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In principle, any gene feature expected to be held (or avoided) in concert for disease genes can be used in this approach. However, given our concern for biasing predictions towards well-studied genes, we built an extensive set of >40,000 features with minimal preference for previously characterized genes. Genomic features were derived from publicly available genome-wide expression data, transcription factor binding information (both observed and predicted), phylogenetic profiles, protein domain organization, and predicted microRNA (miRNA) targets (Table 1). In most cases, preprocessing was required to generate useful predictors. For example, for microarray analysis, we automated downloading, normalization, clustering, and differential gene expression analysis for 1,437 human and murine microarray data sets obtained through the Gene Expression Omnibus (GEO) [23]. As a comparison to assess the impact of biased features on our predictions, we also downloaded GO annotations and used each of the 17,156 terms as descriptive features.




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Complete datasets corresponding to the GNF Body Atlas [55] and Neurocrine Tissue Atlases [56] were downloaded from GEO (GSE7307). The GNF atlas includes 158 microarrays corresponding to 79 different human tissue types (2 technical replicates). The Neurocrine atlas includes 676 microarrays corresponding to 65 human tissue types or cell lines (normal and/or diseased), from 10 donors (a total of 141 conditions were available - such as 'normal prostate' or 'diseased prostate'). Intensities for the GNF arrays were averaged across replicates. For each tissue atlas, the intensity corresponding to the 75th, 90th and 99th percentile across all tissues was identified. Only those probesets mapping uniquely to a single gene were considered, and for those genes mapping to multiple probesets, the probeset with the highest mean expression across all tissues was used. Each combination of percentile threshold and array was included as a single feature, with 1s and 0s assigned to genes whose expression exceeded or fell below the percentile cutoff, respectively. Imputation was performed for any genes not included on the microarrays.


We devised an automated approach to determine gene signatures for a large number of microarrays experiments deposited in GEO. First we identified all experiments performed on the two most commonly used microarray platforms: Affymetrix Human Genome U133A Plus 2.0 and Affymetrix Mouse Genome 430 2.0 Array. Next, for each experiment we downloaded all the corresponding microarrays, normalized them, and performed hierarchical clustering of samples using the hclust function in the R statistical framework [57]. We defined individual sample groups based on multiple cutpoints along the dendrogram (corresponding to tree heights of 0.975, 0.95, 0.925) and evaluated differentially expressed genes between groups using limma [53]. Features consisted of