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A data-driven estimation of the ribosome drop-off rate in S. cerevisiae reveals a correlation with the genes length.

NAR Genom Bioinform · 2024
98/100 3/4
Why this verdict

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

Reproduced on the brainbox compute brainarbeit.com
How its reproducibility compares
98/100
Reproducibility score
1.4 SD above mean
vs. all fields · 1173 studies
🎯 Scores higher than 93% of all assessed papers rank 65 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

REPRODUCED (1:1) the paper's core pipeline-derived result: the global ribosome drop-off rate estimates for both GSE91068 conditions, D1 (Synthetic Defined control) and D3 (Methionine restricted), Table 2. Fresh re-run of the authors' own ribofilio tool end-to-end on the paper's own SRA data on «our HPC» SLURM («job», COMPLETED exit 0, 1:27:39): SRA prefetch/fasterq-dump -> trim_galore 0.6.7 (adapter CTGTAGGCACCATCAAT) -> bowtie2 2.3.5.1 vs Ensembl rel-98 R64-1-1 cDNA (6612 tx) -> samtools 1.11 -> bedtools bamtobed -> ribofilio.py -b 50. SRA read counts (D1FP 41,917,507; D1mRNA 51,165,369; D3FP 42,972,168; D3mRNA 34,390,503) reproduce the repo run-log exactly; D1FP alignment 32.98%. All 12 Table-2 claims match to every printed decimal EXCEPT R^2 in the 4th significant figure (D1 0.4912 vs 0.4907; D3 0.4107 vs 0.4104), consistent with minor aligner-version drift; reproduced D1 also matches the repo's shipped notebooks/D1.regression.log anchor (magnitude identical; anchor records the decay with a negative sign). Outputs bit-identical to the prior (requeued) run. NOT attempted: per-gene-length/per-GO subsets and the gene-length correlation (Table 3, the harder ~20%); other-accession datasets D2/D4/D5-D10 (out of scope).

💻 Code ↗ 🗄 Data: GSE91068

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Assessment versions

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  1. v1 current initial assessment Score 98
    assessed: 2026-06-19 ⛓ 4bbe7b92773a
✎ I am an author of this paper

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Reproduced
2026-06-22
Rubric version
not recorded
Assessed by
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

The paper tests whether ribosome drop-off (premature ribosome detachment from the mRNA before the stop codon) occurs at a quantifiable, non-negligible 'basal' rate in S. cerevisiae under standard (non-stressing) growth conditions, and whether this rate is related to gene length or to stress conditions such as nutrient restriction.

Core claims
  • Ribosome drop-off events occur at a significant rate in S. cerevisiae cultured in standard conditions finding
  • There is a correlation between ribosome drop-off rate and gene length, with longer genes showing lower drop-off rates finding
  • Ribofilio, a novel open-source Python tool embedded in a Snakemake pipeline, estimates ribosome drop-off rate from Ribo-seq data resource
  • The overall ribosome drop-off rate in S. cerevisiae is in the range of 4 × 10-4 events per codon, similar to previously reported E. coli estimates finding
  • The dynamics of ribosome drop-off in S. cerevisiae are significantly different from those in E. coli, reflecting the higher complexity of the eukaryotic genome and its expression finding
  • A basal, sequence-independent ribosome drop-off rate produces a negative exponential distribution of ribosome density along the CDS when translation initiation is the rate-limiting step mechanism
  • Ribofilio estimates the per-bin drop-off rate rb via weighted linear regression of log(Y) on bin position X, then converts it to a per-codon rate rc = 1-(1-rb)^(1/lc) method
  • Ribofilio achieved 81% module test coverage during software validation resource
Experimental setups
Assay System Perturbation Readout Platform
Ribosome profiling (Ribo-seq) S. cerevisiae, log-phase growth, rich (YEPD) medium (datasets D5, D6) none (standard/reference condition) 3' end read counts per transcript bin, used to estimate ribosome drop-off rate
Ribosome profiling (Ribo-seq) S. cerevisiae (datasets D7, D8) amino acid starvation 3' end read counts per transcript bin, ribosome drop-off rate
Ribosome profiling (Ribo-seq) S. cerevisiae (datasets D1/D3) methionine restriction vs Synthetic Defined (SD) control 3' end read counts per transcript bin, ribosome drop-off rate
Ribosome profiling (Ribo-seq) S. cerevisiae (datasets D2/D4) glucose restriction (caloric restriction) vs SD control 3' end read counts per transcript bin, ribosome drop-off rate
RNA-seq S. cerevisiae, matched to each Ribo-seq sample matched to corresponding Ribo-seq condition mRNA read counts per bin, used to normalise Ribo-seq counts
Read trimming and quality filtering Ribo-seq/RNA-seq reads from S. cerevisiae none adapter-trimmed reads ≥20 nucleotides trim_galore v0.6.7
Read alignment to transcriptome and BED conversion S. cerevisiae reads vs Saccharomyces_cerevisiae.R64-1-1.cdna.all.fa none mapped reads, primary alignment percentage, BED-format alignments Bowtie2 v2.4.5; bedtools bamtobed
Key results
  • Drop-off rate per codon (rc) across the eight datasets is broadly in the range of 4 × 10-4 events per codon rc 0.0003-0.0006
  • Longer genes show lower ribosome drop-off rate
  • t-test rejects the null hypothesis rb=0 for all eight datasets p<0.00001 (D1-D4); p=0.0038/0.0029/0.0016/0.0020 (D5-D8)
  • Goodness-of-fit (R2) of the exponential decay model of ribosome density varies substantially across datasets R2 0.0341-0.6887
  • Ribofilio pipeline achieved 81% test coverage in module testing 81%
Key statistics
  • other rc range 0.0003-0.0006 events per codon (drop-off rate per codon across datasets D1-D8 (Table 2))
  • pvalue <0.00001 (t-test for rb=0, datasets D1-D4)
  • pvalue 0.0038 (D5), 0.0029 (D6), 0.0016 (D7), 0.0020 (D8) (t-test for rb=0, datasets D5-D8 (rich medium / starvation))
  • other R2 = 0.4907 (D1), 0.6887 (D2), 0.4104 (D3), 0.4993 (D4), 0.0341 (D5), 0.0462 (D6), 0.5336 (D7), 0.4188 (D8) (coefficient of determination for exponential fit of ribosome density per dataset)
  • other rb per bin: D1=0.0051±0.0003, D5=0.0057±0.0003, D6=0.0053±0.0003 (drop-off rate per bin (bin size=50) with SE, Table 2)
  • other 4 × 10-4 events per codon (previously reported in vivo E. coli drop-off rate (ref. 22), used as comparison)
  • count 81% (Ribofilio module test coverage)
  • other bin size = 50 nucleotides (Ribofilio parameter used to compute Table 2 drop-off rates)

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 paper introduces Ribofilio, a Python pipeline that estimates the ribosome drop-off rate per bin (r_b) from Ribo-seq data by fitting a weighted log-linear regression to binned, RNA-seq-normalised ribosome density profiles across eight publicly available S. cerevisiae datasets. Statistical significance of each estimated r_b is assessed by a t-test against the null hypothesis r_b = 0, with results reported per dataset as point estimates, standard errors, 95% confidence intervals, RMSE, and R². The authors further describe a qualitative correlation between gene length and the estimated drop-off rate.

Replicationunclear Sample sizeEight publicly available Ribo-seq/RNA-seq dataset pairs from three GEO series; within-condition replicates exist in the source data (e.g. D5/D6 are two rich-medium replicates; D7/D8 are two starvation replicates); biological sample sizes of the original experiments are not re-described GroupsStandard growth conditions (D1, D2, D5, D6) vs. nutrient-stress conditions (methionine restriction D3, glucose restriction D4, amino acid starvation D7/D8); comparisons are descriptive, not formally tested Pairingna Randomization/blindingna DispersionCI Exact p-valuesyes Confidence intervalsyes Multiplicity correctionnone stated
Statistical tests used
Test Applied to n Assumptions
Weighted linear regression (log-linear model): log(Y) regressed on bin number X, slope estimated as r_b Estimation of drop-off rate per bin for each of eight Ribo-seq datasets (D1–D8), Table 2 N bins per dataset (bin size fixed at 50 nt; exact N per dataset not stated in text) not stated
t-test (one-sample, null hypothesis r_b = 0) derived from regression slope SE with N−2 degrees of freedom Statistical significance of estimated drop-off rate per dataset, Table 2 p-value column N−2 degrees of freedom; N = number of bins per dataset (not given explicitly) not stated
Approaches that could also have been used
  • The exponential model is fitted by log-transforming Y and applying weighted linear regression to log(Y) vs bin number X
    Could also: Nonlinear least squares (NLS) fitting of the untransformed exponential model Y = A·exp(−r_b·X) directly on the original scale — Log-linearisation changes the error structure: residuals that are homoscedastic on the log scale become heteroscedastic on the original scale, and vice versa; NLS on the original scale allows weights that more directly reflect measurement uncertainty in ribosome read counts, and is a standard alternative when the functional form is known
  • 95% confidence intervals for r_b are derived analytically from the regression slope SE and a t critical value (Equations 2–5), assuming normally distributed residuals
    Could also: Non-parametric bootstrap confidence intervals for r_b obtained by resampling bins or gene subsets — Bootstrap CIs make no distributional assumption about residuals; they can better capture uncertainty when residuals are skewed or when N (number of bins) is small, and serve as a useful robustness check alongside analytic CIs
  • Eight independent t-tests are performed (one per dataset) with no adjustment for the family of comparisons
    Could also: Benjamini-Hochberg FDR correction applied across all eight p-values — When multiple hypothesis tests are conducted on related datasets, FDR correction controls the expected proportion of false discoveries; it is a standard step when interpreting a set of related tests jointly and would not change any of the very small p-values reported here but would make the multiple-testing stance explicit
  • The correlation between gene length and drop-off rate is reported as a qualitative descriptive finding without a stated formal test or effect-size measure
    Could also: Spearman rank correlation coefficient (ρ) with a permutation-based p-value, or Pearson correlation on log-transformed gene lengths, reported with a 95% CI — A formal correlation coefficient with confidence interval and p-value would quantify the strength and precision of the length–drop-off relationship; Spearman rank correlation is robust to the non-normality and heavy right tail typical of gene-length distributions
  • Model adequacy is assessed descriptively using R² and RMSE of the weighted log-linear regression
    Could also: Formal information-criterion comparison (AIC or BIC) between the exponential decay model and nested alternatives such as a constant (no drop-off) or linear model — AIC/BIC penalise model complexity and provide a principled, quantitative basis for concluding that the exponential model captures the data better than simpler alternatives, complementing the descriptive R² value
  • A single fixed bin size of 50 nt is used for all reported analyses (Table 2 header)
    Could also: Sensitivity analysis across a range of bin sizes (e.g. 25, 50, 75, 100 nt) with r_b estimates compared across choices — Binning granularity affects the number of data points N in the regression and the smoothing of read-count variability; reporting how r_b estimates vary with bin size demonstrates robustness of the key parameter to this methodological choice
Software: Ribofilio (Python) · Snakemake · trim_galore 0.6.7 · Bowtie2 2.4.5 · bedtools (bamtobed)

What was reproduced

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

Scope — PMID 38638702 (ribofilio, ribosome drop-off rate in S. cerevisiae)

  • Paper: Awad et al., NAR Genom Bioinform 2024; DOI 10.1093/nargab/lqae036
  • Code: https://github.com/SherineAwad/ribofilio (BSD-3, authors' own tool)
  • Data: GEO GSE91068 (Zou et al.) — the RU's assigned dataset.

The tool & pipeline (from README + pipeline/Snakefile)

ribofilio estimates the ribosome drop-off rate from Ribo-seq footprints. It bins each transcript into l-nt bins, counts footprint 3'-ends per bin (optionally normalized by RNA-seq), and regresses log(Y) on bin number by weighted linear regression: Y(X)=A·e^(−r·X). Output regression.log columns: Dropoff (per bin), Dropoff per codon, RMSE, Rsquare, SE, Margin Error, tscore, pvalue, No.of Bins.

Full Snakemake pipeline per dataset:

  1. trim_galore --gzip --retain_unpaired --trim1 -a CTGTAGGCACCATCAAT (adapter from config_D1D3.yaml)
  2. bowtie2-build yeast.fa yeast ; bowtie2 -x yeast -U trimmed.fq.gz -S out.sam -p4 (yeast.fa = Ensembl release-98 S. cerevisiae R64-1-1 cdna.all)
  3. samtools view -Sb ; bedtools bamtobed → BED
  4. python src/ribofilio.py -t yeast.fa -f FP.bed -r mRNA.bed -b 50 -v 1 -o PREFIX Pinned tool versions (pipeline/env/*.yaml): bowtie2=2.3.5.1, samtools=1.11, trim_galore (paper text says trim_galore 0.6.7, bowtie2 2.4.5 — minor version drift noted).

GSE91068 sample → dataset mapping (from data/Makefile)

  • D1 (Synthetic Defined, control): FP = SRR5090936, mRNA = SRR5090934
  • D3 (Methionine restricted, treatment): FP = SRR5090937, mRNA = SRR5090935

IN SCOPE (pipeline-derived, attempted)

  • D1 global drop-off rate — primary anchor. The repo ships the exact expected output notebooks/D1.regression.log, which equals paper Table 2 D1 (r_c = 0.0003/codon, R² = 0.4907, RMSE = 0.0143, 95% margin 0.0011, 295 bins).
  • D3 global drop-off rate — same GSE91068, treatment condition (secondary).

OUT OF SCOPE (not attempted, with reason)

  • D2/D4 (GSE134152), D5–D10 (GSE13750): other accessions, not this RU's dataset.
  • Per-gene-length and per-GO subset drop-off rates (Table 3, gene-length correlation): many sub-analyses; deferred as the hard last ~20% (80/20 rule). The global D1/D3 values exercise the same core estimator end-to-end.
  • Wet-lab / external biological interpretation: out of scope by definition.

Reproduction strategy

Run the authors' own Snakemake pipeline steps (faithful third-party-of-record = authors' tool on the paper's data) for D1 and D3 on «our HPC» SLURM, compare the produced regression.log to the shipped notebooks/D1.regression.log (D1, exact target) and to paper Table 2 / reported D3 value.

Figures / tables: Table
D1_dropoff_codon
Reported
0.0003 per codon (Table 2 D1; shipped notebooks/D1.regression.log)
Reproduced
0.0003
exact
D1_dropoff_bin
Reported
-0.0051 (shipped log; sign=decay)
Reproduced
0.0051
exact
D1_rmse
Reported
0.0143 (Table 2 D1)
Reproduced
0.0143
exact
D1_rsquare
Reported
0.4907 (Table 2 D1)
Reproduced
0.4912
within tolerance
D1_se
Reported
0.0006 (Table 2 D1)
Reproduced
0.0006
exact
D1_margin
Reported
0.0011 (Table 2 D1)
Reproduced
0.0011
exact
D1_nbins
Reported
295 (shipped log)
Reproduced
295
exact
D3_dropoff_codon
Reported
0.0006 per codon (Table 2 D3)
Reproduced
0.0006
exact
D3_rmse
Reported
0.0762 (Table 2 D3)
Reproduced
0.0762
exact
D3_rsquare
Reported
0.4104 (Table 2 D3)
Reproduced
0.4107
within tolerance
D3_se
Reported
0.0011 (Table 2 D3)
Reproduced
0.0011
exact
D3_margin
Reported
0.0021 (Table 2 D3)
Reproduced
0.0021
exact

Assessments & scoring basis

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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.

880.2 k
tokens (I/O) · 55.3 M incl. cache
470 min
runtime · 1.36 CPU-h
1.9 GB
peak RAM
1
HPC jobs
hummel
machine