Protein Counterfactuals via Diffusion-Guided Latent Optimization
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Abstract (arXiv)
Deep learning models can predict protein properties with unprecedented accuracy but rarely offer mechanistic insight or actionable guidance for engineering improved variants. When a model flags an antibody as unstable, the protein engineer is left without recourse: which mutations would rescue stability while preserving function? We introduce Manifold-Constrained Counterfactual Optimization for Proteins (MCCOP), a framework that computes minimal, biologically plausible sequence edits that flip a model's prediction to a desired target state. MCCOP operates in a continuous joint sequence-structure latent space and employs a pretrained diffusion model as a manifold prior, balancing three objectives: validity (achieving the target property), proximity (minimizing mutations), and plausibility (producing foldable proteins). We evaluate MCCOP on three protein engineering tasks - GFP fluorescence rescue, thermodynamic stability enhancement, and E3 ligase activity recovery - and show that it generates sparser, more plausible counterfactuals than both discrete and continuous baselines. The recovered mutations align with known biophysical mechanisms, including chromophore packing and hydrophobic core consolidation, establishing MCCOP as a tool for both model interpretation and hypothesis-driven protein design. Our code is publicly available at github.com/weroks/mccop.
Key facts. Cited by 3 papers in the map (data through 14 Oct 2026). Builds on 22 of its 44 references that are in the map. First posted on arXiv 11 Mar 2026.
Builds on
22 of 44 references.
References · 22
- 2014Explaining and Harnessing Adversarial Examplescited by 7,364
- 2018Spectral Normalization for Generative Adversarial Networkscited by 1,636
- 2017Counterfactual Explanations without Opening the Black Box: Automated Decisions and the GDPRcited by 1,089
- 2019Explaining Machine Learning Classifiers through Diverse Counterfactual Explanationscited by 614
- 2019Evaluating Protein Transfer Learning with TAPEcited by 231
- 2022Protein structure generation via folding diffusioncited by 115
- 2019Efficient Search for Diverse Coherent Explanationscited by 111
- 2022Diffusion Visual Counterfactual Explanationscited by 96
- 2018Improving DNN Robustness to Adversarial Attacks using Jacobian Regularizationcited by 88
- 2020BERTology Meets Biology: Interpreting Attention in Protein Language Modelscited by 85
- 2022Diffusion Models for Counterfactual Explanationscited by 79
- 2021Beyond Trivial Counterfactual Explanations with Diverse Valuable Explanationscited by 56
10 more
- 2023Adversarial Counterfactual Visual Explanationscited by 47
- 2023Fast Diffusion-Based Counterfactuals for Shortcut Removal and Generationcited by 41
- 2022Diffeomorphic Counterfactuals with Generative Modelscited by 29
- 2024Diffusion-based Iterative Counterfactual Explanations for Fetal Ultrasound Image Quality Assessmentcited by 13
- 2024Diffusion on language model encodings for protein sequence generationcited by 11
- 2024Large Language Models Cannot Explain Themselvescited by 11
- 2025Towards Desiderata-Driven Design of Visual Counterfactual Explainerscited by 7
- 2022ReLSO: A Transformer-based Model for Latent Space Optimization and Generation of Proteinscited by 4
- 2023ExplainableFold: Understanding AlphaFold Prediction with Explainable AIcited by 4
- 2025Graph Diffusion Counterfactual Explanationcited by 4
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Citing papers per year: 2026: 3.
3 papers.
- 2026SCE-LITE-HQ: Smooth visual counterfactual explanations with generative foundation modelscited by 3
- 2026Faithful, Sufficient and Understandable: Rethinking Graph Counterfactual Explanations via Discrete Diffusion Inversioncited by 1
- 2026Towards Fast and Disentangled Counterfactuals for Visual Foundation Modelscited by 0
- 2026Towards Fast and Disentangled Counterfactuals for Visual Foundation Models1 Oct 2026
- 2026Faithful, Sufficient and Understandable: Rethinking Graph Counterfactual Explanations via Discrete Diffusion Inversion12 Aug 2026
- 2026SCE-LITE-HQ: Smooth visual counterfactual explanations with generative foundation models17 Mar 2026
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- 2023Protein Design with Guided Discrete Diffusion
- 2025A Variational Perspective on Generative Protein Fitness Optimization
- 2025Constrained Diffusion for Protein Design with Hard Structural Constraints
- 2026Self-Improvement Imitation with Biologically Guided Search for Protein Design Under Oracle Budgets
- 2025Controllable protein design with particle-based Feynman-Kac steering
- 2024Reinforcement learning on structure-conditioned categorical diffusion for protein inverse folding
- 2022Protein Sequence and Structure Co-Design with Equivariant Translation