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ivision in other vertebrates 42 dna damage repair in the cell cycle edit dna damage is detected and repaired at various points in the cell cycle the g1 s checkpoint g2 m checkpoint and the checkpoint between metaphase and anaphase all monitor for dna damage and halt cell division by inhibiting different cyclin cdk complexes the p53 tumor suppressor protein plays a crucial role at the g1 s checkpoint and the g2 m checkpoint activated p53 proteins result in the expression of many proteins that are important in cell cycle arrest repair and apoptosis at the g1 s checkpoint p53 acts to ensure that the cell is ready for dna replication while at the g2 m checkpoint p53 acts to ensure that the cells have properly duplicated their content before entering mitosis 43 specifically when dna damage is present atm and atr kinases are activated activating various checkpoint kinases 44 these checkpoint kinases phosphorylate p53 which stimulates the production of different enzymes associated with dna repair 45 activated p53 also upregulates p21 which inhibits various cyclin cdk complexes these cyclin cdk complexes phosphorylate the retinoblastoma rb protein a tumor suppressor bound with the e2f family of transcription factors the binding of this rb protein ensures that cells do not enter the s phase prematurely however if it is not able to be phosphorylated by these cyclin cdk complexes the protein will remain and the cell will be halted in the g1 phase of the cell cycle 46 if dna is damaged the cell can also alter the akt pathway in which bad is phosphorylated and dissociated from bcl2 thus inhibiting apoptosis if this pathway is altered by a loss of function mutation in akt or bcl2 then the cell with damaged dna will be forced to undergo apoptosis 47 if the dna damage cannot be repaired activated p53 can induce cell death by apoptosis it can do so by activating the p53 upregulated modulator of apoptosis puma puma is a pro apoptotic protein that rapidly induces apoptosis by inhibiting the anti apoptotic bcl 2 family members 48 degradation edit multicellular organisms replace worn out cells through cell division in some animals however cell division eventually halts in humans this occurs on average after 52 divisions known as the hayflick limit the cell is then referred to as senescent with each division the cells telomeres protective sequences of dna on the end of a chromosome that prevent degradation of the chromosomal dna shorten this shortening has been correlated to negative effects such as age related diseases and shortened lifespans in humans 49 50 cancer cells on the other hand are not thought to degrade in this way if at all an enzyme complex called telomerase present in large quantities in cancerous cells rebuilds the telomeres through synthesis of telomeric dna repeats allowing division to continue indefinitely 51 history edit kurt michel with his phase contrast microscope at the beginning of the 19th century various hypotheses circulated about cell proliferation which became observable in plant and animal organisms as a result of advances in microscopy while the proliferation of cells on the inner side of old cells 52 53 the attachment of vesicles to existing cells 54 or crystallization in the intercellular space 55 were postulated as mechanisms of cell proliferation proponents of cell division itself had to fight for its acceptance for decades the belgian botanist barthélemy charles joseph dumortier must be regarded as the first discoverer of cell division in 1832 he described cell division in simple aquatic plants french conferve as follows translated from french to english the development of the conferve is as simple as its structure it takes place by the attachment of new cells to the old and this attachment always takes place from the end the terminal cell elongates more than the deeper cells then the production of a lateral bisector takes place in the inner fluid which tends to divide the cell into two parts of which the deeper one remains stationary while the terminal part elongates again forms a new inner partition and so on is the production of the middle partition originally double or single it is impossible to determine this but it is always true that it later appears double when united and that when two cells naturally separate each of them is closed at both ends 56 in 1835 the german botanist and physician hugo von mohl described plant cell division in much greater detail in his dissertation on freshwater and seawater algae for his phd thesis in medicine and surgery 57 among the most obscure phenomena of plant life is the manner in which the newly developing cells are formed and so there is no lack of manifold descriptions and explanations of this process and that gaps that were found in the observations were filled in by overly bold conclusions and assumptions translated from german to english in 1838 the german physician and botanist franz julius ferdinand meyen confirmed the mechanism of cell division at the root tips of plants 58 the german polish physician robert remak suspected that he had already discovered animal cell division in the blood of chicken embryos in 1841 59 but it was not until 1852 that he was able to confirm animal cell division for the first time in bird embryos frog larvae and mammals 60 in 1943 cell division was filmed for the first time 61 by kurt michel using a phase contrast microscope 62 see also edit cell fusion gametic fusion cell growth cyclin dependent kinase labile cells cells that constantly divide mitotic catastrophe references edit martin ea hine r 2020 a dictionary of biology 6th ed oxford oxford university press isbn 978 0 19 920462 5 oclc 176818780 griffiths aj 2012 introduction to genetic analysis 10th ed new york w h freeman and co isbn 978 1 4292 2943 2 oclc 698085201 10 2 the cell cycle biology 2e openstax openstax org 28 march 2018 retrieved 2020 11 24 gilbert scott f 2000 meiosis developmental biology 6th edition sinauer 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