526:(ROS), can lead to DNA damage; however, it is yet unclear whether the elevated rate in telomeres is brought about by their inherent susceptibility or a diminished activity of DNA repair systems in these regions. Despite widespread agreement of the findings, widespread flaws regarding measurement and sampling have been pointed out; for example, a suspected species and tissue dependency of oxidative damage to telomeres is said to be insufficiently accounted for. Population-based studies have indicated an interaction between anti-oxidant intake and telomere length. In the Long Island Breast Cancer Study Project (LIBCSP), authors found a moderate increase in breast cancer risk among women with the shortest telomeres and lower dietary intake of beta carotene, vitamin C or E. These results suggest that cancer risk due to telomere shortening may interact with other mechanisms of DNA damage, specifically oxidative stress.
305:). The last primer to be involved in lagging-strand replication sits near the 3'-end of the template (corresponding to the potential 5'-end of the lagging-strand). Originally it was believed that the last primer would sit at the very end of the template, thus, once removed, the DNA-polymerase that substitutes primers with DNA (DNA-Pol δ in eukaryotes) would be unable to synthesize the "replacement DNA" from the 5'-end of the lagging strand so that the template nucleotides previously paired to the last primer would not be replicated. It has since been questioned whether the last lagging strand primer is placed exactly at the 3'-end of the template and it was demonstrated that it is rather synthesized at a distance of about 70–100 nucleotides which is consistent with the finding that DNA in cultured human cell is shortened by 50–100
426:
219:
that determines the number of divisions that a certain cell clone can undergo. Furthermore, it was predicted that a specialized DNA polymerase (originally called a tandem-DNA-polymerase) could extend telomeres in immortal tissues such as germ line, cancer cells and stem cells. It also followed from this hypothesis that organisms with circular genome, such as bacteria, do not have the end replication problem and therefore do not age.
585:
4990:
5489:
1284:
across vertebrates. Phylogeny and life history traits such as body size or the pace of life can also affect telomere dynamics. For example, it has been described across species of birds and mammals. In 2019, a meta-analysis confirmed that the exposure to stressors (e.g. pathogen infection, competition, reproductive effort and high activity level) was associated with shorter telomeres across different animal taxa.
1307:
276:
1122:
1214:
5501:
576:. The literature concerning telomeres as integrative biomarkers of exposure to stress and adversity is dominated by cross-sectional and correlational studies, which makes causal interpretation problematic. A 2020 review argued that the relationship between psychosocial stress and telomere length appears strongest for stress experienced in utero or early life.
1255:(WGS) experiments. Amongst these are TelSeq, Telomerecat and telomereHunter. Length estimation from WGS typically works by differentiating telomere sequencing reads and then inferring the length of telomere that produced that number of reads. These methods have been shown to correlate with preexisting methods of estimation such as PCR and TRF.
359:
42:
612:, that tied telomere shortening with the Hayflick limit. The cloning of the catalytic component of telomerase enabled experiments to test whether the expression of telomerase at levels sufficient to prevent telomere shortening was capable of immortalizing human cells. Telomerase was demonstrated in a 1998 publication in
301:) then being excised and substituted by DNA. The lagging strand, however, is oriented 3'-5' with respect to the replication fork so continuous replication by DNA-polymerase is impossible, which necessitates discontinuous replication involving the repeated synthesis of primers further 5' of the site of initiation (see
636:
A study reported that telomere length of different mammalian species correlates inversely rather than directly with lifespan, and concluded that the contribution of telomere length to lifespan remains controversial. There is little evidence that, in humans, telomere length is a significant biomarker
372:
At the very 3'-end of the telomere there is a 300 base pair overhang which can invade the double-stranded portion of the telomere forming a structure known as a T-loop. This loop is analogous to a knot, which stabilizes the telomere, and prevents the telomere ends from being recognized as breakpoints
1283:
estimates vary greatly within and among species. Age and telomere length often negatively correlate in vertebrates, but this decline is variable among taxa and linked to the method used for estimating telomere length. In contrast, the available information shows no sex differences in telomere length
316:
If coding sequences are degraded in this process, potentially vital genetic code would be lost. Telomeres are non-coding, repetitive sequences located at the termini of linear chromosomes to act as buffers for those coding sequences further behind. They "cap" the end-sequences and are progressively
218:
division, Olovnikov suggested that DNA sequences are lost every time a cell replicates until the loss reaches a critical level, at which point cell division ends. According to his theory of marginotomy, DNA sequences at the ends of telomeres are represented by tandem repeats, which create a buffer
439:
Many organisms have a ribonucleoprotein enzyme called telomerase, which carries out the task of adding repetitive nucleotide sequences to the ends of the DNA. Telomerase "replenishes" the telomere "cap" and requires no ATP. In most multicellular eukaryotic organisms, telomerase is active only in
488:
ranging from 75 to 300 bases, which is essential for telomere maintenance and capping. Multiple proteins binding single- and double-stranded telomere DNA have been identified. These function in both telomere maintenance and capping. Telomeres form large loop structures called telomere loops, or
373:
by the DNA repair machinery. Should non-homologous end joining occur at the telomeric ends, chromosomal fusion would result. The T-loop is maintained by several proteins, collectively referred to as the shelterin complex. In humans, the shelterin complex consists of six proteins identified as
493:. At the very end of the T-loop, the single-stranded telomere DNA is held onto a region of double-stranded DNA by the telomere strand disrupting the double-helical DNA, and base pairing to one of the two strands. This triple-stranded structure is called a
468:. This is because the telomeres act as a sort of time-delay "fuse", eventually running out after a certain number of cell divisions and resulting in the eventual loss of vital genetic information from the cell's chromosome with future divisions.
195:, working with maize. Muller observed that the ends of irradiated fruit fly chromosomes did not present alterations such as deletions or inversions. He hypothesized the presence of a protective cap, which he coined "telomeres", from the Greek
552:
Telomere shortening is associated with aging, mortality, and aging-related diseases in experimental animals. Although many factors can affect human lifespan, such as smoking, diet, and exercise, as persons approach the upper limit of human
417:) to form such G-quadruplexes that accommodate it, rather than a T-loop. G-quadruplexes present an obstacle for enzymes such as DNA-polymerases and are thus thought to be involved in the regulation of replication and transcription.
409:, a special conformation of DNA involving non-Watson-Crick base pairing. There are different subtypes depending on the involvement of single- or double-stranded DNA, among other things. There is evidence for the 3'-overhang in
1239:
Several techniques are currently employed to assess average telomere length in eukaryotic cells. One method is the
Terminal Restriction Fragment (TRF) southern blot. There is a Web-based Analyser of the Length of Telomeres
296:
to initiate replication. On the leading strand (oriented 5'-3' within the replication fork), DNA-polymerase continuously replicates from the point of initiation all the way to the strand's end with the primer (made of
540:
Although telomeres shorten during the lifetime of an individual, it is telomere shortening-rate rather than telomere length that is associated with the lifespan of a species. Critically short telomeres trigger a
1274:
During the last two decades, eco-evolutionary studies have investigated the relevance of life-history traits and environmental conditions on telomeres of wildlife. Most of these studies have been conducted in
291:
of the parent strands. This is a consequence of its unidirectional mode of DNA synthesis: it can only attach new nucleotides to an existing 3'-end (that is, synthesis progresses 5'-3') and thus it requires a
3462:
Fajkus, Petr; Adámik, Matej; Nelson, Andrew D L; Kilar, Agata M; Franek, Michal; Bubeník, Michal; Frydrychová, Radmila Čapková; Votavová, Alena; Sýkorová, Eva; Fajkus, Jiří; Peška, Vratislav (2023-01-11).
2023:
Lanza RP, Cibelli JB, Blackwell C, Cristofalo VJ, Francis MK, Baerlocher GM, et al. (April 2000). "Extension of cell life-span and telomere length in animals cloned from senescent somatic cells".
1259:
is used to quantify the length of telomeres in human white blood cells. A semi-automated method for measuring the average length of telomeres with Flow FISH was published in Nature
Protocols in 2006.
346:), however, are linear and possess telomeres, which are very different from those of the eukaryotic chromosomes in structure and function. The known structures of bacterial telomeres take the form of
5474:
1689:
Olovnikov AM (September 1973). "A theory of marginotomy. The incomplete copying of template margin in enzymic synthesis of polynucleotides and biological significance of the phenomenon".
4913:
1262:
While multiple companies offer telomere length measurement services, the utility of these measurements for widespread clinical or personal use has been questioned. Nobel Prize winner
1248:
assay for telomere length involves determining the
Telomere-to-Single Copy Gene (T/S) ratio, which is demonstrated to be proportional to the average telomere length in a cell.
1132:
3111:
Bodnar AG, Ouellette M, Frolkis M, Holt SE, Chiu CP, Morin GB, et al. (January 1998). "Extension of life-span by introduction of telomerase into normal human cells".
2755:
243:
1291:, and other non-mammalian organisms, show that there is no single universal model of telomere erosion; rather, there is wide variation in relevant dynamics across
3269:
Harris SE, Martin-Ruiz C, von
Zglinicki T, Starr JM, Deary IJ (July 2012). "Telomere length and aging biomarkers in 70-year-olds: the Lothian Birth Cohort 1936".
210:
first recognized that chromosomes could not completely replicate their ends; this is known as the "end replication problem". Building on this, and accommodating
4414:
592:
is the theoretical limit to the number of times a cell may divide until the telomere becomes so short that division is inhibited and the cell enters senescence.
535:
5539:
1522:"A theory of marginotomy: The incomplete copying of template margin in enzymic synthesis of polynucleotides and biological significance of the phenomenon"
588:
The average cell will divide between 50 and 70 times before cell death. As the cell divides the telomeres on the end of the chromosome get smaller. The
320:
The "end replication problem" is exclusive to linear chromosomes as circular chromosomes do not have ends lying without reach of DNA-polymerases. Most
1228:
572:
was associated with a small decrease in telomere length—but that these associations attenuate to no significant association when accounting for
549:. Mice have much longer telomeres, but a greatly accelerated telomere shortening-rate and greatly reduced lifespan compared to humans and elephants.
4979:
4946:
4133:
Remot, Florentin; Ronget, Victor; Froy, Hannah; Rey, Benjamin; Gaillard, Jean-Michel; Nussey, Daniel H.; Lemaître, Jean-François (November 2020).
3877:
Baerlocher GM, Vulto I, de Jong G, Lansdorp PM (December 2006). "Flow cytometry and FISH to measure the average length of telomeres (flow FISH)".
3312:
Lyčka, Martin; Bubeník, Michal; Závodník, Michal; Peska, Vratislav; Fajkus, Petr; Demko, Martin; Fajkus, Jiří; Fojtová, Miloslava (2023-08-21).
645:
Experimentally verified and predicted telomere sequence motifs from more than 9000 species are collected in research community curated database
5155:
4068:
Remot, Florentin; Ronget, Victor; Froy, Hannah; Rey, Benjamin; Gaillard, Jean-Michel; Nussey, Daniel H.; Lemaitre, Jean-François (2021-09-07).
1633:
Blackburn EH, Gall JG (March 1978). "A tandemly repeated sequence at the termini of the extrachromosomal ribosomal RNA genes in
Tetrahymena".
4903:
4544:
1734:"Early and late steps in telomere overhang processing in normal human cells: the position of the final RNA primer drives telomere shortening"
246:
4908:
2794:"Divergence of sperm and leukocyte age-dependent telomere dynamics: implications for male-driven evolution of telomere length in humans"
5725:
5712:
3371:"Characterisation of an unusual telomere motif (TTTTTTAGGG)n in the plant Cestrum elegans (Solanaceae), a species with a large genome"
288:
6038:
5735:
4407:
3612:
Lyčka, Martin; Peska, Vratislav; Demko, Martin; Spyroglou, Ioannis; Kilar, Agata; Fajkus, Jiří; Fojtová, Miloslava (December 2021).
1794:
3060:
Feng J, Funk WD, Wang SS, Weinrich SL, Avilion AA, Chiu CP, et al. (September 1995). "The RNA component of human telomerase".
2646:"Perceived stress and telomere length: A systematic review, meta-analysis, and methodologic considerations for advancing the field"
3222:"Comparative biology of mammalian telomeres: hypotheses on ancestral states and the roles of telomeres in longevity determination"
2947:
Rentscher KE, Carroll JE, Mitchell C (April 2020). "Psychosocial
Stressors and Telomere Length: A Current Review of the Science".
5532:
5464:
5186:
1819:
3569:
Rufer N, et al. (August 1998). "Telomere length dynamics in human lymphocyte subpopulations measured by flow cytometry".
5469:
1371:
234:, discovered the unusual nature of telomeres, with their simple repeated DNA sequences composing chromosome ends. Blackburn,
5766:
5612:
5599:
5313:
5080:
4529:
457:
4400:
4208:"On the comparative biology of mammalian telomeres: Telomere length co-evolves with body mass, lifespan and cancer risk"
625:
demonstrate that the role of telomeres is far from being understood. In 2003, scientists observed that the telomeres of
350:
bound to the ends of linear chromosomes, or hairpin loops of single-stranded DNA at the ends of the linear chromosomes.
31:
1193:
1140:
5303:
4939:
618:
to be capable of extending cell lifespan, and now is well-recognized as capable of immortalizing human somatic cells.
1165:
6033:
5883:
5525:
5160:
1330:
293:
1151:
1136:
425:
5771:
4974:
4676:
1245:
983:
35:
1172:
5505:
5330:
5009:
4969:
4730:
4514:
1361:
1087:
1001:
490:
2343:"Characterization of the yeast telomere nucleoprotein core: Rap1 binds independently to each recognition site"
4094:
2892:"Telomeres as integrative markers of exposure to stress and adversity: a systematic review and meta-analysis"
1015:
132:
115:
6048:
5577:
5552:
5517:
5226:
5004:
1252:
736:
626:
523:
187:
1669:"Elizabeth H. Blackburn, Carol W. Greider, Jack W. Szostak: The Nobel Prize in Physiology or Medicine 2009"
1075:
6007:
5935:
5828:
5493:
5398:
5383:
5211:
5105:
5085:
4932:
4683:
4666:
4636:
4609:
1179:
108:
1241:
5853:
5803:
4671:
4569:
3771:"Telomerecat: A ploidy-agnostic method for estimating telomere length from whole genome sequencing data"
3412:"Allium telomeres unmasked: the unusual telomeric sequence (CTCGGTTATGGG)n is synthesized by telomerase"
1668:
658:
485:
182:
5863:
1279:, i.e. birds and mammals. They have provided evidence for the inheritance of telomere length; however,
633:) seem to lengthen with chronological age, the first observed instance of such behaviour of telomeres.
5985:
5903:
1223:
Preliminary research indicates that disease risk in aging may be associated with telomere shortening,
460:. The steady shortening of telomeres with each replication in somatic (body) cells may have a role in
181:
The existence of a special structure at the ends of chromosomes was independently proposed in 1938 by
6028:
5980:
5893:
5808:
5694:
5556:
5423:
5286:
5150:
5100:
4891:
4816:
4801:
4631:
4579:
4554:
4494:
4146:
3782:
3614:"WALTER: an easy way to online evaluate telomere lengths from terminal restriction fragment analysis"
3178:
3120:
3069:
2903:
2299:
2087:
2032:
1893:
1698:
1533:
1345:
1161:
1099:
1063:
965:
868:
569:
449:
5758:
5629:
5308:
5291:
5271:
5251:
4559:
4368:
2704:
1263:
820:
546:
223:
170:
6043:
5930:
5838:
5813:
5657:
5318:
5064:
4294:
4188:
4115:
3964:
3902:
3594:
3441:
3294:
3202:
3144:
3093:
2990:
Hayflick L, Moorhead PS (December 1961). "The serial cultivation of human diploid cell strains".
2972:
2423:
2323:
2254:
2056:
1612:
1448:
1051:
796:
192:
3410:
Fajkus P, Peška V, Sitová Z, Fulnečková J, Dvořáčková M, Gogela R, et al. (February 2016).
1266:, who was co-founder of one company, promoted the clinical utility of telomere length measures.
2274:
Telomeric 8-oxo-guanine drives rapid premature senescence in the absence of telomere shortening
5908:
5443:
5438:
5393:
5216:
5090:
4621:
4533:
4343:
4286:
4278:
4237:
4229:
4180:
4162:
4107:
4099:
4050:
4032:
3956:
3894:
3859:
3808:
3751:
3702:
3653:
3635:
3586:
3551:
3502:
3484:
3433:
3392:
3351:
3333:
3286:
3251:
3194:
3136:
3085:
3042:
3007:
2964:
2929:
2872:
2823:
2774:
2731:
2675:
2623:
2574:
2521:
2472:
2415:
2374:
2315:
2290:
Shampay J, Szostak JW, Blackburn EH (1984). "DNA sequences of telomeres maintained in yeast".
2246:
2205:
2164:
2123:
2105:
2074:
Whittemore, Kurt; Vera, Elsa; Martínez-Nevado, Eva; Sanpera, Carola; Blasco, Maria A. (2019).
2048:
2005:
1956:
1921:
1862:
1800:
1790:
1763:
1714:
1650:
1604:
1596:
1557:
1549:
1502:
1494:
1440:
1432:
1340:
936:
713:
614:
605:
494:
5873:
5823:
5818:
4070:"Decline in telomere length with increasing age across nonhuman vertebrates: A meta-analysis"
3828:"TelomereHunter–in silico estimation of telomere content and composition from cancer genomes"
2960:
324:, relying on circular chromosomes, accordingly do not possess telomeres. A small fraction of
5992:
5922:
5848:
5798:
5560:
5373:
5340:
5095:
4811:
4710:
4333:
4325:
4268:
4219:
4170:
4154:
4089:
4081:
4040:
4024:
3991:
3948:
3886:
3849:
3839:
3798:
3790:
3741:
3733:
3692:
3684:
3643:
3625:
3578:
3541:
3533:
3492:
3476:
3423:
3382:
3341:
3325:
3278:
3241:
3233:
3220:
Gomes NM, Ryder OA, Houck ML, Charter SJ, Walker W, Forsyth NR, et al. (October 2011).
3186:
3128:
3077:
3034:
2999:
2956:
2919:
2911:
2862:
2854:
2813:
2805:
2764:
2721:
2713:
2665:
2657:
2613:
2605:
2564:
2556:
2511:
2503:
2462:
2454:
2405:
2364:
2354:
2307:
2236:
2195:
2154:
2113:
2095:
2040:
1995:
1987:
1948:
1939:
Lipps HJ, Rhodes D (August 2009). "G-quadruplex structures: in vivo evidence and function".
1911:
1901:
1852:
1753:
1745:
1706:
1642:
1588:
1541:
1424:
1320:
1039:
1027:
948:
597:
573:
515:
453:
211:
207:
60:
5843:
5748:
4866:
4861:
4257:"The association between stressors and telomeres in non-human vertebrates: a meta-analysis"
2792:
Aston KI, Hunt SC, Susser E, Kimura M, Factor-Litvak P, Carrell D, Aviv A (November 2012).
2392:
Griffith JD, Comeau L, Rosenfield S, Stansel RM, Bianchi A, Moss H, de Lange T (May 1999).
518:
and that oxidative stress-mediated DNA damage has a major influence on telomere shortening
5868:
5703:
5675:
5644:
5639:
5206:
5201:
5191:
4703:
4524:
4472:
3921:
3369:
Peška V, Fajkus P, Fojtová M, Dvořáčková M, Hapala J, Dvořáček V, et al. (May 2015).
2592:
Shen J, Gammon MD, Terry MB, Wang Q, Bradshaw P, Teitelbaum SL, et al. (April 2009).
880:
609:
554:
484:-rich, six- to eight-base-pair-long repeats. Eukaryotic telomeres normally terminate with
270:
227:
165:
from progressive degradation and ensure the integrity of linear chromosomes by preventing
3282:
4318:
Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
4150:
3786:
3522:"Human telomeres contain at least three types of G-rich repeat distributed non-randomly"
3464:
3346:
3182:
3124:
3073:
3025:
Hayflick L (March 1965). "The limited in vitro lifetime of human diploid cell strains".
2907:
2303:
2091:
2036:
1897:
1702:
1537:
5975:
5833:
5790:
5618:
5608:
5378:
5355:
5261:
5256:
5196:
5181:
5145:
5030:
4594:
4518:
4338:
4313:
4175:
4045:
3854:
3827:
3803:
3770:
3746:
3721:
3648:
3613:
3497:
3246:
3221:
2924:
2891:
2867:
2842:
2818:
2793:
2726:
2699:
2670:
2645:
2618:
2593:
2569:
2540:
2516:
2491:
2467:
2442:
2369:
2342:
2200:
2183:
2118:
2075:
2000:
1975:
1783:
1758:
1733:
1312:
1224:
695:
604:. Significant discoveries were subsequently made by a group of scientists organized at
601:
589:
302:
284:
231:
4989:
4383:
4374:
4371:, which includes a reference to the impact of stress, and pessimism on telomere length
4364:
3697:
3672:
3546:
3521:
3314:"TeloBase: a community-curated database of telomere sequences across the tree of life"
2410:
2393:
1916:
1881:
1521:
1186:
6022:
5940:
5680:
5603:
5594:
5586:
5276:
5231:
5140:
4955:
4574:
4378:
4298:
4192:
4119:
3445:
3237:
3190:
3038:
3003:
2976:
2273:
1857:
1840:
1710:
1646:
1592:
1545:
907:
759:
565:
336:
310:
235:
3968:
3906:
3598:
3298:
3206:
3163:
3148:
2258:
2060:
1616:
1452:
654:
489:
T-loops. Here, the single-stranded DNA curls around in a long circle, stabilized by
161:. In most, if not all species possessing them, they protect the terminal regions of
5952:
5428:
5413:
5325:
5266:
5165:
5135:
5044:
4549:
4423:
4387:
3465:"Telomerase RNA in Hymenoptera (Insecta) switched to plant/ciliate-like biogenesis"
3097:
2327:
1366:
1280:
748:
690:
406:
330:
239:
215:
17:
2427:
584:
456:. Telomerase can be reactivated and telomeres reset back to an embryonic state by
4135:"No sex differences in adult telomere length across vertebrates: a meta-analysis"
3132:
2341:
Williams TL, Levy DL, Maki-Yonekura S, Yonekura K, Blackburn EH (November 2010).
2044:
5947:
5858:
5720:
5418:
5408:
5403:
4768:
4763:
4626:
4604:
4599:
4467:
2644:
Mathur MB, Epel E, Kind S, Desai M, Parks CG, Sandler DP, Khazeni N (May 2016).
1335:
924:
843:
826:
790:
766:
724:
3794:
3630:
2769:
2750:
2717:
1886:
Proceedings of the
National Academy of Sciences of the United States of America
1306:
5959:
5776:
5345:
5298:
5221:
5130:
4788:
4778:
4773:
4758:
4698:
4688:
4589:
4564:
4538:
4489:
4479:
4427:
3844:
2661:
2507:
1952:
1350:
1325:
1302:
903:
850:
808:
681:
650:
542:
461:
434:
414:
402:
390:
321:
253:
166:
150:
146:
46:
4282:
4233:
4166:
4103:
4036:
3639:
3537:
3488:
3337:
2278:
2109:
1804:
1600:
1553:
1498:
1483:"[Principle of marginotomy in template synthesis of polynucleotides]"
1436:
5878:
5569:
4740:
4484:
4462:
4443:
4392:
4012:
3688:
3081:
2809:
2359:
2159:
2142:
2100:
1991:
1906:
1412:
1356:
1288:
1276:
1256:
959:
772:
637:
of normal aging with respect to important cognitive and physical abilities.
477:
445:
441:
367:
306:
275:
158:
4347:
4329:
4290:
4241:
4184:
4111:
4054:
4028:
3960:
3898:
3890:
3863:
3812:
3755:
3706:
3657:
3506:
3480:
3437:
3396:
3355:
3290:
3255:
3198:
3046:
3011:
2968:
2933:
2876:
2827:
2778:
2735:
2679:
2627:
2578:
2560:
2525:
2476:
2419:
2378:
2241:
2224:
2209:
2168:
2127:
2052:
2009:
1960:
1866:
1767:
1749:
1576:
1482:
1444:
480:
in yeast to many kilobases in humans, and usually is composed of arrays of
41:
3996:
3983:
3590:
3555:
3329:
3140:
3089:
2858:
2319:
2250:
1925:
1718:
1608:
1561:
1506:
1295:, and even within smaller taxonomic groups these patterns appear diverse.
5888:
5652:
5548:
5448:
4510:
4134:
3737:
2458:
1882:"Conservation of the human telomere sequence (TTAGGG)n among vertebrates"
1654:
832:
730:
511:
347:
342:
325:
4158:
3582:
2915:
2594:"Telomere length, oxidative damage, antioxidants and breast cancer risk"
1428:
249:
for the discovery of how chromosomes are protected by telomeres and the
5898:
5388:
5281:
5059:
5054:
4896:
4750:
4661:
4656:
4069:
3939:
von
Zglinicki T (March 2012). "Will your telomeres tell your future?".
2492:"The impact of oxidative DNA damage and stress on telomere homeostasis"
1292:
1251:
Tools have also been developed to estimate the length of telomere from
1150:
if you can. Unsourced or poorly sourced material may be challenged and
783:
622:
519:
481:
476:
Telomere length varies greatly between species, from approximately 300
410:
398:
394:
4273:
4256:
4224:
4207:
4085:
4017:
Philosophical
Transactions of the Royal Society B: Biological Sciences
4013:"Heritability of telomere variation: it is all about the environment!"
3952:
3428:
3411:
3387:
3370:
3313:
2609:
1413:"2009 Nobel Prize in Physiology or Medicine: telomeres and telomerase"
646:
522:. There is a multitude of ways in which oxidative stress, mediated by
5997:
5433:
5025:
4806:
4693:
4457:
4453:
4448:
4255:
Chatelain, Marion; Drobniak, Szymon M.; Szulkin, Marta (2019-11-27).
2311:
918:
892:
707:
465:
250:
157:). Telomeres are a widespread genetic feature most commonly found in
149:
sequences associated with specialized proteins at the ends of linear
138:
121:
358:
5743:
5335:
4871:
4856:
4851:
4846:
4841:
4836:
4831:
4826:
4821:
4796:
4734:
2143:"Senescence and immortalization: role of telomeres and telomerase"
995:
977:
862:
686:
596:
The phenomenon of limited cellular division was first observed by
583:
424:
382:
378:
374:
40:
2277:, Nature, June 30, 2022; Nat Struct Mol Biol 29, 639–652 (2022).
5049:
4924:
386:
5521:
4928:
4396:
5039:
5035:
3164:"Measuring vertebrate telomeres: applications and limitations"
2751:"Telomerase as a Therapeutic Target in Cardiovascular Disease"
1115:
298:
162:
93:
81:
5475:
List of largest biomedical companies by market capitalization
4314:"Ectothermic telomeres: it's time they came in from the cold"
169:
systems from mistaking the very ends of the DNA strand for a
3722:"Estimating telomere length from whole genome sequence data"
514:
studies have shown that telomeres accumulate damage due to
99:
75:
69:
2441:
Burge S, Parkinson GN, Hazel P, Todd AK, Neidle S (2006).
1390:
During replication, multiple DNA-polymerases are involved.
2700:"Telomere dysfunction in ageing and age-related diseases"
1732:
Chow TT, Zhao Y, Mak SS, Shay JW, Wright WE (June 2012).
413:(that possess telomere repeats similar to those found in
362:
Shelterin co-ordinates the T-loop formation of telomeres.
1577:"Telomeres, telomerase, and aging: origin of the theory"
397:. In many species, the sequence repeats are enriched in
27:
Region of repetitive nucleotide sequences on chromosomes
4914:
International System for Human Cytogenetic Nomenclature
4375:
Telomerase and the Consequences of Telomere Dysfunction
4011:
Dugdale, Hannah L.; Richardson, David S. (2018-01-15).
1147:
4206:
Pepke, Michael Le; Eisenberg, Dan T. A. (2021-03-16).
2698:
Rossiello F, Jurk D, Passos JF, di Fagagna F (2022).
2076:"Telomere shortening rate predicts species life span"
1244:), software processing the TRF pictures. A Real-Time
2843:"Telomeres and the natural lifespan limit in humans"
2184:"Telomeres, telomerase, and tumorigenesis--a review"
557:, longer telomeres may be associated with lifespan.
96:
90:
72:
66:
30:
For the use of "telomere" in insect morphology, see
5968:
5921:
5789:
5757:
5734:
5711:
5702:
5693:
5668:
5628:
5585:
5576:
5457:
5364:
5244:
5174:
5123:
5114:
5073:
5018:
4997:
4962:
4884:
4787:
4749:
4719:
4647:
4503:
4434:
1976:"Telomerase Repeated Amplification Protocol (TRAP)"
87:
78:
63:
3162:Nakagawa S, Gemmell NJ, Burke T (September 2004).
2756:Arteriosclerosis, Thrombosis, and Vascular Biology
2749:Hoffmann J, Richardson G, Spyridopoulos I (2021).
2443:"Quadruplex DNA: sequence, topology and structure"
1782:
4095:20.500.11820/91f3fc9e-4a69-4ac4-a8a0-45c93ccbf3b5
2639:
2637:
2271:Barnes, R.P., de Rosa, M., Thosar, S.A., et al.,
1880:Meyne J, Ratliff RL, Moyzis RK (September 1989).
2394:"Mammalian telomeres end in a large duplex loop"
2693:
2691:
2689:
2080:Proceedings of the National Academy of Sciences
649:. Some of the experimentally verified telomere
4365:Telomeres and Telomerase: The Means to the End
2890:Pepper GV, Bateson M, Nettle D (August 2018).
1146:Please review the contents of the section and
287:cannot replicate the sequences present at the
5533:
4940:
4408:
4312:Olsson M, Wapstra E, Friesen C (March 2018).
8:
536:Relationship between telomeres and longevity
317:degraded in the process of DNA replication.
676:Telomeric repeat (5' to 3' toward the end)
5708:
5699:
5582:
5540:
5526:
5518:
5120:
4947:
4933:
4925:
4746:
4415:
4401:
4393:
3673:"Telomere measurement by quantitative PCR"
2279:https://doi.org/10.1038/s41594-022-00790-y
429:Synthesis of chromosome ends by telomerase
4337:
4272:
4223:
4174:
4093:
4044:
3995:
3853:
3843:
3802:
3745:
3696:
3647:
3629:
3545:
3496:
3427:
3386:
3345:
3245:
2923:
2866:
2817:
2768:
2725:
2669:
2617:
2568:
2541:"Does oxidative stress shorten telomeres
2515:
2490:Barnes R, Fouquerel E, Opresko P (2019).
2466:
2409:
2368:
2358:
2240:
2199:
2158:
2117:
2099:
1999:
1915:
1905:
1856:
1757:
1229:senescence-associated secretory phenotype
665:Some known telomere nucleotide sequences
4980:Competitions and prizes in biotechnology
3922:"A Blood Test Offers Clues to Longevity"
2961:10.1146/annurev-publhealth-040119-094239
1789:(5th ed.). New York: W.H. Freeman.
1781:Nelson DL, Lehninger AL, Cox MM (2008).
1411:Varela, E.; Blasco, M. A. (March 2010).
663:
561:Potential effect of psychological stress
510:Apart from the end replication problem,
357:
274:
145: 'part') is a region of repetitive
1841:"Role of shelterin in cancer and aging"
1403:
1383:
3520:Allshire RC, et al. (June 1989).
2841:Steenstrup T, Kark JD, Aviv A (2017).
2225:"Telomeres, telomerase and senescence"
1839:Martínez P, Blasco MA (October 2010).
226:, working as a postdoctoral fellow at
4904:List of organisms by chromosome count
3457:
3455:
2539:Reichert S, Stier A (December 2017).
279:Lagging strand during DNA replication
247:Nobel Prize in Physiology or Medicine
7:
5500:
3283:10.1016/j.neurobiolaging.2010.11.013
2496:Mechanisms of Ageing and Development
1785:Lehninger Principles of Biochemistry
1628:
1626:
206:In the early 1970s, Soviet theorist
2347:The Journal of Biological Chemistry
1974:Mender I, Shay JW (November 2015).
5609:Short tandem repeat/Microsatellite
3984:"Spit test offers guide to health"
49:(grey) capped by telomeres (white)
25:
1007:TGTGGGTGTGGTG (from RNA template)
5499:
5488:
5487:
4988:
3920:Pollack, Andrew (May 18, 2011).
3238:10.1111/j.1474-9726.2011.00718.x
3191:10.1111/j.1365-294X.2004.02291.x
1858:10.1111/j.1474-9726.2010.00596.x
1820:"Bacterial Chromosome Structure"
1305:
1212:
1120:
1009:or G(2-3)(TG)(1-6)T (consensus)
600:, and is now referred to as the
405:, which allows the formation of
59:
5465:Index of biotechnology articles
4525:Macrochromosome/Microchromosome
2598:International Journal of Cancer
2141:Shay JW, Wright WE (May 2005).
1520:Olovnikov, A. M. (1973-09-14).
1470:. Woods Hole. pp. 181–198.
568:found that increased perceived
486:3′ single-stranded-DNA overhang
5613:Trinucleotide repeat disorders
5470:List of biotechnology articles
2949:Annual Review of Public Health
1691:Journal of Theoretical Biology
1671:. Nobel Foundation. 2009-10-05
1526:Journal of Theoretical Biology
1372:Immortal DNA strand hypothesis
1148:add the appropriate references
328:chromosomes (such as those in
154:
1:
5600:Variable number tandem repeat
5314:Genetically modified organism
5081:Biotechnology industrial park
2650:Brain, Behavior, and Immunity
2411:10.1016/S0092-8674(00)80760-6
661:for letter representations).
653:sequences are also listed in
458:somatic cell nuclear transfer
4384:DNA Ends: Just the Beginning
3133:10.1126/science.279.5349.349
3039:10.1016/0014-4827(65)90211-9
3004:10.1016/0014-4827(61)90192-6
2798:Molecular Human Reproduction
2045:10.1126/science.288.5466.665
1711:10.1016/0022-5193(73)90198-7
1647:10.1016/0022-2836(78)90294-2
1635:Journal of Molecular Biology
1593:10.1016/0531-5565(96)00005-8
1546:10.1016/0022-5193(73)90198-7
32:Telomere (insect morphology)
5304:Environmental biotechnology
1468:The Remaking of Chromosomes
1133:reliable medical references
354:Telomere ends and shelterin
6065:
4565:Dinoflagellate chromosomes
4139:Royal Society Open Science
3795:10.1038/s41598-017-14403-y
3631:10.1186/s12859-021-04064-0
3027:Experimental Cell Research
2992:Experimental Cell Research
2896:Royal Society Open Science
2770:10.1161/ATVBAHA.120.315695
2718:10.1038/s41556-022-00842-x
2223:Greider CW (August 1990).
1487:Doklady Akademii Nauk SSSR
1331:DNA damage theory of aging
1105:GGTGTACGGATTTGATTAGGTATGT
1093:GGTGTACGGATTTGATTAGTTATGT
621:Two studies on long-lived
533:
432:
365:
303:lagging strand replication
268:
131:
114:
29:
5567:
5483:
4986:
4975:Timeline of biotechnology
4909:List of sequenced genomes
4677:Chromosomal translocation
4550:A chromosome/B chromosome
4541:(or accessory chromosome)
3845:10.1186/s12859-019-2851-0
2662:10.1016/j.bbi.2016.02.002
2508:10.1016/j.mad.2018.03.013
1953:10.1016/j.tcb.2009.05.002
1818:Maloy S (July 12, 2002).
1575:Olovnikov, A. M. (1996).
1481:Olovnikov, A. M. (1971).
1139:or relies too heavily on
993:
984:Schizosaccharomyces pombe
917:
860:
782:
723:
491:telomere-binding proteins
464:and in the prevention of
185:, studying the fruit fly
36:Telomere (disambiguation)
6039:Repetitive DNA sequences
5331:Microbial biodegradation
5010:Industrial biotechnology
4970:History of biotechnology
4731:Telomere-binding protein
4545:Supernumerary chromosome
1581:Experimental Gerontology
1362:Telomere-binding protein
1112:Research on disease risk
1088:Candida pseudotropicalis
1069:GGTGTACGGATGCAGACTCGCTT
1045:GGTGTACGGATGTCTAACTTCTT
1002:Saccharomyces cerevisiae
283:During DNA replication,
128: 'end' and
5005:Colors of biotechnology
3671:Cawthon RM (May 2002).
3082:10.1126/science.7544491
2360:10.1074/jbc.M110.170167
2101:10.1073/pnas.1902452116
1992:10.21769/bioprotoc.1657
1907:10.1073/pnas.86.18.7049
1738:Genes & Development
1253:whole genome sequencing
1057:GGTGTAGGATGTCACGATCATT
1016:Saccharomyces castellii
524:reactive oxygen species
265:End replication problem
188:Drosophila melanogaster
6008:Protein tandem repeats
5936:Tandemly arrayed genes
5399:Biomedical engineering
5106:Pharmaceutical company
5086:Biotechnology products
4667:Structural alterations
4330:10.1098/rstb.2016.0449
4029:10.1098/rstb.2016.0450
3891:10.1038/nprot.2006.263
3726:Nucleic Acids Research
3677:Nucleic Acids Research
3538:10.1093/nar/17.12.4611
3526:Nucleic Acids Research
3469:Nucleic Acids Research
3318:Nucleic Acids Research
2561:10.1098/rsbl.2017.0463
2447:Nucleic Acids Research
2242:10.1002/bies.950120803
1941:Trends in Cell Biology
1750:10.1101/gad.187211.112
593:
530:Association with aging
430:
363:
280:
260:Structure and function
50:
34:. For other uses, see
4684:Numerical alterations
4672:Chromosomal inversion
4570:Homologous chromosome
3997:10.1038/news.2011.330
3689:10.1093/nar/30.10.e47
3271:Neurobiology of Aging
2859:10.18632/aging.101216
2810:10.1093/molehr/gas028
2160:10.1093/carcin/bgh296
1466:Muller, H.J. (1938).
1076:Candida guillermondii
659:nucleic acid notation
631:Oceanodroma leucorhoa
587:
428:
361:
278:
183:Hermann Joseph Muller
44:
5981:Pathogenicity island
5424:Chemical engineering
5287:Reproductive cloning
5151:Hybridoma technology
5101:Human Genome Project
4892:Extrachromosomal DNA
4580:Satellite chromosome
4555:Lampbrush chromosome
4495:Nuclear organization
3826:Feuerbach L (2019).
3571:Nature Biotechnology
3481:10.1093/nar/gkac1202
2182:Wai LK (July 2004).
1346:Rejuvenation (aging)
1100:Kluyveromyces lactis
966:Ascaris lumbricoides
869:Arabidopsis thaliana
627:Leach's storm-petrel
570:psychological stress
450:embryonic stem cells
5309:Genetic engineering
5292:Therapeutic cloning
5272:Bionic architecture
5252:Animal cell culture
5019:Biological concepts
4585:Centromere position
4560:Polytene chromosome
4530:Circular chromosome
4369:Elizabeth Blackburn
4159:10.1098/rsos.200548
4151:2020RSOS....700548R
3982:Marchant J (2011).
3787:2018NatSR...8.1300F
3583:10.1038/nbt0898-743
3330:10.1093/nar/gkad672
3183:2004MolEc..13.2523N
3125:1998Sci...279..349B
3074:1995Sci...269.1236F
2916:10.1098/rsos.180744
2908:2018RSOS....580744P
2705:Nature Cell Biology
2304:1984Natur.310..154S
2092:2019PNAS..11615122W
2086:(30): 15122–15127.
2037:2000Sci...288..665L
1898:1989PNAS...86.7049M
1703:1973JThBi..41..181O
1538:1973JThBi..41..181O
1429:10.1038/onc.2010.15
1264:Elizabeth Blackburn
666:
655:Telomerase Database
608:by Geron's founder
547:cellular senescence
224:Elizabeth Blackburn
214:'s idea of limited
171:double-strand break
18:Telomere shortening
5931:Gene amplification
5365:Interdisciplinary
5319:Molecular genetics
5065:Selective breeding
4324:(1741): 20160449.
4023:(1741): 20160450.
3926:The New York Times
3832:BMC Bioinformatics
3775:Scientific Reports
3769:Farmery J (2018).
3738:10.1093/nar/gku181
3618:BMC Bioinformatics
2459:10.1093/nar/gkl655
1353:, biological aging
1052:Candida tropicalis
664:
594:
431:
364:
281:
193:Barbara McClintock
51:
6034:Molecular biology
6016:
6015:
5917:
5916:
5785:
5784:
5689:
5688:
5578:Repeated sequence
5553:repeated sequence
5515:
5514:
5444:Nanobiotechnology
5439:Molecular biology
5240:
5239:
5091:Biotechnology law
4922:
4921:
4880:
4879:
4617:Centromere number
4534:Linear chromosome
4386:Nobel Lecture by
4377:Nobel Lecture by
4367:Nobel Lecture by
4274:10.1111/ele.13426
4225:10.1111/mec.15870
4218:(23): 6286–6296.
4212:Molecular Ecology
4086:10.1111/mec.16145
4080:(23): 5917–5932.
4074:Molecular Ecology
3953:10.1136/bmj.e1727
3429:10.1111/tpj.13115
3416:The Plant Journal
3388:10.1111/tpj.12839
3375:The Plant Journal
3324:(D1): D311–D321.
3171:Molecular Ecology
3068:(5228): 1236–41.
2610:10.1002/ijc.24105
1744:(11): 1167–1178.
1423:(11): 1561–1565.
1341:Maximum life span
1221:
1220:
1197:
1109:
1108:
989:TTAC(A)(C)G(1-8)
937:Bombus terrestris
714:Neurospora crassa
606:Geron Corporation
495:displacement loop
454:white blood cells
401:, e.g. TTAGGG in
242:were awarded the
191:, and in 1939 by
16:(Redirected from
6056:
5993:Low copy repeats
5986:Symbiosis island
5923:Gene duplication
5709:
5700:
5583:
5561:gene duplication
5542:
5535:
5528:
5519:
5503:
5502:
5491:
5490:
5341:Pharmacogenomics
5121:
5115:Basic techniques
5096:Green Revolution
5074:General concepts
4992:
4949:
4942:
4935:
4926:
4747:
4711:Polyploidization
4539:Extra chromosome
4454:Genetic material
4417:
4410:
4403:
4394:
4352:
4351:
4341:
4309:
4303:
4302:
4276:
4252:
4246:
4245:
4227:
4203:
4197:
4196:
4178:
4130:
4124:
4123:
4097:
4065:
4059:
4058:
4048:
4008:
4002:
4001:
3999:
3979:
3973:
3972:
3936:
3930:
3929:
3917:
3911:
3910:
3879:Nature Protocols
3874:
3868:
3867:
3857:
3847:
3823:
3817:
3816:
3806:
3766:
3760:
3759:
3749:
3717:
3711:
3710:
3700:
3668:
3662:
3661:
3651:
3633:
3609:
3603:
3602:
3566:
3560:
3559:
3549:
3517:
3511:
3510:
3500:
3459:
3450:
3449:
3431:
3407:
3401:
3400:
3390:
3366:
3360:
3359:
3349:
3309:
3303:
3302:
3277:(7): 1486.e3–8.
3266:
3260:
3259:
3249:
3217:
3211:
3210:
3168:
3159:
3153:
3152:
3119:(5349): 349–52.
3108:
3102:
3101:
3057:
3051:
3050:
3022:
3016:
3015:
2987:
2981:
2980:
2944:
2938:
2937:
2927:
2887:
2881:
2880:
2870:
2838:
2832:
2831:
2821:
2789:
2783:
2782:
2772:
2763:(3): 1047–1061.
2746:
2740:
2739:
2729:
2695:
2684:
2683:
2673:
2641:
2632:
2631:
2621:
2589:
2583:
2582:
2572:
2555:(12): 20170463.
2536:
2530:
2529:
2519:
2487:
2481:
2480:
2470:
2438:
2432:
2431:
2413:
2389:
2383:
2382:
2372:
2362:
2353:(46): 35814–24.
2338:
2332:
2331:
2312:10.1038/310154a0
2287:
2281:
2269:
2263:
2262:
2244:
2220:
2214:
2213:
2203:
2179:
2173:
2172:
2162:
2138:
2132:
2131:
2121:
2103:
2071:
2065:
2064:
2020:
2014:
2013:
2003:
1971:
1965:
1964:
1936:
1930:
1929:
1919:
1909:
1877:
1871:
1870:
1860:
1836:
1830:
1829:
1827:
1826:
1815:
1809:
1808:
1788:
1778:
1772:
1771:
1761:
1729:
1723:
1722:
1686:
1680:
1679:
1677:
1676:
1665:
1659:
1658:
1630:
1621:
1620:
1572:
1566:
1565:
1517:
1511:
1510:
1493:(6): 1496–1499.
1478:
1472:
1471:
1463:
1457:
1456:
1408:
1391:
1388:
1321:Epigenetic clock
1315:
1310:
1309:
1216:
1215:
1207:
1204:
1198:
1196:
1155:
1124:
1123:
1116:
1040:Candida albicans
1033:GGGGTCTGGGTGCTG
1028:Candida glabrata
949:Vespula vulgaris
667:
598:Leonard Hayflick
574:publication bias
516:oxidative stress
506:Oxidative damage
444:, some types of
212:Leonard Hayflick
208:Alexey Olovnikov
142:
135:
125:
118:
106:
105:
102:
101:
98:
95:
92:
89:
84:
83:
80:
77:
74:
71:
68:
65:
21:
6064:
6063:
6059:
6058:
6057:
6055:
6054:
6053:
6019:
6018:
6017:
6012:
5964:
5913:
5781:
5753:
5730:
5704:Retrotransposon
5685:
5676:Inverted repeat
5664:
5649:DNA transposon
5645:Retrotransposon
5640:Gene conversion
5631:
5624:
5621:
5572:
5563:
5546:
5516:
5511:
5479:
5453:
5394:Biopharmacology
5366:
5360:
5236:
5207:Electrophoresis
5202:Kidney dialysis
5192:Crystallization
5170:
5116:
5110:
5069:
5014:
4993:
4984:
4958:
4953:
4923:
4918:
4876:
4783:
4745:
4715:
4704:Paleopolyploidy
4649:
4643:
4499:
4473:Heterochromatin
4436:
4430:
4421:
4361:
4356:
4355:
4311:
4310:
4306:
4261:Ecology Letters
4254:
4253:
4249:
4205:
4204:
4200:
4132:
4131:
4127:
4067:
4066:
4062:
4010:
4009:
4005:
3981:
3980:
3976:
3938:
3937:
3933:
3919:
3918:
3914:
3876:
3875:
3871:
3825:
3824:
3820:
3768:
3767:
3763:
3720:Ding Z (2014).
3719:
3718:
3714:
3670:
3669:
3665:
3611:
3610:
3606:
3568:
3567:
3563:
3532:(12): 4611–27.
3519:
3518:
3514:
3461:
3460:
3453:
3409:
3408:
3404:
3368:
3367:
3363:
3311:
3310:
3306:
3268:
3267:
3263:
3219:
3218:
3214:
3166:
3161:
3160:
3156:
3110:
3109:
3105:
3059:
3058:
3054:
3024:
3023:
3019:
2989:
2988:
2984:
2946:
2945:
2941:
2889:
2888:
2884:
2853:(4): 130–1142.
2840:
2839:
2835:
2791:
2790:
2786:
2748:
2747:
2743:
2697:
2696:
2687:
2643:
2642:
2635:
2591:
2590:
2586:
2549:Biology Letters
2538:
2537:
2533:
2489:
2488:
2484:
2453:(19): 5402–15.
2440:
2439:
2435:
2391:
2390:
2386:
2340:
2339:
2335:
2298:(5973): 154–7.
2289:
2288:
2284:
2270:
2266:
2222:
2221:
2217:
2181:
2180:
2176:
2140:
2139:
2135:
2073:
2072:
2068:
2031:(5466): 665–9.
2022:
2021:
2017:
1973:
1972:
1968:
1938:
1937:
1933:
1892:(18): 7049–53.
1879:
1878:
1874:
1838:
1837:
1833:
1824:
1822:
1817:
1816:
1812:
1797:
1780:
1779:
1775:
1731:
1730:
1726:
1688:
1687:
1683:
1674:
1672:
1667:
1666:
1662:
1632:
1631:
1624:
1574:
1573:
1569:
1519:
1518:
1514:
1480:
1479:
1475:
1465:
1464:
1460:
1410:
1409:
1405:
1400:
1395:
1394:
1389:
1385:
1380:
1311:
1304:
1301:
1272:
1237:
1225:senescent cells
1217:
1213:
1208:
1202:
1199:
1156:
1145:
1141:primary sources
1125:
1121:
1114:
1064:Candida maltosa
1008:
881:Cestrum elegans
643:
610:Michael D. West
582:
563:
555:life expectancy
538:
532:
508:
503:
474:
437:
423:
370:
356:
273:
271:DNA replication
267:
262:
228:Yale University
179:
163:chromosomal DNA
86:
62:
58:
39:
28:
23:
22:
15:
12:
11:
5:
6062:
6060:
6052:
6051:
6049:Non-coding DNA
6046:
6041:
6036:
6031:
6021:
6020:
6014:
6013:
6011:
6010:
6005:
6000:
5995:
5990:
5989:
5988:
5983:
5976:Genomic island
5972:
5970:
5966:
5965:
5963:
5962:
5957:
5956:
5955:
5945:
5944:
5943:
5933:
5927:
5925:
5919:
5918:
5915:
5914:
5912:
5911:
5906:
5901:
5896:
5891:
5886:
5881:
5876:
5871:
5866:
5861:
5856:
5851:
5846:
5841:
5836:
5831:
5826:
5821:
5816:
5811:
5806:
5801:
5795:
5793:
5791:DNA transposon
5787:
5786:
5783:
5782:
5780:
5779:
5774:
5769:
5763:
5761:
5755:
5754:
5752:
5751:
5746:
5740:
5738:
5732:
5731:
5729:
5728:
5723:
5717:
5715:
5706:
5697:
5691:
5690:
5687:
5686:
5684:
5683:
5678:
5672:
5670:
5666:
5665:
5663:
5662:
5661:
5660:
5655:
5647:
5642:
5636:
5634:
5626:
5625:
5623:
5622:
5619:Macrosatellite
5616:
5606:
5597:
5591:
5589:
5587:Tandem repeats
5580:
5574:
5573:
5568:
5565:
5564:
5547:
5545:
5544:
5537:
5530:
5522:
5513:
5512:
5510:
5509:
5497:
5484:
5481:
5480:
5478:
5477:
5472:
5467:
5461:
5459:
5455:
5454:
5452:
5451:
5446:
5441:
5436:
5431:
5426:
5421:
5416:
5411:
5406:
5401:
5396:
5391:
5386:
5384:Bioengineering
5381:
5379:Bioelectronics
5376:
5370:
5368:
5362:
5361:
5359:
5358:
5356:Tissue culture
5353:
5348:
5343:
5338:
5333:
5328:
5323:
5322:
5321:
5316:
5306:
5301:
5296:
5295:
5294:
5289:
5279:
5274:
5269:
5264:
5262:Bioinformatics
5259:
5257:Biofabrication
5254:
5248:
5246:
5242:
5241:
5238:
5237:
5235:
5234:
5229:
5224:
5219:
5214:
5209:
5204:
5199:
5197:Chromatography
5194:
5189:
5184:
5182:Centrifugation
5178:
5176:
5175:Chemical field
5172:
5171:
5169:
5168:
5163:
5158:
5153:
5148:
5146:Flow cytometry
5143:
5138:
5133:
5127:
5125:
5118:
5112:
5111:
5109:
5108:
5103:
5098:
5093:
5088:
5083:
5077:
5075:
5071:
5070:
5068:
5067:
5062:
5057:
5052:
5047:
5042:
5033:
5028:
5022:
5020:
5016:
5015:
5013:
5012:
5007:
5001:
4999:
4995:
4994:
4987:
4985:
4983:
4982:
4977:
4972:
4966:
4964:
4960:
4959:
4954:
4952:
4951:
4944:
4937:
4929:
4920:
4919:
4917:
4916:
4911:
4906:
4901:
4900:
4899:
4888:
4886:
4882:
4881:
4878:
4877:
4875:
4874:
4869:
4864:
4859:
4854:
4849:
4844:
4839:
4834:
4829:
4824:
4819:
4814:
4809:
4804:
4799:
4793:
4791:
4785:
4784:
4782:
4781:
4776:
4771:
4766:
4761:
4755:
4753:
4744:
4743:
4738:
4723:
4721:
4717:
4716:
4714:
4713:
4708:
4707:
4706:
4701:
4696:
4691:
4681:
4680:
4679:
4674:
4664:
4659:
4653:
4651:
4645:
4644:
4642:
4641:
4640:
4639:
4634:
4629:
4624:
4614:
4613:
4612:
4607:
4602:
4597:
4595:Submetacentric
4592:
4582:
4577:
4572:
4567:
4562:
4557:
4552:
4547:
4542:
4536:
4527:
4522:
4521:or heterosome)
4515:Sex chromosome
4507:
4505:
4501:
4500:
4498:
4497:
4492:
4487:
4482:
4477:
4476:
4475:
4470:
4460:
4451:
4446:
4440:
4438:
4432:
4431:
4422:
4420:
4419:
4412:
4405:
4397:
4391:
4390:
4381:
4372:
4360:
4359:External links
4357:
4354:
4353:
4304:
4267:(2): 381–398.
4247:
4198:
4145:(11): 200548.
4125:
4060:
4003:
3974:
3931:
3912:
3885:(5): 2365–76.
3869:
3818:
3761:
3712:
3683:(10): 47e–47.
3663:
3604:
3561:
3512:
3475:(1): 420–433.
3451:
3402:
3361:
3304:
3261:
3212:
3177:(9): 2523–33.
3154:
3103:
3052:
3017:
2998:(3): 585–621.
2982:
2939:
2882:
2833:
2804:(11): 517–22.
2784:
2741:
2712:(2): 135–147.
2685:
2633:
2604:(7): 1637–43.
2584:
2531:
2482:
2433:
2384:
2333:
2282:
2264:
2215:
2174:
2147:Carcinogenesis
2133:
2066:
2015:
1966:
1931:
1872:
1831:
1810:
1795:
1773:
1724:
1681:
1660:
1622:
1587:(4): 443–448.
1567:
1532:(1): 181–190.
1512:
1473:
1458:
1402:
1401:
1399:
1396:
1393:
1392:
1382:
1381:
1379:
1376:
1375:
1374:
1369:
1364:
1359:
1354:
1348:
1343:
1338:
1333:
1328:
1323:
1317:
1316:
1313:Biology portal
1300:
1297:
1271:
1268:
1236:
1233:
1219:
1218:
1211:
1209:
1128:
1126:
1119:
1113:
1110:
1107:
1106:
1103:
1095:
1094:
1091:
1083:
1082:
1079:
1071:
1070:
1067:
1059:
1058:
1055:
1047:
1046:
1043:
1035:
1034:
1031:
1023:
1022:
1019:
1011:
1010:
1005:
998:
991:
990:
987:
980:
973:
972:
969:
962:
956:
955:
952:
944:
943:
940:
932:
931:
928:
921:
915:
914:
911:
900:
899:
896:
888:
887:
884:
876:
875:
872:
865:
858:
857:
854:
847:
840:
839:
836:
816:
815:
812:
804:
803:
800:
787:
780:
779:
776:
763:
756:
755:
752:
744:
743:
740:
727:
721:
720:
717:
710:
703:
702:
699:
684:
678:
677:
674:
671:
642:
639:
602:Hayflick limit
590:Hayflick limit
581:
578:
562:
559:
534:Main article:
531:
528:
507:
504:
502:
499:
473:
470:
452:, and certain
433:Main article:
422:
419:
407:G-quadruplexes
366:Main article:
355:
352:
285:DNA polymerase
269:Main article:
266:
263:
261:
258:
232:Joseph G. Gall
222:In 1975–1977,
178:
175:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
6061:
6050:
6047:
6045:
6042:
6040:
6037:
6035:
6032:
6030:
6027:
6026:
6024:
6009:
6006:
6004:
6001:
5999:
5996:
5994:
5991:
5987:
5984:
5982:
5979:
5978:
5977:
5974:
5973:
5971:
5967:
5961:
5958:
5954:
5951:
5950:
5949:
5946:
5942:
5941:Ribosomal DNA
5939:
5938:
5937:
5934:
5932:
5929:
5928:
5926:
5924:
5920:
5910:
5907:
5905:
5902:
5900:
5897:
5895:
5892:
5890:
5887:
5885:
5882:
5880:
5877:
5875:
5872:
5870:
5867:
5865:
5862:
5860:
5857:
5855:
5852:
5850:
5847:
5845:
5842:
5840:
5837:
5835:
5832:
5830:
5827:
5825:
5822:
5820:
5817:
5815:
5812:
5810:
5807:
5805:
5802:
5800:
5797:
5796:
5794:
5792:
5788:
5778:
5775:
5773:
5770:
5768:
5765:
5764:
5762:
5760:
5756:
5750:
5747:
5745:
5742:
5741:
5739:
5737:
5733:
5727:
5724:
5722:
5719:
5718:
5716:
5714:
5710:
5707:
5705:
5701:
5698:
5696:
5692:
5682:
5681:Direct repeat
5679:
5677:
5674:
5673:
5671:
5667:
5659:
5656:
5654:
5651:
5650:
5648:
5646:
5643:
5641:
5638:
5637:
5635:
5633:
5627:
5620:
5617:
5614:
5610:
5607:
5605:
5604:Minisatellite
5601:
5598:
5596:
5595:Satellite DNA
5593:
5592:
5590:
5588:
5584:
5581:
5579:
5575:
5571:
5566:
5562:
5558:
5554:
5550:
5543:
5538:
5536:
5531:
5529:
5524:
5523:
5520:
5508:
5507:
5498:
5496:
5495:
5486:
5485:
5482:
5476:
5473:
5471:
5468:
5466:
5463:
5462:
5460:
5456:
5450:
5447:
5445:
5442:
5440:
5437:
5435:
5432:
5430:
5427:
5425:
5422:
5420:
5417:
5415:
5412:
5410:
5407:
5405:
5402:
5400:
5397:
5395:
5392:
5390:
5387:
5385:
5382:
5380:
5377:
5375:
5372:
5371:
5369:
5363:
5357:
5354:
5352:
5349:
5347:
5344:
5342:
5339:
5337:
5334:
5332:
5329:
5327:
5324:
5320:
5317:
5315:
5312:
5311:
5310:
5307:
5305:
5302:
5300:
5297:
5293:
5290:
5288:
5285:
5284:
5283:
5280:
5278:
5277:Cell immunity
5275:
5273:
5270:
5268:
5265:
5263:
5260:
5258:
5255:
5253:
5250:
5249:
5247:
5243:
5233:
5232:Sedimentation
5230:
5228:
5225:
5223:
5220:
5218:
5215:
5213:
5210:
5208:
5205:
5203:
5200:
5198:
5195:
5193:
5190:
5188:
5185:
5183:
5180:
5179:
5177:
5173:
5167:
5164:
5162:
5159:
5157:
5154:
5152:
5149:
5147:
5144:
5142:
5141:Cultured meat
5139:
5137:
5134:
5132:
5129:
5128:
5126:
5124:Biology field
5122:
5119:
5113:
5107:
5104:
5102:
5099:
5097:
5094:
5092:
5089:
5087:
5084:
5082:
5079:
5078:
5076:
5072:
5066:
5063:
5061:
5058:
5056:
5053:
5051:
5048:
5046:
5043:
5041:
5037:
5034:
5032:
5029:
5027:
5024:
5023:
5021:
5017:
5011:
5008:
5006:
5003:
5002:
5000:
4996:
4991:
4981:
4978:
4976:
4973:
4971:
4968:
4967:
4965:
4961:
4957:
4956:Biotechnology
4950:
4945:
4943:
4938:
4936:
4931:
4930:
4927:
4915:
4912:
4910:
4907:
4905:
4902:
4898:
4895:
4894:
4893:
4890:
4889:
4887:
4883:
4873:
4870:
4868:
4865:
4863:
4860:
4858:
4855:
4853:
4850:
4848:
4845:
4843:
4840:
4838:
4835:
4833:
4830:
4828:
4825:
4823:
4820:
4818:
4815:
4813:
4810:
4808:
4805:
4803:
4800:
4798:
4795:
4794:
4792:
4790:
4786:
4780:
4777:
4775:
4772:
4770:
4767:
4765:
4762:
4760:
4757:
4756:
4754:
4752:
4748:
4742:
4739:
4736:
4732:
4728:
4725:
4724:
4722:
4718:
4712:
4709:
4705:
4702:
4700:
4697:
4695:
4692:
4690:
4687:
4686:
4685:
4682:
4678:
4675:
4673:
4670:
4669:
4668:
4665:
4663:
4660:
4658:
4655:
4654:
4652:
4650:and evolution
4646:
4638:
4635:
4633:
4630:
4628:
4625:
4623:
4620:
4619:
4618:
4615:
4611:
4608:
4606:
4603:
4601:
4598:
4596:
4593:
4591:
4588:
4587:
4586:
4583:
4581:
4578:
4576:
4575:Isochromosome
4573:
4571:
4568:
4566:
4563:
4561:
4558:
4556:
4553:
4551:
4548:
4546:
4543:
4540:
4537:
4535:
4531:
4528:
4526:
4523:
4520:
4516:
4512:
4509:
4508:
4506:
4502:
4496:
4493:
4491:
4488:
4486:
4483:
4481:
4478:
4474:
4471:
4469:
4466:
4465:
4464:
4461:
4459:
4455:
4452:
4450:
4447:
4445:
4442:
4441:
4439:
4433:
4429:
4425:
4418:
4413:
4411:
4406:
4404:
4399:
4398:
4395:
4389:
4385:
4382:
4380:
4379:Carol Greider
4376:
4373:
4370:
4366:
4363:
4362:
4358:
4349:
4345:
4340:
4335:
4331:
4327:
4323:
4319:
4315:
4308:
4305:
4300:
4296:
4292:
4288:
4284:
4280:
4275:
4270:
4266:
4262:
4258:
4251:
4248:
4243:
4239:
4235:
4231:
4226:
4221:
4217:
4213:
4209:
4202:
4199:
4194:
4190:
4186:
4182:
4177:
4172:
4168:
4164:
4160:
4156:
4152:
4148:
4144:
4140:
4136:
4129:
4126:
4121:
4117:
4113:
4109:
4105:
4101:
4096:
4091:
4087:
4083:
4079:
4075:
4071:
4064:
4061:
4056:
4052:
4047:
4042:
4038:
4034:
4030:
4026:
4022:
4018:
4014:
4007:
4004:
3998:
3993:
3989:
3985:
3978:
3975:
3970:
3966:
3962:
3958:
3954:
3950:
3946:
3942:
3935:
3932:
3927:
3923:
3916:
3913:
3908:
3904:
3900:
3896:
3892:
3888:
3884:
3880:
3873:
3870:
3865:
3861:
3856:
3851:
3846:
3841:
3837:
3833:
3829:
3822:
3819:
3814:
3810:
3805:
3800:
3796:
3792:
3788:
3784:
3780:
3776:
3772:
3765:
3762:
3757:
3753:
3748:
3743:
3739:
3735:
3731:
3727:
3723:
3716:
3713:
3708:
3704:
3699:
3694:
3690:
3686:
3682:
3678:
3674:
3667:
3664:
3659:
3655:
3650:
3645:
3641:
3637:
3632:
3627:
3623:
3619:
3615:
3608:
3605:
3600:
3596:
3592:
3588:
3584:
3580:
3576:
3572:
3565:
3562:
3557:
3553:
3548:
3543:
3539:
3535:
3531:
3527:
3523:
3516:
3513:
3508:
3504:
3499:
3494:
3490:
3486:
3482:
3478:
3474:
3470:
3466:
3458:
3456:
3452:
3447:
3443:
3439:
3435:
3430:
3425:
3422:(3): 337–47.
3421:
3417:
3413:
3406:
3403:
3398:
3394:
3389:
3384:
3381:(4): 644–54.
3380:
3376:
3372:
3365:
3362:
3357:
3353:
3348:
3343:
3339:
3335:
3331:
3327:
3323:
3319:
3315:
3308:
3305:
3300:
3296:
3292:
3288:
3284:
3280:
3276:
3272:
3265:
3262:
3257:
3253:
3248:
3243:
3239:
3235:
3231:
3227:
3223:
3216:
3213:
3208:
3204:
3200:
3196:
3192:
3188:
3184:
3180:
3176:
3172:
3165:
3158:
3155:
3150:
3146:
3142:
3138:
3134:
3130:
3126:
3122:
3118:
3114:
3107:
3104:
3099:
3095:
3091:
3087:
3083:
3079:
3075:
3071:
3067:
3063:
3056:
3053:
3048:
3044:
3040:
3036:
3033:(3): 614–36.
3032:
3028:
3021:
3018:
3013:
3009:
3005:
3001:
2997:
2993:
2986:
2983:
2978:
2974:
2970:
2966:
2962:
2958:
2954:
2950:
2943:
2940:
2935:
2931:
2926:
2921:
2917:
2913:
2909:
2905:
2902:(8): 180744.
2901:
2897:
2893:
2886:
2883:
2878:
2874:
2869:
2864:
2860:
2856:
2852:
2848:
2844:
2837:
2834:
2829:
2825:
2820:
2815:
2811:
2807:
2803:
2799:
2795:
2788:
2785:
2780:
2776:
2771:
2766:
2762:
2758:
2757:
2752:
2745:
2742:
2737:
2733:
2728:
2723:
2719:
2715:
2711:
2707:
2706:
2701:
2694:
2692:
2690:
2686:
2681:
2677:
2672:
2667:
2663:
2659:
2655:
2651:
2647:
2640:
2638:
2634:
2629:
2625:
2620:
2615:
2611:
2607:
2603:
2599:
2595:
2588:
2585:
2580:
2576:
2571:
2566:
2562:
2558:
2554:
2550:
2546:
2544:
2535:
2532:
2527:
2523:
2518:
2513:
2509:
2505:
2501:
2497:
2493:
2486:
2483:
2478:
2474:
2469:
2464:
2460:
2456:
2452:
2448:
2444:
2437:
2434:
2429:
2425:
2421:
2417:
2412:
2407:
2404:(4): 503–14.
2403:
2399:
2395:
2388:
2385:
2380:
2376:
2371:
2366:
2361:
2356:
2352:
2348:
2344:
2337:
2334:
2329:
2325:
2321:
2317:
2313:
2309:
2305:
2301:
2297:
2293:
2286:
2283:
2280:
2276:
2275:
2268:
2265:
2260:
2256:
2252:
2248:
2243:
2238:
2234:
2230:
2226:
2219:
2216:
2211:
2207:
2202:
2197:
2193:
2189:
2185:
2178:
2175:
2170:
2166:
2161:
2156:
2153:(5): 867–74.
2152:
2148:
2144:
2137:
2134:
2129:
2125:
2120:
2115:
2111:
2107:
2102:
2097:
2093:
2089:
2085:
2081:
2077:
2070:
2067:
2062:
2058:
2054:
2050:
2046:
2042:
2038:
2034:
2030:
2026:
2019:
2016:
2011:
2007:
2002:
1997:
1993:
1989:
1986:(22): e1657.
1985:
1981:
1977:
1970:
1967:
1962:
1958:
1954:
1950:
1947:(8): 414–22.
1946:
1942:
1935:
1932:
1927:
1923:
1918:
1913:
1908:
1903:
1899:
1895:
1891:
1887:
1883:
1876:
1873:
1868:
1864:
1859:
1854:
1851:(5): 653–66.
1850:
1846:
1842:
1835:
1832:
1821:
1814:
1811:
1806:
1802:
1798:
1796:9780716771081
1792:
1787:
1786:
1777:
1774:
1769:
1765:
1760:
1755:
1751:
1747:
1743:
1739:
1735:
1728:
1725:
1720:
1716:
1712:
1708:
1704:
1700:
1697:(1): 181–90.
1696:
1692:
1685:
1682:
1670:
1664:
1661:
1656:
1652:
1648:
1644:
1640:
1636:
1629:
1627:
1623:
1618:
1614:
1610:
1606:
1602:
1598:
1594:
1590:
1586:
1582:
1578:
1571:
1568:
1563:
1559:
1555:
1551:
1547:
1543:
1539:
1535:
1531:
1527:
1523:
1516:
1513:
1508:
1504:
1500:
1496:
1492:
1488:
1484:
1477:
1474:
1469:
1462:
1459:
1454:
1450:
1446:
1442:
1438:
1434:
1430:
1426:
1422:
1418:
1414:
1407:
1404:
1397:
1387:
1384:
1377:
1373:
1370:
1368:
1365:
1363:
1360:
1358:
1355:
1352:
1349:
1347:
1344:
1342:
1339:
1337:
1334:
1332:
1329:
1327:
1324:
1322:
1319:
1318:
1314:
1308:
1303:
1298:
1296:
1294:
1290:
1285:
1282:
1278:
1269:
1267:
1265:
1260:
1258:
1254:
1249:
1247:
1243:
1234:
1232:
1230:
1226:
1210:
1206:
1195:
1192:
1188:
1185:
1181:
1178:
1174:
1171:
1167:
1164: –
1163:
1159:
1158:Find sources:
1153:
1149:
1143:
1142:
1138:
1134:
1129:This section
1127:
1118:
1117:
1111:
1104:
1102:
1101:
1097:
1096:
1092:
1090:
1089:
1085:
1084:
1080:
1078:
1077:
1073:
1072:
1068:
1066:
1065:
1061:
1060:
1056:
1054:
1053:
1049:
1048:
1044:
1042:
1041:
1037:
1036:
1032:
1030:
1029:
1025:
1024:
1020:
1018:
1017:
1013:
1012:
1006:
1004:
1003:
999:
997:
992:
988:
986:
985:
981:
979:
975:
974:
970:
968:
967:
963:
961:
958:
957:
953:
951:
950:
946:
945:
941:
939:
938:
934:
933:
929:
927:
926:
922:
920:
916:
912:
910:
909:
908:Chlamydomonas
905:
902:
901:
898:CTCGGTTATGGG
897:
895:
894:
890:
889:
885:
883:
882:
878:
877:
873:
871:
870:
866:
864:
859:
855:
853:
852:
848:
845:
842:
841:
837:
835:
834:
829:
828:
823:
822:
818:
817:
813:
811:
810:
806:
805:
801:
799:
798:
793:
792:
788:
785:
781:
777:
775:
774:
769:
768:
764:
761:
760:Kinetoplastid
758:
757:
753:
751:
750:
749:Dictyostelium
746:
745:
741:
739:
738:
733:
732:
728:
726:
722:
718:
716:
715:
711:
709:
705:
704:
700:
698:
697:
692:
688:
685:
683:
680:
679:
675:
672:
669:
668:
662:
660:
657:website (see
656:
652:
648:
640:
638:
634:
632:
628:
624:
619:
617:
616:
611:
607:
603:
599:
591:
586:
579:
577:
575:
571:
567:
566:Meta-analyses
560:
558:
556:
550:
548:
545:response and
544:
537:
529:
527:
525:
521:
517:
513:
505:
500:
498:
496:
492:
487:
483:
479:
471:
469:
467:
463:
459:
455:
451:
447:
443:
436:
427:
420:
418:
416:
412:
408:
404:
400:
396:
392:
388:
384:
380:
376:
369:
360:
353:
351:
349:
345:
344:
339:
338:
337:Agrobacterium
333:
332:
327:
323:
318:
314:
312:
311:cell division
308:
304:
300:
295:
290:
286:
277:
272:
264:
259:
257:
255:
252:
248:
245:
241:
237:
236:Carol Greider
233:
229:
225:
220:
217:
213:
209:
204:
202:
198:
194:
190:
189:
184:
176:
174:
172:
168:
164:
160:
156:
152:
148:
144:
141:
134:
130:
127:
124:
117:
113:
110:
109:Ancient Greek
104:
56:
48:
43:
37:
33:
19:
6002:
5953:Gene cluster
5721:Alu sequence
5630:Interspersed
5504:
5492:
5429:Microbiology
5414:Biochemicals
5350:
5326:Gene therapy
5267:Biosynthesis
5245:Applications
5166:Spectroscopy
5136:Cell culture
5045:Fermentation
4726:
4616:
4584:
4424:Cytogenetics
4388:Jack Szostak
4321:
4317:
4307:
4264:
4260:
4250:
4215:
4211:
4201:
4142:
4138:
4128:
4077:
4073:
4063:
4020:
4016:
4006:
3987:
3977:
3944:
3940:
3934:
3925:
3915:
3882:
3878:
3872:
3835:
3831:
3821:
3778:
3774:
3764:
3729:
3725:
3715:
3680:
3676:
3666:
3621:
3617:
3607:
3577:(8): 743–7.
3574:
3570:
3564:
3529:
3525:
3515:
3472:
3468:
3419:
3415:
3405:
3378:
3374:
3364:
3321:
3317:
3307:
3274:
3270:
3264:
3232:(5): 761–8.
3229:
3225:
3215:
3174:
3170:
3157:
3116:
3112:
3106:
3065:
3061:
3055:
3030:
3026:
3020:
2995:
2991:
2985:
2952:
2948:
2942:
2899:
2895:
2885:
2850:
2846:
2836:
2801:
2797:
2787:
2760:
2754:
2744:
2709:
2703:
2653:
2649:
2601:
2597:
2587:
2552:
2548:
2542:
2534:
2499:
2495:
2485:
2450:
2446:
2436:
2401:
2397:
2387:
2350:
2346:
2336:
2295:
2291:
2285:
2272:
2267:
2235:(8): 363–9.
2232:
2228:
2218:
2191:
2187:
2177:
2150:
2146:
2136:
2083:
2079:
2069:
2028:
2024:
2018:
1983:
1980:Bio-Protocol
1979:
1969:
1944:
1940:
1934:
1889:
1885:
1875:
1848:
1844:
1834:
1823:. Retrieved
1813:
1784:
1776:
1741:
1737:
1727:
1694:
1690:
1684:
1673:. Retrieved
1663:
1641:(1): 33–53.
1638:
1634:
1584:
1580:
1570:
1529:
1525:
1515:
1490:
1486:
1476:
1467:
1461:
1420:
1416:
1406:
1386:
1286:
1281:heritability
1273:
1261:
1250:
1238:
1222:
1200:
1190:
1183:
1176:
1169:
1157:
1137:verification
1130:
1098:
1086:
1074:
1062:
1050:
1038:
1026:
1014:
1000:
982:
964:
954:TTGCGTCTGGG
947:
942:TTAGGTTGGGG
935:
923:
906:
891:
879:
867:
856:TTAGGG(T/C)
849:
844:Apicomplexan
831:
825:
819:
807:
795:
789:
771:
765:
747:
735:
729:
725:Slime moulds
712:
706:Filamentous
694:
644:
635:
630:
620:
613:
595:
564:
551:
539:
509:
475:
438:
371:
341:
335:
331:Streptomyces
329:
319:
315:
282:
240:Jack Szostak
221:
216:somatic cell
205:
200:
196:
186:
180:
139:
136:
129:
122:
119:
112:
107:; from
54:
52:
6029:Chromosomes
5948:Gene family
5859:Tc1/mariner
5814:EnSpm/CACTA
5419:Biorobotics
5409:Biomimetics
5404:Biomedicine
4637:Polycentric
4627:Monocentric
4610:Holocentric
4605:Acrocentric
4600:Telocentric
4590:Metacentric
4468:Euchromatin
4428:chromosomes
3781:(1): 1300.
2955:: 223–245.
2656:: 158–169.
2545:? A review"
1336:Immortality
1287:Studies on
1270:In wildlife
1235:Measurement
1227:, or SASP (
1131:needs more
925:Bombyx mori
904:Green algae
886:TTTTTTAGGG
827:Stylonychia
814:TTGGG(T/G)
791:Tetrahymena
767:Trypanosoma
682:Vertebrates
580:Lengthening
497:or D-loop.
415:vertebrates
403:vertebrates
322:prokaryotes
151:chromosomes
47:chromosomes
6023:Categories
5960:Pseudogene
5777:retroposon
5695:Transposon
5557:transposon
5374:Bioeconomy
5346:Stem cells
5299:Embryology
5222:Filtration
5212:Extraction
5131:Bioreactor
4789:Centromere
4720:Structures
4699:Polyploidy
4689:Aneuploidy
4490:Nucleosome
4480:Chromosome
3838:(1): 272.
3732:(9): e75.
3624:(1): 145.
3226:Aging Cell
1845:Aging Cell
1825:2008-06-22
1675:2012-06-12
1398:References
1351:Senescence
1326:Centromere
1289:ectotherms
1277:endotherms
1203:March 2018
1173:newspapers
1162:"Telomere"
960:Roundworms
851:Plasmodium
809:Paramecium
651:nucleotide
543:DNA damage
501:Shortening
478:base pairs
462:senescence
446:stem cells
442:germ cells
435:Telomerase
421:Telomerase
307:base pairs
254:telomerase
199:(end) and
167:DNA repair
159:eukaryotes
147:nucleotide
6044:Telomeres
5879:P element
5829:Harbinger
5570:Repeatome
5217:Fed Batch
5117:and tools
4741:Protamine
4648:Processes
4632:Dicentric
4485:Chromatid
4463:Chromatin
4444:Karyotype
4299:208319503
4283:1461-023X
4234:0962-1083
4193:226291119
4167:2054-5703
4120:237328316
4104:0962-1083
4037:0962-8436
3947:: e1727.
3640:1471-2105
3489:0305-1048
3446:206331112
3338:0305-1048
2977:209748557
2502:: 37–45.
2229:BioEssays
2194:(3): 19.
2188:MedGenMed
2110:0027-8424
1805:191854286
1601:0531-5565
1554:0022-5193
1499:0002-3264
1437:1476-5594
1367:G-quartet
1357:Tankyrase
1257:Flow-FISH
1021:TCTGGGTG
913:TTTTAGGG
846:protozoa
838:TTTTGGGG
821:Oxytricha
786:protozoa
773:Crithidia
762:protozoa
673:Organism
641:Sequences
368:Shelterin
326:bacterial
177:Discovery
155:Sequences
6003:Telomere
5969:See also
5909:Zisupton
5889:Polinton
5884:PiggyBac
5839:Helitron
5658:Helitron
5653:Polinton
5549:Genetics
5494:Category
5449:Virology
5351:Telomere
4998:Branches
4885:See also
4727:Telomere
4694:Euploidy
4622:Acentric
4519:allosome
4511:Autosome
4437:concepts
4348:29335373
4291:31773847
4242:33662151
4185:33391781
4112:34437736
4055:29335377
3969:44594597
3961:22415954
3907:20463557
3899:17406480
3864:31138115
3813:29358629
3756:24609383
3707:12000852
3658:33752601
3599:23833545
3507:36546771
3438:26716914
3397:25828846
3356:37602392
3347:10767889
3299:10309423
3291:21194798
3256:21518243
3207:13841086
3199:15315667
3149:35667874
3047:14315085
3012:13905658
2969:31900099
2934:30225068
2877:28394764
2828:22782639
2779:33504179
2736:35165420
2680:26853993
2628:19089916
2579:29212750
2526:29604323
2477:17012276
2420:10338214
2379:20826803
2259:11920124
2210:15520642
2169:15471900
2128:31285335
2061:37387314
2053:10784448
2010:27182535
1961:19589679
1867:20569239
1768:22661228
1617:26381790
1453:11726588
1445:20237481
1417:Oncogene
1299:See also
1081:GGTGTAC
994:Budding
976:Fission
874:TTTAGGG
833:Euplotes
797:Glaucoma
754:AG(1-8)
737:Didymium
731:Physarum
647:TeloBase
623:seabirds
512:in vitro
448:such as
411:ciliates
348:proteins
343:Borrelia
203:(part).
55:telomere
5899:Transib
5874:Novosib
5854:Kolobok
5824:Ginger2
5819:Ginger1
5804:Crypton
5506:Commons
5389:Biology
5282:Cloning
5060:Protein
5055:Plasmid
4963:History
4897:Plasmid
4751:Histone
4662:Meiosis
4657:Mitosis
4339:5784069
4176:7735339
4147:Bibcode
4046:5784070
3855:6540518
3804:5778012
3783:Bibcode
3747:4027178
3649:7986547
3591:9702772
3556:2664709
3498:9841428
3247:3387546
3179:Bibcode
3141:9454332
3121:Bibcode
3113:Science
3098:9440710
3090:7544491
3070:Bibcode
3062:Science
2925:6124068
2904:Bibcode
2868:5425118
2819:3480822
2727:8985209
2671:5590630
2619:2727686
2570:5746531
2543:in vivo
2517:6162185
2468:1636468
2370:2975205
2328:4360698
2320:6330571
2300:Bibcode
2251:2241933
2201:1435592
2119:6660761
2088:Bibcode
2033:Bibcode
2025:Science
2001:4863463
1926:2780561
1894:Bibcode
1759:3371406
1719:4754905
1699:Bibcode
1609:9415101
1562:4754905
1534:Bibcode
1507:5158754
1293:Metazoa
1187:scholar
1152:removed
971:TTAGGC
919:Insects
861:Higher
802:TTGGGG
784:Ciliate
778:TTAGGG
742:TTAGGG
719:TTAGGG
701:TTAGGG
696:Xenopus
615:Science
520:in vivo
482:guanine
399:guanine
289:3' ends
5998:CRISPR
5864:Merlin
5849:ISL2EU
5799:Academ
5632:repeat
5434:Mining
5367:fields
5026:Allele
4458:Genome
4449:Ploidy
4346:
4336:
4297:
4289:
4281:
4240:
4232:
4191:
4183:
4173:
4165:
4118:
4110:
4102:
4053:
4043:
4035:
3988:Nature
3967:
3959:
3905:
3897:
3862:
3852:
3811:
3801:
3754:
3744:
3705:
3698:115301
3695:
3656:
3646:
3638:
3597:
3589:
3554:
3547:318019
3544:
3505:
3495:
3487:
3444:
3436:
3395:
3354:
3344:
3336:
3297:
3289:
3254:
3244:
3205:
3197:
3147:
3139:
3096:
3088:
3045:
3010:
2975:
2967:
2932:
2922:
2875:
2865:
2826:
2816:
2777:
2734:
2724:
2678:
2668:
2626:
2616:
2577:
2567:
2524:
2514:
2475:
2465:
2428:721901
2426:
2418:
2377:
2367:
2326:
2318:
2292:Nature
2257:
2249:
2208:
2198:
2167:
2126:
2116:
2108:
2059:
2051:
2008:
1998:
1959:
1924:
1917:297991
1914:
1865:
1803:
1793:
1766:
1756:
1717:
1655:642006
1653:
1615:
1607:
1599:
1560:
1552:
1505:
1497:
1451:
1443:
1435:
1242:WALTER
1189:
1182:
1175:
1168:
1160:
996:yeasts
978:yeasts
930:TTAGG
893:Allium
863:plants
670:Group
472:Length
466:cancer
393:, and
340:, and
294:primer
251:enzyme
238:, and
45:Human
5904:Zator
5844:IS3EU
5749:LINE2
5744:LINE1
5736:LINEs
5713:SINEs
5669:Other
5458:Lists
5336:Omics
4735:TINF2
4504:Types
4435:Basic
4295:S2CID
4189:S2CID
4116:S2CID
3965:S2CID
3903:S2CID
3595:S2CID
3442:S2CID
3295:S2CID
3203:S2CID
3167:(PDF)
3145:S2CID
3094:S2CID
2973:S2CID
2847:Aging
2424:S2CID
2324:S2CID
2255:S2CID
2057:S2CID
1613:S2CID
1449:S2CID
1378:Notes
1194:JSTOR
1180:books
708:fungi
691:mouse
687:Human
230:with
201:meros
197:telos
153:(see
140:méros
133:μέρος
123:télos
116:τέλος
111:
5894:Sola
5869:MuDR
5809:Dada
5772:MER4
5767:HERV
5759:LTRs
5187:CSTR
5156:HPLC
5050:Gene
5031:Cell
4517:(or
4344:PMID
4287:PMID
4279:ISSN
4238:PMID
4230:ISSN
4181:PMID
4163:ISSN
4108:PMID
4100:ISSN
4051:PMID
4033:ISSN
3957:PMID
3895:PMID
3860:PMID
3809:PMID
3752:PMID
3703:PMID
3654:PMID
3636:ISSN
3587:PMID
3552:PMID
3503:PMID
3485:ISSN
3434:PMID
3393:PMID
3352:PMID
3334:ISSN
3287:PMID
3252:PMID
3195:PMID
3137:PMID
3086:PMID
3043:PMID
3008:PMID
2965:PMID
2930:PMID
2873:PMID
2824:PMID
2775:PMID
2732:PMID
2676:PMID
2624:PMID
2575:PMID
2522:PMID
2473:PMID
2416:PMID
2398:Cell
2375:PMID
2316:PMID
2247:PMID
2206:PMID
2165:PMID
2124:PMID
2106:ISSN
2049:PMID
2006:PMID
1957:PMID
1922:PMID
1863:PMID
1801:OCLC
1791:ISBN
1764:PMID
1715:PMID
1651:PMID
1605:PMID
1597:ISSN
1558:PMID
1550:ISSN
1503:PMID
1495:ISSN
1441:PMID
1433:ISSN
1166:news
1135:for
395:RAP1
391:TPP1
387:POT1
383:TIN2
379:TRF2
375:TRF1
309:per
244:2009
5834:hAT
5726:MIR
5227:PFR
5161:NMR
5040:RNA
5036:DNA
4769:H2B
4764:H2A
4334:PMC
4326:doi
4322:373
4269:doi
4220:doi
4171:PMC
4155:doi
4090:hdl
4082:doi
4041:PMC
4025:doi
4021:373
3992:doi
3949:doi
3945:344
3941:BMJ
3887:doi
3850:PMC
3840:doi
3799:PMC
3791:doi
3742:PMC
3734:doi
3693:PMC
3685:doi
3644:PMC
3626:doi
3579:doi
3542:PMC
3534:doi
3493:PMC
3477:doi
3424:doi
3383:doi
3342:PMC
3326:doi
3279:doi
3242:PMC
3234:doi
3187:doi
3129:doi
3117:279
3078:doi
3066:269
3035:doi
3000:doi
2957:doi
2920:PMC
2912:doi
2863:PMC
2855:doi
2814:PMC
2806:doi
2765:doi
2722:PMC
2714:doi
2666:PMC
2658:doi
2614:PMC
2606:doi
2602:124
2565:PMC
2557:doi
2512:PMC
2504:doi
2500:177
2463:PMC
2455:doi
2406:doi
2365:PMC
2355:doi
2351:285
2308:doi
2296:310
2237:doi
2196:PMC
2155:doi
2114:PMC
2096:doi
2084:116
2041:doi
2029:288
1996:PMC
1988:doi
1949:doi
1912:PMC
1902:doi
1853:doi
1754:PMC
1746:doi
1707:doi
1643:doi
1639:120
1589:doi
1542:doi
1491:201
1425:doi
1246:PCR
1231:).
299:RNA
82:ɪər
6025::
5559:,
5555:,
5551::
4812:C2
4807:C1
4779:H4
4774:H3
4759:H1
4729::
4426::
4342:.
4332:.
4320:.
4316:.
4293:.
4285:.
4277:.
4265:23
4263:.
4259:.
4236:.
4228:.
4216:31
4214:.
4210:.
4187:.
4179:.
4169:.
4161:.
4153:.
4141:.
4137:.
4114:.
4106:.
4098:.
4088:.
4078:31
4076:.
4072:.
4049:.
4039:.
4031:.
4019:.
4015:.
3990:.
3986:.
3963:.
3955:.
3943:.
3924:.
3901:.
3893:.
3881:.
3858:.
3848:.
3836:20
3834:.
3830:.
3807:.
3797:.
3789:.
3777:.
3773:.
3750:.
3740:.
3730:42
3728:.
3724:.
3701:.
3691:.
3681:30
3679:.
3675:.
3652:.
3642:.
3634:.
3622:22
3620:.
3616:.
3593:.
3585:.
3575:16
3573:.
3550:.
3540:.
3530:17
3528:.
3524:.
3501:.
3491:.
3483:.
3473:51
3471:.
3467:.
3454:^
3440:.
3432:.
3420:85
3418:.
3414:.
3391:.
3379:82
3377:.
3373:.
3350:.
3340:.
3332:.
3322:52
3320:.
3316:.
3293:.
3285:.
3275:33
3273:.
3250:.
3240:.
3230:10
3228:.
3224:.
3201:.
3193:.
3185:.
3175:13
3173:.
3169:.
3143:.
3135:.
3127:.
3115:.
3092:.
3084:.
3076:.
3064:.
3041:.
3031:37
3029:.
3006:.
2996:25
2994:.
2971:.
2963:.
2953:41
2951:.
2928:.
2918:.
2910:.
2898:.
2894:.
2871:.
2861:.
2849:.
2845:.
2822:.
2812:.
2802:18
2800:.
2796:.
2773:.
2761:41
2759:.
2753:.
2730:.
2720:.
2710:24
2708:.
2702:.
2688:^
2674:.
2664:.
2654:54
2652:.
2648:.
2636:^
2622:.
2612:.
2600:.
2596:.
2573:.
2563:.
2553:13
2551:.
2547:.
2520:.
2510:.
2498:.
2494:.
2471:.
2461:.
2451:34
2449:.
2445:.
2422:.
2414:.
2402:97
2400:.
2396:.
2373:.
2363:.
2349:.
2345:.
2322:.
2314:.
2306:.
2294:.
2253:.
2245:.
2233:12
2231:.
2227:.
2204:.
2190:.
2186:.
2163:.
2151:26
2149:.
2145:.
2122:.
2112:.
2104:.
2094:.
2082:.
2078:.
2055:.
2047:.
2039:.
2027:.
2004:.
1994:.
1982:.
1978:.
1955:.
1945:19
1943:.
1920:.
1910:.
1900:.
1890:86
1888:.
1884:.
1861:.
1847:.
1843:.
1799:.
1762:.
1752:.
1742:26
1740:.
1736:.
1713:.
1705:.
1695:41
1693:.
1649:.
1637:.
1625:^
1611:.
1603:.
1595:.
1585:31
1583:.
1579:.
1556:.
1548:.
1540:.
1530:41
1528:.
1524:.
1501:.
1489:.
1485:.
1447:.
1439:.
1431:.
1421:29
1419:.
1415:.
1154:.
830:,
824:,
794:,
770:,
734:,
693:,
689:,
389:,
385:,
381:,
377:,
334:,
313:.
256:.
173:.
103:-/
94:iː
53:A
5615:)
5611:(
5602:/
5541:e
5534:t
5527:v
5038:/
4948:e
4941:t
4934:v
4872:T
4867:Q
4862:P
4857:O
4852:N
4847:M
4842:K
4837:J
4832:I
4827:H
4822:F
4817:E
4802:B
4797:A
4737:)
4733:(
4532:/
4513:/
4456:/
4416:e
4409:t
4402:v
4350:.
4328::
4301:.
4271::
4244:.
4222::
4195:.
4157::
4149::
4143:7
4122:.
4092::
4084::
4057:.
4027::
4000:.
3994::
3971:.
3951::
3928:.
3909:.
3889::
3883:1
3866:.
3842::
3815:.
3793::
3785::
3779:8
3758:.
3736::
3709:.
3687::
3660:.
3628::
3601:.
3581::
3558:.
3536::
3509:.
3479::
3448:.
3426::
3399:.
3385::
3358:.
3328::
3301:.
3281::
3258:.
3236::
3209:.
3189::
3181::
3151:.
3131::
3123::
3100:.
3080::
3072::
3049:.
3037::
3014:.
3002::
2979:.
2959::
2936:.
2914::
2906::
2900:5
2879:.
2857::
2851:9
2830:.
2808::
2781:.
2767::
2738:.
2716::
2682:.
2660::
2630:.
2608::
2581:.
2559::
2528:.
2506::
2479:.
2457::
2430:.
2408::
2381:.
2357::
2330:.
2310::
2302::
2261:.
2239::
2212:.
2192:6
2171:.
2157::
2130:.
2098::
2090::
2063:.
2043::
2035::
2012:.
1990::
1984:5
1963:.
1951::
1928:.
1904::
1896::
1869:.
1855::
1849:9
1828:.
1807:.
1770:.
1748::
1721:.
1709::
1701::
1678:.
1657:.
1645::
1619:.
1591::
1564:.
1544::
1536::
1509:.
1455:.
1427::
1240:(
1205:)
1201:(
1191:·
1184:·
1177:·
1170:·
1144:.
629:(
143:)
137:(
126:)
120:(
100:ə
97:l
91:t
88:ˈ
85:,
79:m
76:ə
73:l
70:ɛ
67:t
64:ˈ
61:/
57:(
38:.
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
