For residualized expression, we regressed out covariates from voom-normalized expression using a null model (equation (3)) and applied z-score normalization as described previously12:3\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$\begin{array}{l}E\left(Y\right)={\beta }_{0}+{\beta }_{1}{\mathrm{sex}}+{\beta }_{2}{\mathrm{age}}+{\beta }_{3}{{\mathrm{mito}}\,{\mathrm{rate}}}+{\beta }_{4}{{\mathrm{rRNA}}\,{\mathrm{rate}}}\\\qquad+\,{\beta }_{5}{{\mathrm{total}}\,{\mathrm{assigned}}\,{\mathrm{genes}}}+{\beta }_{6}{{\mathrm{overall}}\,{\mathrm{mapping}}\,{\mathrm{rate}}}+\mathop{\sum }\limits_{i=1}^{k}{\gamma }_{i}{qS}{V}_{i}\end{array}$$\end{document}EY=β0+β1sex+β2age+β3mitorate+β4rRNArate+β5totalassignedgenes+β6overallmappingrate+∑i=1kγiqSVi