Mitosis is the way that in which a cell duplicates its chromosomes to generate two identical nuclei. It is generally followed by cytokinesis which divides the cytoplasm and cell membrane. This results in two identical cells with an equal distribution of organelles and other cellular components. Mitosis and cytokinesis jointly define the mitotic (M) phase of the cell cycle, the division of the mother cell into two sister cells, each with the genetic equivalent of the parent cell. Mitosis occurs most often in eukaryotic cells.
In multicellular organisms, the somatic cells undergo mitosis, while germ cells — cells destined to become sperm in males or ova in females — divide by a related process called meiosis. Prokaryotic cells, which lack a nucleus, divide by a process called binary fission.Process
The process of mitosis is complex and highly regulated. The sequence of events is organized into phases corresponding to the completion of certain phase prerequisites.
Various classes of cells can skip steps depending on the presence of protease inhibitors or if they are in a hurry. These stages are preprophase, prophase, prometaphase, metaphase, anaphase, telophase and cytokinesis. During the process of mitosis the pairs of chromosomes condense and attach to fibers that pull the sister chromatids to opposite sides of the cell. The cell then divides in cytokinesis, to produce two identical daughter cells.
Cytokinesis usually occurs in conjunction with mitosis, "mitosis" is often used interchangeably with the phrase "mitotic phase". However, there are many cells whose mitosis and cytokinesis occur separately, forming single cells with multiple nuclei. This occurs most notably among the fungi and slime moulds, but is found in various different groups. Even in animals, cytokinesis and mitosis may occur independently, for instance during certain stages of fruit fly embryonic development.[1] Errors in mitosis can either kill a cell through apoptosis or cause mutations that may lead to cancer or cell death.
A cell in late metaphase. All chromosomes (blue) but one have arrived at the metaphase plate.
[edit] Phases
The process of Mitosis can be divided into seven stages: Preprophase, Prophase, Prometaphase, Metaphase, Anaphase, Telophase and Cytokinesis. Students often remember the main stages of the cell cycle (Interphase, Prophase, Metaphase, Anaphase and Telophase) by use of the acronym IPMAT.
[edit] Interphase
Main article: Interphase
The cell cycle.The mitotic phase is a relatively short period of the cell cycle. It alternates with the much longer interphase, where the cell prepares itself for cell division. Interphase is divided into three phases, G1 (first gap), S (synthesis), and G2 (second gap). During all three phases, the cell grows by producing proteins and cytoplasmic organelles. However, chromosomes are replicated only during the S phase. Thus, a cell grows (G1), grows as it duplicates its chromosomes (S), grows more and prepares for mitosis (G2), and divides (M).[2]
[edit] Preprophase
Main article: Preprophase
In plant cells only, prophase is preceded by a pre-prophase stage and followed by a post-prophase stage. In plant cells that are highly vacuolated and somewhat amorphoric, the nucleus has to migrate into the center of the cell before mitosis can begin. This is achieved through the formation of a phragmosome, a transverse sheet of cytoplasm that bisects the cell along the future plane of cell division. In addition to phragmosome formation, preprophase is characterized by the formation of a ring of microtubules and actin filaments (called preprophase band) underneath the plasmamembrane around the equatorial plane of the future mitotic spindle and predicting the position of cell plate fusion during telophase. The cells of higher plants (such as the flowering plants) lack centrioles. Instead, spindle microtubules aggregate on the surface of the nuclear envelope during prophase. The preprophase band disappears during nuclear envelope disassembly and spindle formation in prometaphase.[3]
[edit] Prophase
Prophase: The two round objects above the nucleus are the centrosomes. Note the condensed chromatin.Main article: Prophase
Normally, the genetic material in the nucleus is in a loosely bundled coil called chromatin. At the onset of prophase, chromatin condenses together into a highly ordered structure called a chromosome. Since the genetic material has already been duplicated earlier in S phase, the replicated chromosomes have two sister chromatids, bound together at the centromere by the cohesion complex. Chromosomes are visible at high magnification through a light microscope.
Close to the nucleus are two centrosomes. Each centrosome, which was replicated earlier independent of mitosis, acts as a coordinating center for the cell's microtubules. The two centrosomes nucleate microtubules (or microfibrils) (which may be thought of as cellular ropes) by polymerizing soluble tubulin present in the cytoplasm. Molecular motor proteins create repulsive forces that will push the centrosomes to opposite side of the nucleus. The centrosomes are only present in animals. In plants the microtubules form independently.
Some centrosomials contain a pair of centrioles that may help organize microtubule assembly, but they are not essential to formation of the mitotic spindle.[4]
[edit] Prometaphase
Prometaphase: The nuclear membrane has degraded, and microtubules have invaded the nuclear space. These microtubules can attach to kinetochores or they can interact with opposing microtubules.Main article: Prometaphase
The nuclear envelope disassembles and microtubules invade the nuclear space. This is called open mitosis, and it occurs in most multicellular organisms. Fungi and some protists, such as algae or trichomonads, undergo a variation called closed mitosis where the spindle forms inside the nucleus or its microtubules are able to penetrate an intact nuclear envelope.[5][6]
Each chromosome forms two kinetochores at the centromere, one attached at each chromatid. A kinetochore is a complex protein structure that is analogous to a ring for the microtubule hook; it is the point where microtubules attach themselves to the chromosome.[7] Although the kinetochore structure and function are not fully understood, it is known that it contains some form of molecular motor.[8] When a microtubule connects with the kinetochore, the motor activates, using energy from ATP to "crawl" up the tube toward the originating centrosome. This motor activity, coupled with polymerisation and depolymerisation of microtubules, provides the pulling force necessary to later separate the chromosome's two chromatids.[8]
When the spindle grows to sufficient length, usually at least 7 nanometers, kinetochore microtubules begin searching for kinetochores to attach to. A number of nonkinetochore microtubules find and interact with corresponding nonkinetochore microtubules from the opposite centrosome to form the mitotic spindle.[9] Prometaphase is sometimes considered part of prophase.
[edit] Metaphase
Metaphase: The chromosomes have aligned at the metaphase plate.Main article: Metaphase
As microtubules find and attach to kinetochores in prometaphase, the centromeres of the chromosomes convene along the metaphase plate or equatorial plane, an imaginary line that is equidistant from the two centrosome poles.[9] This even alignment is due to the counterbalance of the pulling powers generated by the opposing kinetochores, analogous to a tug-of-war between equally strong people. In certain types of cells, chromosomes do not line up at the metaphase plate and instead move back and forth between the poles randomly, only roughly lining up along the midline. Metaphase comes from the Greek word for "metanosis" μεÏα meaning "after."
Because proper chromosome separation requires that every kinetochore be attached to a bundle of microtubules (spindle fibers) , it is thought that unattached kinetochores generate a signal to prevent premature progression to anaphase[1] without all chromosomes being aligned. The signal creates the mitotic spindle checkpoint.[10]
[edit] Anaphase
Early anaphase: Kinetochore microtubules shortenMain article: Anaphase
When every kinetochore is attached to a cluster of microtubules and the chromosomes have lined up along the metaphase plate, the cell proceeds to anaphase (from the Greek ανα meaning “up,” “against,” “back,” or “re-”).
Two events then occur; First, the proteins that bind sister chromatids together are cleaved, allowing them to separate. These sister chromatids are hereafter independent sister chromosomes. They are pulled apart by shortening kinetochore microtubules and toward the respective centrosomes to which they are attached. This is followed by the elongation of the nonkinetochore microtubules, which pushes the centrosomes (and the set of chromosomes to which they are attached) apart to opposite ends of the cell.
These three stages are sometimes called early, mid and late anaphase. Early anaphase is usually defined as the separation of the sister chromatids. Mid anaphase occurs with the reunification of certain metastic chromatids. Late anaphase is the elongation of the microtubules and the microtubules being pulled further apart. At the end of anaphase, the cell has succeeded in separating identical copies of the genetic material into two distinct populations.
[edit] Telophase
Telophase: The decondensing chromosomes are surrounded by nuclear membranes. Note cytokinesis has already begun, the pinching is known as the cleavage furrow.Main article: Telophase
Telophase (from the Greek ÏÎµÎ»Î¿Ï meaning "end") is a reversal of prophase and prometaphase events. It "cleans up" the after effects of mitosis. At telophase, the nonkinetochore microtubules continue to lengthen, elongating the cell even more. Corresponding sister chromosomes attach at opposite ends of the cell. A new nuclear envelope, using fragments of the parent cell's nuclear membrane, forms around each set of separated sister chromosomes. Both sets of chromosomes, now surrounded by new nuclei, unfold back into chromatin. Mitosis is complete, but cell division is not yet complete.
[edit] Cytokinesis
Main article: Cytokinesis
Cytokinesis is often mistakenly thought to be the final part of telophase, however cytokinesis is a separate process that begins after telophase. Cytokinesis is technically not even a phase of mitosis, but rather a separate process, necessary for completing cell division. In animal cells, a cleavage furrow (pinch) containing a contractile ring develops where the metaphase plate used to be, pinching off the separated nuclei.[11] In both animal and plant cells, cell division is also driven by vesicles derived from the Golgi apparatus, which move along microtubules to the middle of the cell. [12] In plants this structure coalesces into a cell plate at the center of the phragmoplast and develops into a cell wall, separating the two nuclei. The phragmoplast is a microtubule structure typical for higher plants, whereas some green algae use a phycoplast microtubule array during cytokinesis.[13] Each daughter cell has a complete copy of the genome of its parent cell. The end of cytokinesis marks the end of the M-phase.
[edit] Consequences of errors
Although errors in mitosis are rare, the process may go wrong, especially during early cellular divisions in the zygote. Mitotic errors can be especially dangerous to the organism because future offspring from this parent cell will carry the same disorder.
In non-disjunction, a chromosome may fail to separate during anaphase. One daughter cell will receive both sister chromosomes and the other will receive none. This results in the former cell having three chromosomes coding for the same thing (two sisters and a homologue), a condition known as trisomy, and the latter cell having only one chromosome (the homologous chromosome), a condition known as monosomy. These cells are considered aneuploidic cells and these abnormal cells can cause cancer.[14]
Mitosis is a traumatic process. The cell goes through dramatic changes in ultrastructure, its organelles disintegrate and reform in a matter of hours, and chromosomes are jostled constantly by probing microtubules. Occasionally, chromosomes may become damaged. An arm of the chromosome may be broken and the fragment lost, causing deletion. The fragment may incorrectly reattach to another, non-homologous chromosome, causing translocation. It may reattach to the original chromosome, but in reverse orientation, causing inversion. Or, it may be treated erroneously as a separate chromosome, causing chromosomal duplication. The effect of these genetic abnormalities depend on the specific nature of the error. It may range from no noticeable effect, cancer induction, or organism death.
[edit] Endomitosis
Endomitosis is a variant of mitosis without nuclear or cellular division, resulting in cells with many copies of the same chromosome occupying a single nucleus. This process may also be referred to as endoreduplication and the cells as endoploid.[1]
In biology, meiosis (IPA: /maɪËÉÊsɪs/) is the process by which one diploid eukaryotic cell divides to generate four haploid cells often called gametes. The word "meiosis" comes from the Greek meioun, meaning "to make smaller," since it results in a reduction in chromosome number in the gamete cell. Among fungi, spores in which the haploid nuclei are at first disseminated are called meiospores, or more specifically, ascospores in asci (Ascomycota) and basidospores on basidia (Basidiomycota).
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 does not occur in archaea or bacteria, which reproduce via asexual processes such as mitosis or binary fission.
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 haploid cells called gametes. Each gamete contains one complete set of chromosomes, or half of the genetic content of the original cell. These resultant haploid cells can fuse with other haploid cells of the opposite sex or mating type during fertilization to create a new diploid cell, or zygote. 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. In other words, meiosis and sexual reproduction produce genetic variation.
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 genetic interpretation of meiosis.
[edit] Occurrence of meiosis in eukaryotic life cycles
Gametic life cycle.
Zygotic life cycle.
Sporic life cycle.Main article: Biological life cycle
Meiosis occurs in all eukaryotic life cycles involving sexual reproduction, comprising of the constant cyclical process of meiosis and fertilization. This takes place alongside normal mitotic cell division. In multicellular organisms, there is 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 will 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, and indeed its diagram supports this conclusion.
[edit] Process
Because meiosis is a "one-way" process, it cannot 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 cytogenesis. This is a very active period, where cell synthesizes its vast array of proteins, including the enzymes and structural proteins it will need for growth. In G1 stage each of the 46 human chromosomes consists of a single (very long) molecule of DNA
Synthesis (S) phase: The genetic material is replicated: each of its chromosomes duplicates. 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: The cell continues to grow making the cell larger.
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 (G1, S, G2), meiosis I (prophase I, metaphase I, anaphase I, telophase I), and meiosis II (prophase II, metaphase II, anaphase II, telophase II).
[edit] Meiosis I
[edit] Prophase I
[edit] Leptotene
The first stage of prophase I is the leptotene stage, also known as leptonema, from Greek words meaning "thin threads."[1] 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 are indistinguishable from one another.
[edit] Zygotene
The zygotene stage, also known as zygonema, from Greek words meaning "paired threads,"[1] occurs as the chromosomes approximately line up with each other into homologous chromosomes. The combined homologous chromosomes are said to be bivalent. They may also be referred to as a tetrad, a reference to the four sister chromatids. The two chromatids become "zipped" together, forming the synaptonemal complex, in a process known as synapsis.
[edit] Pachytene
The pachytene stage, also known as pachynema, from Greek words meaning "thick threads,"[1] contains the following chromosomal crossover. Nonsister chromatids of homologous chromosomes randomly exchange segments of genetic information over regions of homology. (Sex chromosomes, however, are not identical, and only exchange information over a small region of homology.) Exchange takes place at sites where recombination nodules have formed. The exchange of information between the non-sister chromatids results in a recombination of information; each chromosome has the complete set of information it had before, and there are no gaps formed as a result of the process. Because the chromosomes cannot be distinguished in the synaptonemal complex, the actual act of crossing over is not perceivable through the microscope.
[edit] Diplotene
During the diplotene stage, also known as diplonema, from Greek words meaning "two threads,"[1] the synaptonemal complex degrades and homologous chromosomes separate from one another a little. The chromosomes themselves uncoil a bit, allowing some transcription of DNA. However, the homologous chromosomes of each bivalent remain tightly bound at chiasmata, the regions where crossing over occurred.
[edit] Diakinesis
Chromosomes condense further during the diakinesis stage, from Greek words meaning "moving through."[1] This is the first point in meiosis where the four parts of the tetrads are actually visible. Sites of crossing over entangle together, effectively overlapping, making chiasmata clearly visible. Other than this observation, the rest of the stage closely resembles prometaphase of mitosis; the nucleoli disappears, the nuclear membrane disintegrates into vesicles, and the meiotic spindle begins to form.
[edit] Synchronous processes
During these stages, centrioles are migrating to the two poles of the cell. These centrioles, which were duplicated during interphase, function as microtubule coordinating centers. Centrioles sprout microtubules, essentially cellular ropes and poles, during crossing over. They invade the nuclear membrane after it disintegrates, attaching to the chromosomes at the kinetochore. The kinetochore functions as a motor, pulling the chromosome along the attached microtubule toward the originating centriole, like a train on a track. There are two kinetochores on each tetrad, one for each centrosome. Prophase I is the longest phase in meiosis.
Microtubules that attach to the kinetochores are known as kinetochore microtubules. Other microtubules will interact with microtubules from the opposite centriole. These are called nonkinetochore microtubules.
[edit] 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. The physical basis of the independent assortment of chromosomes is the random orientation of each bivalent along the metaphase plate.
[edit] Anaphase I
Kinetochore microtubules shorten, severing the recombination nodules and pulling homologous chromosomes apart. Since each chromosome only has one kinetochore, whole chromosomes are pulled toward opposing poles, forming two haploid sets. Each chromosome still contains a pair of sister chromatids. Nonkinetochore microtubules lengthen, pushing the centrioles further apart. The cell elongates in preparation for division down the middle. in this phase, DNA becomes visible strands.
[edit] Telophase I
The first meiotic division effectively ends when the centromeres arrive at the poles. Each daughter cell now has half the number of chromosomes but each chromosome consists of a pair of chromatids. This effect produces a variety of responses from the neuro-synrchromatic enzyme, also known as NSE. The microtubules that make up the spindle network disappear, and a new nuclear membrane surrounds each haploid set. The chromosomes uncoil back into chromatin. Cytokinesis, the pinching of the cell membrane in animal cells or the formation of the cell wall in plant cells, occurs, completing the creation of two daughter cells.
Cells enter a period of rest known as interkinesis or interphase II. No DNA replication occurs during this stage. Note that many plants skip telophase I and interphase II, going immediately into prophase II.
[edit] Meiosis II
Prophase II takes an inversely proportional time compared to telophase I. In this prophase we see the disappearance of the nucleoli and the nuclear envelope again as well as the shortening and thickening of the chromatids. Centrioles move to the polar regions and arrange spindle fibres for the second meiotic division. The new equatorial plane is rotated by 90 degrees when compared to meiosis I, perpendicular to the previous plane.
In metaphase II, the centromeres contain two kinetochores, that attach to spindle fibers from the centrosomes (centrioloes) at each pole.
This is followed by anaphase II, where the centromeres are cleaved, allowing microtubules attached to the kinetochores to pull the sister chromatids apart. The sister chromatids by convention are now called sister chromosomes as they move toward opposing poles.
The process ends with telophase II, which is similar to telophase I, and is marked by uncoiling and lengthening of the chromosomes and the disappearance of the microtubules. Nuclear envelopes reform and cleavage or cell wall formation eventually produces a total of four daughter cells, each with a haploid set of chromosomes. Meiosis is now complete.
[edit] Significance of meiosis
Meiosis facilitates stable sexual reproduction. Without the halving of ploidy, or chromosome count, fertilization would result in zygotes that have twice the number of chromosomes than the zygotes from the previous generation. Successive generations would have an exponential increase in chromosome count, resulting in an unwieldy genome that would cripple the reproductive fitness of the species. Polyploidy, the state of having three or more sets of chromosomes, also results in developmental abnormalities or lethality [2]. 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.
[edit] Nondisjunction
The normal separation of chromosomes in Meiosis I or sister chromatids in meiosis II is termed disjunction. When the separation is not normal, it is 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, including:
Down's 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 - an extra Y chromosome in males
[edit] Meiosis in humans
In females, meiosis occurs in precursor cells known as oogonia. Each oogonia that initiates meiosis will divide twice to form a single oocyte and two 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.