Meiosis - Knowledge (XXG)

644:. The cells then proceed to a second division without an intervening round of DNA replication. The sister chromatids are segregated to separate daughter cells to produce a total of four haploid cells. Female animals employ a slight variation on this pattern and produce one large ovum and three small polar bodies. Because of recombination, an individual chromatid can consist of a new combination of maternal and paternal genetic information, resulting in offspring that are genetically distinct from either parent. Furthermore, an individual gamete can include an assortment of maternal, paternal, and recombinant chromatids. This genetic diversity resulting from sexual reproduction contributes to the variation in traits upon which 1752:. Meiosis of primordial germ cells happens at the time of puberty, much later than in females. Tissues of the male testis suppress meiosis by degrading retinoic acid, proposed to be a stimulator of meiosis. This is overcome at puberty when cells within seminiferous tubules called Sertoli cells start making their own retinoic acid. Sensitivity to retinoic acid is also adjusted by proteins called nanos and DAZL. Genetic loss-of-function studies on retinoic acid-generating enzymes have shown that retinoic acid is required postnatally to stimulate spermatogonia differentiation which results several days later in spermatocytes undergoing meiosis, however retinoic acid is not required during the time when meiosis initiates. 1113:. However, a subset of breaks (at least one per chromosome) form crossovers between non-sister (homologous) chromosomes resulting in the exchange of genetic information. The exchange of information between the homologous 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 an ordinary light microscope, and chiasmata are not visible until the next stage. 1759:, meiosis begins immediately after primordial germ cells migrate to the ovary in the embryo. Some studies suggest that retinoic acid derived from the primitive kidney (mesonephros) stimulates meiosis in embryonic ovarian oogonia and that tissues of the embryonic male testis suppress meiosis by degrading retinoic acid. However, genetic loss-of-function studies on retinoic acid-generating enzymes have shown that retinoic acid is not required for initiation of either female meiosis which occurs during embryogenesis or male meiosis which initiates postnatally. 1794:

centromeres and male recombination is higher at the telomeres. On average, 1 million bp (1 Mb) correspond to 1 cMorgan (cm = 1% recombination frequency). The frequency of cross-overs remain uncertain. In yeast, mouse and human, it has been estimated that ≥200 double-strand breaks (DSBs) are formed per meiotic cell. However, only a subset of DSBs (~5–30% depending on the organism), go on to produce crossovers, which would result in only 1-2 cross-overs per human chromosome.

238: 1642:. The diploid organism's germ-line cells undergo meiosis to produce spores. The spores proliferate by mitosis, growing into a haploid organism. The haploid organism's gamete then combines with another haploid organism's gamete, creating the zygote. The zygote undergoes repeated mitosis and differentiation to produce a new diploid organism. The haplodiplontic life cycle can be considered a fusion of the diplontic and haplontic life cycles. 2029: 944: 190: 5517: 1651: 1239:

haploid set. 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. However, cytokinesis does not fully complete resulting in "cytoplasmic bridges" which enable the cytoplasm to be shared between daughter cells until the end of meiosis II. Sister chromatids remain attached during telophase I.

1195:. Over time, the MTOCs merge until two poles have formed, generating a barrel shaped spindle. In human oocytes spindle microtubule nucleation begins on the chromosomes, forming an aster that eventually expands to surround the chromosomes. Chromosomes then slide along the microtubules towards the equator of the spindle, at which point the chromosome kinetochores form end-on attachments to microtubules. 935: 889: 93: 1365:. The independent orientation of homologous chromosome pairs along the metaphase plate during metaphase I and orientation of sister chromatids in metaphase II, this is the subsequent separation of homologs and sister chromatids during anaphase I and II, it allows a random and independent distribution of chromosomes to each daughter cell (and ultimately to gametes); and (2) 1040:, from Greek words meaning "paired threads", which in some organisms is also called the bouquet stage because of the way the telomeres cluster at one end of the nucleus. In this stage the homologous chromosomes become much more closely (~100 nm) and stably paired (a process called synapsis) mediated by the installation of the transverse and central elements of the 1690:(singular: oocyte). Each primary oocyte divides twice in meiosis, unequally in each case. The first division produces a daughter cell, and a much smaller polar body which may or may not undergo a second division. In meiosis II, division of the daughter cell produces a second polar body, and a single haploid cell, which enlarges to become an 1109:, from Greek words meaning "thick threads". is the stage at which all autosomal chromosomes have synapsed. In this stage homologous recombination, including chromosomal crossover (crossing over), is completed through the repair of the double strand breaks formed in leptotene. Most breaks are repaired without forming crossovers resulting in 1713:. During this process, the maturing oocytes resume meiosis and continue until metaphase II of meiosis II, where they are again arrested just before ovulation. If these oocytes are fertilized by sperm, they will resume and complete meiosis. During folliculogenesis in humans, usually one follicle becomes dominant while the others undergo 239: 240: 911:, there are also pervasive translational controls (e.g. selective usage of preformed mRNA), regulating the ultimate meiotic stage-specific protein expression of genes during meiosis. Thus, both transcriptional and translational controls determine the broad restructuring of meiotic cells needed to carry out meiosis. 1230:

the cohesin from the chromosome arms is degraded while the cohesin surrounding the centromere remains protected by a protein named Shugoshin (Japanese for "guardian spirit"), what prevents the sister chromatids from separating. This allows the sister chromatids to remain together while homologs are segregated.

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Meiosis II is the second meiotic division, and usually involves equational segregation, or separation of sister chromatids. Mechanically, the process is similar to mitosis, though its genetic results are fundamentally different. The result is the production of four haploid cells (n chromosomes; 23 in

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holds sister chromatids together from the time of their replication until anaphase. In mitosis, the force of kinetochore microtubules pulling in opposite directions creates tension. The cell senses this tension and does not progress with anaphase until all the chromosomes are properly bi-oriented. In

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Meiotic recombination is mediated through double stranded break, which is catalyzed by Spo11 protein. Also Mre11, Sae2 and Exo1 play role in breakage and recombination. After the breakage happen, recombination take place which is typically homologous. The recombination may go through either a double

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Most monosomic and trisomic human embryos are not viable, but some aneuploidies can be tolerated, such as trisomy for the smallest chromosome, chromosome 21. Phenotypes of these aneuploidies range from severe developmental disorders to asymptomatic. Medical conditions include but are not limited to:

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The first meiotic division effectively ends when the chromosomes arrive at the poles. Each daughter cell now has half the number of chromosomes but each chromosome consists of a pair of chromatids. The microtubules that make up the spindle network disappear, and a new nuclear membrane surrounds each

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Kinetochore microtubules shorten, pulling homologous chromosomes (which each consist of a pair of sister chromatids) to opposite poles. Nonkinetochore microtubules lengthen, pushing the centrosomes farther apart. The cell elongates in preparation for division down the center. Unlike in mitosis, only

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disassembles and homologous chromosomes separate from one another a little. However, the homologous chromosomes of each bivalent remain tightly bound at chiasmata, the regions where crossing-over occurred. The chiasmata remain on the chromosomes until they are severed at the transition to anaphase I

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Meiosis Prophase I in mice. In Leptotene (L), the axial elements (stained by SYCP3) begin to form. In Zygotene (Z), the transverse elements (SYCP1) and central elements of the synaptonemal complex are partially installed (appearing as yellow as they overlap with SYCP3). In Pachytene (P), it is fully

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were once generally regarded as asexual, evidence has been presented that most lineages are anciently sexual and that the majority of asexual groups probably arose recently and independently. Dacks and Rogers proposed, based on a phylogenetic analysis, that facultative sex was likely present in the

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Interphase is followed by meiosis I and then meiosis II. Meiosis I separates replicated homologous chromosomes, each still made up of two sister chromatids, into two daughter cells, thus reducing the chromosome number by half. During meiosis II, sister chromatids decouple and the resultant daughter

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gametes (ova or sperm) that contain one set of 23 chromosomes. When two gametes (an egg and a sperm) fuse, the resulting zygote is once again diploid, with the mother and father each contributing 23 chromosomes. This same pattern, but not the same number of chromosomes, occurs in all organisms that

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Recombination among the 23 pairs of human chromosomes is responsible for redistributing not just the actual chromosomes, but also pieces of each of them. There is also an estimated 1.6-fold more recombination in females relative to males. In addition, average, female recombination is higher at the

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stage, from Greek words meaning "moving through". 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

1291:, which is similar to telophase I, and is marked by decondensation and lengthening of the chromosomes and the disassembly of the spindle. Nuclear envelopes re-form and cleavage or cell plate formation eventually produces a total of four daughter cells, each with a haploid set of chromosomes. 1207:

from both spindle poles attach to their respective kinetochores, the paired 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

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Meiosis occurs in all animals and plants. The result, the production of gametes with half the number of chromosomes as the parent cell, is the same, but the detailed process is different. In animals, meiosis produces gametes directly. In land plants and some algae, there is an

994:, from Greek words meaning "thin threads". In this stage of prophase I, individual chromosomes—each consisting of two replicated sister chromatids—become "individualized" to form visible strands within the nucleus. The chromosomes each form a linear array of loops mediated by 968:). This process facilitates stable pairing between homologous chromosomes and hence enables accurate segregation of the chromosomes at the first meiotic division. The paired and replicated chromosomes are called bivalents (two chromosomes) or tetrads (four 881:

stages, similar in purpose to their analogous subphases in the mitotic cell cycle. Therefore, meiosis includes the stages of meiosis I (prophase I, metaphase I, anaphase I, telophase I) and meiosis II (prophase II, metaphase II, anaphase II, telophase II).

1535: 1527: 1284:, in which the remaining centromeric cohesin, not protected by Shugoshin anymore, is cleaved, allowing the sister chromatids to segregate. The sister chromatids by convention are now called sister chromosomes as they move toward opposing poles. 800:

which are further divided into Karyokinesis I, Cytokinesis I, Karyokinesis II, and Cytokinesis II, respectively. The preparatory steps that lead up to meiosis are identical in pattern and name to interphase of the mitotic cell cycle.

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to produce the meiotic spindle. In mice, approximately 80 MicroTubule Organizing Centers (MTOCs) form a sphere in the ooplasm and begin to nucleate microtubules that reach out towards chromosomes, attaching to the chromosomes at the

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chromosomes are segregated into four daughter cells. For diploid organisms, the daughter cells resulting from meiosis are haploid and contain only one copy of each chromosome. In some species, cells enter a resting phase known as

923:, which are joined as tetrads (2n, 4c), producing two haploid cells (n chromosomes, 23 in humans) which each contain chromatid pairs (1n, 2c). Because the ploidy is reduced from diploid to haploid, meiosis I is referred to as a 834:

of the cell since the centromere number remains the same. The identical sister chromatids have not yet condensed into the densely packaged chromosomes visible with the light microscope. This will take place during prophase I in

1623:. Two organisms of opposing sex contribute their haploid gametes to form a diploid zygote. The zygote undergoes meiosis immediately, creating four haploid cells. These cells undergo mitosis to create the organism. Many 1255:

humans) from the two haploid cells (with n chromosomes, each consisting of two sister chromatids) produced in meiosis I. The four main steps of meiosis II are: prophase II, metaphase II, anaphase II, and telophase II.

412:, which remain held together through sister chromatid cohesion. This S-phase can be referred to as "premeiotic S-phase" or "meiotic S-phase". Immediately following DNA replication, meiotic cells enter a prolonged 241: 4456:

Bowles J, Knight D, Smith C, Wilhelm D, Richman J, Mamiya S, Yashiro K, Chawengsaksophak K, Wilson MJ, Rossant J, Hamada H, Koopman P (April 2006). "Retinoid signaling determines germ cell fate in mice".

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and to have been present early in eukaryotic evolution. Eukaryotes that were once thought to lack meiotic sex have recently been shown to likely have, or once have had, this capability. As one example,

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of chromosomes is the random orientation of each bivalent along with the metaphase plate, with respect to the orientation of the other bivalents along the same equatorial line. The protein complex

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Seemingly there are checkpoints for meiotic cell division too. In S. pombe, Rad proteins, S. pombe Mek1 (with FHA kinase domain), Cdc25, Cdc2 and unknown factor is thought to form a checkpoint.

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that attach to spindle fibers from the centrosomes at opposite poles. The new equatorial metaphase plate is rotated by 90 degrees when compared to meiosis I, perpendicular to the previous plate.

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during prophase I results in new combinations of genetic information within chromosomes. However, such physical exchange does not always occur during meiosis. In the oocytes of the silkworm

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Cycling meiosis and fertilization events results in alternation between haploid and diploid states. The organism phase of the life cycle can occur either during the diploid state (

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We propose to apply the terms Maiosis or Maiotic phase to cover the whole series of nuclear changes included in the two divisions that were designated as Heterotype and Homotype by

1443:, particularly exogenously induced double-strand breaks. DNA repair capability appears to be a key quality control mechanism in the female germ line and a critical determinant of 1667:

generation produces haploid spores instead of gametes. When they germinate, these spores undergo repeated cell division by mitosis, developing into a multicellular haploid

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There are pauses during meiosis in females. Maturing oocytes are arrested in prophase I of meiosis I and lie dormant within a protective shell of somatic cells called the

735:, 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 1018:

which invade the homologous chromosomes, forming inter-axis bridges, and resulting in the pairing/co-alignment of homologues (to a distance of ~400 nm in mice).

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between the two chromosomes in each pair (one received from each parent), producing lots of recombinant chromosomes with unique genetic combinations in every gamete

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again as well as the shortening and thickening of the chromatids. Centrosomes move to the polar regions and arrange spindle fibers for the second meiotic division.

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human cells contain 23 pairs of chromosomes including 1 pair of sex chromosomes (46 total), half of maternal origin and half of paternal origin. Meiosis produces

6063: 5553: 1044:. Synapsis is thought to occur in a zipper-like fashion starting from a recombination nodule. The paired chromosomes are called bivalent or tetrad chromosomes. 4986:

Pérez-Hidalgo L, Moreno S, San-Segundo PA (January 2003). "Regulation of meiotic progression by the meiosis-specific checkpoint kinase Mek1 in fission yeast".

972:), with one chromosome coming from each parent. Prophase I is divided into a series of substages which are named according to the appearance of chromosomes. 956:

Prophase I is by far the longest phase of meiosis (lasting 13 out of 14 days in mice). During prophase I, homologous maternal and paternal chromosomes pair,

2172: 1505:, a haploid multicellular, facultatively sexual green algae, can be induced by heat shock to reproduce by meiotic sex. This induction can be inhibited by 816:: In this very active phase, the cell synthesizes its vast array of proteins, including the enzymes and structural proteins it will need for growth. In G 3344: 1638:(with sporic or intermediate meiosis), the living organism alternates between haploid and diploid states. Consequently, this cycle is also known as the 5845: 5739: 632:, so each homolog now consists of two identical sister chromatids. Then each set of homologs pair with each other and exchange genetic information by 5438: 1318:, a common intestinal parasite, was previously considered to have descended from a lineage that predated the emergence of meiosis and sex. However, 5088: 1217:

meiosis, establishing tension ordinarily requires at least one crossover per chromosome pair in addition to cohesin between sister chromatids (see

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It has been suggested that Yeast CEP1 gene product, that binds centromeric region CDE1, may play a role in chromosome pairing during meiosis-I.

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are present in the oocytes. The arrest of ooctyes at the four genome copy stage was proposed to provide the informational redundancy needed to

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Lin Y, Gill ME, Koubova J, Page DC (December 2008). "Germ cell-intrinsic and -extrinsic factors govern meiotic initiation in mouse embryos".

4196: 4171: 3952: 2794: 2765: 1816:. This results in the production of gametes which have either too many or too few of a particular chromosome, and is a common mechanism for 6557: 5785: 3649:"Inherited differences in crossing over and gene conversion frequencies between wild strains of Sordaria fimicola from "Evolution Canyon"" 4269:

Rosenbusch B (November 2006). "The contradictory information on the distribution of non-disjunction and pre-division in female gametes".

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In order to understand meiosis, a comparison to mitosis is helpful. The table below shows the differences between meiosis and mitosis.

1143:, all developing oocytes develop to this stage and are arrested in prophase I before birth. This suspended state is referred to as the 3088: 1340:, which cause human disease. However, these organisms were shown to have a sexual cycle consistent with a meiotic process. Although 731:

roundworm eggs. The significance of meiosis for reproduction and inheritance, however, was described only in 1890 by German biologist

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Rasmussen SW (April 1977). "The transformation of the Synaptonemal Complex into the 'elimination chromatin' in Bombyx mori oocytes".

1596:(with pre-gametic meiosis), as in humans, the organism is multicellular and diploid, grown by mitosis from a diploid cell called the 6056: 5546: 3274: 1619:(with post-zygotic meiosis), the organism is haploid, by the proliferation and differentiation of a single haploid cell called the 1097: 176: 3269:

Pierce, Benjamin (2009). «Chromosomes and Cell Reproduction». Genetics: A Conceptual Approach, Third Edition. W.H. FREEMAN AND CO.

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Chen JM, Cooper DN, Chuzhanova N, Férec C, Patrinos GP (October 2007). "Gene conversion: mechanisms, evolution and human disease".

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The spelling was changed to "meiosis" by Koernicke (1905) and by Pantel and De Sinety (1906) to follow the usual conventions for

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Holliday junction (dHJ) pathway or synthesis-dependent strand annealing (SDSA). (The second one gives to noncrossover product).

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are arrested at the prophase I stage of meiosis. In humans, as an example, oocytes are formed between three and four months of

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Schematic of the synaptonemal complex at different stages of prophase I and the chromosomes arranged as a linear array of loops.

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that contains two copies of each chromosome, one from each parent. Thus, alternating cycles of meiosis and fertilization enable

447:, Χ) between the homologous chromosomes. In most organisms, these links can help direct each pair of homologous chromosomes to 3976:

Slupphaug G, Kavli B, Krokan HE (October 2003). "The interacting pathways for prevention and repair of oxidative DNA damage".

5416: 1694:. Therefore, in females each primary oocyte that undergoes meiosis results in one mature ovum and two or three polar bodies. 114: 1149:

or dictyate. It lasts until meiosis is resumed to prepare the oocyte for ovulation, which happens at puberty or even later.

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installed except on the sex chromosomes. In Diplotene (D), it disassembles revealing chiasmata. CREST marks the centromeres.

309: 6178: 5473: 1362: 4409:"Two functionally redundant sources of retinoic acid secure spermatogonia differentiation in the seminiferous epithelium" 1002:

assemble forming an "axial element" from which the loops emanate. Recombination is initiated in this stage by the enzyme

6049: 5539: 5488: 5288: 5271: 3163:"Human oocytes. Error-prone chromosome-mediated spindle assembly favors chromosome segregation defects in human oocytes" 2640:"Genetic analysis of chromosome pairing, recombination, and cell cycle control during first meiotic prophase in mammals" 1706: 1589:

life cycle), in which there are two distinct organism phases, one with haploid cells and the other with diploid cells.

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The probability of nondisjunction in human oocytes increases with increasing maternal age, presumably due to loss of

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has now been found to possess a core set of meiotic genes, including five meiosis specific genes. Also evidence for

640:) between the homologs. In the first meiotic division, the homologs are segregated to separate daughter cells by the 458:

During meiosis II, the cohesion between sister chromatids is released and they segregate from one another, as during

354:). Additionally, prior to the division, genetic material from the paternal and maternal copies of each chromosome is 3463:

Akopyants NS, Kimblin N, Secundino N, Patrick R, Peters N, Lawyer P, Dobson DE, Beverley SM, Sacks DL (April 2009).

3114:"Self-organization of MTOCs replaces centrosome function during acentrosomal spindle assembly in live mouse oocytes" 931:

analogous to mitosis, in which the sister chromatids are segregated, creating four haploid daughter cells (1n, 1c).

129: 5996: 5699: 5426: 5266: 4113:"Sex as a response to oxidative stress: a twofold increase in cellular reactive oxygen species activates sex genes" 3892:

Bernstein H, Byerly HC, Hopf FA, Michod RE (September 1985). "Genetic damage, mutation, and the evolution of sex".

3835:"Oocytes can efficiently repair DNA double-strand breaks to restore genetic integrity and protect offspring health" 3696:

Xiang Y, Tsuchiya D, Guo F, Gardner J, McCroskey S, Price A, Tromer EC, Walters JR, Lake CM, Hawley RS (May 2023).

2132: 2097: 1660: 1639: 1549: 763: 378: 63: 5074: 3367:"A phylogenomic inventory of meiotic genes; evidence for sex in Giardia and an early eukaryotic origin of meiosis" 888: 103: 6325: 5694: 5689: 5684: 5679: 5674: 5669: 5664: 5659: 5654: 5256: 4669:

Hassold T, Jacobs P, Kline J, Stein Z, Warburton D (July 1980). "Effect of maternal age on autosomal trisomies".

2400: 2064: 1491: 964:), forming at least one crossover per chromosome. These crossovers become visible as chiasmata (plural; singular 4712:

Tsutsumi M, Fujiwara R, Nishizawa H, Ito M, Kogo H, Inagaki H, Ohye T, Kato T, Fujii T, Kurahashi H (May 2014).

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Overview of chromatides' and chromosomes' distribution within the mitotic and meiotic cycle of a male human cell

6608: 6603: 6379: 6163: 5197: 5098: 1470: 1436: 1396: 1370: 961: 904: 691: 633: 136: 110: 431:, a programmed process in which DNA may be cut and then repaired, which allows them to exchange some of their 4407:

Teletin M, Vernet N, Yu J, Klopfenstein M, Jones JW, Féret B, Kane MA, Ghyselinck NB, Mark M (January 2019).

3336: 3289:"Structure and functions of stable intercellular bridges formed by incomplete cytokinesis during development" 5646: 5478: 3220:"Complete kinetochore tracking reveals error-prone homologous chromosome biorientation in mammalian oocytes" 1608:

in females), which fertilize to form the zygote. The diploid zygote undergoes repeated cellular division by

895: 745: 4510:"Sex-specific timing of meiotic initiation is regulated by Cyp26b1 independent of retinoic acid signalling" 346:. It involves two rounds of division that ultimately result in four cells, each with only one copy of each 6332: 6315: 6285: 6258: 5394: 5389: 5185: 5180: 5124: 1366: 1218: 1209: 849:

phase as seen before mitosis is not present in meiosis. Meiotic prophase corresponds most closely to the G

515: 448: 302: 4803:"Meiosis in protists. Some structural and physiological aspects of meiosis in algae, fungi, and protozoa" 2868:"Regulating double-stranded DNA break repair towards crossover or non-crossover during mammalian meiosis" 1566:

cell division, production of gametes by meiosis and fertilization. At certain stages of the life cycle,

143: 6320: 6218: 5978: 5729: 5498: 2142: 2117: 1721:, and differs from the typical meiosis in that it features a long period of meiotic arrest known as the 1555: 1543: 1455: 1323: 920: 826:: The genetic material is replicated; each of the cell's chromosomes duplicates to become two identical 781: 637: 568: 436: 428: 424: 388:

is followed by two rounds of cell division to produce four daughter cells, each with half the number of

355: 206: 202: 5083: 1824:. Nondisjunction can occur in the meiosis I or meiosis II, phases of cellular reproduction, or during 1469:

that is passed on to progeny. Experimental findings indicate that a substantial benefit of meiosis is

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In vertebrate oogenesis, maintained by cytostatic factor (CSF) has role in switching into meiosis-II.

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Female mammals and birds are born possessing all the oocytes needed for future ovulations, and these

1384: 1334:. Another example of organisms previously thought to be asexual are parasitic protozoa of the genus 1132: 1041: 1007: 999: 759: 667:: haploid cells that can divide vegetatively without undergoing fertilization. Some eukaryotes, like 247: 3465:"Demonstration of genetic exchange during cyclical development of Leishmania in the sand fly vector" 125: 6593: 6208: 5330: 5298: 2386: 2112: 1737: 1559: 1327: 722: 499: 432: 331: 2287: 2239:

Hassold T, Hunt P (April 2001). "To err (meiotically) is human: the genesis of human aneuploidy".

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secretion from the anterior pituitary stimulates a few follicles to mature in a process known as

1380: 1313: 1010:(around 300 per meiosis in mice). This process generates single stranded DNA filaments coated by 965: 740: 440: 3569:

Dacks J, Roger AJ (June 1999). "The first sexual lineage and the relevance of facultative sex".

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The normal separation of chromosomes in meiosis I or sister chromatids in meiosis II is termed

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chromosomes. This attachment is referred to as a bipolar attachment. The physical basis of the

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Kirk DL, Kirk MM (January 1986). "Heat shock elicits production of sexual inducer in Volvox".

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within the fetus and are therefore present at birth. During this prophase I arrested stage (

671:, do not have the ability to carry out meiosis and have acquired the ability to reproduce by 6583: 6460: 6359: 5401: 5283: 5003: 4995: 4958: 4950: 4911: 4871: 4863: 4822: 4814: 4743: 4733: 4678: 4641: 4600: 4577: 4537: 4529: 4474: 4428: 4420: 4376: 4368: 4321: 4278: 4233: 4132: 4124: 4085: 4040: 4032: 3993: 3940: 3909: 3864: 3854: 3802: 3754: 3717: 3709: 3668: 3660: 3621: 3586: 3541: 3533: 3492: 3484: 3435: 3386: 3308: 3300: 3231: 3190: 3182: 3125: 3043: 2996: 2959: 2951: 2879: 2838: 2830: 2700: 2692: 2651: 2598: 2590: 2541: 2531: 2520:"Unified single-cell analysis of testis gene regulation and pathology in five mouse strains" 2490: 2480: 2357: 2318: 2248: 2107: 1921:

Production of gametes (sex cells) in sexually reproducing eukaryotes with diplont life cycle

1848: 1710: 1698: 1510: 1486: 1263: 1171: 1065: 827: 772: 263: 218: 6515: 6510: 2940:"Meiotic chromosomes in motion: a perspective from Mus musculus and Caenorhabditis elegans" 1600:. The organism's diploid germ-line stem cells undergo meiosis to make haploid gametes (the 6352: 6173: 6121: 5483: 5465: 5411: 5355: 5104: 5078: 4902:

Honigberg SM, McCarroll RM, Esposito RE (April 1993). "Regulatory mechanisms in meiosis".

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usually occurs between identical sister chromatids and does not result in genetic changes

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Meiosis begins with a diploid cell, which contains two copies of each chromosome, termed

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Bernstein H, Bernstein C (2010). "Evolutionary origin of recombination during meiosis".

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increased the frequency of mating and the formation of meiotic spores by 4 to 18-fold.

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Although the process of meiosis is related to the more general cell division process of

6243: 6167: 5239: 4963: 4938: 4876: 4851: 4748: 4713: 4682: 4645: 4604: 4542: 4509: 4433: 4408: 4381: 4356: 4137: 4112: 3935:

Bernstein H, Hopf FA, Michod RE (1987). "The Molecular Basis of the Evolution of Sex".

3869: 3834: 3698:"A molecular cell biology toolkit for the study of meiosis in the silkworm Bombyx mori" 3673: 3648: 3546: 3521: 3497: 3464: 3313: 3288: 3195: 3162: 2964: 2939: 2843: 2818: 2705: 2680: 2603: 2570: 2546: 2519: 2495: 2468: 2446: 1842: 1803: 1501: 1361:, which may be beneficial. Meiosis generates gamete genetic diversity in two ways: (1) 687: 502:, with successive generations maintaining the same number of chromosomes. For example, 362:, the haploid cells produced by meiosis from a male and a female will fuse to create a 4827: 4802: 4581: 4045: 4020: 3944: 2028: 1462:

process, and that when it occurs during meiosis it is an adaptation for repairing the

66:. For the process whereby cell nuclei divide to produce two copies of themselves, see 6577: 6223: 5925: 5453: 5384: 5350: 5261: 5227: 5044: 4915: 3016: 2137: 2092: 1836: 736: 714: 710: 491: 490:. Because the number of chromosomes is halved during meiosis, gametes can fuse (i.e. 359: 323: 226: 150: 5025: 4698: 4622: 4341: 4255: 4036: 3065: 2901: 2330: 2268: 2059:

How a cell proceeds to meiotic division in meiotic cell division is not well known.

1650: 943: 478:. In female animals, three of the four meiotic products are typically eliminated by 189: 6198: 6072: 5947: 5520: 5372: 5249: 5244: 5190: 4818: 4494: 3253: 3147: 2783: 1781: 1749: 1745: 1571: 1243: 1163: 859: 483: 444: 250: 4568:

Raikov IB (1995). "Meiosis in protists: recent advances and persisting problems".

3664: 2571:"High-resolution view of the yeast meiotic program revealed by ribosome profiling" 2569:

Brar GA, Yassour M, Friedman N, Regev A, Ingolia NT, Weissman JS (February 2012).

1439:

repair Prophase I arrested oocytes have a high capability for efficient repair of

934: 698:

to another and recombination of these DNA molecules of different parental origin.

4852:"Molecular regulation of the mitosis/meiosis decision in multicellular organisms" 4738: 3997: 2419: 1767:

While the majority of eukaryotes have a two-divisional meiosis (though sometimes

1674:

In both animals and plants, the final stage is for the gametes to fuse to form a

1671:

generation, which then produces gametes directly (i.e. without further meiosis).

1509:

indicating that the induction of meiotic sex by heat shock is likely mediated by

659:

to divide one cell into two identical daughter cells. In some plants, fungi, and

6417: 6412: 6275: 6253: 6248: 6116: 5968: 5937: 4954: 4867: 3713: 2834: 2696: 2518:

Jung M, Wells D, Rusch J, Ahmad S, Marchini J, Myers SR, Conrad DF (June 2019).

2069:. there is a role of MeiRNA binding protein for entry to meiotic cell division. 1772: 1668: 1601: 1506: 1375: 1274: 1192: 1015: 749:, which helped to establish that genetic traits are transmitted on chromosomes. 408:, the DNA of each chromosome is replicated so that it consists of two identical 374: 92: 4021:"Sexual reproduction as a response to H2O2 damage in Schizosaccharomyces pombe" 3839:

Proceedings of the National Academy of Sciences of the United States of America

3522:"The chastity of amoebae: re-evaluating evidence for sex in amoeboid organisms" 3236: 3219: 3130: 3113: 2955: 1353:

The new combinations of DNA created during meiosis are a significant source of

6437: 6427: 6422: 6407: 6347: 6337: 6238: 6213: 6187: 6138: 6128: 6076: 5889: 5773: 5562: 3440: 3415: 3391: 3366: 2884: 2867: 2102: 1726: 1664: 1534: 1526: 1459: 1428: 1400: 1336: 1187: 1144: 969: 802: 797: 718: 707: 405: 397: 389: 370: 347: 327: 214: 198: 194: 4508:

Kumar S, Chatzi C, Brade T, Cunningham TJ, Zhao X, Duester G (January 2011).

4206: 3098: 322: 'lessening', (since it is a reductional division) is a special type of 6389: 6133: 6111: 6092: 6041: 6012: 5962: 5952: 5635: 5631: 5621: 5617: 5613: 5603: 5599: 5589: 5585: 5360: 5335: 5325: 5313: 4478: 4325: 4282: 4238: 4221: 4089: 3913: 3859: 3488: 3186: 3048: 3031: 2594: 2485: 2322: 1718: 1574:

make up the body of the organism and are not involved in gamete production.

1567: 1444: 1416: 1388: 1308: 1167: 1140: 1123: 1053: 878: 870: 793: 656: 535: 519: 479: 393: 343: 210: 5017: 4972: 4885: 4757: 4597:

Emery and Rimoin's Principles and Practice of Medical Genetics and Genomics

4551: 4486: 4442: 4390: 4333: 4290: 4247: 4146: 4128: 4005: 3878: 3806: 3731: 3682: 3612:

Monaghan F, Corcos A (1984-01-01). "On the origins of the Mendelian laws".

3598: 3555: 3537: 3506: 3449: 3400: 3322: 3245: 3204: 3139: 3057: 3008: 2973: 2893: 2852: 2714: 2665: 2612: 2555: 2504: 2371: 2362: 2345: 2260: 1910:

Normally four cells, each with half the number of chromosomes as the parent

4923: 4690: 4097: 4054: 3962: 3921: 3814: 3633: 17: 6375: 6159: 6017: 5957: 5942: 5911: 5897: 5627: 5609: 5595: 5581: 5377: 5278: 4836: 3766: 2656: 2639: 2147: 2122: 1821: 1776: 1771:), a very rare form, one-divisional meiosis, occurs in some flagellates ( 1741: 1722: 1691: 1628: 1478: 1432: 1420: 1027: 957: 874: 866: 838: 809: 683: 625: 420: 413: 358:, creating new combinations of code on each chromosome. Later on, during 79: 4372: 2536: 1369:. The physical exchange of homologous chromosomal regions by homologous 6545: 6399: 6305: 5933: 5906: 5565: 5318: 5071: 5008: 4533: 4424: 4357:"Nanos2 suppresses meiosis and promotes male germ cell differentiation" 3758: 3590: 2404: 1933: 1880: 1825: 1817: 1609: 1563: 1358: 1213: 1136:

to allow homologous chromosomes to move to opposite poles of the cell.

995: 823: 727: 695: 679: 660: 652: 599: 551: 507: 503: 463: 459: 452: 401: 351: 335: 75: 67: 5531: 4999: 3304: 1744:. Meiosis during spermatogenesis is specific to a type of cell called 820:, each of the chromosomes consists of a single linear molecule of DNA. 6455: 6342: 6106: 6102: 6097: 5865: 5781: 5767: 5763: 5759: 5751: 5709: 5704: 5573: 5367: 5340: 5173: 2346:"Sex is not a solution for reproduction: the libertine bubble theory" 2252: 1756: 1687: 1675: 1624: 1620: 1597: 1463: 1424: 1412: 1341: 831: 611: 523: 495: 475: 471: 363: 315: 230: 222: 71: 3416:"Population genetics provides evidence for recombination in Giardia" 3000: 392:

as the original parent cell. The two meiotic divisions are known as

455:

cells that have half the number of chromosomes as the parent cell.

373:(an abnormal number of chromosomes) are the leading known cause of 6520: 6505: 6500: 6495: 6490: 6485: 6480: 6475: 6470: 6445: 6383: 5840: 5835: 5724: 5719: 5714: 3833:

Stringer JM, Winship A, Zerafa N, Wakefield M, Hutt K (May 2020).

2387:"On the Maiotic Phase (Reduction Divisions) in Animals and Plants" 1649: 1562:, consisting of the cyclical process of growth and development by 1294:

Meiosis is now complete and ends up with four new daughter cells.

1011: 1003: 664: 531: 527: 467: 339: 236: 2819:"Recombination, Pairing, and Synapsis of Homologs during Meiosis" 5869: 5860: 5830: 5809: 5747: 5168: 4595:

Hegde MR, Crowley MR (2019-01-01). "Genome and Gene Structure".

1977:

Yes, normally occurs between each pair of homologous chromosomes

1605: 908: 487: 6045: 5535: 5120: 5050:(3rd ed.). Upper Saddle River, NJ: Pearson Prentice Hall. 3520:

Lahr DJ, Parfrey LW, Mitchell EA, Katz LA, Lara E (July 2011).

1913:

Two cells, having the same number of chromosomes as the parent

1246:

or interphase II. No DNA replication occurs during this stage.

830:

attached at a centromere. This replication does not change the

5850: 5814: 5804: 5777: 5755: 5147: 4939:"Mechanism and regulation of meiotic recombination initiation" 4222:"Positioning to get out of meiosis: the asymmetry of division" 2023: 1466: 1086: 86: 1203:

Homologous pairs move together along the metaphase plate: As

907:. In addition to strong meiotic stage-specific expression of 753: 514:

Meiosis occurs in all sexually-reproducing single-celled and

3414:

Cooper MA, Adam RD, Worobey M, Sterling CR (November 2007).

3079:

Nussbaum RL, McInnes RR, Willard HF, Hamosh A (2015-05-21).

1924:

Cellular reproduction, growth, repair, asexual reproduction

1857:– extra X chromosomes in males – i.e. XXY, XXXY, XXXXY, etc. 5110:"Abby Dernburg Seminar: Chromosome Dynamics During Meiosis" 4714:"Age-related decrease of meiotic cohesins in human oocytes" 2063:

seemingly have role in frog Oocyte meiosis. In the fungus

706:

Meiosis was discovered and described for the first time in

275: 269: 1186:

Unlike mitotic cells, human and mouse oocytes do not have

3337:"BioCoach Activity: Concept 11: Meiosis II: Metaphase II" 2917:"Meiosis, genetic recombination, and sexual reproduction" 1678:

in which the original number of chromosomes is restored.

1489:

leading to oxidative DNA damage. Treatment of the yeast

1080: 1074: 281: 3265: 3263: 451:

away from each other during meiosis I, resulting in two

6563:

International System for Human Cytogenetic Nomenclature

3939:. Advances in Genetics. Vol. 24. pp. 323–70. 2781:

Krebs JE, Goldstein ES, Kilpatrick ST (November 2009).

2283: 2040: 2010:

Does not occur in Anaphase I, but occurs in Anaphase II

1958:

Prophase, Prometaphase, Metaphase, Anaphase, Telophase

758:, meaning 'lessening'. It was introduced to biology by 462:. In some cases, all four of the meiotic products form 42: 5116: 3161:

Holubcová Z, Blayney M, Elder K, Schuh M (June 2015).

1938:

In gonads, before gametes (in diplontic life cycles);

1307:

Meiosis appears to be a fundamental characteristic of

766:

in 1905, using the idiosyncratic rendering "maiosis":

3785:

Mira A (September 1998). "Why is meiosis arrested?".

3032:"Meiosis: the chromosomal foundation of reproduction" 1357:

alongside mutation, resulting in new combinations of

1098: 1089: 1077: 284: 278: 1955:

Prophase II, Metaphase II, Anaphase II, Telophase II

1812:. When the segregation is not normal, it is called 1083: 1071: 610:

produces two genetically identical cells, each with

6533: 6436: 6398: 6368: 6296: 6152: 6083: 6005: 5977: 5924: 5888: 5879: 5823: 5795: 5738: 5645: 5572: 5464: 5297: 5154: 1932:Almost all eukaryotes (animals, plants, fungi, and 1451:

Meiosis as an adaptation for repairing germline DNA

1262:, we see the disappearance of the nucleoli and the 1068: 694:involves the transfer of DNA from one bacterium or 366:, a cell with two copies of each chromosome again. 272: 266: 117:. Unsourced material may be challenged and removed. 5043: 3218:Kitajima TS, Ohsugi M, Ellenberg J (August 2011). 2782: 2228:(6th ed.). Hoboken, NY: Pearson. p. 210. 1953:Prophase I, Metaphase I, Anaphase I, Telophase I, 1863:– lacking of one X chromosome in females – i.e. X0 1717:. The process of meiosis in females occurs during 1423:), which may last for decades, four copies of the 752:The term "meiosis" is derived from the Greek word 717:. It was described again in 1883, at the level of 598:produces four genetically unique cells, each with 4897: 4895: 2728: 2726: 2724: 2469:"Regulation of meiotic gene expression in plants" 2420:"Meiosis and mitosis: a terminological criticism" 3365:Ramesh MA, Malik SB, Logsdon JM (January 2005). 3287:Haglund K, Nezis IP, Stenmark H (January 2011). 2681:"Meiotic Recombination: The Essence of Heredity" 743:detected crossovers in meiosis in the fruit fly 70:. For excessive constriction of the pupils, see 5084:Animations from the U. of Arizona Biology Dept. 3828: 3826: 3824: 2812: 2810: 2808: 2806: 2789:(10th ed.). Jones & Barlett Learning. 2751: 2749: 2747: 2745: 2743: 2741: 903:During meiosis, specific genes are more highly 4111:Nedelcu AM, Marcu O, Michod RE (August 2004). 2638:Cohen PE, Pollack SE, Pollard JW (June 2006). 2219: 2217: 1435:. The repair process used appears to involve 1131:, from Greek words meaning "two threads", the 435:. A subset of recombination events results in 6057: 5547: 5132: 2385:Farmer, J.B.; Moore, J.E.S. (February 1904). 1779:) from the gut of the wood-feeding cockroach 1686:In females, meiosis occurs in cells known as 8: 3081:Thompson & Thompson genetics in medicine 651:Meiosis uses many of the same mechanisms as 579:occurs only if needed to repair DNA damage; 233:) with a complete set of paired chromosomes. 3780: 3778: 3776: 3647:Saleem M, Lamb BC, Nevo E (December 2001). 1481:, as indicated by the following examples. 602:the number of chromosomes as in the parent 486:, and only one cell develops to produce an 26: 6395: 6064: 6050: 6042: 5885: 5554: 5540: 5532: 5139: 5125: 5117: 4943:Cold Spring Harbor Perspectives in Biology 4856:Cold Spring Harbor Perspectives in Biology 4402: 4400: 2823:Cold Spring Harbor Perspectives in Biology 2685:Cold Spring Harbor Perspectives in Biology 2445:. University of Cincinnati. Archived from 2391:Quarterly Journal of Microscopical Science 1945:All proliferating cells in all eukaryotes 1242:Cells may enter a period of rest known as 686:, which generally reproduce asexually via 5846:Cellular apoptosis susceptibility protein 5007: 4962: 4875: 4826: 4747: 4737: 4563: 4561: 4541: 4432: 4380: 4237: 4136: 4044: 3868: 3858: 3721: 3672: 3545: 3496: 3439: 3390: 3312: 3235: 3194: 3129: 3047: 2963: 2883: 2842: 2704: 2655: 2602: 2545: 2535: 2494: 2484: 2361: 443:(singular: chiasma, for the Greek letter 253:, played back at 120× the recorded speed. 177:Learn how and when to remove this message 2756:Snustad DP, Simmons MJ (December 2008). 1893: 1533: 1525: 1157:Chromosomes condense further during the 942: 933: 690:. However, a "sexual" process known as 556: 554:, it differs in two important respects: 188: 32:This is an accepted version of this page 4187:Bidlack JE, Jansky S, Stern KR (2011). 3293:Communicative & Integrative Biology 3030:Bolcun-Filas E, Handel MA (July 2018). 2732: 2625: 2164: 1581:life cycle), during the haploid state ( 960:, and exchange genetic information (by 865:Meiosis I and II are each divided into 636:often leading to physical connections ( 439:, which create physical links known as 377:and the most frequent genetic cause of 59:Cell division producing haploid gametes 28: 3083:(8th ed.). Elsevier. p. 19. 1999:Occurs in Telophase I and Telophase II 6553:List of organisms by chromosome count 3626:10.1093/oxfordjournals.jhered.a109868 1174:disintegrates into vesicles, and the 986:The first stage of prophase I is the 213:. The daughter cells divide again in 74:. For the parasitic infestation, see 7: 4782:from the original on 1 November 2012 4638:Encyclopedia of Biological Chemistry 3112:Schuh M, Ellenberg J (August 2007). 2938:Link J, Jantsch V (September 2019). 2467:Zhou A, Pawlowski WP (August 2014). 2290:from the original on 29 January 2018 1748:, which will later mature to become 663:meiosis results in the formation of 655:, the type of cell division used by 209:) during the first division, called 115:adding citations to reliable sources 5235:Heterogametic sex / Homogametic sex 4160:South GR, Whittick A (2009-07-08). 4019:Bernstein C, Johns V (April 1989). 1631:utilize the haplontic life cycle. 5089:Meiosis at Kimball's Biology Pages 4801:Heywood P, Magee PT (March 1976). 4683:10.1111/j.1469-1809.1980.tb00943.x 4646:10.1016/B978-0-12-378630-2.00474-6 4636:Hunter N (2013-01-01). "Meiosis". 4605:10.1016/B978-0-12-812537-3.00004-4 4355:Suzuki A, Saga Y (February 2008). 4189:Stern's Introductory Plant Biology 2866:Baudat F, de Massy B (July 2007). 2817:Zickler D, Kleckner N (May 2015). 2198:"Definition of Reduction division" 1985:Pairing of homologous chromosomes? 1869:– an extra X chromosome in females 1789:Role in human genetics and disease 1725:stage and lacks the assistance of 998:, and the lateral elements of the 862:between meiosis I and meiosis II. 57: 3937:Molecular Genetics of Development 2061:Maturation promoting factor (MPF) 1942:Before spores (in haplodiplontic) 1875:– an extra Y chromosome in males. 1663:such that meiosis in the diploid 1513:leading to increased DNA damage. 1458:can be viewed as fundamentally a 557: 534:. It is an essential process for 427:pair with each other and undergo 5516: 5515: 5309:Evolution of sexual reproduction 4937:Lam I, Keeney S (October 2014). 4570:European Journal of Protistology 4191:. McGraw Hill. pp. 214–29. 2027: 1732:In males, meiosis occurs during 1064: 887: 853:phase of the mitotic cell cycle. 713:in 1876 by the German biologist 614:of chromosomes as in the parent 400:. Before meiosis begins, during 262: 91: 6174:Macrochromosome/Microchromosome 4904:Current Opinion in Cell Biology 4778:. Public Broadcasting Service. 4037:10.1128/jb.171.4.1893-1897.1989 3347:from the original on 2018-02-28 1383:(lacking crossovers). Although 1345:common ancestor of eukaryotes. 585: 369:Errors in meiosis resulting in 102:needs additional citations for 78:. For muscle inflammation, see 5417:Sexual reproduction in animals 4819:10.1128/mmbr.40.1.190-240.1976 3787:Journal of Theoretical Biology 1940:After zygotes (in haplontic); 1273:, the centromeres contain two 805:is divided into three phases: 205:exchange genetic information ( 62:For the figure of speech, see 1: 4582:10.1016/s0932-4739(11)80349-4 4220:Brunet S, Verlhac MH (2010). 3945:10.1016/s0065-2660(08)60012-7 1554:Meiosis occurs in eukaryotic 1399:is absent between the paired 1363:Law of Independent Assortment 1032:Leptotene is followed by the 898:Diagram of the meiotic phases 419:-like stage known as meiotic 5289:Sex as a biological variable 5272:Simultaneous hermaphroditism 4916:10.1016/0955-0674(93)90106-z 4739:10.1371/journal.pone.0096710 4163:An Introduction to Phycology 3998:10.1016/j.mrfmmm.2003.06.002 2403:, Third Edition, June 2001, 628:. First, the cell undergoes 609: 597: 578: 567: 334:organisms that produces the 5856:Maturation promoting factor 5105:Stages of Meiosis animation 5094:Khan Academy, video lecture 4955:10.1101/cshperspect.a016634 4868:10.1101/cshperspect.a002683 3665:10.1093/genetics/159.4.1573 2835:10.1101/cshperspect.a016626 2697:10.1101/cshperspect.a016618 1963:Genetically same as parent? 1701:. At the beginning of each 1612:to grow into the organism. 721:, by the Belgian zoologist 592:chromosome number (ploidy) 6625: 6214:Dinoflagellate chromosomes 5997:Postreplication checkpoint 5427:Penile-vaginal intercourse 5267:Sequential hermaphroditism 3237:10.1016/j.cell.2011.07.031 3131:10.1016/j.cell.2007.06.025 2956:10.1007/s00412-019-00698-5 2473:Frontiers in Plant Science 2284:"Interactive Tree of Life" 2282:Letunic I, Bork P (2006). 2133:Alternation of generations 2098:Coefficient of coincidence 1851:– trisomy of chromosome 18 1845:– trisomy of chromosome 13 1839:– trisomy of chromosome 21 1801: 1661:alternation of generations 1640:alternation of generations 1550:Alternation of generations 1547: 1541: 1437:homologous recombinational 1051: 1025: 979: 754: 678:Meiosis does not occur in 618: 379:developmental disabilities 308: 246:A video of meiosis I in a 229:, forming a diploid cell ( 225:. Two gametes fuse during 64:Meiosis (figure of speech) 61: 6558:List of sequenced genomes 6326:Chromosomal translocation 6199:A chromosome/B chromosome 6190:(or accessory chromosome) 5511: 5257:Testis-determining factor 4226:Human Reproduction Update 3714:10.1093/g3journal/jkad058 3441:10.1016/j.cub.2007.10.020 3392:10.1016/j.cub.2005.01.003 2885:10.1007/s10577-007-1140-3 2679:Hunter N (October 2015). 2401:Oxford English Dictionary 1636:haplodiplontic life cycle 1492:Schizosaccharomyces pombe 1182:Meiotic spindle formation 1006:which creates programmed 591: 586: 561: 518:organisms (which are all 6380:Telomere-binding protein 6194:Supernumerary chromosome 5198:Sex-determination system 4850:Kimble J (August 2011). 4671:Annals of Human Genetics 2989:Nature Reviews. Genetics 2441:Carter JS (2012-10-27). 1485:is an agent that causes 1429:repair damage in the DNA 1397:homologous recombination 1379:, meiosis is completely 1205:kinetochore microtubules 962:homologous recombination 792:Meiosis is divided into 692:horizontal gene transfer 634:homologous recombination 39:latest accepted revision 5101:The Cell-Cycle Ontology 5072:Meiosis Flash Animation 4988:Journal of Cell Science 4807:Bacteriological Reviews 4640:(2nd ed.): 17–23. 4479:10.1126/science.1125691 4361:Genes & Development 4326:10.1126/science.1166340 4090:10.1126/science.3941891 3914:10.1126/science.3898363 3860:10.1073/pnas.2001124117 3614:The Journal of Heredity 3489:10.1126/science.1169464 3187:10.1126/science.aaa9529 3036:Biology of Reproduction 2760:(5th ed.). Wiley. 2595:10.1126/science.1215110 2486:10.3389/fpls.2014.00413 2323:10.1525/bio.2010.60.7.5 2241:Nature Reviews Genetics 1387:are present during the 746:Drosophila melanogaster 6316:Structural alterations 5979:Cell cycle checkpoints 5181:Sexual differentiation 4129:10.1098/rspb.2004.2747 3807:10.1006/jtbi.1998.0761 3538:10.1098/rspb.2011.0289 2758:Principles of Genetics 2363:10.1002/bies.201000125 1974:Crossing over happens? 1655: 1585:life cycle), or both ( 1539: 1531: 1471:recombinational repair 1385:synaptonemal complexes 1287:The process ends with 1219:Chromosome segregation 1210:independent assortment 948: 940: 921:homologous chromosomes 778: 425:homologous chromosomes 254: 234: 203:homologous chromosomes 6333:Numerical alterations 6321:Chromosomal inversion 6219:Homologous chromosome 6006:Other cellular phases 5730:CDK-activating kinase 5223:Temperature-dependent 4514:Nature Communications 4283:10.1093/humrep/del122 4239:10.1093/humupd/dmq044 3049:10.1093/biolre/ioy021 2143:Mitotic recombination 2118:Biological life cycle 1929:Where does it happen? 1887:Comparison to mitosis 1653: 1646:In plants and animals 1544:Biological life cycle 1538:Haplontic life cycle. 1537: 1529: 1456:Genetic recombination 1324:meiotic recombination 1127:stage, also known as 1036:stage, also known as 990:stage, also known as 946: 937: 919:Meiosis I segregates 782:transliterating Greek 768: 429:genetic recombination 245: 207:chromosomal crossover 192: 6541:Extrachromosomal DNA 6229:Satellite chromosome 6204:Lampbrush chromosome 6144:Nuclear organization 2657:10.1210/er.2005-0017 2418:Battaglia E (1985). 2344:Lodé T (June 2011). 2020:Molecular regulation 2002:Occurs in Telophase 1855:Klinefelter syndrome 1738:seminiferous tubules 1617:haplontic life cycle 1594:diplontic life cycle 1530:Diplontic life cycle 1391:stage of meiosis in 1315:Giardia intestinalis 1309:eukaryotic organisms 1280:This is followed by 1133:synaptonemal complex 1042:synaptonemal complex 1008:double strand breaks 1000:synaptonemal complex 925:reductional division 494:) to form a diploid 423:. During this time, 332:sexually-reproducing 111:improve this article 6234:Centromere position 6209:Polytene chromosome 6179:Circular chromosome 5439:Hormonal motivation 5412:Fungal reproduction 5331:Reproductive system 4730:2014PLoSO...996710T 4526:2011NatCo...2..151K 4471:2006Sci...312..596B 4373:10.1101/gad.1612708 4318:2008Sci...322.1685L 4082:1986Sci...231...51K 3990:2003MRFMM.531..231S 3906:1985Sci...229.1277B 3851:2020PNAS..11711513S 3845:(21): 11513–11522. 3799:1998JThBi.194..275M 3583:1999JMolE..48..779D 3481:2009Sci...324..265A 3432:2007CBio...17.1984C 3383:2005CBio...15..185R 3179:2015Sci...348.1143H 2915:O'Connor C (2008). 2872:Chromosome Research 2587:2012Sci...335..552B 2537:10.7554/eLife.43966 2113:Synizesis (biology) 2013:Occurs in Anaphase 1560:sexual reproduction 1328:sexual reproduction 1298:Origin and function 929:equational division 927:. Meiosis II is an 824:Synthesis (S) phase 723:Edouard Van Beneden 558: 500:sexual reproduction 433:genetic information 29:Page version status 6599:Molecular genetics 6589:Cellular processes 5992:Spindle checkpoint 5797:P53 p63 p73 family 5444:Human reproduction 5422:Sexual intercourse 5407:Plant reproduction 5077:2010-08-23 at the 5046:Biological Science 5042:Freeman S (2005). 4772:"How Cells Divide" 4534:10.1038/ncomms1136 4425:10.1242/dev.170225 4271:Human Reproduction 3759:10.1007/BF00329771 3591:10.1007/pl00013156 2735:, pp. 249–250 2424:Annali di Botanica 2226:Biological Science 2224:Freeman S (2011). 2177:Biology LibreTexts 2039:. You can help by 1656: 1540: 1532: 1162:closely resembles 949: 941: 741:Thomas Hunt Morgan 569:shuffles the genes 255: 235: 197:duplicate (during 35: 6571: 6570: 6529: 6528: 6266:Centromere number 6183:Linear chromosome 6039: 6038: 6035: 6034: 5987:Restriction point 5529: 5528: 5449:Lordosis behavior 5164:Sexual dimorphism 5057:978-0-13-140941-5 5000:10.1242/jcs.00232 4655:978-0-12-378631-9 4614:978-0-12-812537-3 4465:(5773): 596–600. 4198:978-0-07-304052-3 4173:978-1-4443-1420-5 3954:978-0-12-017624-3 3900:(4719): 1277–81. 3532:(1715): 2081–90. 3341:The Biology Place 3305:10.4161/cib.13550 2796:978-0-7637-6632-0 2767:978-0-470-38825-9 2644:Endocrine Reviews 2628:, pp. 244–45 2399:as quoted in the 2057: 2056: 2017: 2016: 2007:Centromeres split 1867:Triple X syndrome 1570:produce gametes. 1497:hydrogen peroxide 1483:Hydrogen peroxide 1407:Prophase I arrest 1355:genetic variation 1349:Genetic variation 1303:Origin of meiosis 1105:), also known as 828:sister chromatids 669:bdelloid rotifers 646:natural selection 642:spindle apparatus 622: 621: 511:utilize meiosis. 410:sister chromatids 243: 219:sister chromatids 187: 186: 179: 161: 16:(Redirected from 6616: 6396: 6360:Polyploidization 6188:Extra chromosome 6103:Genetic material 6066: 6059: 6052: 6043: 5886: 5556: 5549: 5542: 5533: 5519: 5518: 5479:Animal sexuality 5402:Sexual selection 5141: 5134: 5127: 5118: 5061: 5049: 5030: 5029: 5011: 4994:(Pt 2): 259–71. 4983: 4977: 4976: 4966: 4934: 4928: 4927: 4899: 4890: 4889: 4879: 4847: 4841: 4840: 4830: 4798: 4792: 4791: 4789: 4787: 4768: 4762: 4761: 4751: 4741: 4709: 4703: 4702: 4666: 4660: 4659: 4633: 4627: 4626: 4592: 4586: 4585: 4565: 4556: 4555: 4545: 4505: 4499: 4498: 4453: 4447: 4446: 4436: 4419:(1): dev170225. 4404: 4395: 4394: 4384: 4352: 4346: 4345: 4312:(5908): 1685–7. 4301: 4295: 4294: 4266: 4260: 4259: 4241: 4217: 4211: 4210: 4184: 4178: 4177: 4157: 4151: 4150: 4140: 4123:(1548): 1591–6. 4108: 4102: 4101: 4065: 4059: 4058: 4048: 4016: 4010: 4009: 3973: 3967: 3966: 3932: 3926: 3925: 3889: 3883: 3882: 3872: 3862: 3830: 3819: 3818: 3782: 3771: 3770: 3742: 3736: 3735: 3725: 3693: 3687: 3686: 3676: 3644: 3638: 3637: 3609: 3603: 3602: 3566: 3560: 3559: 3549: 3517: 3511: 3510: 3500: 3460: 3454: 3453: 3443: 3411: 3405: 3404: 3394: 3362: 3356: 3355: 3353: 3352: 3333: 3327: 3326: 3316: 3284: 3278: 3267: 3258: 3257: 3239: 3215: 3209: 3208: 3198: 3173:(6239): 1143–7. 3158: 3152: 3151: 3133: 3109: 3103: 3102: 3076: 3070: 3069: 3051: 3027: 3021: 3020: 2984: 2978: 2977: 2967: 2935: 2929: 2928: 2921:Nature Education 2912: 2906: 2905: 2887: 2863: 2857: 2856: 2846: 2814: 2801: 2800: 2788: 2778: 2772: 2771: 2753: 2736: 2730: 2719: 2718: 2708: 2676: 2670: 2669: 2659: 2635: 2629: 2623: 2617: 2616: 2606: 2566: 2560: 2559: 2549: 2539: 2515: 2509: 2508: 2498: 2488: 2464: 2458: 2457: 2455: 2454: 2438: 2432: 2431: 2415: 2409: 2398: 2382: 2376: 2375: 2365: 2341: 2335: 2334: 2306: 2300: 2299: 2297: 2295: 2279: 2273: 2272: 2253:10.1038/35066065 2236: 2230: 2229: 2221: 2212: 2211: 2209: 2208: 2194: 2188: 2187: 2185: 2184: 2169: 2108:Oxidative stress 2052: 2049: 2031: 2024: 1894: 1849:Edwards syndrome 1711:folliculogenesis 1511:oxidative stress 1487:oxidative stress 1395:, crossing-over 1326:, indicative of 1264:nuclear envelope 1178:begins to form. 1172:nuclear membrane 1166:of mitosis; the 1146:dictyotene stage 1101: 1096: 1095: 1092: 1091: 1088: 1085: 1082: 1079: 1076: 1073: 1070: 891: 757: 756: 559: 319: 312: 300: 299: 298: 297: 290: 287: 286: 283: 280: 277: 274: 271: 268: 244: 221:to form haploid 193:In meiosis, the 182: 175: 171: 168: 162: 160: 119: 95: 87: 21: 6624: 6623: 6619: 6618: 6617: 6615: 6614: 6613: 6609:1876 in science 6604:Plant sexuality 6574: 6573: 6572: 6567: 6525: 6432: 6394: 6364: 6353:Paleopolyploidy 6298: 6292: 6148: 6122:Heterochromatin 6085: 6079: 6070: 6040: 6031: 6021: 6001: 5973: 5920: 5915: 5901: 5881: 5875: 5819: 5791: 5734: 5641: 5568: 5560: 5530: 5525: 5507: 5494:Differentiation 5484:Human sexuality 5474:Plant sexuality 5460: 5356:Spermatogenesis 5300: 5293: 5156: 5150: 5145: 5079:Wayback Machine 5068: 5058: 5041: 5038: 5033: 4985: 4984: 4980: 4936: 4935: 4931: 4901: 4900: 4893: 4849: 4848: 4844: 4800: 4799: 4795: 4785: 4783: 4770: 4769: 4765: 4711: 4710: 4706: 4668: 4667: 4663: 4656: 4635: 4634: 4630: 4615: 4594: 4593: 4589: 4567: 4566: 4559: 4507: 4506: 4502: 4455: 4454: 4450: 4406: 4405: 4398: 4354: 4353: 4349: 4303: 4302: 4298: 4277:(11): 2739–42. 4268: 4267: 4263: 4219: 4218: 4214: 4199: 4186: 4185: 4181: 4174: 4159: 4158: 4154: 4110: 4109: 4105: 4067: 4066: 4062: 4018: 4017: 4013: 3984:(1–2): 231–51. 3975: 3974: 3970: 3955: 3934: 3933: 3929: 3891: 3890: 3886: 3832: 3831: 3822: 3784: 3783: 3774: 3744: 3743: 3739: 3695: 3694: 3690: 3646: 3645: 3641: 3611: 3610: 3606: 3568: 3567: 3563: 3519: 3518: 3514: 3475:(5924): 265–8. 3462: 3461: 3457: 3413: 3412: 3408: 3364: 3363: 3359: 3350: 3348: 3335: 3334: 3330: 3286: 3285: 3281: 3268: 3261: 3217: 3216: 3212: 3160: 3159: 3155: 3111: 3110: 3106: 3091: 3078: 3077: 3073: 3029: 3028: 3024: 3001:10.1038/nrg2193 2986: 2985: 2981: 2937: 2936: 2932: 2914: 2913: 2909: 2865: 2864: 2860: 2816: 2815: 2804: 2797: 2785:Lewin's Genes X 2780: 2779: 2775: 2768: 2755: 2754: 2739: 2731: 2722: 2691:(12): a016618. 2678: 2677: 2673: 2637: 2636: 2632: 2624: 2620: 2581:(6068): 552–7. 2568: 2567: 2563: 2517: 2516: 2512: 2466: 2465: 2461: 2452: 2450: 2440: 2439: 2435: 2417: 2416: 2412: 2397:(192): 489–558. 2384: 2383: 2379: 2343: 2342: 2338: 2308: 2307: 2303: 2293: 2291: 2281: 2280: 2276: 2238: 2237: 2233: 2223: 2222: 2215: 2206: 2204: 2196: 2195: 2191: 2182: 2180: 2171: 2170: 2166: 2162: 2157: 2153:Mating of yeast 2128:Parthenogenesis 2088: 2053: 2047: 2044: 2037:needs expansion 2022: 1954: 1941: 1939: 1937: 1889: 1873:Jacobs syndrome 1861:Turner syndrome 1806: 1800: 1791: 1765: 1734:spermatogenesis 1703:menstrual cycle 1684: 1648: 1552: 1546: 1524: 1519: 1453: 1409: 1351: 1332:G. intestinalis 1330:, was found in 1320:G. intestinalis 1305: 1300: 1252: 1236: 1227: 1201: 1184: 1176:meiotic spindle 1170:disappear, the 1155: 1139:In human fetal 1119: 1111:gene conversion 1099: 1067: 1063: 1056: 1050: 1030: 1024: 984: 982:Leptotene stage 978: 954: 917: 901: 900: 899: 897: 892: 852: 848: 842: 819: 813: 790: 733:August Weismann 704: 673:parthenogenesis 630:DNA replication 612:the same number 548: 540:spermatogenesis 417: 386:DNA replication 294: 293: 292: 265: 261: 237: 217:, splitting up 183: 172: 166: 163: 120: 118: 108: 96: 83: 60: 55: 54: 53: 52: 51: 50: 34: 22: 15: 12: 11: 5: 6622: 6620: 6612: 6611: 6606: 6601: 6596: 6591: 6586: 6576: 6575: 6569: 6568: 6566: 6565: 6560: 6555: 6550: 6549: 6548: 6537: 6535: 6531: 6530: 6527: 6526: 6524: 6523: 6518: 6513: 6508: 6503: 6498: 6493: 6488: 6483: 6478: 6473: 6468: 6463: 6458: 6453: 6448: 6442: 6440: 6434: 6433: 6431: 6430: 6425: 6420: 6415: 6410: 6404: 6402: 6393: 6392: 6387: 6372: 6370: 6366: 6365: 6363: 6362: 6357: 6356: 6355: 6350: 6345: 6340: 6330: 6329: 6328: 6323: 6313: 6308: 6302: 6300: 6294: 6293: 6291: 6290: 6289: 6288: 6283: 6278: 6273: 6263: 6262: 6261: 6256: 6251: 6246: 6244:Submetacentric 6241: 6231: 6226: 6221: 6216: 6211: 6206: 6201: 6196: 6191: 6185: 6176: 6171: 6170:or heterosome) 6164:Sex chromosome 6156: 6154: 6150: 6149: 6147: 6146: 6141: 6136: 6131: 6126: 6125: 6124: 6119: 6109: 6100: 6095: 6089: 6087: 6081: 6080: 6071: 6069: 6068: 6061: 6054: 6046: 6037: 6036: 6033: 6032: 6030: 6029: 6024: 6019: 6015: 6009: 6007: 6003: 6002: 6000: 5999: 5994: 5989: 5983: 5981: 5975: 5974: 5972: 5971: 5966: 5960: 5955: 5950: 5945: 5940: 5930: 5928: 5922: 5921: 5919: 5918: 5913: 5909: 5904: 5899: 5894: 5892: 5883: 5877: 5876: 5874: 5873: 5863: 5858: 5853: 5848: 5843: 5838: 5833: 5827: 5825: 5821: 5820: 5818: 5817: 5812: 5807: 5801: 5799: 5793: 5792: 5790: 5789: 5771: 5744: 5742: 5736: 5735: 5733: 5732: 5727: 5722: 5717: 5712: 5707: 5702: 5697: 5692: 5687: 5682: 5677: 5672: 5667: 5662: 5657: 5651: 5649: 5643: 5642: 5640: 5639: 5625: 5607: 5593: 5578: 5576: 5570: 5569: 5561: 5559: 5558: 5551: 5544: 5536: 5527: 5526: 5524: 5523: 5512: 5509: 5508: 5506: 5505: 5504: 5503: 5502: 5501: 5496: 5491: 5476: 5470: 5468: 5462: 5461: 5459: 5458: 5457: 5456: 5451: 5446: 5441: 5436: 5431: 5430: 5429: 5414: 5409: 5404: 5399: 5398: 5397: 5392: 5382: 5381: 5380: 5375: 5365: 5364: 5363: 5358: 5348: 5343: 5338: 5333: 5328: 5323: 5322: 5321: 5316: 5305: 5303: 5295: 5294: 5292: 5291: 5286: 5281: 5276: 5275: 5274: 5269: 5259: 5254: 5253: 5252: 5247: 5240:Sex chromosome 5237: 5232: 5231: 5230: 5225: 5220: 5215: 5210: 5205: 5195: 5194: 5193: 5188: 5178: 5177: 5176: 5171: 5160: 5158: 5152: 5151: 5146: 5144: 5143: 5136: 5129: 5121: 5115: 5114: 5113: 5112: 5102: 5096: 5091: 5086: 5081: 5067: 5066:External links 5064: 5063: 5062: 5056: 5037: 5034: 5032: 5031: 4978: 4949:(1): a016634. 4929: 4891: 4862:(8): a002683. 4842: 4813:(1): 190–240. 4793: 4763: 4704: 4661: 4654: 4628: 4613: 4587: 4557: 4500: 4448: 4396: 4347: 4296: 4261: 4212: 4197: 4179: 4172: 4152: 4103: 4076:(4733): 51–4. 4060: 4011: 3968: 3953: 3927: 3884: 3820: 3772: 3737: 3688: 3659:(4): 1573–93. 3639: 3604: 3561: 3512: 3455: 3426:(22): 1984–8. 3406: 3357: 3328: 3279: 3259: 3210: 3153: 3104: 3090:978-1437706963 3089: 3071: 3042:(1): 112–126. 3022: 2995:(10): 762–75. 2979: 2950:(3): 317–330. 2930: 2907: 2858: 2829:(6): a016626. 2802: 2795: 2773: 2766: 2737: 2720: 2671: 2650:(4): 398–426. 2630: 2618: 2561: 2510: 2459: 2433: 2410: 2377: 2336: 2317:(7): 498–505. 2301: 2274: 2231: 2213: 2189: 2173:"4.1: Meiosis" 2163: 2161: 2158: 2156: 2155: 2150: 2145: 2140: 2135: 2130: 2125: 2120: 2115: 2110: 2105: 2100: 2095: 2089: 2087: 2084: 2055: 2054: 2034: 2032: 2021: 2018: 2015: 2014: 2011: 2008: 2004: 2003: 2000: 1997: 1993: 1992: 1989: 1986: 1982: 1981: 1978: 1975: 1971: 1970: 1967: 1964: 1960: 1959: 1956: 1951: 1947: 1946: 1943: 1930: 1926: 1925: 1922: 1919: 1915: 1914: 1911: 1908: 1904: 1903: 1900: 1897: 1888: 1885: 1877: 1876: 1870: 1864: 1858: 1852: 1846: 1843:Patau syndrome 1840: 1814:nondisjunction 1804:Nondisjunction 1802:Main article: 1799: 1798:Nondisjunction 1796: 1790: 1787: 1764: 1761: 1757:female mammals 1683: 1680: 1647: 1644: 1587:haplodiplontic 1542:Main article: 1523: 1522:In life cycles 1520: 1518: 1515: 1502:Volvox carteri 1452: 1449: 1408: 1405: 1350: 1347: 1304: 1301: 1299: 1296: 1251: 1248: 1235: 1232: 1226: 1223: 1200: 1197: 1183: 1180: 1154: 1151: 1118: 1115: 1052:Main article: 1049: 1046: 1026:Main article: 1023: 1020: 980:Main article: 977: 974: 953: 950: 916: 913: 894: 893: 886: 885: 884: 855: 854: 850: 846: 840: 836: 821: 817: 811: 789: 786: 703: 700: 688:binary fission 620: 619: 616: 615: 608: 604: 603: 596: 593: 589: 588: 584: 583: 577: 573: 572: 566: 563: 562:recombination 547: 544: 415: 185: 184: 99: 97: 90: 58: 56: 47:22 August 2024 36: 30: 27: 25: 24: 23: 14: 13: 10: 9: 6: 4: 3: 2: 6621: 6610: 6607: 6605: 6602: 6600: 6597: 6595: 6592: 6590: 6587: 6585: 6582: 6581: 6579: 6564: 6561: 6559: 6556: 6554: 6551: 6547: 6544: 6543: 6542: 6539: 6538: 6536: 6532: 6522: 6519: 6517: 6514: 6512: 6509: 6507: 6504: 6502: 6499: 6497: 6494: 6492: 6489: 6487: 6484: 6482: 6479: 6477: 6474: 6472: 6469: 6467: 6464: 6462: 6459: 6457: 6454: 6452: 6449: 6447: 6444: 6443: 6441: 6439: 6435: 6429: 6426: 6424: 6421: 6419: 6416: 6414: 6411: 6409: 6406: 6405: 6403: 6401: 6397: 6391: 6388: 6385: 6381: 6377: 6374: 6373: 6371: 6367: 6361: 6358: 6354: 6351: 6349: 6346: 6344: 6341: 6339: 6336: 6335: 6334: 6331: 6327: 6324: 6322: 6319: 6318: 6317: 6314: 6312: 6309: 6307: 6304: 6303: 6301: 6299:and evolution 6295: 6287: 6284: 6282: 6279: 6277: 6274: 6272: 6269: 6268: 6267: 6264: 6260: 6257: 6255: 6252: 6250: 6247: 6245: 6242: 6240: 6237: 6236: 6235: 6232: 6230: 6227: 6225: 6224:Isochromosome 6222: 6220: 6217: 6215: 6212: 6210: 6207: 6205: 6202: 6200: 6197: 6195: 6192: 6189: 6186: 6184: 6180: 6177: 6175: 6172: 6169: 6165: 6161: 6158: 6157: 6155: 6151: 6145: 6142: 6140: 6137: 6135: 6132: 6130: 6127: 6123: 6120: 6118: 6115: 6114: 6113: 6110: 6108: 6104: 6101: 6099: 6096: 6094: 6091: 6090: 6088: 6082: 6078: 6074: 6067: 6062: 6060: 6055: 6053: 6048: 6047: 6044: 6028: 6025: 6023: 6016: 6014: 6011: 6010: 6008: 6004: 5998: 5995: 5993: 5990: 5988: 5985: 5984: 5982: 5980: 5976: 5970: 5967: 5964: 5961: 5959: 5956: 5954: 5951: 5949: 5946: 5944: 5941: 5939: 5935: 5932: 5931: 5929: 5927: 5923: 5917: 5910: 5908: 5905: 5903: 5896: 5895: 5893: 5891: 5887: 5884: 5878: 5871: 5867: 5864: 5862: 5859: 5857: 5854: 5852: 5849: 5847: 5844: 5842: 5839: 5837: 5834: 5832: 5829: 5828: 5826: 5822: 5816: 5813: 5811: 5808: 5806: 5803: 5802: 5800: 5798: 5794: 5787: 5783: 5779: 5775: 5772: 5769: 5765: 5761: 5757: 5753: 5749: 5746: 5745: 5743: 5741: 5740:CDK inhibitor 5737: 5731: 5728: 5726: 5723: 5721: 5718: 5716: 5713: 5711: 5708: 5706: 5703: 5701: 5698: 5696: 5693: 5691: 5688: 5686: 5683: 5681: 5678: 5676: 5673: 5671: 5668: 5666: 5663: 5661: 5658: 5656: 5653: 5652: 5650: 5648: 5644: 5637: 5633: 5629: 5626: 5623: 5619: 5615: 5611: 5608: 5605: 5601: 5597: 5594: 5591: 5587: 5583: 5580: 5579: 5577: 5575: 5571: 5567: 5564: 5557: 5552: 5550: 5545: 5543: 5538: 5537: 5534: 5522: 5514: 5513: 5510: 5500: 5497: 5495: 5492: 5490: 5487: 5486: 5485: 5482: 5481: 5480: 5477: 5475: 5472: 5471: 5469: 5467: 5463: 5455: 5454:Pelvic thrust 5452: 5450: 5447: 5445: 5442: 5440: 5437: 5435: 5432: 5428: 5425: 5424: 5423: 5420: 5419: 5418: 5415: 5413: 5410: 5408: 5405: 5403: 5400: 5396: 5393: 5391: 5388: 5387: 5386: 5385:Fertilization 5383: 5379: 5376: 5374: 5371: 5370: 5369: 5366: 5362: 5359: 5357: 5354: 5353: 5352: 5351:Gametogenesis 5349: 5347: 5344: 5342: 5339: 5337: 5334: 5332: 5329: 5327: 5324: 5320: 5317: 5315: 5312: 5311: 5310: 5307: 5306: 5304: 5302: 5296: 5290: 5287: 5285: 5284:parasexuality 5282: 5280: 5277: 5273: 5270: 5268: 5265: 5264: 5263: 5262:Hermaphrodite 5260: 5258: 5255: 5251: 5248: 5246: 5243: 5242: 5241: 5238: 5236: 5233: 5229: 5228:Haplodiploidy 5226: 5224: 5221: 5219: 5216: 5214: 5211: 5209: 5206: 5204: 5201: 5200: 5199: 5196: 5192: 5189: 5187: 5184: 5183: 5182: 5179: 5175: 5172: 5170: 5167: 5166: 5165: 5162: 5161: 5159: 5153: 5149: 5142: 5137: 5135: 5130: 5128: 5123: 5122: 5119: 5111: 5108: 5107: 5106: 5103: 5100: 5097: 5095: 5092: 5090: 5087: 5085: 5082: 5080: 5076: 5073: 5070: 5069: 5065: 5059: 5053: 5048: 5047: 5040: 5039: 5035: 5027: 5023: 5019: 5015: 5010: 5005: 5001: 4997: 4993: 4989: 4982: 4979: 4974: 4970: 4965: 4960: 4956: 4952: 4948: 4944: 4940: 4933: 4930: 4925: 4921: 4917: 4913: 4910:(2): 219–25. 4909: 4905: 4898: 4896: 4892: 4887: 4883: 4878: 4873: 4869: 4865: 4861: 4857: 4853: 4846: 4843: 4838: 4834: 4829: 4824: 4820: 4816: 4812: 4808: 4804: 4797: 4794: 4781: 4777: 4773: 4767: 4764: 4759: 4755: 4750: 4745: 4740: 4735: 4731: 4727: 4724:(5): e96710. 4723: 4719: 4715: 4708: 4705: 4700: 4696: 4692: 4688: 4684: 4680: 4676: 4672: 4665: 4662: 4657: 4651: 4647: 4643: 4639: 4632: 4629: 4624: 4620: 4616: 4610: 4606: 4602: 4598: 4591: 4588: 4583: 4579: 4575: 4571: 4564: 4562: 4558: 4553: 4549: 4544: 4539: 4535: 4531: 4527: 4523: 4519: 4515: 4511: 4504: 4501: 4496: 4492: 4488: 4484: 4480: 4476: 4472: 4468: 4464: 4460: 4452: 4449: 4444: 4440: 4435: 4430: 4426: 4422: 4418: 4414: 4410: 4403: 4401: 4397: 4392: 4388: 4383: 4378: 4374: 4370: 4366: 4362: 4358: 4351: 4348: 4343: 4339: 4335: 4331: 4327: 4323: 4319: 4315: 4311: 4307: 4300: 4297: 4292: 4288: 4284: 4280: 4276: 4272: 4265: 4262: 4257: 4253: 4249: 4245: 4240: 4235: 4231: 4227: 4223: 4216: 4213: 4208: 4204: 4200: 4194: 4190: 4183: 4180: 4175: 4169: 4165: 4164: 4156: 4153: 4148: 4144: 4139: 4134: 4130: 4126: 4122: 4118: 4117:Proc Biol Sci 4114: 4107: 4104: 4099: 4095: 4091: 4087: 4083: 4079: 4075: 4071: 4064: 4061: 4056: 4052: 4047: 4042: 4038: 4034: 4031:(4): 1893–7. 4030: 4026: 4022: 4015: 4012: 4007: 4003: 3999: 3995: 3991: 3987: 3983: 3979: 3972: 3969: 3964: 3960: 3956: 3950: 3946: 3942: 3938: 3931: 3928: 3923: 3919: 3915: 3911: 3907: 3903: 3899: 3895: 3888: 3885: 3880: 3876: 3871: 3866: 3861: 3856: 3852: 3848: 3844: 3840: 3836: 3829: 3827: 3825: 3821: 3816: 3812: 3808: 3804: 3800: 3796: 3793:(2): 275–87. 3792: 3788: 3781: 3779: 3777: 3773: 3768: 3764: 3760: 3756: 3753:(3): 205–21. 3752: 3748: 3741: 3738: 3733: 3729: 3724: 3719: 3715: 3711: 3707: 3703: 3702:G3 (Bethesda) 3699: 3692: 3689: 3684: 3680: 3675: 3670: 3666: 3662: 3658: 3654: 3650: 3643: 3640: 3635: 3631: 3627: 3623: 3619: 3615: 3608: 3605: 3600: 3596: 3592: 3588: 3584: 3580: 3577:(6): 779–83. 3576: 3572: 3565: 3562: 3557: 3553: 3548: 3543: 3539: 3535: 3531: 3527: 3526:Proc Biol Sci 3523: 3516: 3513: 3508: 3504: 3499: 3494: 3490: 3486: 3482: 3478: 3474: 3470: 3466: 3459: 3456: 3451: 3447: 3442: 3437: 3433: 3429: 3425: 3421: 3417: 3410: 3407: 3402: 3398: 3393: 3388: 3384: 3380: 3377:(2): 185–91. 3376: 3372: 3368: 3361: 3358: 3346: 3342: 3338: 3332: 3329: 3324: 3320: 3315: 3310: 3306: 3302: 3298: 3294: 3290: 3283: 3280: 3276: 3275:9780716779285 3272: 3266: 3264: 3260: 3255: 3251: 3247: 3243: 3238: 3233: 3230:(4): 568–81. 3229: 3225: 3221: 3214: 3211: 3206: 3202: 3197: 3192: 3188: 3184: 3180: 3176: 3172: 3168: 3164: 3157: 3154: 3149: 3145: 3141: 3137: 3132: 3127: 3124:(3): 484–98. 3123: 3119: 3115: 3108: 3105: 3100: 3096: 3092: 3086: 3082: 3075: 3072: 3067: 3063: 3059: 3055: 3050: 3045: 3041: 3037: 3033: 3026: 3023: 3018: 3014: 3010: 3006: 3002: 2998: 2994: 2990: 2983: 2980: 2975: 2971: 2966: 2961: 2957: 2953: 2949: 2945: 2941: 2934: 2931: 2926: 2922: 2918: 2911: 2908: 2903: 2899: 2895: 2891: 2886: 2881: 2878:(5): 565–77. 2877: 2873: 2869: 2862: 2859: 2854: 2850: 2845: 2840: 2836: 2832: 2828: 2824: 2820: 2813: 2811: 2809: 2807: 2803: 2798: 2792: 2787: 2786: 2777: 2774: 2769: 2763: 2759: 2752: 2750: 2748: 2746: 2744: 2742: 2738: 2734: 2729: 2727: 2725: 2721: 2716: 2712: 2707: 2702: 2698: 2694: 2690: 2686: 2682: 2675: 2672: 2667: 2663: 2658: 2653: 2649: 2645: 2641: 2634: 2631: 2627: 2622: 2619: 2614: 2610: 2605: 2600: 2596: 2592: 2588: 2584: 2580: 2576: 2572: 2565: 2562: 2557: 2553: 2548: 2543: 2538: 2533: 2529: 2525: 2521: 2514: 2511: 2506: 2502: 2497: 2492: 2487: 2482: 2478: 2474: 2470: 2463: 2460: 2449:on 2012-10-27 2448: 2444: 2437: 2434: 2429: 2425: 2421: 2414: 2411: 2408: 2407: 2402: 2396: 2392: 2388: 2381: 2378: 2373: 2369: 2364: 2359: 2356:(6): 419–22. 2355: 2351: 2347: 2340: 2337: 2332: 2328: 2324: 2320: 2316: 2312: 2305: 2302: 2289: 2285: 2278: 2275: 2270: 2266: 2262: 2258: 2254: 2250: 2247:(4): 280–91. 2246: 2242: 2235: 2232: 2227: 2220: 2218: 2214: 2203: 2199: 2193: 2190: 2178: 2174: 2168: 2165: 2159: 2154: 2151: 2149: 2146: 2144: 2141: 2139: 2138:Brachymeiosis 2136: 2134: 2131: 2129: 2126: 2124: 2121: 2119: 2116: 2114: 2111: 2109: 2106: 2104: 2101: 2099: 2096: 2094: 2093:Fertilisation 2091: 2090: 2085: 2083: 2080: 2077: 2073: 2070: 2068: 2067: 2062: 2051: 2042: 2038: 2035:This section 2033: 2030: 2026: 2025: 2019: 2012: 2009: 2006: 2005: 2001: 1998: 1995: 1994: 1990: 1987: 1984: 1983: 1979: 1976: 1973: 1972: 1968: 1965: 1962: 1961: 1957: 1952: 1949: 1948: 1944: 1935: 1931: 1928: 1927: 1923: 1920: 1917: 1916: 1912: 1909: 1906: 1905: 1901: 1898: 1896: 1895: 1892: 1886: 1884: 1882: 1874: 1871: 1868: 1865: 1862: 1859: 1856: 1853: 1850: 1847: 1844: 1841: 1838: 1837:Down syndrome 1835: 1834: 1833: 1829: 1827: 1823: 1819: 1815: 1811: 1805: 1797: 1795: 1788: 1786: 1784: 1783: 1778: 1774: 1770: 1762: 1760: 1758: 1753: 1751: 1747: 1746:spermatocytes 1743: 1739: 1735: 1730: 1728: 1724: 1720: 1716: 1712: 1708: 1704: 1700: 1695: 1693: 1689: 1681: 1679: 1677: 1672: 1670: 1666: 1662: 1652: 1645: 1643: 1641: 1637: 1632: 1630: 1626: 1622: 1618: 1613: 1611: 1607: 1604:in males and 1603: 1599: 1595: 1590: 1588: 1584: 1580: 1575: 1573: 1572:Somatic cells 1569: 1565: 1561: 1557: 1551: 1545: 1536: 1528: 1521: 1516: 1514: 1512: 1508: 1504: 1503: 1498: 1494: 1493: 1488: 1484: 1480: 1476: 1472: 1468: 1465: 1461: 1457: 1450: 1448: 1446: 1442: 1438: 1434: 1430: 1426: 1422: 1418: 1414: 1406: 1404: 1402: 1398: 1394: 1390: 1386: 1382: 1378: 1377: 1372: 1371:recombination 1368: 1367:Crossing Over 1364: 1360: 1356: 1348: 1346: 1343: 1339: 1338: 1333: 1329: 1325: 1321: 1317: 1316: 1310: 1302: 1297: 1295: 1292: 1290: 1285: 1283: 1278: 1276: 1272: 1267: 1265: 1261: 1256: 1249: 1247: 1245: 1240: 1233: 1231: 1224: 1222: 1220: 1215: 1211: 1206: 1198: 1196: 1194: 1189: 1181: 1179: 1177: 1173: 1169: 1165: 1160: 1152: 1150: 1148: 1147: 1142: 1137: 1134: 1130: 1126: 1125: 1116: 1114: 1112: 1108: 1104: 1103: 1094: 1061: 1055: 1047: 1045: 1043: 1039: 1035: 1029: 1021: 1019: 1017: 1013: 1009: 1005: 1001: 997: 993: 989: 983: 975: 973: 971: 967: 963: 959: 951: 945: 936: 932: 930: 926: 922: 914: 912: 910: 906: 896: 890: 883: 880: 876: 872: 868: 863: 861: 844: 837: 833: 829: 825: 822: 815: 808: 807: 806: 804: 799: 795: 787: 785: 783: 777: 775: 774: 767: 765: 761: 750: 748: 747: 742: 738: 734: 730: 729: 724: 720: 716: 715:Oscar Hertwig 712: 709: 701: 699: 697: 693: 689: 685: 681: 676: 674: 670: 666: 662: 658: 654: 649: 647: 643: 639: 635: 631: 627: 617: 613: 606: 605: 601: 594: 590: 582: 575: 574: 570: 564: 560: 555: 553: 545: 543: 541: 537: 533: 529: 525: 522:), including 521: 517: 516:multicellular 512: 509: 505: 501: 497: 493: 492:fertilization 489: 485: 481: 477: 473: 469: 465: 461: 456: 454: 450: 446: 442: 438: 434: 430: 426: 422: 418: 411: 407: 403: 399: 395: 391: 387: 382: 380: 376: 372: 367: 365: 361: 360:fertilisation 357: 353: 349: 345: 341: 337: 333: 329: 325: 324:cell division 321: 318: 311: 307: 304: 303:Ancient Greek 296: 289: 259: 252: 249: 232: 228: 227:fertilization 224: 220: 216: 212: 208: 204: 200: 196: 191: 181: 178: 170: 167:February 2023 159: 156: 152: 149: 145: 142: 138: 135: 131: 128: – 127: 123: 122:Find sources: 116: 112: 106: 105: 100:This article 98: 94: 89: 88: 85: 81: 77: 73: 69: 65: 48: 44: 40: 33: 19: 6310: 6265: 6233: 6073:Cytogenetics 6026: 5948:Prometaphase 5373:spermatozoon 5345: 5301:reproduction 5250:Y chromosome 5245:X chromosome 5191:Virilization 5186:Feminization 5045: 4991: 4987: 4981: 4946: 4942: 4932: 4907: 4903: 4859: 4855: 4845: 4810: 4806: 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Retrieved 2447:the original 2436: 2427: 2423: 2413: 2405: 2394: 2390: 2380: 2353: 2349: 2339: 2314: 2310: 2304: 2292:. Retrieved 2277: 2244: 2240: 2234: 2225: 2205:. Retrieved 2201: 2192: 2181:. Retrieved 2179:. 2019-10-01 2176: 2167: 2081: 2078: 2074: 2071: 2065: 2058: 2045: 2041:adding to it 2036: 1980:Very rarely 1890: 1878: 1830: 1813: 1809: 1807: 1792: 1782:Cryptocercus 1780: 1773:parabasalids 1766: 1754: 1731: 1696: 1685: 1673: 1657: 1635: 1633: 1616: 1614: 1593: 1591: 1586: 1582: 1578: 1576: 1553: 1507:antioxidants 1500: 1490: 1454: 1410: 1392: 1374: 1352: 1335: 1331: 1319: 1314: 1306: 1293: 1289:telophase II 1288: 1286: 1281: 1279: 1275:kinetochores 1271:metaphase II 1270: 1268: 1259: 1257: 1253: 1244:interkinesis 1241: 1237: 1228: 1204: 1202: 1185: 1164:prometaphase 1158: 1156: 1145: 1138: 1128: 1122: 1120: 1106: 1059: 1057: 1037: 1033: 1031: 991: 987: 985: 955: 928: 924: 918: 902: 864: 860:interkinesis 856: 791: 779: 770: 769: 764:J.E.S. Moore 751: 744: 726: 705: 677: 650: 623: 580: 549: 513: 484:polar bodies 457: 384:In meiosis, 383: 368: 356:crossed over 316: 313: 306: 257: 256: 251:spermatocyte 173: 164: 154: 147: 140: 133: 121: 109:Please help 104:verification 101: 84: 46: 37:This is the 31: 6286:Polycentric 6276:Monocentric 6259:Holocentric 6254:Acrocentric 6249:Telocentric 6239:Metacentric 6117:Euchromatin 6077:chromosomes 5969:Cytokinesis 5938:Preprophase 5882:checkpoints 5036:Cited texts 5009:10261/62904 4413:Development 4025:J Bacteriol 3620:(1): 67–9. 3343:. Pearson. 2202:MedicineNet 2048:August 2020 1996:Cytokinesis 1883:over time. 1810:disjunction 1769:achiasmatic 1763:Flagellates 1750:spermatozoa 1727:centrosomes 1669:gametophyte 1602:spermatozoa 1556:life cycles 1401:chromosomes 1376:Bombyx mori 1282:anaphase II 1260:prophase II 1234:Telophase I 1199:Metaphase I 1193:kinetochore 1188:centrosomes 1121:During the 905:transcribed 839:Growth 2 (G 810:Growth 1 (G 760:J.B. Farmer 739:geneticist 719:chromosomes 390:chromosomes 375:miscarriage 301:; from 195:chromosomes 6594:Cell cycle 6578:Categories 6438:Centromere 6369:Structures 6348:Polyploidy 6338:Aneuploidy 6139:Nucleosome 6129:Chromosome 5890:Interphase 5880:Phases and 5563:Cell cycle 5434:Copulation 5155:Biological 4786:6 December 4520:(1): 151. 3747:Chromosoma 3571:J Mol Evol 3351:2018-02-10 3299:(1): 1–9. 2944:Chromosoma 2530:: e43966. 2453:2018-02-09 2430:: 101–140. 2311:BioScience 2207:2021-05-29 2183:2021-05-29 2160:References 2103:DNA repair 1907:End result 1682:In mammals 1665:sporophyte 1568:germ cells 1558:involving 1548:See also: 1517:Occurrence 1475:DNA damage 1460:DNA repair 1441:DNA damage 1381:achiasmate 1337:Leishmania 1250:Meiosis II 1225:Anaphase I 1159:diakinesis 1153:Diakinesis 970:chromatids 952:Prophase I 803:Interphase 798:meiosis II 708:sea urchin 657:eukaryotes 638:crossovers 520:eukaryotes 437:crossovers 406:cell cycle 398:meiosis II 371:aneuploidy 348:chromosome 328:germ cells 215:meiosis II 199:interphase 137:newspapers 18:Meiosis II 6390:Protamine 6297:Processes 6281:Dicentric 6134:Chromatid 6112:Chromatin 6093:Karyotype 6013:Apoptosis 5963:Telophase 5953:Metaphase 5748:INK4a/ARF 5489:Mechanics 5466:Sexuality 5361:Oogenesis 5336:Sex organ 5326:Germ cell 5314:Anisogamy 4599:: 53–77. 4207:320895017 4166:. Wiley. 3978:Mutat Res 3420:Curr Biol 3371:Curr Biol 3099:908336124 3017:205484180 2927:(1): 174. 2443:"Mitosis" 2350:BioEssays 1777:oxymonads 1742:testicles 1719:oogenesis 1627:and many 1583:haplontic 1579:diplontic 1445:fertility 1417:gestation 1389:pachytene 1141:oogenesis 1129:diplonema 1124:diplotene 1117:Diplotene 1107:pachynema 1060:pachytene 1054:Pachytene 1048:Pachytene 992:leptonema 988:leptotene 976:Leptotene 915:Meiosis I 879:telophase 871:metaphase 794:meiosis I 648:can act. 536:oogenesis 480:extrusion 449:segregate 441:chiasmata 394:meiosis I 344:egg cells 248:crane fly 211:meiosis I 126:"Meiosis" 6534:See also 6376:Telomere 6343:Euploidy 6271:Acentric 6168:allosome 6160:Autosome 6086:concepts 5958:Anaphase 5943:Prophase 5566:proteins 5521:Category 5499:Activity 5395:Internal 5390:External 5279:Intersex 5075:Archived 5026:14608163 5018:12482912 4973:25324213 4886:21646377 4780:Archived 4758:24806359 4718:PLOS ONE 4699:24469567 4623:92480716 4552:21224842 4487:16574820 4443:30487180 4391:18281459 4342:11261341 4334:19074348 4291:16982661 4256:13988521 4248:20833637 4147:15306305 4006:14637258 3879:32381741 3732:36911915 3723:10151401 3683:11779798 3653:Genetics 3599:10229582 3556:21429931 3507:19359589 3450:17980591 3401:15668177 3345:Archived 3323:21509167 3246:21854982 3205:26045437 3140:17693257 3066:38589675 3058:29385397 3009:17846636 2974:30877366 2902:26696085 2894:17674146 2853:25986558 2715:26511629 2666:16543383 2613:22194413 2556:31237565 2505:25202317 2372:21472739 2331:86663600 2288:Archived 2269:22264575 2261:11283700 2148:Dikaryon 2123:Apomixis 2086:See also 2066:S. pombe 1934:protists 1918:Function 1902:Mitosis 1822:monosomy 1723:dictyate 1699:follicle 1629:protozoa 1479:germline 1433:germline 1421:dictyate 1168:nucleoli 1038:zygonema 1034:zygotene 1028:Zygotene 1022:Zygotene 875:anaphase 867:prophase 835:meiosis. 773:Flemming 737:American 696:archaeon 684:bacteria 661:protists 626:homologs 607:mitosis 595:meiosis 576:mitosis 565:meiosis 546:Overview 466:such as 421:prophase 80:Myositis 43:reviewed 6584:Meiosis 6546:Plasmid 6400:Histone 6311:Meiosis 6306:Mitosis 6027:Meiosis 5934:Mitosis 5926:M phase 5907:S phase 5774:cip/kip 5346:Meiosis 5319:Isogamy 4964:4292169 4924:8389567 4877:3140684 4749:4013030 4726:Bibcode 4691:7198887 4576:: 1–7. 4543:3034736 4522:Bibcode 4495:2514848 4467:Bibcode 4459:Science 4434:6340151 4382:2238665 4314:Bibcode 4306:Science 4138:1691771 4098:3941891 4078:Bibcode 4070:Science 4055:2703462 3986:Bibcode 3963:3324702 3922:3898363 3902:Bibcode 3894:Science 3870:7260990 3847:Bibcode 3815:9778439 3795:Bibcode 3674:1461899 3634:6368675 3579:Bibcode 3547:3107637 3498:2729066 3477:Bibcode 3469:Science 3428:Bibcode 3379:Bibcode 3314:3073259 3254:5637615 3196:4477045 3175:Bibcode 3167:Science 3148:5219323 2965:6823321 2844:4448610 2706:4665078 2604:3414261 2583:Bibcode 2575:Science 2547:6615865 2496:4142721 2479:: 413. 2294:23 July 1899:Meiosis 1881:cohesin 1826:mitosis 1818:trisomy 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