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Radioresistance

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in the range that several extremophile species have been observed to be capable of surviving. This is primarily because the Earth's magnetic field shields all its inhabitants from solar cosmic radiation and galactic cosmic rays, which are the two primary sources of ionizing radiation across our solar system, and even including all documented terrestrial sources of ionizing radiation such as radon gas and primordial radionuclides at geographical locations considered to be natural high-level radiation sites, the yearly dose of natural background radiation remains tens of thousands of times smaller than the levels of ionizing radiation that many highly radioresistant organisms can withstand.
277:, and lived, whereas his wife, who got a dose of 5.7 Gy, died. The most likely explanation is that his dose was fractionated into many smaller doses which were absorbed over a length of time while his wife stayed in the house more and was subjected to continuous irradiation without a break so giving the self repair mechanisms in her body less time to repair some of the damage done by the radiation. This resulted in her death. He also eventually died in 1994. In the same way some of the persons who worked in the basement of the wrecked 440:(sterility arises 3 days post-exposure). Other important factors that influence the level of radioresistance include: The length of time during which a dose of radiation is delivered—with doses delivered during longer periods, or at time intervals, being associated with greatly reduced negative effects; The feeding state of individuals—with pre-fed and post-fed individuals being more resistant to radiations compared to starved individuals; The type of radiation used (e.g., tardigrades 200: 36: 3057: 404:
and to different experimental conditions. We should for example consider that because radiation impedes cell division, immature organisms are less resistant to radiations than adults, and adults are sterilized at doses much lower than that necessary to kill them. For example, for the insect parasitoid
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From the perspective of evolutionary history and causation, radioresistance does not appear to be an adaptive trait because there is no documented naturally occurring selection pressure that could have bestowed a fitness advantage to the ability for organisms to withstand doses of ionizing radiation
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One possible explanation for the existence of radioresistance is that it is an example of co-opted adaptation or exaptation, where radioresistance could be an indirect consequence of the evolution of a different, linked adaptation that has been positively selected for by evolution. For example, the
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The comparison in the table below is only meant to give approximate indications of radioresistance for different species and should be taken with great caution. There are generally big differences in radioresistance for one species among experiments, due to the way radiation affects living tissues
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experiments that if a group of cells are irradiated then as the dose increases the number of cells which survive decrease. It has also been found that if a population of cells are given a dose before being set aside (without being irradiated) for a length of time before being irradiated again then
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at 25 days post-exposure is 5,700 R, but to reach 50% mortality at 3 days post-exposure, 45,610 R are needed. 25 days can represent a long survival period for short lived species, such as insects, but would represent a very short survival time for long lived species, such as mammals, so comparing
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sp. producing unusually radiation (and peroxide) resistant spores, have been isolated from spacecraft assembly facilities, and are thought of as candidates that could ride piggyback on spacecraft through interplanetary transfer. Genome analysis of some of these radiation resistant spore producers
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is more resistant to X-ray radiation when in the anhydrobiotic state). The way lethality is measured is also source of variation for the estimated radioresistance of a species. Irradiated specimens are not instantly killed, unless exposed to a very high dose (acute dose). Therefore, irradiated
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to survive extreme doses of ionizing radiation. Exposure to gamma radiation leads to cellular DNA damage including alterations in nitrogenous base-pairing, sugar-phosphate backbone damage, and double-stranded DNA lesions. The extraordinarily efficient cellular repair mechanisms that
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survival of different species after the same amount of time post-exposure also poses some challenges of interpretation. These examples illustrate the many issues associated with comparison of radioresistance among species and the need for caution when doing so.
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cause cellular damage that is virtually identical to damage typically caused by ionizing radiation, and that the cellular repair mechanisms that have evolved to repair this heat or desiccation damage are generalizable to radiation damage as well, allowing
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that is manifested by a complete cessation of activity, including cessation of feeding, and these individuals eventually let themselves starve to death. And an even lower dose of 4,858 R (for 2,650 R/minute) is sufficient to sterilize adult female
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This is a graph showing the effect of fractionation on the ability of gamma rays to cause cell death. The blue line is for cells which were not given a chance to recover while the red line is for cells which were allowed to stand for a time and
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have evolved to repair heat-damage are likely also capable of reversing the effects of DNA damage wrought by ionizing radiation, such as by piecing back together any components of their genome that have been fragmented by the radiation.
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is the same to this of a non-irradiated control group (food was not provided to either groups) (for 6,000 R/minute). However, a lower dose of 102,000 R (for 6,000 R/minute) is sufficient to induce a state of lethargy in
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Many organisms have been found to possess a self-repair mechanism that can be activated by exposure to radiation in some cases. Two examples of this self-repair process in humans are described below.
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Ionizing-radiation-resistant organisms (IRRO) were defined as organisms for which the dose of acute ionizing radiation (IR) required to achieve 90% reduction (D10) is greater than 1,000 gray (Gy)
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Hartman, P; Goldstein, P; Algarra, M; Hubbard, D; Mabery, J (1996). "The nematode Caenorhabditis elegans is up to 39 times more sensitive to gamma radiation generated from 137Cs than from 60Co".
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group of cells has been drawn with and without a rest time for the cells to recover. Other than the recovery time partway through the irradiation, the cells would have been treated identically.
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irradiated with heavy ions have a higher survival than when irradiated with gamma rays, for a same irradiation dose); The physiological state of individuals (e.g., the tardigrade species
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individuals, an upper lethal limit has not been established, rather "nearly all animals were alive with no indication of excess lethality up to 800 Gy, the highest dose... measured."
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Agapov, A. A.; Kulbachinskiy, A. V. (October 2015). "Mechanisms of Stress Resistance and Gene Regulation in the Radioresistant Bacterium Deinococcus radiodurans".
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Murray RGE. 1992. The family Deino- coccaceae. In The Prokaryotes, ed. A Ballows, HG Truper, M Dworkin, W Harder, KH Schleifer 4:3732–44. New York: Springer-Verlag
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Radioresistance is surprisingly high in many organisms, in contrast to previously held views. For example, the study of environment, animals and plants around the
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There is strong evidence that radioresistance can be genetically determined and inherited, at least in some organisms. Heinrich Nöthel, a geneticist from the
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and a human can be killed by the loss of a single tissue in a vital organ. For many short term radiation deaths (3 days to 30 days) the loss of cells forming
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La Duc MT, Nicholson W, Kern R, Venkateswaran K (2003). "Microbial characterization of the Mars Odyssey spacecraft and its encapsulation facility".
1455:"Radioresistance of Deinococcus radiodurans: functions necessary to survive ionizing radiation are also necessary to survive prolonged desiccation" 2838: 422:(for X-ray intensity of 110 R/minute), and haploid (= male) embryos are more resistant than diploid (= female) embryos. The mortality of adults 2962: 2929: 457:
specimens die over a certain period of time and lower irradiation doses correspond to longer survival. This means that the radiation dose LD
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Tirumalai MR, Fox GE (September 2013). "An ICEBs1-like element may be associated with the extreme radiation and desiccation resistance of
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Tirumalai MR, Rastogi R, Zamani N, O'Bryant Williams E, Allen S, Diouf F, Kwende S, Weinstock GM, Venkateswaran KJ, Fox GE (June 2013).
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Horikawa DD; Sakashita T; Katagiri C; Watanabe M; et al. (2006). "Radiation tolerance in the tardigrade Milnesium tardigradum".
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Moustacchi, E (1965). "Induction by physical and chemical agents of mutations for radioresistance in Saccharomyces cerevisiae".
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Radioresistance may be induced by exposure to small doses of ionizing radiation. Several studies have documented this effect in
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desiccation-adaptation hypothesis proposes that the extreme temperatures present in the habitats of hyperthermophiles like
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have built up doses of 10 Gy, these workers received these doses in small fractions so the acute effects were avoided.
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Kempf MJ, Chen F, Kern R, Venkateswaran K (June 2005). "Recurrent isolation of hydrogen peroxide- resistant spores of
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fluctuates with the time at which it is measured. For example, the β radiation dose that causes 50% mortality in the
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Grosch, DS; Sullivan, RL (1954). "The quantitative aspects of permanent and temporary sterility induced in female
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Fornalski KW (2019). "Radiation adaptive response and cancer: from the statistical physics point of view".
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Radiochemistry and Nuclear Chemistry, G. Choppin, J-O. Liljenzin and J. Rydberg, edition three, page 481,
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Friedberg, Errol C.; Friedberg, EC; Walker, GC; Walker, Graham C.; Siede, Wolfram; Wolfram, Siede (1995).
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Fornalski, K.W. (2016). "Radiation and evolution: from Lotka-Volterra equation to balance equation".
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Tirumalai MR, Stepanov VG, Wünsche A, Montazari S, Gonzalez RO, Venkateswaran K, Fox GE (June 2018).
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have thrown some light on the genetic traits that could be responsible for the resistances observed.
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which has high natural radiation levels from uranium deposits, has also shown many radioresistant
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Newcombe DA, Schuerger AC, Benardini JN, Dickinson D, Tanner R, Venkateswaran K (December 2005).
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Link L, Sawyer J, Venkateswaran K, Nicholson W (February 2004). "Extreme spore UV resistance of
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area has revealed an unexpected survival of many species, despite the high radiation levels. A
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Wharton, DRA; Wharton, ML (1959). "The effect of radiation on the longevity of the cockroach,
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Cordeiro, AR; Marques, EK; Veiga-Neto, AJ (1973). "Radioresistance of a natural population of
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Clifton Ling, C.; Endlich, B. (1989). "Radioresistance Induced by Oncogenic Transformation".
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and other processes. Also biophysical models presented general basics for this phenomenon.
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Joiner, M.C. (1994). "Induced Radioresistance: An Overview and Historical Perspective".
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SAFR-032: A Whole Genome Comparison of two Spacecraft Assembly Facility Isolates"
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Shahbazi-Gahrouei, Daryoush; Gholami, Mehrdad; Setayandeh, Samaneh (2013-01-01).
1236: 914: 901: 427: 419: 415: 256: 2462: 2175: 2988: 2900: 1354: 1205: 294: 249: 2769: 2411: 2141: 2085: 1748: 1658: 1541:"DNA repair in the extremely radioresistant bacterium Deinococcus radiodurans" 3071: 1666: 1613: 1566: 1478: 1421: 1412: 1362: 1297: 1098: 170: 2718: 2656: 2244:
Grosch, DS (1954). "Induced lethargy and the radiation control of insects".
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Sullivan, R; Grosch, D (1953). "The radiation tolerance of an adult wasp".
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Sghaier, Haïtham; Ghedira, Kaïs; Benkahla, Alia; Barkallah, Insaf (2008).
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are more resistant to gamma-ray radiation when in the hydrated state, and
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isolates obtained from an ultraclean Spacecraft Assembly Facility".
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Weidhaas, J.B.; Eisenmann, D.M.; Holub, J.M.; Nallur, S.V. (2006).
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Level of ionizing radiation that organisms are able to withstand
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Gioia J, Yerrapragada S, Qin X, et al. (September 2007).
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for haploid embryo during cleavage (1–3 hours of age) is 200
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carried out the most extensive study about radioresistance
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Ghosh S, Osman S, Vaishampayan P, Venkateswaran K (2010).
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Krisko, A.; Magali, L.; Radman, M.; Meselson, M. (2012).
1017: 548: 504: 2558:"Radiation Notes: Radiation Damage and Dose Measurement" 1590:"Oxidative stress resistance in Deinococcus radiodurans" 375: 2391: 2389: 2387: 2331: 2329: 2327: 2271: 2269: 2267: 2215: 2213: 2176:"Effects of X-Rays upon Haploid and Diploid Embryos of 1091: 3045: 305:) and the cells in the digestive system (wall of the 2384: 2324: 2264: 2210: 1941: 2784: 2340:, as affected by dose, age, sex, and food intake". 1644: 60:. Unsourced material may be challenged and removed. 2687: 2685: 1511: 208:In the graph on left, a dose/survival curve for a 1452: 312: 269:Devair Alves Ferreira received a large dose (7.0 3069: 1453:Mattimore, V.; Battista, J. R. (February 1996). 1151:Ito H; Watanabe H; Takeshia M; Iizuka H (1983). 904:drug studied for its ability to protect against 2839:Proceedings of the National Academy of Sciences 2682: 2623: 1264:"Cosmic Influence on the Sun-Earth Environment" 336: 2335: 2275: 2239: 2237: 2235: 2219: 289:the radiation has less of an ability to cause 2923: 2173: 1238:Extremophiles: Microbiology and Biotechnology 398: 2525: 2523: 2521: 2519: 2517: 2067: 2232: 1588:Slade, Dea; Radman, Miroslav (March 2011). 1587: 383:is also a term sometimes used in medicine ( 2930: 2916: 2757:International Journal of Radiation Biology 2440: 2399:International Journal of Radiation Biology 1594:Microbiology and Molecular Biology Reviews 1396:"A review on natural background radiation" 1056: 194: 2886: 2869: 2859: 2727: 2717: 2665: 2655: 2514: 2470: 2150: 2140: 2044: 2034: 1981: 1971: 1918: 1858: 1621: 1556: 1486: 1429: 1411: 1370: 1305: 1287: 1261: 1183: 1168: 1118: 994: 984: 418:, but about 4 hours later it is of 7,000 120:Learn how and when to remove this message 391:cells which are difficult to treat with 293:. The human body contains many types of 198: 2937: 1328: 1234: 954: 14: 3070: 2889:International Journal of Low Radiation 2831: 2753: 2500: 2498: 2496: 2494: 2492: 2490: 2243: 1782:from a spacecraft assembly facility". 1538: 1022:living in a radioactive environment". 2911: 1157:Agricultural and Biological Chemistry 376:Radioresistance in radiation oncology 58:adding citations to reliable sources 29: 2999:Radiation-induced cognitive decline 2536:Australian Broadcasting Corporation 2487: 24: 2747: 1558:10.1111/j.1365-2958.1994.tb00397.x 1262:Mukherjee, Saumitra (2008-12-03). 333:, in a series of 14 publications. 25: 3099: 1092:Moseley BEB; Mattingly A (1971). 231:, plants, insects, as well as in 3055: 2174:Clark, AM; Mitchell, CJ (1952). 1851:10.1128/AEM.71.12.8147-8156.2005 1710:10.1046/j.1462-2920.2003.00496.x 884:has been reported for wild type 157:study in a hill in the state of 34: 2588: 2570: 2550: 2434: 2167: 2108: 2061: 1998: 1935: 1875: 1818: 1771: 1724: 1689: 1638: 1581: 1532: 1503: 1446: 1387: 1335:Life Sciences in Space Research 1322: 183:, can withstand large doses of 45:needs additional citations for 2246:Journal of Economic Entomology 1329:Kennedy, Ann R. (2014-04-01). 1255: 1235:Anitori, Roberto Paul (2012). 1228: 1177: 1144: 1135: 1085: 1050: 1011: 960: 313:Inheritance of radioresistance 13: 1: 3035:Radiation dose reconstruction 3004:Radiation-induced lung injury 2531:"Cockroaches & Radiation" 1471:10.1128/jb.178.3.633-637.1996 1111:10.1128/JB.105.3.976-983.1971 426:exposed to a dose of 180,250 2609:10.1016/0921-8777(96)00012-2 2280:by X-Rays and β radiation". 2036:10.1371/journal.pone.0066012 1973:10.1371/journal.pone.0000928 1400:Advanced Biomedical Research 1268:Sensors (Basel, Switzerland) 1241:. Horizon Scientific Press. 1071:10.1016/0027-5107(65)90052-7 1036:10.1016/0027-5107(73)90233-9 917:drug, that protects against 337:Evolution of radioresistance 7: 1539:Minton, K. W. (July 1994). 891: 10: 3104: 2901:10.1504/IJLR.2016.10002388 1514:DNA Repair and Mutagenesis 1355:10.1016/j.lssr.2014.02.004 1206:10.1103/PhysRevE.99.022139 858:Thermococcus gammatolerans 399:Radioresistance comparison 3012: 2981: 2945: 2770:10.1080/09553009414550111 2463:10.1093/genetics/30.2.197 2412:10.1080/09553000600972956 2142:10.1186/s12866-018-1191-y 2086:10.1007/s00792-013-0559-z 1749:10.1007/s00248-003-1029-4 1659:10.1134/S0006297915100016 3083:Radiation health effects 2994:Radiation-induced cancer 1647:Biochemistry. Biokhimiia 1413:10.4103/2277-9175.115821 919:acute radiation syndrome 906:acute radiation syndrome 470:Lethal radiation doses ( 319:Freie Universität Berlin 2719:10.1073/pnas.1119762109 2657:10.1073/pnas.0603791103 1459:Journal of Bacteriology 1170:10.1271/bbb1961.47.1239 986:10.1186/1471-2164-9-297 837:Deinococcus radiodurans 348:Deinococcus radiodurans 331:Drosophila melanogaster 240:and human cells and in 195:Induced radioresistance 176:Deinococcus radiodurans 173:, such as the bacteria 143:are able to withstand. 2861:10.1073/pnas.84.4.1045 2698:Proc Natl Acad Sci USA 2636:Proc Natl Acad Sci USA 2441:Heidenthal, G (1945). 1831:Appl Environ Microbiol 1804:10.1089/ast.2005.5.391 1545:Molecular Microbiology 205: 187:on the order of 5,000 2338:Periplaneta americana 1911:10.1089/ast.2009.0396 1606:10.1128/MMBR.00015-10 1020:Drosophila willistoni 814:Milnesium tardigradum 454:Macrobiotus areolatus 450:Milnesium tardigradum 446:Richtersius coronifer 442:Milnesium tardigradum 284:It has been found in 202: 3030:Radiation protection 2258:10.1093/jee/49.5.629 2021:(6:e66012): e66012. 955:Notes and references 934:Background radiation 169:and plants. Certain 54:improve this article 2973:Biological timeline 2939:Radiation poisoning 2852:1987PNAS...84.1045N 2832:Nothel, H. (1987). 2801:1989RadR..120..267L 2710:2012PNAS..109.2354K 2648:2006PNAS..103.9946W 2354:1959RadR...11..600W 2294:1954RadR....1..294G 2184:Biological Bulletin 2027:2013PLoSO...866012T 1964:2007PLoSO...2..928G 1903:2010AsBio..10..325G 1843:2005ApEnM..71.8147N 1796:2005AsBio...5..391K 1347:2014LSSR....1...10K 1280:2008Senso...8.7736M 1198:2019PhRvE..99b2139F 944:Radiotrophic fungus 793:Habrobracon hebetor 475: 407:Habrobracon hebetor 3062:Nuclear technology 3020:Dose fractionation 2788:Radiation Research 2342:Radiation Research 2282:Radiation Research 2072:SAFR-032 spores". 939:Radiation hormesis 469: 463:American cockroach 206: 185:ionizing radiation 151:Chernobyl disaster 137:ionizing radiation 3043: 3042: 1837:(12): 8147–8156. 1698:Environ Microbiol 1653:(10): 1201–1216. 1274:(12): 7736–7752. 1186:Physical Review E 1059:Mutation Research 1024:Mutation Research 878: 877: 325:using the common 286:radiation biology 130: 129: 122: 104: 69:"Radioresistance" 16:(Redirected from 3095: 3060: 3059: 3058: 3051: 2932: 2925: 2918: 2909: 2908: 2904: 2883: 2873: 2863: 2828: 2781: 2742: 2741: 2731: 2721: 2704:(7): 2354–2357. 2689: 2680: 2679: 2669: 2659: 2627: 2621: 2620: 2592: 2586: 2585: 2580:. Archived from 2574: 2568: 2567: 2565: 2564: 2554: 2548: 2547: 2545: 2544: 2527: 2512: 2502: 2485: 2484: 2474: 2438: 2432: 2431: 2393: 2382: 2381: 2333: 2322: 2321: 2273: 2262: 2261: 2241: 2230: 2229: 2217: 2208: 2207: 2171: 2165: 2164: 2154: 2144: 2112: 2106: 2105: 2070:Bacillus pumilus 2065: 2059: 2058: 2048: 2038: 2011:SAFR-032 Spores" 2009:Bacillus pumilus 2002: 1996: 1995: 1985: 1975: 1958:(9:e928): e928. 1946:Bacillus pumilus 1939: 1933: 1932: 1922: 1888: 1879: 1873: 1872: 1862: 1822: 1816: 1815: 1780:Bacillus pumilus 1775: 1769: 1768: 1733:Bacillus pumilus 1728: 1722: 1721: 1693: 1687: 1686: 1642: 1636: 1635: 1625: 1585: 1579: 1578: 1560: 1536: 1530: 1529: 1517: 1507: 1501: 1500: 1490: 1450: 1444: 1443: 1433: 1415: 1391: 1385: 1384: 1374: 1326: 1320: 1319: 1309: 1291: 1289:10.3390/s8127736 1259: 1253: 1252: 1232: 1226: 1225: 1181: 1175: 1174: 1172: 1148: 1142: 1139: 1133: 1132: 1122: 1089: 1083: 1082: 1054: 1048: 1047: 1015: 1009: 1008: 998: 988: 964: 929:Radiosensitivity 727:Common fruit fly 683:German cockroach 661:Escherichia coli 476: 468: 275:Goiânia accident 135:is the level of 125: 118: 114: 111: 105: 103: 62: 38: 30: 21: 3103: 3102: 3098: 3097: 3096: 3094: 3093: 3092: 3068: 3067: 3066: 3056: 3054: 3046: 3044: 3039: 3025:Radioresistance 3008: 2977: 2941: 2936: 2809:10.2307/3577713 2750: 2748:Further reading 2745: 2690: 2683: 2642:(26): 9946–51. 2628: 2624: 2593: 2589: 2576: 2575: 2571: 2562: 2560: 2556: 2555: 2551: 2542: 2540: 2529: 2528: 2515: 2503: 2488: 2439: 2435: 2394: 2385: 2362:10.2307/3570814 2334: 2325: 2302:10.2307/3570374 2274: 2265: 2242: 2233: 2218: 2211: 2196:10.2307/1538443 2172: 2168: 2113: 2109: 2066: 2062: 2003: 1999: 1940: 1936: 1886: 1880: 1876: 1823: 1819: 1776: 1772: 1729: 1725: 1694: 1690: 1643: 1639: 1586: 1582: 1537: 1533: 1526: 1508: 1504: 1451: 1447: 1392: 1388: 1327: 1323: 1260: 1256: 1249: 1233: 1229: 1182: 1178: 1149: 1145: 1140: 1136: 1090: 1086: 1055: 1051: 1016: 1012: 965: 961: 957: 915:radioprotective 902:radioprotective 894: 883: 779: 714: 645: 620: 597: 515: 496: 489: 460: 413: 401: 381:Radioresistance 378: 339: 315: 309:) cause death. 257:gene expression 197: 133:Radioresistance 126: 115: 109: 106: 63: 61: 51: 39: 28: 23: 22: 15: 12: 11: 5: 3101: 3091: 3090: 3085: 3080: 3065: 3064: 3041: 3040: 3038: 3037: 3032: 3027: 3022: 3016: 3014: 3010: 3009: 3007: 3006: 3001: 2996: 2991: 2989:Radiation burn 2985: 2983: 2979: 2978: 2976: 2975: 2970: 2965: 2960: 2955: 2949: 2947: 2943: 2942: 2935: 2934: 2927: 2920: 2912: 2906: 2905: 2884: 2829: 2782: 2749: 2746: 2744: 2743: 2681: 2622: 2603:(3): 201–208. 2587: 2584:on 2006-07-16. 2569: 2549: 2513: 2486: 2457:(2): 197–205. 2433: 2383: 2348:(4): 600–615. 2323: 2288:(3): 294–320. 2263: 2252:(5): 629–631. 2231: 2209: 2190:(2): 170–177. 2166: 2107: 2080:(5): 767–774. 2060: 1997: 1934: 1874: 1817: 1790:(3): 391–405. 1770: 1743:(2): 159–163. 1723: 1704:(10): 977–85. 1688: 1637: 1600:(1): 133–191. 1580: 1531: 1524: 1502: 1465:(3): 633–637. 1445: 1386: 1321: 1254: 1247: 1227: 1176: 1163:(6): 1239–47. 1143: 1134: 1084: 1049: 1010: 958: 956: 953: 952: 951: 946: 941: 936: 931: 926: 908: 893: 890: 881: 876: 875: 870: 867: 864: 861: 853: 852: 849: 846: 843: 840: 832: 831: 826: 823: 820: 817: 809: 808: 805: 802: 799: 796: 788: 787: 784: 781: 777: 774: 771: 765: 764: 759: 756: 753: 750: 742: 741: 738: 735: 732: 729: 723: 722: 719: 716: 712: 709: 706: 700: 699: 694: 691: 688: 685: 679: 678: 673: 670: 667: 664: 656: 655: 650: 647: 643: 640: 637: 631: 630: 625: 622: 618: 615: 612: 606: 605: 602: 599: 595: 592: 589: 583: 582: 579: 576: 573: 570: 564: 563: 560: 557: 554: 551: 545: 544: 541: 538: 535: 532: 526: 525: 520: 517: 513: 510: 507: 501: 500: 499:Class/Kingdom 497: 494: 491: 487: 483: 480: 458: 411: 400: 397: 377: 374: 362:D. radiodurans 353:D. radiodurans 338: 335: 314: 311: 255:and increased 196: 193: 128: 127: 42: 40: 33: 26: 18:Radioresistant 9: 6: 4: 3: 2: 3100: 3089: 3086: 3084: 3081: 3079: 3076: 3075: 3073: 3063: 3053: 3052: 3049: 3036: 3033: 3031: 3028: 3026: 3023: 3021: 3018: 3017: 3015: 3011: 3005: 3002: 3000: 2997: 2995: 2992: 2990: 2987: 2986: 2984: 2980: 2974: 2971: 2969: 2966: 2964: 2961: 2959: 2956: 2954: 2951: 2950: 2948: 2944: 2940: 2933: 2928: 2926: 2921: 2919: 2914: 2913: 2910: 2902: 2898: 2895:(3): 222–33. 2894: 2890: 2885: 2881: 2877: 2872: 2867: 2862: 2857: 2853: 2849: 2846:(4): 1045–9. 2845: 2841: 2840: 2835: 2830: 2826: 2822: 2818: 2814: 2810: 2806: 2802: 2798: 2795:(2): 267–79. 2794: 2790: 2789: 2783: 2779: 2775: 2771: 2767: 2763: 2759: 2758: 2752: 2751: 2739: 2735: 2730: 2725: 2720: 2715: 2711: 2707: 2703: 2699: 2695: 2688: 2686: 2677: 2673: 2668: 2663: 2658: 2653: 2649: 2645: 2641: 2637: 2633: 2626: 2618: 2614: 2610: 2606: 2602: 2598: 2591: 2583: 2579: 2573: 2559: 2553: 2538: 2537: 2532: 2526: 2524: 2522: 2520: 2518: 2511: 2510:0-7506-7463-6 2507: 2501: 2499: 2497: 2495: 2493: 2491: 2482: 2478: 2473: 2468: 2464: 2460: 2456: 2452: 2448: 2446: 2437: 2429: 2425: 2421: 2417: 2413: 2409: 2406:(12): 843–8. 2405: 2401: 2400: 2392: 2390: 2388: 2379: 2375: 2371: 2367: 2363: 2359: 2355: 2351: 2347: 2343: 2339: 2332: 2330: 2328: 2319: 2315: 2311: 2307: 2303: 2299: 2295: 2291: 2287: 2283: 2279: 2272: 2270: 2268: 2259: 2255: 2251: 2247: 2240: 2238: 2236: 2227: 2223: 2216: 2214: 2205: 2201: 2197: 2193: 2189: 2185: 2181: 2179: 2170: 2162: 2158: 2153: 2148: 2143: 2138: 2134: 2130: 2129:BMC Microbiol 2126: 2124: 2120: 2111: 2103: 2099: 2095: 2091: 2087: 2083: 2079: 2075: 2074:Extremophiles 2071: 2064: 2056: 2052: 2047: 2042: 2037: 2032: 2028: 2024: 2020: 2016: 2012: 2010: 2001: 1993: 1989: 1984: 1979: 1974: 1969: 1965: 1961: 1957: 1953: 1949: 1947: 1938: 1930: 1926: 1921: 1920:2027.42/85129 1916: 1912: 1908: 1904: 1900: 1897:(3): 325–35. 1896: 1892: 1885: 1878: 1870: 1866: 1861: 1856: 1852: 1848: 1844: 1840: 1836: 1832: 1828: 1821: 1813: 1809: 1805: 1801: 1797: 1793: 1789: 1785: 1781: 1774: 1766: 1762: 1758: 1754: 1750: 1746: 1742: 1738: 1734: 1727: 1719: 1715: 1711: 1707: 1703: 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ASM Press. 1516: 1515: 1506: 1498: 1494: 1489: 1484: 1480: 1476: 1472: 1468: 1464: 1460: 1456: 1449: 1441: 1437: 1432: 1427: 1423: 1419: 1414: 1409: 1405: 1401: 1397: 1390: 1382: 1378: 1373: 1368: 1364: 1360: 1356: 1352: 1348: 1344: 1340: 1336: 1332: 1325: 1317: 1313: 1308: 1303: 1299: 1295: 1290: 1285: 1281: 1277: 1273: 1269: 1265: 1258: 1250: 1248:9781904455981 1244: 1240: 1239: 1231: 1223: 1219: 1215: 1211: 1207: 1203: 1199: 1195: 1192:(2): 022139. 1191: 1187: 1180: 1171: 1166: 1162: 1158: 1154: 1147: 1138: 1130: 1126: 1121: 1116: 1112: 1108: 1105:(3): 976–83. 1104: 1101: 1100: 1099:J. Bacteriol. 1095: 1088: 1080: 1076: 1072: 1068: 1065:(5): 403–12. 1064: 1060: 1053: 1045: 1041: 1037: 1033: 1029: 1025: 1021: 1014: 1006: 1002: 997: 992: 987: 982: 978: 974: 970: 963: 959: 950: 947: 945: 942: 940: 937: 935: 932: 930: 927: 924: 920: 916: 912: 909: 907: 903: 899: 896: 895: 889: 887: 880:∗ While an LD 874: 871: 868: 865: 862: 860: 859: 855: 854: 850: 847: 844: 841: 839: 838: 834: 833: 830: 827: 824: 821: 818: 816: 815: 811: 810: 806: 803: 800: 797: 795: 794: 790: 789: 785: 782: 775: 772: 770: 767: 766: 763: 760: 757: 754: 751: 749: 748: 744: 743: 739: 736: 733: 730: 728: 725: 724: 720: 717: 710: 707: 705: 702: 701: 698: 695: 692: 689: 686: 684: 681: 680: 677: 674: 671: 668: 665: 663: 662: 658: 657: 654: 651: 648: 641: 638: 636: 633: 632: 629: 626: 623: 616: 613: 611: 608: 607: 603: 600: 593: 590: 588: 585: 584: 580: 577: 574: 571: 569: 566: 565: 561: 558: 555: 552: 550: 547: 546: 542: 539: 536: 533: 531: 528: 527: 524: 521: 518: 511: 508: 506: 503: 502: 498: 492: 490: 484: 481: 478: 477: 473: 467: 464: 455: 451: 447: 443: 439: 434: 429: 425: 421: 417: 409: 408: 396: 394: 390: 386: 382: 373: 370: 366: 363: 360:species like 359: 354: 349: 343: 334: 332: 328: 324: 320: 310: 308: 304: 300: 296: 292: 287: 282: 280: 276: 273:) during the 272: 267: 264: 262: 258: 254: 251: 247: 243: 242:animal models 239: 236: 235: 230: 226: 222: 218: 213: 211: 201: 192: 190: 186: 182: 178: 177: 172: 171:extremophiles 168: 164: 160: 156: 152: 147: 144: 142: 138: 134: 124: 121: 113: 102: 99: 95: 92: 88: 85: 81: 78: 74: 71: –  70: 66: 65:Find sources: 59: 55: 49: 48: 43:This article 41: 37: 32: 31: 19: 3088:Radiodurants 3078:Radiobiology 3024: 2892: 2888: 2843: 2837: 2792: 2786: 2764:(1): 79–84. 2761: 2755: 2701: 2697: 2639: 2635: 2625: 2600: 2596: 2590: 2582:the original 2572: 2561:. 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83: 76: 64: 52:Please help 47:verification 44: 2968:Experiments 2445:Habrobracon 2278:Habrobracon 2178:Habrobracon 2121:FO-36b and 2119:B. safensis 1737:Microb Ecol 1551:(1): 9–15. 482:Lethal dose 358:Deinococcus 303:bone marrow 299:blood cells 246:cytoplasmic 181:tardigrades 3072:Categories 3013:Treatments 2982:Conditions 2563:2018-06-16 2543:2006-05-13 2222:Nucleotics 2135:(57): 57. 2123:B. pumilus 949:Kojic acid 913:a similar 886:C. elegans 778:50/30 days 747:C. elegans 713:50/30 days 644:50/30 days 619:50/30 days 596:50/30 days 514:50/30 days 438:H. hebetor 433:H. hebetor 424:H. hebetor 307:intestines 291:cell death 261:DNA repair 80:newspapers 2963:Accidents 2597:Mutat Res 1948:SAFR-032" 1667:1608-3040 1614:1098-5557 1567:0950-382X 1479:0021-9193 1422:2277-9175 1406:(1): 65. 1363:2214-5524 1341:: 10–43. 1298:1424-8220 921:, during 851:Bacteria 776:1,000 (LD 758:≫ 500-800 704:Shellfish 327:fruit fly 323:mutations 279:Chernobyl 238:mammalian 155:Brazilian 141:organisms 2738:22308443 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1983:1976550 1960:Bibcode 1899:Bibcode 1860:1317311 1839:Bibcode 1792:Bibcode 1623:3063356 1575:7984097 1497:8550493 1431:3814895 1372:4170231 1343:Bibcode 1307:3790986 1276:Bibcode 1194:Bibcode 1129:4929286 1079:5878261 1044:4796403 996:2441631 979:: 297. 911:CBLB502 873:Archaea 711:200 (LD 697:Insects 628:Reptile 523:Mammals 512:3.5 (LD 250:nuclear 163:insects 94:scholar 3048:Portal 2878:  2871:304358 2868:  2823:  2815:  2776:  2736:  2726:  2674:  2664:  2615:  2508:  2479:  2469:  2426:  2418:  2376:  2368:  2316:  2308:  2202:  2159:  2149:  2100:  2092:  2053:  2043:  1990:  1980:  1927:  1867:  1857:  1810:  1763:  1755:  1716:  1681:  1673:  1665:  1630:  1620:  1612:  1573:  1565:  1522:  1495:  1488:177705 1485:  1477:  1438:  1428:  1420:  1379:  1369:  1361:  1314:  1304:  1296:  1245:  1220:  1212:  1127:  1120:248526 1117:  1077:  1042:  1003:  993:  898:Ex-Rad 869:  866:  863:30,000 848:  845:  842:15,000 825:  822:  804:  801:  783:  773:  769:Amoeba 752:  737:  734:  718:  708:  693:  687:  669:  649:  642:20 (LD 639:  624:  617:15 (LD 614:  601:  591:  587:Rabbit 578:  572:4.5–12 559:  553:  519:  509:  389:cancer 387:) for 96:  89:  82:  75:  67:  2953:Acute 2813:JSTOR 2424:S2CID 2366:JSTOR 2306:JSTOR 2200:JSTOR 2098:S2CID 1887:(PDF) 1761:S2CID 1679:S2CID 1218:S2CID 819:5,000 798:1,800 594:8 (LD 575:8.6–9 568:Mouse 530:Human 295:cells 229:algae 217:yeast 167:worms 139:that 101:JSTOR 87:books 2876:PMID 2821:PMID 2774:PMID 2734:PMID 2672:PMID 2613:PMID 2506:ISBN 2477:PMID 2416:PMID 2374:PMID 2314:PMID 2157:PMID 2090:PMID 2051:PMID 1988:PMID 1925:PMID 1865:PMID 1808:PMID 1753:PMID 1714:PMID 1671:PMID 1663:ISSN 1628:PMID 1610:ISSN 1571:PMID 1563:ISSN 1520:ISBN 1493:PMID 1475:ISSN 1436:PMID 1418:ISSN 1377:PMID 1359:ISSN 1312:PMID 1294:ISSN 1243:ISBN 1210:PMID 1125:PMID 1075:PMID 1040:PMID 1001:PMID 653:Fish 534:4–10 472:Gray 448:and 248:and 73:news 2897:doi 2866:PMC 2856:doi 2805:doi 2793:120 2766:doi 2724:PMC 2714:doi 2702:109 2662:PMC 2652:doi 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