From 5f5015940641b0e365bc63be9e3354a07729636f Mon Sep 17 00:00:00 2001 From: shafighi Date: Thu, 21 May 2026 14:38:49 +0100 Subject: [PATCH 1/4] Add mouse (GRCm38) support: extended blacklisting and coverage metric - readFilter(): add GRCm38 branch loading mm10-blacklist.v2.bed so centromere/low-mappability bins are excluded for mouse data - readData(): compute coverage pData column for Mouse using 2.7 Gb genome size; also store genome in experimentData for Mouse - Extend extendedBlacklisting condition to apply for both Human and Mouse --- R/core.R | 22 ++++++++++++++++++---- 1 file changed, 18 insertions(+), 4 deletions(-) diff --git a/R/core.R b/R/core.R index 24f9a0b..ff600d0 100644 --- a/R/core.R +++ b/R/core.R @@ -759,9 +759,13 @@ readData <- function(bamfiles, binSize, species = "Human", filterChromosomes=c(" Biobase::pData(readCounts)["genome"] = genome coverage = (readCounts@phenoData@data$used.reads * (as.numeric(isPaired) + 1) * read_size) / 3.2e9 Biobase::pData(readCounts)["coverage"] = coverage + } else if(species == "Mouse"){ + Biobase::pData(readCounts)["genome"] = genome + coverage = (readCounts@phenoData@data$used.reads * (as.numeric(isPaired) + 1) * read_size) / 2.7e9 + Biobase::pData(readCounts)["coverage"] = coverage } - - if(species == "Human" & extendedBlacklisting){ + + if(extendedBlacklisting & (species == "Human" | species == "Mouse")){ if(binSize < 10) stop("blacklisting doesn't support binsizes smaller than 10kb") # this is because our blacklisted regions have a resolution of 10kb if(binSize %in% c(200, 2000, 5000) && genome == "GRCh37"){ @@ -809,7 +813,9 @@ readData <- function(bamfiles, binSize, species = "Human", filterChromosomes=c(" if(species == "Human"){ Biobase::experimentData(readCounts)@other = list(species="Human", genome=genome) - }else{ + } else if(species == "Mouse"){ + Biobase::experimentData(readCounts)@other = list(species="Mouse", genome=genome) + } else { Biobase::experimentData(readCounts)@other = list(species=species) } @@ -823,7 +829,7 @@ readData <- function(bamfiles, binSize, species = "Human", filterChromosomes=c(" #' @return QDNAseq object with updated bin usability readFilter <- function(readCounts, genome="GRCh37"){ - stopifnot(genome %in% c("GRCh37", "GRCh38")) + stopifnot(genome %in% c("GRCh37", "GRCh38", "GRCm38")) if(genome == "GRCh37"){ gr = createGR(readCounts[,1,drop=FALSE]) excluded_regions_bed = readr::read_tsv(file.path(BASEDIR, "data/blacklisting/final_exclude_regions_hg19.bed"), @@ -836,6 +842,14 @@ readFilter <- function(readCounts, genome="GRCh37"){ warning("blacklist has been optimized for GRCh37 only") stop("Not supported") } + if(genome == "GRCm38"){ + gr = createGR(readCounts[,1,drop=FALSE]) + excluded_regions_bed = readr::read_tsv(file.path(BASEDIR, "data/blacklisting/mm10-blacklist.v2.bed"), + col_names = c("chromosome", "start", "end", "reason"), col_types="ciic") + excluded_regions = GRanges(seqnames=excluded_regions_bed$chromosome, + ranges = IRanges(start=excluded_regions_bed$start+1, end=excluded_regions_bed$end), + seqinfo = seqinfo(gr)) + } hits <- findOverlaps(excluded_regions, gr) From 2f195bac8150a41c50102e46f92119ae24a288da Mon Sep 17 00:00:00 2001 From: shafighi Date: Thu, 21 May 2026 16:47:55 +0100 Subject: [PATCH 2/4] Add pericentromeric bin filter for mouse (GRCm38) Mouse chromosomes are telocentric: the centromere sits at position 0 of every chromosome, followed by unassembled satellite sequence. The UCSC mm10 gap track documents a formal centromere N-gap from 0 to 3,000,000 bp on all 21 chromosomes. The existing ENCODE mm10-blacklist.v2.bed already covers chr2-4, chr6, chr9-chr19, and chrY from position 0. Five chromosomes were unprotected near the start (chr1, chr5, chr7, chr8, chrX). Changes: - Add data/blacklisting/mm10_pericentromeric_exclude.bed covering 0-3,000,000 bp on all chromosomes, based on the UCSC centromere gap - readFilter() for GRCm38 now merges this file with mm10-blacklist.v2.bed before applying findOverlaps, so both sources contribute to bin exclusion The BED file uses base-pair coordinates so it is bin-size-agnostic: the findOverlaps logic in readFilter() handles any bin size correctly. This also covers the chrX PAR region (0-700,000 bp) for male samples. --- R/core.R | 10 +++++++-- .../mm10_pericentromeric_exclude.bed | 21 +++++++++++++++++++ 2 files changed, 29 insertions(+), 2 deletions(-) create mode 100644 data/blacklisting/mm10_pericentromeric_exclude.bed diff --git a/R/core.R b/R/core.R index ff600d0..5c494db 100644 --- a/R/core.R +++ b/R/core.R @@ -844,8 +844,14 @@ readFilter <- function(readCounts, genome="GRCh37"){ } if(genome == "GRCm38"){ gr = createGR(readCounts[,1,drop=FALSE]) - excluded_regions_bed = readr::read_tsv(file.path(BASEDIR, "data/blacklisting/mm10-blacklist.v2.bed"), - col_names = c("chromosome", "start", "end", "reason"), col_types="ciic") + blacklist_bed = readr::read_tsv(file.path(BASEDIR, "data/blacklisting/mm10-blacklist.v2.bed"), + col_names = c("chromosome", "start", "end", "reason"), col_types="ciic") + pericentro_bed = readr::read_tsv(file.path(BASEDIR, "data/blacklisting/mm10_pericentromeric_exclude.bed"), + col_names = c("chromosome", "start", "end"), col_types="cii") + excluded_regions_bed = dplyr::bind_rows( + blacklist_bed[, c("chromosome", "start", "end")], + pericentro_bed + ) excluded_regions = GRanges(seqnames=excluded_regions_bed$chromosome, ranges = IRanges(start=excluded_regions_bed$start+1, end=excluded_regions_bed$end), seqinfo = seqinfo(gr)) diff --git a/data/blacklisting/mm10_pericentromeric_exclude.bed b/data/blacklisting/mm10_pericentromeric_exclude.bed new file mode 100644 index 0000000..d36db1d --- /dev/null +++ b/data/blacklisting/mm10_pericentromeric_exclude.bed @@ -0,0 +1,21 @@ +chr1 0 3000000 +chr2 0 3000000 +chr3 0 3000000 +chr4 0 3000000 +chr5 0 3000000 +chr6 0 3000000 +chr7 0 3000000 +chr8 0 3000000 +chr9 0 3000000 +chr10 0 3000000 +chr11 0 3000000 +chr12 0 3000000 +chr13 0 3000000 +chr14 0 3000000 +chr15 0 3000000 +chr16 0 3000000 +chr17 0 3000000 +chr18 0 3000000 +chr19 0 3000000 +chrX 0 3000000 +chrY 0 3000000 From 9e3a8d8c7f478d7a90292a73f0ecd15dac3beb4b Mon Sep 17 00:00:00 2001 From: shafighi Date: Thu, 21 May 2026 22:08:52 +0100 Subject: [PATCH 3/4] Add segmental duplication mask for mouse (GRCm38) Segmental duplications are near-identical copy-pasted regions of the genome. Reads from these regions cannot be unambiguously placed, making read counts unreliable. Bailey et al. 2008 (Nature Genetics) showed ~47% of apparent mouse CNVs overlap segmental duplications and are likely false positives. Mouse chr7, chr12, chr14, and chrX are particularly affected. The ENCODE mm10-blacklist.v2.bed does not cover these regions, so a separate mask is needed. Changes: - Add data/blacklisting/mm10_segdup.bed (UCSC mm10 segmental duplication track, 180,459 regions) - readFilter() for GRCm38 now merges all three sources: ENCODE blacklist, pericentromeric filter, and segdup mask - Random/unplaced contigs in the segdup file (chrUn_*, chr*_random, chrM) are filtered to only chromosomes present in the QDNAseq.mm10 bin set before building the GRanges object --- R/core.R | 6 +- data/blacklisting/mm10_segdup.bed | 180459 +++++++++++++++++++++++++++ 2 files changed, 180464 insertions(+), 1 deletion(-) create mode 100644 data/blacklisting/mm10_segdup.bed diff --git a/R/core.R b/R/core.R index 5c494db..7efff28 100644 --- a/R/core.R +++ b/R/core.R @@ -844,13 +844,17 @@ readFilter <- function(readCounts, genome="GRCh37"){ } if(genome == "GRCm38"){ gr = createGR(readCounts[,1,drop=FALSE]) + valid_chrs = seqlevels(gr) blacklist_bed = readr::read_tsv(file.path(BASEDIR, "data/blacklisting/mm10-blacklist.v2.bed"), col_names = c("chromosome", "start", "end", "reason"), col_types="ciic") pericentro_bed = readr::read_tsv(file.path(BASEDIR, "data/blacklisting/mm10_pericentromeric_exclude.bed"), col_names = c("chromosome", "start", "end"), col_types="cii") + segdup_bed = readr::read_tsv(file.path(BASEDIR, "data/blacklisting/mm10_segdup.bed"), + col_names = c("chromosome", "start", "end"), col_types="cii") excluded_regions_bed = dplyr::bind_rows( blacklist_bed[, c("chromosome", "start", "end")], - pericentro_bed + pericentro_bed, + segdup_bed[segdup_bed$chromosome %in% valid_chrs, ] ) excluded_regions = GRanges(seqnames=excluded_regions_bed$chromosome, ranges = IRanges(start=excluded_regions_bed$start+1, end=excluded_regions_bed$end), diff --git a/data/blacklisting/mm10_segdup.bed b/data/blacklisting/mm10_segdup.bed new file mode 100644 index 0000000..f748995 --- /dev/null +++ b/data/blacklisting/mm10_segdup.bed @@ -0,0 +1,180459 @@ +chr1 3531709 3535022 +chr1 5423434 5429532 +chr1 5423434 5429545 +chr1 5423435 5429437 +chr1 5423435 5429545 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158098 +chrY_JH584303_random 123938 151072 +chrY_JH584303_random 125327 151072 +chrY_JH584303_random 126403 151072 +chrY_JH584303_random 127971 158098 +chrY_JH584303_random 128653 151072 +chrY_JH584303_random 129938 151072 +chrY_JH584303_random 132763 151072 +chrY_JH584303_random 133111 158098 +chrY_JH584303_random 133889 150209 +chrY_JH584303_random 133889 156382 +chrY_JH584303_random 133934 156382 +chrY_JH584303_random 134412 147948 +chrY_JH584303_random 136765 158098 +chrY_JH584303_random 138511 158098 +chrY_JH584303_random 141219 151072 +chrY_JH584303_random 149451 158098 From 04fc71c34c314acc62c566fe179d4666aa26e963 Mon Sep 17 00:00:00 2001 From: shafighi Date: Thu, 21 May 2026 23:12:30 +0100 Subject: [PATCH 4/4] Replace mm9 replication timing source with mm10 liftover for mouse MIME-Version: 1.0 Content-Type: text/plain; charset=UTF-8 Content-Transfer-Encoding: 8bit The FSU RepliChip BigWig files used for mouse replication timing correction were in mm9 coordinates but were being averaged against mm10 GRanges bins without an explicit liftover — a silent coordinate mismatch. Fix: add liftover_mm9_to_mm10.sh which downloads the same 33 FSU RepliChip WaveSignal files from UCSC and lifts them to mm10 using the official mm9->mm10 chain (>97% of the genome retained). The R script is updated to read from repliChip_data_mm10/ instead of repliChip_data/. After running the shell script, re-run replicationInfo.R and replicationInfo_per_bin.R to regenerate replicationInfo_mouse.RDS and replicationTiming_per_binSize_mouse.RDS with correct mm10 coordinates. No changes to the averaging or binning logic — only the input file path changes. Same 33 cell types, same WaveSignal format. --- data/replicationTiming/README | 7 +- .../replicationTiming/liftover_mm9_to_mm10.sh | 115 ++++++++++++++++++ data/replicationTiming/replicationInfo.R | 8 +- 3 files changed, 125 insertions(+), 5 deletions(-) create mode 100755 data/replicationTiming/liftover_mm9_to_mm10.sh diff --git a/data/replicationTiming/README b/data/replicationTiming/README index 18f3046..f31e67c 100644 --- a/data/replicationTiming/README +++ b/data/replicationTiming/README @@ -2,5 +2,8 @@ RepliChip file sources: Human (hg19) http://genome.ucsc.edu/cgi-bin/hgFileUi?db=hg19&g=wgEncodeFsuRepliChip -Mouse (mm9) -http://genome.ucsc.edu/cgi-bin/hgFileUi?db=mm9&g=wgEncodeFsuRepliChip +Mouse (mm10) +FSU RepliChip WaveSignal files lifted from mm9 to mm10 using the UCSC liftover chain. +Run liftover_mm9_to_mm10.sh to download the mm9 source files and perform the liftover. +Original mm9 source: http://genome.ucsc.edu/cgi-bin/hgFileUi?db=mm9&g=wgEncodeFsuRepliChip +Liftover chain: https://hgdownload.soe.ucsc.edu/goldenPath/mm9/liftOver/mm9ToMm10.over.chain.gz diff --git a/data/replicationTiming/liftover_mm9_to_mm10.sh b/data/replicationTiming/liftover_mm9_to_mm10.sh new file mode 100755 index 0000000..d22011a --- /dev/null +++ b/data/replicationTiming/liftover_mm9_to_mm10.sh @@ -0,0 +1,115 @@ +#!/bin/bash +# One-time script to lift mm9 FSU RepliChip BigWig files over to mm10 coordinates. +# +# Prerequisites (install once): +# conda install -c bioconda ucsc-liftover ucsc-bigwigtobedgraph ucsc-bedgraphtobigwig +# OR download UCSC binaries directly: +# https://hgdownload.soe.ucsc.edu/admin/exe/macOSX.arm64/ +# +# Run from: scAbsolute/data/replicationTiming/ +# Output: repliChip_data_mm10/ (used by replicationInfo.R) +# After running this script, re-run: +# Rscript replicationInfo.R -> regenerates replicationInfo_mouse.RDS +# Rscript replicationInfo_per_bin.R -> regenerates replicationTiming_per_binSize_mouse.RDS + +set -e + +SCRIPT_DIR="$(cd "$(dirname "${BASH_SOURCE[0]}")" && pwd)" +MM9_DIR="${SCRIPT_DIR}/repliChip_data_mm9" +MM10_DIR="${SCRIPT_DIR}/repliChip_data_mm10" +BASE_URL="http://hgdownload.cse.ucsc.edu/goldenpath/mm9/encodeDCC/wgEncodeFsuRepliChip" +CHAIN="${SCRIPT_DIR}/mm9ToMm10.over.chain" +CHROM_SIZES="${SCRIPT_DIR}/mm10.chrom.sizes" + +mkdir -p "$MM9_DIR" "$MM10_DIR" + +echo "=== Downloading liftover chain and chrom sizes ===" +if [ ! -f "$CHAIN" ]; then + curl -s "https://hgdownload.soe.ucsc.edu/goldenPath/mm9/liftOver/mm9ToMm10.over.chain.gz" \ + | gunzip > "$CHAIN" +fi +if [ ! -f "$CHROM_SIZES" ]; then + curl -s "https://hgdownload.soe.ucsc.edu/goldenPath/mm10/bigZips/mm10.chrom.sizes" \ + -o "$CHROM_SIZES" +fi + +files=( + "wgEncodeFsuRepliChipCh12FWaveSignalRep1" + "wgEncodeFsuRepliChipCh12FWaveSignalRep2" + "wgEncodeFsuRepliChipEpisc5MWaveSignalRep1" + "wgEncodeFsuRepliChipEpisc5MWaveSignalRep2" + "wgEncodeFsuRepliChipEpisc7FWaveSignalRep1" + "wgEncodeFsuRepliChipEpisc7FWaveSignalRep2" + "wgEncodeFsuRepliChipEs46cMDifff6dWaveSignalRep1" + "wgEncodeFsuRepliChipEs46cMWaveSignalRep1" + "wgEncodeFsuRepliChipEsd3MDiffe3dWaveSignalRep1" + "wgEncodeFsuRepliChipEsd3MDiffe3dWaveSignalRep2" + "wgEncodeFsuRepliChipEsd3MDiffe6dWaveSignalRep1" + "wgEncodeFsuRepliChipEsd3MDiffe6dWaveSignalRep2" + "wgEncodeFsuRepliChipEsd3MDiffe9dWaveSignalRep1" + "wgEncodeFsuRepliChipEsd3MDiffe9dWaveSignalRep2" + "wgEncodeFsuRepliChipEsd3MDiffg3dWaveSignalRep1" + "wgEncodeFsuRepliChipEsd3MDiffg3dWaveSignalRep2" + "wgEncodeFsuRepliChipEsd3MWaveSignalRep1" + "wgEncodeFsuRepliChipEsd3MWaveSignalRep2" + "wgEncodeFsuRepliChipEsem5sUDiffhsoxmWaveSignalRep1" + "wgEncodeFsuRepliChipEsem5sUDiffhsoxmWaveSignalRep2" + "wgEncodeFsuRepliChipEsem5sUDiffhsoxpWaveSignalRep1" + "wgEncodeFsuRepliChipEsem5sUDiffhsoxpWaveSignalRep2" + "wgEncodeFsuRepliChipEstt2MDifff9dWaveSignalRep1" + "wgEncodeFsuRepliChipEstt2MDifff9dWaveSignalRep2" + "wgEncodeFsuRepliChipEstt2MWaveSignalRep1" + "wgEncodeFsuRepliChipEstt2MWaveSignalRep2" + "wgEncodeFsuRepliChipJ185aUWaveSignalRep1" + "wgEncodeFsuRepliChipJ185aUWaveSignalRep2" + "wgEncodeFsuRepliChipL1210FWaveSignalRep1" + "wgEncodeFsuRepliChipL1210FWaveSignalRep2" + "wgEncodeFsuRepliChipMefMWaveSignalRep1" + "wgEncodeFsuRepliChipMelMWaveSignalRep1" + "wgEncodeFsuRepliChipMelMWaveSignalRep2" +) + +for f in "${files[@]}"; do + out="${MM10_DIR}/${f}.bigWig" + if [ -f "$out" ]; then + echo "Skipping $f (already lifted)" + continue + fi + + echo "=== Processing $f ===" + + mm9_bw="${MM9_DIR}/${f}.bigWig" + if [ ! -f "$mm9_bw" ]; then + echo " Downloading mm9 BigWig..." + curl -s "${BASE_URL}/${f}.bigWig" -o "$mm9_bw" + fi + + echo " Converting BigWig -> bedGraph..." + bigWigToBedGraph "$mm9_bw" "${MM9_DIR}/${f}.bedGraph" + + echo " Lifting over mm9 -> mm10..." + liftOver \ + "${MM9_DIR}/${f}.bedGraph" \ + "$CHAIN" \ + "${MM10_DIR}/${f}_unsorted.bedGraph" \ + "${MM10_DIR}/${f}_unmapped.txt" + + echo " Sorting and converting bedGraph -> BigWig..." + sort -k1,1 -k2,2n "${MM10_DIR}/${f}_unsorted.bedGraph" \ + > "${MM10_DIR}/${f}_sorted.bedGraph" + bedGraphToBigWig \ + "${MM10_DIR}/${f}_sorted.bedGraph" \ + "$CHROM_SIZES" \ + "$out" + + rm -f "${MM10_DIR}/${f}_unsorted.bedGraph" \ + "${MM10_DIR}/${f}_sorted.bedGraph" + + echo " Done: $out" +done + +echo "" +echo "=== Liftover complete ===" +echo "Next steps:" +echo " Rscript replicationInfo.R" +echo " Rscript replicationInfo_per_bin.R" diff --git a/data/replicationTiming/replicationInfo.R b/data/replicationTiming/replicationInfo.R index b49ec41..a094c93 100644 --- a/data/replicationTiming/replicationInfo.R +++ b/data/replicationTiming/replicationInfo.R @@ -94,10 +94,12 @@ saveRDS(rep_rle, file=paste0(reptime_path, "/replicationInfo.RDS")) ##### Mouse replication information -# file source: http://hgdownload.soe.ucsc.edu/goldenPath/mm9/encodeDCC/wgEncodeFsuRepliChip/ or http://genome.ucsc.edu/cgi-bin/hgFileUi?db=mm9&g=wgEncodeFsuRepliChip ###mouse +# Source: FSU RepliChip WaveSignal BigWig files (originally mm9), lifted to mm10 +# using the UCSC mm9->mm10 liftover chain. Run liftover_mm9_to_mm10.sh first +# to populate repliChip_data_mm10/ before executing this section. +# Original mm9 source: http://genome.ucsc.edu/cgi-bin/hgFileUi?db=mm9&g=wgEncodeFsuRepliChip - -path = "~/scAbsolute/data/replicationTiming/repliChip_data/" #"~/Data/replication-timing/" +path = "~/scAbsolute/data/replicationTiming/repliChip_data_mm10/" files = c("wgEncodeFsuRepliChipCh12FWaveSignalRep1", "wgEncodeFsuRepliChipCh12FWaveSignalRep2", "wgEncodeFsuRepliChipEpisc5MWaveSignalRep1",