Cell-level QC

Will Macnair - Neuroscience and Rare Diseases, Roche Innovation Center, Basel, Switzerland

February 28, 2026

Sample QC metrics

An overview of quality control (QC) metrics for all samples, both before and after filtering. The QC metrics include:

  • no. of cells : Total number of barcodes called as cells for each sample.

  • no. of UMIs : Sum of counts across all features for each cell with log-transformed values displayed.

  • no. of genes : Number of unique features with non-zero counts for each cell with log-transformed values displayed.

  • mito pct. : The proportion of reads that mapped to genes in the mitochondrial genome. High proportions are indicative of poor-quality cells (compromised membranes allow individual RNA molecules to escape while retaining mitochondria, leading to an increased relative abundance of mitochondrial transcript) or nuclei (incomplete removal of cytoplasm). Logit-transformed values are displayed as applying the logit (inverse logistic) transformation to mitochondrial proportions yields approximately Gaussian distributions.

  • spliced pct. : The proportion of spliced reads. In single-nuclei RNA sequencing, high spliced proportions may indicate inadequate removal of cytoplasmic material from the nuclei. Logit-transformed values are displayed as applying the logit (inverse logistic) transformation to spliced proportions results in approximately Gaussian distributions.

suppressWarnings({
  cat("### Pre-QC\n")
  print(plot_qc_ranges_marginals(qc_dt, b_lvls, qc_names, qc_lu, cuts_dt, batch_var))
  cat("\n\n")
  cat("### After light QC\n")
  print(plot_qc_ranges_marginals(kept_dt, b_lvls, qc_names, qc_lu, cuts_dt, batch_var))
  cat("\n\n")
})

Pre-QC

After light QC

QC metrics scatterplots

Pairwise relationships of QC metrics at individual cell level.

for (b in b_lvls) {
  qc_tmp      = qc_dt[ batch_var == b ]
  if (nrow(qc_tmp) == 1)
    next
  cat('### ', b, '\n')
  print(plot_qc_metric_scatter(qc_tmp, qc_names, qc_lu, cuts_dt[ batch_var == b ], b))
  cat('\n\n')
}

GSM5292146

GSM5292147

GSM5292156

GSM5292157

GSM5292158

GSM5292200

SRR27882183

SRR27882182

SRR27882181

SRR27882171

SRR27882170

SRR27882169

SRR27882094

SRR27882093

SRR27882092

SRR27882091

SRR27882090

SRR27882089

SRR27882088

SRR27882086

SRR27882085

SRR27882084

SRR27882158

SRR27882157

SRR27882156

SRR27882154

SRR27882153

SRR27882152

SRR27882151

SRR27882150

SRR27882149

SRR27882148

GSM5292143

GSM5292144

GSM5292145

GSM5292148

GSM5292149

GSM5292150

GSM5292151

GSM5292152

GSM5292153

GSM5292154

GSM5292155

GSM5292194

GSM5292195

GSM5292196

GSM5292197

GSM5292198

GSM5292199

SRR27882082

SRR27882081

SRR27882080

SRR27882079

SRR27882078

SRR27882077

SRR27882075

SRR27882074

SRR27882073

SRR27882180

SRR27882179

SRR27882178

SRR27882177

SRR27882175

SRR27882174

SRR27882173

SRR27882172

GSM5292174

GSM5292175

GSM5292176

GSM5292177

GSM5292178

GSM5292179

GSM5292180

GSM5292181

GSM5292182

GSM5292183

GSM5292184

GSM5292185

GSM5292186

GSM5292187

GSM5292188

GSM5292193

GSM5292201

SRR27882193

SRR27882192

SRR27882191

SRR27882190

SRR27882189

SRR27882188

SRR27882187

SRR27882186

SRR27882185

SRR27882165

SRR27882164

SRR27882163

SRR27882162

SRR27882161

SRR27882160

SRR27882159

GSM5292164

GSM5292169

GSM5292173

GSM5292190

GSM5292191

GSM5292192

GSM5292202

GSM5292203

GSM5292204

GSM5292205

GSM5292206

GSM5292207

GSM5292208

SRR27882214

SRR27882212

SRR27882203

SRR27882202

SRR27882201

SRR27882200

SRR27882199

SRR27882198

SRR27882197

SRR27882196

SRR27882194

GSM5292159

GSM5292160

GSM5292161

GSM5292162

GSM5292163

GSM5292165

GSM5292166

GSM5292167

GSM5292168

GSM5292170

GSM5292171

GSM5292172

GSM5292189

SRR27882168

SRR27882167

SRR27882166

SRR27882216

SRR27882215

SRR27882213

SRR27882211

SRR27882210

SRR27882209

SRR27882208

SRR27882207

SRR27882205

SRR27882204

How many cells and samples retained?

Table summarizing the number of cells before and after qc filtering. Some samples may have been excluded from further analysis due to insufficient number of cells.

calc_qc_summary(qc_dt, kept_dt, cuts_dt, qc_lu, batch_var) %>% knitr::kable()
sample_id excluded N pre-QC N post-QC N excluded pct. excluded pct. excluded by no. of UMIs pct. excluded by no. of genes pct. excluded by mito. pct. pct. excluded by spliced pct.
SRR27882190 FALSE 10912 2929 7983 73.2 71.1 69.3 7.7 1.5
GSM5292179 FALSE 2254 961 1293 57.4 25.0 24.4 50.9 2.5
SRR27882203 FALSE 2729 1197 1532 56.1 14.0 17.0 47.1 2.7
GSM5292169 FALSE 2973 1358 1615 54.3 35.2 33.1 48.3 0.3
GSM5292195 FALSE 3655 1678 1977 54.1 14.0 12.9 47.3 5.4
SRR27882162 FALSE 1384 715 669 48.3 5.6 7.6 45.4 1.4
SRR27882168 FALSE 6587 3476 3111 47.2 13.2 13.6 42.0 0.6
GSM5292204 FALSE 4374 2361 2013 46.0 4.0 4.0 44.5 0.5
GSM5292206 FALSE 3361 2023 1338 39.8 23.4 18.2 21.9 1.5
SRR27882197 FALSE 3717 2377 1340 36.1 17.0 15.4 24.9 0.9
GSM5292182 FALSE 1843 1203 640 34.7 8.8 8.6 31.8 1.6
GSM5292159 FALSE 5205 3558 1647 31.6 12.2 10.3 27.4 4.0
GSM5292185 FALSE 4434 3109 1325 29.9 9.4 8.0 24.8 0.5
GSM5292200 FALSE 6843 4813 2030 29.7 10.9 9.5 24.6 5.6
SRR27882165 FALSE 8289 5898 2391 28.8 10.5 9.6 22.7 7.0
GSM5292198 FALSE 7768 5645 2123 27.3 8.4 6.3 18.4 21.9
SRR27882170 FALSE 8945 6577 2368 26.5 10.3 9.3 18.1 19.0
GSM5292193 FALSE 6079 4480 1599 26.3 8.6 8.3 20.0 7.9
GSM5292173 FALSE 3788 2795 993 26.2 12.9 11.4 18.7 0.4
GSM5292190 FALSE 5413 4015 1398 25.8 10.3 9.5 19.1 1.8
GSM5292189 FALSE 9446 7059 2387 25.3 15.9 12.8 8.7 12.2
SRR27882183 FALSE 17079 12810 4269 25.0 24.3 19.3 1.3 0.2
SRR27882200 FALSE 8994 6792 2202 24.5 7.0 7.2 20.1 0.2
SRR27882160 FALSE 8908 6883 2025 22.7 10.6 8.9 15.6 1.1
GSM5292196 FALSE 7221 5609 1612 22.3 10.4 7.4 13.5 11.5
SRR27882192 FALSE 5120 4005 1115 21.8 8.1 8.8 16.6 1.8
GSM5292161 FALSE 6486 5099 1387 21.4 7.8 7.0 16.9 3.6
GSM5292192 FALSE 5053 4020 1033 20.4 13.5 12.5 13.9 0.8
GSM5292177 FALSE 4070 3245 825 20.3 12.5 10.3 11.1 3.4
SRR27882199 FALSE 21084 16821 4263 20.2 9.5 8.6 14.9 1.2
SRR27882172 FALSE 14988 12215 2773 18.5 9.4 7.9 10.8 11.7
SRR27882081 FALSE 8095 6617 1478 18.3 8.1 7.7 13.0 11.0
GSM5292165 FALSE 10904 9008 1896 17.4 7.1 6.7 14.6 0.6
SRR27882171 FALSE 12692 10498 2194 17.3 8.0 6.6 10.6 6.6
SRR27882214 FALSE 6549 5414 1135 17.3 6.7 6.7 12.9 2.8
SRR27882188 FALSE 6182 5118 1064 17.2 12.1 11.2 8.2 0.6
GSM5292194 FALSE 12425 10328 2097 16.9 9.0 5.7 9.0 9.0
SRR27882175 FALSE 5298 4402 896 16.9 10.5 9.1 10.1 7.0
SRR27882073 FALSE 5822 4842 980 16.8 10.8 8.8 9.2 1.0
GSM5292207 FALSE 3668 3055 613 16.7 12.4 9.8 2.6 5.6
SRR27882189 FALSE 27458 22887 4571 16.6 16.3 12.2 0.8 0.2
GSM5292171 FALSE 2663 2220 443 16.6 9.3 8.6 10.5 2.9
GSM5292197 FALSE 9407 7912 1495 15.9 7.0 5.9 11.1 5.0
SRR27882169 FALSE 11735 9922 1813 15.4 7.9 6.5 7.2 9.8
SRR27882077 FALSE 14215 12041 2174 15.3 4.3 3.2 10.9 13.8
GSM5292199 FALSE 8616 7299 1317 15.3 5.8 4.0 9.1 12.2
SRR27882205 FALSE 7728 6587 1141 14.8 7.5 7.2 9.8 1.4
SRR27882082 FALSE 11720 10025 1695 14.5 7.5 5.8 7.9 10.4
GSM5292176 FALSE 7542 6447 1095 14.5 11.7 9.0 3.5 1.6
SRR27882216 FALSE 8536 7303 1233 14.4 5.4 5.1 11.2 1.1
SRR27882163 FALSE 11192 9593 1599 14.3 8.5 7.2 9.6 3.6
SRR27882150 FALSE 8670 7438 1232 14.2 9.7 7.9 1.7 5.8
GSM5292203 FALSE 5097 4376 721 14.1 7.0 6.2 10.0 0.7
SRR27882158 FALSE 7103 6162 941 13.2 8.7 7.5 3.5 5.9
SRR27882161 FALSE 5906 5140 766 13.0 8.0 7.2 7.1 3.3
SRR27882094 FALSE 10333 9005 1328 12.9 8.7 7.2 1.9 5.5
SRR27882181 FALSE 13748 11984 1764 12.8 3.3 3.1 10.3 10.2
SRR27882159 FALSE 6881 6001 880 12.8 7.2 6.4 8.6 2.6
SRR27882182 FALSE 10809 9441 1368 12.7 9.4 8.7 6.1 2.2
SRR27882180 FALSE 15818 13840 1978 12.5 6.2 4.9 7.8 8.5
SRR27882092 FALSE 11841 10395 1446 12.2 7.3 5.7 1.9 6.7
SRR27882157 FALSE 9256 8147 1109 12.0 5.3 4.3 1.3 8.6
GSM5292156 FALSE 7433 6589 844 11.4 7.6 6.2 3.7 2.9
SRR27882201 FALSE 12137 10770 1367 11.3 4.8 5.1 7.8 1.7
SRR27882156 FALSE 8944 7966 978 10.9 7.4 6.0 1.0 5.0
GSM5292175 FALSE 7430 6634 796 10.7 5.2 4.3 5.3 5.7
SRR27882093 FALSE 11990 10731 1259 10.5 7.3 5.9 1.2 4.5
GSM5292150 FALSE 12051 10826 1225 10.2 8.0 6.2 1.7 3.4
SRR27882174 FALSE 9724 8734 990 10.2 7.3 5.6 3.7 1.9
GSM5292162 FALSE 9671 8709 962 9.9 3.7 2.7 5.6 7.3
SRR27882177 FALSE 10668 9643 1025 9.6 4.3 4.2 6.5 6.1
SRR27882086 FALSE 8392 7604 788 9.4 5.8 4.9 0.6 5.0
GSM5292181 FALSE 6573 5955 618 9.4 6.6 5.3 1.0 4.6
SRR27882079 FALSE 10190 9244 946 9.3 4.9 4.2 5.9 6.1
SRR27882075 FALSE 14749 13388 1361 9.2 5.2 4.3 4.7 5.0
SRR27882091 FALSE 12066 10962 1104 9.1 4.8 3.6 0.7 5.8
GSM5292178 FALSE 6630 6036 594 9.0 4.2 3.9 5.2 4.1
SRR27882080 FALSE 10818 9864 954 8.8 5.7 5.0 2.5 5.2
GSM5292158 FALSE 8334 7621 713 8.6 6.9 5.3 2.3 2.0
GSM5292152 FALSE 5029 4599 430 8.6 5.4 4.3 2.4 4.3
SRR27882166 FALSE 6434 5888 546 8.5 5.4 5.2 4.4 1.3
GSM5292184 FALSE 5549 5078 471 8.5 5.1 4.4 4.1 1.5
GSM5292160 FALSE 12538 11482 1056 8.4 5.4 4.2 2.6 3.7
SRR27882154 FALSE 10678 9792 886 8.3 5.8 4.3 1.0 3.4
SRR27882211 FALSE 8684 7974 710 8.2 4.4 5.3 4.1 2.1
SRR27882085 FALSE 7350 6764 586 8.0 5.2 4.8 1.9 3.9
SRR27882191 FALSE 5197 4782 415 8.0 4.6 4.8 4.9 2.3
SRR27882185 FALSE 12895 11881 1014 7.9 3.8 2.9 2.9 5.6
SRR27882179 FALSE 9922 9135 787 7.9 3.9 3.0 4.2 6.1
GSM5292167 FALSE 6821 6291 530 7.8 6.8 5.2 1.6 0.7
GSM5292168 FALSE 3676 3391 285 7.8 4.6 4.5 4.9 0.1
GSM5292188 FALSE 5879 5424 455 7.7 6.2 5.2 1.5 1.6
SRR27882090 FALSE 8007 7400 607 7.6 5.5 5.0 0.7 2.8
SRR27882153 FALSE 8628 7981 647 7.5 5.2 4.5 1.0 3.4
GSM5292155 FALSE 6601 6114 487 7.4 4.1 2.8 1.5 5.2
GSM5292146 FALSE 10398 9646 752 7.2 3.7 2.6 2.2 5.0
SRR27882207 FALSE 7070 6561 509 7.2 4.9 4.3 1.3 3.3
GSM5292191 FALSE 8859 8233 626 7.1 2.6 2.2 1.0 5.9
GSM5292187 FALSE 7203 6690 513 7.1 5.6 4.9 1.5 1.5
SRR27882198 FALSE 4879 4533 346 7.1 3.1 2.8 4.2 5.0
SRR27882209 FALSE 8418 7826 592 7.0 3.8 3.6 2.7 4.0
GSM5292148 FALSE 11077 10320 757 6.8 3.9 3.5 2.6 4.1
SRR27882167 FALSE 9423 8783 640 6.8 4.6 3.8 3.2 2.0
SRR27882149 FALSE 9245 8612 633 6.8 4.2 3.4 0.3 3.5
GSM5292186 FALSE 8565 7984 581 6.8 4.0 2.9 2.4 3.5
GSM5292151 FALSE 10250 9564 686 6.7 5.1 3.2 0.7 2.5
GSM5292149 FALSE 14344 13397 947 6.6 3.8 2.8 2.4 4.1
GSM5292180 FALSE 7656 7150 506 6.6 3.0 2.6 4.4 2.9
SRR27882196 FALSE 4869 4556 313 6.4 3.8 3.2 1.9 3.5
SRR27882178 FALSE 12062 11304 758 6.3 4.2 3.4 2.2 3.1
GSM5292170 FALSE 4021 3766 255 6.3 4.7 4.4 1.4 1.5
SRR27882164 FALSE 14729 13826 903 6.1 3.9 2.9 2.5 1.9
GSM5292143 FALSE 10348 9732 616 6.0 4.2 3.4 1.6 2.7
SRR27882187 FALSE 9207 8650 557 6.0 3.9 3.2 2.7 1.5
SRR27882212 FALSE 8641 8122 519 6.0 4.7 4.1 1.0 2.0
GSM5292147 FALSE 12168 11447 721 5.9 3.6 2.5 1.8 3.8
GSM5292157 FALSE 8897 8385 512 5.8 3.0 2.1 0.9 3.6
SRR27882208 FALSE 8891 8382 509 5.7 3.7 3.8 2.6 1.3
SRR27882074 FALSE 8758 8261 497 5.7 4.4 3.7 1.7 1.6
GSM5292163 FALSE 13090 12359 731 5.6 3.7 2.4 1.1 2.6
SRR27882194 FALSE 4994 4720 274 5.5 2.7 2.2 2.8 3.8
GSM5292201 FALSE 6508 6155 353 5.4 2.9 2.4 2.9 3.2
GSM5292183 FALSE 1720 1627 93 5.4 3.0 3.2 0.2 2.2
GSM5292153 FALSE 7785 7373 412 5.3 4.5 3.5 0.9 1.2
GSM5292144 FALSE 13721 13012 709 5.2 3.7 3.1 1.3 2.0
SRR27882088 FALSE 8509 8067 442 5.2 2.2 1.8 0.2 3.6
SRR27882204 FALSE 13518 12846 672 5.0 3.6 2.7 0.8 2.0
GSM5292174 FALSE 7198 6838 360 5.0 3.8 3.1 1.1 1.5
SRR27882152 FALSE 8563 8149 414 4.8 3.6 2.9 0.5 1.5
GSM5292205 FALSE 8031 7654 377 4.7 2.4 1.9 0.9 3.2
SRR27882084 FALSE 7167 6829 338 4.7 2.5 2.3 1.1 3.2
SRR27882173 FALSE 9927 9493 434 4.4 2.8 2.6 1.7 2.2
SRR27882151 FALSE 9167 8772 395 4.3 2.8 2.2 0.4 2.0
GSM5292172 FALSE 5249 5023 226 4.3 1.9 1.7 2.1 1.3
GSM5292166 FALSE 9080 8695 385 4.2 3.1 2.1 0.9 1.7
SRR27882213 FALSE 8590 8244 346 4.0 1.8 1.6 1.8 2.7
SRR27882210 FALSE 5861 5640 221 3.8 1.6 1.6 1.6 2.6
GSM5292145 FALSE 6349 6116 233 3.7 2.9 3.4 0.4 0.0
SRR27882089 FALSE 8051 7762 289 3.6 2.0 1.4 0.2 2.0
SRR27882186 FALSE 9149 8826 323 3.5 2.2 1.8 1.5 1.4
SRR27882202 FALSE 7345 7090 255 3.5 1.8 1.7 1.6 1.2
SRR27882148 FALSE 7712 7452 260 3.4 2.0 1.9 0.7 1.6
SRR27882215 FALSE 6847 6617 230 3.4 1.9 1.7 1.2 1.8
GSM5292202 FALSE 7689 7465 224 2.9 2.0 1.7 0.6 1.0
SRR27882193 FALSE 6295 6120 175 2.8 1.6 1.4 1.1 1.1
GSM5292154 FALSE 6756 6572 184 2.7 2.1 1.7 0.2 0.6
SRR27882078 FALSE 10122 9856 266 2.6 2.1 1.7 0.6 0.9
GSM5292164 FALSE 11633 11341 292 2.5 1.4 1.0 0.1 1.4
GSM5292208 FALSE 8830 8687 143 1.6 1.0 0.7 0.1 0.8

What were the reasons for excluding barcodes?

An upset plot is generated for each sample to visualize the reasons for barcode exclusion. The plot shows the overlap between different exclusion criteria, with vertical bars representing the size of each intersection. Plots are omitted for samples where no barcodes were excluded.

for (bb in b_lvls) {
  qc_tmp      = qc_dt[ batch_var == bb ]
  if (nrow(qc_tmp) == 1 | sum(qc_tmp$keep) == nrow(qc_tmp))
    next
  cat('### ', bb, '\n')
  suppressMessages( print(plot_upset_of_exclusions(qc_tmp, qc_names, qc_lu, cuts_dt)) )
  cat('\n\n')
}

GSM5292146

GSM5292147

GSM5292156

GSM5292157

GSM5292158

GSM5292200

SRR27882183

SRR27882182

SRR27882181

SRR27882171

SRR27882170

SRR27882169

SRR27882094

SRR27882093

SRR27882092

SRR27882091

SRR27882090

SRR27882089

SRR27882088

SRR27882086

SRR27882085

SRR27882084

SRR27882158

SRR27882157

SRR27882156

SRR27882154

SRR27882153

SRR27882152

SRR27882151

SRR27882150

SRR27882149

SRR27882148

GSM5292143

GSM5292144

GSM5292145

GSM5292148

GSM5292149

GSM5292150

GSM5292151

GSM5292152

GSM5292153

GSM5292154

GSM5292155

GSM5292194

GSM5292195

GSM5292196

GSM5292197

GSM5292198

GSM5292199

SRR27882082

SRR27882081

SRR27882080

SRR27882079

SRR27882078

SRR27882077

SRR27882075

SRR27882074

SRR27882073

SRR27882180

SRR27882179

SRR27882178

SRR27882177

SRR27882175

SRR27882174

SRR27882173

SRR27882172

GSM5292174

GSM5292175

GSM5292176

GSM5292177

GSM5292178

GSM5292179

GSM5292180

GSM5292181

GSM5292182

GSM5292183

GSM5292184

GSM5292185

GSM5292186

GSM5292187

GSM5292188

GSM5292193

GSM5292201

SRR27882193

SRR27882192

SRR27882191

SRR27882190

SRR27882189

SRR27882188

SRR27882187

SRR27882186

SRR27882185

SRR27882165

SRR27882164

SRR27882163

SRR27882162

SRR27882161

SRR27882160

SRR27882159

GSM5292164

GSM5292169

GSM5292173

GSM5292190

GSM5292191

GSM5292192

GSM5292202

GSM5292203

GSM5292204

GSM5292205

GSM5292206

GSM5292207

GSM5292208

SRR27882214

SRR27882212

SRR27882203

SRR27882202

SRR27882201

SRR27882200

SRR27882199

SRR27882198

SRR27882197

SRR27882196

SRR27882194

GSM5292159

GSM5292160

GSM5292161

GSM5292162

GSM5292163

GSM5292165

GSM5292166

GSM5292167

GSM5292168

GSM5292170

GSM5292171

GSM5292172

GSM5292189

SRR27882168

SRR27882167

SRR27882166

SRR27882216

SRR27882215

SRR27882213

SRR27882211

SRR27882210

SRR27882209

SRR27882208

SRR27882207

SRR27882205

SRR27882204

R session info

Details of the R package versions used are given below.

devtools::session_info()
## ─ Session info ───────────────────────────────────────────────────────────────
##  setting  value
##  version  R version 4.4.3 (2025-02-28)
##  os       Red Hat Enterprise Linux 8.10 (Ootpa)
##  system   x86_64, linux-gnu
##  ui       X11
##  language (EN)
##  collate  en_US.UTF-8
##  ctype    en_US.UTF-8
##  tz       Europe/Zurich
##  date     2026-03-04
##  pandoc   3.8.2.1 @ /home/macnairw/packages/scprocess/.snakemake/conda/4fef11cadd34f9d2d13a0d6139d09340_/bin/ (via rmarkdown)
##  quarto   NA
## 
## ─ Packages ───────────────────────────────────────────────────────────────────
##  package              * version   date (UTC) lib source
##  abind                * 1.4-8     2024-09-12 [1] CRAN (R 4.4.3)
##  assertthat           * 0.2.1     2019-03-21 [1] CRAN (R 4.4.3)
##  beachmat               2.22.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.3)
##  beeswarm               0.4.0     2021-06-01 [1] CRAN (R 4.4.3)
##  Biobase              * 2.66.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  BiocGenerics         * 0.52.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  BiocIO                 1.16.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  BiocManager            1.30.27   2025-11-14 [1] CRAN (R 4.4.3)
##  BiocNeighbors          2.0.0     2024-10-29 [1] Bioconductor 3.20 (R 4.4.3)
##  BiocParallel         * 1.40.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  BiocSingular           1.22.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.3)
##  BiocStyle            * 2.34.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  Biostrings             2.74.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  bitops                 1.0-9     2024-10-03 [1] CRAN (R 4.4.3)
##  bluster                1.16.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.3)
##  bookdown               0.45      2025-10-03 [1] CRAN (R 4.4.3)
##  bslib                  0.9.0     2025-01-30 [1] CRAN (R 4.4.3)
##  ca                     0.71.1    2020-01-24 [1] CRAN (R 4.4.3)
##  cachem                 1.1.0     2024-05-16 [1] CRAN (R 4.4.3)
##  callr                  3.7.6     2024-03-25 [1] CRAN (R 4.4.3)
##  cellranger             1.1.0     2016-07-27 [1] CRAN (R 4.4.3)
##  circlize             * 0.4.16    2024-02-20 [1] CRAN (R 4.4.3)
##  cli                    3.6.5     2025-04-23 [1] CRAN (R 4.4.3)
##  clue                   0.3-66    2024-11-13 [1] CRAN (R 4.4.3)
##  cluster                2.1.8.1   2025-03-12 [1] CRAN (R 4.4.3)
##  codetools              0.2-20    2024-03-31 [1] CRAN (R 4.4.3)
##  colorspace             2.1-2     2025-09-22 [1] CRAN (R 4.4.3)
##  ComplexHeatmap       * 2.22.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  crayon                 1.5.3     2024-06-20 [1] CRAN (R 4.4.3)
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##  data.table           * 1.17.8    2025-07-10 [1] CRAN (R 4.4.3)
##  DelayedArray         * 0.32.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  devtools               2.4.6     2025-10-03 [1] CRAN (R 4.4.3)
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##  forcats              * 1.0.1     2025-09-25 [1] CRAN (R 4.4.3)
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##  GenomeInfoDb         * 1.42.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  GenomeInfoDbData       1.2.13    2026-02-17 [1] Bioconductor
##  GenomicAlignments      1.42.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.3)
##  GenomicRanges        * 1.58.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  GetoptLong             1.0.5     2020-12-15 [1] CRAN (R 4.4.3)
##  getPass                0.2-4     2023-12-10 [1] CRAN (R 4.4.3)
##  ggbeeswarm           * 0.7.2     2023-04-29 [1] CRAN (R 4.4.3)
##  ggh4x                * 0.3.1     2025-05-30 [1] CRAN (R 4.4.3)
##  ggplot.multistats    * 1.0.1     2024-09-25 [1] CRAN (R 4.4.3)
##  ggplot2              * 4.0.1     2025-11-14 [1] CRAN (R 4.4.3)
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##  git2r                  0.35.0    2024-10-20 [1] CRAN (R 4.4.3)
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##  htmltools              0.5.8.1   2024-04-04 [1] CRAN (R 4.4.3)
##  httpuv                 1.6.16    2025-04-16 [1] CRAN (R 4.4.3)
##  httr                   1.4.7     2023-08-15 [1] CRAN (R 4.4.3)
##  igraph                 2.1.4     2025-01-23 [1] CRAN (R 4.4.3)
##  IRanges              * 2.40.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
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##  jsonlite               2.0.0     2025-03-27 [1] CRAN (R 4.4.3)
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##  magrittr             * 2.0.4     2025-09-12 [1] CRAN (R 4.4.3)
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##  Matrix               * 1.7-4     2025-08-28 [1] CRAN (R 4.4.3)
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##  pkgconfig              2.0.3     2019-09-22 [1] CRAN (R 4.4.3)
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##  ps                     1.9.1     2025-04-12 [1] CRAN (R 4.4.3)
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##  R.methodsS3            1.8.2     2022-06-13 [1] CRAN (R 4.4.3)
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##  R.utils                2.13.0    2025-02-24 [1] CRAN (R 4.4.3)
##  R6                     2.6.1     2025-02-15 [1] CRAN (R 4.4.3)
##  RColorBrewer         * 1.1-3     2022-04-03 [1] CRAN (R 4.4.3)
##  Rcpp                   1.1.0     2025-07-02 [1] CRAN (R 4.4.3)
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##  readr                * 2.1.6     2025-11-14 [1] CRAN (R 4.4.3)
##  readxl               * 1.4.5     2025-03-07 [1] CRAN (R 4.4.3)
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##  rtracklayer            1.66.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.3)
##  S4Arrays             * 1.6.0     2024-10-29 [1] Bioconductor 3.20 (R 4.4.3)
##  S4Vectors            * 0.44.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
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##  scDblFinder          * 1.23.4    2025-08-22 [1] Bioconductor 3.22 (R 4.4.3)
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##  scuttle              * 1.16.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.3)
##  seriation            * 1.5.8     2025-08-20 [1] CRAN (R 4.4.3)
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##  stringi                1.8.7     2025-03-27 [1] CRAN (R 4.4.3)
##  stringr              * 1.6.0     2025-11-04 [1] CRAN (R 4.4.3)
##  SummarizedExperiment * 1.36.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
##  tibble               * 3.3.0     2025-06-08 [1] CRAN (R 4.4.3)
##  tidyr                * 1.3.1     2024-01-24 [1] CRAN (R 4.4.3)
##  tidyselect             1.2.1     2024-03-11 [1] CRAN (R 4.4.3)
##  tidyverse            * 2.0.0     2023-02-22 [1] CRAN (R 4.4.3)
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##  UpSetR               * 1.4.0     2019-05-22 [1] CRAN (R 4.4.3)
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##  viridisLite          * 0.4.2     2023-05-02 [1] CRAN (R 4.4.3)
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##  workflowr            * 1.7.2     2025-08-18 [1] CRAN (R 4.4.3)
##  xfun                   0.54      2025-10-30 [1] CRAN (R 4.4.3)
##  xgboost                3.1.2.1   2026-01-06 [1] local
##  XML                    3.99-0.20 2025-11-08 [1] CRAN (R 4.4.3)
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##  zlibbioc               1.52.0    2024-10-29 [1] Bioconductor 3.20 (R 4.4.2)
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##  * ── Packages attached to the search path.
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