeres. However, the identical sister chromatids are in the chromatin form because spiralisation and condensation into denser chromosomes have not taken place yet. It will take place in prophase I in meiosis.
- Growth 2 (G2) phase: G2 phase is absent in Meiosis
Interphase is immediately followed by meiosis I and meiosis II. Meiosis I consists of
segregating the
homologous chromosomes from each other, then dividing the diploid cell into two haploid cells each containing one of the segregates. Meiosis II consists of decoupling each chromosome's sister strands (
chromatids), segregating the DNA into two sets of strands (each set containing one of each homologue), and dividing both haploid, duplicated cells to produce four haploid, unduplicated cells. Meiosis I and II are both divided into
prophase,
metaphase,
anaphase, and
telophase subphases, similar in purpose to their analogous subphases in the mitotic cell cycle. Therefore, meiosis encompasses the interphase (G
1, S, G
2), meiosis I (prophase I, metaphase I, anaphase I, telophase I), and meiosis II (prophase II, metaphase II, anaphase II, telophase II).
Meiosis generates genetic diversity in two ways: (1) independent assortment of chromosomes at both of the meiotic divisions allows genetic differences among gametes; and (2) physical exchange of chromosomal regions by homologous recombination during prophase I results in new genetic combinations within chromosomes.
Meiosis I
In meiosis I, the chromosomes in a diploid cell separate, producing two diploid cells (
23, N).
Prophase I
Homologous chromosomes pair and form synapsis, a step unique to meiosis. The paired chromosomes are called bivalents or tetrads, which have two chromosomes and four chromatids, with one chromosome coming from each parent. At this stage, non-sister chromatids may cross-over at points called chiasmata.
Leptotene
The first stage of prophase I is the
leptotene stage, also known as
leptonema, from Greek words meaning "thin threads." During this stage, individual chromosomes begin to condense into long strands within the nucleus. However the two sister chromatids are still so tightly bound that they're indistinguishable from one another.
The chromosomes in the leptotene stage show a specific arrangement where the telomeres are oriented towards the nuclear membrane. Hence this stage is called, "bouquet stage".
Zygotene
The
zygotene stage, also known as
zygonema, from Greek words meaning "paired threads,"
Microtubules that attach to the kinetochores are known as
kinetochore microtubules. Other microtubules will interact with microtubules from the opposite centriole. These are also
nonkinetochore microtubules.
Meiosis-phases
Metaphase I
Homologous pairs move together along the phase plate:
as
kinetochore microtubules from both centrioles attach to their respective kinetochores, the homologous chromosomes align along an equatorial plane that bisects the spindle, due to continuous counterbalancing forces exerted on the bivalents by the microtubules emanating from the two kinetochores of homologous chromosomes. The physical basis of the independent assortment of chromosomes is the random orientation of each bivalent along the metaphase plate.
Anaphase I
Kinetochore microtubules shorten, severing the recombination nodules and pulling homologous chromosomes apart. Since each chromosome only has one functional unit of a pair of kinetochores. Polyploidy is poorly tolerated in animal species. Plants, however, regularly produce fertile, viable polyploids. Polyploidy has been implicated as an important mechanism in plant speciation.
Most importantly, however, meiosis produces genetic variety in gametes that propagate to offspring. Recombination and independent assortment allow for a greater diversity of genotypes in the population. As a system of creating diversity, meiosis allows a species to maintain stability under environmental changes.
Nondisjunction
The normal separation of chromosomes in Meiosis I or sister chromatids in meiosis II is termed
disjunction. When the separation isn't normal, it's called
nondisjunction. This results in the production of gametes which have either more or less of the usual amount of genetic material, and is a common mechanism for
trisomy or
monosomy. Nondisjunction can occur in the meiosis I or meiosis II, phases of cellular reproduction, or during
mitosis.
This is a cause of several medical conditions in humans:
Down Syndrome - trisomy of chromosome 21
Patau Syndrome - trisomy of chromosome 13
Edward Syndrome - trisomy of chromosome 18
Klinefelter Syndrome - extra X chromosomes in males - ie XXY, XXXY, XXXXY
Turner Syndrome - atypical X chromosome dosage in females - ie XO, XXX, XXXX
XYY Syndrome Supermale - an extra Y chromosome in males
Meiosis in humans
In females, meiosis occurs in cells known as oogonia (singular: oogonium). Each oogonium that initiates meiosis will divide twice to form a single oocyte and three polar bodies. However, before these divisions occur, these cells stop at the diplotene stage of meiosis I and lay dormant within a protective shell of somatic cells called the follicle. Follicles begin growth at a steady pace in a process known as folliculogenesis, and a small number enter the menstrual cycle. Menstruated oocytes continue meiosis I and arrest at meiosis II until fertilization. The process of meiosis in females occurs during oogenesis, and differs from the typical meiosis in that it features a long period of meiotic arrest known as the Dictyate stage and lacks the assistance of centrosomes.
In males, meiosis occurs in precursor cells known as spermatogonia that divide twice to become sperm. These cells continuously divide without arrest in the seminiferous tubules of the testicles. Sperm is produced at a steady pace. The process of meiosis in males occurs during spermatogenesis.
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