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Deep Supervised and Convolutional Generative Stochastic Network for Protein Secondary Structure Prediction

Jian Zhou, Olga G. Troyanskaya

arXiv:1403.1347v1q-bio.QMcs.CEcs.LG

TL;DR

Protein secondary-structure prediction must assign local structural states despite difficult long-range dependencies and limited homologous information. The paper develops a supervised convolutional GSN that samples labels conditioned on protein inputs using hierarchical representations, achieving superior 8-state prediction accuracy while retaining limitations for rare states and hard-coded spatial connectivity.

  • Problem

    The paper addresses 8-state secondary-structure prediction for proteins without close homologs, where local labels depend on complex long-range interactions.

  • Method

    The method extends GSNs to supervised conditional sampling and adds convolutional hierarchical representations that combine local and global information.

  • Results

    The model demonstrates superior accuracy on 8-state protein secondary-structure prediction over previous methods.

  • Takeaways & Limitations

    The experiments suggest supervised GSNs can effectively model structured prediction and may apply to other protein structure-prediction tasks.

  • Takeaways & Limitations

    The convolutional structure is hard-coded and may not optimally capture spatial organization, especially for long-range interactions.

Abstract

from arXiv · show

Predicting protein secondary structure is a fundamental problem in protein structure prediction. Here we present a new supervised generative stochastic network (GSN) based method to predict local secondary structure with deep hierarchical representations. GSN is a recently proposed deep learning technique (Bengio & Thibodeau-Laufer, 2013) to globally train deep generative model. We present the supervised extension of GSN, which learns a Markov chain to sample from a conditional distribution, and applied it to protein structure prediction. To scale the model to full-sized, high-dimensional data, like protein sequences with hundreds of amino acids, we introduce a convolutional architecture, which allows efficient learning across multiple layers of hierarchical representations. Our architecture uniquely focuses on predicting structured low-level labels informed with both low and high-level representations learned by the model. In our application this corresponds to labeling the secondary structure state of each amino-acid residue. We trained and tested the model on separate sets of non-homologous proteins sharing less than 30% sequence identity. Our model achieves 66.4% Q8 accuracy on the CB513 dataset, better than the previously reported best performance 64.9% (Wang et al., 2011) for this challenging secondary structure prediction problem.

1. Introduction

The paper targets 8-state secondary-structure prediction for proteins without close homologs, where local labels depend on complex long-range interactions. It introduces supervised and convolutional GSN techniques to learn hierarchical representations while combining local and global information.

  • Protein secondary-structure prediction is difficult for sequences lacking close homologs with known three-dimensional structures.
  • 8-state prediction is presented as a challenging, less-addressed problem for proteins without close homologs.
  • Long-range interactions, such as hydrogen bonds between distant β-strands, make global sequence information important for local structure labels.
  • The supervised GSN learns a Markov chain for sampling outputs conditioned on inputs without explicitly marginalizing hidden variables.
  • The convolutional architecture learns hierarchical representations for full-sized protein sequences and communicates high- and low-level features across spatial locations.

2. Preliminaries

GSNs learn data distributions through noisy reconstruction and Markov-chain sampling, while supervised GSNs adapt this framework to conditional structured prediction. The supervised model samples labels conditioned on inputs and trains reconstruction distributions without hidden-variable marginalization.

  • A GSN learns a deep generative model by reconstructing corrupted inputs and sampling from a learned Markov chain.
  • GSNs avoid intractable inference and explicit marginalization over latent variables by learning to sample directly from the data distribution.
  • The supervised GSN captures the conditional dependency structure of outputs given inputs by sampling from P(Y |X).
  • Training estimates Pθ(Y | ˜Y , X) from corrupted labels and inputs using a reconstruction cost interpretable as regularized conditional log likelihood.
  • Prediction initializes labels arbitrarily and repeatedly samples through the learned Markov chain conditioned on X.

3. Algorithms

Convolutional GSNs use shared one-dimensional filters, pooling, noise, and recurrent bidirectional sampling to build hierarchical representations for sequential data. In supervised use, label channels are reconstructed while feature channels remain available as inputs.

  • Convolutional GSN replaces fully connected spatial connectivity with one-dimensional convolutions suited to sequential protein data.
  • A convolutional layer uses shared filters over local windows, producing feature maps from visible input channels.
  • Noise enters through a stochastic activation function, and gradients through stochastic neurons use a straight-through estimator.
  • Pooling reduces upper-layer feature-map size and supports higher-level representations by averaging adjacent features.
  • Unlike feed-forward convolutional networks, recurrent convolutional GSN connections let different layers and spatial positions communicate through alternating upward and downward passes.
  • For supervised prediction, the architecture corrupts and reconstructs label channels while leaving feature channels uncorrupted as inputs to hidden activations.

4. Experimental Results

The experiments evaluate the model on non-homologous protein datasets, compare architectural and training choices, and report strong 8-state secondary-structure prediction performance. Results also identify challenges for rare structure states and show benefits from deeper models and kick-start training.

  • Features and dataset: The study targets 8-state secondary-structure and solvent-accessibility prediction using multi-task learning on protein sequences.The dataset contains 57 feature and label channels, with sequences padded to a maximum length of 700 amino acids.
  • Performance: 72.1 ± 0.6% Q8 accuracy was achieved on the Cull PDB test sequences with the best tested 3-layer convolutional architecture.The architecture uses convolutional and mean-pooling layers with 80 channels in each convolutional layer.
  • Performance: 0.664 Q8 accuracy was achieved on CB513, exceeding the previous CNF/Raptor-SS8 result of 0.649.The CB513 evaluation trained on Cull PDB sequences filtered to remove sequences sharing more than 25% identity with CB513.
  • Analysis of architecture: The 3-layer model slightly outperforms the 2-layer model, while omitting kick-start training reduces convergence speed and prediction accuracy.The authors attribute kick-start’s benefit to learning a better reconstruction distribution from arbitrary initial states; rare states G, S, and especially I remain difficult.

5. Conclusions

The paper combines supervised GSNs with convolutional architectures for structured prediction, capturing local information while incorporating high-level and distant features. The authors report success on protein secondary structure prediction but identify hard-coded connectivity as a limitation for modeling long-range interactions.

  • The method combines supervised generative stochastic networks with convolutional architectures to learn hierarchical representations for full-sized protein sequences.This combination is presented as an effective approach for structured prediction.
  • The architecture targets low-level structured predictions using local information informed by high-level and distant features.The authors suggest this design may extend beyond bioinformatics to tasks such as scene parsing and image segmentation.
  • The current convolutional structure is hard-coded and may not optimally capture protein sequence organization, especially for structures formed by long-range interactions.The authors propose adaptive, dynamic connectivity as a possible direction for improvement.
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