query: when do the sister chromatids separate in meiosis

pos: Sister chromatids Sister chromatids A sister chromatid refers to the identical copies (chromatids) formed by the DNA replication of a chromosome, with both copies joined together by a common centromere. In other words, a sister chromatid may also be said to be 'one-half' of the duplicated chromosome. A pair of sister chromatids is called a dyad. A full set of sister chromatids is created during the synthesis (S) phase of interphase, when all the chromosomes in a cell are replicated. The two sister chromatids are separated from each other into two different cells during mitosis or during the second division of meiosis.

pos: Nondisjunction Nondisjunction Nondisjunction is the failure of homologous chromosomes or sister chromatids to separate properly during cell division. There are three forms of nondisjunction: failure of a pair of homologous chromosomes to separate in meiosis I, failure of sister chromatids to separate during meiosis II, and failure of sister chromatids to separate during mitosis. Nondisjunction results in daughter cells with abnormal chromosome numbers (aneuploidy). Calvin Bridges and Thomas Hunt Morgan are credited with discovering nondisjunction in "Drosophila melanogaster" sex chromosomes in the spring of 1910, while working in the Zoological Laboratory of Columbia University. In general, nondisjunction can occur in any

pos: Homologous chromosome the same in mitosis as they do in meiosis. Prior to every single mitotic division a cell undergoes, the chromosomes in the parent cell replicate themselves. The homologous chromosomes within the cell will ordinarily not pair up and undergo genetic recombination with each other. Instead, the replicants, or sister chromatids, will line up along the metaphase plate and then separate in the same way as meiosis II - by being pulled apart at their centromeres by nuclear mitotic spindles. If any crossing over does occur between sister chromatids during mitosis, it does not produce any new recombinant genotypes. Homologous pairing

pos: Sister chromatids Sister chromatids A sister chromatid refers to the identical copies (chromatids) formed by the DNA replication of a chromosome, with both copies joined together by a common centromere. In other words, a sister chromatid may also be said to be 'one-half' of the duplicated chromosome. A pair of sister chromatids is called a dyad. A full set of sister chromatids is created during the synthesis (S) phase of interphase, when all the chromosomes in a cell are replicated. The two sister chromatids are separated from each other into two different cells during mitosis or during the second division of meiosis.

pos: Chromatid during mitosis or meiosis. SCEs appear to primarily reflect DNA recombinational repair processes responding to DNA damage (see articles Sister chromatids and Sister chromatid exchange). "Non-sister chromatids", on the other hand, refers to either of the two chromatids of paired homologous chromosomes, that is, the pairing of a paternal chromosome and a maternal chromosome. In chromosomal crossovers, non-sister (homologous) chromatids form chiasmata to exchange genetic material during the prophase I of meiosis (See Homologous recombination). Chromatid A chromatid (Greek "khrōmat-" 'color' + "-id") is one copy of a newly copied chromosome which is still joined to the original chromosome by

pos: Chromosome segregation Chromosome segregation Chromosome segregation is the process in eukaryotes by which two sister chromatids formed as a consequence of DNA replication, or paired homologous chromosomes, separate from each other and migrate to opposite poles of the nucleus. This segregation process occurs during both mitosis and meiosis. During mitosis chromosome segregation occurs routinely as a step in cell division (see mitosis diagram). As indicated in the mitosis diagram, mitosis is preceded by a round of DNA replication, so that each chromosome forms two copies called chromatids. These chromatids separate to opposite poles, a process facilitated by a protein complex referred to

pos: Meiosis diploid cell, which contains two copies of each chromosome, termed homologs. First, the cell undergoes DNA replication, so each homolog now consists of two identical sister chromatids. Then each set of homologs pair with each other and exchange DNA by homologous recombination leading to physical connections (crossovers) between the homologs. In the first meiotic division, the homologs are segregated to separate daughter cells by the spindle apparatus. 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

pos: Meiosis of sister chromatids. Mechanically, the process is similar to mitosis, though its genetic results are fundamentally different. The end result is production of four haploid cells (n chromosomes, 23 in 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. In prophase II we see the disappearance of the nucleoli and the nuclear envelope again as well as the shortening and thickening of the chromatids. Centrosomes move to the polar regions and arrange spindle

pos: Nondisjunction II. In oocytes, one sister chromatid is segregated into the second polar body, while the other stays inside the egg. During spermatogenesis, each meiotic division is symmetric such that each primary spermatocyte gives rise to 2 secondary spermatocytes after meiosis I, and eventually 4 spermatids after meiosis II. Meiosis II-nondisjunction may also result in aneuploidy syndromes, but only to a much smaller extent than do segregation failures in meiosis I. Division of somatic cells through mitosis is preceded by replication of the genetic material in S phase. As a result, each chromosome consists of two sister chromatids held together at

pos: Origin and function of meiosis than the repair mechanism provided by meiosis. They primarily do not require a second homologous chromosome for the recombination that promotes a more extensive repair. Thus, despite the efficiency of recombinational repair involving sister chromatids, the repair still needs to be improved, and another type of repair is required. Moreover, due to the more extensive homologous recombinational repair in meiosis in comparison to the repair in mitosis, meiosis as a repair mechanism can accurately remove any damage that arises at any stage of the cell cycle more than mitotic repair mechanism can do and was, therefore, naturally selected. In contrast,

pos: Meiosis pair in addition to cohesin between sister chromatids. Kinetochore microtubules shorten, pulling homologous chromosomes (which 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 the cohesin from the chromosome arms is degraded while the cohesin surrounding the centromere remains protected. This allows the sister chromatids to remain together while homologs are segregated. 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

pos: Cohesin has been used to describe conditions affecting the cohesin complex. These conditions include: Cohesin Cohesin is a protein complex that regulates the separation of sister chromatids during cell division, either mitosis or meiosis. Cohesins hold sister chromatids together after DNA replication until anaphase when removal of cohesin leads to separation of sister chromatids. Cohesin is a multi-subunit protein complex, made up of four core subunits: two SMC proteins (SMC1 and 3), an alpha-kleisin (orthologues of yeast Scc1), and an orthologue of the yeast Scc3 protein (e.g. STAG1-3 in humans and SA1-3 in mice). Smc1 and Smc3 are members of the

pos: Cohesin Cohesin Cohesin is a protein complex that regulates the separation of sister chromatids during cell division, either mitosis or meiosis. Cohesins hold sister chromatids together after DNA replication until anaphase when removal of cohesin leads to separation of sister chromatids. Cohesin is a multi-subunit protein complex, made up of four core subunits: two SMC proteins (SMC1 and 3), an alpha-kleisin (orthologues of yeast Scc1), and an orthologue of the yeast Scc3 protein (e.g. STAG1-3 in humans and SA1-3 in mice). Smc1 and Smc3 are members of the Structural Maintenance of Chromosomes (SMC) family. SMC proteins have two main structural characteristics:

pos: Chromatid dyad. Once sister chromatids have separated (during the anaphase of mitosis or the anaphase II of meiosis during sexual reproduction), they are again called chromosomes. Although having the same genetic mass as the individual chromatids that made up its parent, the daughter "molecules" are called chromosomes in a similar way that one child of a pair of twins is not referred to as a single twin. The DNA sequence of two sister chromatids is completely identical (apart from very rare DNA copying errors). Sister chromatid exchange (SCE) is the exchange of genetic information between two sister chromatids. SCEs can occur

pos: Chiasma (genetics) Chiasma (genetics) In genetics, a chiasma (pl. chiasmata) is the point of contact, the physical link, between two (non-sister) chromatids belonging to homologous chromosomes. At a given chiasma, an exchange of genetic material can occur between both chromatids, what is called a chromosomal crossover, but this is much more frequent during meiosis than mitosis. In meiosis, absence of a chiasma generally results in improper chromosomal segregation and aneuploidy. The phenomenon of genetic chiasmata ("chiasmatypie") was discovered and described in 1909 by Frans Alfons Janssens, a Professor at the University of Leuven in Belgium. When each tetrad, which is composed of

pos: Anaphase lag Anaphase lag Anaphase lag is a consequence of an event during cell division where sister chromatids do not properly separate from each other because of improper spindle formation. The chromosome or chromatid does not properly migrate during anaphase and the daughter cells will lose some genetic information. It is one of many causes of aneuploidy. This event can occur during both meiosis and mitosis with unique repercussions. In either case, anaphase lag will cause one daughter cell to receive a complete set of chromosomes while the other lacks one paired set of chromosomes, creating a form of monosomy. Whether the

pos: Establishment of sister chromatid cohesion Establishment of sister chromatid cohesion Sister chromatid cohesion refers to the process by which sister chromatids are paired and held together during certain phases of the cell cycle. Establishment of sister chromatid cohesion is the process by which chromatin-associated cohesin protein becomes competent to physically bind together the sister chromatids. In general, cohesion is established during S phase as DNA is replicated, and is lost when chromosomes segregate during mitosis and meiosis. Some studies have suggested that cohesion aids in aligning the kinetochores during mitosis by forcing the kinetochores to face opposite cell poles. Cohesin first associates with the chromosomes

pos: Chiasma (genetics) increase in mean cell chiasma frequency. Chiasma frequency was scored at the later diplotene-diakinesis stages of meiosis. These results suggest that X-rays induce DNA damages, likely including double-strand breaks, and these damages are repaired by a crossover pathway leading to chiasma formation. Chiasma (genetics) In genetics, a chiasma (pl. chiasmata) is the point of contact, the physical link, between two (non-sister) chromatids belonging to homologous chromosomes. At a given chiasma, an exchange of genetic material can occur between both chromatids, what is called a chromosomal crossover, but this is much more frequent during meiosis than mitosis. In meiosis, absence of

pos: Kinesin 13 make full contact with the tubulin subunit. More MCAK molecules collectively bind to the curved region supporting the theory that they do not actively peel away the microtubule but they wait patiently for it to adopt this curved conformation. They stabilize the curved conformation by binding to the end of the microtubule and then catalyze depolymerization. The major function of mitosis is to separate replicated sister chromatids, and this is accomplished in part during anaphase A when "kinetochore microtubules (or kMTs)" that link the sister chromatids to opposite spindle poles shorten by depolymerization, exerting forces on the chromatids that pull

pos: Cell growth DNA molecule. During mitosis the double chromosomes are split to produce 92 "single chromosomes", half of which go into each daughter cell. During meiosis, there are two chromosome separation steps which assure that each of the four daughter cells gets one copy of each of the 23 types of chromosome. Though cell reproduction that uses mitosis can reproduce eukaryotic cells, eukaryotes bother with the more complicated process of meiosis because sexual reproduction such as meiosis confers a selective advantage. Notice that when meiosis starts, the two copies of sister chromatids number 2 are adjacent to each other. During this time,

pos: Isochromosome may appear monocentric with only one centromere, it is isodicentric with two centromeres very close to each other; resulting in a potential loss of genetic material found on the other arms. Misdivision of the centromere can also produce monocentric isochromosomes, but they are not as common as dicentric isochromosomes. A more common mechanism in the formation of isochromosomes is through the breakage and fusion of sister chromatids, most likely occurring in early anaphase of mitosis or meiosis. A double-stranded break in the pericentric region of the chromosome is repaired when the sister chromatids, each containing a centromere, are fused together.

pos: Isochromosome Isochromosomes can be created during mitosis and meiosis through a misdivision of the centromere or U-type strand exchange. Under normal separation of sister chromatids in metaphase, the centromere will divide longitudinally, or parallel to the long axis of the chromosome. An isochromosome is created when the centromere is divided transversely, or perpendicular to the long axis of the chromosome. The division is usually not occurring in the centromere itself, but in an area surrounding the centromere, also known as a pericentric region. It is proposed that these sites of exchange contain homologous sequences between sister chromatids. Although the resulting chromosome

pos: Kinetochore Kinetochore A kinetochore (, ) is a disc-shaped protein structure associated with duplicated chromatids in eukaryotic cells where the spindle fibers attach during cell division to pull sister chromatids apart. The kinetochore assembles on the centromere and links the chromosome to microtubule polymers from the mitotic spindle during mitosis and meiosis. Its proteins also help to hold the sister chromatids together and play a role in chromosome editing. "Monocentric" organisms, including vertebrates, fungi, and most plants, have a single centromeric region on each chromosome which assembles a single, localized kinetochore. "Holocentric" organisms, such as nematodes and some plants, assemble a

pos: Kinetochore be "bona fide" components of both structures. Two groups of proteins seem to be particularly important: kinesins which work like depolymerases, such as KinI kinesins; and proteins bound to MT (+) ends, +TIPs, promoting polymerization, perhaps antagonizing the depolymerases effect. Kinetochore A kinetochore (, ) is a disc-shaped protein structure associated with duplicated chromatids in eukaryotic cells where the spindle fibers attach during cell division to pull sister chromatids apart. The kinetochore assembles on the centromere and links the chromosome to microtubule polymers from the mitotic spindle during mitosis and meiosis. Its proteins also help to hold the sister chromatids

pos: Sister chromatid exchange timing as inter-homolog recombination, but with reduced (2- to 3-fold) yields of Holliday junction joint molecules. This study, and comparable evidence from other organisms (e.g. Peacock), indicates that inter-sister recombination occurs frequently during meiosis, and up to one-third of all recombination events occur between sister chromatids, although mainly by a pathway that does not involve Holliday junction intermediates. Sister chromatid exchange Sister chromatid exchange (SCE) is the exchange of genetic material between two identical sister chromatids. It was first discovered by using the Giemsa staining method on one chromatid belonging to the sister chromatid complex before anaphase in mitosis. The

pos: Meiosis 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. Meiosis I segregates homologous chromosomes, 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 "reductional division". Meiosis II is an "equational division" analogous to mitosis, in which the sister chromatids are segregated, creating four haploid daughter cells (1n,

pos: Nondisjunction form of cell division that involves ordered distribution of chromosomal material. Higher animals have three distinct forms of such cell divisions: Meiosis I and meiosis II are specialized forms of cell division occurring during generation of gametes (eggs and sperm) for sexual reproduction, mitosis is the form of cell division used by all other cells of the body. Ovulated eggs become arrested in metaphase II until fertilization triggers the second meiotic division. Similar to the segregation events of mitosis, the pairs of sister chromatids resulting from the separation of bivalents in meiosis I are further separated in anaphase of meiosis

pos: Katanin step in mitosis and meiosis. It has been shown that katanin is responsible for severing microtubules during M-phase in "Xenopus laevis". The disassembly of microtubules from their interphase structures is necessary to prepare the cell and the mitotic spindle for cell division. This regulation is indirect: MAP proteins, which protect the microtubules from being severed during interphase, dissociate and allow katanin to act. In addition, katanin is responsible for severing microtubules at the mitotic spindles when disassembly is required to segregate sister chromatids during anaphase. Similar results have been obtained in relation to katanin’s activity during meiosis in "C. elegans".

pos: Meiomitosis Meiomitosis In cell biology, Meiomitosis is an aberrant cellular division pathway through which daughter cells following mitosis inherit partially expressed meiotic machinery. During normal mitosis, sister chromatids separate into two identical daughter cells. In meiosis is a diploid cell (2N) duplicates its DNA once and undergoes two rounds of cell division to produce four haploid (1N), germ cells that promotes the recombination of large expanses of chromosomal DNA, potentially similar in size to those lost or gained in cancer. In meiotic division, homologous chromosomes are held together by cohesions, and after resolution of chiasma (the recombination crossover points with the

pos: Chromosome segregation ability depends on the microtubule motor dynein that regulates the movement of chromosomes to the poles of the meiotic spindle. Chromosome segregation Chromosome segregation is the process in eukaryotes by which two sister chromatids formed as a consequence of DNA replication, or paired homologous chromosomes, separate from each other and migrate to opposite poles of the nucleus. This segregation process occurs during both mitosis and meiosis. During mitosis chromosome segregation occurs routinely as a step in cell division (see mitosis diagram). As indicated in the mitosis diagram, mitosis is preceded by a round of DNA replication, so that each chromosome

pos: Meiomitosis to tumor and even from cell to cell. Research suggests that genomic instability affects the cancer cells' ability to metastasize. Cells able to effectively metastasize are more genomically stable than the rest of the local tumor bulk. Meiomitosis In cell biology, Meiomitosis is an aberrant cellular division pathway through which daughter cells following mitosis inherit partially expressed meiotic machinery. During normal mitosis, sister chromatids separate into two identical daughter cells. In meiosis is a diploid cell (2N) duplicates its DNA once and undergoes two rounds of cell division to produce four haploid (1N), germ cells that promotes the recombination of

pos: Spindle apparatus Spindle apparatus In cell biology, the spindle apparatus (or mitotic spindle) refers to the cytoskeletal structure of eukaryotic cells that forms during cell division to separate sister chromatids between daughter cells. It is referred to as the mitotic spindle during mitosis, a process that produces genetically identical daughter cells, or the meiotic spindle during meiosis, a process that produces gametes with half the number of chromosomes of the parent cell. Besides chromosomes, the spindle apparatus is composed of hundreds of proteins. Microtubules comprise the most abundant components of the machinery. Attachment of microtubules to chromosomes is mediated by kinetochores, which

pos: Spindle apparatus are determined by localization of Tricellular junctions localized at cell vertices. The spatial distribution of cortical clues leads to the force field that determine final spindle apparatus orientation and the subsequent orientation of cell division. Spindle apparatus In cell biology, the spindle apparatus (or mitotic spindle) refers to the cytoskeletal structure of eukaryotic cells that forms during cell division to separate sister chromatids between daughter cells. It is referred to as the mitotic spindle during mitosis, a process that produces genetically identical daughter cells, or the meiotic spindle during meiosis, a process that produces gametes with half the number of

pos: Meiosis cut and then repaired, which allows them to exchange some of their genetic information. A subset of recombination events results in crossovers, which create physical links known as chiasmata (singular: chiasma, for the Greek letter Chi (X)) between the homologous chromosomes. In most organisms, these links are essential to direct each pair of homologous chromosomes to segregate away from each other during Meiosis I, resulting in two haploid cells that have half the number of chromosomes as the parent cell. During Meiosis II, the cohesion between sister chromatids is released and they segregate from one another, as during mitosis. In

pos: Cohesin coupling, chromosome architecture and rearrangement, DNA replication etc. The anaphase promoting complex associated to Cdc20 (APC/C-cdc20) cleaves the securin (anaphase inhibitor), which holds the sister chromatids together. Securin is cleaved at anaphase, following APC/C-cdc20 mediated degradation, and it renders separase (a protease, inhibited by the association with securin) to cleave the kleisin subunit. An alpha-kleisin is associated with the cohesin complex, linking both SMC 3 and SMC 1 together, with the exact kleisin varying between mitosis and meiosis (Scc1 and Rec8 respectively), and its cleavage ultimately leads to the removal of cohesin from chromosomes. Dissociation of sister chromatids cohesion defines

pos: Prophase the spindle apparatus at locations called foci. The mitotic spindle is of great importance in the process of mitosis and will eventually segregate the sister chromatids in metaphase. The nucleoli begin to break down in prophase, resulting in the discontinuation of ribosome production. This indicates a redirection of cellular energy from general cellular metabolism to cellular division. The nuclear envelope stays intact during this process. Meiosis involves two rounds of chromosome segregation and thus undergoes prophase twice, resulting in prophase I and prophase II. Prophase I the most complex phase in all of meiosis because homologous chromosomes must pair and

pos: Meiosis attachment is referred to as a bipolar attachment. The physical basis of the independent assortment of chromosomes is the random orientation of each bivalent along the metaphase plate, with respect to the orientation of the other bivalents along the same equatorial line. The protein complex cohesin 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 meiosis, establishing tension requires at least one crossover per chromosome

pos: MAP2K1 to 3-fold) yields of joint molecules. MAP2K1 is also known as MEK1 (see Mitogen-activated protein kinase kinase). MEK1 is a meiotic chromosome-axis-associated kinase that is thought to slow down, but not entirely block, sister chromatid recombination. Loss of MEK1 allows inter-sister DSB repair and also inter-sister Holliday junction intermediates to increase. Despite the normal activity of MEK1 in reducing inter-sister chromatid recombination, such recombination still occurs frequently during normal budding yeast meiosis (although not as frequently as during mitosis), and up to one-third of all recombination events are between sister chromatids. MAP2K1 has been shown to interact with C-Raf, Phosphatidylethanolamine

pos: Spindle checkpoint phase in the cell cycle have been fulfilled are called checkpoints. All along the cell cycle, there are different checkpoints. The checkpoint ensuring that chromosome segregation is correct is termed spindle assembly checkpoint (SAC), spindle checkpoint or mitotic checkpoint. During mitosis or meiosis, the spindle checkpoint prevents anaphase onset until all chromosomes are properly attached to the spindle. To achieve proper segregation, the two kinetochores on the sister chromatids must be attached to opposite spindle poles (bipolar orientation). Only this pattern of attachment will ensure that each daughter cell receives one copy of the chromosome. When cells are ready to

pos: Shugoshin N terminal protein domain cohesion during cell division. It does this by preventing the cohesin complex which regulates chromatid separation from prematurely dissociating. Shugoshin protein is thought to act by protecting two proteins, named Rec8 and Rad21 at the centromeres from protein degradation by the enzyme, separase. This results in the sister chromatids remaining tethered. Shugoshin also acts as a spindle checkpoint component. It senses tension between sister chromatids during mitosis, and it degrades when they separate preventing cell cycle arrest and chromosome loss.Human shugoshin is diffusible and mediates formation of kinetochore-microtubules during bipolar spindle assembly. Recent studies indicate that the structure of the

pos: Condensin II’s contribution to resolving sister chromatids initiates as early as in S phase. Eukaryotic cells have two additional classes of SMC protein complexes. Cohesin contains SMC1 and SMC3 and is involved in sister chromatid cohesion. The SMC5/6 complex contains SMC5 and SMC6 and is implicated in recombinational repair. Condensin Condensins are large protein complexes that play a central role in chromosome assembly and segregation during mitosis and meiosis. Many eukaryotic cells possess two different types of condensin complexes, known as condensin I and condensin II, each of which is composed of five subunits. Condensins I and II share the same

pos: Cohesin of a hand-cuff model, the model is inconsistent with a number of experimental observations, and is generally considered to entrap chromatin as a monomer. The topology and structure of these subunits has been best characterized in budding yeast (Haering et al. 2002, 2004), but the sequence conservation of these proteins and biochemical and electron microscopic observations imply that cohesin complexes in other species are very similar in their structure, . The cohesin ring has many functions: 1. It is used to keep the sister chromatids connected with each other during metaphase ensuring that during mitosis (and meiosis), each sister chromatid

pos: Cell nucleus in open mitosis, the cell divides to form two cells. In order for this process to be possible, each of the new daughter cells must have a full set of genes, a process requiring replication of the chromosomes as well as segregation of the separate sets. This occurs by the replicated chromosomes, the sister chromatids, attaching to microtubules, which in turn are attached to different centrosomes. The sister chromatids can then be pulled to separate locations in the cell. In many cells, the centrosome is located in the cytoplasm, outside the nucleus; the microtubules would be unable to attach to

pos: Prophase the synaptonemal complex dissolves. It is at this stage where meiotic arrest occurs in many species. In the fifth and final phase of prophase I, diakinesis (from the Greek for “double movement”), full chromatin condensation has occurred and all four sister chromatids can be seen in bivalents with microscopy. The rest of the phase remesemble the early stages of mitotic prometaphase, as the meiotic prophase ends with the spindle apparatus beginning to form, and the nuclear membrane beginning to break down. Prophase II of meiosis is very similar to prophase of mitosis. The most noticeable difference is that prophase II

pos: Nondisjunction the centromere. In the anaphase of mitosis, sister chromatids separate and migrate to opposite cell poles before the cell divides. Nondisjunction during mitosis leads to one daughter receiving both sister chromatids of the affected chromosome while the other gets none. This is known as a chromatin bridge or an anaphase bridge. Mitotic nondisjunction results in somatic mosaicism, since only daughter cells originating from the cell where the nondisjunction event has occurred will have an abnormal number of chromosomes. Nondisjunction during mitosis can contribute to the development of some forms of cancer, e.g. retinoblastoma (see below). Chromosome nondisjunction in mitosis can

pos: Kinetochore kinetochore along the entire length of a chromosome. Kinetochores start, control, and supervise the striking movements of chromosomes during cell division. During mitosis, which occurs after chromosomes are duplicated in S phase, two sister chromatids are held together by a centromere. Each chromatid has its own kinetochore, which face in opposite directions and attach to opposite poles of the mitotic spindle apparatus. Following the transition from metaphase to anaphase, the sister chromatids separate from each other, and the individual kinetochores on each chromatid drive their movement to the spindle poles that will define the two new daughter cells. The kinetochore

pos: Synapsis by a protein complex along the whole length of the chromosomes called the synaptonemal complex. Sex chromosomes also undergo synapsis; however, the synaptonemal protein complex that holds the homologous chromosomes together is only present at one end of each sex chromosome. This is not to be confused with mitosis. Mitosis also has prophase, but does not ordinarily do pairing of two homologous chromosomes. When the non-sister chromatids intertwine, segments of chromatids with similar sequence may break apart and be exchanged in a process known as genetic recombination or "crossing-over". This exchange produces a chiasma, a region that is shaped like

neg: Sister chromatids is largely limited to interaction between nearby sister chromatids that are present in a cell subsequent to DNA replication but prior to cell division. Due to the special nearby relationship they share, sister chromatids are not only preferred over distant homologous chromatids as substrates for recominational repair, but have the capacity to repair more DNA damage than do homologs. Studies with the budding yeast "Saccharomyces cerevisiae" indicate that inter-sister recombination occurs frequently during meiosis, and up to one-third of all recombination events occur between sister chromatids. Sister chromatids A sister chromatid refers to the identical copies (chromatids) formed by the

neg: Homologous chromosome separated in meiosis I, the sister chromatids from each pair are separated. The two haploid(because the chromosome no. has reduced to half. Earlier two sets of chromosomes were present, but now each set exists in two different daughter cells that have arisen from the single diploid parent cell by meiosis I) daughter cells resulting from meiosis I undergo another cell division in meiosis II but without another round of chromosomal replication. The sister chromatids in the two daughter cells are pulled apart during anaphase II by nuclear spindle fibers, resulting in four haploid daughter cells. Homologous chromosomes do not function

neg: REC8 REC8 Meiotic recombination protein REC8 homolog is a protein that in humans is encoded by the "REC8" gene. Rec8 is a meiosis-specific component of the cohesin complex that binds sister chromatids in preparation for the two divisions of meiosis. Rec8 is sequentially removed from sister chromatids. It is removed from the arms of chromosomes in the first division - separating homologous chromosomes from each other. However, Rec8 is maintained at centromeres so that sister chromatids are kept joined until anaphase of meiosis II, at which point removal of remaining cohesin leads to the separation of sister chromatids. This gene encodes

neg: Establishment of sister chromatid cohesion Sororin also has chromatin binding activity independent of its ability to mediate cohesion. Cohesion proteins SMC1ß, SMC3, REC8 and STAG3 appear to participate in the cohesion of sister chromatids throughout the meiotic process in human oocytes. SMC1ß, REC8 and STAG3 are meiosis specific cohesin proteins. The STAG3 protein is essential for female meiosis and fertility. A growing body of evidence ties establishment of cohesion to DNA replication. As mentioned above, functional coupling of these two processes prevents the cell from having to later distinguish which chromosomes are sisters by ensuring that the sister chromatids are never separate after replication. Another

neg: Sister chromatids Compare sister chromatids to homologous chromosomes, which are the two "different" copies of a chromosome that diploid organisms (like humans) inherit, one from each parent. Sister chromatids are by and large identical (since they carry the same alleles, also called variants or versions, of genes) because they derive from one original chromosome. An exception is towards the end of meiosis, after crossing over has occurred, because sections of each sister chromatid may have been exchanged with corresponding sections of the homologous chromatids with which they are paired during meiosis. Homologous chromosomes might or might not be the same as each

neg: Meiosis further divided into Karyokinesis I and Cytokinesis I and 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. Interphase is divided into three phases: Interphase is followed by meiosis I and then meiosis II. Meiosis I separates 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 chromosomes are segregated into four daughter cells. For diploid organisms, the daughter cells resulting

neg: Meiosis fibers for the second meiotic division. In metaphase II, the centromeres contain two kinetochores 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. This is followed by anaphase II, in which the remaining centromeric cohesin is cleaved allowing the sister chromatids to segregate. 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 decondensation and lengthening

neg: Parthenogenesis in squamata sexual species, identical duplicate sister chromosomes, produced through premeiotic replication, pair and separate during meiosis I in true parthenotes. Pairing of identical sister chromosomes, in comparison to the alternative of pairing homologous chromosomes, maintains heterozygosity in obligate parthenotes. Meiosis II involves the separation of sister chromatids in both sexual and parthenogenetic species. This method of parthenogenesis is observed in obligate parthenotes, such as lizards in the genus "Cnemidophorus" and "Lacerta", and also in certain facultative parthenotes like the Burmese python. Another mechanism typically observed in facultative parthenote reptiles is terminal fusion, in which a haploid polar body produced as a

neg: Meiosis of the male testis suppress meiosis by degrading retinoic acid, a stimulator of meiosis. This is overcome at puberty when cells within seminiferous tubules called Sertoli cells start making their own retinoic acid. The normal separation of chromosomes in meiosis I or sister chromatids in meiosis II is termed "disjunction". When the segregation is not normal, it is called "nondisjunction". 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 trisomy or monosomy. Nondisjunction can occur in the meiosis I or meiosis II, phases of

neg: Meiotic recombination checkpoint spindle pole bodies can separate. The event is triggered by the activation of M-Cdk in late prophase I. Then the spindle assembly checkpoint examines the attachment of microtubules at kinetochores and APC can initiate the metaphase I. The special chromosome separation in meiosis, homologous chromosomes separation in meiosis I and chromatids separation in meiosis II, requires special tension between homologous chromatids and non-homologous chromatids for distinguishing microtubule attachment and it relies on the programmed DNA double strand break (DSB) and repair in prophase I. Therefore meiotic recombination checkpoint can be a kind of DNA damage response at specific time spot.

neg: Sister chromatid exchange preferred over distant homologous chromatids as substrates for recombinational repair, but have the capacity to repair more DNA damage than do homologs. The genomes of diploid organisms in natural populations are highly polymorphic for insertions and deletions. During meiosis double-strand breaks (DSBs) that form within such polymorphic regions must be repaired by inter-sister chromatid exchange, rather than by inter-homolog exchange. A molecular-level study of recombination during budding yeast meiosis has shown that recombination events initiated by DSBs in regions that lack corresponding sequences in the non-sister homolog are efficiently repaired by inter-sister chromatid recombination. This recombination occurs with the same

neg: Homology directed repair region to be repaired. In a diploid cell in G1 phase of the cell cycle, such a molecule is present in the form of the homologous chromosome. However, in the G2 stage of the cell cycle (following DNA replication), a second homologous DNA molecule is also present: the sister chromatid. Evidence indicates that, due to the special nearby relationship they share, sister chromatids are not only preferred over distant homologous chromatids as substrates for recombinational repair, but have the capacity to repair more DNA damage than do homologs. During meiosis up to one-third of all homology directed repair events occur

neg: Meiosis chromosome pairs along the metaphase plate during metaphase I & orientation of sister chromatids in metaphase II, this is the subsequent separation of homologs and sister chromatids during anaphase I & II, it allows a random and independent distribution of chromosomes to each daughter cell (and ultimately to gametes); and (2) Crossing Over. The physical exchange of homologous chromosomal regions by homologous recombination during prophase I results in new combinations of DNA within chromosomes. During meiosis, specific genes are more highly transcribed. In addition to strong meiotic stage-specific expression of mRNA, there are also pervasive translational controls (e.g. selective usage

neg: Klinefelter syndrome chromosome is through a nondisjunction event during meiosis II in the egg. Nondisjunction occurs when sister chromatids on the sex chromosome, in this case an X and an X, fail to separate. An XX egg is produced, which when fertilized with a Y sperm, yields an XXY offspring. This XXY chromosome arrangement is one of the most common genetic variations from the XY karyotype, occurring in about one in 500 live male births. See also Triple X syndrome. In mammals with more than one X chromosome, the genes on all but one X chromosome are not expressed; this is known

neg: Chromosome segregation as cohesin. Upon proper segregation, a complete set of chromatids ends up in each of two nuclei, and when cell division is completed, each DNA copy previously referred to as a chromatid is now called a chromosome. Chromosome segregation occurs at two separate stages during meiosis called prophase I and prophase II (see meiosis diagram). In a diploid cell there are two sets of homologous chromosomes of different parental origin (e.g. a paternal and a maternal set). During the phase of meiosis labeled “interphase s” in the meiosis diagram there is a round of DNA replication, so that each of

neg: Homologous chromosome crossing over occurs and throughout the process of chromosomal segregation during meiosis. Both the non-crossover and crossover types of recombination function as processes for repairing DNA damage, particularly double-strand breaks. At the diplotene stage of prophase I the synaptonemal complex disassembles before which will allow the homologous chromosomes to separate, while the sister chromatids stay associated by their centromeres. In metaphase I of meiosis I, the pairs of homologous chromosomes, also known as bivalents or tetrads, line up in a random order along the metaphase plate. The random orientation is another way for cells to introduce genetic variation. Meiotic spindles

neg: Meiomitosis other chromosome pair), the homologous chromosomes are segregated together. In these cells chiasma occurs and sister chromatids are partly held together by cohesions; when cell division occurs, the sister chromatids are sheared and mis-segregated which results in an aneuploid state. Melanoma may be prone to meiomitosis due to the high frequency of testis antigen that is expressed in tumor cells, two of which are known meiosis proteins. Melanoma tumor cells that replicate via meiomitosis may become immortal due to the expression of telomerase, but this comes at the cost of meiotic proteins and genomic instability. Instability is different from tumor

neg: Chromomere and their effects on genes near breakpoints. Chromomeres display different properties and behaviours when associated with lampbrush chromosomes. Although found all across the lampbrush chromosome, they are not organized in a clear pattern along as they are in normal pachytene chromosomes of meiosis. The two sister chromatids of a lampbrush chromosome separate fully, forming lateral loops that extend from chromomeres, and act as transcription complexes. The lateral loops are areas where the chromosomes are transcriptionally active. The loops define where the chromomere will be positioned. The chromomere is an organization of regions of non-transcribed chromatin. These regions of chromatin that

neg: Meiosis process of pairing the homologous chromosomes is called synapsis. At this stage, non-sister chromatids may cross-over at points called chiasmata (plural; singular chiasma). Prophase I has historically been divided into a series of substages which are named according to the appearance of chromosomes. The first stage of prophase I is the "leptotene" stage, also known as "leptonema", from Greek words meaning "thin threads".In this stage of prophase I, individual chromosomes—each consisting of two sister chromatids—become "individualized" to form visible strands within the nucleus. The two sister chromatids closely associate and are visually indistinguishable from one another. During leptotene, lateral elements

neg: Meiosis chromosome consists of a pair of chromatids. 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. Sister chromatids remain attached during telophase I. Cells may enter a period of rest known as interkinesis or interphase II. No DNA replication occurs during this stage. Meiosis II is the second meiotic division, and usually involves equational segregation, or separation

neg: Meiosis of chromosomes as the original parent cell. The two meiotic divisions are known as Meiosis I and Meiosis II. Before meiosis begins, during S phase of the cell cycle, the DNA of each chromosome is replicated so that it consists of two identical sister chromatids, 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 G2-like stage known as meiotic prophase. During this time, homologous chromosomes pair with each other and undergo genetic recombination, a programmed process in which DNA is

neg: Homologous chromosome separating the homologous chromosomes in meiosis I and then the sister chromatids in meiosis II. The process of meiosis I is generally longer than meiosis II because it takes more time for the chromatin to replicate and for the homologous chromosomes to be properly oriented and segregated by the processes of pairing and synapsis in meiosis I. During meiosis, genetic recombination (by random segregation) and crossing over produces daughter cells that each contain different combinations of maternally and paternally coded genes. This recombination of genes allows for the introduction of new allele pairings and genetic variation. Genetic variation among organisms

neg: Immature ovum the total quantity of chromatids 46. This is twice the number of chromosome pairs (2N). Oogonia are created in early embryonic life. All have turned into primary oocytes at late fetal age. The primary oocyte is defined by its process of ootidogenesis, which is meiosis. It has duplicated its DNA, so that each chromosome has two chromatids, i.e. 92 chromatids all in all (4C). When meiosis I is completed, one secondary oocyte and one polar body is created. Primary oocytes have been created in late fetal life. This is the stage where immature ova spend most of their lifetime, more

neg: Spindle checkpoint in one part of the cell, this information is extended all along the cytoplasm, and can overcome the signal "wait to enter in anaphase" associated to a second spindle containing unattached kinetochores. As it has been previously noted, sister chromatids stay associated from S phase (when DNA is replicated to generate two identic copies, the two chromatids) until anaphase. At this point, the two sister chromatids separate and travel to opposite poles in the dividing cell. Genetic and biochemical studies in yeast and in egg's extracts in "Xenopus laevis" identified a polyprotein complex as an essential player in sister chromatids

neg: Parthenogenesis because at anaphase II the sister chromatids are separated and whatever heterozygosity is present is due to crossing over. In the case of endomitosis after meiosis the offspring is completely homozygous and has only half the mother's genetic material. This can result in parthenogenetic offspring being unique from each other and from their mother. In apomictic parthenogenesis, the offspring are clones of the mother and hence (except for aphids) are usually female. In the case of aphids, parthenogenetically produced males and females are clones of their mother except that the males lack one of the X chromosomes (XO). When meiosis

neg: Chiasma (genetics) two pairs of sister chromatids, begins to split, the only points of contact are at the chiasmata. The chiasmata become visible during the diplotene stage of prophase I of meiosis, but the actual "crossing-overs" of genetic material are thought to occur during the previous pachytene stage. Sister chromatids also form chiasmata between each other (also known as a chi structure), but because their genetic material is identical, it does not cause any noticeable change in the resulting daughter cells. In humans, there seems to be one chiasma per chromosome arm, and in mammals, the number of chromosome arms is a

neg: Chromosome segregation DNA replication. Thus the two chromatids comprising each chromosome separate into different nuclei, so that each nucleus gets a single set of chromatids (now called chromosomes) and each nucleus becomes included in a haploid gamete (see stages following prophase II in the meiosis diagram). This segregation process is also facilitated by cohesin. Failure of proper segregation during prophase II can also lead to aneuploid gametes. Aneuploid gametes can undergo fertilization to form aneuploid zygotes and hence to serious adverse consequences for progeny. Meiotic chromosomal crossover (CO) recombination facilitates the proper segregation of homologous chromosomes. This is because, at the end

neg: Homologous chromosome emanating from opposite spindle poles attach to each of the homologs (each pair of sister chromatids) at the kinetochore. In anaphase I of meiosis I the homologous chromosomes are pulled apart from each other. The homologs are cleaved by the enzyme separase to release the cohesin that held the homologous chromosome arms together. This allows the chiasmata to release and the homologs to move to opposite poles of the cell. The homologous chromosomes are now randomly segregated into two daughter cells that will undergo meiosis II to produce four haploid daughter germ cells. After the tetrads of homologous chromosomes are

neg: Leptotene stage Leptotene stage The leptotene stage, also known as the leptonema, is the first of five substages of prophase I in meiosis. The term "leptonema" derives from Greek words meaning "thin threads". A cell destined to become a gamete enters the leptotene stage after its chromosomes are duplicated during interphase. During the leptotene stage those duplicated chromosomes—each consisting of two sister chromatids—condense from diffuse chromatin into long, thin strands that are more visible within the nucleoplasm. The next stage of prophase I in meiosis is the zygotene stage. During this stage, the chromosomes attach themselves by their ends (telomeres) to the

neg: Establishment of sister chromatid cohesion Tying establishment to DNA replication allows the cell to institute cohesion as soon as the sister chromatids are formed. This solves the problem of how the cell might properly identify and pair sister chromatids by ensuring that the sister chromatids are never separate once replication has occurred. The Eco1/Ctf7 gene (yeast) was one of the first genes to be identified as specifically required for the establishment of cohesion. Eco1 must be present in S phase to establish cohesion, but its continued presence is not required to maintain cohesion. Eco1 interacts with many proteins directly involved in DNA replication, including the

neg: Crossover value fraction when the distance between the markers in question is short. Crossover value In genetics, the crossover value is the linked frequency of chromosomal crossover between two gene loci (markers). For a fixed set of genetic and environmental conditions, recombination in a particular region of a linkage structure (chromosome) tends to be constant and the same is then true for the crossover value which is used in the production of genetic map. Crossover implies the exchange of chromosomal segments between non-sister chromatids, in meiosis during the production of gametes. The effect is to assort the alleles on parental chromosomes, so

neg: Parthenogenesis mother's genetic material and heterozygosity is mostly preserved (if the mother has two alleles for a locus, it is likely that the offspring will get both). This is because in anaphase I the homologous chromosomes are separated. Heterozygosity is not completely preserved when crossing over occurs in central fusion. In the case of pre-meiotic doubling, recombination -if it happens- occurs between identical sister chromatids. If "terminal fusion" (restitutional meiosis of anaphase II or the fusion of its products) occurs, a little over half the mother's genetic material is present in the offspring and the offspring are mostly homozygous. This is

neg: Chromosomal crossover Chromosomal crossover Chromosomal crossover (or crossing over) is the exchange of genetic material between 2 homologous chromosomes non-sister chromatids that results in recombinant chromosomes during sexual reproduction. It is one of the final phases of genetic recombination, which occurs in the "pachytene" stage of prophase I of meiosis during a process called synapsis. Synapsis begins before the synaptonemal complex develops and is not completed until near the end of prophase I. Crossover usually occurs when matching regions on matching chromosomes break and then reconnect to the other chromosome. Crossing over was described, in theory, by Thomas Hunt Morgan. He relied

neg: Chromosome segregation the chromosomes initially present is now composed of two copies called chromatids. These chromosomes (paired chromatids) then pair with the homologous chromosome (also paired chromatids) present in the same nucleus (see prophase I in the meiosis diagram). The process of alignment of paired homologous chromosomes is called synapsis (see Synapsis). During synapsis, genetic recombination usually occurs. Some of the recombination events occur by crossing over (involving physical exchange between two chromatids), but most recombination events involve information exchange but not physical exchange between two chromatids (see Synthesis-dependent strand annealing (SDSA)). Following recombination, chromosome segregation occurs as indicated by the stages

neg: Schizosaccharomyces pombe RTS1, a site-specific replication pause site MPS1, and a novel type of chromosomal imprint, marking one of the sister chromatids at the mating-type locus mat1. In addition, work on the silenced donor region has led to great advances in understanding formation and maintenance of heterochromatin. "Schizosaccharomyces pombe" is a facultative sexual microorganism that can undergo mating when nutrients are limiting. Exposure of "S. pombe" to hydrogen peroxide, an agent that causes oxidative stress leading to oxidative DNA damage, strongly induces mating and formation of meiotic spores. This finding suggests that meiosis, and particularly meiotic recombination, may be an adaptation for

neg: Immature ovum specifically in prophase I of meiosis. The halt is called dictyate. Most degenerate by atresia, but a few go through ovulation, and that's the trigger to the next step. Thus, an immature ovum can spend up to ~55 years as a primary oocyte (the last ovulation before menopause). The secondary oocyte is the cell that is formed by meiosis I in oogenesis. Thus, it has only one of each pair of homologous chromosomes. In other words, it is haploid. However, each chromosome still has two chromatids, making a total of 46 chromatids (1N but 2C). The secondary oocyte continues the

neg: Breakage-fusion-bridge cycle Breakage-fusion-bridge cycle Breakage-fusion-bridge (BFB) cycle (also breakage-rejoining-bridge cycle) is a mechanism of chromosomal instability, discovered by Barbara McClintock in the late 1930s. The BFB cycle begins when the end region of a chromosome, called its telomere, breaks off. When that chromosome subsequently replicates it forms two sister chromatids which both lack a telomere. Since telomeres appear at the end of chromatids, and function to prevent their ends from fusing with other chromatids, the lack of a telomere on these two sister chromatids causes them to fuse with one another. During anaphase the sister chromatids will form a bridge where the

neg: Crossover value Crossover value In genetics, the crossover value is the linked frequency of chromosomal crossover between two gene loci (markers). For a fixed set of genetic and environmental conditions, recombination in a particular region of a linkage structure (chromosome) tends to be constant and the same is then true for the crossover value which is used in the production of genetic map. Crossover implies the exchange of chromosomal segments between non-sister chromatids, in meiosis during the production of gametes. The effect is to assort the alleles on parental chromosomes, so that the gametes carry "recombinations" of genes different from either parent.

neg: Bivalent (genetics) highlighting the robustness of the chiasma and the cohesion that hold the bivalents together. Bivalent (genetics) A bivalent, sometimes referred to as a tetrad, is the association of a pair of homologous chromosomes physically held together by at least one DNA crossover. This physical attachment allows for alignment and segregation of the homologous chromosomes in the first meiotic division. The formation of a bivalent occurs during the first division of meiosis (in the pachynema stage of meiotic prophase 1). In most organisms, each replicated chromosome (composed of two identical sister chromatids) elicits formation of DNA double-strand breaks during the leptotene

neg: Bivalent (genetics) Bivalent (genetics) A bivalent, sometimes referred to as a tetrad, is the association of a pair of homologous chromosomes physically held together by at least one DNA crossover. This physical attachment allows for alignment and segregation of the homologous chromosomes in the first meiotic division. The formation of a bivalent occurs during the first division of meiosis (in the pachynema stage of meiotic prophase 1). In most organisms, each replicated chromosome (composed of two identical sister chromatids) elicits formation of DNA double-strand breaks during the leptotene phase. These breaks are repaired by homologous recombination, that uses the homologous chromosome as

neg: Synaptonemal complex Synaptonemal complex The synaptonemal complex (SC) is a protein structure that forms between homologous chromosomes (two pairs of sister chromatids) during meiosis and is thought to mediate chromosome pairing, synapsis, and recombination. It is now evident that the synaptonemal complex is not required for genetic recombination in some organisms. For instance, in protozoan ciliates such as "Tetrahymena thermophila" and "Paramecium tetraurelia" genetic crossover does not appear to require synaptonemal complex formation. Research has shown that not only does the SC form after genetic recombination but mutant yeast cells unable to assemble a synaptonemal complex can still engage in the exchange

neg: Chromosome segregation of meiotic prophase I, CO recombination provides a physical link that holds homologous chromosome pairs together. These linkages are established by chiasmata, which are the cytological manifestations of CO recombination. Together with cohesion linkage between sister chromatids, CO recombination may help ensure the orderly segregation of the paired homologous chromosomes to opposite poles. In support of this, a study of aneuploidy in single spermatozoa by whole genome sequencing found that, on average, human sperm cells with aneuploid autosomes exhibit significantly fewer crossovers than normal cells. After the first chromosome segregation in meiosis I is complete, there is further chromosome segregation

neg: Meiosis of homology. Sex chromosomes, however, are not wholly identical, and only exchange information over a small region of homology. At the sites where exchange happens, chiasmata form. 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, and chiasmata are not visible until the next stage. During the "diplotene" stage, also

neg: Bivalent (genetics) a template for repair. The search for the homologous target, helped by numerous proteins collectively referred as the synaptonemal complex, cause the two homologs to pair, between the leptotene and the pachytene phases of meiosis I. Resolution of the DNA recombination intermediate into a crossover exchanges DNA segments between the two homologous chromosomes at a site called a chiasma (plural: chiasmata). This physical strand exchange and the cohesion between the sister chromatids along each chromosome ensure robust pairing of the homologs in diplotene phase. The structure, visible by microscopy, is called a bivalent. A bivalent is the association of two

neg: Separase of STAG1 or STAG2 in the cohesin complex. The second step involves cleavage of the cohesin subunit SCC1 (RAD21) by separase, which initiates the final separation of sister chromatids. In "S. cerevisiae", Esp1 is coded by ESP1 and is regulated by the securin Pds1. The two sister chromatids are initially bound together by the cohesin complex until the beginning of anaphase, at which point the mitotic spindle pulls the two sister chromatids apart, leaving each of the two daughter cells with an equivalent number of sister chromatids. The proteins that bind the two sister chromatids, disallowing any premature sister chromatid

neg: Condensin is present in the cytoplasm during interphase, and gains access to chromosomes only after the nuclear envelope breaks down at the end of prophase. During prometaphase and metaphase, both condensin I and condensin II contribute to the assembly of condensed chromosomes, in which two sister chromatids are fully resolved. The two complexes apparently stay associated with chromosomes after the sister chromatids separate from each other in anaphase. At least one of the subunits of condensin I is known to be a direct target of a cyclin-dependent kinase (Cdk). Recent studies have shown that condensins participate in a wide variety of

neg: Chromatid identical, and said to be homozygous; however, if mutation(s) occur, they will present slight differences, in which case they are heterozygous. The pairing of chromatids should not be confused with the ploidy of an organism, which is the number of homologous versions of a chromosome. Chromonema is the fibre-like structure in prophase in the primary stage of DNA condensation. In metaphase, they are called chromatids. Chromatids may be sister or non-sister chromatids. A sister chromatid is either one of the two chromatids of the same chromosome joined together by a common centromere. A pair of sister chromatids is called a

neg: Breakage-fusion-bridge cycle centromere in one of the sister chromatids will be pulled in one direction of the dividing cell, while the centromere of the other will be pulled in the opposite direction. Being pulled in opposite directions will cause the two sister chromatids to break apart from each other, but not necessarily at the site that they fused. This results in the two daughter cells receiving an uneven chromatid. Since the two resulting chromatids lack telomeres, when they replicate the BFB cycle will repeat, and will continue every subsequent cell division until those chromatids receive a telomere, usually from a different chromatid

neg: Meiosis 1c). Prophase I is typically the longest phase of meiosis. During prophase I, homologous chromosomes pair and exchange DNA (homologous recombination). This often results in chromosomal crossover. This process is critical for pairing between homologous chromosomes and hence for accurate segregation of the chromosomes at the first meiosis division. The new combinations of DNA created during crossover are a significant source of genetic variation, and result in new combinations of alleles, which may be beneficial. The paired and replicated chromosomes are called bivalents or tetrads, which have two chromosomes and four chromatids, with one chromosome coming from each parent. The

neg: Chromosome once (S phase), and the copy is joined to the original by a centromere, resulting either in an X-shaped structure (pictured to the right) if the centromere is located in the middle of the chromosome or a two-arm structure if the centromere is located near one of the ends. The original chromosome and the copy are now called sister chromatids. During metaphase the X-shape structure is called a metaphase chromosome. In this highly condensed form chromosomes are easiest to distinguish and study. In animal cells, chromosomes reach their highest compaction level in anaphase during chromosome segregation. Chromosomal recombination during meiosis

neg: River Out of Eden to shrink until we find the "most recent common ancestor" (MRCA) of all humans alive in 1995 "via this particular gene pathway". In theory, one can also trace human ancestry via a single chromosome, as a chromosome contains a set of genes and is passed down from parents to children via independent assortment from only one of the two parents. But genetic recombination (chromosomal crossover) mixes genes from non-sister chromatids from both parents during meiosis, thus muddling the ancestry path. However, the mitochondrial DNA (mtDNA) is immune to sexual mixing, unlike the nuclear DNA whose chromosomes are shuffled and recombined

neg: Synaptonemal complex genetics, involved dosing C. elegans worms with that toxin and then verifying the results in mouse studies). by 3D-Structured Illumination, photograph by Dr. Chung-Ju Rachel Wang University of California Berkeley, Department of Molecular and Cell Biology Berkeley, CA, USA, second place winner of the 2009 Olympus Bioscapes Digital Imaging Competition. Synaptonemal complex The synaptonemal complex (SC) is a protein structure that forms between homologous chromosomes (two pairs of sister chromatids) during meiosis and is thought to mediate chromosome pairing, synapsis, and recombination. It is now evident that the synaptonemal complex is not required for genetic recombination in some organisms. For

neg: Most recent common ancestor that are passed down from parents to children via independent assortment from only one of the two parents, genetic recombination (chromosomal crossover) mixes genes from non-sister chromatids from both parents during meiosis, thus changing the genetic composition of the chromosome. Different types of MRCAs are estimated to have lived at different times in the past. These "time to MRCA" ("TMRCA") estimates are also computed differently depending on the type of MRCA being considered. Patrilineal and matrilineal MRCAs (Mitochondrial Eve and Y-chromosomal Adam) are traced by single gene markers, thus their TMRCA are computed based on DNA test results and established

neg: Synapsis the purging of the detrimental. Following synapsis, a type of recombination referred to as synthesis dependent strand annealing (SDSA) occurs frequently. SDSA recombination involves information exchange between paired non-sister homologous chromatids, but not physical exchange. SDSA recombination does not cause crossing-over. Both the non-crossover and crossover types of recombination function as processes for repairing DNA damage, particularly double-strand breaks (see Genetic recombination). The central function of synapsis is therefore the identification of homologues by pairing, an essential step for a successful meiosis. The processes of DNA repair and chiasma formation that take place following synapsis have consequences at many levels,