External Citation Audit
Research Question
Which non-ingested papers are cited inline in entity, concept, and method pages — and are those citations accurate?
Background
Wiki Rule 6: “Everything in entity/concept/method pages should trace back to a source page.” A deep lint on 2026-05-08 found ~80 inline references to ~54 unique external papers across ~25 wiki pages. These references were introduced during ingests when claims from ingested sources included secondary citations. This audit page catalogs every instance so the curator can verify accuracy and decide which papers to ingest.
Status: Pending curator review. Once verified, external citations in wiki pages should be rewritten to attribute claims solely to ingested source pages.
External Papers Cited
Each entry lists: citation as it appears, DOI (where found), the ingested source that cited it, and all wiki pages where it appears.
1. William et al. 2002, Science
Topic: SHM can occur outside germinal centres (murine EF)
DOI: 10.1126/science.1073924
Cited via: Wei2007 - DN Memory B Cells in SLE, Sanz2025 - Human Atypical B Cells Overview
Wiki locations:
- Double-Negative B Cell — line 51 (bare: “cited but not yet ingested”)
- Extrafollicular Response — line 19
- Germinal Center — line 22
- Somatic Hypermutation — lines 21, 37
2. Wirths & Lanzavecchia 2005, Eur J Immunol
Topic: CD27⁻ PBL memory population identified by R123 extrusion
DOI: 10.1002/eji.200535364
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- Double-Negative B Cell — line 62 (bare: “cited but not yet ingested”)
3. Anolik et al. 2003, Eur J Immunol
Topic: CD20 downregulation upon B cell activation via lipid rafts
DOI: 10.1002/eji.200323515
Cited via: not formally via any ingested source (separate Anolik paper from the ingested 2004)
Wiki locations:
- CD20 — line 21 (bare: “cited but not yet ingested”)
4. Qi et al. 2006, Science
Topic: Antigen-bearing DCs activate B cells in extrafollicular zones
DOI: 10.1126/science.1125703
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- Extrafollicular Response — line 20
5. Litinskiy et al. 2002, Nat Immunol
Topic: DCs induce IgG/IgA class switching via BLyS/APRIL without CD40
DOI: 10.1038/ni829
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- Extrafollicular Response — line 21
- Class Switch Recombination — line 18
6. Cappione et al. 2005, JCI
Topic: Defective GC exclusion of autoreactive B cells in SLE
DOI: 10.1172/JCI24179
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- Germinal Center — line 21
7. Klein et al. 1998, J Exp Med
Topic: CD27 as marker of human memory B cells
DOI: 10.1084/jem.188.9.1679
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- CD27 — line 19
- Memory B Cell — line 19
8. Weller et al. 2001
Topic: IgM/IgD CD27⁺ B cells develop via GC-independent, CD40-independent pathways
DOI: 10.1073/pnas.021600598 (verify)
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- Memory B Cell — line 20
9. Maclennan 1994, Annu Rev Immunol
Topic: Germinal centres (foundational review)
DOI: 10.1146/annurev.immunol.12.1.117
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- Germinal Center — line 19
10. Grewal & Flavell 1998
Topic: CD40–CD154 signalling
DOI: 10.1146/annurev.immunol.16.1.111 (verify)
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- Germinal Center — line 19
11. Toellner et al. 2002, J Exp Med
Topic: T cell-independent GCs produce low-level SHM
DOI: 10.1084/jem.20011112
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- Germinal Center — line 23
12. Kobata et al. 1995
Topic: CD27–CD70 interactions regulate B cell activation DOI: not retrieved — search: Kobata CD27 CD70 B cell 1995 Cited via: Wei2007 - DN Memory B Cells in SLE Wiki locations:
- CD27 — line 21
13. Jacquot et al. 1997
Topic: CD27 and plasma cell differentiation DOI: not retrieved — search: Jacquot CD27 plasma cell 1997 Cited via: Wei2007 - DN Memory B Cells in SLE Wiki locations:
- CD27 — line 21
14. Ehrhardt et al. 2005, J Exp Med
Topic: FcRH4 marks tissue-based memory B cells
DOI: 10.1084/jem.20050879
Cited via: Wei2007 - DN Memory B Cells in SLE
Wiki locations:
- FcRH4 — line 21, 29
15. Masilamani et al. 2003, Eur J Immunol
Topic: CD21 (CR2) shedding upon B cell activation
DOI: 10.1002/eji.200323843
Cited via: Tipton2015 - ASC Diversity and Origin in SLE
Wiki locations:
16. Wehr et al. 2004
Topic: CD21^lo activated B cell population in SLE DOI: not confirmed — search: Wehr CD21 low B cells primary immunodeficiency 2004 Cited via: Tipton2015 - ASC Diversity and Origin in SLE Wiki locations:
17. Hanten et al. 2008
Topic: CD23 downregulation by IFN-α DOI: not retrieved — search: Hanten CD23 IFN-alpha 2008 Cited via: Tipton2015 - ASC Diversity and Origin in SLE Wiki locations:
- CD23 — line 18
18. Delespesse et al. 1989
Topic: CD23 regulation DOI: not retrieved — search: Delespesse CD23 IgE receptor 1989 Cited via: Tipton2015 - ASC Diversity and Origin in SLE Wiki locations:
- CD23 — line 18
19. Dogan et al. 2009, J Exp Med
Topic: IgM memory as GC-independent first memory layer
DOI: 10.1084/jem.20091087
Cited via: Tipton2015 - ASC Diversity and Origin in SLE
Wiki locations:
- Germinal Center — line 28
- Memory B Cell — line 47
- IgM — line 20
20. Dominguez et al. 2015, Nat Immunol
Topic: T-bet and ZEB2 cooperate for terminal effector CD8⁺ T cell differentiation
DOI: 10.1038/ni.3248
Cited via: Jenks2018 - DN2 B Cells and EF Pathway in SLE
Wiki locations:
21. Rubtsova et al. 2015, J Immunol
Topic: Murine T-bet⁺ age-associated B cells
DOI: 10.4049/jimmunol.1501209
Cited via: Jenks2018 - DN2 B Cells and EF Pathway in SLE
Wiki locations:
- CD11c — line 20
22. Rubtsova et al. 2017, JCI
Topic: T-bet⁺ B cells localise to T-B border and drive autoimmunity
DOI: 10.1172/JCI91250
Cited via: Jenks2018 - DN2 B Cells and EF Pathway in SLE
Wiki locations:
- T-bet — line 20
23. Russell et al. 2015
Topic: ETS1 deficiency leads to extrafollicular autoreactive B cells DOI: not confirmed — search: Russell ETS1 extrafollicular autoreactive B cells 2015 Cited via: Jenks2018 - DN2 B Cells and EF Pathway in SLE Wiki locations:
- IRF4 — line 19
24. Scharer et al. 2016, Sci Rep
Topic: ATAC-seq protocol for B cells
DOI: 10.1038/srep27030
Cited via: Jenks2018 - DN2 B Cells and EF Pathway in SLE
Wiki locations:
- ATAC-seq — line 18
25. Kang et al. 2024
Topic: CD27 downregulation by CD70, TLR, cytokines DOI: not retrieved — search: Kang CD27 downregulation B cell 2024 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- CD27 — line 28
26. Yang et al. 2022
Topic: T-bet not absolutely required for ABC/CD11c⁺ B cells DOI: not retrieved — search: Yang T-bet dispensable ABC B cell 2022 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- T-bet — line 23
27. Du et al. 2019
Topic: CD11c inducible without IFN-γ or T-bet DOI: not retrieved — search: Du CD11c B cell T-bet independent 2019 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- T-bet — line 23
28. Nellore et al. 2023
Topic: T-bet⁺/FcRL5⁺ memory partition; influenza vaccination DOI: not confirmed — search: Nellore T-bet FcRL5 memory influenza 2023 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- T-bet — line 25
- FCRL5 — line 22
- Memory B Cell — line 35
29. Portugal et al. 2015, eLife
Topic: FCRL5⁺ atypical memory B cells in malaria
DOI: 10.7554/eLife.07218
Cited via: Jenks2018 - DN2 B Cells and EF Pathway in SLE
Wiki locations:
- FCRL5 — line 18
30. Brown et al. 2022, Nature
Topic: TLR7 gain-of-function mutation causes human SLE
DOI: 10.1038/s41586-022-04642-z
Cited via: Sanz2025 - Human Atypical B Cells Overview
Wiki locations:
- TLR7 — line 21
- Germinal Center — line 36
- Extrafollicular Response — line 40
31. Zumaquero et al. 2019
Topic: TLR7 obligatory role in naive B cell → DN2 differentiation
DOI: 10.1038/s41467-019-11290-x (verify — Nat Commun)
Cited via: Sanz2025 - Human Atypical B Cells Overview
Wiki locations:
- TLR7 — line 22
32. Zhu et al. 2024
Topic: CD21lo cells have increased TLR7 sensitivity DOI: not retrieved — search: Zhu CD21 low TLR7 sensitivity B cell 2024 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
33. Dai et al. 2024, Nature
Topic: ZEB2 drives B cell autoimmunity; ZEB2 represses Mef2b
DOI: 10.1038/s41586-024-07613-w (verify)
Cited via: Sanz2025 - Human Atypical B Cells Overview
Wiki locations:
- ZEB2 — lines 21, 22
- Germinal Center — line 35
- Extrafollicular Response — line 41
34. Gao et al. 2024
Topic: ABC sustain GC responses; Mef2b repression DOI: not retrieved — search: Gao ABC B cell germinal center 2024 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- Germinal Center — lines 35, 37
35. Zhang et al. 2019
Topic: ABC as APCs; abnormal TFH regulation DOI: not retrieved — search: Zhang ABC APC TFH B cell 2019 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- Germinal Center — line 37
36. Jenks et al. 2021
Topic: B cell endotypes in SLE (EF vs. GC/memory clusters)
DOI: 10.1172/JCI150888 (verify)
Cited via: Sanz2025 - Human Atypical B Cells Overview
Wiki locations:
- Extrafollicular Response — line 37
37. Faliti et al. 2024
Topic: Durable DN2/DN3 cells >1 year post-SARS-CoV-2 vaccination DOI: not confirmed — search: Faliti DN2 DN3 SARS-CoV-2 vaccination memory 2024 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- DN3 B Cell — line 23
- Memory B Cell — line 34
- Extrafollicular Response — line 37
38. Ma et al. 2024
Topic: EF responses in cancer; worse clinical outcomes DOI: not retrieved — search: Ma extrafollicular B cell cancer immunotherapy 2024 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- Extrafollicular Response — line 38
39. Kaneko et al. 2020, Cell
Topic: Loss of Bcl-6⁺ Tfh and germinal centers in fatal COVID-19
DOI: 10.1016/j.cell.2020.08.025
Cited via: Woodruff2020 - EF B Cell Responses in COVID-19
Wiki locations:
- Extrafollicular Response — line 48
- Germinal Center — line 47
40. Ho et al. 2008
Topic: IFN-γ synergizes with TLR to enhance ATF3 DOI: not retrieved — search: Ho IFN-gamma TLR ATF3 2008 Cited via: Scharer2019 - Epigenetic Programming in SLE B Cells Wiki locations:
- ATF3 — line 23
41. Gomez-Martin et al. 2010
Topic: EGR and autoimmunity; SLE susceptibility DOI: not retrieved — search: Gomez-Martin EGR autoimmunity SLE 2010 Cited via: Scharer2019 - Epigenetic Programming in SLE B Cells Wiki locations:
- EGR — line 24
42. Oh et al. 2015
Topic: EGR in B cell biology DOI: not retrieved — search: Oh EGR B cell 2015 Cited via: Scharer2019 - Epigenetic Programming in SLE B Cells Wiki locations:
- EGR — line 24
43. Gururajan et al. 2008
Topic: EGR and plasma cell differentiation DOI: not retrieved — search: Gururajan EGR plasma cell 2008 Cited via: Scharer2019 - Epigenetic Programming in SLE B Cells Wiki locations:
- EGR — line 24
44. Myouzen et al. 2010
Topic: EGR2 polymorphisms in SLE susceptibility DOI: not retrieved — search: Myouzen EGR2 SLE polymorphism 2010 Cited via: Scharer2019 - Epigenetic Programming in SLE B Cells Wiki locations:
- EGR — line 24
45. Wong et al. 2020, Immunity
Topic: Affinity-restricted memory B cells dominate flavivirus recall
DOI: 10.1016/j.immuni.2020.09.001
Cited via: Singh2026 - DENV-Specific Memory B Cell Subsets
Wiki locations:
- Somatic Hypermutation — line 44
- Singh2026 source page — line 117
46. Turner et al. 2021, Nature
Topic: Prolonged GC reactions post-SARS-CoV-2 vaccination (~8 months)
DOI: 10.1038/s41586-021-03647-4
Cited via: Singh2026 - DENV-Specific Memory B Cell Subsets
Wiki locations:
- Germinal Center — line 50
- Singh2026 source page — line 126
47. Arce et al. 2001, J Immunol
Topic: Pre-GC (Bm2’) cells expanded in children with SLE
DOI: 10.4049/jimmunol.167.4.2361
Cited via: Anolik2004 - Rituximab and B Cell Abnormalities in SLE
Wiki locations:
- Bm Classification — line 21
48. Woodruff et al. 2020, 2022
Topic: Naïve-derived DN2 in COVID-19; self-limited EF autoreactivity
DOI: 2020: 10.1038/s41590-020-00814-z (ingested as Woodruff2020); 2022: not confirmed — search: Woodruff self-limited autoreactivity SARS-CoV-2 B cells 2022
Cited via: Sanz2025 - Human Atypical B Cells Overview
Wiki locations:
- DN2 B Cell — line 32
- DN3 B Cell — lines 19, 21
49. Allard-Chamard et al. 2023, Cell Rep
Topic: DN3 tissue infiltration in fibrosis and COVID-19
DOI: 10.1016/j.celrep.2023.112630
Cited via: Sanz2025 - Human Atypical B Cells Overview
Wiki locations:
- DN3 B Cell — line 22
50. Szelinski et al. 2022
Topic: CD27⁻ CXCR5⁻ CD19⁻low pre-plasmablast population in SLE DOI: not retrieved — search: Szelinski DN3 pre-plasmablast SLE CD27 CXCR5 2022 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- DN3 B Cell — line 20
51. Csomos et al. 2022
Topic: Partial RAG deficiency; T-bet⁺ B cells with autoreactivity DOI: not confirmed — search: Csomos RAG deficiency T-bet B cells autoreactivity 2022 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- Somatic Hypermutation — line 39
52. Holla et al. 2019
Topic: Malaria AtB respond to membrane-associated antigens DOI: not confirmed — search: Holla malaria atypical B cell membrane antigen 2019 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- DN2 B Cell — line 52
53. MacLennan et al. 2003
Topic: EF SHM and CSR evidence
DOI: 10.1111/j.0105-2896.2003.00066.x (verify — Immunol Rev)
Cited via: Sanz2025 - Human Atypical B Cells Overview
Wiki locations:
- Somatic Hypermutation — line 37
54. Wing et al. 2023
Topic: DN2 cells in rheumatoid synovium as main ASC precursor DOI: not retrieved — search: Wing DN2 rheumatoid synovium ASC precursor 2023 Cited via: Sanz2025 - Human Atypical B Cells Overview Wiki locations:
- DN2 B Cell — line 31
- Plasmablast — line 32
Summary
| Category | Count |
|---|---|
| Unique external papers | 54 |
| Total inline citations across wiki pages | ~80 |
| Wiki pages affected | ~25 |
| DOIs confirmed | 26 |
| DOIs needing verification | 7 |
| DOIs not retrieved | 21 |
Top-priority verifications (bare external cites — Pattern A):
- William et al. 2002 —
10.1126/science.1073924— in Double-Negative B Cell - Wirths & Lanzavecchia 2005 —
10.1002/eji.200535364— in Double-Negative B Cell - Anolik et al. 2003 —
10.1002/eji.200323515— in CD20
Open Questions
- Should verified external citations be rewritten to attribute solely to the ingested source, or should the external papers be ingested?
- Are any of these external papers high-priority for ingestion (e.g., William et al. 2002 for murine EF SHM)?
Sources Used
All 8 ingested source pages were scanned. External citations traced to their ingested source of origin.