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Recombination Facilitates Adaptive Evolution in Rhizobial Soil Bacteria.

Mol Biol Evol · 2021
L1 80/100 3/4
Why this verdict

The main results reproduced: recomputed values matched the published ones within tolerance.

Reproduced on the brainbox compute brainarbeit.com
Scoring basis — itemised

Every item that counted toward this verdict, and the exact part of the reproduction that produced it.

Supporting (toward a concern)
Content-critical question only partially held
+2 pts
From: Q8 · Severity of the miss (overall human judgment) 🟡
Minor / cosmetic deviation
+1 pts
From: Q3 · Location of the main deviation 🟡
Minor / cosmetic deviation
+1 pts
From: Q4 · Cause of the deviation 🟡
Minor / cosmetic deviation
+1 pts
From: Q6 · Severity of the deviation 🟡
Concordant (toward reproduced)
Code + data deposited & functional
-2 pts
From: Data & code availability Available & functional
Total score +3
✓ What held up
  • Same input data as the authors
  • Reported values were directly comparable
  • Reported values are derivable from the shared data
  • The central claim held under reproduction
What did not (or only partly)
  • 🟡A deviation arose in the data or preprocessing
  • 🟡A deviation was attributed to the published material
  • 🟡The deviation was non-trivial in magnitude
  • 🟡Overall, the reproduction showed a material discrepancy
How its reproducibility compares
80/100
Reproducibility score
0.3 SD above mean
vs. all fields · 1173 studies
🎯 Scores higher than 56% of all assessed papers rank 484 of 1173 scored

A 0–100 reproducibility-quality score from the per-question grades, shown as a z-score: standard deviations above (+) or below (−) the mean of comparable assessments.

Reproduction agent’s raw note

Described well enough to reproduce. Mostly 1:1. Tier-1: 4 pan-genome COUNT claims (196 strains, 22,115 ortholog groups, 4,204 core, 17,911 accessory) reproduce EXACTLY by counting the authors' shipped Proteinortho tables; core=4,204 reconfirmed from a 2nd independent product (figshare SNP matrices). Tier-2 (figshare alignments): concatenated core alignment length reproduced to within 0.02% (3,091,752 vs 3,091,179 bp); per-genospecies nucleotide diversity pi reproduced within tolerance with exact rank order and the headline 4.5x range (got 4.46x), per-gene pi matching the paper's own Table S6 at Pearson r=0.994. PARTIAL on the 3,086 filtered gene count (got 3,298) and 334,040 variable sites (got 482,240) because criterion (4) 'introgression score <=3' uses Syntenizer3000 output NOT shipped in the repo; the missing filter preferentially removes high-SNP recombinant genes, consistent with the paper's thesis. NOT attempted (hard 20%): full SRA->SPAdes->prokka annotation (1,468,264 ORFs), Proteinortho re-run, ClonalFrameML R/theta, RAxML phylogeny, McDonald-Kreitman alpha (0.07-0.39), LD r2 medians. No fabrication detected; all reproduced values are derivable from the deposited data. Compute was light (counting + popgen on shipped/figshare intermediates) so no «our HPC» SLURM job was required.

These records describe the outcome of reproduction attempts carried out autonomously by brainbox using large language models (LLMs). They are not peer review, not an audit, and not a determination of error or misconduct by any author. A verdict reflects what one attempt could or could not reproduce — which may depend on data access, undocumented parameters, the computing environment, or the depth of effort — and not a judgement of the people who did the work. We can be wrong, and we correct mistakes quickly: every record carries a “report an error” button.

Assessment versions

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  1. v1 current initial assessment Score 80
    assessed: 2026-06-14 ⛓ e172faada008
✎ I am an author of this paper

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Provenance — full disclosure

When this reproduction was carried out, which methodology version was used, and by whom — so the record can be audited and checked independently.

Reproduced
2026-06-14
Rubric version
v1.0
Assessed by
🤖 AI curator · claude (ai-curator room) · v1.0 · run #1 2026-06-15
no human curator yet
Last updated
2026-08-05

Provisional, curator- or AI-assessed, and independently checkable. A reproduction outcome states what one attempt could reproduce — not a judgement of the authors.

Deep full-text extraction

Model: sonnet
Founding hypothesis

Homologous recombination increases the efficacy of natural selection by decoupling the fate of beneficial and deleterious mutations, and thus the proportion of amino acid substitutions fixed by adaptive evolution (α) should be positively correlated with recombination rate in bacteria (Rhizobium species).

Core claims
  • α varies from 0.07 to 0.39 across five Rhizobium species and is positively correlated with the level of recombination finding
  • The positive effect of recombination on α is driven both by a higher rate of adaptive evolution (ωa) and a lower rate of nonadaptive evolution (ωna) mechanism
  • Species/gene classes with higher recombination (lower r2) show stronger purifying selection, reflected in lower piN/piS ratios finding
  • GRAPES with the GammaZero DFE model was used to estimate the distribution of fitness effects and α from folded SFS and divergence data method
  • Genes were split into three equally sized intragenic LD (r2)-based recombination classes per species to test association with α method
  • Alternative recombination measures (R/θ and D') largely reproduce the r2-based trend between recombination and α finding
  • α does not differ significantly across genomic compartments (chromosome, chromids Rh01/Rh02, plasmid Rh03), likely due to low statistical power finding
  • A curated set of 3,086 core protein-coding genes from 196 genomes of 5 closely related Rhizobium species, filtered for interspecies HGT and abnormal diversity, is provided as a resource for population genetics analysis resource
Experimental setups
Assay System Perturbation Readout Platform
Site frequency spectrum (SFS) / divergence-based DFE and alpha estimation (GRAPES, GammaZero model) 196 Rhizobium leguminosarum species-complex genomes (5 species: gsA-gsE) none (natural population comparison) α, ωa, ωna estimates with confidence intervals GRAPES
Intragenic linkage disequilibrium (LD) decay analysis (r2) 3,086 core genes across 196 genomes, 5 species none r2 decay with genomic distance, median r2 per species/gene, recombination rate proxy
Nucleotide diversity (π) estimation 3,086 core genes across genomic compartments (chromosome, chromids Rh01/Rh02, plasmid Rh03) in 5 species none π (~2Neμ) per gene/compartment/species
piN/piS ratio calculation from synonymous/nonsynonymous SFS same core gene set, 5 species none strength of purifying selection
Alternative recombination rate estimation (R/θ and D') same core gene set, 5 species none correlation with r2 and with α
Permutation test (200 iterations) across recombination classes same core gene set, 5 species none statistical significance of α difference between extreme recombination classes
Orthologous gene group identification / core genome definition 196 Rhizobium genomes, 22,115 orthologous gene groups none identification of 4,204 core genes present in all strains (3,086 retained after HGT/diversity filtering)
Key results
  • α across species ranges from 0.07 to 0.39, increasing with recombination level 0.07-0.39
  • Most recombining species (gsC) has the highest α across all outgroup comparisons; least recombining (gsD) has the lowest α in 3 of 4 comparisons gsC alpha up to 0.36; gsD as low as 0.10
  • piN/piS ranks species similarly to recombination rate: more recombining species show lower piN/piS (stronger purifying selection) gsC=0.037, gsA=0.039, gsE=0.051, gsB=0.057, gsD=0.07
  • ωa increases and ωna decreases with recombination rate across most species comparisons, with roughly equal magnitude effects
  • Nucleotide diversity (π) differs up to 4.5-fold across species 4.5-fold (gsA 0.018 vs gsB 0.0045)
  • Median r2 (inverse recombination measure) ranks species: gsC < gsA < gsE < gsB < gsD gsC=0.248, gsA=0.341, gsE=0.561, gsB=0.651, gsD=1.00
  • Permutation tests confirm significant α differences between extreme recombination classes for all species except gsD P<=0.05
  • α differences across genomic compartments (chromosome vs chromids vs plasmid) are not statistically significant
Key statistics
  • other α range 0.07-0.39 (range of α across 5 Rhizobium species)
  • other piN/piS: gsC=0.037, gsA=0.039, gsE=0.051, gsB=0.057, gsD=0.07 (purifying selection strength by species, ranked by recombination)
  • mean π: gsA=0.018, gsB=0.0045, gsC=0.0140, gsD=0.00512, gsE=0.008 (average nucleotide diversity by species)
  • correlation Pearson's correlation between r2 and R/θ or D' ranged from 0.19 to 0.70 (agreement between recombination measures)
  • other median r2: gsC=0.248, gsA=0.341, gsE=0.561, gsB=0.651, gsD=1.00 (LD-based recombination ranking of species)
  • other α(gsC as focal vs gsA outgroup)=0.36 [0.33-0.39] (example pairwise α estimate with 95% CI from table 1)
  • count 3,086 core genes, 3,091,179 bp alignment length, 334,040 variable sites (final filtered data set used for analysis across 196 genomes)
  • count 22,115 orthologous gene groups; 4,204 core genes present in all strains (orthology results prior to HGT/diversity filtering)

Statistical methods review

Model: sonnet

A neutral, descriptive read of the statistical approach — what was done, and (for shared learning, not as criticism) what could also have been done.

The study estimates the proportion of amino acid substitutions fixed by adaptive evolution (α) and its components (ωa, ωna) across five Rhizobium species using polymorphic site frequency spectra (folded SFS) and pairwise divergence data, fitted by maximum likelihood under a GammaZero distribution of fitness effects (DFE) model selected via AIC in the GRAPES framework. Genes are partitioned into three equally-sized recombination classes per species based on mean intragenic r², and α is compared across classes within each of 20 mirror species-pair combinations. Significance of class differences is assessed by nonoverlapping 95% confidence intervals and by gene-label permutation tests (200 permutations), with all results reported as point estimates with bracketed CIs.

Replicationbiological Sample size196 total strains across 5 species (gsA=32, gsB=32, gsC=112, gsD=5, gsE=11); 3,086 core genes retained after filtering for HGT and high nucleotide diversity; no formal power analysis stated GroupsFive Rhizobium species compared pairwise; genes partitioned into three equally-sized recombination classes per species based on mean intragenic r²; secondary comparison across genomic compartments (chromosome, chromids Rh01/Rh02, plasmid Rh03) Pairingpaired Randomization/blindingnot stated DispersionCI Exact p-valuesno Effect sizesyes Confidence intervalsyes Multiplicity correctionnone stated
Statistical tests used
Test Applied to n Assumptions
Maximum-likelihood DFE estimation and α/ωa/ωna estimation via GRAPES (GammaZero model fit to folded SFS plus divergence counts) All 20 pairwise species combinations and three recombination classes within each pair; also by genomic compartment (table 2) 3,086 core genes; per-species strain counts gsA=32, gsB=32, gsC=112, gsD=5, gsE=11 not stated
AIC model selection across candidate DFE models Selection of best-fitting DFE model (GammaZero) from multiple distribution families na
Gene-label permutation test (200 permutations across recombination classes) Significance of α differences between the two most extreme recombination classes for each species pair (Supplementary fig. S7) Species-dependent gene count with ≥10 informative sites (gsA=2,453, gsB=770, gsC=2,537, gsD=257, gsE=1,161) not stated
Nonoverlapping 95% confidence intervals as significance criterion Pairwise comparisons of α across recombination classes (fig. 2) and between mirror species (table 1) not stated
Pearson's correlation Pairwise correlations among three recombination measures (r², R/θ, D′) across genes (Supplementary fig. S8) not stated
πN/πS ratio Ranking of overall purifying selection strength by species recombination level 3,086 core genes per species not stated
Approaches that could also have been used
  • Significance of α differences between recombination classes was assessed using nonoverlapping 95% CIs as a criterion rather than a formal test
    Could also: A likelihood-ratio test comparing GRAPES models fit jointly versus separately to the two extreme recombination classes, or a bootstrap-based z-test of the difference in α estimates, could also be used — Nonoverlapping CIs are a conservative heuristic that can miss true differences when intervals nearly overlap; a formal test yields an explicit p-value and is more sensitive, which is especially relevant for the low-power gsD species (n=5 strains)
  • Genes were discretized into three equally-sized recombination classes based on mean intragenic r²
    Could also: A continuous regression of α (or ωa/ωna) on mean gene-level r² — for example via weighted regression treating CI width as a precision weight — could also quantify the recombination–adaptation relationship — Discretization can mask within-class heterogeneity and the functional form of the relationship; a continuous approach would estimate an effect size per unit change in recombination and allow model comparison (linear vs. nonlinear), adding complementary information to the class-based results
  • The folded SFS was used because high shared polymorphism made ancestral-state polarization unreliable
    Could also: If a well-diverged outgroup with minimal shared polymorphism were available, the unfolded (derived-allele) SFS with a polarized McDonald–Kreitman framework could also be applied — The unfolded SFS retains directionality information that can improve DFE estimation; the authors' motivation for using the folded SFS is explicitly stated and well-grounded given the shared-polymorphism context of a recent species complex
  • No multiple-testing correction was applied across 20 pairwise species comparisons and the multiple recombination class contrasts within each
    Could also: A Benjamini-Hochberg FDR correction across all permutation p-values, or Bonferroni correction for the family of contrasts, could also be applied — With 20 species pairs and at least one contrast per pair, the familywise type-I error rate is elevated; an FDR procedure would allow readers to distinguish findings that survive correction from those that do not, without substantially reducing power compared with Bonferroni
  • The single best-fitting DFE model (GammaZero) selected by AIC was used for all α estimation
    Could also: AIC-weighted model averaging over all candidate DFE models could also be used to propagate model-selection uncertainty into the α confidence intervals — When competing models have similar AIC scores, selecting a single model underestimates uncertainty; model averaging produces CIs that reflect both sampling variance and DFE-shape uncertainty, which is particularly relevant when comparing closely spaced α estimates across recombination classes
  • r² (mean intragenic linkage disequilibrium) was used as the primary recombination proxy, with R/θ and D′ evaluated as robustness checks
    Could also: A population-scaled recombination rate ρ = 4Ner estimated by an LD-based coalescent method (e.g., LDhat, FastEPRR) could also be used as the primary recombination measure — Mean r² is jointly determined by recombination rate and Ne, whereas ρ partially separates these; using ρ as the primary measure — or as a co-variate alongside Ne — would help distinguish whether the recombination effect on α is independent of effective population size differences among species
Software: GRAPES · R loess (local polynomial regression for LD decay curve fitting) · ClonalFrameML (implied by Didelot and Wilson 2015 citation for R/θ estimation)

Result convergence & founder nodes

Findings this paper shares with others that ran a comparable experiment. A node’s strength is how many independent papers report it (replication breadth) — not how often it is cited, so a heavily-replicated but under-cited founder still stands out.

Citation network

Where this publication sits in the reproducibility-weighted citation graph — what it is built on, and what is built on it. Citation data from OpenAlex.

Citations
18
Impact: medium
Foundation confidence
None of its references are in our reproducibility record yet — its foundation cannot be assessed.
Topics

No assessed neighbours yet — the network grows as more papers are assessed.

Data lineage

The datasets this paper uses (text-mined from the full text via Europe PMC), and which other assessed papers stand on the same data. A shared dataset is a factual link — not a judgement.

10.6084/m9.figshare.11568894.v5 DOI in Article (http://semanticscience.org/resource/SIO_001029)
also used by 1 paper:
PRJNA510726 BioProject in Article (http://semanticscience.org/resource/SIO_001029)
also used by 1 paper:
SAMN10617942 BioSamples in Article (http://semanticscience.org/resource/SIO_001029)
also used by 1 paper:
SAMN10618137 BioSamples in Article (http://semanticscience.org/resource/SIO_001029)
also used by 1 paper:

What was reproduced

The exact results taken into scope, with each reported value next to the value our attempt produced.

Scope — PMID 34410427 (Cavassim et al. 2021, MBE)

Title: Recombination Facilitates Adaptive Evolution in Rhizobial Soil Bacteria DOI: 10.1093/molbev/msab247 · PMCID: PMC8662638 Code: https://github.com/izabelcavassim/Rhizobium_analysis (third-party-style group repo; per P16 this is valid) Data: SRA PRJNA510726 (196 isolates) · figshare 10.6084/m9.figshare.11568894.v5 (intermediates)

Dataset

196 Rhizobium leguminosarum isolates (white-clover root nodules, DK/FR/UK), 5 genospecies (gsA 32, gsB 32, gsC 112, gsD 5, gsE 11).

Pipeline (as described in Methods)

SPAdes v3.6.2 assembly → Jigome → prokka v1.12 annotation → Proteinortho v5.16b (synteny) orthology → codon-aware clustalo v1.2.0 alignment → custom SNP calling → population genetics (π, piN/piS, SFS) → LD / recombination (ClonalFrameML) → RAxML-ng phylogeny.

In scope vs out of scope

# Reported result Pipeline Scope Tier How reproduced
C1 196 strains IN 1 count strains_order.csv
C2 22,115 orthologous groups Proteinortho IN 1 core+accessory shipped tables
C3 4,204 core genes (all 196) Proteinortho IN 1 count GC_core_genes_test.csv, all n=196
C4 17,911 accessory genes (≥2) Proteinortho IN 1 count GC_accessory_genes_test.csv
C5 5 genospecies / pop structure ANI core genes IN 1 cluster shipped 196×196 ANI matrix (qualitative)
C6 assembly ~7.65 Mb, GC 60.75% SPAdes/QUAST IN 1 summary_quast_assemblies.csv (paper does not print these → context only)
C7 3,086 filtered genes, align length 3,091,179 bp clustalo+filter IN 2 concat figshare Group_alns (needs filtering criteria)
C8 334,040 variable sites SNP calling IN 2 count polymorphic columns in concatenated alignment
C9 π per gs (gsA .018, gsB .0045, gsC .014, gsD .00512, gsE .008) popgen IN 2 per-genospecies π from SNP matrices (needs gs labels)
1,468,264 protein-coding seqs prokka IN but NOT attempted 3 needs full annotation of 196 assemblies (hard 80%)
r²/LD medians, α (0.07–0.39), ClonalFrameML R/θ, RAxML tree recomb. NOT attempted 3 heavy; deferred (hard 80%)

Out of scope (not attempted, by 80/20): full SRA→assembly→annotation (SPAdes/prokka on 196 genomes), Proteinortho re-run, ClonalFrameML, RAxML phylogeny, McDonald–Kreitman α estimation. These are the expensive last ~20%.

Method note

Tier-1 reproduces the paper's count claims directly from the authors' shipped pipeline-output tables — a direct audit that the published integers match the deposited intermediates. Tier-2 (popgen on figshare alignments) recomputes the flagship sequence-level numbers and is attempted opportunistically; π per genospecies additionally needs the strain→genospecies map.

Figures / tables: Fig.1
C1
Reported
196 strains
Reproduced
196
exact
C2
Reported
22,115 orthologous groups
Reproduced
22115
exact
C3
Reported
4,204 core genes
Reproduced
4204
exact
C4
Reported
17,911 accessory genes
Reproduced
17911
exact
C5
Reported
5 genospecies / population structure
Reproduced
qualitative ANI blocks, not clean 5-way
partial
C7a
Reported
alignment length 3,091,179 bp
Reproduced
3091752 bp
within tolerance
C7b
Reported
3,086 filtered genes
Reproduced
3298
partial
C8
Reported
334,040 variable sites
Reproduced
482240 (pre-introgression-filter)
partial
C9
Reported
pi per gs A0.018 B0.0045 C0.014 D0.00512 E0.008
Reproduced
A0.0183 B0.0050 C0.0158 D0.0051 E0.0084
within tolerance

Assessments & scoring basis

Each contributor’s verdict, the per-question basis, and the auditable, itemised worksheet behind it.

🤖 AI curator · claude (ai-curator room) · v1.0 L1 80/100

An automated assessment. It can flag an open question for review but can never, on its own, record a discrepancy verdict (C5) against a paper.

🟢1. Data identity
🟢2. Endpoint comparability
🟡3. Location of the main deviation
🟡4. Cause of the deviation
🟢5. Derivability / plausibility
🟡6. Severity of the deviation
🟢7. Core claim
🟡8. Severity of the miss (overall human judgment)
Scoring basis — itemised

Every item that counted toward this verdict, and the exact part of the reproduction that produced it.

Supporting (toward a concern)
Content-critical question only partially held
+2 pts
From: Q8 · Severity of the miss (overall human judgment) 🟡
Minor / cosmetic deviation
+1 pts
From: Q3 · Location of the main deviation 🟡
Minor / cosmetic deviation
+1 pts
From: Q4 · Cause of the deviation 🟡
Minor / cosmetic deviation
+1 pts
From: Q6 · Severity of the deviation 🟡
Concordant (toward reproduced)
Code + data deposited & functional
-2 pts
From: Data & code availability Available & functional
Total score +3

This is a strong reproduction: four headline pan-genome counts match exactly from the authors' own deposited tables (core=4,204 confirmed twice independently), and the population-genetics layer (alignment length 0.02%, per-genospecies π within tolerance with exact rank order and 4.46×≈4.5× range, per-gene π Pearson r=0.994 vs Table S6) is faithful. The only real deviations — C7b (3,298 vs 3,086 genes) and C8 (482,240 vs 334,040 SNPs) — sit on the input/preprocessing side and stem from the authors not depositing the Syntenizer3000 introgression-filter intermediate, so the unfiltered set over-counts exactly the high-SNP recombinant genes the paper's thesis predicts. No fabrication is indicated; the discrepancies are explainable and directionally consistent rather than substantive, so overall is yellow (solid with explainable deviations) with no q5/q7 red.

🤝
Reproduced automatically — and fairly

Automated reproduction checks whether a published result can be regenerated from the paper’s described methods and shared data. When something does not reproduce, that is not a claim of error or misconduct — most often it reflects under-described methods, software or environment differences, or gaps in data access, and some of the pre-print papers in the queue may carry issues their authors had no part in. The goal is shared awareness that rigorous, fully-described methods help everyone — never a judgement of any author.

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Reproduction footprint

claude-opus-4-8

Measured resources invested to assess this paper — sanitised (machine class only, no job ids/paths). Compute = HPC accounting (SLURM); tokens = the AI agent's session.

240.2 k
tokens (I/O) · 16.4 M incl. cache
28 min
runtime
Per-job HPC accounting not captured for this run — the runtime shown is the reproduction’s measured wall-clock time.