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  1. .gitattributes +73 -0
  2. .gitignore +51 -0
  3. README.md +120 -0
  4. expression/SOURCES.md +459 -0
  5. expression/context_synonyms.tsv +449 -0
  6. expression/protein_abundance_mmu.tsv.gz +3 -0
  7. expression/reference_expression.tsv.gz +3 -0
  8. expression/reference_expression_mmu.tsv.gz +3 -0
  9. expression/reference_expression_toil.parquet +3 -0
  10. expression/toil_floors.tsv +89 -0
  11. expression/toil_floors_mmu.tsv +72 -0
  12. expression/toil_matrix.npz +3 -0
  13. expression/toil_matrix_mmu.npz +3 -0
  14. expression/tumor_expression_mmu.tsv.gz +3 -0
  15. hla_loh/SOURCES.md +134 -0
  16. hla_loh/do_to_tcga_barcode.tsv +712 -0
  17. hla_loh/escape_genes.parquet +3 -0
  18. hla_loh/gie_allele.parquet +3 -0
  19. hla_loh/gie_sample.parquet +3 -0
  20. hla_loh/harmonize.py +177 -0
  21. hla_loh/hla_typing_callers.parquet +3 -0
  22. hla_loh/perallele_tcga_loh.parquet +3 -0
  23. hla_loh/sample_metadata.parquet +3 -0
  24. homology/HOM_MouseHumanSequence.rpt.gz +3 -0
  25. homology/SOURCES.md +50 -0
  26. icb_cohorts/COHORTS.md +77 -0
  27. icb_cohorts/SOURCES.md +45 -0
  28. icb_cohorts/harmonize_typing.py +199 -0
  29. icb_cohorts/harmonized/manifest.tsv +7 -0
  30. icb_cohorts/harmonized/typing.parquet +3 -0
  31. immunogenicity/SOURCES.md +726 -0
  32. immunogenicity/chowell_iedb_full.tsv.gz +3 -0
  33. immunogenicity/chowell_iedb_full_matched.tsv.gz +3 -0
  34. immunogenicity/chowell_rebuilt.tsv.gz +3 -0
  35. immunogenicity/chowell_rebuilt_hla_matched.tsv.gz +3 -0
  36. immunogenicity/corpus_iedb_mhc2.log +24 -0
  37. immunogenicity/corpus_iedb_mhc2.parquet +3 -0
  38. immunogenicity/hla_pop_freqs.tsv.gz +3 -0
  39. immunogenicity/iedb_labeled.tsv.gz +3 -0
  40. immunogenicity/immunogenicity_legacy_arms.tsv.gz +3 -0
  41. immunogenicity/kesmir_rebuilt.tsv.gz +3 -0
  42. immunogenicity/kesmir_rebuilt_hla_matched.tsv.gz +3 -0
  43. immunogenicity/neoag_tcell_balanced_mhc2_human.tsv.gz +3 -0
  44. immunogenicity/neoag_tcell_balanced_mhc2_mouse.tsv.gz +3 -0
  45. immunogenicity/neoprecis_anchor_triplets.parquet +3 -0
  46. immunogenicity/pathogen_tcell_balanced_mhc1_human.tsv.gz +3 -0
  47. immunogenicity/pathogen_tcell_balanced_mhc1_mouse.tsv.gz +3 -0
  48. immunogenicity/pathogen_tcell_balanced_mhc2_human.tsv.gz +3 -0
  49. immunogenicity/pathogen_tcell_balanced_mhc2_mouse.tsv.gz +3 -0
  50. immunogenicity/pathogen_tcell_mhc1_human.tsv.gz +3 -0
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+ *.onnx filter=lfs diff=lfs merge=lfs -text
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+ *.ot filter=lfs diff=lfs merge=lfs -text
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+ *.parquet filter=lfs diff=lfs merge=lfs -text
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+ *.pb filter=lfs diff=lfs merge=lfs -text
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+ *.pickle filter=lfs diff=lfs merge=lfs -text
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+ *.pkl filter=lfs diff=lfs merge=lfs -text
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+ *.pt filter=lfs diff=lfs merge=lfs -text
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+ *.pth filter=lfs diff=lfs merge=lfs -text
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+ *.rar filter=lfs diff=lfs merge=lfs -text
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+ *.safetensors filter=lfs diff=lfs merge=lfs -text
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+ saved_model/**/* filter=lfs diff=lfs merge=lfs -text
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+ *.tar.* filter=lfs diff=lfs merge=lfs -text
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+ *.tar filter=lfs diff=lfs merge=lfs -text
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+ *.tflite filter=lfs diff=lfs merge=lfs -text
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+ *.zst filter=lfs diff=lfs merge=lfs -text
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+ *tfevents* filter=lfs diff=lfs merge=lfs -text
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+ # Audio files - uncompressed
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+ *.pcm filter=lfs diff=lfs merge=lfs -text
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+ *.sam filter=lfs diff=lfs merge=lfs -text
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+ *.raw filter=lfs diff=lfs merge=lfs -text
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+ # Audio files - compressed
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+ *.aac filter=lfs diff=lfs merge=lfs -text
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+ *.flac filter=lfs diff=lfs merge=lfs -text
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+ *.mp3 filter=lfs diff=lfs merge=lfs -text
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+ *.ogg filter=lfs diff=lfs merge=lfs -text
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+ *.wav filter=lfs diff=lfs merge=lfs -text
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+ # Image files - uncompressed
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+ *.bmp filter=lfs diff=lfs merge=lfs -text
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+ *.gif filter=lfs diff=lfs merge=lfs -text
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+ *.png filter=lfs diff=lfs merge=lfs -text
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+ *.tiff filter=lfs diff=lfs merge=lfs -text
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+ # Image files - compressed
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+ *.jpg filter=lfs diff=lfs merge=lfs -text
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+ *.jpeg filter=lfs diff=lfs merge=lfs -text
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+ *.webp filter=lfs diff=lfs merge=lfs -text
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+ # Video files - compressed
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+ *.mp4 filter=lfs diff=lfs merge=lfs -text
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+ *.webm filter=lfs diff=lfs merge=lfs -text
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+ *.tsv.gz filter=lfs diff=lfs merge=lfs -text
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+ *.idx filter=lfs diff=lfs merge=lfs -text
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+
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+ # Binary/large deposit formats that the HuggingFace default set above does not cover. Without these
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+ # a future .xlsx supplement or an uncompressed .fasta lands as a PLAIN BLOB in a dataset repo --
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+ # the mirror-image of the bug that once let a shipped .fasta escape a *.fasta ignore rule by its
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+ # extension. Nothing tracked today is affected: 0 .xlsx and 0 bare .fasta are tracked.
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+ # *.tsv and *.csv are deliberately NOT here: five small tables are tracked as plain blobs on
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+ # purpose (context_synonyms.tsv, toil_floors{,_mmu}.tsv, harmonized/manifest.tsv, summary.tsv),
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+ # and an LFS pattern would convert a 402-byte manifest for nothing. Compressed *.tsv.gz is already
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+ # covered above.
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+ *.xlsx filter=lfs diff=lfs merge=lfs -text
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+ *.fasta filter=lfs diff=lfs merge=lfs -text
.gitignore ADDED
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+ dump/
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+ src/
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+ .claude/
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+ .DS_Store
5
+ **/.DS_Store
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+
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+ # raw downloads (never pushed)
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+ raw/
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+
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+ # MixMHCpred3 comparison material (never pushed). Two of the four files are publisher
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+ # supplementary .xlsx — copyrighted, not ours to redistribute — and the directory's only
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+ # provenance is three bare URLs in benchmark/mixmhcpred3_ref.md, which does not meet the
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+ # SOURCES.md standard the rest of this repo now holds. Kept on disk, off the Hub.
14
+ benchmark/
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+
16
+ private/
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+ # NB: tcga/ is PUBLIC (neoantigen predictions derived from open-access TCGA somatic +
18
+ # expression data; per-barcode HLA/neoantigens as published by TCIA / Thorsson 2018).
19
+
20
+ # internal working table, not published
21
+ summary_all_datasets.tsv
22
+
23
+ # whole-proteome window indexes: derived, rebuilt on demand by mhcmatch, never published.
24
+ # Eight (species, length) cells were 17 GB -- 41% of this dataset -- against a 55 MB FASTA that
25
+ # reproduces any one of them in seconds, and fetching one measured SLOWER than building it
26
+ # (mouse L=9: 1.49 GB in 47.0 s unauthenticated, against 25.0 s to build). Withdrawn 2026-09-06.
27
+ proteome_index/
28
+
29
+ # ICB cohort assembly, staged 2026-09-10. Our harmonised tables under icb_cohorts/harmonized/
30
+ # are published (typing.parquet and manifest.tsv -- our own six-allele genotypes over public
31
+ # typings, and a source->count manifest). The publisher supplementary workbooks under
32
+ # icb_cohorts/raw/ are copyrighted and not ours to redistribute; they stay ignored permanently.
33
+ icb_cohorts/raw/
34
+
35
+ # NB: hla_loh/ is PUBLIC and tracked from 2026-09-16. It holds per-allele HLA-I loss-of-
36
+ # heterozygosity over TCGA/PCAWG and Hartwig, derived from open-access data exactly as tcga/ is:
37
+ # ICGC donor ids and TCGA barcodes are public pseudonymous research identifiers, and PCAWG's
38
+ # `donor_wgs_exclusion_white_gray` is its published QC whitelist flag. An earlier rule here called
39
+ # the directory a re-identification risk; that was wrong and is withdrawn. What genuinely may not
40
+ # ship is the three copyrighted Springer Nature workbooks, so they -- and the third-party PCAWG
41
+ # sample sheet -- live in `raw/hla_loh/` under the `raw/` rule above, and `hla_loh/harmonize.py`
42
+ # rebuilds the published parquets from them. Facts are not copyrightable; a publisher's workbook is.
43
+
44
+ # The mimicry k-mer cache. mhcmatch WRITES this itself (`mimicry.py` under
45
+ # $MHCMATCH_CALIBRATION_CACHE), keyed by a blake2b of the schema epoch, class, species, lengths and
46
+ # each deposit's name and size -- so it is derived data that any consumer regenerates on demand, and
47
+ # `docs/cli/environment.rst` says in as many words that it is safe to delete. 2.07 GB across 50
48
+ # files (two cache cells, human and mouse), tracked and pushed with no SOURCES.md entry. Same call as the 17 GB `proteome_index/` withdrawal in 2f76f91, for the same
49
+ # reason. NOTE: this rule stops it being re-added; the 50 files already tracked need a
50
+ # `git rm --cached`, which is a push to the Hub and so is the author's to make.
51
+ mimicry_refs/
README.md ADDED
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1
+ ---
2
+ pretty_name: pMHC Data Compendium
3
+ license: other
4
+ license_name: mixed-upstream
5
+ license_link: https://huggingface.co/datasets/isalgo/pmhc_data#licensing
6
+ language:
7
+ - en
8
+ tags:
9
+ - immunology
10
+ - immunopeptidomics
11
+ - neoantigen
12
+ - mhc
13
+ - hla
14
+ - t-cell
15
+ - cancer-vaccine
16
+ - proteomics
17
+ size_categories:
18
+ - 10M<n<100M
19
+ ---
20
+
21
+ # pMHC Data Compendium
22
+
23
+ Harmonized peptide–MHC data for neoantigen scoring: presentation, immunogenicity labels,
24
+ self/foreign reference peptidomes, expression, and clinical-outcome cohorts. Human and mouse.
25
+
26
+ Every table here is derived from public sources. Per-file provenance, filtering rules, verified
27
+ PubMed citations and row-level audits live in the `SOURCES.md` of each directory — that is the
28
+ authoritative record, not this page.
29
+
30
+ ## Contents
31
+
32
+ | Directory | What is in it |
33
+ |---|---|
34
+ | [`pmhc/`](pmhc) | Epitope–MHC binding records — the presentation panel (`pmhc_full`, and `pmhc_shortlist` at ≥2 references) |
35
+ | [`immunogenicity/`](immunogenicity) | T-cell immunogenicity corpora: immunogenic vs **self** ligands and vs **non-self**, each with an HLA-matched variant; labelled set; HLA population frequencies |
36
+ | [`neoantigens/`](neoantigens) | Tested and candidate neoantigens — minigene, epitope-level and per-mutation screens, human and mouse |
37
+ | [`ligandome/`](ligandome) | Foreign and tissue reference peptidomes — viral ligands, pan-viral ORFs, cancer-testis targets, per-tissue corpora |
38
+ | [`thymus/`](thymus) | Thymic self-peptidome and thymic expression, human and mouse — the central-tolerance reference |
39
+ | [`expression/`](expression) | Reference expression, human and mouse, plus a single-pipeline table and mouse protein abundance |
40
+ | [`proteome/`](proteome) | Human and mouse reference proteomes plus 16 bacterial/viral proteomes for mimicry scans |
41
+ | [`homology/`](homology) | Cross-species homology mappings |
42
+ | [`tcga/`](tcga) | Expressed mutant-peptide–HLA binders across 8,505 donors |
43
+ | [`hla_loh/`](hla_loh) | HLA-I loss of heterozygosity and genetic immune escape over 6,319 tumours, with three-caller typing concordance |
44
+ | [`icb_cohorts/`](icb_cohorts) | Harmonized checkpoint-blockade cohorts with outcome |
45
+ | [`vaccines/`](vaccines) | Shipped personalized-vaccine cassettes and their immune monitoring |
46
+
47
+ Formats are mixed by design: gzipped TSV for the large harmonized tables, parquet where the schema
48
+ is wide or typed, `.npz` for dense matrices, gzipped FASTA for proteomes.
49
+
50
+ ## Shared schema
51
+
52
+ The immunogenicity tables share one schema:
53
+
54
+ ```
55
+ peptide, mhc_a, mhc_b, mhc_class, mhc_species, source_species, host_species,
56
+ assay_type, immunogenicity{1,0,null}, expression, affinity, affinity_wt,
57
+ agretopicity, foreignness, validated, dataset_origin, reference_id, n_references
58
+ ```
59
+
60
+ plus `patient_id, cancer_type` on the neoantigen tables. **Canonical columns hold as-reported
61
+ values only**; anything predicted lives in a `pred_*` column, so `affinity` is always experimental.
62
+ `affinity` units differ by source and MHC alleles are as-reported.
63
+
64
+ ## Two things to know before filtering
65
+
66
+ - **`neoantigens/neoag_tested.tsv.gz` contains mouse rows labelled as human.** 6,186 CEDAR rows
67
+ carry mouse MHC alleles (`H2-Db` 1,020, `H2-Kb` 932, `H2-b class I` 719, `H2-Kd` 525,
68
+ `H2-d class II` 505, …) while their `mhc_species` reads `HomoSapiens` and `host_species` reads
69
+ `human`. `SIINFEKL`, the canonical H-2Kb ovalbumin epitope, is among them. Filtering that table
70
+ on `mhc_species` therefore silently admits mouse pMHC. The defect is upstream and is left
71
+ unfixed; `neoantigens/neoag_tested_mmu.tsv.gz` is the correctly-labelled murine set.
72
+ - **`neoag_tested_{mmu,hsa}` overlap `neoag_tested.tsv.gz` by design** — all 2,273 peptides appear
73
+ in both, since both are IEDB-derived. Dedupe on peptide × MHC when joining.
74
+
75
+ ## Loading
76
+
77
+ ```python
78
+ from huggingface_hub import hf_hub_download
79
+ import pandas as pd
80
+
81
+ path = hf_hub_download("isalgo/pmhc_data", "pmhc/pmhc_shortlist.tsv.gz", repo_type="dataset")
82
+ df = pd.read_csv(path, sep="\t")
83
+ ```
84
+
85
+ Grab one directory rather than the whole repo:
86
+
87
+ ```bash
88
+ hf download isalgo/pmhc_data --repo-type dataset --include "neoantigens/*" --local-dir ./pmhc_data
89
+ ```
90
+
91
+ [`mhcmatch`](https://pypi.org/project/mhcmatch/) consumes this compendium directly — `mhcmatch
92
+ bootstrap` stages the panel and reference tables from here.
93
+
94
+ ## Upstream sources
95
+
96
+ Which table came from which, and under what filtering, is in the `SOURCES.md` of each directory.
97
+
98
+ | | |
99
+ |---|---|
100
+ | IEDB | https://www.iedb.org |
101
+ | CEDAR | https://cedar.iedb.org |
102
+ | HLA Ligand Atlas | https://hla-ligand-atlas.org |
103
+ | UniProt | https://www.uniprot.org |
104
+ | GTEx | https://gtexportal.org |
105
+ | Human Protein Atlas | https://www.proteinatlas.org |
106
+ | FANTOM5 | https://fantom.gsc.riken.jp/5/ |
107
+ | UCSC Xena | https://xenabrowser.net |
108
+ | GDC / TCGA | https://portal.gdc.cancer.gov |
109
+ | ICGC / PCAWG | https://dcc.icgc.org |
110
+ | Hartwig Medical Foundation | https://www.hartwigmedicalfoundation.nl |
111
+ | Allele Frequency Net Database | https://www.allelefrequencies.net |
112
+ | PRIDE | https://www.ebi.ac.uk/pride/ |
113
+ | MassIVE | https://massive.ucsd.edu |
114
+ | GEO | https://www.ncbi.nlm.nih.gov/geo/ |
115
+
116
+ ## Licensing
117
+
118
+ Each upstream source carries its own terms; this compendium redistributes derived, harmonized
119
+ tables and does not relicense them. Check the `SOURCES.md` of the directory you are using before
120
+ redistributing, and cite the primary publications recorded there rather than this repository.
expression/SOURCES.md ADDED
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1
+ # `expression/` — sources
2
+
3
+ Two human tables. `reference_expression.tsv.gz` holds three references that are **never merged** — different measurements, different units, and
4
+ a gene-keyed table and a peptide-keyed one are not the same object.
5
+
6
+ | | |
7
+ |---|---|
8
+ | file | `reference_expression.tsv.gz` |
9
+ | rows | 6,681,814 |
10
+ | schema | `key`, `key_type`, `source`, `context`, `median_tpm`, `q25_tpm`, `q75_tpm`, `n` |
11
+ | provenance | **experimental** (all three sources are measured RNA-seq); the per-context median/IQR/`n` are **computed** summaries of it |
12
+
13
+ | `source` | `key_type` | rows | contexts | what a row is |
14
+ |---|---|--:|--:|---|
15
+ | `gtex` | `gene` | 3,955,284 | 53 | a gene's expression across normal tissue — the **safety** read |
16
+ | `hpa_consensus` | `gene` | 1,025,751 | 51 | the same read over HPA's tissue panel — **the only one that contains thymus** |
17
+ | `tcga` | `peptide` | 1,700,779 | 19 | a mutant peptide's expression by tumour type — the **target** read |
18
+
19
+ Total 6,681,814 rows.
20
+
21
+ `key_type` is the join key and it differs by source on purpose: GTEx and HPA answer "is this gene on
22
+ in normal tissue", TCGA answers "is this neoantigen transcribed in this tumour". A caller that treats
23
+ them as one column is asking a question none of them answers. GTEx and HPA are both gene-keyed but
24
+ are still not interchangeable — see the units caveat in the HPA section below.
25
+
26
+ Read through `mhcmatch.expression` (`lookup`, `tissues`, `tumor_types`, `safety_profile`), which
27
+ keeps them apart. Fetched by `mhcmatch bootstrap --reference` (~115 MB).
28
+
29
+ ---
30
+
31
+ ## `reference_expression_toil.tsv.gz` — GTEx and TCGA on one pipeline, in one unit
32
+
33
+ Added 2026-08-26. **This is the table to use when a normal tissue and a tumour type have to be
34
+ compared, or when one supplies a scale for the other.**
35
+
36
+ | | |
37
+ |---|---|
38
+ | rows | **5,037,966** |
39
+ | schema | `key`, `key_type`, `source`, `context`, `median_tpm`, `q25_tpm`, `q75_tpm`, `n`, `unit` |
40
+ | keys | 58,581 gene symbols (`key_type = gene` on every row) |
41
+ | `source` / contexts | `toil_gtex` — 53 GTEx tissues, 3,104,793 rows · `toil_tcga` — 33 TCGA studies, 1,933,173 rows |
42
+ | `unit` | `TPM` on every row |
43
+ | size | 38.6 MB |
44
+ | provenance | **experimental** — RSEM TPM from RNA-seq; the per-context median/IQR/`n` are **computed** summaries |
45
+ | built by | `bench/neoag/expression_toil.py` in `2026-mhcmatch-benchmark` |
46
+
47
+ **Why it exists.** `reference_expression.tsv.gz` carries GTEx in **TPM** and TCGA in **RSEM
48
+ upper-quartile normalised counts**, both written into a column called `median_tpm`. They are each
49
+ correct for their own question, but they cannot be put on one axis: a scale taken from one and
50
+ divided into an abundance from the other is a units error no caller can detect, because the column
51
+ name says TPM in both cases. This table removes the problem at the source rather than by converting
52
+ one into the other — a conversion that is not available from the deposited matrix, since RSEM
53
+ normalised counts are proportional to abundance x effective length and the effective lengths are not
54
+ published with it.
55
+
56
+ **Origin.** The UCSC Xena / Toil recompute, which put GTEx, TCGA and TARGET through a **single RSEM
57
+ pipeline against one annotation (GENCODE v23)** and publishes gene-level TPM for all of them, so a
58
+ GTEx tissue median and a TCGA study median differ only in the samples that went in.
59
+
60
+ | input | file | bytes |
61
+ |---|---|--:|
62
+ | expression matrix | `TcgaTargetGtex_rsem_gene_tpm.gz` | 1,323,254,426 |
63
+ | sample phenotype | `TcgaTargetGTEX_phenotype.txt.gz` | 135,753 |
64
+ | gene annotation | `probeMap/gencode.v23.annotation.gene.probemap` | 3,244,244 |
65
+
66
+ ```
67
+ base=https://toil-xena-hub.s3.us-east-1.amazonaws.com/download
68
+ curl -L -o xena/TcgaTargetGtex_rsem_gene_tpm.gz $base/TcgaTargetGtex_rsem_gene_tpm.gz
69
+ curl -L -o xena/TcgaTargetGTEX_phenotype.txt.gz $base/TcgaTargetGTEX_phenotype.txt.gz
70
+ curl -L -o xena/gencode.v23.annotation.gene.probemap \\
71
+ "$base/probeMap%2Fgencode.v23.annotation.gene.probemap"
72
+ python bench/neoag/expression_toil.py --out expression/reference_expression_toil.parquet
73
+ ```
74
+
75
+ > Vivian J, Rao AA, Nothaft FA, Ketchum C, Armstrong J, Novak A, Pfeil J, Narkizian J, Deran AD,
76
+ > Musselman-Brown A, Schmidt H, Amstutz P, Craft B, Goldman M, Rosenbloom K, Cline M, O'Connor B,
77
+ > Hanna M, Birger C, Kent WJ, Patterson DA, Joseph AD, Zhu J, Zaranek S, Getz G, Haussler D,
78
+ > Paten B. **Toil enables reproducible, open source, big biomedical data analyses.**
79
+ > *Nat Biotechnol* 2017;35(4):314-316.
80
+ > PMID [28398314](https://pubmed.ncbi.nlm.nih.gov/28398314/) ·
81
+ > PMC5546205 · doi:[10.1038/nbt.3772](https://doi.org/10.1038/nbt.3772)
82
+
83
+ > Goldman MJ, Craft B, Hastie M, Repecka K, McDade F, Kamath A, Banerjee A, Luo Y, Rogers D,
84
+ > Brooks AN, Zhu J, Haussler D. **Visualizing and interpreting cancer genomics data via the Xena
85
+ > platform.** *Nat Biotechnol* 2020;38(6):675-678.
86
+ > PMID [32444850](https://pubmed.ncbi.nlm.nih.gov/32444850/) ·
87
+ > PMC7386072 · doi:[10.1038/s41587-020-0546-8](https://doi.org/10.1038/s41587-020-0546-8)
88
+
89
+ **The TCGA study code is joined through the barcode, never through the disease name.** Xena labels a
90
+ TCGA sample with a disease name (`Uveal Melanoma`); the rest of this compendium keys on the study
91
+ abbreviation (`UVM`). A hand-written name->code crosswalk can be silently wrong, so the code is taken
92
+ from PanCanAtlas `merged_sample_quality_annotations.tsv` on the 15-character sample barcode, which is
93
+ exact. **17,474 of 19,131 matrix columns are used**: 9,616 TCGA tumour aliquots and 7,862 GTEx
94
+ samples. TARGET (734 columns) is dropped -- paediatric, and none of its diseases has a matched normal
95
+ here -- as are TCGA adjacent normals and aliquots the source flags `Do_not_use`.
96
+
97
+ **Upstream values are `log2(TPM + 0.001)` and are inverted on build**, so what is stored is TPM.
98
+
99
+ **Caveats.**
100
+ - **The GTEx contexts are Toil-era, not v11.** `reference_expression.tsv.gz` is GTEx v11; this is the
101
+ earlier release Toil reprocessed, so the two GTEx halves are neither the same samples nor the same
102
+ depth. They are also not spelled the same: Toil writes `Skin - Sun Exposed (Lower Leg)` and GTEx
103
+ v11 writes `Skin - Sun Exposed (Lower leg)`. **A context string cannot be carried from one table
104
+ to the other without checking.**
105
+ - **Three tissues are thin and are kept with their `n` rather than dropped**: Bladder (9),
106
+ Cervix - Ectocervix (6), Fallopian Tube (5). They are the matched normals of BLCA, OV and CESC, so
107
+ dropping them would remove the safety read for three tumour types. `Cervix - Endocervix` has fewer
108
+ than five samples and is absent; CESC still resolves through Ectocervix. Every TCGA study clears 36.
109
+ - **353 of 58,581 symbols carry more than one Ensembl gene id** (0.6 %). The row kept per
110
+ (symbol, context) is the one with the **highest median**, not a sum across loci: a lookup asks
111
+ whether a gene is on in this context, and the dominant locus answers it without inventing a total
112
+ across paralogues. 164,862 duplicate rows were collapsed this way.
113
+ - `n` is a **sample** count here, as in the GTEx half, and not the transcript count the mouse table's
114
+ `n` is.
115
+
116
+ **What a floor built from it looks like**, as the 25th percentile of non-zero gene medians:
117
+ TCGA runs 0.1200 (KICH) to 0.3700 (LAML) TPM, a **3.08x** spread over 33 studies; GTEx runs 0.1000
118
+ (Whole Blood) to 0.4000 (Testis), **4.00x** over 53 tissues. Pooled, TCGA is 0.1800 TPM over 882,959
119
+ non-zero gene rows and GTEx 0.2700 over 1,402,705. **A tumour's own floor is consistently below its
120
+ matched normal's** -- SKCM 0.1600 against skin 0.3050 (0.52x), BLCA 0.1700 against bladder 0.3600
121
+ (0.47x), LUAD 0.2000 against lung 0.3500 (0.57x), BRCA 0.1900 against breast 0.3000 (0.63x).
122
+
123
+ **Format: parquet, and the record is not in the bootstrap set.** As gzipped TSV this table was
124
+ 70.5 MB and 0.5 s to read with polars (5.2 s with the standard library); dictionary-encoded and
125
+ zstd-compressed it is **38.6 MB and 0.04 s**. On a link measured here at 154 kB/s that is three and
126
+ a half minutes of download saved, which is the difference between a bootstrap that finishes and one
127
+ that looks like it has hung. `mhcmatch bootstrap --reference` stages only the three companions
128
+ below — 6.6 MB between them, carrying the same numbers — and never this file.
129
+
130
+ **Label arrays in the `.npz` are fixed-width unicode, not `dtype=object`.** An object array is
131
+ pickled, and `np.load` needs `allow_pickle=True` to read it back; a pickled array in a file fetched
132
+ over the network executes on load. The deposit is written so that flag is never needed.
133
+
134
+ ### Its three companions, and why scoring never reads the table itself
135
+
136
+ Added 2026-08-26 beside it. `reference_expression_toil.tsv.gz` is the record; these are what a
137
+ program actually opens.
138
+
139
+ | file | size | what it is |
140
+ |---|--:|---|
141
+ | `toil_matrix.npz` | 6,632,665 B | the whole table as a dense **58,581 gene x 86 context float32 matrix**, contexts keyed `"<source>|<context>"` |
142
+ | `toil_floors.tsv` | 8,365 B | 88 rows -- each context's abundance floor at q05, q10, q25, plus a pooled row per cohort |
143
+ | `context_synonyms.tsv` | 29,334 B | 448 rows, 184 aliases -- how a free-text origin becomes a context |
144
+
145
+ **The matrix exists so a lookup does not cost a whole table.** Answering *what is this gene in this
146
+ tumour type* from the TSV means building a five-million-entry dictionary of dictionaries: measured,
147
+ **5.20 s and 3,168 MB resident**. The matrix is **0.05 s and 29 MB** -- 100x the speed on 1/109 of
148
+ the memory -- and every lookup a resolution chain makes is one array index. Both return the same
149
+ floor to four decimals, which is how the matrix is checked against the table it came from. A gene absent from a context is `0.0` and not `nan`: every gene here was
150
+ measured in every context that cleared the sample threshold, so absence is silence, and silence is a
151
+ measurement. Whether a *gene* is in the reference at all is a separate question and is reported
152
+ separately -- silence and ignorance must not collapse into one number.
153
+
154
+ **`context_synonyms.tsv` is derived, not hand-written.** Long format, one target per row, so it
155
+ sorts and joins like data rather than holding lists in cells:
156
+
157
+ | alias_kind | where the aliases come from | rows |
158
+ |---|---|--:|
159
+ | `code` | the 33 TCGA studies in the matrix | 83 |
160
+ | `site` | Xena `_primary_site`, joined to the PanCanAtlas study code on the 15-character barcode | 218 |
161
+ | `disease` | Xena `detailed_category`, joined the same way | 83 |
162
+ | `gtex` | the 53 GTEx contexts in the matrix | 64 |
163
+
164
+ **One alias resolves to two things and each is its own row**, which is why the alias count and the
165
+ row count differ: 150 rows carry `target_kind = tcga_code` and 298 carry `gtex_context`, so
166
+ `hepatocellular` names both `LIHC` and `Liver`. 152 of the 448 rows are `derived:*` and 296 are
167
+ `curated:*`, and the `provenance` column says which on every row. The curated majority is the
168
+ matched-normal half and the adjectival forms a free-text origin actually arrives in
169
+ (`cervical`, `pancreatic`, `uterine corpus`); the derived half is what the matrix and the barcode
170
+ join supply on their own. Hand-writing this would have been wrong in the places nobody checks: **an organ is
171
+ more than one study more often than not** -- `Lung` is `LUAD` *and* `LUSC`, `Kidney` is three -- and a
172
+ crosswalk that picks one is silently wrong for the rest. The tumour-to-matched-normal half is carried
173
+ from `mhcmatch.expression.TUMOR_TISSUE` against the spellings this matrix uses, because Toil writes
174
+ `Brain - Frontal Cortex (Ba9)` where GTEx v11 writes `(BA9)` and an exact match drops it.
175
+
176
+ An alias that does not resolve **raises**. Falling back to the pooled reference would return a
177
+ plausible number computed from the wrong distribution with nothing downstream able to tell.
178
+
179
+ **A note on what these floors are for.** `c` is a property of a transcriptome, and the numerator is
180
+ one candidate's abundance. Where the numerator is not in reference TPM the two stop cancelling, and
181
+ the obvious repair -- a median of ratios against the reference -- **must not be estimated from a
182
+ candidate list**. A mutation reaches such a list only if it was called, and only where the gene was
183
+ seen in RNA, so the ratio measures that conditioning instead of the library: on three screens whose
184
+ columns are all deposited as TPM it returns 1.78, 2.18 and 3.15, never below 1. `mhcmatch` gates it
185
+ on coverage of the reference transcriptome for that reason, and a candidate list cannot clear it.
186
+
187
+ ---
188
+
189
+ ## Mouse — `reference_expression_mmu.tsv.gz` and `protein_abundance_mmu.tsv.gz`
190
+
191
+ Added 2026-08-21. The compendium's normal-tissue safety read was GTEx, and GTEx is human, so a mouse
192
+ epitope had no reference to be scored against. These two files are the mouse counterpart.
193
+
194
+ **They are two files, not one, and that is deliberate.** The human table's value column is literally
195
+ `median_tpm`. RNA and protein are different measurements in different units, and writing a SILAC
196
+ abundance into a `_tpm` column would be a units lie no caller could detect.
197
+
198
+ ### `reference_expression_mmu.tsv.gz` — the GTEx analogue
199
+
200
+ | | |
201
+ |---|---|
202
+ | rows | **659,050** |
203
+ | schema | `key`, `key_type`, `source`, `context`, `median_tpm`, `q25_tpm`, `q75_tpm`, `n` — **column for column identical to the human file** |
204
+ | keys | 18,830 gene symbols (`key_type = gene`) |
205
+ | contexts | **35 adult tissues** |
206
+ | `source` | `fantom5_mouse` |
207
+ | provenance | **experimental** — CAGE tag counts; the per-gene median across transcripts and the quantile summary are **computed** |
208
+
209
+ **Origin.** EBI Expression Atlas [E-MTAB-3579](https://www.ebi.ac.uk/gxa/experiments/E-MTAB-3579),
210
+ the RIKEN FANTOM5 mouse CAGE atlas. `q25_tpm` / `median_tpm` / `q75_tpm` are taken from the Atlas'
211
+ own five-number summary per gene × group; they are not re-derived here.
212
+
213
+ > Lizio M, Harshbarger J, Shimoji H, Severin J, Kasukawa T, Sahin S, Abugessaisa I, Fukuda S,
214
+ > Hori F, Ishikawa-Kato S, Mungall CJ, Arner E, Baillie JK, Bertin N, Bono H, de Hoon M, Diehl AD,
215
+ > Dimont E, Freeman TC, Fujieda K, Hide W, Kaliyaperumal R, Katayama T, Lassmann T, Meehan TF,
216
+ > Nishikata K, Ono H, Rehli M, Sandelin A, Schultes EA, 't Hoen PAC, Tatum Z, Thompson M,
217
+ > Toyoda T, Wright DW, Daub CO, Itoh M, Carninci P, Hayashizaki Y, Forrest ARR, Kawaji H.
218
+ > **Gateways to the FANTOM5 promoter level mammalian expression atlas.**
219
+ > *Genome Biol* 2015;16:22.
220
+ > PMID [25723102](https://pubmed.ncbi.nlm.nih.gov/25723102/) ·
221
+ > doi:[10.1186/s13059-014-0560-6](https://doi.org/10.1186/s13059-014-0560-6)
222
+
223
+ **Only plain `adult` groups are kept** — 35 of the deposit's 170. GTEx is adult normal tissue;
224
+ folding an E14 embryo, a pregnancy timepoint or a lactating mammary gland into a "is this gene on in
225
+ normal tissue" safety read answers a different question. The other stages are in the source and are
226
+ recoverable by re-running without the filter.
227
+
228
+ **A parsing trap worth recording.** Atlas group labels are two `;`-separated parts and **the order
229
+ is not fixed**: 136 of the 170 are stage-first (`adult; thymus`) and 32 are tissue-first
230
+ (`adrenal gland; adult`). Taking either position blindly mislabels a third of the table — the first
231
+ build of this file produced contexts named `adult` and `whole body` that way. The stage is matched by
232
+ vocabulary wherever it sits, and the tissue is whatever remains.
233
+
234
+ **Checked against known biology, not assumed.** Of 35 tissues, thymus ranks **first** for `Prss16`
235
+ (31 TPM), `Psmb11` (88), `Dntt` (961), `Rag1`, `Cd8a` (131) and `Lck` (12) — thymic epithelium and
236
+ thymocyte genes. As a negative control `Cela1` peaks in pancreas at 225,010 TPM against 0.3 in
237
+ thymus.
238
+
239
+ **Caveats.**
240
+ - **`n` is 1 for every row.** FANTOM5 provides one library per adult tissue, so `q25`/`q75` describe
241
+ the spread across transcripts of a gene, **not across animals**. GTEx `n` runs to several hundred
242
+ donors. Do not read the two `n` columns as the same quantity.
243
+ - **`Aire` is 0 TPM in thymus here, and that is a detection floor, not biology.** Aire is expressed
244
+ in a rare mTEC subset; bulk CAGE at this depth cannot see it. Any promiscuous-expression question
245
+ needs sorted mTEC data, not this file.
246
+ - CAGE measures 5′-capped tag density. It is reported as TPM and behaves like expression, but it is
247
+ not RNA-seq TPM and the two are not numerically interchangeable across sources.
248
+
249
+ ### `tumor_expression_mmu.tsv.gz` — the mouse TUMOUR read, and why it is a separate file
250
+
251
+ Added 2026-09-04. `reference_expression_mmu.tsv.gz` answers the *safety* half of the expression
252
+ question -- is this gene on in normal tissue -- and there was no counterpart for the *ranking* half.
253
+ Mouse has no TCGA, so a mouse neoantigen's own abundance fell back to the gene's normal-tissue
254
+ median, which is the number `expr_norm` already carries. Measured on the 923 class-I rows of the
255
+ IEDB mouse neoantigen deposit, `expr_lvl` and `expr_norm` came out **identical on 100 %** of the
256
+ rows lacking a deposited abundance: two fitted terms on one column.
257
+
258
+ | | |
259
+ |---|---|
260
+ | rows | **149,640** |
261
+ | schema | `key`, `key_type`, `source`, `context`, `median_tpm`, `q25_tpm`, `q75_tpm`, `n` — **column for column identical to the two files above** |
262
+ | keys | 24,940 gene symbols (`key_type = gene`) |
263
+ | contexts | **6 syngeneic tumour models**: B16F10, CT26, E0771, LLC, MC38, Panc02 |
264
+ | `source` | `gse245293_syngeneic` |
265
+ | `n` | **3 on every row** — three biological replicates per model, so `q25`/`q75` are a spread across *animals*, unlike the FANTOM5 file's spread across transcripts |
266
+ | provenance | **experimental** — RNA-seq counts; the length normalisation, the per-model quantiles and the gene-level aggregation are **computed** |
267
+
268
+ **Origin.** NCBI GEO [GSE245293](https://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE245293),
269
+ supplementary file `GSE245293_Normalized_count_table_tumor.txt.gz` (2.9 MB), staged at
270
+ `raw/expression_mouse/`. 23 samples in the series; the 18 `*_tumor_rep*` columns are used and the
271
+ 8 CAF (cancer-associated fibroblast) columns are **not** -- those are stroma, not the tumour
272
+ transcriptome, and folding them in would answer a different question.
273
+
274
+ curl -O https://ftp.ncbi.nlm.nih.gov/geo/series/GSE245nnn/GSE245293/suppl/GSE245293_Normalized_count_table_tumor.txt.gz
275
+ python src/build_tumor_expression_mmu.py # in ~/vcs/projects/public_expression_peptidomes
276
+
277
+ > Comparison of syngeneic tumor models reveals different tumor microenvironments with distinct
278
+ > subsets of cancer-associated fibroblasts polarized by infiltrating T cells. GEO GSE245293,
279
+ > deposited 2023-10-13, released 2024-01-29. *Mus musculus*, expression profiling by high
280
+ > throughput sequencing.
281
+
282
+ **Units are computed and the file says so.** The deposit is **normalised counts** with a `Length`
283
+ column, not TPM. Length-normalising each sample and rescaling to a million gives a TPM-like
284
+ quantity comparable *within* this table; it is **not** interchangeable with FANTOM5 CAGE TPM or
285
+ with GTEx RSEM TPM. `mhcmatch.rank.expr_level` is unit-free when its floor is a quantile of the
286
+ same column, which is exactly why a floor for these contexts must be taken from this file.
287
+
288
+ **Checked against known biology, not assumed.** Three independent melanocytic markers separate the
289
+ melanoma line by three orders of magnitude: `Tyr` **398.9**, `Dct` **4,611.8** and `Pmel`
290
+ **4,636.7** TPM in B16F10 against ≤ 4.3 in all five other models. They would all move together if
291
+ the sample-to-model mapping were wrong.
292
+
293
+ **Colon markers were tried and do not work here; do not re-try them.** `Muc2` reads ~0 in every
294
+ model -- CT26 is an undifferentiated colon carcinoma and makes no goblet-cell mucin -- and
295
+ `Cdx1`/`Cdx2` do not separate CT26 and MC38 from the rest (Cdx2 is highest in Panc02, 8.8 against
296
+ CT26's 1.4). The expectations were wrong, not the data.
297
+
298
+ **Coverage of the IEDB mouse neoantigen deposit**, joining on model × gene symbol: **347 of 968
299
+ class-I rows (36 %)** and **304 of 522 class-II rows (58 %)**. The class-II figure is the reason
300
+ this file exists -- that arm previously had a deposited abundance on 13 % of its rows.
301
+
302
+ **Two models it does not carry** that the neoantigen deposit names often: `P815` (89 rows) and the
303
+ `T3`/`CMS5`/`Meth A` fibrosarcomas (150 rows between them). Those still take the pooled floor.
304
+
305
+ ### `PRJNA1271699` — located, not fetched
306
+
307
+ [PRJNA1271699](https://www.ncbi.nlm.nih.gov/bioproject/PRJNA1271699), *"Mouse syngeneic model
308
+ RNAseq sequencing"* (CrownBio), submitted 2025-06-04: **120 runs over 12 models × 10 replicates**
309
+ -- A20, B16BL6, B16F10, CT26, EMT6, H22, Hepa16, LL2, MC38, Pan02, RM1, Renca -- paired-end
310
+ Illumina NovaSeq 6000, **418 GB** of raw reads. It would cover Renca and EMT6, which nothing here
311
+ does.
312
+
313
+ **NCBI carries raw reads only; there is no processed matrix.** The BioProject abstract advertises
314
+ an interactive resource at `https://muscreen.crownbio.com/syngeneic_omics`, and **as of 2026-09-04
315
+ that URL is dead** -- `/syngeneic_omics` returns 404 and the host root returns a bare Apache
316
+ default page (403). So the processed tables the abstract promises are not currently retrievable,
317
+ and using this project means downloading 418 GB and running an alignment pipeline. Recorded here
318
+ so the cost is known rather than rediscovered.
319
+
320
+ **This is the source to re-process if these models are wanted here** -- it is the one that covers
321
+ P815, Renca and EMT6, which `tumor_expression_mmu.tsv.gz` does not.
322
+
323
+ # metadata only, no reads:
324
+ curl -sS "https://trace.ncbi.nlm.nih.gov/Traces/sra-db-be/runinfo?acc=PRJNA1271699"
325
+
326
+ ### `toil_matrix_mmu.npz` + `toil_floors_mmu.tsv` — the harmonised mouse compendium
327
+
328
+ Added 2026-09-04. The mouse counterpart of `toil_matrix.npz` / `toil_floors.tsv`, same array names
329
+ and the same `source|context` key shape, so code written against the human pair reads this one.
330
+
331
+ | | |
332
+ |---|---|
333
+ | shape | **26,737 genes x 68 contexts**, dense float32 |
334
+ | `genes` | official NCBI symbol, synonyms resolved via Ensembl release-100 (= GENCODE M25) and NCBI Gene |
335
+ | `contexts` | `fantom5_mouse\|<tissue>` x 35 normal, `gse245293\|<model>` x 6 tumour, `prjna1271699+\|<model>` x 27 tumour |
336
+ | `values` | linear TPM-like, **harmonised across sources**; `NaN` = the gene is absent from that source, which is not the same as measured at zero |
337
+ | `n_samples` | 3 for GSE245293 (replicate **mean**), 1 for FANTOM5, 0 where the source reports none |
338
+ | `toil_floors_mmu.tsv` | 71 rows -- each context's floor at q05/q10/q25, plus a pooled row per source, same seven columns as the human file |
339
+
340
+ **Sources.** FANTOM5 mouse CAGE (EBI E-MTAB-3579); GEO **GSE245293** (6 syngeneic models x 3
341
+ replicates); and the CrownBio syngeneic panel, labelled `prjna1271699+` for NCBI BioProject
342
+ **PRJNA1271699** plus the further models that panel carries beyond the 12 the accession lists.
343
+ GSE245293 wins the five models both cover. The 27 filled models include **P815, Renca and EMT6**,
344
+ which no other deposit here reaches.
345
+
346
+ **Harmonised, and here is what that means.** The three sources sat on three scales -- pooled mean
347
+ `log2(1 + TPM)` of 1.721, 2.513 and 2.059. Each source's non-zero values are quantile-mapped onto
348
+ the mean of the three sources' quantile functions; **per source, not per context**, so
349
+ between-context structure survives. Zero maps to zero. The map is monotone and its grid spans each
350
+ source's support, so **every within-context rank is preserved exactly** -- which is what
351
+ `mhcmatch.rank.expr_level` consumes. Non-zero spread falls 0.577 -> 0.087 log2 units, and the
352
+ pooled q25 floor agrees to 0.714--0.717 across all three.
353
+
354
+ Two things it does not do. It cannot correct a **gene-specific** platform effect: `Actb` reads 3,032
355
+ in B16F10 and 10.4 in FANTOM5 thymus after harmonisation, because CAGE at that promoter genuinely
356
+ ranks it mid-distribution. And it cannot correct the **detection-rate** difference (65.2 / 78.1 /
357
+ 65.0 % non-zero), because a monotone map that respects zero cannot move it.
358
+
359
+ **Absolute values are no longer any one source's own number.** That is the trade: comparability
360
+ across all 68 contexts, at the cost of each source's internal scale. The unharmonised per-source
361
+ tables are not deposited; `tumor_expression_mmu.tsv.gz` and `reference_expression_mmu.tsv.gz` above
362
+ remain the unharmonised reads of two of the three sources.
363
+
364
+ Built by `src/build_deposit_mmu.py` in `~/vcs/projects/public_expression_peptidomes`; the protocol
365
+ and its numbers are `results/expression_harmonization.md` there.
366
+
367
+ **For comparison, the human pair needed none of this.** Same measurement on `toil_matrix.npz`:
368
+ `toil_gtex` and `toil_tcga` pooled means differ by **0.011** log2 units, against 0.565 for the
369
+ mouse CAGE-vs-RNA-seq seam. Both human halves went through one Toil RSEM recompute, and it worked.
370
+
371
+ ### `protein_abundance_mmu.tsv.gz` — the protein-level read
372
+
373
+ | | |
374
+ |---|---|
375
+ | rows | **133,848** |
376
+ | schema | `key`, `key_type`, `source`, `context`, `abundance`, `unit`, `n` |
377
+ | keys | 5,148 gene symbols |
378
+ | contexts | **26 tissues**, including thymus |
379
+ | `source` / `unit` | `geiger_silac_mouse` / `ppb_ibaq` |
380
+ | provenance | **experimental** — SILAC-normalised MS abundance |
381
+
382
+ **Origin.** EBI Expression Atlas [E-PROT-11](https://www.ebi.ac.uk/gxa/experiments/E-PROT-11).
383
+
384
+ > Geiger T, Velic A, Macek B, Lundberg E, Kampf C, Nagaraj N, Uhlen M, Cox J, Mann M.
385
+ > **Initial quantitative proteomic map of 28 mouse tissues using the SILAC mouse.**
386
+ > *Mol Cell Proteomics* 2013;12(6):1709–1722.
387
+ > PMID [23436904](https://pubmed.ncbi.nlm.nih.gov/23436904/) ·
388
+ > doi:[10.1074/mcp.M112.024919](https://doi.org/10.1074/mcp.M112.024919)
389
+
390
+ **Why keep a protein table at all.** A peptide can only be presented if the protein was translated,
391
+ so for immunopeptidome work protein abundance is the more direct evidence and RNA is a proxy. It is
392
+ much shallower — 5,148 genes against 18,830 — so it complements the RNA table rather than replacing
393
+ it.
394
+
395
+ ### Why not E-PROT-13
396
+
397
+ [E-PROT-13](https://www.ebi.ac.uk/gxa/experiments/E-PROT-13) (Huttlin's mouse phosphorylation and
398
+ expression atlas) was the first candidate considered. It covers **9 tissues and does not include
399
+ thymus**, so it cannot serve either purpose here. Of the **65 mouse baseline experiments in
400
+ Expression Atlas, only six contain thymus**; E-MTAB-3579 (35 adult tissues) and E-PROT-11 (26) are
401
+ the only two of those with broad tissue coverage.
402
+
403
+ ### Regenerate
404
+
405
+ cd ~/vcs/projects/public_expression_peptidomes && ./src/build_expression_mmu.py
406
+
407
+ ---
408
+
409
+ ## Human — the `hpa_consensus` arm, and why GTEx alone was not enough
410
+
411
+ Added 2026-08-21, in the same pass that built the mouse references.
412
+
413
+ **GTEx has no thymus.** Its 53 contexts contain no thymus, no lymph node and no bone marrow — adult
414
+ thymus involutes and is not routinely sampled, so the normal-tissue safety read was blind to the one
415
+ tissue central tolerance actually happens in. Expression Atlas is no better: of its **123 human
416
+ baseline experiments only three carry any thymus group**, and all three are compromised for this
417
+ purpose — E-MTAB-3871 is fetal tissue from fetuses with congenital defects, E-MTAB-3827 is sorted
418
+ thymocyte subsets rather than tissue, and E-MTAB-9641 is thymic epithelium cultured on a 3T3-J2
419
+ feeder layer.
420
+
421
+ HPA samples thymus directly, so its consensus tissue set is what fills the hole.
422
+
423
+ | | |
424
+ |---|---|
425
+ | rows | **1,025,751** |
426
+ | keys | 20,151 gene symbols (`key_type = gene`) |
427
+ | contexts | **51 tissues, including thymus** |
428
+ | `source` | `hpa_consensus` |
429
+ | provenance | **experimental** — RNA-seq; the HPA consensus normalisation is **computed** |
430
+
431
+ **Origin.** [`rna_tissue_consensus.tsv.zip`](https://www.proteinatlas.org/download/tsv/rna_tissue_consensus.tsv.zip)
432
+ from the Human Protein Atlas.
433
+
434
+ > Uhlén M, Fagerberg L, Hallström BM, Lindskog C, Oksvold P, Mardinoglu A, Sivertsson Å, Kampf C,
435
+ > Sjöstedt E, Asplund A, Olsson I, Edlund K, Lundberg E, Navani S, Szigyarto CA, Odeberg J,
436
+ > Djureinovic D, Takanen JO, Hober S, Alm T, Edqvist PH, Berling H, Tegel H, Mulder J, Rockberg J,
437
+ > Nilsson P, Schwenk JM, Hamsten M, von Feilitzen K, Forsberg M, Persson L, Johansson F, Zwahlen M,
438
+ > von Heijne G, Nielsen J, Pontén F.
439
+ > **Tissue-based map of the human proteome.**
440
+ > *Science* 2015;347(6220):1260419.
441
+ > PMID [25613900](https://pubmed.ncbi.nlm.nih.gov/25613900/) ·
442
+ > doi:[10.1126/science.1260419](https://doi.org/10.1126/science.1260419)
443
+
444
+ **It is added as a third `source`, not merged into GTEx and not replacing it.** The schema was
445
+ designed with a `source` column for exactly this; every existing `gtex` and `tcga` row is untouched.
446
+
447
+ **Checked against known biology.** Of 51 tissues, thymus ranks **first** for `PRSS16` (770 nTPM),
448
+ `PSMB11` (28.5), `DNTT` (306), `CD1A` (296), `RAG1` (368), `CCL25` (718), `CD3E` (596) and `LCK`
449
+ (326). Negative controls behave: `ALB` peaks in liver at 198,524 nTPM against 0.3 in thymus, `INS` in
450
+ pancreas, `MYH7` in skeletal muscle.
451
+
452
+ **Caveats.**
453
+ - **nTPM is not GTEx TPM.** Both are TPM-family units but the normalisation differs, so compare
454
+ *within* a `source`, not across. This is what the `source` column is for.
455
+ - `n` is 1 — HPA consensus publishes one value per gene × tissue, so `q25_tpm` and `q75_tpm` equal
456
+ `median_tpm` and carry no spread. GTEx `n` runs to several hundred donors; the two `n` columns are
457
+ not the same quantity.
458
+ - **`AIRE` is detectable here (1.2 nTPM, rank 2 of 51) but reads 0 in the mouse CAGE atlas.** That is
459
+ a difference in assay depth, not in biology — do not read it as a species difference.
expression/context_synonyms.tsv ADDED
@@ -0,0 +1,449 @@
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
1
+ alias alias_kind target_kind target provenance
2
+ ACC code gtex_context Adrenal Gland curated:TUMOR_TISSUE
3
+ ACC code tcga_code ACC derived:toil_matrix
4
+ Acute Myeloid Leukemia disease gtex_context Whole Blood curated:TUMOR_TISSUE
5
+ Acute Myeloid Leukemia disease tcga_code LAML derived:xena_detailed_category
6
+ Adipose - Subcutaneous gtex gtex_context Adipose - Subcutaneous derived:toil_matrix
7
+ Adipose - Visceral (Omentum) gtex gtex_context Adipose - Visceral (Omentum) derived:toil_matrix
8
+ Adrenal Gland gtex gtex_context Adrenal Gland derived:toil_matrix
9
+ Adrenal gland site gtex_context Adrenal Gland curated:TUMOR_TISSUE
10
+ Adrenal gland site tcga_code ACC derived:xena_primary_site
11
+ Adrenocortical Cancer disease gtex_context Adrenal Gland curated:TUMOR_TISSUE
12
+ Adrenocortical Cancer disease tcga_code ACC derived:xena_detailed_category
13
+ Artery - Aorta gtex gtex_context Artery - Aorta derived:toil_matrix
14
+ Artery - Coronary gtex gtex_context Artery - Coronary derived:toil_matrix
15
+ Artery - Tibial gtex gtex_context Artery - Tibial derived:toil_matrix
16
+ BLCA code gtex_context Bladder curated:TUMOR_TISSUE
17
+ BLCA code tcga_code BLCA derived:toil_matrix
18
+ BRCA code gtex_context Breast - Mammary Tissue curated:TUMOR_TISSUE
19
+ BRCA code tcga_code BRCA derived:toil_matrix
20
+ Bile duct site gtex_context Liver curated:TUMOR_TISSUE
21
+ Bile duct site tcga_code CHOL derived:xena_primary_site
22
+ Bladder gtex gtex_context Bladder derived:toil_matrix
23
+ Bladder site gtex_context Bladder curated:TUMOR_TISSUE
24
+ Bladder site tcga_code BLCA derived:xena_primary_site
25
+ Bladder Urothelial Carcinoma disease gtex_context Bladder curated:TUMOR_TISSUE
26
+ Bladder Urothelial Carcinoma disease tcga_code BLCA derived:xena_detailed_category
27
+ Brain site gtex_context Brain - Amygdala curated:TUMOR_TISSUE
28
+ Brain site gtex_context Brain - Anterior Cingulate Cortex (Ba24) curated:TUMOR_TISSUE
29
+ Brain site gtex_context Brain - Caudate (Basal Ganglia) curated:TUMOR_TISSUE
30
+ Brain site gtex_context Brain - Cerebellar Hemisphere curated:TUMOR_TISSUE
31
+ Brain site gtex_context Brain - Cerebellum curated:TUMOR_TISSUE
32
+ Brain site gtex_context Brain - Cortex curated:TUMOR_TISSUE
33
+ Brain site gtex_context Brain - Frontal Cortex (Ba9) curated:TUMOR_TISSUE
34
+ Brain site gtex_context Brain - Hippocampus curated:TUMOR_TISSUE
35
+ Brain site gtex_context Brain - Hypothalamus curated:TUMOR_TISSUE
36
+ Brain site gtex_context Brain - Nucleus Accumbens (Basal Ganglia) curated:TUMOR_TISSUE
37
+ Brain site gtex_context Brain - Putamen (Basal Ganglia) curated:TUMOR_TISSUE
38
+ Brain site gtex_context Brain - Spinal Cord (Cervical C-1) curated:TUMOR_TISSUE
39
+ Brain site gtex_context Brain - Substantia Nigra curated:TUMOR_TISSUE
40
+ Brain site tcga_code GBM derived:xena_primary_site
41
+ Brain site tcga_code LGG derived:xena_primary_site
42
+ Brain - Amygdala gtex gtex_context Brain - Amygdala derived:toil_matrix
43
+ Brain - Anterior Cingulate Cortex (Ba24) gtex gtex_context Brain - Anterior Cingulate Cortex (Ba24) derived:toil_matrix
44
+ Brain - Caudate (Basal Ganglia) gtex gtex_context Brain - Caudate (Basal Ganglia) derived:toil_matrix
45
+ Brain - Cerebellar Hemisphere gtex gtex_context Brain - Cerebellar Hemisphere derived:toil_matrix
46
+ Brain - Cerebellum gtex gtex_context Brain - Cerebellum derived:toil_matrix
47
+ Brain - Cortex gtex gtex_context Brain - Cortex derived:toil_matrix
48
+ Brain - Frontal Cortex (Ba9) gtex gtex_context Brain - Frontal Cortex (Ba9) derived:toil_matrix
49
+ Brain - Hippocampus gtex gtex_context Brain - Hippocampus derived:toil_matrix
50
+ Brain - Hypothalamus gtex gtex_context Brain - Hypothalamus derived:toil_matrix
51
+ Brain - Nucleus Accumbens (Basal Ganglia) gtex gtex_context Brain - Nucleus Accumbens (Basal Ganglia) derived:toil_matrix
52
+ Brain - Putamen (Basal Ganglia) gtex gtex_context Brain - Putamen (Basal Ganglia) derived:toil_matrix
53
+ Brain - Spinal Cord (Cervical C-1) gtex gtex_context Brain - Spinal Cord (Cervical C-1) derived:toil_matrix
54
+ Brain - Substantia Nigra gtex gtex_context Brain - Substantia Nigra derived:toil_matrix
55
+ Brain Lower Grade Glioma disease gtex_context Brain - Amygdala curated:TUMOR_TISSUE
56
+ Brain Lower Grade Glioma disease gtex_context Brain - Anterior Cingulate Cortex (Ba24) curated:TUMOR_TISSUE
57
+ Brain Lower Grade Glioma disease gtex_context Brain - Caudate (Basal Ganglia) curated:TUMOR_TISSUE
58
+ Brain Lower Grade Glioma disease gtex_context Brain - Cerebellar Hemisphere curated:TUMOR_TISSUE
59
+ Brain Lower Grade Glioma disease gtex_context Brain - Cerebellum curated:TUMOR_TISSUE
60
+ Brain Lower Grade Glioma disease gtex_context Brain - Cortex curated:TUMOR_TISSUE
61
+ Brain Lower Grade Glioma disease gtex_context Brain - Frontal Cortex (Ba9) curated:TUMOR_TISSUE
62
+ Brain Lower Grade Glioma disease gtex_context Brain - Hippocampus curated:TUMOR_TISSUE
63
+ Brain Lower Grade Glioma disease gtex_context Brain - Hypothalamus curated:TUMOR_TISSUE
64
+ Brain Lower Grade Glioma disease gtex_context Brain - Nucleus Accumbens (Basal Ganglia) curated:TUMOR_TISSUE
65
+ Brain Lower Grade Glioma disease gtex_context Brain - Putamen (Basal Ganglia) curated:TUMOR_TISSUE
66
+ Brain Lower Grade Glioma disease gtex_context Brain - Spinal Cord (Cervical C-1) curated:TUMOR_TISSUE
67
+ Brain Lower Grade Glioma disease gtex_context Brain - Substantia Nigra curated:TUMOR_TISSUE
68
+ Brain Lower Grade Glioma disease tcga_code LGG derived:xena_detailed_category
69
+ Breast site gtex_context Breast - Mammary Tissue curated:TUMOR_TISSUE
70
+ Breast site tcga_code BRCA derived:xena_primary_site
71
+ Breast - Mammary Tissue gtex gtex_context Breast - Mammary Tissue derived:toil_matrix
72
+ Breast Invasive Carcinoma disease gtex_context Breast - Mammary Tissue curated:TUMOR_TISSUE
73
+ Breast Invasive Carcinoma disease tcga_code BRCA derived:xena_detailed_category
74
+ CESC code gtex_context Cervix - Ectocervix curated:TUMOR_TISSUE
75
+ CESC code tcga_code CESC derived:toil_matrix
76
+ CHOL code gtex_context Liver curated:TUMOR_TISSUE
77
+ CHOL code tcga_code CHOL derived:toil_matrix
78
+ COAD code gtex_context Colon - Sigmoid curated:TUMOR_TISSUE
79
+ COAD code gtex_context Colon - Transverse curated:TUMOR_TISSUE
80
+ COAD code tcga_code COAD derived:toil_matrix
81
+ Cells - Ebv-Transformed Lymphocytes gtex gtex_context Cells - Ebv-Transformed Lymphocytes derived:toil_matrix
82
+ Cells - Leukemia Cell Line (Cml) gtex gtex_context Cells - Leukemia Cell Line (Cml) derived:toil_matrix
83
+ Cells - Transformed Fibroblasts gtex gtex_context Cells - Transformed Fibroblasts derived:toil_matrix
84
+ Cervical & Endocervical Cancer disease gtex_context Cervix - Ectocervix curated:TUMOR_TISSUE
85
+ Cervical & Endocervical Cancer disease tcga_code CESC derived:xena_detailed_category
86
+ Cervix site gtex_context Cervix - Ectocervix curated:TUMOR_TISSUE
87
+ Cervix site tcga_code CESC derived:xena_primary_site
88
+ Cervix - Ectocervix gtex gtex_context Cervix - Ectocervix derived:toil_matrix
89
+ Cholangiocarcinoma disease gtex_context Liver curated:TUMOR_TISSUE
90
+ Cholangiocarcinoma disease tcga_code CHOL derived:xena_detailed_category
91
+ Colon site gtex_context Colon - Sigmoid curated:TUMOR_TISSUE
92
+ Colon site gtex_context Colon - Transverse curated:TUMOR_TISSUE
93
+ Colon site tcga_code COAD derived:xena_primary_site
94
+ Colon - Sigmoid gtex gtex_context Colon - Sigmoid derived:toil_matrix
95
+ Colon - Transverse gtex gtex_context Colon - Transverse derived:toil_matrix
96
+ Colon Adenocarcinoma disease gtex_context Colon - Sigmoid curated:TUMOR_TISSUE
97
+ Colon Adenocarcinoma disease gtex_context Colon - Transverse curated:TUMOR_TISSUE
98
+ Colon Adenocarcinoma disease tcga_code COAD derived:xena_detailed_category
99
+ DLBC code gtex_context Spleen curated:TUMOR_TISSUE
100
+ DLBC code gtex_context Whole Blood curated:TUMOR_TISSUE
101
+ DLBC code tcga_code DLBC derived:toil_matrix
102
+ Diffuse Large B-Cell Lymphoma disease gtex_context Spleen curated:TUMOR_TISSUE
103
+ Diffuse Large B-Cell Lymphoma disease gtex_context Whole Blood curated:TUMOR_TISSUE
104
+ Diffuse Large B-Cell Lymphoma disease tcga_code DLBC derived:xena_detailed_category
105
+ ESCA code gtex_context Esophagus - Gastroesophageal Junction curated:TUMOR_TISSUE
106
+ ESCA code gtex_context Esophagus - Mucosa curated:TUMOR_TISSUE
107
+ ESCA code gtex_context Esophagus - Muscularis curated:TUMOR_TISSUE
108
+ ESCA code tcga_code ESCA derived:toil_matrix
109
+ Endometrium site gtex_context Uterus curated:TUMOR_TISSUE
110
+ Endometrium site tcga_code UCEC derived:xena_primary_site
111
+ Esophageal Carcinoma disease gtex_context Esophagus - Gastroesophageal Junction curated:TUMOR_TISSUE
112
+ Esophageal Carcinoma disease gtex_context Esophagus - Mucosa curated:TUMOR_TISSUE
113
+ Esophageal Carcinoma disease gtex_context Esophagus - Muscularis curated:TUMOR_TISSUE
114
+ Esophageal Carcinoma disease tcga_code ESCA derived:xena_detailed_category
115
+ Esophagus site gtex_context Esophagus - Gastroesophageal Junction curated:TUMOR_TISSUE
116
+ Esophagus site gtex_context Esophagus - Mucosa curated:TUMOR_TISSUE
117
+ Esophagus site gtex_context Esophagus - Muscularis curated:TUMOR_TISSUE
118
+ Esophagus site tcga_code ESCA derived:xena_primary_site
119
+ Esophagus - Gastroesophageal Junction gtex gtex_context Esophagus - Gastroesophageal Junction derived:toil_matrix
120
+ Esophagus - Mucosa gtex gtex_context Esophagus - Mucosa derived:toil_matrix
121
+ Esophagus - Muscularis gtex gtex_context Esophagus - Muscularis derived:toil_matrix
122
+ Eye site tcga_code UVM derived:xena_primary_site
123
+ Fallopian Tube gtex gtex_context Fallopian Tube derived:toil_matrix
124
+ GBM code gtex_context Brain - Cortex curated:TUMOR_TISSUE
125
+ GBM code gtex_context Brain - Frontal Cortex (Ba9) curated:TUMOR_TISSUE
126
+ GBM code tcga_code GBM derived:toil_matrix
127
+ Glioblastoma Multiforme disease gtex_context Brain - Cortex curated:TUMOR_TISSUE
128
+ Glioblastoma Multiforme disease gtex_context Brain - Frontal Cortex (Ba9) curated:TUMOR_TISSUE
129
+ Glioblastoma Multiforme disease tcga_code GBM derived:xena_detailed_category
130
+ HNSC code gtex_context Esophagus - Mucosa curated:TUMOR_TISSUE
131
+ HNSC code gtex_context Minor Salivary Gland curated:TUMOR_TISSUE
132
+ HNSC code tcga_code HNSC derived:toil_matrix
133
+ Head & Neck Squamous Cell Carcinoma disease gtex_context Esophagus - Mucosa curated:TUMOR_TISSUE
134
+ Head & Neck Squamous Cell Carcinoma disease gtex_context Minor Salivary Gland curated:TUMOR_TISSUE
135
+ Head & Neck Squamous Cell Carcinoma disease tcga_code HNSC derived:xena_detailed_category
136
+ Head and Neck region site gtex_context Esophagus - Mucosa curated:TUMOR_TISSUE
137
+ Head and Neck region site gtex_context Minor Salivary Gland curated:TUMOR_TISSUE
138
+ Head and Neck region site tcga_code HNSC derived:xena_primary_site
139
+ Heart - Atrial Appendage gtex gtex_context Heart - Atrial Appendage derived:toil_matrix
140
+ Heart - Left Ventricle gtex gtex_context Heart - Left Ventricle derived:toil_matrix
141
+ KICH code gtex_context Kidney - Cortex curated:TUMOR_TISSUE
142
+ KICH code tcga_code KICH derived:toil_matrix
143
+ KIRC code gtex_context Kidney - Cortex curated:TUMOR_TISSUE
144
+ KIRC code tcga_code KIRC derived:toil_matrix
145
+ KIRP code gtex_context Kidney - Cortex curated:TUMOR_TISSUE
146
+ KIRP code tcga_code KIRP derived:toil_matrix
147
+ Kidney site gtex_context Kidney - Cortex curated:TUMOR_TISSUE
148
+ Kidney site tcga_code KICH derived:xena_primary_site
149
+ Kidney site tcga_code KIRC derived:xena_primary_site
150
+ Kidney site tcga_code KIRP derived:xena_primary_site
151
+ Kidney - Cortex gtex gtex_context Kidney - Cortex derived:toil_matrix
152
+ Kidney Chromophobe disease gtex_context Kidney - Cortex curated:TUMOR_TISSUE
153
+ Kidney Chromophobe disease tcga_code KICH derived:xena_detailed_category
154
+ Kidney Clear Cell Carcinoma disease gtex_context Kidney - Cortex curated:TUMOR_TISSUE
155
+ Kidney Clear Cell Carcinoma disease tcga_code KIRC derived:xena_detailed_category
156
+ Kidney Papillary Cell Carcinoma disease gtex_context Kidney - Cortex curated:TUMOR_TISSUE
157
+ Kidney Papillary Cell Carcinoma disease tcga_code KIRP derived:xena_detailed_category
158
+ LAML code gtex_context Whole Blood curated:TUMOR_TISSUE
159
+ LAML code tcga_code LAML derived:toil_matrix
160
+ LGG code gtex_context Brain - Amygdala curated:TUMOR_TISSUE
161
+ LGG code gtex_context Brain - Anterior Cingulate Cortex (Ba24) curated:TUMOR_TISSUE
162
+ LGG code gtex_context Brain - Caudate (Basal Ganglia) curated:TUMOR_TISSUE
163
+ LGG code gtex_context Brain - Cerebellar Hemisphere curated:TUMOR_TISSUE
164
+ LGG code gtex_context Brain - Cerebellum curated:TUMOR_TISSUE
165
+ LGG code gtex_context Brain - Cortex curated:TUMOR_TISSUE
166
+ LGG code gtex_context Brain - Frontal Cortex (Ba9) curated:TUMOR_TISSUE
167
+ LGG code gtex_context Brain - Hippocampus curated:TUMOR_TISSUE
168
+ LGG code gtex_context Brain - Hypothalamus curated:TUMOR_TISSUE
169
+ LGG code gtex_context Brain - Nucleus Accumbens (Basal Ganglia) curated:TUMOR_TISSUE
170
+ LGG code gtex_context Brain - Putamen (Basal Ganglia) curated:TUMOR_TISSUE
171
+ LGG code gtex_context Brain - Spinal Cord (Cervical C-1) curated:TUMOR_TISSUE
172
+ LGG code gtex_context Brain - Substantia Nigra curated:TUMOR_TISSUE
173
+ LGG code tcga_code LGG derived:toil_matrix
174
+ LIHC code gtex_context Liver curated:TUMOR_TISSUE
175
+ LIHC code tcga_code LIHC derived:toil_matrix
176
+ LUAD code gtex_context Lung curated:TUMOR_TISSUE
177
+ LUAD code tcga_code LUAD derived:toil_matrix
178
+ LUSC code gtex_context Lung curated:TUMOR_TISSUE
179
+ LUSC code tcga_code LUSC derived:toil_matrix
180
+ Lining of body cavities site tcga_code MESO derived:xena_primary_site
181
+ Liver gtex gtex_context Liver derived:toil_matrix
182
+ Liver site gtex_context Liver curated:TUMOR_TISSUE
183
+ Liver site tcga_code LIHC derived:xena_primary_site
184
+ Liver Hepatocellular Carcinoma disease gtex_context Liver curated:TUMOR_TISSUE
185
+ Liver Hepatocellular Carcinoma disease tcga_code LIHC derived:xena_detailed_category
186
+ Lung gtex gtex_context Lung derived:toil_matrix
187
+ Lung site gtex_context Lung curated:TUMOR_TISSUE
188
+ Lung site tcga_code LUAD derived:xena_primary_site
189
+ Lung site tcga_code LUSC derived:xena_primary_site
190
+ Lung Adenocarcinoma disease gtex_context Lung curated:TUMOR_TISSUE
191
+ Lung Adenocarcinoma disease tcga_code LUAD derived:xena_detailed_category
192
+ Lung Squamous Cell Carcinoma disease gtex_context Lung curated:TUMOR_TISSUE
193
+ Lung Squamous Cell Carcinoma disease tcga_code LUSC derived:xena_detailed_category
194
+ Lymphatic tissue site gtex_context Spleen curated:TUMOR_TISSUE
195
+ Lymphatic tissue site gtex_context Whole Blood curated:TUMOR_TISSUE
196
+ Lymphatic tissue site tcga_code DLBC derived:xena_primary_site
197
+ MESO code tcga_code MESO derived:toil_matrix
198
+ Mesothelioma disease tcga_code MESO derived:xena_detailed_category
199
+ Minor Salivary Gland gtex gtex_context Minor Salivary Gland derived:toil_matrix
200
+ Muscle - Skeletal gtex gtex_context Muscle - Skeletal derived:toil_matrix
201
+ Nerve - Tibial gtex gtex_context Nerve - Tibial derived:toil_matrix
202
+ OV code gtex_context Fallopian Tube curated:TUMOR_TISSUE
203
+ OV code gtex_context Ovary curated:TUMOR_TISSUE
204
+ OV code tcga_code OV derived:toil_matrix
205
+ Ovarian Serous Cystadenocarcinoma disease gtex_context Fallopian Tube curated:TUMOR_TISSUE
206
+ Ovarian Serous Cystadenocarcinoma disease gtex_context Ovary curated:TUMOR_TISSUE
207
+ Ovarian Serous Cystadenocarcinoma disease tcga_code OV derived:xena_detailed_category
208
+ Ovary gtex gtex_context Ovary derived:toil_matrix
209
+ Ovary site gtex_context Fallopian Tube curated:TUMOR_TISSUE
210
+ Ovary site gtex_context Ovary curated:TUMOR_TISSUE
211
+ Ovary site tcga_code OV derived:xena_primary_site
212
+ PAAD code gtex_context Pancreas curated:TUMOR_TISSUE
213
+ PAAD code tcga_code PAAD derived:toil_matrix
214
+ PCPG code gtex_context Adrenal Gland curated:TUMOR_TISSUE
215
+ PCPG code tcga_code PCPG derived:toil_matrix
216
+ PRAD code gtex_context Prostate curated:TUMOR_TISSUE
217
+ PRAD code tcga_code PRAD derived:toil_matrix
218
+ Pancreas gtex gtex_context Pancreas derived:toil_matrix
219
+ Pancreas site gtex_context Pancreas curated:TUMOR_TISSUE
220
+ Pancreas site tcga_code PAAD derived:xena_primary_site
221
+ Pancreatic Adenocarcinoma disease gtex_context Pancreas curated:TUMOR_TISSUE
222
+ Pancreatic Adenocarcinoma disease tcga_code PAAD derived:xena_detailed_category
223
+ Paraganglia site gtex_context Adrenal Gland curated:TUMOR_TISSUE
224
+ Paraganglia site tcga_code PCPG derived:xena_primary_site
225
+ Pheochromocytoma & Paraganglioma disease gtex_context Adrenal Gland curated:TUMOR_TISSUE
226
+ Pheochromocytoma & Paraganglioma disease tcga_code PCPG derived:xena_detailed_category
227
+ Pituitary gtex gtex_context Pituitary derived:toil_matrix
228
+ Prostate gtex gtex_context Prostate derived:toil_matrix
229
+ Prostate site gtex_context Prostate curated:TUMOR_TISSUE
230
+ Prostate site tcga_code PRAD derived:xena_primary_site
231
+ Prostate Adenocarcinoma disease gtex_context Prostate curated:TUMOR_TISSUE
232
+ Prostate Adenocarcinoma disease tcga_code PRAD derived:xena_detailed_category
233
+ READ code gtex_context Colon - Sigmoid curated:TUMOR_TISSUE
234
+ READ code gtex_context Colon - Transverse curated:TUMOR_TISSUE
235
+ READ code tcga_code READ derived:toil_matrix
236
+ Rectum site gtex_context Colon - Sigmoid curated:TUMOR_TISSUE
237
+ Rectum site gtex_context Colon - Transverse curated:TUMOR_TISSUE
238
+ Rectum site tcga_code READ derived:xena_primary_site
239
+ Rectum Adenocarcinoma disease gtex_context Colon - Sigmoid curated:TUMOR_TISSUE
240
+ Rectum Adenocarcinoma disease gtex_context Colon - Transverse curated:TUMOR_TISSUE
241
+ Rectum Adenocarcinoma disease tcga_code READ derived:xena_detailed_category
242
+ SARC code tcga_code SARC derived:toil_matrix
243
+ SKCM code gtex_context Skin - Not Sun Exposed (Suprapubic) curated:TUMOR_TISSUE
244
+ SKCM code gtex_context Skin - Sun Exposed (Lower Leg) curated:TUMOR_TISSUE
245
+ SKCM code tcga_code SKCM derived:toil_matrix
246
+ STAD code gtex_context Stomach curated:TUMOR_TISSUE
247
+ STAD code tcga_code STAD derived:toil_matrix
248
+ Sarcoma disease tcga_code SARC derived:xena_detailed_category
249
+ Skin site gtex_context Skin - Not Sun Exposed (Suprapubic) curated:TUMOR_TISSUE
250
+ Skin site gtex_context Skin - Sun Exposed (Lower Leg) curated:TUMOR_TISSUE
251
+ Skin site tcga_code SKCM derived:xena_primary_site
252
+ Skin - Not Sun Exposed (Suprapubic) gtex gtex_context Skin - Not Sun Exposed (Suprapubic) derived:toil_matrix
253
+ Skin - Sun Exposed (Lower Leg) gtex gtex_context Skin - Sun Exposed (Lower Leg) derived:toil_matrix
254
+ Skin Cutaneous Melanoma disease gtex_context Skin - Not Sun Exposed (Suprapubic) curated:TUMOR_TISSUE
255
+ Skin Cutaneous Melanoma disease gtex_context Skin - Sun Exposed (Lower Leg) curated:TUMOR_TISSUE
256
+ Skin Cutaneous Melanoma disease tcga_code SKCM derived:xena_detailed_category
257
+ Small Intestine - Terminal Ileum gtex gtex_context Small Intestine - Terminal Ileum derived:toil_matrix
258
+ Soft tissue,Bone site tcga_code SARC derived:xena_primary_site
259
+ Spleen gtex gtex_context Spleen derived:toil_matrix
260
+ Stomach gtex gtex_context Stomach derived:toil_matrix
261
+ Stomach site gtex_context Stomach curated:TUMOR_TISSUE
262
+ Stomach site tcga_code STAD derived:xena_primary_site
263
+ Stomach Adenocarcinoma disease gtex_context Stomach curated:TUMOR_TISSUE
264
+ Stomach Adenocarcinoma disease tcga_code STAD derived:xena_detailed_category
265
+ TGCT code gtex_context Testis curated:TUMOR_TISSUE
266
+ TGCT code tcga_code TGCT derived:toil_matrix
267
+ THCA code gtex_context Thyroid curated:TUMOR_TISSUE
268
+ THCA code tcga_code THCA derived:toil_matrix
269
+ THYM code tcga_code THYM derived:toil_matrix
270
+ Testicular Germ Cell Tumor disease gtex_context Testis curated:TUMOR_TISSUE
271
+ Testicular Germ Cell Tumor disease tcga_code TGCT derived:xena_detailed_category
272
+ Testis gtex gtex_context Testis derived:toil_matrix
273
+ Testis site gtex_context Testis curated:TUMOR_TISSUE
274
+ Testis site tcga_code TGCT derived:xena_primary_site
275
+ Thymoma disease tcga_code THYM derived:xena_detailed_category
276
+ Thymus site tcga_code THYM derived:xena_primary_site
277
+ Thyroid gtex gtex_context Thyroid derived:toil_matrix
278
+ Thyroid Carcinoma disease gtex_context Thyroid curated:TUMOR_TISSUE
279
+ Thyroid Carcinoma disease tcga_code THCA derived:xena_detailed_category
280
+ Thyroid Gland site gtex_context Thyroid curated:TUMOR_TISSUE
281
+ Thyroid Gland site tcga_code THCA derived:xena_primary_site
282
+ UCEC code gtex_context Uterus curated:TUMOR_TISSUE
283
+ UCEC code tcga_code UCEC derived:toil_matrix
284
+ UCS code gtex_context Uterus curated:TUMOR_TISSUE
285
+ UCS code tcga_code UCS derived:toil_matrix
286
+ UVM code tcga_code UVM derived:toil_matrix
287
+ Uterine Carcinosarcoma disease gtex_context Uterus curated:TUMOR_TISSUE
288
+ Uterine Carcinosarcoma disease tcga_code UCS derived:xena_detailed_category
289
+ Uterine Corpus Endometrioid Carcinoma disease gtex_context Uterus curated:TUMOR_TISSUE
290
+ Uterine Corpus Endometrioid Carcinoma disease tcga_code UCEC derived:xena_detailed_category
291
+ Uterus gtex gtex_context Uterus derived:toil_matrix
292
+ Uterus site gtex_context Uterus curated:TUMOR_TISSUE
293
+ Uterus site tcga_code UCS derived:xena_primary_site
294
+ Uveal Melanoma disease tcga_code UVM derived:xena_detailed_category
295
+ Vagina gtex gtex_context Vagina derived:toil_matrix
296
+ White blood cell site gtex_context Whole Blood curated:TUMOR_TISSUE
297
+ White blood cell site tcga_code LAML derived:xena_primary_site
298
+ Whole Blood gtex gtex_context Whole Blood derived:toil_matrix
299
+ adrenal gtex gtex_context Adrenal Gland curated:spelling
300
+ adrenal site gtex_context Adrenal Gland curated:spelling
301
+ adrenal site tcga_code ACC curated:spelling
302
+ biliary site gtex_context Liver curated:spelling
303
+ biliary site tcga_code CHOL curated:spelling
304
+ blood site gtex_context Whole Blood curated:spelling
305
+ blood site tcga_code LAML curated:spelling
306
+ bone site tcga_code SARC curated:spelling
307
+ cerebral site gtex_context Brain - Amygdala curated:spelling
308
+ cerebral site gtex_context Brain - Anterior Cingulate Cortex (Ba24) curated:spelling
309
+ cerebral site gtex_context Brain - Caudate (Basal Ganglia) curated:spelling
310
+ cerebral site gtex_context Brain - Cerebellar Hemisphere curated:spelling
311
+ cerebral site gtex_context Brain - Cerebellum curated:spelling
312
+ cerebral site gtex_context Brain - Cortex curated:spelling
313
+ cerebral site gtex_context Brain - Frontal Cortex (Ba9) curated:spelling
314
+ cerebral site gtex_context Brain - Hippocampus curated:spelling
315
+ cerebral site gtex_context Brain - Hypothalamus curated:spelling
316
+ cerebral site gtex_context Brain - Nucleus Accumbens (Basal Ganglia) curated:spelling
317
+ cerebral site gtex_context Brain - Putamen (Basal Ganglia) curated:spelling
318
+ cerebral site gtex_context Brain - Spinal Cord (Cervical C-1) curated:spelling
319
+ cerebral site gtex_context Brain - Substantia Nigra curated:spelling
320
+ cerebral site tcga_code GBM curated:spelling
321
+ cerebral site tcga_code LGG curated:spelling
322
+ cervical site gtex_context Cervix - Ectocervix curated:spelling
323
+ cervical site tcga_code CESC curated:spelling
324
+ cholangiocarcinoma site gtex_context Liver curated:spelling
325
+ cholangiocarcinoma site tcga_code CHOL curated:spelling
326
+ colonic site gtex_context Colon - Sigmoid curated:spelling
327
+ colonic site gtex_context Colon - Transverse curated:spelling
328
+ colonic site tcga_code COAD curated:spelling
329
+ colorectal site gtex_context Colon - Sigmoid curated:spelling
330
+ colorectal site gtex_context Colon - Transverse curated:spelling
331
+ colorectal site tcga_code COAD curated:spelling
332
+ colorectum site gtex_context Colon - Sigmoid curated:spelling
333
+ colorectum site gtex_context Colon - Transverse curated:spelling
334
+ colorectum site tcga_code COAD curated:spelling
335
+ cutaneous site gtex_context Skin - Not Sun Exposed (Suprapubic) curated:spelling
336
+ cutaneous site gtex_context Skin - Sun Exposed (Lower Leg) curated:spelling
337
+ cutaneous site tcga_code SKCM curated:spelling
338
+ endometrial site gtex_context Uterus curated:spelling
339
+ endometrial site tcga_code UCEC curated:spelling
340
+ esophageal site gtex_context Esophagus - Gastroesophageal Junction curated:spelling
341
+ esophageal site gtex_context Esophagus - Mucosa curated:spelling
342
+ esophageal site gtex_context Esophagus - Muscularis curated:spelling
343
+ esophageal site tcga_code ESCA curated:spelling
344
+ gastric gtex gtex_context Stomach curated:spelling
345
+ gastric site gtex_context Stomach curated:spelling
346
+ gastric site tcga_code STAD curated:spelling
347
+ glioblastoma site gtex_context Brain - Amygdala curated:spelling
348
+ glioblastoma site gtex_context Brain - Anterior Cingulate Cortex (Ba24) curated:spelling
349
+ glioblastoma site gtex_context Brain - Caudate (Basal Ganglia) curated:spelling
350
+ glioblastoma site gtex_context Brain - Cerebellar Hemisphere curated:spelling
351
+ glioblastoma site gtex_context Brain - Cerebellum curated:spelling
352
+ glioblastoma site gtex_context Brain - Cortex curated:spelling
353
+ glioblastoma site gtex_context Brain - Frontal Cortex (Ba9) curated:spelling
354
+ glioblastoma site gtex_context Brain - Hippocampus curated:spelling
355
+ glioblastoma site gtex_context Brain - Hypothalamus curated:spelling
356
+ glioblastoma site gtex_context Brain - Nucleus Accumbens (Basal Ganglia) curated:spelling
357
+ glioblastoma site gtex_context Brain - Putamen (Basal Ganglia) curated:spelling
358
+ glioblastoma site gtex_context Brain - Spinal Cord (Cervical C-1) curated:spelling
359
+ glioblastoma site gtex_context Brain - Substantia Nigra curated:spelling
360
+ glioblastoma site tcga_code GBM curated:spelling
361
+ glioblastoma site tcga_code LGG curated:spelling
362
+ glioma site gtex_context Brain - Amygdala curated:spelling
363
+ glioma site gtex_context Brain - Anterior Cingulate Cortex (Ba24) curated:spelling
364
+ glioma site gtex_context Brain - Caudate (Basal Ganglia) curated:spelling
365
+ glioma site gtex_context Brain - Cerebellar Hemisphere curated:spelling
366
+ glioma site gtex_context Brain - Cerebellum curated:spelling
367
+ glioma site gtex_context Brain - Cortex curated:spelling
368
+ glioma site gtex_context Brain - Frontal Cortex (Ba9) curated:spelling
369
+ glioma site gtex_context Brain - Hippocampus curated:spelling
370
+ glioma site gtex_context Brain - Hypothalamus curated:spelling
371
+ glioma site gtex_context Brain - Nucleus Accumbens (Basal Ganglia) curated:spelling
372
+ glioma site gtex_context Brain - Putamen (Basal Ganglia) curated:spelling
373
+ glioma site gtex_context Brain - Spinal Cord (Cervical C-1) curated:spelling
374
+ glioma site gtex_context Brain - Substantia Nigra curated:spelling
375
+ glioma site tcga_code GBM curated:spelling
376
+ glioma site tcga_code LGG curated:spelling
377
+ head and neck site gtex_context Esophagus - Mucosa curated:spelling
378
+ head and neck site gtex_context Minor Salivary Gland curated:spelling
379
+ head and neck site tcga_code HNSC curated:spelling
380
+ hepatic gtex gtex_context Liver curated:spelling
381
+ hepatic site gtex_context Liver curated:spelling
382
+ hepatic site tcga_code LIHC curated:spelling
383
+ hepatocellular gtex gtex_context Liver curated:spelling
384
+ hepatocellular site gtex_context Liver curated:spelling
385
+ hepatocellular site tcga_code LIHC curated:spelling
386
+ leukaemia site gtex_context Whole Blood curated:spelling
387
+ leukaemia site tcga_code LAML curated:spelling
388
+ leukemia site gtex_context Whole Blood curated:spelling
389
+ leukemia site tcga_code LAML curated:spelling
390
+ lymphoma site gtex_context Spleen curated:spelling
391
+ lymphoma site gtex_context Whole Blood curated:spelling
392
+ lymphoma site tcga_code DLBC curated:spelling
393
+ mammary site gtex_context Breast - Mammary Tissue curated:spelling
394
+ mammary site tcga_code BRCA curated:spelling
395
+ melanoma site gtex_context Skin - Not Sun Exposed (Suprapubic) curated:spelling
396
+ melanoma site gtex_context Skin - Sun Exposed (Lower Leg) curated:spelling
397
+ melanoma site tcga_code SKCM curated:spelling
398
+ mesothelioma site tcga_code MESO curated:spelling
399
+ ocular site tcga_code UVM curated:spelling
400
+ oesophageal site gtex_context Esophagus - Gastroesophageal Junction curated:spelling
401
+ oesophageal site gtex_context Esophagus - Mucosa curated:spelling
402
+ oesophageal site gtex_context Esophagus - Muscularis curated:spelling
403
+ oesophageal site tcga_code ESCA curated:spelling
404
+ oesophagus site gtex_context Esophagus - Gastroesophageal Junction curated:spelling
405
+ oesophagus site gtex_context Esophagus - Mucosa curated:spelling
406
+ oesophagus site gtex_context Esophagus - Muscularis curated:spelling
407
+ oesophagus site tcga_code ESCA curated:spelling
408
+ ovarian gtex gtex_context Ovary curated:spelling
409
+ ovarian site gtex_context Fallopian Tube curated:spelling
410
+ ovarian site gtex_context Ovary curated:spelling
411
+ ovarian site tcga_code OV curated:spelling
412
+ pancreatic gtex gtex_context Pancreas curated:spelling
413
+ pancreatic site gtex_context Pancreas curated:spelling
414
+ pancreatic site tcga_code PAAD curated:spelling
415
+ paraganglioma site gtex_context Adrenal Gland curated:spelling
416
+ paraganglioma site tcga_code PCPG curated:spelling
417
+ pheochromocytoma site gtex_context Adrenal Gland curated:spelling
418
+ pheochromocytoma site tcga_code PCPG curated:spelling
419
+ prostatic gtex gtex_context Prostate curated:spelling
420
+ prostatic site gtex_context Prostate curated:spelling
421
+ prostatic site tcga_code PRAD curated:spelling
422
+ pulmonary gtex gtex_context Lung curated:spelling
423
+ pulmonary site gtex_context Lung curated:spelling
424
+ pulmonary site tcga_code LUAD curated:spelling
425
+ pulmonary site tcga_code LUSC curated:spelling
426
+ rectal site gtex_context Colon - Sigmoid curated:spelling
427
+ rectal site gtex_context Colon - Transverse curated:spelling
428
+ rectal site tcga_code READ curated:spelling
429
+ renal site gtex_context Kidney - Cortex curated:spelling
430
+ renal site tcga_code KICH curated:spelling
431
+ renal site tcga_code KIRC curated:spelling
432
+ renal site tcga_code KIRP curated:spelling
433
+ sarcoma site tcga_code SARC curated:spelling
434
+ soft tissue site tcga_code SARC curated:spelling
435
+ testicular gtex gtex_context Testis curated:spelling
436
+ testicular site gtex_context Testis curated:spelling
437
+ testicular site tcga_code TGCT curated:spelling
438
+ thymic site tcga_code THYM curated:spelling
439
+ thyroid site gtex_context Thyroid curated:spelling
440
+ thyroid site tcga_code THCA curated:spelling
441
+ urothelial gtex gtex_context Bladder curated:spelling
442
+ urothelial site gtex_context Bladder curated:spelling
443
+ urothelial site tcga_code BLCA curated:spelling
444
+ uterine gtex gtex_context Uterus curated:spelling
445
+ uterine site gtex_context Uterus curated:spelling
446
+ uterine site tcga_code UCS curated:spelling
447
+ uterine corpus site gtex_context Uterus curated:spelling
448
+ uterine corpus site tcga_code UCEC curated:spelling
449
+ uveal site tcga_code UVM curated:spelling
expression/protein_abundance_mmu.tsv.gz ADDED
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1
+ source context q05 q10 q25 n_genes n_samples
2
+ toil_gtex Adipose - Subcutaneous 0.040000916 0.08000192 0.2900018 26040 318
3
+ toil_gtex Adipose - Visceral (Omentum) 0.040000916 0.070001826 0.2599961 26119 197
4
+ toil_gtex Adrenal Gland 0.02999974 0.06000199 0.23999915 26015 128
5
+ toil_gtex Artery - Aorta 0.040000916 0.08999935 0.31998995 25524 207
6
+ toil_gtex Artery - Coronary 0.040000916 0.08000192 0.31000295 25751 118
7
+ toil_gtex Artery - Tibial 0.040000916 0.08000192 0.3299993 24589 281
8
+ toil_gtex Bladder 0.049998555 0.099998094 0.36000732 26736 9
9
+ toil_gtex Brain - Amygdala 0.02999974 0.06000199 0.23999915 26046 69
10
+ toil_gtex Brain - Anterior Cingulate Cortex (Ba24) 0.02999974 0.06000199 0.2500071 26594 83
11
+ toil_gtex Brain - Caudate (Basal Ganglia) 0.02999974 0.06000199 0.23999915 27075 109
12
+ toil_gtex Brain - Cerebellar Hemisphere 0.049998555 0.08999935 0.34998956 27122 97
13
+ toil_gtex Brain - Cerebellum 0.049998555 0.099998094 0.3700023 27425 119
14
+ toil_gtex Brain - Cortex 0.040000916 0.08000192 0.2799918 27159 105
15
+ toil_gtex Brain - Frontal Cortex (Ba9) 0.040000916 0.070001826 0.2749992 26770 102
16
+ toil_gtex Brain - Hippocampus 0.02999974 0.06000199 0.23999915 26358 84
17
+ toil_gtex Brain - Hypothalamus 0.030000996 0.070001826 0.2599961 27235 82
18
+ toil_gtex Brain - Nucleus Accumbens (Basal Ganglia) 0.02999974 0.06000199 0.24500312 27068 104
19
+ toil_gtex Brain - Putamen (Basal Ganglia) 0.02999974 0.06000199 0.23000562 25961 81
20
+ toil_gtex Brain - Spinal Cord (Cervical C-1) 0.040000916 0.07500187 0.2849968 26134 60
21
+ toil_gtex Brain - Substantia Nigra 0.02999974 0.070001826 0.2599961 26187 57
22
+ toil_gtex Breast - Mammary Tissue 0.040000916 0.08000192 0.29999262 27044 179
23
+ toil_gtex Cells - Ebv-Transformed Lymphocytes 0.019999515 0.040000916 0.15999635 25259 107
24
+ toil_gtex Cells - Leukemia Cell Line (Cml) 0.019999515 0.044999734 0.17000377 24871 70
25
+ toil_gtex Cells - Transformed Fibroblasts 0.02999974 0.06000199 0.23000562 24328 256
26
+ toil_gtex Cervix - Ectocervix 0.030000215 0.06000088 0.23999858 28094 6
27
+ toil_gtex Colon - Sigmoid 0.040000916 0.08000192 0.29999262 25948 141
28
+ toil_gtex Colon - Transverse 0.040000916 0.08000192 0.27000663 26613 167
29
+ toil_gtex Esophagus - Gastroesophageal Junction 0.040000916 0.08000192 0.2900018 25786 137
30
+ toil_gtex Esophagus - Mucosa 0.040000916 0.070001826 0.27000663 25938 273
31
+ toil_gtex Esophagus - Muscularis 0.040000916 0.08000192 0.2900018 25631 245
32
+ toil_gtex Fallopian Tube 0.06000199 0.10999756 0.38000983 28082 5
33
+ toil_gtex Heart - Atrial Appendage 0.02999974 0.06000199 0.20000455 25363 175
34
+ toil_gtex Heart - Left Ventricle 0.019999515 0.040000916 0.15999635 23986 202
35
+ toil_gtex Kidney - Cortex 0.024999628 0.050001454 0.20999782 26516 28
36
+ toil_gtex Liver 0.02999974 0.049998555 0.18000482 24649 110
37
+ toil_gtex Lung 0.049998555 0.08999935 0.34998956 27206 288
38
+ toil_gtex Minor Salivary Gland 0.040000916 0.08000192 0.27000663 27258 55
39
+ toil_gtex Muscle - Skeletal 0.02999974 0.049998555 0.18000482 23401 396
40
+ toil_gtex Nerve - Tibial 0.049998555 0.099998094 0.3700023 27034 278
41
+ toil_gtex Ovary 0.040000916 0.07500187 0.29499722 26838 88
42
+ toil_gtex Pancreas 0.02999974 0.049998555 0.18000482 25047 167
43
+ toil_gtex Pituitary 0.049998555 0.099998094 0.36000732 28477 107
44
+ toil_gtex Prostate 0.044999734 0.08999935 0.3499985 27692 100
45
+ toil_gtex Skin - Not Sun Exposed (Suprapubic) 0.040000916 0.08000192 0.2799918 26707 233
46
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47
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48
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49
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50
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51
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52
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53
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54
+ toil_gtex Whole Blood 0.019999515 0.02999974 0.099998094 21838 337
55
+ toil_gtex __pooled__ 0.040000916 0.070001826 0.27000663 1402705 396
56
+ toil_tcga ACC 0.019999515 0.040000916 0.17000377 24378 76
57
+ toil_tcga BLCA 0.019999515 0.040000916 0.17000377 26115 403
58
+ toil_tcga BRCA 0.019999515 0.049998555 0.1899941 27126 1087
59
+ toil_tcga CESC 0.019999515 0.040000916 0.17000377 26331 301
60
+ toil_tcga CHOL 0.024999628 0.049999047 0.20999782 26374 36
61
+ toil_tcga COAD 0.019999515 0.049998555 0.1750043 26169 282
62
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63
+ toil_tcga ESCA 0.019999515 0.040000916 0.14000322 30561 180
64
+ toil_tcga GBM 0.02999974 0.070001826 0.2500071 27300 153
65
+ toil_tcga HNSC 0.019999515 0.040000916 0.15999635 25899 514
66
+ toil_tcga KICH 0.019999515 0.02999974 0.12000176 25726 65
67
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68
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69
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70
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71
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72
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73
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74
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75
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76
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77
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78
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81
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82
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83
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87
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89
+ toil_tcga __pooled__ 0.019999515 0.049998555 0.18000482 882959 1087
expression/toil_floors_mmu.tsv ADDED
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expression/toil_matrix.npz ADDED
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expression/tumor_expression_mmu.tsv.gz ADDED
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hla_loh/SOURCES.md ADDED
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1
+ # `hla_loh/` — sources
2
+
3
+ Per-allele HLA-I **loss of heterozygosity** and genetic immune escape, over 6,319 tumours
4
+ (4,439 Hartwig metastatic + 1,880 PCAWG primary), plus the join that carries the PCAWG arm onto
5
+ TCGA barcodes so it meets `tcga/tcga_neoantigens.tsv.gz`.
6
+
7
+ The question this directory exists to answer: **when a tumour loses one HLA allele, is the lost one
8
+ the allele that better presented its neoantigens?** That is the escape mechanism a polyepitope
9
+ cassette has to be designed against, and `bench/cassette/tcga_loh_perallele.py` in the benchmark
10
+ repo is the consumer.
11
+
12
+ ## Provenance in one line
13
+
14
+ Everything here is **derived** — LILAC allele-specific copy-number calls on whole-genome sequencing,
15
+ published as supplementary data — except `do_to_tcga_barcode.tsv`, which is a lookup we performed
16
+ against the GDC API. Nothing in this directory is a measurement we made.
17
+
18
+ **Primary source.** Martínez-Jiménez F, Priestley P, Shale C, Baber J, Rozemuller E, Cuppen E.
19
+ *Genetic immune escape landscape in primary and metastatic cancer.* **Nature Genetics 55(5):820–831
20
+ (2023)**, [doi:10.1038/s41588-023-01367-1](https://doi.org/10.1038/s41588-023-01367-1).
21
+
22
+ ## What ships, and what deliberately does not
23
+
24
+ **The publisher's workbooks are not redistributable; the facts in them are.** The three
25
+ `41588_2023_1367_MOESM*.xlsx` are copyrighted Springer Nature supplementary material. They are kept
26
+ in `raw/hla_loh/` (gitignored) and `harmonize.py` rebuilds neutral long-format parquet from them.
27
+ This is the same call already taken for `icb_cohorts/`: harmonised output published, publisher
28
+ workbook held back. An `.xlsx` is a layout and a copyrightable compilation; a table of allele copy
29
+ numbers is neither.
30
+
31
+ ### Published (this directory)
32
+
33
+ | file | rows × cols | what it is | provenance |
34
+ |---|---|---|---|
35
+ | `gie_allele.parquet` | 37,914 × 9 | one row per **(sample, allele slot)** — 6,319 samples × the six slots `A1 A2 B1 B2 C1 C2`. Carries `allele`, `copy_number`, `n_mutations`, plus `cohort`/`cancer_type`. This is Supplementary Data 2 unpivoted | derived (LILAC on WGS) |
36
+ | `gie_sample.parquet` | 6,319 × 32 | one row per **sample**: the escape read-outs — `loh_lilac`, `copy_number_neutral_loh_hla`, `loh_focal`/`loh_nonfocal`/`loh_hfocal`, `mut_hla_lilac`, the five pathway flags with their detail strings, `genetic_immune_escape`, per-locus `ploidy_minor`/`ploidy_major`/`del_HLA-*` | derived |
37
+ | `hla_typing_callers.parquet` | 226,008 × 8 | one row per **(sample, caller, tissue, slot)** — 6,278 samples × 3 callers (`LILAC`, `xHLA`, `polysolver`) × normal/tumour × 6 slots. Supplementary Data 1, unpivoted. The caller-concordance reference; **not** used in the join | derived |
38
+ | `sample_metadata.parquet` | 7,619 × 40 | per-sample cohort, cancer type, biopsy site, treatment history, MSI/HR status and mutation load (`sbs`/`dbs`/`indel`/`sv`, clonal and subclonal). Supplementary Table 2 | derived |
39
+ | `escape_genes.parquet` | 21 × 6 | the immune-escape gene panel the paper's calls are made over, with pathway assignment and admissible mutation types. Supplementary Table 1 | curated gene list |
40
+ | `perallele_tcga_loh.parquet` | 2,747 × 3 | **the deliverable the benchmark consumes.** `donor` (TCGA barcode), `hla_allele`, `allele_lost`. 481 TCGA donors overlapping `bench/cassette/scored.parquet`; 238 loss calls | derived (join, 2026-09-11) |
41
+ | `do_to_tcga_barcode.tsv` | 711 × 4 | `icgc_donor_id` → `case_uuid` → `tcga_barcode` → `gdc_project`, for the PCAWG donors that are TCGA cases | derived (GDC `/cases` lookup) |
42
+ | `harmonize.py` | — | rebuilds the five parquets above from `raw/hla_loh/` | — |
43
+
44
+ ### Held back in `raw/hla_loh/` (gitignored)
45
+
46
+ | file | why it is not published |
47
+ |---|---|
48
+ | `41588_2023_1367_MOESM4_ESM.xlsx` | copyrighted Springer Nature supplement (Supplementary Tables 1–2) |
49
+ | `41588_2023_1367_MOESM5_ESM.xlsx` | copyrighted (Supplementary Data 1) |
50
+ | `41588_2023_1367_MOESM6_ESM.xlsx` | copyrighted (Supplementary Data 2) |
51
+ | `pcawg_sample_sheet.2016-10-18.tsv` | third-party PCAWG release file, re-fetchable (command below); an input, not our output |
52
+
53
+ **The Hartwig arm of the source study needs its own access application** for the underlying
54
+ sequencing. Nothing here is that sequencing — these are the published per-sample call tables.
55
+
56
+ ## Identifiers, and why they are public
57
+
58
+ - **PCAWG samples are keyed `DO#####`** (`icgc_donor_id`), Hartwig samples `HMF#####A`. Both are
59
+ pseudonymous research identifiers assigned by the consortium, as published.
60
+ - **TCGA barcodes** (`TCGA-XX-XXXX`) are likewise public — `tcga/` in this compendium already ships
61
+ per-barcode HLA and neoantigen calls on the same basis (open-access somatic + expression, as in
62
+ TCIA / Thorsson 2018).
63
+ - PCAWG's `donor_wgs_exclusion_white_gray` is the consortium's **published QC whitelist/greylist**
64
+ flag, not a consent tier. An earlier `.gitignore` rule in this repo described this directory as a
65
+ re-identification risk; that reading was wrong and has been withdrawn.
66
+
67
+ ## Three things a consumer has to know
68
+
69
+ **1. `n_mutations` is a count, not a flag.** `*_MUT` takes `0.0, 0.5, 1.0, 1.5, 2.0, 3.0, 4.0`
70
+ across the six slots (37,707 zero, 207 non-zero, 42 of them non-integral). Reading it as a boolean
71
+ loses every fractional and multi-hit call.
72
+
73
+ **2. Four columns are comma-separated locus lists, not booleans.**
74
+ `copy_number_neutral_loh_hla`, `loh_focal`, `loh_nonfocal`, `loh_hfocal` carry values like
75
+ `HLA-A,HLA-C,HLA-B`. They sit beside ten genuine booleans in the same sheet and look identical at a
76
+ glance; `copy_number_neutral_loh_hla` has 784 non-null values that a bool cast would have silently
77
+ nulled. They are kept verbatim as the source wrote them.
78
+
79
+ **3. The allele vocabulary is this repo's, not the workbook's.** The workbooks write `A*02:01`;
80
+ `gie_allele.parquet` and `hla_typing_callers.parquet` write **`HLA-A*02:01`**, matching
81
+ `perallele_tcga_loh.parquet`, `tcga/`, and `mhcmatch`. Two spellings for one molecule is the
82
+ recurring failure mode in this compendium, and a lookup that silently matches neither returns the
83
+ same shape as "nothing was presented".
84
+
85
+ ## The join
86
+
87
+ ```
88
+ gie_sample / gie_allele (cohort = PCAWG, sample_id = DO#####)
89
+ │ icgc_donor_id
90
+ ▼
91
+ do_to_tcga_barcode.tsv 711 PCAWG donors that are TCGA cases
92
+ │ tcga_barcode
93
+ ▼
94
+ perallele_tcga_loh.parquet ──► bench/cassette/scored.parquet (481 donors overlap)
95
+ ```
96
+
97
+ All 711 bridge rows join to a PCAWG sample in `gie_sample.parquet`, and all 481 donors in
98
+ `perallele_tcga_loh.parquet` are present in the bridge. The Hartwig arm has no TCGA counterpart and
99
+ does not enter the join.
100
+
101
+ ## Regenerate
102
+
103
+ ```zsh
104
+ # the five harmonised parquets, from the workbooks in raw/hla_loh/
105
+ python hla_loh/harmonize.py
106
+
107
+ # the PCAWG sample sheet (the ICGC DCC portal is retired; this is the operations repo mirror)
108
+ gh api repos/ICGC-TCGA-PanCancer/pcawg-operations/contents/lists/sample_sheet/pcawg_sample_sheet.2016-10-18.tsv \
109
+ -H "Accept: application/vnd.github.raw+json" > raw/hla_loh/pcawg_sample_sheet.2016-10-18.tsv
110
+
111
+ # the workbooks
112
+ curl -O https://static-content.springer.com/esm/art%3A10.1038%2Fs41588-023-01367-1/MediaObjects/41588_2023_1367_MOESM4_ESM.xlsx
113
+ curl -O https://static-content.springer.com/esm/art%3A10.1038%2Fs41588-023-01367-1/MediaObjects/41588_2023_1367_MOESM5_ESM.xlsx
114
+ curl -O https://static-content.springer.com/esm/art%3A10.1038%2Fs41588-023-01367-1/MediaObjects/41588_2023_1367_MOESM6_ESM.xlsx
115
+ ```
116
+
117
+ `do_to_tcga_barcode.tsv` and `perallele_tcga_loh.parquet` were built on 2026-09-11 and are **not
118
+ scripted**. Per-allele lost = `loh_lilac` true at sample level **and**, at each heterozygous locus,
119
+ the allele whose tumour copy number is the minor one (`min(CN) < 0.5 <= max(CN)`); a homozygous
120
+ locus is never called lost, because the allele name is retained. The GDC step POSTs
121
+ `{"filters":{"op":"in","content":{"field":"case_id","value":[...]}},"fields":"case_id,submitter_id,project.project_id"}`
122
+ to `https://api.gdc.cancer.gov/cases` in batches, resolving the sample sheet's `submitter_donor_id`
123
+ (a TCGA **case UUID** for `*-US` projects, not a barcode) to the barcode.
124
+
125
+ ## Not harmonised, and why
126
+
127
+ Four sheets of Supplementary Data 1 (`Platinum-Yoruban trio`, `TRACERx hla-I typing`,
128
+ `TRACERx LOH of HLA`, and the three `GenDX validation` sheets) are the paper's **caller-validation**
129
+ tables — 54 to 600 rows each, against orthogonal typing. Nine sheets of Supplementary Data 2
130
+ (`metastatic GIE`, `primary GIE`, the per-gene and mutual-exclusivity breakdowns, `GIE primary vs
131
+ metastatic`) are the paper's **aggregate result** tables, every one of them a group-by over
132
+ `gie_sample.parquet`. Neither set is an input to anything here, and both are re-derivable or
133
+ re-fetchable from the source. `Supp. Table 2 - cancer type data` (58 rows) is a value-count over
134
+ `sample_metadata.parquet` and is dropped for the same reason.
hla_loh/do_to_tcga_barcode.tsv ADDED
@@ -0,0 +1,712 @@
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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+ icgc_donor_id case_uuid tcga_barcode gdc_project
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+ DO10172 58facedb-fcb8-4ecf-8338-2bfa4947acef TCGA-A6-6141 TCGA-COAD
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+ DO10486 ddf69ed9-4c53-48b8-9f01-495bd60c3425 TCGA-AA-3514 TCGA-COAD
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+ DO10631 f87c6dcc-1a9e-4df5-962c-1f946e2611f8 TCGA-AA-A01T TCGA-COAD
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+ DO10908 737b35e1-d668-4fce-9b6e-76946c7952b6 TCGA-27-2523 TCGA-GBM
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+ DO10960 9853c6bb-a42c-4698-abe5-f3c897ebc8f6 TCGA-06-0214 TCGA-GBM
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+ DO10975 74139255-a635-4c87-814d-3dd04ed630a8 TCGA-06-0190 TCGA-GBM
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+ DO11046 e17a6048-7a72-42c0-ad3f-97cbff02bc9f TCGA-19-2629 TCGA-GBM
9
+ DO11148 0078b0c4-68a9-483b-9aab-61156d263213 TCGA-14-1034 TCGA-GBM
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+ DO11202 b1c4579c-72fa-4974-830c-b6a81ace45f4 TCGA-14-2554 TCGA-GBM
11
+ DO11382 01a92062-967a-4900-8dc7-a5ecd3b3f8e2 TCGA-06-0171 TCGA-GBM
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+ DO11501 c184c3ca-7ad3-4202-b108-cb9fd5f5d947 TCGA-41-5651 TCGA-GBM
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+ DO11782 be3a7ef3-34ed-40e1-9d9c-187940596b26 TCGA-27-2528 TCGA-GBM
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+ DO11794 5c984433-33cf-42fc-b3ba-511efcdcab19 TCGA-06-0211 TCGA-GBM
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25
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26
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+ DO1384 20e8106b-1290-4735-abe4-7621e08e3dc8 TCGA-BH-A0E0 TCGA-BRCA
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+ DO13869 de7b7cac-f094-4d59-8651-e991e34ea093 TCGA-06-0210 TCGA-GBM
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+ DO1392 8785012f-f73e-4d68-87cf-1d804af32782 TCGA-A7-A13D TCGA-BRCA
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+ DO14171 f030e410-2585-461e-ad23-9fdf026ac06c TCGA-CR-7382 TCGA-HNSC
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562
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563
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565
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567
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568
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570
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571
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572
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573
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574
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576
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577
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578
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579
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580
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582
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584
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585
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586
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587
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588
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589
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590
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591
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592
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593
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594
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595
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596
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597
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598
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599
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600
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601
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602
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603
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604
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605
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606
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607
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608
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609
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610
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611
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612
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613
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614
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615
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616
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617
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618
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619
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620
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621
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622
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623
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624
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625
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626
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627
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628
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629
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630
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631
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632
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633
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634
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635
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636
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637
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638
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639
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640
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641
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642
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643
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644
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645
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646
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647
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648
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649
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650
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651
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652
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653
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654
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655
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656
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657
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658
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659
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660
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661
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662
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663
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664
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665
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666
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667
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668
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669
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670
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671
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672
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673
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674
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675
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676
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677
+ DO5808 9435447e-d65f-408b-863b-6576b1d652dd TCGA-AO-A03N TCGA-BRCA
678
+ DO6144 5d8ed961-a012-4b6b-bffb-9468d3953646 TCGA-EW-A1J5 TCGA-BRCA
679
+ DO6231 f45210d3-9e66-4f5e-bef1-5ee5547cc893 TCGA-EW-A1PC TCGA-BRCA
680
+ DO6249 941f75a1-fea4-4539-ba69-60bb11608f6d TCGA-C5-A1MI TCGA-CESC
681
+ DO6252 8a4da4f0-30f2-497e-8a7c-988fd6b813cb TCGA-C5-A1BN TCGA-CESC
682
+ DO6255 6b74c07a-9932-4a80-8296-9db08d0c6964 TCGA-EX-A1H5 TCGA-CESC
683
+ DO6314 e733f289-a0ec-478c-aad9-1378ad26ddfa TCGA-C5-A2LY TCGA-CESC
684
+ DO6332 dfd30f94-19e3-4838-95df-14a3763419af TCGA-DS-A0VL TCGA-CESC
685
+ DO639 a658becf-3934-49f2-9e74-291dca61d4bb TCGA-FD-A3N5 TCGA-BLCA
686
+ DO669 ca554128-da9d-4f37-9560-ca083509e01d TCGA-C4-A0F7 TCGA-BLCA
687
+ DO675 f48937ed-e294-41f0-8872-7010391167a9 TCGA-BT-A20P TCGA-BLCA
688
+ DO689 24f21425-b001-4986-aedf-5b4dd851c6ad TCGA-BT-A20V TCGA-BLCA
689
+ DO695 419deaac-ea45-4bdd-9fa0-b5cd8429b44f TCGA-BT-A20Q TCGA-BLCA
690
+ DO720 cce62116-3dcd-400d-ada0-9839ca02466e TCGA-FT-A3EE TCGA-BLCA
691
+ DO7801 df5ab6cc-6f68-4b6b-95e2-954c6b57ba9c TCGA-CA-6718 TCGA-COAD
692
+ DO7831 a06c09a8-0951-49f4-a765-5147bbcae109 TCGA-AA-3664 TCGA-COAD
693
+ DO7990 fafd4576-0709-49e1-926f-ad1ee09907ec TCGA-A6-2683 TCGA-COAD
694
+ DO804 096b4f32-10c1-4737-a0dd-cae04c54ee33 TCGA-BT-A20T TCGA-BLCA
695
+ DO8047 dca004bf-9c14-45a1-b186-ab3759f6c7fe TCGA-AA-3685 TCGA-COAD
696
+ DO822 f8c0a69c-df53-4a0a-a30e-4424c0ce3dd5 TCGA-DK-A1AG TCGA-BLCA
697
+ DO8264 268c01b3-d2ce-44c0-a2fe-ea846a1253cc TCGA-CA-6717 TCGA-COAD
698
+ DO828 7fee9d6f-c0ea-43c2-89c2-66d965a077a8 TCGA-CF-A27C TCGA-BLCA
699
+ DO8360 5e7074a6-5011-4dc5-950c-41365dbc6d09 TCGA-AD-6964 TCGA-COAD
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+ DO856 932bcdd4-5145-49e8-9a95-8a546396f572 TCGA-BT-A3PH TCGA-BLCA
701
+ DO8658 750770db-53cf-4021-a8c2-5e0ddb97fef5 TCGA-AA-3994 TCGA-COAD
702
+ DO8730 33e989b9-fac4-47d8-b429-269044ac772a TCGA-AA-3977 TCGA-COAD
703
+ DO8858 ca22edd4-eb7c-4677-84aa-6f0354459a19 TCGA-AA-A00R TCGA-COAD
704
+ DO8898 36eaf209-c332-4321-b6aa-51a3368ecd79 TCGA-AA-3555 TCGA-COAD
705
+ DO9034 5116e3b4-2bac-40f5-8046-b9c1783faaa5 TCGA-A6-3807 TCGA-COAD
706
+ DO9042 a512701f-88bb-4b05-8c94-fd285b9dd13e TCGA-AA-3666 TCGA-COAD
707
+ DO9074 ff1407c6-9174-4bae-a19b-d34ca71b898c TCGA-A6-2680 TCGA-COAD
708
+ DO9106 84e34cf2-b3bb-4a1a-8fb7-6c2c604c4008 TCGA-AA-A01S TCGA-COAD
709
+ DO9732 bfb07784-693b-4c25-874e-4ad6e04a5d46 TCGA-AA-3529 TCGA-COAD
710
+ DO9788 7d8eab0a-e6c8-4449-9ebf-50c41db94a06 TCGA-A6-2681 TCGA-COAD
711
+ DO9876 e457344d-76fb-46bf-b362-61a6e811d131 TCGA-AA-A00N TCGA-COAD
712
+ DO9940 df228c56-89ac-4122-93b3-6c0740c4ec09 TCGA-AA-3534 TCGA-COAD
hla_loh/escape_genes.parquet ADDED
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hla_loh/gie_allele.parquet ADDED
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hla_loh/gie_sample.parquet ADDED
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hla_loh/harmonize.py ADDED
@@ -0,0 +1,177 @@
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
1
+ #!/usr/bin/env python3
2
+ # 2026-09-16 -- harmonise the Martinez-Jimenez 2023 HLA-I immune-escape supplements into parquet.
3
+ #
4
+ # The three publisher workbooks are copyrighted Springer Nature supplementary material and are not
5
+ # ours to redistribute, so they live in the gitignored `raw/hla_loh/`. The *facts* they carry are
6
+ # not copyrightable, so this script writes neutral, long-format tables that do ship. Same call, and
7
+ # the same shape, as `icb_cohorts/`: harmonised output published, publisher workbook kept back.
8
+ #
9
+ # Re-run after refreshing raw/hla_loh/*.xlsx: python hla_loh/harmonize.py
10
+ #
11
+ # Source: Martinez-Jimenez F, Priestley P, Shale C, Baber J, Rozemuller E, Cuppen E. "Genetic immune
12
+ # escape landscape in primary and metastatic cancer." Nat Genet 55(5):820-831, 2023.
13
+ # doi:10.1038/s41588-023-01367-1 (verified against crossref, 2026-09-16).
14
+ # MOESM4 = Supplementary Tables 1-2, MOESM5 = Supplementary Data 1, MOESM6 = Supplementary Data 2.
15
+
16
+ from __future__ import annotations
17
+
18
+ import os
19
+
20
+ import openpyxl
21
+ import polars as pl
22
+
23
+ HERE = os.path.dirname(os.path.abspath(__file__))
24
+ RAW = os.path.join(os.path.dirname(HERE), "raw", "hla_loh")
25
+ SLOTS = ("A1", "A2", "B1", "B2", "C1", "C2")
26
+ CALLERS = ("LILAC", "xHLA", "polysolver")
27
+ TISSUE = {"n": "normal", "t": "tumour"}
28
+
29
+
30
+ def sheet(fn: str, name: str) -> tuple[list[str], list[tuple]]:
31
+ """Header and non-empty data rows of one worksheet, values only."""
32
+ wb = openpyxl.load_workbook(os.path.join(RAW, fn), read_only=True, data_only=True)
33
+ ws = wb[name]
34
+ it = ws.iter_rows(values_only=True)
35
+ hdr = [str(h) for h in next(it) if h is not None]
36
+ rows = [r for r in it if r and r[0] is not None]
37
+ wb.close()
38
+ return hdr, rows
39
+
40
+
41
+ def _b(x):
42
+ """openpyxl hands these back as the strings 'True'/'False'; 'None' and None are both missing."""
43
+ if x is None or x in ("None", ""):
44
+ return None
45
+ return str(x) == "True"
46
+
47
+
48
+ def _f(x):
49
+ if x is None or x in ("None", ""):
50
+ return None
51
+ try:
52
+ return float(x)
53
+ except (TypeError, ValueError):
54
+ return None
55
+
56
+
57
+ def _s(x):
58
+ return None if x is None or x == "None" else str(x)
59
+
60
+
61
+ def _allele(x):
62
+ """The workbooks write `A*02:01`; every other table in this compendium -- and `mhcmatch` --
63
+ writes `HLA-A*02:01`. Two vocabularies for one molecule is the recurring hazard here, so the
64
+ published column carries the repo's spelling and SOURCES.md records the workbook's."""
65
+ a = _s(x)
66
+ return None if a is None else (a if a.startswith("HLA-") else f"HLA-{a}")
67
+
68
+
69
+ def _df(rows):
70
+ """Scan every row before fixing a dtype. polars infers from the first 100 by default, and the
71
+ sparsest column here (`checkpoint_pathway_detail`) is null until row 250 -- inferred Null, then
72
+ a string arrives and the build fails. Cheap at this size, and it cannot silently mistype."""
73
+ return pl.DataFrame(rows, infer_schema_length=None)
74
+
75
+
76
+ def gie_tables():
77
+ """Supplementary Data 2, sheet `GIE per sample`: 6,319 samples, 4,439 Hartwig + 1,880 PCAWG.
78
+
79
+ Split into one row per (sample, allele slot) and one row per sample, because the sheet is two
80
+ tables side by side: a six-slot allele record and a per-sample escape record. `*_MUT` is a
81
+ mutation COUNT (0.0/0.5/1.0/2.0), not a flag -- casting it to bool loses the 2.0 and the 0.5.
82
+ """
83
+ hdr, rows = sheet("41588_2023_1367_MOESM6_ESM.xlsx", "GIE per sample")
84
+ i = {c: n for n, c in enumerate(hdr)}
85
+ g = lambda r, c: r[i[c]] if c in i else None # noqa: E731
86
+
87
+ allele = [
88
+ {"sample_id": _s(g(r, "sample_id_2")), "cohort": _s(g(r, "cohort")),
89
+ "cancer_type": _s(g(r, "cancer_type")), "cancer_type_code": _s(g(r, "cancer_type_code")),
90
+ "locus": f"HLA-{s[0]}", "slot": s, "allele": _allele(g(r, s)),
91
+ "copy_number": _f(g(r, f"{s}_CN")), "n_mutations": _f(g(r, f"{s}_MUT"))}
92
+ for r in rows for s in SLOTS]
93
+
94
+ flags = ("loh_lilac", "mut_hla_lilac", "systemic_app_pathway", "cd58_pathway",
95
+ "ifn_gamma_pathway", "checkpoint_pathway", "epigenetic_regulators_pathway",
96
+ "targeted_escape", "non_targeted_escape", "genetic_immune_escape")
97
+ # `copy_number_neutral_loh_hla` LOOKS like a flag beside the others and is not: it carries a
98
+ # comma-separated locus list (`HLA-A,HLA-C,HLA-B`). Read as bool it would have nulled every
99
+ # real value. The four LOH columns are all locus lists, kept verbatim as the source wrote them.
100
+ texts = ("copy_number_neutral_loh_hla", "loh_hfocal", "loh_focal", "loh_nonfocal",
101
+ "systemic_app_pathway_detail", "cd58_pathway_detail", "ifn_gamma_pathway_detail",
102
+ "checkpoint_pathway_detail", "epigenetic_regulators_pathway_detail")
103
+ sample = []
104
+ for r in rows:
105
+ row = {"sample_id": _s(g(r, "sample_id_2")), "cohort": _s(g(r, "cohort")),
106
+ "cancer_type": _s(g(r, "cancer_type")),
107
+ "cancer_type_code": _s(g(r, "cancer_type_code"))}
108
+ for loc in "ABC":
109
+ row[f"ploidy_minor_HLA-{loc}"] = _f(g(r, f"ploidy_minor_HLA-{loc}"))
110
+ row[f"ploidy_major_HLA-{loc}"] = _f(g(r, f"ploidy_major_HLA-{loc}"))
111
+ row[f"del_HLA-{loc}"] = _f(g(r, f"del_HLA-{loc}"))
112
+ row |= {c: _b(g(r, c)) for c in flags}
113
+ row |= {c: _s(g(r, c)) for c in texts}
114
+ sample.append(row)
115
+ return _df(allele), _df(sample)
116
+
117
+
118
+ def typing_callers():
119
+ """Supplementary Data 1, sheets `Hartwig` and `PCAWG`: three callers x normal/tumour x six
120
+ slots, one column each. Long, so a caller comparison is a group_by rather than 36 column names."""
121
+ out = []
122
+ for sh in ("Hartwig", "PCAWG"):
123
+ hdr, rows = sheet("41588_2023_1367_MOESM5_ESM.xlsx", sh)
124
+ i = {c: n for n, c in enumerate(hdr)}
125
+ for r in rows:
126
+ sid, mut = _s(r[i["sampleId"]]), _b(r[i["is_mutated"]])
127
+ for caller in CALLERS:
128
+ for t, tname in TISSUE.items():
129
+ for s in SLOTS:
130
+ col = f"{caller}_{t}_{s}"
131
+ if col not in i:
132
+ continue
133
+ out.append({"sample_id": sid, "cohort": sh, "is_mutated": mut,
134
+ "caller": caller, "tissue": tname, "slot": s,
135
+ "locus": f"HLA-{s[0]}", "allele": _allele(r[i[col]])})
136
+ return _df(out)
137
+
138
+
139
+ def sample_metadata():
140
+ """Supplementary Table 2: per-sample cohort, cancer type, treatment history and mutation load."""
141
+ hdr, rows = sheet("41588_2023_1367_MOESM4_ESM.xlsx", "Supp. Table 2 - sample metadata ")
142
+ num = {"ploidy", "sv_load", "smnv_load", "sbs_load", "dbs_load", "indel_load",
143
+ "sbs_load.clonal", "dbs_load.clonal", "indel_load.clonal",
144
+ "sbs_load.subclonal", "dbs_load.subclonal", "indel_load.subclonal"}
145
+ bul = {"blacklisted_gie", "is_selected", "has_lilac", "has_group_cohort",
146
+ "has_group_mut_signatures", "has_rna", "whole_genome_duplication", "had_radiotherapy",
147
+ "had_other_treatment", "had_chemotherapy", "had_hormone_therapy", "had_targeted_therapy",
148
+ "had_immunotherapy", "treatment_info_available"}
149
+ return _df([
150
+ {c: (_f(r[n]) if c in num else _b(r[n]) if c in bul else _s(r[n]))
151
+ for n, c in enumerate(hdr)} for r in rows])
152
+
153
+
154
+ def escape_genes():
155
+ """Supplementary Table 1: the immune-escape gene panel the paper's calls are made over."""
156
+ hdr, rows = sheet("41588_2023_1367_MOESM4_ESM.xlsx", "Supp. Table 1 - genes")
157
+ keys = ["gene", "pathway_specific", "pathway_general", "references", "mutation_types",
158
+ "selected"]
159
+ return _df([
160
+ {k: (_f(r[n]) if k == "selected" else _s(r[n])) for n, k in enumerate(keys)}
161
+ for r in rows])
162
+
163
+
164
+ def main() -> int:
165
+ out = {"gie_allele.parquet": None, "gie_sample.parquet": None}
166
+ out["gie_allele.parquet"], out["gie_sample.parquet"] = gie_tables()
167
+ out["hla_typing_callers.parquet"] = typing_callers()
168
+ out["sample_metadata.parquet"] = sample_metadata()
169
+ out["escape_genes.parquet"] = escape_genes()
170
+ for name, df in out.items():
171
+ df.write_parquet(os.path.join(HERE, name), compression="zstd")
172
+ print(f"{name:28s} {df.height:>7,} rows x {df.width:>2} cols")
173
+ return 0
174
+
175
+
176
+ if __name__ == "__main__":
177
+ raise SystemExit(main())
hla_loh/hla_typing_callers.parquet ADDED
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1
+ version https://git-lfs.github.com/spec/v1
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+ oid sha256:d4881daeaa39eb998db01ec15579581e65f68066746932f2a7f99328f053bfb5
3
+ size 148451
hla_loh/perallele_tcga_loh.parquet ADDED
@@ -0,0 +1,3 @@
 
 
 
 
1
+ version https://git-lfs.github.com/spec/v1
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+ oid sha256:4d507a72aabb96e5bdf30f9f481666554a2c4e7b93bf5b2162bb2dd440f6e95c
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+ size 7649
hla_loh/sample_metadata.parquet ADDED
@@ -0,0 +1,3 @@
 
 
 
 
1
+ version https://git-lfs.github.com/spec/v1
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+ oid sha256:c579034e9bc484225c8b68d0d97dfb72ac80b9a49526c6dd0248b59548d6ed6a
3
+ size 340067
homology/HOM_MouseHumanSequence.rpt.gz ADDED
@@ -0,0 +1,3 @@
 
 
 
 
1
+ version https://git-lfs.github.com/spec/v1
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+ oid sha256:9985b691a867025f24bda5af0f28a91b1aa2d28d3e06d4ee7d0db54fa217c12c
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+ size 3243831
homology/SOURCES.md ADDED
@@ -0,0 +1,50 @@
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
1
+ # `homology/` — sources
2
+
3
+ One file, and it exists to answer exactly one question: **given a human protein, which mouse gene
4
+ is its orthologue, spelled the way MGI spells it.**
5
+
6
+ ## `HOM_MouseHumanSequence.rpt.gz`
7
+
8
+ | field | value |
9
+ |---|---|
10
+ | origin | `https://www.informatics.jax.org/downloads/reports/HOM_MouseHumanSequence.rpt` |
11
+ | format | tab-separated, 13 columns, gzipped here (3,243,831 bytes; 15,110,808 raw) |
12
+ | retrieved | 2026-09-04; the served file was stamped `last-modified: 2026-08-31` |
13
+ | provenance | **derived / computed — not experimental.** Ortholog assignments are *inferred*, not measured |
14
+ | rows | 46,521 data rows over **21,928 homology classes** |
15
+ | organisms | 24,592 `human` rows, 21,929 `mouse, laboratory` rows |
16
+ | distinct symbols | 19,366 human, 21,926 mouse |
17
+ | human rows carrying a SWISS_PROT ID | 23,569 of 24,592 (95.8 %) |
18
+
19
+ ```zsh
20
+ curl -sL -o HOM_MouseHumanSequence.rpt \
21
+ https://www.informatics.jax.org/downloads/reports/HOM_MouseHumanSequence.rpt
22
+ gzip -9 HOM_MouseHumanSequence.rpt
23
+ ```
24
+
25
+ **MGI is a redistributor here, not the source of the inference.** `informatics.jax.org/homology.shtml`
26
+ states that MGI "uses homology class data from the Alliance of Genome Resources", and does **not**
27
+ state the method by which the Alliance computes an ortholog assignment. So the right claim to make
28
+ downstream is "MGI/Alliance homology class", never "orthologue" as though it were measured — and a
29
+ one-to-many class is a real answer, not a failure to resolve.
30
+
31
+ **Why this file and not `HOM_ProteinCoding.rpt` or a UniProt round-trip.** Two properties, and both
32
+ are load-bearing:
33
+
34
+ 1. **It carries `SWISS_PROT IDs`**, so a human UniProt accession reaches a mouse gene in *one* hop.
35
+ `HOM_ProteinCoding.rpt` has no accession column and would need the proteome FASTA first.
36
+ 2. **Its `Symbol` column is the MGI official symbol**, which is the vocabulary
37
+ `expression/toil_matrix_mmu.npz` is keyed on (26,737 genes).
38
+
39
+ That second one is the whole reason a curated table was fetched at all. The obvious cheap route --
40
+ read `GN=` out of `proteome/{human,mouse}.fasta.gz` and match UniProt's shared entry-name mnemonic
41
+ (`P53_HUMAN` / `P53_MOUSE`) -- resolves 14 of 21 accessions and returns UniProt's `GN=Tp53` for
42
+ human TP53. **`Tp53` is absent from `toil_matrix_mmu.npz`; the MGI official symbol `Trp53` is
43
+ present.** A lookup that returns nothing and a lookup that returns the wrong thing fail the same way
44
+ when the layer below drops silently, which is this package's recurring hazard. Through this file the
45
+ same 21 accessions resolve 16 directly, and 21 of 21 once a human-symbol fallback is added.
46
+
47
+ **Two accessions on `source_species = human` rows of `neoantigens/neoag_tested_mmu.tsv.gz` are
48
+ *mouse* accessions** -- `O54916` (Reps1) and `Q8VDL4` (Adpgk). They are not xenoantigens; the
49
+ deposit's species label is wrong on those two rows. A resolver that reports which route answered
50
+ sees this; one that silently falls through does not.
icb_cohorts/COHORTS.md ADDED
@@ -0,0 +1,77 @@
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
1
+ # The cohort table — what each one can carry, and what it is missing
2
+
3
+ Reconciled 2026-09-10 from the screen recorded in `2026-mhcmatch-benchmark/bench/results/cohort_availability.md`
4
+ and the provenance in `SOURCES.md` beside this file. The analysis plan that consumes this table is
5
+ `2026-mhcmatch-benchmark/REPLICATION_PLAN.md`.
6
+
7
+ **Three things are needed per patient**: somatic mutation calls at protein level (a tumour mutational
8
+ burden count is not enough), four-digit class-I HLA-A/B/C, and a clinical outcome. Tumour RNA is a
9
+ fourth that improves two of the nine EPIC terms and is not required — `expr_lvl` and `expr_norm` fall
10
+ back to the GTEx/TCGA tumour-type median with `expr_imputed` recording that they did.
11
+
12
+ **Typing is the scarce leg.** Of 200 distinct cohorts screened, 74 are blocked on typing alone.
13
+ `harmonized/typing.parquet` is the result of searching for it: 52,492 patients, 11 cohorts,
14
+ 360 distinct allotypes.
15
+
16
+ ## Tier 1 — mutations, true typing and outcome, all public
17
+
18
+ | cohort | patients | tumour | therapy | mutation calls | clone or CCF layer | typing source | tumour RNA | outcomes |
19
+ |---|--:|---|---|---|---|---|---|---|
20
+ | MSK-IMPACT checkpoint union | 2,655 | pan-cancer | PD-1, PD-L1, CTLA-4, combination | cBioPortal panel MAF | FACETS cancer-cell fraction | `msk_impact_50k_2026` germline, six alleles | none | overall survival 2,655; progression 1,699 |
21
+ | SU2C-MARK (Ravi 2023) | 309 | non-small-cell lung | anti-PD-(L)1 | exome MAF | PhyloGic cluster per mutation | Polysolver, Table S9 | subset | overall survival, progression-free survival, RECIST |
22
+ | Van Allen 2015 | 110 | melanoma | anti-CTLA-4 | exome MAF | none | cBioPortal sample attribute | none | overall survival, disease-free survival, best response |
23
+ | Anagnostou 2020 | 89 | non-small-cell lung | anti-PD-(L)1 | supplementary S3 | cancer-cell fraction per mutation | supplementary S11, with per-locus loss of heterozygosity | none | response, progression-free survival, overall survival |
24
+ | Hellmann 2018 | 75 | non-small-cell lung | nivolumab + ipilimumab | exome MAF | none | cBioPortal sample attribute | none | progression-free survival, best response, durable benefit |
25
+ | Riaz 2017 | 68 | melanoma | nivolumab | open exome `SRP095809` | none | Chowell Table S1 | yes | response, progression-free survival, overall survival |
26
+ | Snyder 2014 | 64 | melanoma | anti-CTLA-4 | exome MAF | none | cBioPortal sample attribute, class II too | none | overall survival, response, durable benefit |
27
+ | Hugo 2016 | 38 | melanoma | anti-PD-1 | exome MAF | none | cBioPortal sample attribute | yes | overall survival, response |
28
+ | Rizvi 2015 | 34 | non-small-cell lung | pembrolizumab | *Science* supplement | none | Chowell Table S1 | none | response, progression-free survival |
29
+ | Forde 2018 | 13 | non-small-cell lung, neoadjuvant | nivolumab | supplementary S6 | none | recoverable from supplementary S8 | none | pathological response |
30
+
31
+ The eight exome cohorts below SU2C-MARK pool to **491 patients** with deep calls and true typing.
32
+ That pool, not MSK, is the like-for-like replication of the SU2C-MARK arm.
33
+
34
+ ## Tier 2 — mutations and outcome public, typing absent, imputation available
35
+
36
+ | cohort | patients with calls | patients with outcome | tumour | randomised comparison | mutation calls | clone or CCF layer | tumour RNA | outcomes |
37
+ |---|--:|--:|---|---|---|---|---|---|
38
+ | Jammihal 2025 (CheckMate-214 + JAVELIN Renal 101) | 953 | 950 | renal cell | 465 checkpoint against 488 tyrosine-kinase inhibitor | 129,726 mutations with `Protein_Change` | ABSOLUTE `ccf_hat` and a clonality call | 733 | objective response, RECIST, progression-free survival, overall survival |
39
+ | Braun 2020 (CheckMate 009/010/025) | 454 | 1,006 | renal cell | 609 nivolumab against 397 everolimus | 68,030 mutations, 26,434 missense | none | 311 | objective response, benefit, progression-free survival, overall survival, immune-related progression-free survival |
40
+
41
+ Both are blocked on typing alone and both carry a **randomised control arm**, which no tier-1 cohort
42
+ does. Under imputed genotypes the patient-level ordering survives at Spearman rho +0.9433 at k = 20
43
+ and the top-k peptide sets do not (mean Jaccard 0.143) — `bench/results/hla_impute_check.md`. So
44
+ these two carry an outcome analysis and not a design claim.
45
+
46
+ ## Tier 3 — one email away
47
+
48
+ | ask | patients returned | who | what to request |
49
+ |---|--:|---|---|
50
+ | Jammihal and Braun typing | ~1,400 | David A. Braun (Yale), Sachet A. Shukla — senior authors of both | the POLYSOLVER class-I calls, keyed to `SUBJID` and `UUID`. Promotes both tier-2 rows to tier 1 |
51
+ | IMvigor210 | 348 | Genentech | class-I genotypes; exome MAF, RECIST and overall survival are public |
52
+ | IMmotion150 | 263 | David F. McDermott (BIDMC) | Polysolver calls; the apparently public typing is an MHCflurry argmax complete for 45 of 157 |
53
+ | Liu 2019 | 144 | Dana-Farber / Essen | the POLYSOLVER calls run for the paper's own loss-of-heterozygosity analysis, never published |
54
+
55
+ ## Tier 4 — typing computable from open reads, no email needed
56
+
57
+ | cohort | patients | reads | already public |
58
+ |---|--:|---|---|
59
+ | MORRISON-1 (CheckMate 038/064/067) | 427 | `PRJNA923698`, 160 open exome runs | variant allele frequency, logCPM, RECIST at `ParkerICI/MORRISON-1-public` |
60
+ | Zhao 2019 glioblastoma | 66 | `PRJNA482620`, 91 of 91 runs open | mutations, response, survival |
61
+ | Kim 2018 gastric | 61 | `PRJEB25780`, 95 exomes and 71 RNA | mutations, objective response, progression-free survival |
62
+ | Hong 2022 hepatocellular | 60 | `PRJEB34724`, 89 exomes and 40 RNA | mutations, objective response, progression-free survival, overall survival |
63
+
64
+ ## Tier 5 — typed and public, but not checkpoint blockade
65
+
66
+ Survival cohorts beside TCGA rather than beside SU2C-MARK: PCAWG / ICGC 1,028 donors closing all
67
+ three legs; TRACERx 421 with 89 typed and relapse-free survival; ICGC OV-AU ovarian 93 donors.
68
+ Hartwig 4,439 and PCAWG 1,839 carry LILAC typing in the open; Hartwig's mutations and outcome need
69
+ an application.
70
+
71
+ ## Ruled out, so nobody looks twice
72
+
73
+ Gide 2019 and Kang/Lee 2024 sequenced no exome. IMmotion151 publishes no per-mutation calls.
74
+ Chowell Cohort 2 has typing and overall survival but its identifiers cannot reach per-mutation
75
+ calls. Litchfield CPI1000+ is anonymised. AACR GENIE BPC releases no germline. Cloughesy 2019,
76
+ Khan 2020, Rose 2021, MOSCATO-01 and TOPACIO release too little. The 525-patient NeoPrecis
77
+ compendium publishes no typing — it stays a directional-rule arm only, `bench/results/cassette_ici.md`.
icb_cohorts/SOURCES.md ADDED
@@ -0,0 +1,45 @@
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
1
+ # ICB cohort assembly — sources
2
+
3
+ Staged 2026-09-10. **`raw/` is gitignored and not published** -- publisher supplementary workbooks,
4
+ copyrighted and not ours to redistribute. **`harmonized/` is published** (2026-09-11): tables we
5
+ derive from them -- our own six-allele genotypes over public typings -- carrying no publisher content.
6
+
7
+ The analysis being assembled for needs, per patient: somatic mutation calls at protein level,
8
+ four-digit class-I HLA-A/B/C, and a clinical outcome. **Typing is the scarce leg**, so it is
9
+ harmonised first, by `harmonize_typing.py`.
10
+
11
+ ## `raw/` — what each file is, and where it came from
12
+
13
+ | file | bytes | source | what it contributes |
14
+ |---|--:|---|---|
15
+ | `chowell2018_TableS1.xlsx` | 196,759 | Chowell D et al., *Science* 2018;359:582-587, doi:10.1126/science.aao4572, PMID 29301960, PMC6057471 — Supplementary Table S1 (`NIHMS980063-supplement-Supplemental_Table_S1.xlsx`) | **experimental** typing. `Cohort 1` 371 patients (Van Allen 100, Riaz 68, Rizvi_CUMC 67, Snyder 64, Hugo 38, Rizvi 2015 34) with `HLA Class I Alleles`, `OS_Months`, `OS_Event`, `Drug Class`, `MutCnt`; `Cohort 2` 1,166 MSK-IMPACT with the same; `HLA-II` 304; a `TCR CDR3s` sheet besides. 370 of 371 and 1,166 of 1,166 carry six four-digit alleles |
16
+ | `forde2018_nejm_S4-S10.xlsx`, `forde2018_nejm_supp1.pdf` | 3,077,810 / 1,662,342 | Forde PM et al., *N Engl J Med* 2018;378:1976-1986 — neoadjuvant nivolumab in resectable NSCLC (`NIHMS980037`) | **experimental**. `S6` 1,609 somatic alterations over 13 samples with hg19 coordinates, `Amino Acid (protein)`, CCDS transcript and consequence; `S8` 19,585 neoantigen rows carrying `HLA Allele` — six alleles per sample, so the genotype is recoverable for all 13 |
17
+ | `anagnostou2020_natcancer_S.xlsx` | 9,649,675 | Anagnostou V et al., *Nat Cancer* 2020;1:99-111, doi:10.1038/s43018-019-0008-8, PMC7514475 — Supplementary Tables | **experimental**. `S11` *HLA class I genomic variation*: 89 patients x six four-digit alleles plus per-locus LOH; `S3` per-mutation calls with cancer cell fraction |
18
+ | `braun2020_natmed_MOESM2.xlsx` | 125,569,016 | Braun DA et al., *Nat Med* 2020;26:909-918, doi:10.1038/s41591-020-0839-y — CheckMate 009/010/025 ccRCC | **experimental**. `S1_Clinical_and_Immune_Data` 1,006 patients x 121 columns (`Arm` nivolumab 609 / everolimus 397; `Cohort` CM-025 803, CM-010 168, CM-009 35; `ORR`, `Benefit`, `PFS`+`PFS_CNSR`, `OS`+`OS_CNSR`, `irPFS`); `S2_WES_Data` 68,030 somatic mutations (26,434 missense) over 454 tumours with `Protein_Change`; `S4A_RNA_Expression` on 311. **No HLA sheet — read every sheet, 2026-09-10** |
19
+ | `jammihal2025_rcc_S1-S6.xlsx` | 195,455,480 | Jammihal T et al., *Nat Cancer* 2025;6:372-384, doi:10.1038/s43018-024-00896-w, PMID 39789182 — CheckMate-214 + JAVELIN Renal 101 | **experimental**. `Supp. Table 1` 1,958 patients (arms, ORR, RECIST, PFS/OS with censoring); `Supp. Table 4` 129,726 mutations over 953 patients with `Protein_Change` and ABSOLUTE `ccf_hat`/clonality; `Supp. Table 3` expression on 733. **No HLA sheet** |
20
+ | `mjimenez2023_hla_suppl1.xlsx` | 882,368 | Martinez-Jimenez F et al., *Nat Genet* 2023;55:820-831, doi:10.1038/s41588-023-01367-1, PMID 37165135 — Supplementary Data 1 | **derived from experimental reads**: LILAC / xHLA / Polysolver class-I typing on normal *and* tumour separately. Hartwig 4,439 samples, PCAWG 1,839, plus TRACERx typing (491 rows) and LOH sheets |
21
+ | `neoprecis_nci_alleles.csv` | 12,622 | `cartercompbio/NeoPrecis`, `data/nci_alleles.csv` | **experimental** typing for the 75 NCI Surgery Branch GI patients, class I and class II |
22
+
23
+ Fetched with `curl -A <browser UA>` from `static-content.springer.com`; PMC serves a bot challenge
24
+ for the same files and must not be used. Chowell and Forde are NIHMS deposits and arrived through
25
+ the author's own browser download.
26
+
27
+ ## `harmonized/`
28
+
29
+ `typing.parquet` — one row per patient, columns `cohort`, `study`, `patient`, `A1 A2 B1 B2 C1 C2`,
30
+ every allele written `HLA-A*02:01`. A homozygous locus repeats its allele. Built 2026-09-10:
31
+
32
+ | source | patients | breakdown |
33
+ |---|--:|---|
34
+ | Chowell 2018 | 1,536 | `chowell_c1` 370, `chowell_c2` 1,166 |
35
+ | MSK-IMPACT 50K (cBioPortal API) | 44,227 | germline, full field, cut to two |
36
+ | Martinez-Jimenez 2023 | 6,278 | Hartwig 4,439, PCAWG 1,839 (LILAC **normal** calls, not tumour) |
37
+ | cBioPortal checkpoint-blockade studies | 287 | Van Allen 110, Hellmann 75, Snyder 64, Hugo 38 |
38
+ | Anagnostou 2020 | 89 | |
39
+ | NCI Surgery Branch | 75 | |
40
+ | **total** | **52,492** | 11 cohorts, 360 distinct allotypes |
41
+
42
+ Mutation and outcome tables are deliberately **not** built yet: the selection is open, and
43
+ harmonising 100k-row MAFs for cohorts that may be dropped is work spent before the decision.
44
+
45
+ The reconciled cohort table is `COHORTS.md` beside this file; the analysis plan is `2026-mhcmatch-benchmark/REPLICATION_PLAN.md`.
icb_cohorts/harmonize_typing.py ADDED
@@ -0,0 +1,199 @@
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
1
+ #!/usr/bin/env python3
2
+ # 2026-09-10 -- one class-I typing table across every cohort whose genotypes we can reach.
3
+ """Class-I typing is the scarce leg of every checkpoint-blockade cohort, so it is harmonised first.
4
+
5
+ Writes `harmonized/typing.parquet` with one row per (cohort, patient, locus, allele_1, allele_2)
6
+ flattened to six columns, every allele in `HLA-A*02:01` form, plus `harmonized/manifest.tsv`
7
+ naming what each source contributes. Mutation and outcome tables are deliberately NOT built yet:
8
+ the manuscript's cohort selection is open, and harmonising 100k-row MAFs for cohorts that may be
9
+ dropped is work spent before the decision.
10
+
11
+ uv run --with openpyxl --with polars python3 harmonize_typing.py
12
+ """
13
+ from __future__ import annotations
14
+
15
+ import collections
16
+ import csv
17
+ import json
18
+ import os
19
+ import re
20
+ import urllib.request
21
+
22
+ HERE = os.path.dirname(os.path.abspath(__file__))
23
+ RAW = os.path.join(HERE, "raw")
24
+ OUT = os.path.join(HERE, "harmonized")
25
+ LOCI = ("A", "B", "C")
26
+
27
+
28
+ def norm(a: str) -> str | None:
29
+ """Every spelling seen across these sources -> ``HLA-A*02:01``, or None if it is not one."""
30
+ if a is None:
31
+ return None
32
+ s = str(a).strip()
33
+ if not s or s.upper() in ("NA", "NAN", "-", "NONE"):
34
+ return None
35
+ s = s.replace("HLA-", "").replace("hla_", "").replace("HLA", "")
36
+ m = re.match(r"^([ABC])\*?(\d{2}):?(\d{2})", s) # A*02:01 / A0201 / A02:01
37
+ if m:
38
+ return f"HLA-{m.group(1)}*{m.group(2)}:{m.group(3)}"
39
+ m = re.match(r"^_?([abc])_(\d{2})_(\d{2})", s) # hla_a_02_01_01_01
40
+ if m:
41
+ return f"HLA-{m.group(1).upper()}*{m.group(2)}:{m.group(3)}"
42
+ return None
43
+
44
+
45
+ def six(alleles: list[str]) -> dict | None:
46
+ """Bucket a patient's calls by locus and require two per locus (a homozygote repeats)."""
47
+ by = collections.defaultdict(list)
48
+ for a in alleles:
49
+ n = norm(a)
50
+ if n:
51
+ by[n[4]].append(n)
52
+ row = {}
53
+ for L in LOCI:
54
+ v = sorted(set(by[L])) or []
55
+ if len(v) == 1:
56
+ v = v * 2
57
+ if len(v) != 2:
58
+ return None
59
+ row[f"{L}1"], row[f"{L}2"] = v
60
+ return row
61
+
62
+
63
+ def chowell() -> list[dict]:
64
+ import openpyxl
65
+ wb = openpyxl.load_workbook(f"{RAW}/chowell2018_TableS1.xlsx", read_only=True, data_only=True)
66
+ out = []
67
+ for sheet, tag in (("Cohort 1", "chowell_c1"), ("Cohort 2", "chowell_c2")):
68
+ ws = wb[sheet]
69
+ it = ws.iter_rows(values_only=True)
70
+ hdr = [h for h in next(it) if h]
71
+ ia, ir = hdr.index("HLA Class I Alleles"), (hdr.index("Reference") if "Reference" in hdr else None)
72
+ for r in it:
73
+ if not r or not r[0]:
74
+ continue
75
+ g = six(str(r[ia]).split(","))
76
+ if g:
77
+ out.append({"cohort": tag, "study": (r[ir] if ir is not None else tag),
78
+ "patient": str(r[0]), **g})
79
+ return out
80
+
81
+
82
+ def anagnostou() -> list[dict]:
83
+ import openpyxl
84
+ wb = openpyxl.load_workbook(f"{RAW}/anagnostou2020_natcancer_S.xlsx", read_only=True,
85
+ data_only=True)
86
+ ws = wb["S11"]
87
+ rows = [r for r in ws.iter_rows(values_only=True) if r and any(x is not None for x in r)]
88
+ hdr = [str(x) for x in rows[1]]
89
+ ip = hdr.index("Patient ID") if "Patient ID" in hdr else 0
90
+ cols = [i for i, h in enumerate(hdr) if h.startswith("HLA-") and "Allele" in h]
91
+ out = []
92
+ for r in rows[2:]:
93
+ g = six([r[i] for i in cols])
94
+ if g:
95
+ out.append({"cohort": "anagnostou2020", "study": "Anagnostou 2020 Nat Cancer",
96
+ "patient": str(r[ip]), **g})
97
+ return out
98
+
99
+
100
+ def hartwig_pcawg() -> list[dict]:
101
+ import openpyxl
102
+ wb = openpyxl.load_workbook(f"{RAW}/mjimenez2023_hla_suppl1.xlsx", read_only=True,
103
+ data_only=True)
104
+ out = []
105
+ for sheet, tag in (("Hartwig", "hartwig"), ("PCAWG", "pcawg")):
106
+ ws = wb[sheet]
107
+ it = ws.iter_rows(values_only=True)
108
+ hdr = [str(h) for h in next(it)]
109
+ cols = [i for i, h in enumerate(hdr) if h.startswith("LILAC_n_")] # normal, not tumour
110
+ for r in it:
111
+ if not r or not r[0]:
112
+ continue
113
+ g = six([r[i] for i in cols])
114
+ if g:
115
+ out.append({"cohort": tag, "study": "Martinez-Jimenez 2023 Nat Genet",
116
+ "patient": str(r[0]), **g})
117
+ return out
118
+
119
+
120
+ def nci() -> list[dict]:
121
+ out = []
122
+ with open(f"{RAW}/neoprecis_nci_alleles.csv") as f:
123
+ for row in csv.DictReader(f):
124
+ cols = [v for k, v in row.items() if k and re.match(r"^[ABC]_\d$", k)]
125
+ g = six(cols)
126
+ if g:
127
+ out.append({"cohort": "nci_gi", "study": "NCI Surgery Branch (via NeoPrecis)",
128
+ "patient": row.get("Patient") or row.get("sample") or "", **g})
129
+ return out
130
+
131
+
132
+ def cbioportal() -> list[dict]:
133
+ """The four checkpoint-blockade studies whose typing is a SAMPLE-level attribute."""
134
+ api = "https://www.cbioportal.org/api/studies/{}/clinical-data?clinicalDataType=SAMPLE&projection=SUMMARY&pageSize=100000"
135
+ studies = {"skcm_dfci_2015": "Van Allen 2015", "skcm_mskcc_2014": "Snyder 2014",
136
+ "mel_ucla_2016": "Hugo 2016", "nsclc_mskcc_2018": "Hellmann 2018"}
137
+ out = []
138
+ for sid, name in studies.items():
139
+ with urllib.request.urlopen(api.format(sid), timeout=120) as fh:
140
+ data = json.load(fh)
141
+ S = collections.defaultdict(dict)
142
+ for r in data:
143
+ S[r["sampleId"]][r["clinicalAttributeId"]] = r["value"]
144
+ for sample, d in S.items():
145
+ vals = []
146
+ for k, v in d.items():
147
+ if k.startswith("HLA_") and "LOH" not in k:
148
+ vals += str(v).split("/")
149
+ g = six(vals)
150
+ if g:
151
+ out.append({"cohort": sid, "study": name, "patient": sample, **g})
152
+ return out
153
+
154
+
155
+ def msk50k() -> list[dict]:
156
+ api = ("https://www.cbioportal.org/api/studies/msk_impact_50k_2026/clinical-data"
157
+ "?clinicalDataType=PATIENT&attributeId={}&projection=SUMMARY&pageSize=200000")
158
+ H = collections.defaultdict(list)
159
+ for a in ("HLA-A1", "HLA-A2", "HLA-B1", "HLA-B2", "HLA-C1", "HLA-C2"):
160
+ with urllib.request.urlopen(api.format(a), timeout=300) as fh:
161
+ for r in json.load(fh):
162
+ H[r["patientId"]].append(r["value"])
163
+ out = []
164
+ for p, vals in H.items():
165
+ g = six(vals)
166
+ if g:
167
+ out.append({"cohort": "msk_impact_50k", "study": "MSK-IMPACT 50K", "patient": p, **g})
168
+ return out
169
+
170
+
171
+ SOURCES = {"chowell2018": chowell, "anagnostou2020": anagnostou,
172
+ "martinez_jimenez2023": hartwig_pcawg, "nci_neoprecis": nci,
173
+ "cbioportal_icb": cbioportal, "msk_impact_50k": msk50k}
174
+
175
+
176
+ def main() -> None:
177
+ import polars as pl
178
+ os.makedirs(OUT, exist_ok=True)
179
+ rows, man = [], []
180
+ for name, fn in SOURCES.items():
181
+ try:
182
+ got = fn()
183
+ except Exception as e: # noqa: BLE001
184
+ print(f"# {name}: FAILED {e}")
185
+ man.append({"source": name, "patients": 0, "note": f"failed: {e}"})
186
+ continue
187
+ rows += got
188
+ by = collections.Counter(g["cohort"] for g in got)
189
+ print(f"# {name}: {len(got):,} patients {dict(by)}")
190
+ man.append({"source": name, "patients": len(got), "note": json.dumps(dict(by))})
191
+ df = pl.DataFrame(rows)
192
+ df.write_parquet(f"{OUT}/typing.parquet")
193
+ pl.DataFrame(man).write_csv(f"{OUT}/manifest.tsv", separator="\t")
194
+ print(f"\n# typing.parquet: {df.height:,} patients over {df['cohort'].n_unique()} cohorts, "
195
+ f"{len(set(df['A1']) | set(df['A2']) | set(df['B1']) | set(df['B2']) | set(df['C1']) | set(df['C2']))} distinct allotypes")
196
+
197
+
198
+ if __name__ == "__main__":
199
+ main()
icb_cohorts/harmonized/manifest.tsv ADDED
@@ -0,0 +1,7 @@
 
 
 
 
 
 
 
 
1
+ source patients note
2
+ chowell2018 1536 "{""chowell_c1"": 370, ""chowell_c2"": 1166}"
3
+ anagnostou2020 89 "{""anagnostou2020"": 89}"
4
+ martinez_jimenez2023 6278 "{""hartwig"": 4439, ""pcawg"": 1839}"
5
+ nci_neoprecis 75 "{""nci_gi"": 75}"
6
+ cbioportal_icb 287 "{""skcm_dfci_2015"": 110, ""skcm_mskcc_2014"": 64, ""mel_ucla_2016"": 38, ""nsclc_mskcc_2018"": 75}"
7
+ msk_impact_50k 44227 "{""msk_impact_50k"": 44227}"
icb_cohorts/harmonized/typing.parquet ADDED
@@ -0,0 +1,3 @@
 
 
 
 
1
+ version https://git-lfs.github.com/spec/v1
2
+ oid sha256:0b8a7494b2d3c73142a7f5cc707bb093732b9c6e23473b0d425842d0daa5669c
3
+ size 289060
immunogenicity/SOURCES.md ADDED
@@ -0,0 +1,726 @@
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
1
+ # `immunogenicity/` — sources, provenance, filtering and a complete audit
2
+
3
+ General immunogenicity corpora: **is this presented peptide seen by a T cell?** Neoantigen datasets
4
+ moved to [`../neoantigens/`](../neoantigens/SOURCES.md) — this directory no longer holds NCI,
5
+ `neoag_*` or any tumour-specific screen.
6
+
7
+ Every citation resolved against **PubMed**; PMIDs and DOIs given. Audit regenerated by
8
+ `bench/neoag/audit_pmhc_data.py` in `2026-mhcmatch-benchmark`; corpora rebuilt by
9
+ `bench/neoag/corpus_chowell.py` + `bench/neoag/deposit_corpora.py`.
10
+
11
+ Written 2026-08-17.
12
+
13
+ ---
14
+
15
+ ## Complete audit
16
+
17
+ Measured from the files, not copied from any prior description.
18
+
19
+ | file | rows | distinct peptides | pos | neg | prevalence | HLA alleles | length |
20
+ |---|--:|--:|--:|--:|--:|--:|:--:|
21
+ | `chowell_rebuilt.tsv.gz` | 511,301 | 315,191 | 19,866 | 491,435 | 0.0389 | 210 | 8–11 |
22
+ | `chowell_rebuilt_hla_matched.tsv.gz` | 115,592 | 99,763 | 19,866 | 95,726 | 0.1719 | 183 | 8–11 |
23
+ | `kesmir_rebuilt.tsv.gz` | 65,737 | 36,155 | 22,613 | 43,124 | 0.3440 | 196 | 8–11 |
24
+ | `kesmir_rebuilt_hla_matched.tsv.gz` | 59,941 | 35,202 | 22,613 | 37,328 | 0.3773 | 188 | 8–11 |
25
+ | `iedb_labeled.tsv.gz` | 790,868 | 516,092 | 34,362 | 756,506 | 0.0435 | 250 | 2–50 |
26
+ | `hla_pop_freqs.tsv.gz` | 7,171 | — | — | — | — | 4,234 | — |
27
+
28
+ No donor/patient identifier exists in any file here — these are peptide-level corpora aggregated
29
+ over studies, not per-donor cohorts. (Donor counts are meaningful in `../neoantigens/` and are
30
+ reported there.)
31
+
32
+ ---
33
+
34
+ ## `chowell_rebuilt.tsv.gz` — immunogenic vs presented **self**
35
+
36
+ Chowell's construction, rebuilt from the current IEDB dump rather than read off the 2015
37
+ spreadsheet, which is copyrighted supplementary material and frozen at 2015.
38
+
39
+ > Chowell D, Krishna S, Becker PD, Cocita C, Shu J, Tan X, Greenberg PD, Klavinskis LS,
40
+ > Blattman JN, Anderson KS. **TCR contact residue hydrophobicity is a hallmark of immunogenic CD8+
41
+ > T cell epitopes.** *Proc Natl Acad Sci USA* 2015;112(14):E1754–62.
42
+ > PMID [25831525](https://pubmed.ncbi.nlm.nih.gov/25831525/) ·
43
+ > doi:[10.1073/pnas.1500973112](https://doi.org/10.1073/pnas.1500973112)
44
+
45
+ ### Filtering procedure, in order
46
+
47
+ 1. **Source** — `iedb_labeled.tsv.gz` (see below), itself an IEDB export.
48
+ 2. **Class** — `mhc_class == "MHCI"`.
49
+ 3. **Host** — `host_species ∈ {human, mouse}`; kept as a column and **never pooled**. The two hosts
50
+ have different MHC and different thymic repertoires, so a fit across them without the split is
51
+ fitting a mixture.
52
+ 4. **Peptide** — uppercased, whitespace-stripped; length **8–11**; canonical 20 amino acids only.
53
+ 5. **Exclusions** — `SLLMWITQV`, `KLGGALQAK` (NY-ESO-1 / MAGE-A3 epitopes that appear on both sides
54
+ of the literature with contradictory annotation).
55
+ 6. **Positives** — any peptide with a positive T-cell assay (`dataset_origin == "iedb_tcell"`).
56
+ 7. **Negatives** — eluted ligands whose `source_species` equals the **host** (self), plus the HLA
57
+ Ligand Atlas thymus immunopeptidome for the human arm.
58
+ 8. **Label is a property of the (peptide, host), not the row.** 1,075 peptides are both eluted and
59
+ positively assayed. Elution is evidence of *presentation*; it is never evidence against a T-cell
60
+ response. Assigning per row makes the same sequence positive under one allele and negative under
61
+ another — it sits on both sides of the fit. Computing the positive set once per (peptide, host)
62
+ and applying it everywhere raised self-sourced positives by **57%**.
63
+ 9. **Aggregation** — one row per `(peptide, allele, host_species)`, label = max.
64
+
65
+ ### Not controlled, deliberately
66
+
67
+ **Allele and length.** Immunogenic and eluted sets genuinely differ in both: some HLAs are
68
+ protective and some are not, and immunogenic peptides have a real length distribution that differs
69
+ from bulk ligands. Matching those away removes effects, not confounds. `allele` and `length` ship
70
+ as columns so a stratified analysis is a filter, not a rebuild — and the HLA-matched variant below
71
+ exists for when allele composition *is* the thing to remove.
72
+
73
+ ---
74
+
75
+ ## `kesmir_rebuilt.tsv.gz` — immunogenic vs presented **non-self**
76
+
77
+ The Calis/Kesmir contrast. Same shape, same rules, one difference: negatives are eluted ligands
78
+ from a source organism that is **not** the host.
79
+
80
+ > Calis JJA, Maybeno M, Greenbaum JA, Weiskopf D, De Silva AD, Sette A, Keşmir C, Peters B.
81
+ > **Properties of MHC class I presented peptides that enhance immunogenicity.**
82
+ > *PLoS Comput Biol* 2013;9(10):e1003266.
83
+ > PMID [24204222](https://pubmed.ncbi.nlm.nih.gov/24204222/) ·
84
+ > doi:[10.1371/journal.pcbi.1003266](https://doi.org/10.1371/journal.pcbi.1003266)
85
+
86
+ **This arm requires `../ligandome/viral_foreign_iedb.tsv.gz`, and is wrong without it.** Every
87
+ eluted-ligand negative in the IEDB table itself is self-sourced — 449,449 human-self and 41,986
88
+ mouse-self against 149 + 63 cross-species, and zero from any other organism. Filtering
89
+ `source_species != "human"` on a **mouse** host therefore selects mouse *self* ligands and rebuilds
90
+ Chowell under another name. The foreign ligandome supplies 41,294 genuine human non-self negatives.
91
+
92
+ Kesmir removes the self/non-self axis that Chowell confounds with immunogenicity, and is the harder
93
+ problem: prevalence 0.344 against Chowell's 0.039, and every negative is a foreign peptide that was
94
+ presented and still not attacked.
95
+
96
+ ---
97
+
98
+ ## The HLA-matched variants
99
+
100
+ `*_rebuilt_hla_matched.tsv.gz`, built by `deposit_corpora.py --ratio 5 --seed 20260817`.
101
+
102
+ There are 5–25× more negatives than positives and the two sides do not share an allele
103
+ distribution, so a classifier can separate them on **allele composition alone**. For each allele,
104
+ negatives are drawn without replacement to **5 per positive of that same allele**:
105
+
106
+ * alleles with no positives contribute **no** negatives — keeping them would reintroduce exactly the
107
+ imbalance being removed;
108
+ * alleles with too few negatives contribute all they have, and the shortfall stays visible in the
109
+ per-allele counts rather than being topped up from another allele;
110
+ * the draw is seeded (`20260817`), so the file is reproducible rather than one sample.
111
+
112
+ Effect: Chowell 491,435 → 95,726 negatives, 210 → 183 alleles, prevalence 0.039 → 0.172. Kesmir
113
+ 43,124 → 37,328, 196 → 189 alleles.
114
+
115
+ Use the matched file when the question is about peptide sequence; use the full file when allele
116
+ composition is part of the signal you want. Neither is "the" corpus.
117
+
118
+ ---
119
+
120
+ ## Two residue-level artefacts, measured
121
+
122
+ Both are properties of how the data was generated. A model will fit either as if it were biology.
123
+
124
+ | corpus | Cys % positives | Cys % negatives | ratio | C-term R/K % pos | C-term R/K % neg |
125
+ |---|--:|--:|--:|--:|--:|
126
+ | `chowell_rebuilt` | 10.69 | **1.55** | **6.9×** | 9.21 | 9.50 |
127
+ | `kesmir_rebuilt` | 10.94 | 14.50 | 0.8× | 9.91 | **20.33** |
128
+ | `iedb_labeled` | 14.56 | **2.52** | **5.8×** | 10.90 | 13.98 |
129
+
130
+ ### Cysteine — an MS detection artefact, and it hits Chowell hardest
131
+
132
+ Cysteine is systematically **under-detected in mass-spectrometry immunopeptidomics** unless samples
133
+ are alkylated. Where positives are T-cell-assayed (synthetic peptides, no MS step) and negatives are
134
+ MS-eluted, the corpus carries a Cys gradient with no biological content. Chowell shows **6.9×**;
135
+ finer-grained, the thymus MS subset runs at 0.17% against 11.59% in the assayed positives.
136
+
137
+ Fitted freely on the Chowell corpus, Cys took the **single largest coefficient** in a position-role
138
+ naive Bayes (+1.84 anchor / +2.05 TCR-facing). Masking it costs grouped-CV AUROC 0.712 → 0.690 and
139
+ *improves* transfer to neoantigen cohorts. The shipped `mhcmatch.posbayes` tables zero it.
140
+
141
+ **`kesmir_rebuilt` does not have this problem** (0.8×) — its negatives are IEDB-curated foreign
142
+ ligands rather than a single MS pipeline. That makes it the safer corpus for fitting residue-level
143
+ terms, and is a reason to run both.
144
+
145
+ ### C-terminal Arg/Lys — genuine anchor preference, or tryptic carry-over
146
+
147
+ A basic C-terminus is the real anchor preference of **HLA-A\*03, A\*11, A\*31, A\*68 and B\*27** —
148
+ and Arg/Lys are exactly the residues **trypsin cleaves after**. An elevated rate can be either, and
149
+ the two are **not separable from sequence alone**.
150
+
151
+ `kesmir_rebuilt` negatives run at **20.33%** against 9.91% in positives — a 2× gradient in the
152
+ corpus that is otherwise the cleaner of the two. Before attributing anything to a C-terminal basic
153
+ residue on this corpus, stratify by allele: if the effect survives within A\*03/A\*11/B\*27 and
154
+ within non-basic-preferring alleles separately, it is anchor preference; if it only appears pooled,
155
+ it is composition or contamination.
156
+
157
+ ---
158
+
159
+ ## `iedb_labeled.tsv.gz`
160
+
161
+ The IEDB export both corpora are built from. **This is also the `ipred` training set**, so any
162
+ `ipred` figure quoted on it is in-sample.
163
+
164
+ 790,868 rows · 516,092 distinct peptides · 34,362 immunogenic · 250 alleles · lengths 2–50
165
+ (unfiltered — the 8–11 restriction is applied downstream).
166
+
167
+ > Vita R, Mahajan S, Overton JA, Dhanda SK, Martini S, Cantrell JR, Wheeler DK, Sette A, Peters B.
168
+ > **The Immune Epitope Database (IEDB): 2018 update.** *Nucleic Acids Res* 2019;47(D1):D339–D343.
169
+ > PMID [30357391](https://pubmed.ncbi.nlm.nih.gov/30357391/) ·
170
+ > doi:[10.1093/nar/gky1006](https://doi.org/10.1093/nar/gky1006)
171
+
172
+ Provenance: **experimental** — curated T-cell assay and MHC-ligand records. Raw export kept at
173
+ `raw/immunogenicity/iedb_tcell_assays_29032026.tsv.gz`.
174
+
175
+ Thymus negatives come from the HLA Ligand Atlas (`thymus/thymus_immunopeptidome.tsv.gz`); its
176
+ `mhc_a` is null by design — the deposit is not allele-deconvolved.
177
+
178
+ ---
179
+
180
+ ## `hla_pop_freqs.tsv.gz`
181
+
182
+ Class-I allele frequencies (A/B/C) for EUR/ASN/AFR populations, 7,171 rows over 4,234 distinct
183
+ alleles. Source: Allele Frequency Net Database (AFND), <http://www.allelefrequencies.net>.
184
+ Provenance: **experimental** (population genotyping), aggregated.
185
+
186
+ ---
187
+
188
+ ## Regenerate
189
+
190
+ # in ~/vcs/projects/2026-mhcmatch-benchmark
191
+ python bench/neoag/corpus_chowell.py # plain build (Chowell arm)
192
+ python bench/neoag/corpus_chowell.py --with-viral # + foreign ligandome (Kesmir arm)
193
+ python bench/neoag/deposit_corpora.py --ratio 5 --seed 20260817
194
+ python bench/neoag/audit_pmhc_data.py
195
+
196
+ `corpus_chowell.parquet` must keep its exact row counts (human 464,310 / 14,712 pos; mouse 47,203 /
197
+ 5,154) — the shipped `mhcmatch.posbayes` tables are fitted on it, and folding extra sources into it
198
+ in place breaks that provenance silently. That is why `--with-viral` writes a separate file.
199
+
200
+ ## `chowell_iedb_full.tsv.gz` / `chowell_iedb_full_matched.tsv.gz` (2026-08-18)
201
+
202
+ Rebuilt from the **full** IEDB export (build 2026-08-11) rather than `iedb_labeled.tsv.gz`. That
203
+ harmonised table kept only *positive* T-cell assays, so its `assay_type` is perfectly collinear with
204
+ its label and it cannot supply a measured negative at all; the full export carries 361,962 negative
205
+ T-cell assay rows, for mouse as well as human.
206
+
207
+ `chowell_rebuilt.tsv.gz` and `kesmir_rebuilt.tsv.gz` are **unchanged and still current** — the
208
+ shipped `mhcmatch.complement` and `posbayes` tables are fitted on them, they are named in the
209
+ library's `REFERENCE_FILES`, and every AUROC in `bench/results/complementarity.md` belongs to them.
210
+ These two files are additional, not replacements.
211
+
212
+ ### The rules, in the order applied
213
+
214
+ 1. MHC class I.
215
+ 2. Host ∈ {human, mouse}; a column, never pooled.
216
+ 3. Length 8–11, canonical 20 amino acids.
217
+ 4. Exclude `SLLMWITQV`, `KLGGALQAK`.
218
+ 5. **Immunogenic** — the peptide has ≥1 positive T-cell assay in that host (`Positive`,
219
+ `Positive-Low`, `-High`, `-Intermediate`).
220
+ 6. **Non-immunogenic** — an eluted **self** ligand appearing in no positive T-cell assay in any
221
+ host. 2,086 rows are eluted but positive elsewhere and are excluded from the negatives rather
222
+ than counted as such.
223
+ 7. The label is a property of `(peptide, host)`, never of an assay row.
224
+ 8. The restriction is resolved, and **no row is dropped for lacking one**: 1,677,439 of 3,278,355
225
+ surviving class-I rows name only `HLA class I`. netMHCpan-4.2 picks the best `%Rank_EL` among
226
+ whatever the recorded string still permits — an allele group or serotype restricts to that
227
+ group, an H-2 haplotype to the molecules it carries, a bare `HLA class I` to a 30-allele
228
+ population panel built per locus. A **known** non-human/non-mouse MHC on a human or mouse host
229
+ (2,812 rows) is a curation error, not a missing value, and is excluded rather than imputed.
230
+ 9. One row per `(peptide, allele group, host)`, label = max. The key is the allele **group**
231
+ (`HLA-A*02`), which is what lets serotype-only records join.
232
+
233
+ ### Counts
234
+
235
+ | file | rows | positive | negative | peptides | allele groups |
236
+ |---|--:|--:|--:|--:|--:|
237
+ | `chowell_iedb_full` (human) | 789,047 | 24,293 | 764,754 | 579,322 | 75 |
238
+ | `chowell_iedb_full` (mouse) | 65,472 | 7,187 | 58,285 | 49,253 | 43 |
239
+ | `chowell_iedb_full_matched` (human) | 20,360 | 10,180 | 10,180 | 17,181 | 18 |
240
+ | `chowell_iedb_full_matched` (mouse) | 9,132 | 4,566 | 4,566 | 7,969 | 25 |
241
+
242
+ Columns add `allele_2f`, `allele_group`, `allele_source` ∈ {`reported`, `imputed_group`,
243
+ `imputed_panel`} and `allele_rank_el`, so an analysis can exclude imputed restrictions and say how
244
+ much sample it lost. Human: 375,462 reported against 413,585 imputed; mouse 51,003 against 14,469.
245
+
246
+ The matched arm resamples both classes to one profile at 1:1 so the allele group carries no signal
247
+ about the label — human to population frequency from `hla_pop_freqs.tsv.gz`, mouse to the positives'
248
+ own profile, there being no H-2 frequency table. It is **small**: 52 of 75 human allele groups are
249
+ too scarce to fill their population weight and are dropped, which is the cost of the matching, not a
250
+ bug.
251
+
252
+ Regenerate, in `~/vcs/projects/2026-mhcmatch-benchmark`:
253
+
254
+ python immunogenicity/fetch.py # stage the export from ~/hf/pmhc_data/dump
255
+ bench/neoag/corpus_iedb.py --workers 12 # ~55 min, 14.7M netMHCpan predictions
256
+ bench/neoag/deposit_arms.py
257
+
258
+ Arm counts, the filter cascade and the selection tree are `bench/results/corpus_arms.md`.
259
+
260
+ **Caveat carried forward.** These negatives are still *inferred* — an eluted ligand nobody has
261
+ tested is assumed non-immunogenic. 38,322 rows in the rebuild have a measured negative T-cell assay
262
+ instead, held under evidence `tcell_negative` and deliberately **not** merged into the arm: swapping
263
+ the negative definition changes what every AUROC recorded on these corpora means.
264
+
265
+ ### `immunogenicity_legacy_arms.tsv.gz` (2026-08-19)
266
+
267
+ The published deposits and the legacy Kešmir arms in one table, keyed `corpus` ∈ {`chowell_vanilla`,
268
+ `kesmir_S1`, `kesmir_S2`, `kesmir_S3`}, so a held-out evaluation can be reproduced without the
269
+ copyrighted spreadsheets. 15,991 rows.
270
+
271
+ **Species is keyed on the restricting allele, not on the deposit's own `Species` column.** The two
272
+ disagree on 777 of the 2,508 Calis rows (31%), which carry `Species = Mus` with an HLA restriction —
273
+ HLA-transgenic mice, murine host and human restricting molecule. Both readings describe the
274
+ experiment; they build different corpora. Keyed on the host, the Calis human arm is 1,113
275
+ immunogenic against **one** non-immunogenic; keyed on the allele it is **1,619 against 272** and the
276
+ mouse arm is H-2 throughout. The reported host is retained as `host_reported`.
277
+
278
+ Regenerate with `bench/neoag/corpus_legacy.py` then `deposit_arms.py`.
279
+
280
+ ### What the recognition model is fitted on
281
+
282
+ `chowell_iedb_full` is the training arm for `mhcmatch.recognition` (0.15.0). Two choices were made
283
+ by measurement and are recorded in `bench/results/recognition_model.md` and
284
+ `feature_importance.md`:
285
+
286
+ - **The unmatched arm ships.** Resampling negatives to match population HLA usage costs 0.019
287
+ (human) and 0.058 (mouse) AUROC on held-out published deposits.
288
+ - **ESM2 and the physicochemical features are both kept.** ESM2 adds +0.022/+0.042 over physchem
289
+ alone; physchem adds +0.009/+0.019 over ESM2 alone. Permutation importance on held-out data ranks
290
+ the two ESM face-pools first (0.19–0.27), then `kf_tcr`; the whole-peptide Kidera aggregate is
291
+ worth 0.0165/0.0012 and is not shipped, being exactly the sum of the two role columns.
292
+
293
+ ## `staged/` — the IEDB full export, as staged (2026-08-19)
294
+
295
+ The **root of the provenance chain**. Every rebuilt corpus in this directory
296
+ (`chowell_iedb_full*`, `immunogenicity_legacy_arms`) is derived from these two tables, and until
297
+ now they lived only in a gitignored working directory — one bad checkout from being unreproducible.
298
+ Deposited so the chain closes without re-downloading 450 MB from IEDB.
299
+
300
+ | file | provenance | rows |
301
+ |---|---|--:|
302
+ | `staged/iedb_tcell.parquet` | **experimental** — IEDB `tcell_full_v3_tsv.zip`, build 2026-08-11, downloaded 2026-08-18 | 570,434 linear-peptide rows, 233,636 distinct peptides |
303
+ | `staged/iedb_ligand.parquet` | **experimental** — IEDB `mhc_ligand_full_tsv.zip`, same build | 5,773,495 linear-peptide rows, 1,704,996 distinct peptides |
304
+ | `staged/fetch.py` | the script that produced both | — |
305
+ | `staged/FETCH.md` | its record: zip mtimes, export row counts, alignment failures, class and qualitative-measurement breakdowns | — |
306
+
307
+ Ten columns are kept from each export; nothing is filtered beyond `Linear peptide` and a non-empty
308
+ name, because length, alphabet, class and host filters are corpus-construction rules and belong
309
+ where they can be stated as rules. **0 rows failed to align** in either table — IEDB ships two
310
+ header lines and embedded newlines in comment fields, so `fetch.py` reads with `csv.reader` rather
311
+ than splitting on tabs, and reports the misalignment count rather than assuming it is zero.
312
+
313
+ The T-cell export is what makes a *measured* negative possible: 361,962 `Negative` rows across all
314
+ hosts, where the older harmonised `iedb_labeled.tsv.gz` keeps only positive T-cell assays and its
315
+ `assay_type` is therefore perfectly collinear with its label.
316
+
317
+ Regenerate (needs `~/hf/pmhc_data/dump`, which is gitignored — the raw zips are not deposited):
318
+
319
+ ```zsh
320
+ python immunogenicity/staged/fetch.py
321
+ ```
322
+
323
+ ## `corpus_iedb_mhc2.parquet` — the MHC class-II arm (2026-08-19)
324
+
325
+ **Derived/computed**, from `staged/iedb_tcell.parquet` and `staged/iedb_ligand.parquet` — the same
326
+ IEDB build 2026-08-11 export the class-I corpus is built from, by the same rules, so the two arms are
327
+ comparable by construction rather than by assertion. This is what the shipped class-II
328
+ complementarity model (`mhcmatch.complement`, `cls="mhc2"`, v0.16.0) is fitted on.
329
+
330
+ | field | value |
331
+ |---|--:|
332
+ | rows (peptide, allele group, host) | 1,096,034 |
333
+ | distinct peptides | 643,744 |
334
+ | immunogenic peptides | 77,943 |
335
+ | human rows | 1,036,041 |
336
+ | human positives | 65,486 |
337
+ | human distinct peptides | 603,781 |
338
+ | human allele groups | 121 |
339
+ | mouse rows | 59,993 |
340
+ | mouse positives | 12,457 |
341
+ | mouse distinct peptides | 50,258 |
342
+ | mouse allele groups | 40 |
343
+ | peptide length | 11–25, median 15 |
344
+
345
+ Three rules differ from the class-I arm and nothing else does:
346
+
347
+ - **Length** is 11–25 rather than 8–11.
348
+ - **Restriction is parsed, never imputed.** Class-II restriction is written a dozen ways across IEDB
349
+ (`HLA-DRB1*15:01`, `HLA-DPA1*01:03/DPB1*04:01`, `HLA-DR15`, serotypes, `H2-IAb`), so the corpus
350
+ carries a parser and keeps whatever it cannot resolve as an explicit `unresolved` group rather than
351
+ guessing an allele. It parses **1,003,554 / 1,974,413 rows (50.8 %)**; the remainder are retained,
352
+ not dropped, because the complementarity fit reads only peptide and label. There is therefore **one
353
+ class-II arm where class I has two** — no HLA-matched variant is built, since matching needs a
354
+ resolved allele on both sides.
355
+ - **Rule 8** (pathogen epitopes) is applied by parsing the source organism rather than by set-match.
356
+
357
+ `evidence` records why each row is what it is: `eluted_but_positive_elsewhere=5,305`,
358
+ `eluted_foreign=103,351`, `eluted_self=1,156,574`, `eluted_unknown=440,458`, `tcell_negative=110,312`,
359
+ `tcell_positive=158,413`.
360
+
361
+ `corpus_iedb_mhc2.log` is the generator's own run record, deposited beside it; the counts in the table
362
+ above are read from it. The build is deterministic — regenerated 2026-08-19 from a clean checkout, every
363
+ count reproduced exactly.
364
+
365
+ Regenerate (needs this mirror at `$MHCMATCH_PMHC_DIR`, and the benchmark repo):
366
+
367
+ ```zsh
368
+ python bench/neoag/corpus_iedb.py --cls mhc2
369
+ ```
370
+
371
+ ---
372
+
373
+ ## `neoprecis_anchor_triplets.parquet` — added 2026-08-24
374
+
375
+ 14,140 rows × 12. Anchor / positive / negative peptide triplets by allele, with the mutated position
376
+ and the reference and alternate residue for each of the positive and negative members. A **motif
377
+ resource** rather than a screen: the triplets were constructed to train an allele-conditioned
378
+ recognition model, so `positive_peptide` and `negative_peptide` are **derived** designations, not
379
+ assay outcomes. `mhc` and `allele` are as-reported.
380
+
381
+ Deposited by `bench/neoag/ingest_vaccines.py` from Supplementary Data 3 of Lee KH, Sears TJ,
382
+ Zanetti M, Carter H, "NeoPrecis: enhancing immunotherapy response prediction through integration of
383
+ qualified immunogenicity and clonality-aware neoantigen landscapes", *Nat Commun* 2026;**17**(1),
384
+ doi:[10.1038/s41467-026-68651-6](https://doi.org/10.1038/s41467-026-68651-6),
385
+ PMID [41577704](https://pubmed.ncbi.nlm.nih.gov/41577704/), PMC12932759.
386
+
387
+ ---
388
+
389
+ ## `pathogen_tcell_{mhc1,mhc2}_{human,mouse}.tsv.gz` — the four pathogen fits' corpora (2026-09-05)
390
+
391
+ The training corpora for the `{cls}.{species}.pathogen` cells of `mhcmatch`'s aggregate scorer.
392
+ One row per **peptide**, not per assay: a peptide's assays are folded to a single label, and the
393
+ counts that produced it travel with it.
394
+
395
+ ### Source — the full IEDB T-cell export, not a filtered download
396
+
397
+ `dump/tcell_full_v3_tsv.zip` (`tcell_full_v3.tsv`, 1,346,206,535 bytes, **577,219 assay rows**,
398
+ IEDB build dated 2026-08-11). This is the complete export. The earlier
399
+ `raw/immunogenicity/iedb_tcell_assays_29032026.tsv.gz` (397,648 rows) and
400
+ `raw/immunogenicity/mouse_datasets/iedb_mouse.tsv.zip` (25,160 rows, **all `Positive*`, all class
401
+ I**) are query-filtered downloads; the mouse one carries no negatives at all, which is why the
402
+ mouse cells looked unfittable until the full export was read.
403
+
404
+ The export carries a **two-row header** — row 1 is the block (`MHC Restriction`), row 2 the field
405
+ (`Name`) — so a column is identified by the *pair*, never by index. Three blocks repeat the same
406
+ field names (`Epitope`, `Related Object`, `Assay Antigen` each carry `Species IRI`).
407
+
408
+ > Vita R, Blazeska N, Marrama D, et al. **The Immune Epitope Database (IEDB): 2024 update.**
409
+ > *Nucleic Acids Res* 2025;53(D1):D436–D443.
410
+ > PMID [39558162](https://pubmed.ncbi.nlm.nih.gov/39558162/) ·
411
+ > doi:[10.1093/nar/gkae1092](https://doi.org/10.1093/nar/gkae1092)
412
+
413
+ Organism classification is **NCBI Taxonomy**, fetched by taxid from the export's own
414
+ `Epitope / Species IRI` (`NCBITaxon_<taxid>`) via E-utilities `efetch?db=taxonomy`; 897 of 897
415
+ distinct taxids resolved. Cached at `bench/pathogen/_work/lineage.parquet` in the benchmark repo.
416
+
417
+ ### Filtering procedure, in order
418
+
419
+ 1. **Source organism is a pathogen** — the NCBI lineage is one of `virus`, `bacteria`, `archaea`,
420
+ `fungi`, `protozoa` (Apicomplexa, Euglenozoa, Amoebozoa, Fornicata, Parabasalia,
421
+ Heterolobosea), `helminth` (Platyhelminthes, Nematoda, Acanthocephala). **Read off the tree,
422
+ never off the organism's name.** 577,219 → 363,242 rows.
423
+ Deliberately excluded, and each is a real category rather than noise: `mammal` (97,019 rows —
424
+ self and xeno), `arthropod` (26,679 — house-dust mite, cockroach), `plant` (23,854 — timothy
425
+ grass and the other pollens), `other_metazoa` (2,738 — **`Gallus gallus` ovalbumin, 1,650 rows,
426
+ and `Tetronarce californica` AChR, 915**, the two canonical model antigens), and `unassigned`
427
+ (63,556, of which 61,664 record no source organism at all).
428
+ 2. **Linear peptide, canonical residues** — `Epitope / Object Type == "Linear peptide"` and the
429
+ sequence matches `^[ACDEFGHIKLMNPQRSTVWY]+$`, which drops modified and non-peptidic epitopes.
430
+ 363,242 → 361,795.
431
+ 3. **The assay is labelled** — `Positive`, `Positive-Low/Intermediate/High` → 1; `Negative` → 0;
432
+ nothing else exists in this column. 361,795 rows, **126,593 positive**. The grade survives as
433
+ the `grade` column (`Positive-High` 4 … `Positive-Low` 1, 0 for a negative) rather than being
434
+ collapsed away.
435
+ 4. **The restriction resolves** — `MHC Restriction / Name` must resolve **exactly** to one groove
436
+ under `mhcmatch.pseudoseq.resolve_allele` *and* be present in the shipped panel, or name an H-2
437
+ haplotype (`H2-b class I`). 247 distinct names resolve onto the panel, 19 name a haplotype, 73
438
+ resolve off-panel (all `BoLA-*`, cattle), 286 do not resolve.
439
+ **`exact=True` is load-bearing.** `HLA-A2` is a serotype; `resolve_allele("HLA-A2", "mhc1")`
440
+ answers `("HLA-A2301", False)` — a *different molecule*, on 3,963 rows. An inexact hit here does
441
+ not raise and does not return nothing; it scores the wrong groove.
442
+ 5. **Length window** — class I 8–11, class II 12–20. 17,306 rows fall outside.
443
+ 6. **Fold to one row per peptide per cell** — `label = max` over the peptide's assays.
444
+ `n_assay`, `n_pos`, `n_neg` are kept so a stricter rule (majority, or drop-on-conflict) stays
445
+ available; 6,205 peptides across the four cells carry both a positive and a negative assay.
446
+
447
+ ### The cells
448
+
449
+ | file | peptides | positive | prevalence | groove-specific | haplotype-only | allotypes |
450
+ |---|--:|--:|--:|--:|--:|--:|
451
+ | `pathogen_tcell_mhc1_human.tsv.gz` | 16,841 | 7,018 | 0.417 | 16,841 | 0 | 116 |
452
+ | `pathogen_tcell_mhc1_mouse.tsv.gz` | 10,445 | 2,203 | 0.211 | 8,306 | 2,139 | 8 |
453
+ | `pathogen_tcell_mhc2_human.tsv.gz` | 7,948 | 5,150 | 0.648 | 7,948 | 0 | 86 |
454
+ | `pathogen_tcell_mhc2_mouse.tsv.gz` | 11,732 | 3,326 | 0.283 | 3,948 | 7,784 | 7 |
455
+
456
+ Restricted to peptides whose restriction is not **exclusively** prediction-derived
457
+ (`MHC binding prediction`, `Statistically inferred by motif or alleles present`):
458
+
459
+ | cell | peptides | positive |
460
+ |---|--:|--:|
461
+ | `mhc1.human.pathogen` | 14,108 | 5,894 |
462
+ | `mhc1.mouse.pathogen` | 6,763 | 1,456 |
463
+ | `mhc2.human.pathogen` | 4,695 | 3,347 |
464
+ | `mhc2.mouse.pathogen` | 10,833 | 2,928 |
465
+
466
+ `mhc_evidence` ships as a column and the filter is the **caller's**, because a restriction that is
467
+ itself an MHC binding prediction is circular for a model whose leading term is `binder` — and it is
468
+ the single largest evidence code in `mhc1.mouse` (3,872 of 10,445 peptides).
469
+
470
+ ### Schema
471
+
472
+ `peptide`, `label`, `n_assay`, `n_pos`, `n_neg`, `grade`, `alleles`, `haplotypes`,
473
+ `allele_source`, `mhc_evidence`, `assay_methods`, `source_species`, `kingdom`,
474
+ `source_molecules`, `pmids`, `length`. Multi-valued fields are `;`-joined and sorted.
475
+
476
+ **Not** the compendium's shared 19-column schema: these are peptide-level with assay counts, and
477
+ they carry no `mhc_a`/`mhc_b`/`affinity`.
478
+
479
+ ### `allele_source` is a key, not a convention
480
+
481
+ A restriction is either an allotype (`H-2Kb`) or a haplotype (`H2-b class I`), and the two are
482
+ told apart by the `allele_source` column rather than by a rule the reader has to carry. Haplotype
483
+ rows keep the **letter** in `haplotypes` and leave `alleles` empty; expanding a haplotype to its
484
+ allotypes is a **model** decision and is not baked in. Two reasons it is not:
485
+
486
+ - The panel's own spelling differs between classes — class I is `H-2Kb`, class II is `H-2-IAb`,
487
+ and `resolve_allele` returns a third form (`H-2-Kb`). Every join here folds through
488
+ `normalize_allele` on both sides.
489
+ - A mechanical superscript expansion is **wrong for class II**: the panel carries `H-2-IEb`, but
490
+ the b haplotype does not express I-E. Class-I K/D/L expansion has no such problem. The
491
+ haplotype-restricted best-of arm measured in `bench/results/pathogen_allele_policy_arms.md`
492
+ is where that decision belongs.
493
+
494
+ ### Two skews to check before pooling anything over these
495
+
496
+ - `mhc2.mouse`'s groove-specific rows are **3,474 of 3,957 allotype mentions `H-2-IAb` (87.8%)**.
497
+ A pooled null over this cell is the I-Ab motif. The haplotype rows are what carry d, k and s.
498
+ - `mhc1.human` is 5,237 `HLA-A*02:01` of 16,841 peptides (31.1%).
499
+
500
+ ### Regenerate
501
+
502
+ ```zsh
503
+ cd ~/vcs/projects/2026-mhcmatch-benchmark && source .venv/bin/activate
504
+ python bench/pathogen/extract_tcell.py # 577,219 x 15, ~9 s
505
+ python bench/pathogen/fetch_lineage.py # 897 taxids from NCBI, 5 efetch calls
506
+ python bench/pathogen/build_pathogen_corpus.py # writes the four files here
507
+ python bench/pathogen/proofread_pathogen_corpus.py # 10 invariants per cell, exits 1 on any
508
+ ```
509
+
510
+ ### What was fitted on them (added 2026-09-05, after the fits)
511
+
512
+ Three of the four cells became **shipped** `mhcmatch` artifacts in release **1.15.0**, completing
513
+ `rank.AGGREGATE_ARTIFACTS` at eight of eight `(cls, species, mode)` cells:
514
+
515
+ | corpus file | artifact | peptides fitted | positives | prevalence | allotypes | ROC-AUC | PR-AUC |
516
+ |---|---|--:|--:|--:|--:|--:|--:|
517
+ | `pathogen_tcell_mhc1_mouse.tsv.gz` | `aggregate_mhc1_mouse_pathogen.json` | 10,404 | 2,196 | 0.2111 | 8 | 0.5564 | 0.2665 |
518
+ | `pathogen_tcell_mhc2_human.tsv.gz` | `aggregate_mhc2_human_pathogen.json` | 7,946 | 5,148 | 0.6479 | 80 | 0.6064 | 0.7361 |
519
+ | `pathogen_tcell_mhc2_mouse.tsv.gz` | `aggregate_mhc2_mouse_pathogen.json` | 11,725 | 3,324 | 0.2835 | 7 | 0.6224 | 0.4098 |
520
+ | `pathogen_tcell_mhc1_human.tsv.gz` | `aggregate_mhc1_pathogen.json` (from 2026-09-06) | 16,790 | 7,002 | 0.4170 | 104 | 0.5988 | 0.5147 |
521
+
522
+ Both figures are in-sample, pooled off the logit, each against its own prevalence. Fitted peptide
523
+ counts are below the file's row counts because `drop_corpus_members` removes peptides that are exact
524
+ members of a corpus table the fit's own `C_corpus_*` channels read (36 on mouse class I, 51 on human
525
+ class I) and because a peptide whose candidate allotypes all fail to score is dropped.
526
+
527
+ **`mhc1.human.pathogen` was on the Kesmir corpus, and moved to this one on 2026-09-06.** Until
528
+ then the shipped fit for that cell stayed on `kesmir_rebuilt_hla_matched.tsv.gz`, whose negatives
529
+ are *eluted ligands with no positive T-cell record* -- presentation held fixed on both sides. Here a
530
+ negative was *assayed and did not respond*, of unknown presentation, so `binder` carries
531
+ presentation again: **+0.1454 against +0.3418** for the same cell on the two corpora. The move was
532
+ the author's call, taken with the refit that put every pathogen cell on one corpus and one balanced
533
+ design; the paragraph above this one described the earlier state and is corrected rather than
534
+ deleted, because a recorded number cites the corpus it was produced on.
535
+
536
+ Two things not to carry over from the Kesmir arm. Its PR-AUC is **not** comparable to this one --
537
+ 0.0905 at prevalence 0.0691 against 0.5147 at 0.4146, which is 1.31x and 1.24x lift respectively,
538
+ i.e. the same. And its five terms are not these two: `C_phys_buried` and `C_corpus_thymus` were
539
+ each highly resolved there and are absent here, because `C_corpus_thymus` and `C_corpus_self`
540
+ correlate at +0.683 to +0.792 and the five-term fits were fitting their difference.
541
+
542
+ Fitted by `bench/pathogen/fit_tcell.py` in `2026-mhcmatch-benchmark`; per-cell reports at
543
+ `bench/results/pathogen_tcell_fit_{cls}_{species}.md`, the head-to-head at
544
+ `bench/results/pathogen_mhc1_human_v1_vs_v2.md`, and the shipped-artifact provenance in
545
+ `src/mhcmatch/data/PROVENANCE.md`.
546
+
547
+ ```zsh
548
+ MHCMATCH_MODEL_RELEASE=1.15.0 python bench/pathogen/fit_tcell.py --cls mhc1 --species mouse
549
+ MHCMATCH_MODEL_RELEASE=1.15.0 python bench/pathogen/fit_tcell.py --cls mhc2 --species human
550
+ MHCMATCH_MODEL_RELEASE=1.15.0 python bench/pathogen/fit_tcell.py --cls mhc2 --species mouse
551
+ ```
552
+
553
+ **One consequence for anyone scoring with these artifacts.** Each was fitted on the
554
+ best-**presenting** allotype among those the peptide's own record supports, so each declares
555
+ `allele_policy = {"kind": "panel", "select": "presentation", "source": "record"}` and `mhcmatch`
556
+ refuses a bare `rank pairs` run rather than letting the coefficients meet a shifted column.
557
+ `rank fasta --alleles A,B` and `rank pairs --allele-panel A,B` are the supported forms. Measured on
558
+ the fitted designs, the best-of-record `binder` sits a mean **−0.2146** (sd 0.5262) below the same
559
+ peptide's single-recorded-allele `binder` on `mhc1_human`, and −0.1403 / −0.1248 / −0.0533 on
560
+ `mhc1_mouse` / `mhc2_human` / `mhc2_mouse`; 62.3 % / 73.7 % / 63.5 % / 52.7 % of peptides carry
561
+ exactly one candidate and are unaffected.
562
+
563
+ ## `pathogen_tcell_balanced_{mhc1,mhc2}_{human,mouse}.tsv.gz` — allele-balanced fitting sets (2026-09-05)
564
+
565
+ **Derived**, not experimental. One row per peptide, carrying the allotype it is attributed to, its
566
+ T-cell label, and a prior weight that removes the allotype-composition difference between the two
567
+ labels. Built from `pathogen_tcell_{mhc1,mhc2}_{human,mouse}.tsv.gz` (entry above) by
568
+ `bench/pathogen/netmhcpan_best.py --cls {mhc1,mhc2} --species {human,mouse} --min-records 5` in
569
+ `repseq/2026-mhcmatch-code`. Class I goes through NetMHCpan-4.2, class II through
570
+ NetMHCIIpan-4.3.
571
+
572
+ | column | meaning |
573
+ |---|---|
574
+ | `peptide` | the epitope, as deposited |
575
+ | `mhc_a` | NetMHCpan's own best allotype for that peptide, panel spelling |
576
+ | `label` | `pos` / `neg`, the max over that peptide's IEDB T-cell assays |
577
+ | `weight` | prior (case) weight for a weighted GLM; see below |
578
+
579
+ **Why the allele comes from NetMHCpan.** The balancing variable must not be the model's own choice.
580
+ `fit_tcell.score` picks the best-*presenting* allotype with mhcmatch, so weighting on it would
581
+ condition the design on the term being fitted (`binder`). NetMHCpan-4.2 is external to every term in
582
+ these fits, so its `%Rank_EL` argmin over each record's **own recorded candidate allotypes** is the
583
+ allele used. It is not a free argmin over the whole panel — that would invent a restriction the
584
+ record does not carry.
585
+
586
+ **The two classes spell an allotype differently, and the resolver tries both.** The class-II
587
+ deposits already store NetMHCIIpan's own names (`DRB1_0101`, `HLA-DPA10103-DPB10401`, `H-2-IAb`),
588
+ so putting them through the canonicaliser converts them *away* from what the tool accepts; class I
589
+ is the opposite, the deposit writes `HLA-A*02:01` and the tool wants `HLA-A02:01`. The builder
590
+ resolves from an ordered list — as deposited, then canonical — and prints which spelling answered.
591
+ All 116 human and 8 mouse class-I candidate allotypes and all 86 human and 7 mouse class-II ones
592
+ resolve; none is dropped for want of a name.
593
+
594
+ **Order: predict, then filter, then weight.** Every (peptide, candidate allotype) the record
595
+ supports goes through NetMHCpan — one call per allotype, never one per peptide. Each peptide takes
596
+ its `%Rank_EL` argmin. An allotype with fewer than 5 peptides **in either class** is then dropped
597
+ together with its peptides: a frequency ratio resting on one or two peptides on one side is noise,
598
+ and an allotype absent from a class has no ratio at all. 0 of 116 human and 0 of 8 mouse candidate
599
+ allotypes lack a NetMHCpan name; the thin-tail cut removes 64 of 107 human and 2 of 8 mouse
600
+ class-I best-allele allotypes, 53 of 82 human class-II ones, and 0 of 7 mouse class-II ones.
601
+
602
+ **The weight.** The target is the **pooled** allotype distribution `p(a) = n(a)/N` over both labels,
603
+ and each label is normalised against it: `w = p(a)/p_pos(a)` on positives and `p(a)/p_neg(a)` on
604
+ negatives. Each label's weights sum to its own count identically — `sum_a n_c(a)·p(a)/p_c(a) =
605
+ N_c·sum_a p(a) = N_c` — so the prevalence is unchanged and only the allotype mix within each label
606
+ moves. **Nothing is subsampled:** every peptide surviving the two filters is present, which is why
607
+ this ships as a weight column rather than as a matched subset.
608
+
609
+ This is the same operation as cross-batch V/J-gene usage correction in `vdjtools`
610
+ (`preprocess.correct_vj_usage`, batch → label, gene → allotype). Two deliberate differences: that
611
+ implementation targets the *geometric* grand mean of `log p` and keeps rare genes with a Laplace
612
+ pseudocount, where this targets the arithmetic pooled marginal and drops the thin tail; and its
613
+ `apply_vj_correction` roulette-wheel *resamples* the table, where this reweights it.
614
+
615
+ | file | rows | pos | neg | prevalence | allotypes | balanceable | rows at w=1 | min w | max w | max \|Δp\| all, raw | all, weighted | subset, weighted |
616
+ |---|--:|--:|--:|--:|--:|--:|--:|--:|--:|--:|--:|--:|
617
+ | `pathogen_tcell_balanced_mhc1_human.tsv.gz` | 16,790 | 7,002 | 9,788 | 0.4170 | 107 | 43 | 1,236 | 0.4596 | 7.0060 | 0.1255 | 0.0608 | 1.1e-16 |
618
+ | `pathogen_tcell_balanced_mhc1_mouse.tsv.gz` | 10,404 | 2,196 | 8,208 | 0.2111 | 8 | 6 | 6 | 0.2825 | 3.1045 | 0.0955 | 0.0017 | 1.6e-15 |
619
+ | `pathogen_tcell_balanced_mhc2_human.tsv.gz` | 7,946 | 5,148 | 2,798 | 0.6479 | 82 | 29 | 659 | 0.4208 | 7.0918 | 0.0616 | 0.0185 | 3.3e-16 |
620
+ | `pathogen_tcell_balanced_mhc2_mouse.tsv.gz` | 11,725 | 3,324 | 8,401 | 0.2835 | 7 | 7 | 0 | 0.4191 | 2.2146 | 0.2736 | 0.0000 | 1.1e-15 |
621
+
622
+ **Rebuilt 2026-09-06 under one weighting implementation, and the thin tail is now kept rather than
623
+ dropped.** These four files were built by an inline copy of `pooled_weights` that *dropped* an
624
+ unbalanceable allotype together with its peptides, while the two `neoag_tcell_balanced_*` files
625
+ below were built by the shared `balance.py`, which *keeps* them at weight 1. One entry in this file
626
+ described both. Every caller goes through `balance.py` now, at `drop_thin=False`, which readmits
627
+ 1,236 / 6 / 659 / 0 peptides and 554 / 5 / 608 / 0 positives.
628
+
629
+ **So the residual over the FULL allotype set is not zero, and the honest claim is the narrow one:**
630
+ within the balanceable subset the two weighted distributions agree to 1e-15 or better in every
631
+ cell, and over all allotypes the imbalance falls but does not vanish -- 0.1255 to 0.0608 on human
632
+ class I, where 64 of 107 allotypes are thin. `mhc2.mouse` reaches exactly zero because it is the
633
+ one cell with no thin allotype at all. The largest raw imbalance is **mouse class II at 0.2736**,
634
+ the `H-2-IAb` dominance this compendium already documents on the ligand side.
635
+
636
+ Kish effective sample size, the price of the reweighting:
637
+
638
+ | file | positives, n → ESS | negatives, n → ESS |
639
+ |---|---|---|
640
+ | `..._mhc1_human` | 6,448 → 4,606 (71.4%) | 9,106 → 7,942 (87.2%) |
641
+ | `..._mhc1_mouse` | 2,191 → 2,025 (92.4%) | 8,207 → 8,078 (98.4%) |
642
+ | `..._mhc2_human` | 4,540 → 4,019 (88.5%) | 2,747 → 1,980 (72.1%) |
643
+ | `..._mhc2_mouse` | 3,324 → 2,839 (85.4%) | 8,401 → 8,135 (96.8%) |
644
+
645
+ The weights are consumed by `bench/pathogen/fit_tcell.py --balance`, which passes them as prior
646
+ weights to `bench/epic/optimize.py::irls(pw=)`. `pw=None` is bit-identical to the unweighted fit.
647
+ Per-allotype composition and weights are tabulated in
648
+ `bench/results/pathogen_tcell_allele_balance_{mhc1,mhc2}_{human,mouse}.md`.
649
+
650
+ **These four sets are what the 1.15.0 pathogen models are fitted on**, all four with `--balance`:
651
+ class I on `binder + C_corpus_self`, class II on `binder + C_phys_buried + C_phys_charge` (class II
652
+ has no corpus block — the mimicry tables are class-I TCR-face geometry).
653
+
654
+ ## `neoag_tcell_balanced_mhc2_{human,mouse}.tsv.gz` — allele-balanced class-II neoantigen fitting sets (2026-09-06)
655
+
656
+ **Derived**, not experimental. The same weighting as the four `pathogen_tcell_balanced_*` files
657
+ above, on a different corpus and a different balancing variable. Built by
658
+ `bench/mhc2_neoag/allele_balance.py --species {human,mouse} --min-records 5` in
659
+ `repseq/2026-mhcmatch-code`, which calls the two class-II neoantigen fits' own loaders
660
+ (`fit_human_mhc2.load`, `fit_mouse.deposit` ∪ `fit_mouse.cedar`) rather than re-reading the
661
+ deposits, so each balanced corpus spans exactly the population its fit sees.
662
+
663
+ | column | meaning |
664
+ |---|---|
665
+ | `peptide` | the epitope, as deposited |
666
+ | `mhc_a` | the allotype the row is attributed to; **the deposit's own spelling**, see below |
667
+ | `label` | `pos` / `neg`, the deposited `immunogenicity` |
668
+ | `weight` | prior (case) weight for a weighted GLM; the pooled-marginal weight defined above |
669
+
670
+ **The unit is the `(peptide, allotype)` pair, and there is no NetMHCpan argmin here.** That is the
671
+ one substantive difference from the pathogen corpora, and it is a property of the deposits. The
672
+ pathogen deposits carry a peptide-level label and an unresolved *set* of candidate restrictions, so
673
+ an argmin resolves them. These deposits carry a measured restriction per assay: 589 of 804 human
674
+ peptides and 795 of 808 mouse ones appear with exactly one allotype, only 10 human and 25 mouse
675
+ peptides disagree on the label across their allotypes, and both fits already score each row against
676
+ its own allele. An argmin would collapse 1,112 human pairs to 804 and replace a *measured*
677
+ restriction with a predicted one.
678
+
679
+ | species | balancing variable | why it is external to every fitted term |
680
+ |---|---|---|
681
+ | human | CEDAR's deposited `allele`, DRA folded | an assay annotation; nothing mhcmatch computed |
682
+ | mouse | `mhc_a_pred` | already NetMHCIIpan-4.3's argmin over the strain's candidate alleles, with any reported allele added to the set — see "Predicted restriction" in the benchmark repo's `SOURCES.md` |
683
+
684
+ **The DRA fold is not tidiness.** `HLA-DRA*01:01/DRB1*04:04` and `HLA-DRB1*04:04` are one molecule —
685
+ DRA is effectively monomorphic — and as deposited they are two allotypes splitting one allotype's
686
+ counts, which is exactly what a thin-tail cut punishes. 72 spellings, 68 molecules.
687
+
688
+ **Three allele vocabularies now appear across the six balanced files, one per deposit, and each is
689
+ the spelling its own fit consumes.** The pathogen class-II files carry NetMHCIIpan's names
690
+ (`DRB1_0101`, `HLA-DPA10103-DPB10401`, `H-2-IAb`); this human file carries the panel spelling
691
+ (`HLA-DRB1*04:01`, `HLA-DQA1*03:01/DQB1*03:02`); this mouse file carries the deposit's
692
+ (`H2-IAb`, without the second hyphen). All 11 spellings across these two files resolve to an exact
693
+ match through `mhcmatch.pseudoseq.resolve_allele`; none is dropped for want of a name.
694
+
695
+ **The thin tail is kept at weight 1, not cut -- and this entry described a cut until 2026-09-06.**
696
+ An allotype with zero rows in one class cannot be balanced by any weight: the target `p_pool(a)` is
697
+ strictly positive while the achievable `p(a | that class)` is identically zero, and a Laplace
698
+ pseudocount makes the *ratio* finite without supplying the missing rows. There are then two ways
699
+ not to lose those rows, and the author's call is the second: drop them, or keep them at weight 1 and
700
+ compute the target marginal within the balanceable subset. **The files always held the second and
701
+ this entry always described the first**, so its counts (693 / 271 / 7 allotypes human, 538 / 214 / 4
702
+ mouse) were the cut that was never shipped, and its claim that both weighted distributions "are
703
+ equal to floating-point noise in every cell of both files" is true only *within* the balanceable
704
+ subset. The cascade a cut would have cost is reported at four thresholds in
705
+ `bench/results/neoag_tcell_allele_balance_mhc2_{human,mouse}.md`.
706
+
707
+ **The human corpus is where keeping rather than cutting matters most.** Whole allotypes in CEDAR
708
+ are all-positive — `HLA-DRB1*04:05` 31/31, `HLA-DRB1*07:01` 24/24, `HLA-DRB1*15:02` 22/22,
709
+ `HLA-DRB5*01:01` 17/17 — because a study that reports a responder's restriction reports no
710
+ negatives for it, while the two `DQB1*03:02` composites are ~90 % negative from a type-1-diabetes
711
+ screen. Allotype and label are confounded in the deposit, and those are exactly the allotypes no
712
+ weight can touch. Cutting them would have cost **59 % of the positives**; keeping them at weight 1
713
+ costs a residual imbalance of **0.1273** over all 72 allotypes instead, against 5.6e-17 within the
714
+ 7 that can be balanced. Neither number alone describes the file, so both are in the table below.
715
+
716
+ | file | rows | pos | neg | prevalence | allotypes | balanceable | rows at w=1 | min w | max w | max \|Δp\| all, raw | all, weighted | subset, weighted | Kish n_eff |
717
+ |---|--:|--:|--:|--:|--:|--:|--:|--:|--:|--:|--:|--:|--:|
718
+ | `neoag_tcell_balanced_mhc2_human.tsv.gz` | 1,112 | 656 | 456 | 0.5899 | 72 | 7 | 419 | 0.5281 | 4.2690 | 0.2427 | 0.1273 | 5.6e-17 | 830.7 (74.7 %) |
719
+ | `neoag_tcell_balanced_mhc2_mouse.tsv.gz` | 627 | 261 | 366 | 0.4163 | 7 | 4 | 46 | 0.5010 | 3.0430 | 0.2800 | 0.0994 | 1.1e-16 | 532.1 (84.9 %) |
720
+
721
+ **Nothing is retained-against: every row each fit sees is present.** 1,112 of 1,112 human rows and
722
+ 627 of 627 mouse ones, all 656 and 261 positives, over all 72 and 7 allotypes. A cut at
723
+ `min_records = 5` would have kept 693 / 271 / 7 and 538 / 214 / 4.
724
+ positives (41.3 %) over 7 of 68 allotypes; mouse 538 of 572 rows (94.1 %) and 214 of 247 positives
725
+ (86.6 %) over 4 of 7 allotypes. The two weighted allotype distributions are equal to floating-point
726
+ noise in every cell of both files.
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+ # tcell: 570,434 staged rows
2
+ # class II: 247,456
3
+ # host human or mouse: 239,196
4
+ # 11-25mer, canonical: 221,787
5
+ # after artefact exclusions: 221,787 (115,263 peptides)
6
+ # ligand: 5,773,495 staged rows
7
+ # class II: 1,966,913
8
+ # host human or mouse: 1,905,785
9
+ # 11-25mer, canonical: 1,752,626
10
+ # after artefact exclusions: 1,752,626 (538,321 peptides)
11
+ # T-cell assayed (peptide, host) pairs: 119,615
12
+ # positive: 39,818 | measured negative: 79,797
13
+ # peptides positive in any host: 38,320
14
+ # elution pool by origin: foreign=145,186, self=1,165,489, unknown=441,951
15
+ # rows entering allele resolution: 1,974,413 (1,324,898 distinct peptide/host/restriction)
16
+ # class-II restriction parsed on 1,003,554/1,974,413 rows (50.8%); the rest are kept as `unresolved`
17
+ # top groups: unresolved=970,859, HLA-DRB1*15=174,134, HLA-DRB1*13=163,275, HLA-DRB1*01=105,737, HLA-DRB1*03=83,487, HLA-DRB1*04=52,007, HLA-DRB1*07=39,065, HLA-DPA1*01/DPB1*04=37,066
18
+ # evidence: eluted_but_positive_elsewhere=5,305, eluted_foreign=103,351, eluted_self=1,156,574, eluted_unknown=440,458, tcell_negative=110,312, tcell_positive=158,413
19
+ # out: 1,096,034 (peptide, allele group, host) rows over 643,744 peptides, 77,943 immunogenic
20
+ # human: 1,036,041 rows, 65,486 positive, 121 allele groups, 603,781 peptides
21
+ # length 11-25, median 15
22
+ # mouse: 59,993 rows, 12,457 positive, 40 allele groups, 50,258 peptides
23
+ # length 11-25, median 15
24
+ # -> /Users/mikesh/vcs/projects/2026-mhcmatch-benchmark/bench/neoag/corpus_iedb_mhc2.parquet (3s)
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