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Mitosis
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Not to be confused with Meiosis, Miosis, Myiasis, Myositis, or Myosotis.
Mitosis in an animal cell (phases ordered counter-clockwise).
Mitosis divides the chromosomes in a cell nucleus.
Onion (Allium) cells in different phases of the cell cycle enlarged 800 diameters.
a. non-dividing cells
b. nuclei preparing for division (spireme-stage)
c. dividing cells showing mitotic figures
e. pair of daughter-cells shortly after division
In cell biology, mitosis is a part of the cell cycle when replicated chromosomes are separated into two new nuclei. In general, mitosis (division of the nucleus) is preceded by the S stage of interphase (during which the DNA is replicated) and is often accompanied or followed by cytokinesis, which divides the cytoplasm, organelles and cell membrane into two new cells containing roughly equal shares of these cellular components.[1] Mitosis and cytokinesis together define the mitotic (M) phase of an animal cell cycle—the division of the mother cell into two daughter cells genetically identical to each other.
The process of mitosis is divided into stages corresponding to the completion of one set of activities and the start of the next. These stages are prophase, prometaphase, metaphase, anaphase, and telophase. During mitosis, the chromosomes, which have already duplicated, condense and attach to spindle fibers that pull one copy of each chromosome to opposite sides of the cell.[2] The result is two genetically identical daughter nuclei. The rest of the cell may then continue to divide by cytokinesis to produce two daughter cells.[3] Producing three or more daughter cells instead of normal two is a mitotic error called tripolar mitosis or multipolar mitosis (direct cell triplication / multiplication).[4] Other errors during mitosis can induce apoptosis (programmed cell death) or cause mutations. Certain types of cancer can arise from such mutations.[5]
Mitosis occurs only in eukaryotic cells. Prokaryotic cells, which lack a nucleus, divide by a different process called binary fission. Mitosis varies between organisms.[6] For example, animal cells undergo an "open" mitosis, where the nuclear envelope breaks down before the chromosomes separate, whereas fungi undergo a "closed" mitosis, where chromosomes divide within an intact cell nucleus.[7] Most animal cells undergo a shape change, known as mitotic cell rounding, to adopt a near spherical morphology at the start of mitosis. Most human cells are produced by mitotic cell division. Important exceptions include the gametes – sperm and egg cells – which are produced by meiosis.
Contents
[hide]
1 Discovery
2 Overview
3 Phases of cell cycle and mitosis
3.1 Interphase
3.2 Mitosis
3.2.1 Preprophase (plant cells)
3.2.2 Prophase
3.2.3 Prometaphase
3.2.4 Metaphase
3.2.5 Anaphase
3.2.6 Telophase
3.3 Cytokinesis
4 Significance
5 Cell shape changes during mitosis
6 Errors and variations
7 Mitotic recombination
8 Timeline in pictures
9 See also
10 References
11 Further reading
12 External links
Discovery[edit]
Cell division was first discovered by the German botanist Hugo von Mohl in 1835 as he worked with the green alga Cladophora glomerata.[8] The German zoologist Otto Bütschli might have claimed the discovery of the process presently known as "mitosis",[9] a term coined by Walther Flemming in 1882.[10]
Mitosis was discovered in frog, rabbit, and cat cornea cells in 1873 and described for the first time by the Polish histologist Wacław Mayzel in 1875.[11][12] The term is derived from the Greek word μίτος mitos "warp thread".[13][14]
Overview[edit]
Types of mitosis
Play media
Time-lapse video of mitosis in a Drosophila melanogaster embryo
The primary result of mitosis and cytokinesis is the transfer of a parent cell's genome into two daughter cells. The genome is composed of a number of chromosomes—complexes of tightly coiled DNA that contain genetic information vital for proper cell function. Because each resultant daughter cell should be genetically identical to the parent cell, the parent cell must make a copy of each chromosome before mitosis. This occurs during the S phase of interphase.[15] Chromosome duplication results in two identical sister chromatids bound together by cohesin proteins at the centromere.
When mitosis begins, the chromosomes condense and become visible. In some eukaryotes, for example animals, the nuclear envelope, which segregates the DNA from the cytoplasm, disintegrates into small vesicles. The nucleolus, which makes ribosomes in the cell, also disappears. Microtubules project from opposite ends of the cell, attach to the centromeres, and align the chromosomes centrally within the cell. The microtubules then contract to pull the sister chromatids of each chromosome apart.[16] Sister chromatids at this point are called daughter chromosomes. As the cell elongates, corresponding daughter chromosomes are pulled toward opposite ends of the cell and condense maximally in late anaphase. A new nuclear envelope forms around the separated daughter chromosomes, which decondense to form interphase nuclei.
During mitotic progression, typically after the anaphase onset, the cell may undergo cytokinesis. In animal cells, a cell membrane pinches inward between the two developing nuclei to produce two new cells. In plant cells, a cell plate forms between the two nuclei. Cytokinesis does not always occur; coenocytic (a type of multinucleate condition) cells undergo mitosis without cytokinesis.
Phases of cell cycle and mitosis[edit]

Interphase[edit]
Main article: Interphase
The mitotic phase is a relatively short period of the cell cycle. It alternates with the much longer interphase, where the cell prepares itself for the process of cell division. Interphase is divided into three phases: G1 (first gap), S (synthesis), and G2 (second gap). During all three parts of interphase, the cell grows by producing proteins and cytoplasmic organelles. However, chromosomes are replicated only during the S phase. Thus, a cell grows (G1), continues to grow as it duplicates its chromosomes (S), grows more and prepares for mitosis (G2), and finally divides (M) before restarting the cycle.[15] All these phases in the cell cycle are highly regulated by cyclins, cyclin-dependent kinases, and other cell cycle proteins. The phases follow one another in strict order and there are "checkpoints" that give the cell cues to proceed from one phase to another. Cells may also temporarily or permanently leave the cell cycle and enter G0 phase to stop dividing. This can occur when cells become overcrowded (density-dependent inhibition) or when they differentiate to carry out specific functions for the organism, as is the case for human heart muscle cells and neurons. Some G0 cells have the ability to re-enter the cell cycle.
Mitosis[edit]
Preprophase (plant cells)[edit]
Main article: Preprophase
In plant cells only, prophase is preceded by a pre-prophase stage. In highly vacuolated plant cells, the nucleus has to migrate into the center of the cell before mitosis can begin. This is achieved through the formation of a phragmosome, a transverse sheet of cytoplasm that bisects the cell along the future plane of cell division. In addition to phragmosome formation, preprophase is characterized by the formation of a ring of microtubules and actin filaments (called preprophase band) underneath the plasma membrane around the equatorial plane of the future mitotic spindle. This band marks the position where the cell will eventually divide. The cells of higher plants (such as the flowering plants) lack centrioles; instead, microtubules form a spindle on the surface of the nucleus and are then organized into a spindle by the chromosomes themselves, after the nuclear envelope breaks down.[17] The preprophase band disappears during nuclear envelope breakdown and spindle formation in prometaphase.[18]:58–67
Prophase[edit]
Main article: Prophase
During prophase, which occurs after G2 interphase, the cell prepares to divide by tightly condensing its chromosomes and initiating mitotic spindle formation. During interphase, the genetic material in the nucleus consists of loosely packed chromatin. At the onset of prophase, chromatin fibers condense into discrete chromosomes that are typically visible at high magnification through a light microscope. In this stage, chromosomes are long, thin and thread-like. Each chromosome has two chromatids. The two chromatids are joined at the centromere.
Gene transcription ceases during prophase and does not resume until late anaphase to early G1 phase.[19][20][21] The nucleolus also disappears during early prophase.[22]
Condensing chromosomes. Interphase nucleus (left), condensing chromosomes (middle) and condensed chromosomes (right).
Close to the nucleus of animal cells are structures called centrosomes, consisting of a pair of centrioles surrounded by a loose collection of proteins. The centrosome is the coordinating center for the cell's microtubules. A cell inherits a single centrosome at cell division, which is duplicated by the cell before a new round of mitosis begins, giving a pair of centrosomes. The two centrosomes polymerize tubulin to help form a microtubule spindle apparatus. Motor proteins then push the centrosomes along these microtubules to opposite sides of the cell. Although centrosomes help organize microtubule assembly, they are not essential for the formation of the spindle apparatus, since they are absent from plants,[17] and are not absolutely required for animal cell mitosis.[23]
Prometaphase[edit]
Main article: Prometaphase
At the beginning of prometaphase in animal cells, phosphorylation of nuclear lamins causes the nuclear envelope to disintegrate into small membrane vesicles. As this happens, microtubules invade the nuclear space. This is called open mitosis, and it occurs in some multicellular organisms. Fungi and some protists, such as algae or trichomonads, undergo a variation called closed mitosis where the spindle forms inside the nucleus, or the microtubules penetrate the intact nuclear envelope.[24][25]
In late prometaphase, kinetochore microtubules begin to search for and attach to chromosomal kinetochores.[26] A kinetochore is a proteinaceous microtubule-binding structure that forms on the chromosomal centromere during late prophase.[26][27] A number of polar microtubules find and interact with corresponding polar microtubules from the opposite centrosome to form the mitotic spindle.[28] Although the kinetochore structure and function are not fully understood, it is known that it contains some form of molecular motor.[29] When a microtubule connects with the kinetochore, the motor activates, using energy from ATP to "crawl" up the tube toward the originating centrosome. This motor activity, coupled with polymerisation and depolymerisation of microtubules, provides the pulling force necessary to later separate the chromosome's two chromatids.[29]
Metaphase[edit]
A cell in late metaphase. All chromosomes (blue) but one have arrived at the metaphase plate.
Main article: Metaphase
After the microtubules have located and attached to the kinetochores in prometaphase, the two centrosomes begin pulling the chromosomes towards opposite ends of the cell. The resulting tension causes the chromosomes to align along the metaphase plate or equatorial plane, an imaginary line that is centrally located between the two centrosomes (at approximately the midline of the cell).[28] To ensure equitable distribution of chromosomes at the end of mitosis, the metaphase checkpoint guarantees that kinetochores are properly attached to the mitotic spindle and that the chromosomes are aligned along the metaphase plate.[30] If the cell successfully passes through the metaphase checkpoint, it proceeds to anaphase.
Anaphase[edit]
Main article: Anaphase
During anaphase A, the cohesins that bind sister chromatids together are cleaved, forming two identical daughter chromosomes.[31] Shortening of the kinetochore microtubules pulls the newly formed daughter chromosomes to opposite ends of the cell. During anaphase B, polar microtubules push against each other, causing the cell to elongate.[32] In late anaphase, chromosomes also reach their overall maximal condensation level, to help chromosome segregation and the re-formation of the nucleus.[33] In most animal cells, anaphase A precedes anaphase B, but some vertebrate egg cells demonstrate the opposite order of events.[31]
Telophase[edit]
Main article: Telophase
Telophase (from the Greek word τελος meaning "end") is a reversal of prophase and prometaphase events. At telophase, the polar microtubules continue to lengthen, elongating the cell even more. If the nuclear envelope has broken down, a new nuclear envelope forms using the membrane vesicles of the parent cell's old nuclear envelope. The new envelope forms around each set of separated daughter chromosomes (though the membrane does not enclose the centrosomes) and the nucleolus reappears. Both sets of chromosomes, now surrounded by new nuclear membrane, begin to "relax" or decondense. Mitosis is complete. Each daughter nucleus has an identical set of chromosomes. Cell division may or may not occur at this time depending on the organism.
Cytokinesis[edit]
Cilliate undergoing cytokinesis, with the cleavage furrow being clearly visible
Main article: Cytokinesis
Cytokinesis is not a phase of mitosis but rather a separate process, necessary for completing cell division. In animal cells, a cleavage furrow (pinch) containing a contractile ring develops where the metaphase plate used to be, pinching off the separated nuclei.[34] In both animal and plant cells, cell division is also driven by vesicles derived from the Golgi apparatus, which move along microtubules to the middle of the cell.[35] In plants, this structure coalesces into a cell plate at the center of the phragmoplast and develops into a cell wall, separating the two nuclei. The phragmoplast is a microtubule structure typical for higher plants, whereas some green algae use a phycoplast microtubule array during cytokinesis.[18]:64–7, 328–9 Each daughter cell has a complete copy of the genome of its parent cell. The end of cytokinesis marks the end of the M-phase.
There are many cells where mitosis and cytokinesis occur separately, forming single cells with multiple nuclei. The most notable occurrence of this is among the fungi, slime molds, and coenocytic algae, but the phenomenon is found in various other organisms. Even in animals, cytokinesis and mitosis may occur independently, for instance during certain stages of fruit fly embryonic development.[36]
Significance[edit]
Mitosis is important for the maintenance of the chromosomal set; each cell formed receives chromosomes that are alike in composition and equal in number to the chromosomes of the parent cell.
Mitosis occurs in the following circumstances:
Development and growth
The number of cells within an organism increases by mitosis. This is the basis of the development of a multicellular body from a single cell, i.e., zygote and also the basis of the growth of a multicellular body.
Cell replacement
In some parts of body, e.g. skin and digestive tract, cells are constantly sloughed off and replaced by new ones. New cells are formed by mitosis and so are exact copies of the cells being replaced. In like manner, red blood cells have short lifespan (only about 4 months) and new RBCs are formed by mitosis.
Regeneration
Some organisms can regenerate body parts. The production of new cells in such instances is achieved by mitosis. For example, starfish regenerate lost arms through mitosis.
Asexual reproduction
Some organisms produce genetically similar offspring through asexual reproduction. For example, the hydra reproduces asexually by budding. The cells at the surface of hydra undergo mitosis and form a mass called a bud. Mitosis continues in the cells of the bud and this grows into a new individual. The same division happens during asexual reproduction or vegetative propagation in plants.
Cell shape changes during mitosis[edit]
Cell shape changes through mitosis for a typical animal cell cultured on a flat surface. The cell undergoes mitotic cell rounding during spindle assembly and then divides via cytokinesis. The actomyosin cortex is depicted in red, DNA/chromosomes purple, microtubules green, and membrane and retraction fibers in black. Rounding also occurs in live tissue, as described in the text.
Main article: Mitotic cell rounding
In animal tissue, most cells round up to a near-spherical shape during mitosis.[37][38][39] In epithelia and epidermis, an efficient rounding process is correlated with proper mitotic spindle alignment and subsequent correct positioning of daughter cells.[38][39][40][41] Moreover, researchers have found that if rounding is heavily suppressed it may result in spindle defects, primarily pole splitting and failure to efficiently capture chromosomes.[42] Therefore, mitotic cell rounding is thought to play a protective role in ensuring accurate mitosis.[41][43]
Rounding forces are driven by reorganization of F-actin and myosin (actomyosin) into a contractile homogeneous cell cortex that 1) rigidifies the cell periphery[43][44][45] and 2) facilitates generation of intracellular hydrostatic pressure (up to 10 fold higher than interphase).[46][47][48] The generation of intracellular pressure is particularly critical under confinement, such as would be important in a tissue scenario, where outward forces must be produced to round up against surrounding cells and/or the extracellular matrix. Generation of pressure is dependent on formin-mediated F-actin nucleation[48] and Rho kinase (ROCK)-mediated myosin II contraction,[44][46][48] both of which are governed upstream by signaling pathways RhoA and ECT2[44][45] through the activity of Cdk1.[48] Due to its importance in mitosis, the molecular components and dynamics of the mitotic actomyosin cortex is an area of active research.
Errors and variations[edit]
An abnormal (tripolar) mitosis (12 o'clock position) in a precancerous lesion of the stomach (H&E stain)
Errors can occur during mitosis, especially during early embryonic development in humans.[49] Mitotic errors can create aneuploid cells that have too few or too many of one or more chromosomes, a condition associated with cancer.[50][51] Early human embryos, cancer cells, infected or intoxicated cells can also suffer from pathological division into three or more daughter cells (tripolar or multipolar mitosis), resulting in severe errors in their chromosomal complements.[4]
In nondisjunction, sister chromatids fail to separate during anaphase.[52] One daughter cell receives both sister chromatids from the nondisjoining chromosome and the other cell receives none. As a result, the former cell gets three copies of the chromosome, a condition known as trisomy, and the latter will have only one copy, a condition known as monosomy. On occasion, when cells experience nondisjunction, they fail to complete cytokinesis and retain both nuclei in one cell, resulting in binucleated cells.[53]
Anaphase lag occurs when the movement of one chromatid is impeded during anaphase.[52] This may be caused by a failure of the mitotic spindle to properly attach to the chromosome. The lagging chromatid is excluded from both nuclei and is lost. Therefore, one of the daughter cells will be monosomic for that chromosome.
Endoreduplication (or endoreplication) occurs when chromosomes duplicate but the cell does not subsequently divide. This results in polyploid cells or, if the chromosomes duplicates repeatedly, polytene chromosomes.[52][54] Endoreduplication is found in many species and appears to be a normal part of development.[54] Endomitosis is a variant of endoreduplication in which cells replicate their chromosomes during S phase and enter, but prematurely terminate, mitosis. Instead of being divided into two new daughter nuclei, the replicated chromosomes are retained within the original nucleus.[36][55] The cells then re-enter G1 and S phase and replicate their chromosomes again.[55] This may occur multiple times, increasing the chromosome number with each round of replication and endomitosis. Platelet-producing megakaryocytes go through endomitosis during cell differentiation.[56][57]
Mitotic recombination[edit]
Mitotic cells irradiated with X-rays in the G1 phase of the cell cycle repair recombinogenic DNA damages primarily by recombination between homologous chromosomes.[58] Mitotic cells irradiated in the G2 phase repair such damages preferentially by sister-chromatid recombination.[58] Mutations in genes encoding enzymes employed in recombination cause cells to have increased sensitivity to being killed by a variety of DNA damaging agents.[59] These findings suggest that mitotic recombination is an adaptation for repairing DNA damages including those that are potentially lethal.
Timeline in pictures[edit]
Mitotic cells can be visualized microscopically by staining them with fluorescent antibodies and dyes.
Early prophase: Polar microtubules, shown as green strands, have established a matrix around the currently intact nucleus, with the condensing chromosomes in blue. The red nodules are the centromeres.
Early prometaphase: The nuclear membrane has just disassembled, allowing the microtubules to quickly interact with the kinetochores, which assemble on the centromeres of the condensing chromosomes.
Metaphase: The centrosomes have moved to the poles of the cell and have established the mitotic spindle. The chromosomes have congressed at the metaphase plate.
Anaphase: Kinetochore microtubules pull the two sets of chromosomes apart, and lengthening polar microtubules push the halves of the dividing cell further apart, while chromosomes are condensed maximally.
Telophase: Reversal of prophase and prometaphase events and thus completing the cell cycle.
See also[edit]
Aneuploidy
Binary fission
Chromosome abnormality
Cytoskeleton
Meiosis
Mitogen
Mitosis Promoting Factor
Mitotic bookmarking
Motor protein
References[edit]
Jump up ^ Carter, J. Stein (2014-01-14). "Mitosis". biology.clc.uc.edu. Archived from the original on 2012-10-27.
Jump up ^ "Cell Division: Stages of Mitosis | Learn Science at Scitable". www.nature.com. Archived from the original on 2015-11-14. Retrieved 2015-11-16.
Jump up ^ Maton A, Hopkins JJ, LaHart S, Quon Warner D, Wright M, Jill D (1997). Cells: Building Blocks of Life. New Jersey: Prentice Hall. pp. 70–4. ISBN 0-13-423476-6.
^ Jump up to: a b Kalatova B, Jesenska R, Hlinka D, Dudas M (January 2015). "Tripolar mitosis in human cells and embryos: occurrence, pathophysiology and medical implications". Acta Histochemica. 117 (1): 111–25. doi:10.1016/j.acthis.2014.11.009. PMID 25554607.
Jump up ^ Kops GJ, Weaver BA, Cleveland DW (October 2005). "On the road to cancer: aneuploidy and the mitotic checkpoint". Nature Reviews. Cancer. 5 (10): 773–85. doi:10.1038/nrc1714. PMID 16195750.
Jump up ^ Raikov, IB (1994). "The diversity of forms of mitosis in protozoa: A comparative review". European Journal of Protistology. 30 (3): 253–69. doi:10.1016/S0932-4739(11)80072-6.
Jump up ^ De Souza CP, Osmani SA (September 2007). "Mitosis, not just open or closed". Eukaryotic Cell. 6 (9): 1521–7. doi:10.1128/EC.00178-07. PMC 2043359  . PMID 17660363.
Jump up ^ Karl Mägdefrau (1994), "Mohl, Hugo von", Neue Deutsche Biographie (NDB) (in German), 17, Berlin: Duncker & Humblot, pp. 690–691 ; (full text online)
Jump up ^ Fokin, S. I. (2013). "Otto Bütschli (1848–1920) Where we will genuflect?" (PDF). Protistology. 8 (1): 22–35.
Jump up ^ Sharp LW (1921). Introduction To Cytology. New York: McGraw Hill Book Company Inc. p. 143.
Jump up ^ Janusz Komender (2008). "Kilka słów o doktorze Wacławie Mayzlu i jego odkryciu" [On Waclaw Mayzel and his observation of mitotic division] (PDF). Postępy Biologii Komórki (in Polish). Polskie Towarzystwo Anatomiczne, Polskie Towarzystwo Biologii Komórki. 35 (3): 405–407. Archived (PDF) from the original on 2012-10-27.
Jump up ^ Iłowiecki M (1981). Dzieje nauki polskiej. Warszawa: Wydawnictwo Interpress. p. 187. ISBN 83-223-1876-6.
Jump up ^ "mitosis". Online Etymology Dictionary.
Jump up ^ μίτος. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
^ Jump up to: a b Blow JJ, Tanaka TU (November 2005). "The chromosome cycle: coordinating replication and segregation. Second in the cycles review series". EMBO Reports. 6 (11): 1028–34. doi:10.1038/sj.embor.7400557. PMC 1371039  . PMID 16264427.
Jump up ^ Zhou J, Yao J, Joshi HC (September 2002). "Attachment and tension in the spindle assembly checkpoint". Journal of Cell Science. 115 (Pt 18): 3547–55. doi:10.1242/jcs.00029. PMID 12186941.
^ Jump up to: a b Lloyd C, Chan J (February 2006). "Not so divided: the common basis of plant and animal cell division". Nature Reviews Molecular Cell Biology. 7 (2): 147–52. doi:10.1038/nrm1831. PMID 16493420.
^ Jump up to: a b Raven PH, Evert RF, Eichhorn SE (2005). Biology of Plants (7th ed.). New York: W. H. Freeman and Co. ISBN 0716710072.
Jump up ^ Prasanth KV, Sacco-Bubulya PA, Prasanth SG, Spector DL (March 2003). "Sequential entry of components of the gene expression machinery into daughter nuclei". Molecular Biology of the Cell. 14 (3): 1043–57. doi:10.1091/mbc.E02-10-0669. PMC 151578  . PMID 12631722.
Jump up ^ Kadauke S, Blobel GA (April 2013). "Mitotic bookmarking by transcription factors". Epigenetics & Chromatin. 6 (1): 6. doi:10.1186/1756-8935-6-6. PMC 3621617  . PMID 23547918.
Jump up ^ Prescott DM, Bender MA (March 1962). "Synthesis of RNA and protein during mitosis in mammalian tissue culture cells". Experimental Cell Research. 26 (2): 260–8. doi:10.1016/0014-4827(62)90176-3. PMID 14488623.
Jump up ^ Olson MO (2011). The Nucleolus. Volume 15 of Protein Reviews. Berlin: Springer Science & Business Media. p. 15. ISBN 9781461405146.
Jump up ^ Basto R, Lau J, Vinogradova T, Gardiol A, Woods CG, Khodjakov A, Raff JW (June 2006). "Flies without centrioles". Cell. 125 (7): 1375–86. doi:10.1016/j.cell.2006.05.025. PMID 16814722.
Jump up ^ Heywood P (June 1978). "Ultrastructure of mitosis in the chloromonadophycean alga Vacuolaria virescens". Journal of Cell Science. 31: 37–51. PMID 670329.
Jump up ^ Ribeiro KC, Pereira-Neves A, Benchimol M (June 2002). "The mitotic spindle and associated membranes in the closed mitosis of trichomonads". Biology of the Cell. 94 (3): 157–72. doi:10.1016/S0248-4900(02)01191-7. PMID 12206655.
^ Jump up to: a b Chan GK, Liu ST, Yen TJ (November 2005). "Kinetochore structure and function". Trends in Cell Biology. 15 (11): 589–98. doi:10.1016/j.tcb.2005.09.010. PMID 16214339.
Jump up ^ Cheeseman IM, Desai A (January 2008). "Molecular architecture of the kinetochore-microtubule interface". Nature Reviews Molecular Cell Biology. 9 (1): 33–46. doi:10.1038/nrm2310. PMID 18097444.
^ Jump up to: a b Winey M, Mamay CL, O'Toole ET, Mastronarde DN, Giddings TH, McDonald KL, McIntosh JR (June 1995). "Three-dimensional ultrastructural analysis of the Saccharomyces cerevisiae mitotic spindle". The Journal of Cell Biology. 129 (6): 1601–15. doi:10.1083/jcb.129.6.1601. PMC 2291174  . PMID 7790357.
^ Jump up to: a b Maiato H, DeLuca J, Salmon ED, Earnshaw WC (November 2004). "The dynamic kinetochore-microtubule interface". Journal of Cell Science. 117 (Pt 23): 5461–77. doi:10.1242/jcs.01536. PMID 15509863.
Jump up ^ Chan GK, Yen TJ (2003). "The mitotic checkpoint: a signaling pathway that allows a single unattached kinetochore to inhibit mitotic exit". Progress in Cell Cycle Research. 5: 431–9. PMID 14593737.
^ Jump up to: a b FitzHarris G (March 2012). "Anaphase B precedes anaphase A in the mouse egg". Current Biology. 22 (5): 437–44. doi:10.1016/j.cub.2012.01.041. PMID 22342753.
Jump up ^ Miller KR (2000). "Anaphase". Biology (5 ed.). Pearson Prentice Hall. pp. 169–70. ISBN 978-0-13-436265-6.
Jump up ^ "Chromosome condensation through mitosis". Science Daily. Retrieved 12 June 2007.
Jump up ^ Glotzer M (March 2005). "The molecular requirements for cytokinesis". Science. 307 (5716): 1735–9. doi:10.1126/science.1096896. PMID 15774750.
Jump up ^ Albertson R, Riggs B, Sullivan W (February 2005). "Membrane traffic: a driving force in cytokinesis". Trends in Cell Biology. 15 (2): 92–101. doi:10.1016/j.tcb.2004.12.008. PMID 15695096.
^ Jump up to: a b Lilly MA, Duronio RJ (April 2005). "New insights into cell cycle control from the Drosophila endocycle". Oncogene. 24 (17): 2765–75. doi:10.1038/sj.onc.1208610. PMID 15838513.
Jump up ^ Sauer, F. C. (1935). "Mitosis in the neural tube". Journal of Comparative Neurology. 62 (2): 377–405. doi:10.1002/cne.900620207.
^ Jump up to: a b Meyer EJ, Ikmi A, Gibson MC (March 2011). "Interkinetic nuclear migration is a broadly conserved feature of cell division in pseudostratified epithelia". Current Biology. 21 (6): 485–91. doi:10.1016/j.cub.2011.02.002. PMID 21376598.
^ Jump up to: a b Luxenburg C, Pasolli HA, Williams SE, Fuchs E (March 2011). "Developmental roles for Srf, cortical cytoskeleton and cell shape in epidermal spindle orientation". Nature Cell Biology. 13 (3): 203–14. doi:10.1038/Ncb2163. PMC 3278337  . PMID 21336301.
Jump up ^ Nakajima Y, Meyer EJ, Kroesen A, McKinney SA, Gibson MC (August 2013). "Epithelial junctions maintain tissue architecture by directing planar spindle orientation". Nature. 500 (7462): 359–62. doi:10.1038/nature12335. PMID 23873041.
^ Jump up to: a b Cadart C, Zlotek-Zlotkiewicz E, Le Berre M, Piel M, Matthews HK (April 2014). "Exploring the function of cell shape and size during mitosis". Developmental Cell. 29 (2): 159–69. doi:10.1016/j.devcel.2014.04.009. PMID 24780736.
Jump up ^ Lancaster OM, Le Berre M, Dimitracopoulos A, Bonazzi D, Zlotek-Zlotkiewicz E, Picone R, Duke T, Piel M, Baum B (May 2013). "Mitotic rounding alters cell geometry to ensure efficient bipolar spindle formation". Developmental Cell. 25 (3): 270–83. doi:10.1016/j.devcel.2013.03.014. PMID 23623611.
^ Jump up to: a b Lancaster OM, Baum B (October 2014). "Shaping up to divide: coordinating actin and microtubule cytoskeletal remodelling during mitosis". Seminars in Cell & Developmental Biology. 34: 109–15. doi:10.1016/j.semcdb.2014.02.015. PMID 24607328.
^ Jump up to: a b c Maddox AS, Burridge K (January 2003). "RhoA is required for cortical retraction and rigidity during mitotic cell rounding". The Journal of Cell Biology. 160 (2): 255–65. doi:10.1083/jcb.200207130. PMC 2172639  . PMID 12538643.
^ Jump up to: a b Matthews HK, Delabre U, Rohn JL, Guck J, Kunda P, Baum B (August 2012). "Changes in Ect2 localization couple actomyosin-dependent cell shape changes to mitotic progression". Developmental Cell. 23 (2): 371–83. doi:10.1016/j.devcel.2012.06.003. PMC 3763371  . PMID 22898780.
^ Jump up to: a b Stewart MP, Helenius J, Toyoda Y, Ramanathan SP, Muller DJ, Hyman AA (January 2011). "Hydrostatic pressure and the actomyosin cortex drive mitotic cell rounding". Nature. 469 (7329): 226–30. doi:10.1038/nature09642. PMID 21196934.
Jump up ^ Fischer-Friedrich E, Hyman AA, Jülicher F, Müller DJ, Helenius J (August 2014). "Quantification of surface tension and internal pressure generated by single mitotic cells". Scientific Reports. 4 (6213): 6213. doi:10.1038/srep06213. PMC 4148660  . PMID 25169063.
^ Jump up to: a b c d Ramanathan SP, Helenius J, Stewart MP, Cattin CJ, Hyman AA, Muller DJ (February 2015). "Cdk1-dependent mitotic enrichment of cortical myosin II promotes cell rounding against confinement". Nature Cell Biology. 17 (2): 148–59. doi:10.1038/ncb3098. PMID 25621953.
Jump up ^ Mantikou E, Wong KM, Repping S, Mastenbroek S (December 2012). "Molecular origin of mitotic aneuploidies in preimplantation embryos". Biochimica et Biophysica Acta. 1822 (12): 1921–30. doi:10.1016/j.bbadis.2012.06.013. PMID 22771499.
Jump up ^ Draviam VM, Xie S, Sorger PK (April 2004). "Chromosome segregation and genomic stability". Current Opinion in Genetics & Development. 14 (2): 120–5. doi:10.1016/j.gde.2004.02.007. PMID 15196457.
Jump up ^ Santaguida, Stefano; Amon, Angelika (2015-08-01). "Short- and long-term effects of chromosome mis-segregation and aneuploidy". Nature Reviews. Molecular Cell Biology. 16 (8): 473–485. doi:10.1038/nrm4025. ISSN 1471-0080. PMID 26204159. Archived from the original on 2017-11-07.
^ Jump up to: a b c Iourov I, Vorsanova S, Yurov Y (2006). "Chromosomal Variations in Mammalian Neuronal Cells: Known Facts and Attractive Hypotheses". In Jeon KJ. International Review Of Cytology: A Survey of Cell Biology. 249. Waltham, MA: Academic Press. p. 146. ISBN 9780080463506.
Jump up ^ Shi Q, King RW (October 2005). "Chromosome nondisjunction yields tetraploid rather than aneuploid cells in human cell lines". Nature. 437 (7061): 1038–42. doi:10.1038/nature03958. PMID 16222248.
^ Jump up to: a b Edgar BA, Orr-Weaver TL (May 2001). "Endoreplication cell cycles: more for less". Cell. 105 (3): 297–306. doi:10.1016/S0092-8674(01)00334-8. PMID 11348589.
^ Jump up to: a b Lee HO, Davidson JM, Duronio RJ (November 2009). "Endoreplication: polyploidy with purpose". Genes & Development. 23 (21): 2461–77. doi:10.1101/gad.1829209. PMC 2779750  . PMID 19884253.
Jump up ^ Italiano JE, Shivdasani RA (June 2003). "Megakaryocytes and beyond: the birth of platelets". Journal of Thrombosis and Haemostasis. 1 (6): 1174–82. doi:10.1046/j.1538-7836.2003.00290.x. PMID 12871316.
Jump up ^ Vitrat N, Cohen-Solal K, Pique C, Le Couedic JP, Norol F, Larsen AK, Katz A, Vainchenker W, Debili N (May 1998). "Endomitosis of human megakaryocytes are due to abortive mitosis". Blood. 91 (10): 3711–23. PMID 9573008.
^ Jump up to: a b Kadyk LC, Hartwell LH (1992). "Sister chromatids are preferred over homologs as substrates for recombinational repair in Saccharomyces cerevisiae". Genetics. 132 (2): 387–402. PMC 1205144  . PMID 1427035.
Jump up ^ Harris Bernstein, Carol Bernstein and Richard E. Michod (2011). Meiosis as an Evolutionary Adaptation for DNA Repair. Chapter 19 in DNA Repair. Inna Kruman editor. InTech Open Publisher. DOI: 10.5772/25117 "Archived copy". Archived from the original on 2013-06-16. Retrieved 2013-07-29.
Further reading[edit]
Morgan DL (2007). The cell cycle: principles of control. London: Published by New Science Press in association with Oxford University Press. ISBN 0-9539181-2-2.
Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). "Mitosis". Molecular Biology of the Cell (4th ed.). Garland Science. Retrieved 2006-01-22.
Campbell N, Reece J (December 2001). "The Cell Cycle". Biology (6th ed.). San Francisco: Benjamin Cummings/Addison-Wesley. pp. 217–224. ISBN 0-8053-6624-5.
Cooper G (2000). "The Events of M Phase". The Cell: A Molecular Approach (2nd ed.). Sinaeur Associates, Inc. Retrieved 2006-01-22.
Freeman S (2002). "Cell Division". Biological Science. Upper Saddle River, NJ: Prentice Hall. pp. 155–174. ISBN 0-13-081923-9.
Lodish H, Berk A, Zipursky L, Matsudaira P, Baltimore D, Darnell J (2000). "Overview of the Cell Cycle and Its Control". Molecular Cell Biology (4th ed.). W. H. Freeman. Retrieved 2006-01-22.
External links[edit]
Wikimedia Commons has media related to Mitosis.
A Flash animation comparing Mitosis and Meiosis
Khan Academy, lecture
Studying Mitosis in Cultured Mammalian Cells
General K-12 classroom resources for Mitosis
The Cell-Cycle Ontology
WormWeb.org: Interactive Visualization of the C. elegans Cell Lineage – Visualize the entire cell lineage tree and all of the cell divisions of the nematode C. elegans
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text:
PDCA
From Wikipedia, the free encyclopedia
Jump to: navigation, search
For other uses, see PDCA (disambiguation).
PDCA (plan–do–check–act or plan–do–check–adjust) is an iterative four-step management method used in business for the control and continual improvement of processes and products.[1] It is also known as the Deming circle/cycle/wheel, the Shewhart cycle, the control circle/cycle, or plan–do–study–act (PDSA). Another version of this PDCA cycle is OPDCA.[citation needed] The added "O" stands for observation or as some versions say: "Observe the current condition." This emphasis on observation and current condition has currency with the literature on lean manufacturing and the Toyota Production System.[2]:129–158
Contents
[hide]
1 Meaning
1.1 Plan
1.2 Do
1.3 Check
1.4 Act
2 About
3 See also
4 References
5 Further reading
Meaning[edit]
This section contains instructions, advice, or how-to content. The purpose of Wikipedia is to present facts, not to train. Please help improve this article either by rewriting the how-to content or by moving it to Wikiversity, Wikibooks or Wikivoyage. (October 2017)
The PDCA cycle[3]
Continuous quality improvement with PDCA
Plan[edit]
During the plan phase, establish the objectives and processes necessary to deliver results in accordance with the expected output (the target or goals). By establishing output expectations, the completeness and accuracy of the specification is also a part of the targeted improvement. When possible start on a small scale to test possible effects.
Do[edit]
During the do phase, implement the plan, execute the process, make the product. Collect data for charting and analysis in the following check and act steps.
Check[edit]
In the check phase, study the actual results (measured and collected in do phase above) and compare against the expected results (targets or goals from the plan phase) to ascertain any differences. Look for deviation in implementation from the plan and also look for the appropriateness and completeness of the plan to enable the execution, i.e., the doing. Charting data can make this much easier to see trends over several PDCA cycles and in order to convert the collected data into information. Information is needed for the next step: act.
Act[edit]
If the check phase shows that the plan phase which was implemented in do phase is an improvement to the prior standard (baseline), then that becomes the new standard (baseline) for how the organization should act going forward (new standards are thus said to be enACTed). Instead, if the check phase shows that the plan phase which was implemented in do phase is not an improvement, then the existing standard (baseline) will remain in place. In either case, if the check phase showed something different than expected (whether better or worse), then there is some more learning to be done... and that will suggest potential future PDCA cycles. Note that some who teach PDCA assert that the act phase involves making adjustments or corrective actions, but generally it would be counter to PDCA thinking to propose and decide upon alternative changes without using a proper plan phase, or to make them the new standard (baseline) without going through do and check steps.
About[edit]
PDCA was made popular by W. Edwards Deming, who is considered by many to be the father of modern quality control; however, he always referred to it as the "Shewhart cycle". Later in Deming's career, he modified PDCA to "Plan, Do, Study, Act" (PDSA) because he felt that "check" emphasized inspection over analysis.[4] The PDSA cycle was used to create the model of know-how transfer process,[5] and other models.[6]
The concept of PDCA is based on the scientific method, as developed from the work of Francis Bacon (Novum Organum, 1620). The scientific method can be written as "hypothesis–experiment–evaluation" or as "plan–do–check". Walter A. Shewhart described manufacture under "control"—under statistical control—as a three-step process of specification, production, and inspection.[7]:45 He also specifically related this to the scientific method of hypothesis, experiment, and evaluation. Shewhart says that the statistician "must help to change the demand [for goods] by showing [...] how to close up the tolerance range and to improve the quality of goods."[7]:48 Clearly, Shewhart intended the analyst to take action based on the conclusions of the evaluation. According to Deming, during his lectures in Japan in the early 1950s, the Japanese participants shortened the steps to the now traditional plan, do, check, act.[8]:88 Deming preferred plan, do, study, act because "study" has connotations in English closer to Shewhart's intent than "check".[9]
Multiple iterations of the PDCA cycle are repeated until the problem is solved.
A fundamental principle of the scientific method and PDCA is iteration—once a hypothesis is confirmed (or negated), executing the cycle again will extend the knowledge further. Repeating the PDCA cycle can bring its users closer to the goal, usually a perfect operation and output.[9]
Another fundamental function of PDCA is the "hygienic" separation of each phase, for if not properly separated measurements of effects due to various simultaneous actions (causes) risk becoming confounded.[10]
PDCA (and other forms of scientific problem solving) is also known as a system for developing critical thinking. At Toyota this is also known as "Building people before building cars".[11] Toyota and other lean manufacturing companies propose that an engaged, problem-solving workforce using PDCA in a culture of critical thinking is better able to innovate and stay ahead of the competition through rigorous problem solving and the subsequent innovations.[11]
Deming continually emphasized iterating towards an improved system, hence PDCA should be repeatedly implemented in spirals of increasing knowledge of the system that converge on the ultimate goal, each cycle closer than the previous.[citation needed] One can envision an open coil spring, with each loop being one cycle of the scientific method, and each complete cycle indicating an increase in our knowledge of the system under study. This approach is based on the belief that our knowledge and skills are limited, but improving. Especially at the start of a project, key information may not be known; the PDCA—scientific method—provides feedback to justify guesses (hypotheses) and increase knowledge. Rather than enter "analysis paralysis" to get it perfect the first time, it is better to be approximately right than exactly wrong. With improved knowledge, one may choose to refine or alter the goal (ideal state). The aim of the PDCA cycle is to bring its users closer to whatever goal they choose.[2]:160
When PDCA is used for complex projects or products with a certain controversy, checking with external stakeholders should happen before the Do stage, since changes to projects and products that are already in detailed design can be costly; this is also seen as Plan-Check-Do-Act.[citation needed]
Rate of change, that is, rate of improvement, is a key competitive factor in today's world.[citation needed] PDCA allows for major "jumps" in performance ("breakthroughs" often desired in a Western approach), as well as kaizen (frequent small improvements).[citation needed] In the United States a PDCA approach is usually associated with a sizable project involving numerous people's time,[citation needed] and thus managers want to see large "breakthrough" improvements to justify the effort expended. However, the scientific method and PDCA apply to all sorts of projects and improvement activities.[2]:76
See also[edit]
COBIT
Decision cycle
DMAIC
Lean manufacturing
Learning cycle
Lesson study
OODA loop
Performance management
Quality storyboard
Robert S. Kaplan (closed loop management system)
Six sigma
Theory of constraints
Total security management
Software development process
References[edit]
Jump up ^ Tague, Nancy R. (2005) [1995]. "Plan–Do–Study–Act cycle". The quality toolbox (2nd ed.). Milwaukee: ASQ Quality Press. pp. 390–392. ISBN 0873896394. OCLC 57251077. Retrieved 2017-10-21.
^ Jump up to: a b c Rother, Mike (2010). Toyota kata: managing people for improvement, adaptiveness, and superior results. New York: McGraw-Hill. ISBN 0071635238. OCLC 318409119.
Jump up ^ "Taking the first step with PDCA". 2 February 2009. Retrieved 17 March 2011.
Jump up ^ Aguayo, Rafael (1990). Dr. Deming: the American who taught the Japanese about quality. A Lyle Stuart book. Secaucus, NJ: Carol Pub. Group. p. 76. ISBN 0818405198. OCLC 22347078. Also published by Simon & Schuster, 1991.
Jump up ^ Dubickis, Mikus; Gaile-Sarkane, Elīna (December 2017). "Transfer of know-how based on learning outcomes for development of open innovation". Journal of Open Innovation: Technology, Market, and Complexity. 3 (1): 4. doi:10.1186/s40852-017-0053-4.
Jump up ^ Dubberly, Hugh (2008) [2004]. "How do you design?: a compendium of models". dubberly.com. Retrieved 2017-10-21.
^ Jump up to: a b Shewhart, Walter Andrew (1986) [1939]. Statistical method from the viewpoint of quality control. New York: Dover. ISBN 0486652327. OCLC 13822053. Reprint. Originally published: Washington, DC: Graduate School of the Department of Agriculture, 1939.
Jump up ^ Deming, W. Edwards (1986). Out of the crisis. Cambridge, MA: Massachusetts Institute of Technology, Center for Advanced Engineering Study. ISBN 0911379010. OCLC 13126265.
^ Jump up to: a b Moen, Ronald; Norman, Clifford. "Evolution of the PDCA cycle" (PDF). westga.edu. Paper delivered to the Asian Network for Quality Conference in Tokyo on September 17, 2009. Retrieved 1 October 2011.
Jump up ^ Berengueres, José (2007). The Toyota production system re-contextualized. Tokyo: José Berengueres. p. 74. ISBN 1847534775. OCLC 906982187.
^ Jump up to: a b Liker, Jeffrey K. (2004). The Toyota way: 14 management principles from the world's greatest manufacturer. New York: McGraw-Hill. ISBN 0071392319. OCLC 54005437.
Further reading[edit]
Wikimedia Commons has media related to PDCA.
Kolesar, Peter J. (2005) [1994]. "What Deming told the Japanese in 1950". In Wood, John C.; Wood, Michael C. W. Edwards Deming: critical evaluations in business and management. 2. New York: Routledge. pp. 87–107. ISBN 9780415323888. OCLC 55738077. Reprint. Originally published: Quality Management Journal 2(1) (1994): 9–24.
Langley, Gerald J.; Moen, Ronald D.; Nolan, Kevin M.; Nolan, Thomas W.; Norman, Clifford L.; Provost, Lloyd P. (2009) [1996]. The improvement guide: a practical approach to enhancing organizational performance (2nd ed.). San Francisco: Jossey-Bass. ISBN 9780470192412. OCLC 236325893.
Shewhart, Walter Andrew (1980) [1931]. Economic control of quality of manufactured product. Milwaukee: American Society for Quality. ISBN 0873890760. OCLC 7543940. 50th anniversary commemorative reissue. Originally published: New York: Van Nostrand, 1931.
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text:
Electrical brain stimulation
From Wikipedia, the free encyclopedia
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Chronic subcortical electrode implant in a laboratory rat used to deliver electrical stimulation to the brain.
Electrical brain stimulation (EBS), also referred to as focal brain stimulation (FBS), is a form of electrotherapy and technique used in research and clinical neurobiology to stimulate a neuron or neural network in the brain through the direct or indirect excitation of its cell membrane by using an electric current. It is used for research or for therapeutical purposes.
Contents
[hide]
1 History
2 Process
3 Effects
4 Therapeutic applications
5 References
History[edit]
Electrical brain stimulation was first used in the first half of the 19th century by pioneering researchers such as Luigi Rolando[citation needed](1773–1831) and Pierre Flourens[citation needed](1794–1867), to study the brain localization of function, following the discovery by Italian physician Luigi Galvani (1737–1798) that nerves and muscles were electrically excitable. The stimulation of the surface of the cerebral cortex by using brain stimulation was used to investigate the motor cortex in animals by researchers such as Eduard Hitzig (1838–1907), Gustav Fritsch (1838–1927), David Ferrier (1842–1928) and Friedrich Goltz (1834–1902). The human cortex was also stimulated electrically by neurosurgeons and neurologists such as Robert Bartholow (1831–1904) and Fedor Krause (1857–1937).
In the following century, the technique was improved by the invention of the stereotactic method by British neurosurgeon pioneer Victor Horsley (1857–1916), and by the development of chronic electrode implants by Swiss neurophysiologist Walter Rudolf Hess (1881–1973), José Delgado (1915-2011) and others, by using electrodes manufactured by straight insulated wire that could be inserted deep into the brain of freely-behaving animals, such as cats and monkeys. This approach was used by Dr.James Olds (1922–1976) and colleagues to discover brain stimulation reward and the pleasure center. American-Canadian neurosurgeon Wilder Penfield (1891–1976) and colleagues at the Montreal Neurological Institute used extensively electrical stimulation of the brain cortex in awake neurosurgical patients to investigate the motor and sensory homunculus (the representation of the body in the brain cortex according to the distribution of motor and sensory territories).
EBS remains inextricably entwined with the work of Dr. Robert Galbraith Heath, Dr. Jose Delgado and Dr. Wilder Penfield. It's of interest that during cerebral localization studies, neurosurgeon Penfield could not elicit emotional reactions in humans, either by observing spontaneous epilepsy or by electrically stimulating the surface of the cerebral cortex. Neurophysiologist Dr. Jose Delgado noted a few exceptions to this rule. In contrast, EBS, via deeply implanted electrodes in localized areas of the brain (deep brain stimulation; DBS), elicited both pleasurable and aversive responses in laboratory animals and man as previously described.[1][2][3]
EBS could elicit the ritualistic, motor responses of sham rage in cats by stimulation of the anterior hypothalamus, as well as more complex emotional and behavioral components of "true rage" in both experimental animals by stimulaton of the lateral hypothalamus, and in human subjects by stimulating various deep areas of the brain. EBS in human patients with epilepsy could trigger seizures in the surface of the brain and pathologic aggression and rage with stimulation of the amygdala.[3][4]
Process[edit]
Two-photon excitation microscopy has shown that microstimulation activates neurons sparsely around the electrode even at low currents (as low as 10 μA) up to distances as far as four millimeters away. This happens without particularly selecting other neurons much nearer the electrode's tip. This is due to activation of neurons being determined by whether they do or do not have axons or dendrites that pass within a radius of 15 μm near the tip of the electrode. As the current is increased the volume around the tip that activates neuron axons and dendrites increases and with this the number of neurons activated. Activation is most likely to be due to direct depolarization rather than synaptic activation.[5]
Effects[edit]
A comprehensive review of EBS research compiled a list of many different acute impacts of stimulation depending on the brain region targeted. Following are some examples of the effects documented:[6]
Sensory: Feelings of body tingling, swaying, movement, suffocation, burning, shock, warmth, paresthesia, feeling of falling, oscillopsia, dysesthesia, levitation, sounds, phosphenes, hallucinations, micropsia, diplopia, etc.
Motor: Eye movements, locomotion, speech arrest, automatisms, laughter, palilalia, chewing, urge to move, crying without feeling sad, etc.
Autonomic: Blushing, mydriasis, change in blood pressure and breathing, apnea, nausea, tachycardia, sweating, etc.
Emotional: Anxiety, mirth, feeling of unreality, fear, happiness, anger, sadness, transient acute depression, hypomania, etc.
Cognitive: Acalculia, paraphasia, anomic aphasia, recalling memories, "going into a trance", "out of this world", conduction aphasia, hemispatial neglect, alexia, déjà vu, reliving past experiences, agraphia, apraxia, etc.
EBS in face-sensitive regions of the fusiform gyrus caused a patient to report that the faces of the people in the room with him had "metamorphosed" and became distorted: "Your nose got saggy, went to the left. [...] Only your face changed, everything else was the same."[7]
Therapeutic applications[edit]
Stereotactic apparatus used to insert an electrode into the basal ganglia of the brain, for Parkinson's disease surgery.
Examples of therapeutic EBS are:
Cranial electrotherapy stimulation (CES)
Deep brain stimulation (DBS)
Transcranial direct current stimulation (tDCS)
Electroconvulsive therapy (ECT)
Low Field Magnetic Stimulation (LFMS)
Functional electrical stimulation (FES)
Magnetic seizure therapy (MST)
Vagus nerve stimulation (VNS)
Deep transcranial magnetic stimulation (Deep TMS)
Strong electric currents may cause a localized lesion in the nervous tissue, instead of a functional reversible stimulation. This property has been used for neurosurgical procedures in a variety of treatments, such as for Parkinson's disease, focal epilepsy and psychosurgery. Sometimes the same electrode is used to probe the brain for finding defective functions, before passing the lesioning current (electrocoagulation).
References[edit]
Jump up ^ Penfield, Wilder (1974). Speech and Brain Mechanisms. New York: Atheneum.
Jump up ^ Delgado, Jose (1986). Physical Control of the Mind: Toward a Psychocivilized Society. New York: Harper and Row.
^ Jump up to: a b Faria, Miguel A. "Violence, mental illness, and the brain - A brief history of psychosurgery: Part 2 - From the limbic system and cingulotomy to deep brain stimulation". Surg Neurol Int 01-Jun-2013;4:75. Retrieved April 7, 2014.
Jump up ^ Mark, Vernon (1970). Violence and the Brain. New York: Harper and Row.
Jump up ^ Histed, MH; Bonin, V; Reid, RC. (2009). "Direct activation of sparse, distributed populations of cortical neurons by electrical microstimulation". Neuron. 63 (4): 508–22. doi:10.1016/j.neuron.2009.07.016. PMC 2874753  . PMID 19709632.
Jump up ^ Aslihan Selimbeyoglu; Josef Parvizi (30 May 2010). "Electrical stimulation of the human brain: perceptual and behavioral phenomena reported in the old and new literature". Frontiers in Human Neuroscience. 4 (46). doi:10.3389/fnhum.2010.00046. ISSN 1662-5161. Retrieved 26 July 2014.
Jump up ^ Josef Parvizi; Corentin Jacques; Brett L. Foster; Nathan Withoft; Vinitha Rangarajan; Kevin S. Weiner; Kalanit Grill-Spector (24 Oct 2012). "Electrical Stimulation of Human Fusiform Face-Selective Regions Distorts Face Perception". Journal of Neuroscience. 32 (43): 14915–14920. doi:10.1523/JNEUROSCI.2609-12.2012. ISSN 1529-2401. PMC 3517886  . PMID 23100414. Retrieved 26 July 2014.
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