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Genes in which the expression varied between different samples were selected with the FindVariableGenes function in the SEURAT package [47]. Genes were first divided into 20 bins based on their average expression levels across all samples. The log-transformed values of the variance divided by the mean, as a measure of dispersion, were then calculated in each bin and z-scored. A set of variable genes were then obtained by setting the threshold of the dispersion value > 0.8 and the average log2-transformed FPKM value > 3 (Additional file 3: Figure S2).

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