Cited by 31Microtubule and mitosis dynamicsGenomics and Chromatin DynamicsDNA Repair Mechanisms

The DOI matches its Crossref record; 2 of the 2 references it cites were found in a registry or an index.

Stable kinetochore‐microtubule attachment requires loop‐dependent Ndc80‐Ndc80 binding

Open access
Polley, Soumitra, Müschenborn, Helen, Terbeck, Melina, De Antoni, Anna, Vetter, Ingrid R, Dogterom, Marileen, Musacchio, Andrea, Volkov, Vladimir A, Huis in 't Veld, Pim J·2023·The EMBO Journal

Published version of a preprint

Abstract

During cell division, kinetochores link chromosomes to spindle microtubules. The Ndc80 complex, a crucial microtubule binder, populates each kinetochore with dozens of copies. Whether adjacent Ndc80 complexes cooperate to promote microtubule binding remains unclear. Here we demonstrate that the Ndc80 loop, a short sequence that interrupts the Ndc80 coiled-coil at a conserved position, folds into a more rigid structure than previously assumed and promotes direct interactions between full-length Ndc80 complexes on microtubules. Mutations in the loop impair these Ndc80-Ndc80 interactions, prevent the formation of force-resistant kinetochore-microtubule attachments, and cause cells to arrest in mitosis for hours. This arrest is not due to an inability to recruit the kinetochore-microtubule stabilizing SKA complex and cannot be overridden by mutations in the Ndc80 tail that strengthen microtubule attachment. Thus, loop-mediated organization of adjacent Ndc80 complexes is crucial for stable end-on kinetochore-microtubule attachment and spindle assembly checkpoint satisfaction.

Overview

The claims and statistics of this paper haven’t been checked yet.

Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.

A few free checks a day · papers already verified open instantly · results are public

Integrity signals

Registry

Crossref · 10.15252/embj.2022112504

Cites

2 of 2 references verified0 citation errors · 0 not checkable

Statistics

Verification

Claims the paper makes

The claims and statistics of this paper haven’t been checked yet.

Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.

A few free checks a day · papers already verified open instantly · results are public

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

The self-check needs the paper’s full text.

Citations

Cites

Flagged first, then by citations

Research works printed but not linked: 81

  1. Akiyoshi B, Sarangapani KK, Powers AF, Nelson CR, Reichow SL, Arellano‐Santoyo H, Gonen T, Ranish JA, Asbury CL, Biggins S (2010) Tension directly stabilizes reconstituted kinetochore‐microtubule attachments. Nature 468: 576–579 · DOI
  2. Alex A, Piano V, Polley S, Stuiver M, Voss S, Ciossani G, Overlack K, Voss B, Wohlgemuth S, Petrovic A et al (2019) Electroporated recombinant proteins as tools for in vivo functional complementation, imaging, and chemical biology. Elife 8: e48287 · DOI
  3. Alushin GM, Ramey VH, Pasqualato S, Ball DA, Grigorieff N, Musacchio A, Nogales E (2010) The Ndc80 kinetochore complex forms oligomeric arrays along microtubules. Nature 467: 805–810 · DOI
  4. Alushin GM, Musinipally V, Matson D, Tooley J, Stukenberg PT, Nogales E (2012) Multimodal microtubule binding by the Ndc80 kinetochore complex. Nat Struct Mol Biol 19: 1161–1167 · DOI
  5. Arlt C, Götze M, Ihling CH, Hage C, Schäfer M, Sinz A (2016) Integrated workflow for structural proteomics studies based on cross‐linking/mass spectrometry with an MS/MS cleavable cross‐linker. Anal Chem 88: 7930–7937 · DOI
  6. Boeckmann B, Marcet‐Houben M, Rees JA, Forslund K, Huerta‐Cepas J, Muffato M, Yilmaz P, Xenarios I, Bork P, Lewis SE et al (2015) Quest for orthologs entails quest for tree of life: in search of the gene stream. Genome Biol Evol 7: 1988–1999 · DOI
  7. Chakraborty M, Tarasovetc EV, Zaytsev AV, Godzi M, Figueiredo AC, Ataullakhanov FI, Grishchuk EL (2019) Microtubule end conversion mediated by motors and diffusing proteins with no intrinsic microtubule end‐binding activity. Nat Commun 10: 1673 · DOI
  8. Cheeseman IM, Chappie JS, Wilson‐Kubalek EM, Desai A (2006) The conserved KMN network constitutes the core microtubule‐binding site of the kinetochore. Cell 127: 983–997 · DOI
  9. Chen Y, Riley DJ, Chen PL, Lee WH (1997) HEC, a novel nuclear protein rich in leucine heptad repeats specifically involved in mitosis. Mol Cell Biol 17: 6049–6056 · DOI
  10. Chivers CE, Crozat E, Chu C, Moy VT, Sherratt DJ, Howarth M (2010) A streptavidin variant with slower biotin dissociation and increased mechanostability. Nat Methods 7: 391–393 · DOI
  11. Ciferri C, Pasqualato S, Screpanti E, Varetti G, Santaguida S, Dos Reis G, Maiolica A, Polka J, De Luca JG, De Wulf P et al (2008) Implications for kinetochore‐microtubule attachment from the structure of an engineered Ndc80 complex. Cell 133: 427–439 · DOI
  12. DeLuca JG, Gall WE, Ciferri C, Cimini D, Musacchio A, Salmon ED (2006) Kinetochore microtubule dynamics and attachment stability are regulated by Hec1. Cell 127: 969–982 · DOI
  13. DeLuca KF, Lens SM, DeLuca JG (2011) Temporal changes in Hec1 phosphorylation control kinetochore‐microtubule attachment stability during mitosis. J Cell Sci 124: 622–634 · DOI
  14. DeLuca KF, Meppelink A, Broad AJ, Mick JE, Peersen OB, Pektas S, Lens SMA, DeLuca JG (2018) Aurora A kinase phosphorylates Hec1 to regulate metaphase kinetochore‐microtubule dynamics. J Cell Biol 217: 163–177 · DOI
  15. Emsley P, Cowtan K (2004) Coot: model‐building tools for molecular graphics. Acta Crystallogr D Biol Crystallogr 60: 2126–2132 · DOI · Paper page
  16. Etemad B, Kuijt TEF, Kops GJPL (2015) Kinetochore‐microtubule attachment is sufficient to satisfy the human spindle assembly checkpoint. Nat Commun 6: 8987 · DOI
  17. Fairhead M, Veggiani G, Lever M, Yan J, Mesner D, Robinson CV, Dushek O, van der Merwe PA, Howarth M (2014) SpyAvidin hubs enable precise and ultrastable orthogonal nanoassembly. J Am Chem Soc 136: 12355–12363 · DOI
  18. Gell C, Bormuth V, Brouhard GJ, Cohen DN, Diez S, Friel CT, Helenius J, Nitzsche B, Petzold H, Ribbe J et al (2010) Microtubule dynamics reconstituted in vitro and imaged by single‐molecule fluorescence microscopy. Methods Cell Biol 95: 221–245 · DOI
  19. Goddard TD, Huang CC, Meng EC, Pettersen EF, Couch GS, Morris JH, Ferrin TE (2018) UCSF ChimeraX: meeting modern challenges in visualization and analysis. Protein Sci 27: 14–25 · DOI · Paper page
  20. Graham M, Combe C, Kolbowski L, Rappsilber J (2019) xiView: a common platform for the downstream analysis of crosslinking mass spectrometry data. bioRxivhttps://doi.org/10.1101/561829 [PREPRINT] · DOI
  21. Guimaraes GJ, Dong Y, McEwen BF, Deluca JG (2008) Kinetochore‐microtubule attachment relies on the disordered N‐terminal tail domain of Hec1. Curr Biol 18: 1778–1784 · DOI
  22. Hauf S, Cole RW, LaTerra S, Zimmer C, Schnapp G, Walter R, Heckel A, van Meel J, Rieder CL, Peters JM (2003) The small molecule Hesperadin reveals a role for Aurora B in correcting kinetochore‐microtubule attachment and in maintaining the spindle assembly checkpoint. J Cell Biol 161: 281–294 · DOI
  23. Helgeson LA, Zelter A, Riffle M, MacCoss MJ, Asbury CL, Davis TN (2018) Human Ska complex and Ndc80 complex interact to form a load‐bearing assembly that strengthens kinetochore‐microtubule attachments. Proc Natl Acad Sci USA 115: 2740–2745 · DOI
  24. Hirakawa H, Ishikawa S, Nagamune T (2015) Ca2+‐independent sortase‐A exhibits high selective protein ligation activity in the cytoplasm of Escherichia coli. Biotechnol J 10: 1487–1492 · DOI
  25. Hsu KS, Toda T (2011) Ndc80 internal loop interacts with Dis1/TOG to ensure proper kinetochore‐spindle attachment in fission yeast. Curr Biol 21: 214–220 · DOI
  26. Huis in 't Veld PJ, Jeganathan S, Petrovic A, Singh P, John J, Krenn V, Weissmann F, Bange T, Musacchio A (2016) Molecular basis of outer kinetochore assembly on CENP‐T. Elife 5: e21007 · DOI
  27. Huis in 't Veld PJ, Volkov VA, Stender ID, Musacchio A, Dogterom M (2019) Molecular determinants of the Ska‐Ndc80 interaction and their influence on microtubule tracking and force‐coupling. Elife 8: e49539 · DOI
  28. Hyman A, Drechsel D, Kellogg D, Salser S, Sawin K, Steffen P, Wordeman L, Mitchison T (1991) Preparation of modified tubulins. Methods Enzymol 196: 478–485 · DOI
  29. Janczyk PL, Skorupka KA, Tooley JG, Matson DR, Kestner CA, West T, Pornillos O, Stukenberg PT (2017) Mechanism of Ska recruitment by Ndc80 complexes to kinetochores. Dev Cell 41: 438–449 · DOI
  30. Jehl P, Manguy J, Shields DC, Higgins DG, Davey NE (2016) ProViz—a web‐based visualization tool to investigate the functional and evolutionary features of protein sequences. Nucleic Acids Res 44: W11–W15 · DOI
  31. Jenni S, Harrison SC (2018) Structure of the DASH/Dam1 complex shows its role at the yeast kinetochore‐microtubule interface. Science 360: 552–558 · DOI · Paper page
  32. Kapoor TM, Lampson MA, Hergert P, Cameron L, Cimini D, Salmon ED, McEwen BF, Khodjakov A (2006) Chromosomes can congress to the metaphase plate before biorientation. Science 311: 388–391 · DOI
  33. Kiewisz R, Fabig G, Conway W, Baum D, Needleman D, Müller‐Reichert T (2022) Three‐dimensional structure of kinetochore‐fibers in human mitotic spindles. Elife 11: e75459 · DOI
  34. Kim S, Yu H (2015) Multiple assembly mechanisms anchor the KMN spindle checkpoint platform at human mitotic kinetochores. J Cell Biol 208: 181–196 · DOI
  35. Krenn V, Musacchio A (2015) The Aurora B kinase in chromosome bi‐orientation and spindle checkpoint signaling. Front Oncol 5: 225 · DOI
  36. Kucharski TJ, Hards R, Vandal SE, Abad MA, Jeyaprakash AA, Kaye E, Al‐Rawi A, Ly T, Godek KM, Gerber SA et al (2022) Small changes in phospho‐occupancy at the kinetochore–microtubule interface drive mitotic fidelity. J Cell Biol 221: e202107107 · DOI
  37. Lampert F, Mieck C, Alushin GM, Nogales E, Westermann S (2013) Molecular requirements for the formation of a kinetochore‐microtubule interface by Dam1 and Ndc80 complexes. J Cell Biol 200: 21–30 · DOI
  38. Lampson MA, Grishchuk EL (2017) Mechanisms to avoid and correct erroneous kinetochore‐microtubule attachments. Biology (Basel) 6: 1
  39. Letunic I, Bork P (2021) Interactive Tree Of Life (iTOL) v5: an online tool for phylogenetic tree display and annotation. Nucleic Acids Res 49: W293–W296 · DOI
  40. Liebschner D, Afonine PV, Baker ML, Bunkóczi G, Chen VB, Croll TI, Hintze B, Hung L‐W, Jain S, McCoy AJ et al (2019) Macromolecular structure determination using X‐rays, neutrons and electrons: recent developments in Phenix. Acta Crystallogr D Struct Biol 75: 861–877 · DOI · Paper page
  41. Long AF, Udy DB, Dumont S (2017) Hec1 tail phosphorylation differentially regulates mammalian kinetochore coupling to polymerizing and depolymerizing microtubules. Curr Biol 27: 1692–1699 · DOI
  42. Magidson V, O'Connell CB, Lončarek J, Paul R, Mogilner A, Khodjakov A (2011) The spatial arrangement of chromosomes during prometaphase facilitates spindle assembly. Cell 146: 555–567 · DOI
  43. Maiolica A, Cittaro D, Borsotti D, Sennels L, Ciferri C, Tarricone C, Musacchio A, Rappsilber J (2007) Structural analysis of multiprotein complexes by cross‐linking, mass spectrometry, and database searching. Mol Cell Proteomics 6: 2200–2211 · DOI
  44. Maleki AN, Huis in 't Veld PJ, Akhmanova A, Dogterom M, Volkov VA (2023) Estimation of microtubule‐generated forces using a DNA origami nanospring. J Cell Sci 136: jcs260154
  45. Maure JF, Komoto S, Oku Y, Mino A, Pasqualato S, Natsume K, Clayton L, Musacchio A, Tanaka TU (2011) The Ndc80 loop region facilitates formation of kinetochore attachment to the dynamic microtubule plus end. Curr Biol 21: 207–213 · DOI
  46. McCleland ML, Gardner RD, Kallio MJ, Daum JR, Gorbsky GJ, Burke DJ, Stukenberg PT (2003) The highly conserved Ndc80 complex is required for kinetochore assembly, chromosome congression, and spindle checkpoint activity. Genes Dev 17: 101–114 · DOI
  47. Miller SA, Johnson ML, Stukenberg PT (2008) Kinetochore attachments require an interaction between unstructured tails on microtubules and Ndc80(Hec1). Curr Biol 18: 1785–1791 · DOI
  48. Monda JK, Cheeseman IM (2018) The kinetochore‐microtubule interface at a glance. J Cell Sci 131: jcs214577 · DOI
  49. Musacchio A, Desai A (2017) A molecular view of kinetochore assembly and function. Biology (Basel) 6: 1
  50. O'Toole E, Morphew M, McIntosh JR (2020) Electron tomography reveals aspects of spindle structure important for mechanical stability at metaphase. Mol Biol Cell 31: 184–195 · DOI
  51. Pan D, Brockmeyer A, Mueller F, Musacchio A, Bange T (2018) Simplified protocol for cross‐linking mass spectrometry using the MS‐cleavable cross‐linker DSBU with efficient cross‐link identification. Anal Chem 90: 10990–10999 · DOI
  52. Powers AF, Franck AD, Gestaut DR, Cooper J, Gracyzk B, Wei RR, Wordeman L, Davis TN, Asbury CL (2009) The Ndc80 kinetochore complex forms load‐bearing attachments to dynamic microtubule tips via biased diffusion. Cell 136: 865–875 · DOI
  53. Roscioli E, Germanova TE, Smith CA, Embacher PA, Erent M, Thompson AI, Burroughs NJ, McAinsh AD (2020) Ensemble‐Level organization of human kinetochores and evidence for distinct tension and attachment sensors. Cell Rep 31: 107535 · DOI
  54. Santaguida S, Tighe A, D'Alise AM, Taylor SS, Musacchio A (2010) Dissecting the role of MPS1 in chromosome biorientation and the spindle checkpoint through the small molecule inhibitor reversine. J Cell Biol 190: 73–87 · DOI
  55. Saurin AT (2018) Kinase and phosphatase cross‐talk at the kinetochore. Front Cell Dev Biol 6: 62 · DOI
  56. Scarborough EA, Davis TN, Asbury CL (2019) Tight bending of the Ndc80 complex provides intrinsic regulation of its binding to microtubules. Elife 8: e44489 · DOI
  57. Schindelin J, Arganda‐Carreras I, Frise E, Kaynig V, Longair M, Pietzsch T, Preibisch S, Rueden C, Saalfeld S, Schmid B et al (2012) Fiji: an open‐source platform for biological‐image analysis. Nat Methods 9: 676–682 · DOI · Paper page
  58. Shrestha RL, Draviam VM (2013) Lateral to end‐on conversion of chromosome‐microtubule attachment requires kinesins CENP‐E and MCAK. Curr Biol 23: 1514–1526 · DOI
  59. Shrestha RL, Conti D, Tamura N, Braun D, Ramalingam RA, Cieslinski K, Ries J, Draviam VM (2017) Aurora‐B kinase pathway controls the lateral to end‐on conversion of kinetochore‐microtubule attachments in human cells. Nat Commun 8: 150 · DOI
  60. Sundin LJ, Guimaraes GJ, Deluca JG (2011) The NDC80 complex proteins Nuf2 and Hec1 make distinct contributions to kinetochore‐microtubule attachment in mitosis. Mol Biol Cell 22: 759–768 · DOI
  61. Suzuki A, Badger BL, Salmon ED (2015) A quantitative description of Ndc80 complex linkage to human kinetochores. Nat Commun 6: 8161 · DOI
  62. Tanaka K, Mukae N, Dewar H, van Breugel M, James EK, Prescott AR, Antony C, Tanaka TU (2005) Molecular mechanisms of kinetochore capture by spindle microtubules. Nature 434: 987–994 · DOI · Paper page
  63. Tang G, Peng L, Baldwin PR, Mann DS, Jiang W, Rees I, Ludtke SJ (2007) EMAN2: an extensible image processing suite for electron microscopy. J Struct Biol 157: 38–46 · DOI
  64. Tang NH, Takada H, Hsu KS, Toda T (2013) The internal loop of fission yeast Ndc80 binds Alp7/TACC‐Alp14/TOG and ensures proper chromosome attachment. Mol Biol Cell 24: 1122–1133 · DOI
  65. Tauchman EC, Boehm FJ, DeLuca JG (2015) Stable kinetochore–microtubule attachment is sufficient to silence the spindle assembly checkpoint in human cells. Nat Commun 6: 10036 · DOI
  66. Tooley JG, Miller SA, Stukenberg PT (2011) The Ndc80 complex uses a tripartite attachment point to couple microtubule depolymerization to chromosome movement. Mol Biol Cell 22: 1217–1226 · DOI
  67. Valverde R, Ingram J, Harrison SC (2016) Conserved tetramer junction in the kinetochore Ndc80 complex. Cell Rep 17: 1915–1922 · DOI
  68. Varma D, Chandrasekaran S, Sundin LJ, Reidy KT, Wan X, Chasse DA, Nevis KR, DeLuca JG, Salmon ED, Cook JG (2012) Recruitment of the human Cdt1 replication licensing protein by the loop domain of Hec1 is required for stable kinetochore‐microtubule attachment. Nat Cell Biol 14: 593–603 · DOI
  69. Volkov VA, Huis in 't Veld PJ, Dogterom M, Musacchio A (2018) Multivalency of NDC80 in the outer kinetochore is essential to track shortening microtubules and generate forces. Elife 7: e36764 · DOI
  70. Wang HW, Long S, Ciferri C, Westermann S, Drubin D, Barnes G, Nogales E (2008) Architecture and flexibility of the yeast Ndc80 kinetochore complex. J Mol Biol 383: 894–903 · DOI
  71. Waterhouse AM, Procter JB, Martin DM, Clamp M, Barton GJ (2009) Jalview Version 2—a multiple sequence alignment editor and analysis workbench. Bioinformatics 25: 1189–1191 · DOI
  72. Wei RR, Al‐Bassam J, Harrison SC (2007) The Ndc80/HEC1 complex is a contact point for kinetochore‐microtubule attachment. Nat Struct Mol Biol 14: 54–59 · DOI
  73. Weissmann F, Petzold G, VanderLinden R, Huis in 't Veld PJ, Brown NG, Lampert F, Westermann S, Stark H, Schulman BA, Peters JM (2016) biGBac enables rapid gene assembly for the expression of large multisubunit protein complexes. Proc Natl Acad Sci USA 113: E2564–E2569
  74. Wigge PA, Jensen ON, Holmes S, Souès S, Mann M, Kilmartin JV (1998) Analysis of the Saccharomyces spindle pole by matrix‐assisted laser desorption/ionization (MALDI) mass spectrometry. J Cell Biol 141: 967–977 · DOI
  75. Wimbish RT, DeLuca KF, Mick JE, Himes J, Jiménez‐Sánchez I, Jeyaprakash AA, DeLuca JG (2020) The Hec1/Ndc80 tail domain is required for force generation at kinetochores, but is dispensable for kinetochore‐microtubule attachment formation and Ska complex recruitment. Mol Biol Cell 31: 1453–1473 · DOI
  76. Yoo TY, Choi J‐M, Conway W, Yu C‐H, Pappu RV, Needleman DJ (2018) Measuring NDC80 binding reveals the molecular basis of tension‐dependent kinetochore‐microtubule attachments. Elife 7: e36392
  77. Zahm JA, Jenni S, Harrison SC (2023) Structure of the Ndc80 complex and its interactions at the yeast kinetochore–microtubule interface. Open Biol 13: 220378
  78. Zaytsev AV, Mick JE, Maslennikov E, Nikashin B, DeLuca JG, Grishchuk EL (2015) Multisite phosphorylation of the NDC80 complex gradually tunes its microtubule‐binding affinity. Mol Biol Cell 26: 1829–1844 · DOI
  79. Zhang G, Kelstrup CD, Hu XW, Kaas Hansen MJ, Singleton MR, Olsen JV, Nilsson J (2012) The Ndc80 internal loop is required for recruitment of the Ska complex to establish end‐on microtubule attachment to kinetochores. J Cell Sci 125: 3243–3253
  80. Zhang Q, Sivakumar S, Chen Y, Gao H, Yang L, Yuan Z, Yu H, Liu H (2017) Ska3 phosphorylated by Cdk1 binds Ndc80 and recruits Ska to kinetochores to promote mitotic progression. Curr Biol 27: 1477–1484 · DOI
  81. Zhu T, Dou Z, Qin B, Jin C, Wang X, Xu L, Zhaoyang W, Zhu L, Liu F, Gao X et al (2013) Phosphorylation of microtubule‐binding protein Hec1 by mitotic kinase Aurora B specifies spindle checkpoint kinase Mps1 signaling at the kinetochore. J Biol Chem 288: 36149–36159 · DOI

Metadata

Authors
Polley, S., Müschenborn, H., Terbeck, M., De Antoni, A., Vetter, I. R., Dogterom, M., Musacchio, A., Volkov, V. A., & Huis in 't Veld, P. J.
Affiliations
  • Max Planck Institute of Molecular Physiology (DE)
  • Delft University of Technology (NL)
  • Queen Mary University of London (GB)
Corresponding author
  • Musacchio, Andrea
  • Volkov, Vladimir A
Venue
The EMBO Journal
Published
2023-05-19
Volume
42
Issue
13
Pages
e112504-e112504
ISSN
  • 0261-4189 (ISSN-L)
  • 1460-2075
OpenAlex
W4377094368
Cited by
31
References
2
Citations by year
2026: 8 · 2025: 6 · 2024: 8 · 2023: 9
FWCI
4.49
Type
journal article
Language
en
Open access
Locations
  • The EMBO Journal (publishedVersion, cc-by-nc-nd)
  • bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by-nc)
  • PubMed Central (publishedVersion, cc-by-nc-nd)
  • PubMed (publishedVersion)
  • Data Archiving and Networked Services (DANS) (submittedVersion, cc-by-nc-nd)
  • Queen Mary Research Online (Queen Mary University of London) (submittedVersion, cc-by-nc-nd)
  • Research Repository (Delft University of Technology) (submittedVersion)
Funders
  • EC | European Research Council (609822, 951430)
  • QMUL Startup Grant (SBC8VOL2)
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Microtubule and mitosis dynamics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Genomics and Chromatin Dynamics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › DNA Repair Mechanisms
Keywords
Ndc80 complex, kinetochore-microtubule attachment, coiled coil, microtubule binding, spindle assembly checkpoint, mitotic arrest, chromosome segregation, mitosis
MeSH terms
Animals, Microtubule-Associated Proteins, Microtubules, Mitosis, Protein Binding, Kinetochores, Chromosome Segregation

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.

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.

What counts as a claim ›How we measure confidence ›How we check this ›

Are you an author, or do you see a problem with this page? Request a review