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p53 Deficiency Rescues the Adverse Effects of Telomere Loss and Cooperates with Telomere Dysfunction to Accelerate Carcinogenesis
Open accessAbstract
Maintenance of telomere length and function is critical for the efficient proliferation of eukaryotic cells. Here, we examine the interactions between telomere dysfunction and p53 in cells and organs of telomerase-deficient mice. Coincident with severe telomere shortening and associated genomic instability, p53 is activated, leading to growth arrest and/or apoptosis. Deletion of p53 significantly attenuated the adverse cellular and organismal effects of telomere dysfunction, but only during the earliest stages of genetic crisis. Correspondingly, the loss of telomere function and p53 deficiency cooperated to initiate the transformation process. Together, these studies establish a key role for p53 in the cellular response to telomere dysfunction in both normal and neoplastic cells, question the significance of crisis as a tumor suppressor mechanism, and identify a biologically relevant stage of advanced crisis, termed genetic catastrophe.
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TL;DR
In telomerase-deficient (mTR−/−) mice, critically short telomeres activate p53, and deleting p53 rescues the resulting apoptosis and cell-cycle arrest — but only during early "crisis." Loss of telomere function and p53 deficiency cooperate to initiate transformation, while telomerase reconstitution mainly benefits later stages of tumorigenesis.
Key findings
- Critical telomere shortening activates p53: p53 protein was detectable in G6 mTR−/− testis but not in mTR+/+ or G6 mTR−/− p53−/− littermates; >80% of G4 and G6 MEFs stained positive for p53 at passage 6 versus 21% of mTR+/+ MEFs. p21CIP1/WAF1 rose 3- to 4-fold in G6 versus G2 mTR−/− p53+/+ MEFs, and p53-dependent CAT reporter activity increased ~2-fold.
- p53 loss rescued telomere-dysfunction phenotypes early but not late: 15% of G4–G6 mTR−/− mice (86/565) were p53−/− versus 26% of G7–G8 mice (24/91) (p = 0.015); all G5 mTR−/− p53−/− testes had normal histology, but 3 of 7 G6 and all five G7–G8 samples showed germ cell depletion. G6 mTR−/− p53−/− MEFs showed a 3-fold increase in chromosome fusions (1.65, SD 2.59 vs 0.42, SD 0.71 per metaphase; p = 0.012), with no change in telomere length (75% reduction regardless of p53 status).
- Telomere dysfunction cooperated with p53 deficiency to initiate transformation: adding mTR suppressed Myc/RAS focus formation in G6 mTR−/− p53−/− MEFs, but mTR-transfected foci had higher subcloning efficiency (23/26, 89%) than vector foci (19/32, 59%; p = 0.018). Tumor formation in SCID mice was similar between groups.
- The authors propose dividing crisis into p53-dependent "early crisis" and p53-independent "genetic catastrophe," questioning crisis as a tumor suppressor mechanism.
Methods
- Mouse genetics: mTR+/− mice crossed with p53+/− mice to generate successive generations (G1–G8) of mTR−/− animals with p53+/+, +/−, and −/− genotypes in a mixed WW6/C57BL/6 background; p53 status maintained heterozygous during matings.
- Cellular/organ assays: immunoblotting and immunohistochemistry for p53 and p21, pG13CAT reporter assays, testis histology with TUNEL, BrdU/propidium iodide FACS cell-cycle profiles, flow FISH telomere length, and metaphase spreads for chromosome fusions.
- Transformation assays: Myc/RAS cotransfection of early-passage MEFs with empty vector or an mTR genomic fragment, scoring focus formation, subcloning efficiency, and tumor formation after subcutaneous injection into SCID mice.
Limitations
- Stated: the mechanisms by which tumorigenic potential is maintained in late stages of tumorigenesis despite telomere dysfunction "remain to be determined"; whether loss of TRF2 binding, short telomeres, or fusion-bridge-breakage cycles mediate the p53 induction described also remains undetermined.
- Evident from the text: the G7–G8 genotype comparison rests on small numbers (24 of 91 mice), and all mice are in a mixed genetic background, which the authors note required cousin mating to avoid substrains.
- Evident from the text: the transformation data are from cultured MEFs and SCID xenografts, and tumor weights/efficiency between mTR and vector lines were not significantly different, so the in vivo relevance of the initiation advantage is inferred rather than directly demonstrated.
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40 Own interpretation · 39 Own result · 21 Not checked yet · 6 Own method · 4 No basis given · 1 Check pending
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“Correspondingly, the loss of telomere function and p53 deficiency cooperated to initiate the transformation process.”
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“Coincident with severe telomere shortening and associated genomic instability, p53 is activated, leading to growth arrest and/or apoptosis.”
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“This activity was strictly dependent upon p53, since p53−/− and G6 mTR−/− p53−/− MEF cultures showed only background levels of CAT activity.”
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“We found that p53 mediates the adverse effects of critically shortened telomeres.”
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“As shown in Figure 3A, DNA content analyses of G6 mTR−/− p53+/+ MEFs revealed a profile consistent with a biphasic block as evidenced by marked accumulation of cells in G1 and G2/M phases.”
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“Indeed, analysis of metaphase spreads derived from G6 mTR−/− p53+/+ MEFs showed a rate of 0.42 (SD 0.71) chromosomal fusions per metaphase, a rate consistent with previous studies (Figure 3D) (Blasco et al. 1997; Greenberg et al. 1999; Rudolph et al. 1999”
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“Furthermore, p53-independent cell death becomes a more consistent feature in G7 and G8, since all of the five G7–G8 mTR−/− p53−/− testis samples showed significant germ cell depletion microscopically (Figure 2D).”
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“However, a relative increase in p53−/− mice was evident in G7 and G8 mTR−/− background (24% and 38%, respectively).”
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“Gross testis size was consistently rescued to near normal in G6 mTR−/− p53−/− mice compared to G6 mTR−/− p53+/+ littermates (see Figure 2A for representative comparison).”
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“Furthermore, loss of telomere function and p53 deficiency can cooperate to enhance transformation.”
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“Transfection of this mTR genomic fragment restored telomerase activity in mTR−/− cells (Figure 4B).”
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“Modest attenuation of the cell cycle block was evident in the G6 mTR−/− p53+/− cultures, suggesting functional haploinsufficiency.”
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“However, loss of p53 does not appear to eliminate the cell cycle blocks completely, since G6 mTR−/− p53−/− MEFs still exhibited fewer cells in S phase (33% vs. 55%) and more cells in G2/M (14% vs. 7%) than mTR+/+ p53−/− MEFs.”
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“Similar results were obtained for primary thymocytes, irrespective of p53 status (Figure 3C).”
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“Endogenous p21CIP1/WAF1 levels were increased 3- to 4-fold in late-generation G6 mTR−/− p53+/+ MEFs compared to early-generation G2 mTR−/− p53+/+ MEFs (Figure 1C).”
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“Most of the G5–G7 mTR−/− p53−/− mice showed improved histology compared to their age-matched controls with competent p53 function.”
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“In mTR−/− mice, a decrease in testis size that correlated with germ cell depletion by histology was first evident in G5 mTR−/− animals and became more prevalent in G6 mTR−/− mice (Lee et al. 1998”
What the sources say
- Not checked yetcites Lee et al. 1998 source
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“However, when these established cell lines were injected into SCID mice, mTR lines and vector lines formed tumors with similar efficiency (Figure 4D).”
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“In the G6 mTR−/− p53+/− intercrosses, we observed an unusually high rate of productive mating pairs (11 of 22 mating pairs) compared to G6 mTR−/− p53+/+ matings (Lee et al. 1998”
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- Not checked yetcites Lee et al. 1998 source
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“we demonstrated that the addition of mTR consistently suppressed the rate of Myc/RAS focus formation in G6 mTR−/− p53−/− MEFs (Table 2)”
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“MEFs from mTR+/+ mice and from late-generation mTR−/− mice exhibited marked differences in the proportion of cells expressing p53 as determined by immunohistochemistry.”
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“Of note, the G8 mTR−/− p53−/− mouse that exhibited a relatively normal histology (1+ germ cell depletion, Table 1) had significantly longer telomere length by flow FISH determination (data not shown) than the other G8 mTR−/− p53−/− mouse.”
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“After normalization for transfection efficiency, CAT activity was increased approximately 2-fold in G6 mTR−/− MEFs compared to mTR+/+ p53+/+ controls at identical passage number (Figure 1D).”
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“TUNEL assays demonstrated that the restoration of germ cells in the G5–G7 mTR−/− p53−/− testis is accompanied by a dramatic reduction of apoptosis to levels observed in mTR+/+ testis (Figure 2C).”
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“As anticipated, p21CIP1/WAF1 was not detected in G6 mTR−/− p53−/− cells, indicating that p21CIP1/WAF1 induction in mTR−/− cells is largely p53 dependent.”
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“Furthermore, the average weight of mTR tumors was not significantly different from that of the vector tumors.”
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“while restoration of telomerase activity in mTR+/+ p53−/− MEFs had no significant effect (data not shown).”
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“As shown in Figure 3B, all of the G6 mTR−/− MEF cultures regardless of p53 status exhibited an equivalent and marked decline in mean telomere signal relative to mTR+/+ MEFs (75% reduction).”
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“p53−/− mice were underrepresented as a percentage of total mice in late-generation mTR−/− background (19%, 15%, and 14% for G4, G5, and G6 respectively) (Figure 1E).”
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“Myc/RAS foci transfected with mTR exhibited a significant enhancement in subcloning efficiency—23 of 26 (89%) mTR foci were successfully established as permanent cell lines compared with only 19 of 32 (59%) vector foci (Figures 4C, p value = 0.018).”
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“As telomere function worsens in later generations of mTR deficiency, germ cell death by p53-independent mechanisms becomes evident.”
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“Significantly, we also observed productive matings arising from G7 mTR−/− p53+/− pairs (5 of 17 mating pairs) to yield G8 mTR−/− offspring of all three p53 genotypes.”
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“Thus, telomere shortening in the absence of telomerase plays a critical role in signaling entry into senescence”
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“These results indicate that the oncogenic potential of a would-be cancer cell bearing short telomeres may depend upon its genetic profile.”
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“In contrast, SV40 large T antigen efficiently transformed cells from late-generation mTR−/− mice despite evidence for telomere dysfunction (Blasco et al. 1997; Greenberg et al. 1999”
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“Together, these studies establish a key role for p53 in the cellular response to telomere dysfunction in both normal and neoplastic cells”
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“identify a biologically relevant stage of advanced crisis, termed genetic catastrophe.”
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“The inactivation of Rb and/or p53 may account for the efficient transformation of cells with significant telomere loss by SV40 large T antigen.”
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“In cancer-prone mice rendered doubly null for the INK4a tumor suppressor and mTR, shortened telomeres markedly reduced both cancer incidence and transformation by cellular oncogenes (Greenberg et al., 1999”
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- Not checked yetcites Greenberg et al., 1999 source
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“Taken together, the differences in the subcloning efficiency of the mTR versus vector cells and their similarity in tumorigenic potential following prolonged passage in culture argue that telomerase provides a significant advantage in steps following the initiation of cellular transformation.”
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“However, once telomerase-deficient cells have adapted to the existing telomere dysfunction, they grow as efficiently as telomerase-expressing cells.”
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“Similar to well-characterized DNA damage checkpoints, the specific response to telomere dysfunction can be either growth arrest or apoptosis depending upon cell type, p53 pathway status, and physiological context.”
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“These findings indicate that germ cell apoptosis coincident with telomere loss is mediated by p53.”
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“Loss of telomere function in late-generation (G4 onward) mTR−/− mice has been shown to adversely affect the morphology and physiology of diverse cell types and organ systems (Lee et al. 1998; Rudolph et al. 1999”
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“Therefore, cell death in the context of extreme telomere dysfunction, genetic instability, and p53 deficiency in G7 and G8 mTR−/− mice may represent a more accurate model of the late stage of crisis experienced by human cells that bypass senescence (see Discussion).”
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“Nevertheless, we have established that p53 plays a key role in the cellular response to telomere dysfunction in vivo in both normal and neoplastic cells, a role that is particularly prominent during the early stages of genomic crisis.”
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“We propose that late crisis is not a checkpoint but instead represents the consequence of unimpeded proliferation in the absence of two powerful genome-stabilizing factors: functional telomeres and p53.”
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- Rests on an assumption: Human cells require inactivation of Rb and/or p53 to bypass the major checkpoint that limits replicative lifespan: cellular senescence.
- Rests on an assumption: Rb/p53 inactivation allows continued cell division beyond senescence, progressive telomere shortening, and entry into crisis, from which only rare survivors emerge.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Together these data establish p53 as an important effector in a pathway activated by loss of telomere function.”
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Rests on an assumption: The mTR−/− mouse is a valid model for studying the consequences of telomere loss in vivo.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The rescue of proliferation, cellularity, and cell death is therefore due to loss of p53 checkpoint functions.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Together, these data indicate that critical telomere shortening and the associated chromosomal instability correlate with an increase in the level and the activation of the p53 protein.”
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
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A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Cell death in early crisis is p53 dependent.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Furthermore, this p53-mediated pathway can dramatically affect the consequence of telomere shortening during carcinogenesis.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Rests on an assumption: The mTR−/− mouse is a valid model for studying the consequences of telomere loss in vivo.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These data indicate that as telomere function worsens in G7 and G8, p53+/+ and p53+/− mice are at a significant survival disadvantage compared to p53−/− mice.”
hedgeddirectn = 565 ·- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
In the paper
“Most significantly, in contrast to late-generation mTR−/− p53−/− MEFs, addition of mTR to the Myc/RAS cotransfections of late-generation mTR−/− INK4a−/− MEFs enhanced the rate of focus formation relative to empty vector controls.”
What the sources say
- Not checked yetcites Greenberg et al. 1999 source
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
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A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Thus, the activation of telomerase enhances cell survival possibly through avoidance of the p53-independent death of genetic catastrophe (Figure 5).”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Our in vivo and cell culture data suggest that the period defined as crisis in cultured cells can be divided into two phases: “early crisis” and “late crisis,” the latter we term “genetic catastrophe” (Figure 5).”
hedged- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Rests on an assumption: Human cells require inactivation of Rb and/or p53 to bypass the major checkpoint that limits replicative lifespan: cellular senescence.
- Rests on an assumption: Rb/p53 inactivation allows continued cell division beyond senescence, progressive telomere shortening, and entry into crisis, from which only rare survivors emerge.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In other cells, increased aneuploidy or fusion-bridge-breakage may facilitate genetic changes that promote survival, such as oncogene activation, tumor suppressor deletion, reactivation of telomere maintenance mechanisms, or adaptation to telomere dysfunction (e.g., Myc/RAS-expressing G6 mTR−/− p53−/− MEFs) (Figure 5).”
hedgedconditional- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The contrasting effect of telomere loss on cellular transformation phenotypes in cells that have intact or inactivated DNA damage pathways suggested that the p53 pathway may alter the cellular response to telomere loss in both normal and neoplastic cells.”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Cells that cannot reestablish telomere maintenance or tolerate genomic instability (e.g., germ cells in G7 and G8) will die.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This unanticipated result indicates that telomere dysfunction cooperates with p53 deficiency to facilitate cellular transformation.”
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Rests on an assumption: Focus formation reflects the earliest stages of oncogenic transformation.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The mTR−/− INK4a−/− study provides strong in vivo support for the hypothesis that maintenance of telomere function is required for efficient tumorigenesis.”
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“However, in cancer cells harboring intact DNA damage response pathways such as those with an INK4a−/− p53+/+ genotype, pharmacological inhibition of telomerase and subsequent telomere shortening would be predicted to be more therapeutically efficacious.”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- It may state two separate claims as one. 36
- The quote only partly states this claim. 1
- Probably not a substantial claim. 16
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Deletion of p53 significantly attenuated the adverse cellular and organismal effects of telomere dysfunction, but only during the earliest stages of genetic crisis.”
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In contrast, mTR+/+ p53+/+ MEF cultures demonstrated 35% of cells in S phase and only 13% in G2/M.”
boosted- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In mTR+/+ testis, p53 status did not affect testicular volume or significantly alter the cytoarchitecture of seminiferous tubules (Figure 2A and Figure 2B).”
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Whereas only 21% of the mTR+/+ MEFs at passage 6 stained positive for p53, more than 80% of MEFs derived from G4 and G6 exhibited strong immunoreactivity to p53 (Figure 1B, compare [a] with [b] and [c]).”
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“In male germ cells and lymphocytes, telomere loss elicits a strong apoptotic response, whereas MEFs experience a potent biphasic cell cycle block.”
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: The mTR−/− mouse is a valid model for studying the consequences of telomere loss in vivo.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Whereas all G5 mTR−/− p53−/− testes exhibited normal histology, 3 of 7 G6 mTR−/− p53−/− samples still showed varying degrees of pathology (Table 1).”
plain- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Accordingly, BrdU incorporation assay showed that 36% of the G6 mTR−/− p53+/+ cells were in G1 phase, 3% were in S phase, and 40% were in G2/M (Figure 3A, top panel).”
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Immunoprecipitation and Western blot analysis readily detected p53 in lysates from G6 mTR−/− testis, but not in lysates derived from age-matched mTR+/+ testis or from G6 mTR−/− p53−/− littermate controls (Figure 1A).”
plain- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
In the paper
“This induction of p53 coincides with the dramatic increase in apoptosis (as assessed by TUNEL) and decreased cellularity in the G6 mTR−/− testis (Lee et al. 1998”
What the sources say
- Not checked yetcites Lee et al. 1998 source
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Significantly, loss of p53 rescues the cell cycle arrest in MEFs and the apoptotic elimination of male germ cells.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: The mTR−/− mouse is a valid model for studying the consequences of telomere loss in vivo.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“loss of p53 abrogated this block.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
The quote only partly states this claim.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Significantly, 15% of the G4–G6 mTR−/− mice (86 out of 565 mice) were p53−/− compared to 26% of the G7–G8 mTR−/− mice (24 of 91 mice) (p = 0.015 by Fisher’s two-sided t test) (Figure 1E).”
plaincorrelationaln = 565 ·- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“By comparison, G6 mTR−/− p53−/− MEFs exhibited 1.65 (SD 2.59) fusions per metaphase, a 3-fold increase (p = 0.012).”
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In the case of MEFs, this arrest occurs in concert with the stabilization and activation of the p53 protein, evident in p53-dependent reporter assays and activation of p21CIP1/WAF1.”
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: The mTR−/− mouse is a valid model for studying the consequences of telomere loss in vivo.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Similar differences were observed in low-density seeding experiments (data not shown).”
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“In human cells, a critical telomere length is eventually reached that induces cellular senescence, thus limiting the replicative capacity of cultured cells at the Hayflick limit (Hayflick and Moorhead 1961”
What the sources say
- Not checked yetcites Hayflick and Moorhead 1961 source
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may state two separate claims as one.
In the paper
“In primary cells deficient for Rb and p53, continued cellular growth beyond the Hayflick limit results in severe telomere shortening, marked genetic instability, and massive cell death—a period referred to as crisis (Counter et al. 1992”
What the sources say
- Not checked yetcites Counter et al. 1992 source
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Ectopically expressed hTERT maintains telomere length and allows unlimited growth (Bodnar et al. 1998; Counter et al. 1998; Vaziri and Benchimol 1998”
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“The notion that telomere function may indeed be required for immortalization is supported by the frequent activation of telomerase or ALT in human cancers (Kim et al. 1994; Bryan et al. 1995”
What the sources say
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“The telomere signal that activates this senescent program operates through the RB and p53 pathways, since cell division beyond the Hayflick limit can occur after antisense neutralization (Hara et al. 1991”
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
In the paper
“Telomerase is comprised of essential catalytic (telomerase reverse transcriptase [TERT]) and RNA components (telomerase RNA [TR]) (reviewed in Greider 1998”
What the sources say
- Not checked yetcites Greider 1998 source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“Strikingly, as mTR−/− mice advance with age and generation, an increase in cancer incidence is observed presumably due to increased genetic instability caused by telomere shortening, repeated chromosomal fusion-bridge-breakage cycles, and chromosomal loss (Rudolph et al. 1999”
What the sources say
- Not checked yetcites Rudolph et al. 1999 source
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Due to the very long telomeres of Mus musculus, critical reduction in telomere length and chromosomal instability is only seen following extensive passage of mTR−/− cells (Niida et al. 1998”
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“The RNA subunit provides the template for telomere repeat synthesis (Greider and Blackburn 1989; Yu et al. 1990”
What the sources say
- Not checked yetcites Greider and Blackburn 1989
- Not checked yetcites Yu et al. 1990
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“although immortalization of some epithelial cells requires not only TERT expression but also compromise of the RB pathway (Kiyono et al. 1998”
What the sources say
- Not checked yetcites Kiyono et al. 1998 source
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Although loss of telomere and p53 function facilitates initiation of transformation, the reconstitution of telomerase activity greatly improved the capacity of transformed foci to yield immortal cultures.”
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Because G2 mTR−/− and G6 mTR−/− MEFs both lack telomerase activity, we conclude that telomere shortening or associated chromosome fusions and breakage rather than telomerase deficiency per se signals to p53 and accounts for p21CIP1/WAF1 induction.”
boosteddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Thus, the rampant genetic instability of the G7 and G8 mTR−/− p53−/− cells is associated with loss of cell viability despite p53 deficiency, implying that advanced genomic catastrophe gives way to a p53-independent process.”
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: The mTR−/− mouse is a valid model for studying the consequences of telomere loss in vivo.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“These results indicate that telomere dyfunction acts synergistically with p53 deficiency to initiate malignant transformation, presumably by enhancing genetic instability.”
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Continued proliferation in the absence of the p53 checkpoint may lead to an increased loss of telomere sequences and therefore account for the increased frequency of chromosome fusions.”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“For these matings, the p53 status was maintained in the heterozygous state to avoid potential broad genomic effects arising from p53 deficiency.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The ability of exogenous telomerase to reduce focus formation in p53 null cells with short dysfunctional telomeres suggests that genetic instability associated with telomere loss can serve as an engine driving initiation of transformation, but only in the absence of functional p53 (Figure 5).”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: Focus formation reflects the earliest stages of oncogenic transformation.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Early crisis accounts for efficient germ cell depletion in G6 mTR−/− p53+/+, whereas the altered histology of G7 and G8 mTR−/− p53−/− testis provides experimental support for late crisis.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: The mTR−/− mouse is a valid model for studying the consequences of telomere loss in vivo.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Together, these data indicate that p53 deficiency restores neither telomere length nor function in the telomerase-deficient mouse.”
hedged- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“This mating scheme also produced p53+/+ and p53−/− littermate controls for each mTR−/− generation.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“Telomere shortening in mTR−/− mice results in loss of telomere function and chromosome end-to-end fusion (Blasco et al. 1997; Rudolph et al. 1999”
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The observation that mTR expression facilitates subcloning efficiency of Myc/RAS-transformed late-generation mTR−/− cells supports the importance of telomere maintenance mechanisms during later stages of carcinogenesis.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Induction of double strand breaks in DNA by ionizing radiation or genotoxic chemotherapy leads to the rapid stabilization of p53 through phosphorylation of its N terminus (Shieh et al. 1997; Siliciano et al. 1997”
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Together with previous work, our data indicated that, first, during the early stages of tumorigenesis, cells suffer telomere shortening, and second, that telomerase activation is advantageous primarily in later stages of tumorigenesis.”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
- Rests on an assumption: Many human tumors that are telomerase positive still have short telomeres.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Cells with short telomeres and intact p53-mediated DNA damage responses may be more efficiently “culled” from the population (via apoptosis or arrest); therefore, p53 null cells may accumulate more end-to-end fusions than their p53+/+ counterparts.”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Once both telomere function and p53 have been impaired, the fate of the cell is unpredictable and depends on other stochastic genetic changes.”
plainconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“This stabilization results in either cell cycle arrest via transcriptional induction of the p21CIP1/WAF1 CDK inhibitor or apoptosis (reviewed in Levine 1997”
What the sources say
- Not checked yetcites Levine 1997 source
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Loss of p53 can circumvent this checkpoint, leading eventually to p53-independent apoptosis, perhaps as a consequence of uncontrolled chromosome fusion-bridge-breakage cycles and progressive genomic instability.”
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Thus, both the apoptotic response and cell cycle arrest induced by telomere dysfunction are mediated by p53.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“For telomere length determination of MEFs, littermate MEF cultures were assayed simultaneously to reduce experimental variability.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“At least two independent determinations of the same cultures were performed.”
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“Day to day variations in the linearity of the flow cytometer were controlled by the use of FITC-labeled fluorescence beads (Flow Cytometry Standards Corporation).”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
What the paper did — it cites no source for this, so there is none to check. The quote shows it is in the paper.
In the paper
“Three independent age-matched sets of animals were used.”
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In mTR−/− INK4a−/− mice, it is likely that telomere dysfunction activates p53 via a DNA damage signaling pathway and results in growth arrest or apoptosis, limiting the efficiency of carcinogenesis”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper relies on this without support.
The signals and pathways mediating the cellular responses associated with telomere dysfunction have yet to be defined in mammalian cells.
“The signals and pathways mediating these cellular responses associated with telomere dysfunction have yet to be defined in mammalian cells.”
The paper relies on this without support.
The RAD9 and MEC1 DNA damage response pathways in yeast are similar to the p53 checkpoint in mammalian cells.
“The RAD9 and MEC1 DNA damage response pathways in yeast are thought to be similar to the p53 checkpoint in mammalian cells.”
The paper relies on this without support.
The mTR−/− mouse is a valid model for studying the consequences of telomere loss in vivo.
“The mTR−/− mouse serves as a model to study the consequences of telomere loss in vivo.”
The paper relies on this without support.
Focus formation reflects the earliest stages of oncogenic transformation.
“Focus formation is thought to reflect the earliest stages of oncogenic transformation.”
The paper relies on this without support.
Human cells require inactivation of Rb and/or p53 to bypass the major checkpoint that limits replicative lifespan: cellular senescence.
“Human cells require inactivation of Rb and/or p53 to bypass the major checkpoint that limits replicative lifespan: cellular senescence.”
The paper relies on this without support.
Rb/p53 inactivation allows continued cell division beyond senescence, progressive telomere shortening, and entry into crisis, from which only rare survivors emerge.
“Rb/p53 inactivation allows continued cell division beyond senescence, progressive telomere shortening, and entry into crisis, from which only rare survivors will emerge.”
The paper relies on this without support.
Many human tumors that are telomerase positive still have short telomeres.
“The fact that many human tumors that are telomerase positive still have short telomeres supports the idea that telomerase activation occurs late in tumorigenesis.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Although enabling cellular growth and survival and masking degenerative phenotypes, these mTerc -/ -p53 mutant mice experience a dramatic increase in cancer incidence, shift in tumor spectrum toward epithelial cancers, and accumulation of marked chromosomal aberrations targeting cancerrelevant loci for amplification and deletion (4-6).”
From Ink4a / Arf tumor suppressor does not modulate the degenerative conditions or tumor spectrum of the telomerase-deficient mouse (2007) · cites this paper as [4]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Specifically, p53 deficiency results in nearcomplete rescue of adverse cellular consequences associated with telomere dysfunction, including cellular growth arrest and apoptosis, progressive organ system atrophy, and physiological compromise.”
From Ink4a / Arf tumor suppressor does not modulate the degenerative conditions or tumor spectrum of the telomerase-deficient mouse (2007) · cites this paper as [4]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Telomere dysfunction appears to engage the p53-dependent double-stranded break DNA damage pathway.”
From Ink4a / Arf tumor suppressor does not modulate the degenerative conditions or tumor spectrum of the telomerase-deficient mouse (2007) · cites this paper as [4]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“When the p53 checkpoint is compromised, unchecked genomic instability caused by telomere dysfunction accelerates tumorigenesis as seen in late-generation mTerc -/ -p53 mutant mice (4, 5).”
From Ink4a / Arf tumor suppressor does not modulate the degenerative conditions or tumor spectrum of the telomerase-deficient mouse (2007) · cites this paper as [4]
Experimental — accuracy not yet measuredIn the citing paper
“By contrast, p53 inactivation alone is sufficient to avoid telomere-induced senescence and apoptosis in mice49,50.”
From Telomeres in cancer: tumour suppression and genome instability (2017) · cites this paper as [49]
In the citing paper
“In the absence of telomerase, continued cell division results in telomere shortening, loss of ‘capping’ function, and p53 activation12-15.”
From Telomere dysfunction induces metabolic and mitochondrial compromise (2011) · cites this paper as [15]
In the citing paper
“p53 deficiency markedly attenuates the degenerative phenotypes associated with telomere dysfunction, whereas constitutively activated p53 accelerates ageing phenotypes15-18.”
From Telomere dysfunction induces metabolic and mitochondrial compromise (2011) · cites this paper as [15]
In the citing paper
“Critically short telomeres induce cellular and tissue defects through checkpoint signaling pathways that converge on the p53 tumor suppressor protein, a well-known cellular stress sensor that enforces cell cycle arrest or cell death in response to diverse stimuli including activated oncogenes, DNA damage and hypoxia (69)”
From Telomeres and telomerase in cancer (2010) · cites this paper as [69]
In the citing paper
“The downstream effector pathways of telomere-induced senescence are largely p53-dependent96, but there may be involvement of the retinoblastoma protein (RB) pathway in some settings97.”
From The telomere syndromes (2012) · cites this paper as [96]
In the citing paper
“Importantly, removal of p53 function rescued many of the cellular defects despite the presence of critically short telomeres, including growth arrest in cell culture, testicular atrophy, intestinal and germ cell apoptosis46.”
From Telomere dysfunction and tumour suppression: the senescence connection (2008) · cites this paper as [46]
In the citing paper
“recent reports of the effect of the mTR deletion in different genetic backgrounds (17, 18) suggest that the role of telomerase in cancer development could be highly context dependent (19).”
From Inhibition of human telomerase in immortal human cells leads to progressive telomere shortening and cell death (1999) · cites this paper as [18]
Citations
Cites
The serial cultivation of human diploid cell strains
Hayflick, L., & Moorhead, P. S. (1961) · Experimental Cell Research · doi:10.1016/0014-4827(61)90192-6 · Cited by 8,508
p53, the cellular gatekeeper for growth and division
Levine, A. J. (1997) · Cell · doi:10.1016/s0092-8674(00)81871-1 · Cited by 7,693
Specific Association of Human Telomerase Activity with Immortal Cells and Cancer
Kim, N. W., Piatyszek, M. A., Prowse, K. R., Harley, C. B., West, M. D., Ho, P. L. C., Coviello, G. M., Wright, W. E., Weinrich, S. L., & Shay, J. W. (1994) · Science · doi:10.1126/science.7605428 · Cited by 7,091
Oncogenic ras provokes premature cell senescence associated with accumulation of p53 and p16INK4a
Serrano, M., Lin, A. W., McCurrach, M. E., Beach, D., & Lowe, S. W. (1997) · Cell · doi:10.1016/s0092-8674(00)81902-9 · Cited by 5,218 · cited 1×
“INK4a2/2 centration of 10 mM for 3.5 hr. Cells were processed as previouslymice lack functional p19ARF, a modulator of p53 function described (Serrano et al., 1997).activated by inappropiate oncogene activation, but re- tain intact DNA damage–induced p53 activation (Kamijo MEF Transformation…”
MethodInfluentialExtension of Life-Span by Introduction of Telomerase into Normal Human Cells
Bodnar, A. G., Ouellette, M., Frolkis, M., Holt, S. E., Chiu, C.-P., Morin, G. B., Harley, C. B., Shay, J. W., Lichtsteiner, S., & Wright, W. E. (1998) · Science · doi:10.1126/science.279.5349.349 · Cited by 4,975
p53-dependent apoptosis modulates the cytotoxicity of anticancer agents
Lowe, S. W., Ruley, H. E., Jacks, T., & Housman, D. E. (1993) · Cell · doi:10.1016/0092-8674(93)90719-7 · Cited by 2,911
Tumorigenic conversion of primary embryo fibroblasts requires at least two cooperating oncogenes
Land, H., Parada, L. F., & Weinberg, R. A. (1983) · Nature · doi:10.1038/304596a0 · Cited by 2,804
Requirement for p53 and p21 to Sustain G 2 Arrest After DNA Damage
Bunz, F., Dutriaux, A., Lengauer, C., Waldman, T., Zhou, S., Brown, J. P., Sedivy, J. M., Kinzler, K. W., & Vogelstein, B. (1998) · Science · doi:10.1126/science.282.5393.1497 · Cited by 2,803
Mice Lacking p21CIP1/WAF1 undergo normal development, but are defective in G1 checkpoint control
Deng, C., Zhang, P., Wade Harper, J., Elledge, S. J., & Leder, P. (1995) · Cell · doi:10.1016/0092-8674(95)90039-x · Cited by 2,113
Telomere Shortening and Tumor Formation by Mouse Cells Lacking Telomerase RNA
Blasco, M. A., Lee, H.-W., Hande, M. P., Samper, E., Lansdorp, P. M., DePinho, R. A., & Greider, C. W. (1997) · Cell · doi:10.1016/s0092-8674(01)80006-4 · Cited by 2,108
Self-citation
Cited by
Sorted by citations
Minotti, G., Menna, P., Salvatorelli, E., Cairo, G., & Gianni, L. (2004) · Pharmacological Reviews · doi:10.1124/pr.56.2.6 · Cited by 3,868
“It has been well established that telomerase is not active in most somatic tissues; therefore, telomeres shorten gradually with age, both in vitro and in vivo, and such erosion is sensed by the cells as a clock to switch toward a p53-mediated senescence program (Chin et al., 1999).”
BackgroundShelterin: the protein complex that shapes and safeguards human telomeres
de Lange, T. (2005) · Genes & Development · doi:10.1101/gad.1346005 · Cited by 3,056
“For instance, p53-deficient mice better tolerate the consequences of telomere shortening in the later generations of telomerase-deficient mice (Chin et al. 1999).”
Death and anti-death: tumour resistance to apoptosis
Igney, F. H., & Krammer, P. H. (2002) · Nature reviews. Cancer · doi:10.1038/nrc776 · Cited by 2,073
The essence of senescence: Figure 1.
Kuilman, T., Michaloglou, C., Mooi, W. J., & Peeper, D. S. (2010) · Genes & Development · doi:10.1101/gad.1971610 · Cited by 2,038
“In the absence of p53, however, telomere dysfunction instead promotes tumorigenesis (Chin et al. 1999).”
BackgroundLowe, S. W., & Lin, A. W. (2000) · Carcinogenesis · doi:10.1093/carcin/21.3.485 · Cited by 1,889
Metadata
- Authors
- Chin, L., Artandi, S. E., Shen, Q., Tam, A., Lee, S.-L., Gottlieb, G. J., Greider, C. W., & DePinho, R. A.
- Affiliations
- Harvard University (US)
- Dana-Farber Cancer Institute (US)
- Quest Diagnostics (United States) (US)
- Johns Hopkins University (US)
- Corresponding author
- DePinho, Ronald A
- Venue
- Cell
- Publisher
- Elsevier BV
- Published
- 1999-05
- Volume
- 97
- Issue
- 4
- Pages
- 527-538
- ISSN
- 0092-8674 (ISSN-L)
- OpenAlex
- W2027779909
- Cited by
- 979
- References
- 44
- Citations by year
- 2026: 10 · 2025: 12 · 2024: 18 · 2023: 21 · 2022: 15 · 2021: 22 · 2020: 15 · 2019: 26 · 2018: 24 · 2017: 23 · 2016: 32 · 2015: 34 · 2014: 23 · 2013: 40 · 2012: 51
- FWCI
- 36.05
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- Cell (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health (5r01hd028317-05, 5r01hd034880-04, 5k08ar002104-04)
- National Institute of Arthritis and Musculoskeletal and Skin Diseases (K08AR02104-01)
- Eunice Kennedy Shriver National Institute of Child Health and Human Development (R01HD34880, R01 HD28317)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Telomeres, Telomerase, and Senescence
- Health Sciences › Medicine › Oncology › Cancer-related Molecular Pathways
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Aging › Genetics, Aging, and Longevity in Model Organisms
- Keywords
- p53, telomere dysfunction, telomere shortening, telomerase deficiency, telomere crisis, carcinogenesis, genomic instability, tumor suppression, apoptosis, growth arrest, telomere length
- MeSH terms
- Animals, Male, Mice, Inbred C57BL, Neoplasms, Phenotype, Spermatozoa, Testis, Tumor Suppressor Protein p53, Telomere, Apoptosis, Telomerase, Mice
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
A language model drew the claims from the paper on 2026-10-06, under version 3 of the claim definition (model deepseek-v4.1-flash, prompt extract-v3).
A second model labelled each claim on 2026-10-06 (model jev-1.13-20260917, definition revision gold-guideline-v0.2).
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
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