# No Generation without Representation: Efficient Causal Protein Language Models Enable Zero-Shot Fitness Estimation

arXiv: https://arxiv.org/abs/2602.01845 · HTML: https://mlsanity.com/paper/2602.01845/

## Key facts

- No citing papers in the map yet (data through 14 Oct 2026).
- Builds on 25 of its 82 references that are in the map.
- First posted on arXiv 2 Feb 2026.

## Builds on (most cited first, 25 of 25)

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- [Understanding protein function with a multimodal retrieval-augmented foundation model](https://mlsanity.com/paper/2508.04724/) (2025)
- [ProtSAE: Disentangling and Interpreting Protein Language Models via Semantically-Guided Sparse Autoencoders](https://mlsanity.com/paper/2509.05309/) (2025)

## Abstract (arXiv)

Protein language models (PLMs) face a fundamental divide: masked language models (MLMs) excel at fitness prediction while causal models enable generation, forcing practitioners to maintain separate architectures. We introduce Proust, a 309M-parameter causal PLM that bridges this gap through architectural innovations adapted from recent LLM research, including grouped-query attention with shared K/V projections, cross-layer value residuals, and depthwise causal convolutions. Trained on 33B tokens in 40 B200 GPU-hours, Proust achieves Spearman ρ = 0.390 on ProteinGym substitutions, competitive with MLMs requiring 50--200× the compute. On indels, Proust sets a new state-of-the-art, outperforming models up to 20× larger. On EVEREST viral fitness benchmarks, it approaches structure-aware methods using sequence alone. These powerful representations position Proust in a sweet spot as it also retains native generative capabilities that MLMs lack by design. Interpretability analysis reveals that per-position entropy variance predicts, to an extent, when retrieval augmentation helps and hurts. Such insights can grow in both quantity and quality at scale and inform capabilities such as test-time scaling. Code and weights are available at https://github.com/Furkan9015/proust-inference
