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tcren

tcren — structure-based prediction of TCR–epitope recognition

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TCRen predicts which epitopes a T-cell receptor recognises from a single TCR–peptide–MHC structure (experimental or modelled). It extracts the TCR–peptide contact map and scores every candidate peptide with a residue-level statistical potential derived from contact preferences in TCR:pMHC crystal structures — answering not "what fancy complex can a model draw?" but "is this binding physically plausible?".

This is a documented, tested, CLI-driven Python library. TCR chains are annotated with the sibling arda; MHC chains are mapped and the groove partitioned against a curated reference; structures are oriented into one canonical frame; and the original contact maps, potential, and scores are reproduced numerically (validated against committed oracles to floating-point precision).

Where the original tcren scored TCR:peptide contacts alone, this version also scores the TCR:MHC and peptide:MHC interfaces, which a full picture of TCR:pMHC binding mechanics and any ΔΔG estimate both need.

What it is evaluated on

Five benchmark blocks, chosen to cover the problems users actually bring to a modelled TCR:pMHC complex. They are the backbone of the accompanying manuscript, and they are why the API looks the way it does.

block the question entry points
Combinatorial peptide libraries which peptides does this receptor read? tcren features, tcren assess (peptide_score), tcren.ddg.neoantigen_ddg
A functionally validated repertoire screen which receptors read this epitope? tcren features, tcren recognize
A balanced epitope panel, template-stratified the same, where no related complex has been solved as above, with template availability reported rather than inferred
Molecular dynamics with measured kinetics may a single static structure be scored at all, and what does its energy reach? the three interface energies, tcren.potts contact marginals
Model-confidence diagnostics which confidently modelled complexes are not real? tcren assess, tcren.score.confidence_residual, tcren.reliability.af_band

The last is the one most users reach for first: you already have an AlphaFold model and want to know whether to trust it. Everything in the score set is frozen on a hold-out that ships inside the wheel and refits from a manifest that ships with it, so nothing is estimated from the rows you score and no number depends on what was scored beside it.

What it does

Scope. tcren is for αβ TCR : peptide–MHC complexes and nothing else: a TRA and a TRB chain, a peptide of standard amino acids, and a class I or class II groove. γδ receptors, single-chain constructs, pMHC with no TCR and non-peptidic ligands are out of scope — derive-potential drops them rather than deriving from them, and there is no flag to widen it.

From one TCR–peptide–MHC structure (crystal or model), each task is one command or one call:

task command library
Score candidate epitopes for a TCR tcren score score_peptides
Rank peptides for a fixed receptor — the poly-alanine-referenced energy tcren assess --peptide score.peptide_score
Percentile-rank a peptide vs background tcren rank percentile_rank
ΔΔG of mutations (alanine scan / neoantigen) tcren ddg alanine_scan, neoantigen_ddg
Predict a CPL response matrix from a template tcren cpl response_matrix, mutation_effect, position_scan, equimolar_effect
Binder vs non-binder for a TCR model tcren features + tcren assess score.binder_score, score.score_table
Which part of the structure says so — the five channels tcren assess score.channel_scores
Is the reported confidence warranted? tcren assess score.confidence_residual
Every interface descriptor, 164 in six families (four by default) tcren features recognition_table(include=...), descriptors
All interface descriptors, one row per structure tcren recognize recognition_features
S — geometry, footprint shape and energy in native-sd units tcren recognize --features reliability.s_score
Is this model worth believing? — the score set, S and the generator diagnostic tcren assess score.score_table, reliability.s_score, af_band
Refit the frozen model from its shipped manifest tcren fit-holdout score.holdout_model, score.holdout_manifest
The contact map as a probability model — energy, partition function, per-pair contact probability tcren potts fit / score / contacts potts.fit_potts, score_sites, contact_probabilities
Three-interface energy Φ, poly-Ala ΔΦ, interface geometry tcren scoring run_pipeline
Annotate chains + region markup tcren annotate classify_chains, annotate_mhc
Interface contact table (5/8/12 Å) tcren contacts ContactMap, multi_contacts
Orient into the canonical MHC frame tcren superimpose / orient superimpose, canonicalize_structure
Graft a TCR onto another pMHC (chimera) tcren substitute-tcr substitute_tcr
Wrong-TCR decoy set (recognition negatives) tcren shuffle make_decoys, graft_tcr
Substitute a peptide + refine its pose tcren refine substitute_peptide, refine_peptide
Surface topology of the pMHC face — is this epitope featureless? tcren surface surface_map, surface_stats, surface_distance
Backbone dynamics — does the peptide hold its TCR-facing conformation? peptide_stability, stability_table
Repack side chains into their preferred rotamers tcren refine --repack repack
DOPE interface energy (ΔΔG e_native) tcren energy interface_energy
Interface mechanics — koff proxies (stiffness / rupture) tcren recognize --mechanics, or tcren mechanics alone interface_mechanics
Re-derive the statistical potential tcren derive-potential derive_tcren
Steric-clash / wrong-register QC interface_clashes, check_register
2D complementarity map + 3D pocket/CDR view render_complementarity_map, view_pocket_cdr
Publication PyMOL figures, with a labelled axis gizmo viz.pymol.render, overlay_scene, groove_scene, interface_scene

Scope — ranking, not affinity. TCRen ranks peptide/TCR specificity for a given receptor (and the ddg matrix is a fast triage, not a free energy). It is not an affinity model: on the ATLAS SPR benchmark neither the raw contact energy nor its poly-alanine difference predicts Kd/ΔG/koff/kon (|ρ|≤0.3). The one affinity-adjacent quantity a structure predicts is the off-rate koff, via interface mechanics (tcren mechanics) — not the contact sum.

Install

pip install tcren          # from PyPI — binary wheels ship the C++ extension; pulls in arda-mapper
tcren build-mhc-ref        # once: builds the MHC allele reference from IMGT (not bundled in the wheel)

tcren build-mhc-ref is a required one-time step after a pip install. The curated MHC allele reference is built from IMGT on demand rather than shipped in the wheel, and every command that annotates a structure needs it.

For development (a repo-local .venv via uv, an editable install, and the reference data fetched into data/):

bash setup.sh                    # uv venv + editable install + arda + fetch data/ (no conda)
source .venv/bin/activate

setup.sh needs only uv and a C++ compiler (macOS: xcode-select --install); it never touches conda. Pass --tests to run the fast suite after install.

tcren ships five small pybind11/C++ extensions, built on install by scikit-build-core (which fetches cmake+ninja automatically): tcren._align (MHC-pseudosequence fitting alignment; a Biopython fallback runs if unbuilt), tcren._refine (DOPE atom-level Monte-Carlo peptide refinement), tcren._relax (DOPE interface energy for tcren energy / ΔΔG), tcren._fold (CCD loop closure) and tcren._geom (interface geometry, clash detection and contact stability). TCR annotation is provided by arda, a runtime dependency published to PyPI as arda-mapper (it imports as arda); uv/setup.sh pull it automatically, and from arda-mapper >= 2.5.7 it auto-fetches both its own reference and a static mmseqs2 binary on first use — so no conda/bioconda and no ARDA_HOME to set (override the binary with $ARDA_MMSEQS). setup.sh also runs tcren fetch-data to populate data/ with the reference structure sets (Native2026, Canonical2026) used by orient/superimpose (set TCREN_NO_FETCH=1 to skip).

Command line

# Score structures: the three interface contact energies (TCRen for TCR↔peptide, MJ for
# TCR↔MHC and peptide↔MHC) and their total Φ. One row per structure.
tcren scoring -s complex.pdb.gz -o scores.csv

# Inputs: a file, a directory, a .tar.gz, a quoted glob, a .txt manifest (one path per line),
# a comma-separated list, or a repeated -s. Mix freely.
tcren scoring -s a.pdb.gz -s b.pdb.gz -o scores.csv
tcren scoring -s 'models/*.pdb.gz' -o scores.csv
tcren scoring -s models/ --delta --geometry -t 8 -o scores.csv   # a directory, 8 workers
tcren scoring -s models.txt -o scores.csv

# --delta adds the poly-alanine reference ΔΦ per interface (ΔΦ_TCR:MHC is identically 0).
# Use ΔΦ, not Φ, when each candidate carries its OWN generated pose: raw Φ then partly reads
# the pose the predictor chose rather than the peptide.
tcren scoring -s 'models/*.pdb.gz' --delta -o scores.csv

# --geometry adds the interface descriptors and Q, the directional decorrelated
# interface-quality score (native-crystal calibrated, so it is defined for a single structure).
tcren scoring -s complex.pdb.gz --delta --geometry -o scores.csv

# Configurable per-interface potential: swap a bundled name (tcren2|karnaukhov2022|mj|keskin),
# a CSV, or None for any interface; default reproduces the built-in per-interface families exactly.
tcren scoring -s complex.pdb -o scores.csv --tcr-mhc-potential keskin

# Opt-in TCR framework regions: --regions {all,cdr,cdr+fr} chooses which TCR regions
# contribute on the TCR side (cdr = CDR1-3 only; cdr+fr adds FR1-3; all = unfiltered, default).
tcren score -s complex.pdb -c candidates.txt -o ranked.csv --regions cdr+fr

# Surface topology: the pMHC face a TCR meets, BEFORE any TCR is there. A height field over
# the groove with hydropathy and charge painted on, per structure, plus the scalars that make
# "featureless" a number: relief, peak_to_valley, frac_above_ridge.
tcren surface -s complex.pdb -o surface.csv

# --compare writes the pairwise map distance (SURFMAP's Manhattan metric) so epitopes cluster;
# --svg writes one figure per structure; --cells writes the long per-cell table.
tcren surface -s models/ -o surface.csv --compare dist.csv --svg figs/ --channel h
tcren surface -s models/ -o surface.csv --channel phobic --scale kd   # or --scale mj

# Three opt-in reweightings of the SAME energy sum, all off by default so nothing moves
# unless asked:
#   --drop-untyped      ignore contacts that are only proximity (no h-bond / salt bridge /
#                       stacking / hydrophobic / polar chemistry)
#   --position-weights  weight by where a contact sits on the peptide (central | tcr_facing);
#                       a clash at an anchor the TCR never touches is not a clash at P5
#   --soft              replace the hard 5 A cutoff with a contact PROBABILITY averaged over
#                       side-chain rotamers, Boltzmann-weighted under DOPE
tcren score -s complex.pdb -c candidates.txt -o ranked.csv --drop-untyped
tcren score -s complex.pdb -c candidates.txt -o ranked.csv --position-weights central
tcren score -s complex.pdb -c candidates.txt -o ranked.csv --soft

# Opt-in intra-peptide term: every interface energy sums over contacts between two DIFFERENT
# chains, so a candidate held in the template's conformation by its own side chains costs the
# same as one that is not. --intra-weight w adds score = Φ + w·E_intra (5 Å, |i-j| >= 3, MJ).
# Sparse by design: an extended class-I 9-mer makes zero to two internal contacts. w=0 = off.
tcren score -s complex.pdb -c candidates.txt -o ranked.csv --intra-weight 0.5
tcren scoring -s complex.pdb -o scores.csv --intra-weight 0.5   # reports Phi_pep_int separately

# Percentile-rank the native (or candidate) peptide's TCRen energy against a random pMHC
# background — small rank_pct = the peptide scores among the best binders.
tcren rank -s complex.pdb -o rank.csv

# Fast ΔΔG of peptide point mutations (virtual-matrix path: no atoms move, no re-docking).
# Requires --native (the peptide) and exactly one mode: --alanine-scan or --mutant.
# ddG = E(native) - E(mutant), and lower energy binds better, so POSITIVE = stabilising.
tcren ddg -s complex.pdb --native EPITOPE --alanine-scan -o ddg.csv

# Predict a combinatorial-peptide-library (CPL) response matrix from ONE template TCR:pMHC
# structure: every peptide position x all 20 residues, threaded on the template's own contact map.
# Every cell sums BOTH peptide-bearing interfaces (TCRen over TCR:peptide + Miyazawa-Jernigan over
# peptide:MHC), because the assay reads activation, which needs presentation as well as engagement.
tcren cpl -s complex.pdb -o cpl_matrix.csv
# Two reference states, both emitted, and a cell means nothing except against one of them:
#   effect_equimolar  vs the 1/20 mixture  -> the CPL background; compare against a measured matrix
#   effect_wild_type  vs the template residue -> the mutation-scan / neoantigen question
# Positive is favourable on both. Three narrower questions off the same matrix:
tcren cpl -s complex.pdb --position 5                  # every substitution at position 5, best first
tcren cpl -s complex.pdb --position 5 --mutation W     # just that one cell
tcren cpl -s complex.pdb --position 5 --to-mixture     # cost of giving position 5 up to the mixture

# Rank candidate receptors against a fixed pMHC. `assess` is the score set: is the pose real, is it
# a binder, and which part of the structure says so. Every read-out is defined for ONE structure --
# the transform, the class means and the covariance are frozen on a hold-out that ships in the wheel.
tcren features -s candidates/ -o feats.tsv
tcren assess   --features feats.tsv -o scores.tsv
tcren assess   --features cpl_feats.tsv --peptide -o cpl.tsv   # when the PEPTIDE is what varies

# The fit-free predecessor tier on the same table: Q (interface geometry), T (footprint shape) and
# their composition with the contact energy, S. Still shipped, still reported, and it COMPOSES with
# the score set rather than being replaced by it.
tcren recognize --features feats.tsv -o qts.tsv

# One TSV per structure: every interface descriptor (geometry + energies).
tcren recognize -s my_pdbs/ -o recognize.tsv          # descriptors, one row per PDB

# End-to-end candidate-epitope scoring from a structure
tcren score -s complex.pdb -c candidates.txt -o ranked.csv

# Wrong-TCR decoys: keep each ORIENTED complex's pMHC, graft on 10 other complexes' TCRs (within
# MHC class, no real pairing). Real-vs-decoy trains a label-free TCR-recognition classifier.
tcren orient -s natives/ -o oriented/          # inputs must share the canonical MHC frame
tcren shuffle -s oriented/ -o shuffled/ --n 10

# Substitute a peptide and refine its pose (knowledge-based MC scored by the DOPE atom-level
# statistical potential — independent of the TCRen/MJ scoring potentials, restrained to the input).
# Not physics relaxation — use Rosetta FlexPepDock for that.
tcren refine -s complex.pdb -o refined/ --substitute KQWLVWLFL

# Structures: any of .pdb / .cif / .pdb.gz / .cif.gz, a directory, or a .tar.gz batch
tcren contacts -s batch.tar.gz -o contacts.csv --interface tcr_peptide

# Per-residue markup: TCR (CDR/FR) + MHC groove (helix/floor) + peptide in one table.
# --regions all|tcr|mhc|peptide filters; --pseudo also marks NetMHCpan groove residues (MPS).
tcren annotate -s complex.cif.gz -o markup.csv --regions mhc --pseudo

# Superimpose structure(s) onto the canonical frame, by MHC, against the canonical database
# (data/Canonical2026, fetched at install). Detects MHC class + species and averages the
# superposition over every database structure of that class/species. Chains -> A=Vα B=Vβ
# C=peptide D=MHCα E=MHCβ/β2m. -s takes a file / directory / .tar.gz / glob; -o is a directory,
# or a single structure file (one input) whose extension must match --mmCIF/--compress; -t threads.
tcren superimpose -s complex.pdb -o oriented.pdb           # single file
tcren superimpose -s 'data/*.pdb' -o oriented/ -t 8        # glob -> directory, threaded

# Build a canonical database from native complexes (how Canonical2026 is produced). Annotation
# is one batched mmseqs call; -t threads only the structural alignment + write.
tcren orient -s data/Native2026 -o data/Canonical2026 -t 8

# Structure outputs are plain .pdb by default; add --mmCIF for .cif and --compress for .gz.
tcren superimpose -s complex.pdb -o oriented/ --mmCIF --compress   # -> oriented/<id>.cif.gz

# Fetch recent TCR-pMHC structures from RCSB -> data/pdb_recent (mmCIF .cif.gz, 5-chain validated)
tcren fetch-recent --discover --after 2024-01-01

# Build the MHC reference once (IMGT/HLA + mouse H-2; cached, not committed)
tcren build-mhc-ref

tcren info
tcren --install-completion        # shell tab-completion (bash/zsh)

tcren orient and tcren superimpose need the reference sets in data/ (Native2026, Canonical2026); setup.sh fetches them at install via tcren fetch-data (re-run it any time).

One table per structure: descriptors, energies and the score set

Two commands, two jobs. tcren features reads structures and writes descriptors; tcren recognize turns descriptors into scores. The feature pass is the expensive half, so it runs once and the scoring pass can be repeated for nothing.

tcren features  -s my_pdbs/ -o feats.tsv                   # the four default families (--all adds potts and kinetics)
tcren features  -s my_pdbs/ -o shape.tsv -i topology       # one family -- and only it is computed
tcren recognize --features feats.tsv -o scores.tsv         # Q, T, S

The 164 descriptors are catalogued in six families, four of them computed by default (potts and kinetics are opt-in), split by what each is invariant under — which is also the axis along which they carry independent evidence:

family what it is invariance
placement where the receptor sits in the groove frame — angles, TCRdock parameters, ride height, shift, offset, the CDR3 loop frames frame-dependent
interface how much contact and of what chemical kind — buried area, contact counts and types, hydrogen bonds, clashes SE(3)-invariant
topology the shape of the contact set, free of its size — coverage entropy, Hill numbers, Betti numbers, persistence entropy, canonical preference SE(3)-invariant
energetics statistical-potential interface energies Phi and their references dPhi — poly-alanine, and the smoothed background form SE(3)-invariant
potts the contact map's own energy against a Boltzmann distribution over it — neg_energy, log_z, log_lik (off unless asked) SE(3)-invariant
kinetics the interface as a spring network — stiffness, rupture, coupling residues (off unless asked)

tcren recognize -s my_pdbs/ reads the structures itself, skipping the feature file:

tcren recognize -s my_pdbs/ -o recognize.tsv               # descriptors only
tcren recognize -s my_pdbs/ -o scored.tsv --mechanics      # + the spring-network kinetics terms
what you want columns in recognize.tsv
(a) energyPhi per interface (TCRen on TCR:peptide, MJ on presentation) + references dPhi + loop parts Phi_tcr_pep, Phi_tcr_mhc, Phi_pep_mhc, dPhi_tcr_pep, dPhi_pep_mhc, Phi_cdr12, Phi_cdr3a, Phi_cdr3b, dPhi_{pep,tcr,tra,trb}_soft, varPhi_{pep,tcr}_soft
(a′) intra-peptide (--full) — the peptide's contacts with itself, which every interface sum omits Phi_pep_int, n_pep_int
(b) geometry — every docking + interface descriptor pitch, crossing, crossing_signed, dock_d, dock_torsion, dock_{tcr,mhc}_u{y,z}, extent, chain_balance, burial, n_contacts_{tp,tm}, n_pep_contacted, ct_{tp,tm}_*
(c) scores — written by tcren recognize --features, not by -s. No training set and no binding label enters any of them Q — interface quality; T — footprint shape; S — the blocks combined with the contact energy. See tcren.cohort, tcren.reliability. The two-class read-outs are tcren assess, below

Is this model worth believing? — tcren assess

A co-folding model will seat any TCR against any peptide, binding or not. assess reads the coordinates it produced and answers four separate questions about them, and every answer is defined for a single structure: the transform, the class means and the covariance are all frozen on a hold-out that ships with the package, so nothing is estimated from the rows you pass and a score does not move depending on what was scored beside it.

tcren features -s models/ -o feats.tsv                 # the expensive pass, once
tcren assess   --features feats.tsv -o scores.tsv      # arithmetic over that table

The score set — five read-outs of one frozen object. Higher is better throughout.

read-out tier what is estimated what it answers
peptide_score 0 nothing; the direction is fixed by the potential which peptide does this receptor read?
pose_score 1 a covariance over hold-out binders — no negative, no label is this the kind of interface real complexes make?
confidence_residual 1 the same covariance, read as a conditional mean is the reported confidence warranted?
binder_score 2 class means and covariances, from hold-out binder labels binder or not?
channel_scores 2 the same object, marginalized to one descriptor family which part of the structure says so?

binder_iptm is binder_score + logit(ipTM): two log-odds added, no coefficient to fit, and still defined for one structure. It is the recommended read when a confidence is available.

The five channels are named in physics and geometry terms — placement (where the receptor sits in the groove frame), interface (how much interface it makes, of what chemistry), shape (the footprint free of its size), energetics (the contact chemistry in kT), mechanics (the interface as a network of breakable springs). A marginal of a Gaussian is a sub-block of its covariance — exact, closed form, no re-fit — so attributing a score to a part of the structure costs an index and nothing else. They do not sum to binder_score and should not: the whole model also reads the correlations between channels. Sometimes a channel is the better instrument: on template-free cohorts of the 22-cohort VDJdb panel channel_shape reads 0.637 median ROC-AUC against the full posterior's 0.615.

Pass --peptide when the peptide is what varies across the structures being compared, as in a combinatorial library or a mutational scan. Otherwise the five descriptors computed without the receptor are marginalized out, because they are constant across every structure of one epitope on one allele and a model reading them reaches the cohort's name without reading an interface.

assess also emits, on the same rows: the fit-free predecessor tier S; the rank and percentile within the set with the expected mean score at a recall budget; and — when the table carries ipTM — the generator diagnostic, which band the model falls in, how often models in that band turned out to be non-binders, and what S still separates inside it. On the balanced 22-cohort VDJdb panel the top ipTM decile is 26.2% [18.7, 35.5] non-binders.

The coefficients are frozen, and the inputs they were frozen against are named. That is the contract the withdrawn cohort-refit posterior could not offer:

tcren fetch-data                                       # the structure sets the manifest names
tcren features -s <those structures> -o hold.tsv
tcren fit-holdout --features hold.tsv -o refit.npz     # matches the shipped model bit for bit

tcren.score.holdout_manifest() returns the 8,292 structures with their dataset, epitope, label and ipTM. From Python the whole set is one call:

import polars as pl
from tcren import score_table

scores = score_table(pl.read_csv("feats.tsv", separator="\t"))

(c) physics of the interaction. The koff proxies fold into the same table with --mechanics; only the mutation scan, which is per-residue rather than per-structure, needs its own command:

tcren recognize -s models/ --mechanics -t 0 -o out.tsv    # every per-structure descriptor, one table
tcren ddg       -s complex.pdb -o ddg.csv     # per-residue alanine / neoantigen ΔΔG (fast virtual matrix)

--mechanics is how to ask for the stiffness tensor, steered rupture and coupling residues on a cohort. tcren mechanics still exists and gives the same numbers, but as a second command it repeats the parse and both mmseqs searches to return a second table — CSV, keyed pdb.id rather than complex.id — that then has to be joined. Inside recognize the structures are already annotated, so the flag costs only the mechanics arithmetic (12 crystals: 19.0 s → 19.5 s, against 22.5 s for the two commands).

(Per the affinity scope caveat above, structures predict the off-rate koff via the mechanics columns, not Kd/ΔG/kon.) From Python:

from tcren.recognition import recognition_features
from tcren.reliability import s_score

feats = recognition_features("complex.pdb")    # dict of the 40 descriptors (RECOGNITION_FEATURES)
score = s_score({k: [v] for k, v in feats.items()})[0]     # one structure is enough

Library

from tcren import run_pipeline, parse_structure, import_structure, ContactMap, score_peptides
from tcren.annotation import classify_chains
from tcren.potential import tcren

# One call: annotate -> superimpose -> contacts -> per-interface energies + total
res = run_pipeline("complex.pdb")              # res.scores, res.markup, res.contacts, res.oriented
res = run_pipeline("complex.pdb", reference_aa="A")  # + delta_* : the poly-alanine ΔΦ per interface

# Oracle facade: one structure -> a bundle of ready-to-tabulate frames for the paper
# notebooks (scores, percentile rank, ΔΔG alanine scan, markup, contacts). Configurable
# per-interface potentials and TCR-region selection are forwarded to every milestone.
from tcren import summarize_structure
bundle = summarize_structure("complex.pdb", alanine=True)   # bundle["scores"], ["rank"], ["ddg"], …

# …or the individual steps:
s = parse_structure("complex.pdb.gz")          # also .cif/.cif.gz; import_structure trims the C-gene
classify_chains(s, organism="human")           # TRA/TRB via arda, peptide, MHC
cm = ContactMap.from_structure(s)              # 5 Å contacts + interface partitioning
ranked = score_peptides(cm, ["KQWLVWLFL", "RLLHPHHPL"], tcren())

# Opt-in intra-peptide term: the contacts the peptide makes with ITSELF, which every
# interface energy omits. Off by default (intra_weight=0 leaves every score untouched).
from tcren import intra_peptide_energy
from tcren.potential import mj
cm = ContactMap.from_structure(s, peptide_internal=True)
intra_peptide_energy(cm, mj())                                  # the native peptide's own energy
intra_peptide_energy(cm, mj(), peptide="KQWLVWLFL")             # a candidate on the same pose
score_peptides(cm, cands, tcren(), intra_weight=0.5, intra_potential=mj())
res = run_pipeline("complex.pdb", intra_weight=0.5)             # + scores["peptide_internal"]

Beyond the contact sum

Five things a TCR:pMHC interface does are invisible to a sum over a contact list, and each has its own instrument here: the one-body / pair split of a potential (tcren.potential), peptide backbone stability under Monte Carlo (tcren.mechanics.dynamics), discrete side-chain repacking (tcren.energetics.rotamers), footprint shape (tcren.topology.footprint), the pMHC surface a TCR meets before it binds (tcren.topology.surface) and ring-stacking geometry (tcren.stacking). What each measures, and what it was measured on, is in Beyond the contact sum.

CPL response matrices from one template structure

A positional-scanning combinatorial peptide library fixes position i to residue a and leaves every other position an equimolar 1/20 mixture, so a measured cell is an ensemble mean, R[i,a] = E[response | x_i = a]. tcren.cpl predicts that matrix from a single template complex — each of the twenty residues threaded through the template's own contact map, nothing re-docked, nothing fitted to any assay.

from tcren import (ContactMap, parse_structure, response_matrix,
                   mutation_effect, position_scan, equimolar_effect)
from tcren.annotation import classify_chains
from tcren.mhc import annotate_mhc

s = parse_structure("3HG1.pdb", pdb_id="3HG1")
classify_chains(s, organism="human")
annotate_mhc(s)                       # REQUIRED: without it peptide:MHC is empty and anchors zero out
rm = response_matrix(ContactMap.from_structure(s, cutoff=5.0))

rm.to_frame()                         # the whole matrix, one row per (position, amino acid) cell
position_scan(rm, 5)                  # every substitution at position 5, best first
mutation_effect(rm, 5, "W")           # one cell
equimolar_effect(rm, 5)               # cost of giving position 5 up to the 1/20 mixture

Every cell sums both peptide-bearing interfaces — TCRen over TCR:peptide plus Miyazawa–Jernigan over peptide:MHC — because the assay reads activation, which needs the peptide presented as well as the receptor engaged. A position the receptor never touches is an anchor; its TCR term is constant along the row, so the sum degrades to presentation alone rather than to a special case.

Two reference states, and a cell is meaningless except against one of them. A raw Φ carries a large per-position offset that says only how many contacts the position makes:

reference cell value use it for
"equimolar" (default) mean_b Φ(x_{i→b}) − Φ(x_{i→a}) comparing against a measured CPL matrix — the mixture is the assay's own background
"wild_type" Φ(x_{i→wt}) − Φ(x_{i→a}) mutation scan / neoantigen ranking off the residue the template carries

They differ by a per-position constant — how far the template's residue sits above its column mean. Positive is favourable on both, since lower energy is the better binder. Under "wild_type" the template's own cell is identically zero; under "equimolar" it is an ordinary measurement.

Batch inputs, gzip, archives

from tcren.structure import iter_structures
for pdb_id, structure in iter_structures("batch.tar.gz"):   # file | directory | .tar.gz
    classify_chains(structure, organism="human")
    ...

Canonical orientation, contacts, docking geometry

from tcren.mhc import annotate_mhc
from tcren.docking import canonicalize_structure, superimpose, docking_angles
from tcren.contacts import multi_contacts, ContactDefinition

annotate_mhc(s)
oriented, info = canonicalize_structure(s)     # frame: z=MHC→TCR, y=peptide, x=thin; chains A–E
oriented, info = superimpose(s)                # orient onto data/Canonical2026 by MHC (class+species ensemble)
layers = multi_contacts(s, ContactDefinition(d1=5, d2=8, d3=12))   # heavy-atom / Cβ / Cα
d = docking_angles(s)                          # crossing (~20–70° αβ) + incident angle

2D complementarity maps & region-pair contacts

from tcren.project2d import (project_structure, residue_markup_table, contacts_table,
                             region_pair_summary)
from tcren.viz import render_complementarity_map, view_pocket_cdr

proj = project_structure(s)                                   # canonical groove plane
svg  = render_complementarity_map(residue_markup_table(s, proj),
                                  contacts=contacts_table(s, threshold=5.0))
region_pair_summary(s, kind="closest")        # contacts per region pair + bond types (cb/ca too)
view_pocket_cdr(s).show()                      # interactive 3D pocket + CDR overlay (py3Dmol)

Publication figures

tcren.viz.pymol drives a headless PyMOL to ray-trace figure panels of oriented complexes. Three scenes cover the usual views, and every panel carries a labelled axis gizmo in its corner:

Figures need the viz extra (pip install "tcren[viz]") for Pillow, plus a pymol binary on PATH — PyMOL is a separate install, not a Python dependency.

from tcren.viz.pymol import render, overlay_scene, groove_scene, interface_scene
render(groove_scene("1ao7", "data/Canonical2026"), "groove.png")            # peptide in the cleft
render(groove_scene("1ao7", "data/Canonical2026", surface=True), "s.png")   # + molecular surface
render(overlay_scene(ids, "data/Canonical2026"), "overlay.png")             # ensemble, side-on
render(interface_scene("1ao7", "data/Canonical2026", cdr), "iface.png")     # peptide + CDR loops

A canonically-oriented structure is only interpretable if the reader can tell which way the frame points, and x/y/z does not tell them — so the arrows are named for what they mean:

axis label direction
x width groove width, across the cleft (α1↔α2)
y N→C groove axis, toward the peptide C-terminus
z TCR docking normal, MHC floor → TCR

The triad is thin, arrow-headed, and turns with the camera. An axis pointing at the viewer foreshortens to a dot and its label drops to the lower left of it, the usual convention for an axis normal to the page. These are the three directions the docking-geometry literature uses (SwiftTCR, TCR3d); only the principal-component ranking differs, because tcren.docking.frame fits the whole complex where those fit the MHC groove alone.

Colour by which residues carry the score. Φ is a sum over residue–residue contacts, so it decomposes exactly: a residue's share is the sum of φ(a_i, a_j) over the contacts it makes. The total says how large the score is; this says what it is made of.

from tcren.viz.pymol import residue_importance, importance_scene
imp = residue_importance(structure)                 # phi + n_contacts, per residue
render(importance_scene("1ao7", CANON, imp), "importance.png")                     # energy share
render(importance_scene("1ao7", CANON, imp, by="n_contacts",
                        spectrum="white_red"), "contacts.png")                     # geometric share

CDR3 and peptide residues become sticks on a ramp, everything else stays pale. Blue is favourable and red unfavourable — the ramp is centred on zero rather than fitted to the range, so those words keep their meaning even when every contact in an interface is stabilising. Each contact is attributed to both residues it joins, so the per-residue values sum to twice Φ: an attribution, not a partition.

render() is deliberately not a tcren subcommand: a figure is a handful of styling choices that want editing, not a fixed flag set. Pass any PyMOL script body as the scene.

Explore it interactively with the marimo app — pick a structure and scene, swing the camera and watch the gizmo follow, restyle it, colour by importance with the numbers beside the render, and rotate a live 3Dmol.js view with the mouse:

pip install "tcren[marimo]"
marimo run notebooks/pymol_interactive.py       # or `marimo edit` to change the code

Worked examples of every view, with images: Figure gallery.

Modules

module what it does
tcren.structure parse/write .pdb/.cif(.gz)/.tar.gz; the Atom/Residue/Chain/Structure model; iter_structures
tcren.annotation chain typing — TCR loci/CDRs via arda, peptide, MHC; αβ/γδ C-gene call
tcren.mhc map MHC chains to allele/class/role; partition the groove (helices/floor); NetMHCpan pseudosequence
tcren.contacts / contactmap closest-atom 5 Å contacts, Cα distances, multi-layer (5/8/12 Å) contact tables, interface partitioning
tcren.potential Potential (TCRen/MJ/Keskin/MJ1996 + MJ partition energies); decompose / hydrophobicity_fit — the one-body vs pair split; derive_tcren (classic/AM/LOO) with non-redundancy filtering
tcren.stacking ring-stacking geometry (centroid distance, interplanar angle, vertical/lateral offset) — the directional signal a contact potential cannot see
tcren.energetics the interface energy sum (scoring), ΔΔG on mutation (mutation), rotamer-averaged contacts (rotamers); scoring_rank percentile-ranks a peptide against a background
tcren.cpl CPL response-matrix prediction from one template complex; equimolar and wild-type references; per-position and per-cell queries
tcren.binder the pre-energy check that an interface is a plausible dock at all — a rule over contact count and docking geometry, not a model
tcren.recognition / descriptors the descriptor catalogue: 164 columns in six families, what each means, its units and its known defects (DESCRIPTORS, STATUS), plus the 40-column interface block this layer computes itself
tcren.score the score set — one frozen object, five read-outs (peptide_score, pose_score, confidence_residual, binder_score, channel_scores), each defined for a single structure
tcren.cohort / reliability the fit-free predecessor tier: Q, T, S, the AlphaFold band table and the screening cut
tcren.docking canonical frame, superimpose onto the canonical DB, docking angles, reverse-dock detection
tcren.refine peptide substitution + refinement (DOPE MC; CCD/OpenMM/ProMod3/FlexPepDock engines); register QC
tcren.clashes / mechanics steric-clash report; interface spring-network stiffness + rupture model; peptide backbone dynamics
tcren.topology footprint shape: coverage entropy / Hill numbers over the CDR-loop × target partition, canonical germline-MHC vs CDR3-peptide preference, α/β contact imbalance, and the footprint's topology (patches, holes, H₀ persistence); the pMHC surface as a height field and the gap between the two faces — no potential, no reference, orientation-free
tcren.project2d / viz project the interface onto the groove plane; SVG complementarity maps + 3D pocket/CDR views
tcren.pipeline / oracle one-call structure scoring (run_pipeline → Φ, ΔΦ per interface; summarize_structure)
tcren.paper Nat Comput Sci 2022 reproduction (HF bootstrap, batch annotation, legacy comparison)

Data

Structures live in the Hugging Face dataset isalgo/tcren_structures, all gzipped:

folder contents
Native2022 the 2022 paper set (oracle)
Native2026 the comprehensive 2026 TCR:pMHC set the current potential is derived from
Canonical2026 Native2026 re-oriented into the canonical frame (tcren orient)

tcren reads .pdb/.cif/.pdb.gz/.cif.gz and .tar.gz batches; an installed library lazily fetches the canonical reference structures from the Hub when orienting a new complex.

Where it all lives: tcren.paths.tcren_home(). That one root holds the MHC allele reference (database/mhc/, written by tcren build-mhc-ref), its mmseqs index (data/mhc_cache/) and the structure sets (data/). It resolves to $TCREN_HOME when set; otherwise to the source checkout, recognised by its pyproject.toml, so a development install uses the repo's own data/; otherwise to $XDG_CACHE_HOME/tcren (in practice ~/.cache/tcren), which an installed wheel can write and which survives an upgrade. $TCREN_DATA_DIR overrides the data/ subdirectory alone.

That data/ holds Native2026 (+ Canonical2026, gitignored, fetched on demand), PDB_date.tsv, and TCRen_potential.csv — the 2022 (karnaukhov2022) matrix, kept for reproducing published results; the current default is the bundled tcren2 (use -p karnaukhov2022 for the old one). Canonical2026's orient_metadata.json ships inside the package (src/tcren/data/), because the fetch brings down structures only and an installed library has no repo data/.

Notebooks

Runnable examples under notebooks/ (rendered in the docs):

  • complementarity_map_2d — 2D interface maps, multiple structural + map views of 1ao7
  • contact_thresholds_and_bondtypes — region-pair contact counts (closest/Cβ/Cα) + bond types
  • canonical_frame_figures — canonical-frame QC across the Native2026 set
  • pymol_canonical_figures — ray-traced PyMOL panels (overlay, groove, interface) by class/species
  • mhc_pseudosequence_mps — NetMHCpan MHC pseudosequence (MPS) residues vs. peptide contacts
  • example_gil_a02_rs_motif — GILGFVFTL/HLA-A*02 and the public CDR3β Arg–Ser motif
  • pocket_cdr_3d — 3D peptide-binding pocket with the CDR loops overlaid (py3Dmol)
  • tcren_analysis — potential heatmaps (TCRen / MJ / Keskin) and contact distributions
  • natcompsci2022/ — full reproduction of the Nat Comput Sci 2022 analyses

Two of them run the score set end to end, from fetching structures to ranking them:

  • score_vdjdb_panel — receptor ranking for a fixed epitope on the balanced 22-cohort VDJdb panel (1,089 TCRmodel2 models), reported one cohort at a time with template-covered and template-free apart, and the five channel scores per cohort
  • rank_peptides_cpl — peptide ranking for a fixed receptor on the combinatorial-peptide-library set (7 clones, 2,103 models): per-clone ROC, the graded activation read-out, and a whole response matrix predicted from one template

Four marimo apps ship alongside them (pip install 'tcren[marimo]', then marimo run <file>):

  • surface_topology.py — elevation / charge / hydropathy maps over the groove, and the featureless-vs-bulged epitope comparison against the structures the literature names
  • pymol_interactive.py — a PyMOL render explorer over the canonical scenes (overlay, groove, interface, residue importance)
  • confident_negatives.py — a generator's confidence read together with the coordinates
  • potts_contact_map.py — the predicted contact-frequency map beside the contacts a structure made

Performance

Per-stage wall time (best of n) on a TCR-pMHC complex (1ao7), Apple M-series, single thread (RUN_BENCHMARK=1 pytest -k benchmark -s to reproduce the core stages):

stage time notes
parse a gzipped structure ~17 ms .pdb.gz / .cif.gz
contact map (5 Å, cKDTree) ~9 ms per structure
score 1000 candidate peptides ~11 ms ~10 µs/peptide (vectorised)
ΔΔG alanine scan (9-mer) ~11 ms virtual-matrix; no atoms move
the score set, all six read-outs ~5 ms for one structure, ~34 µs/structure over 1,089 frozen transform + two Gaussians; no structure re-read
peptide refine (2000-step DOPE MC) ~320 ms knowledge-based rigid-body refinement
annotate (MHC map, 1 structure) ~670 ms one mmseqs2 search
annotate (TCR + MHC), batched ~0.2 s/structure one mmseqs2 call for the whole set; vs ~1.5 s/structure unbatched
superimpose onto the canonical DB (per query) ~2.8 s aligns to every same-class DB structure
peak RSS value notes
single-structure pipeline (no orient) ~200 MB parse → annotate → contacts → score → refine
+ superimpose (loads canonical DB) ~780 MB holds Canonical2026 in RAM; skip with --no-superimpose

Annotation is the only network/compute-heavy step and is always batched (one mmseqs2 search over all chains; mmseqs2 parallelises internally — never per-structure, never Python-threaded). Threads are used only for the embarrassingly-parallel, mmseqs-free stages (structural alignment, write, rendering): tcren orient -t N. Screening a peptide/TCR panel is embarrassingly parallel — references are annotated and oriented once, so the hot loop is just refine + contacts + score per complex.

Tests

pytest -m "not slow"          # unit + fast regression (the CI gate)
pytest                        # add the arda/mmseqs-backed regression tests
RUN_BENCHMARK=1 pytest -k benchmark -s

Project state

  • CHANGELOG.md — what has landed, per release, with the measurement for each.
  • STATUS.md — where the modules stand, and the known caveats.
  • ROADMAP.md — where it is going, and what each direction is waiting on.
  • BENCHMARKS.md — achieved accuracy.

Citing

TCRen is free for academic and non-commercial use. If you use it, please cite our latest Nature Computational Science 2024 paper:

Karnaukhov VK, Shcherbinin DS, Chugunov AO, Chudakov DM, Efremov RG, Zvyagin IV, Shugay M. Structure-based prediction of T cell receptor recognition of unseen epitopes using TCRen. Nat Comput Sci. 2024 Jul;4(7):510-521. doi: 10.1038/s43588-024-00653-0. Epub 2024 Jul 10. PMID: 38987378.

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Predicting TCR:pMHC binding from structural data

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