1. Quick Start: miRNA-oriented enrichment analysis with miREA

Introduction

miREA performs microRNA-oriented enrichment analysis by modeling miRNA-gene interaction (MGI) networks directly, rather than collapsing them to miRNA- or gene-level sets. Conventional node-centric enrichment approaches treat miRNAs and genes as independent units, which discards the inherent many-to-many structure of miRNA-gene regulation and can bias functional interpretation. miREA instead models the regulatory edges themselves.

The package implements:

  • Four node-based methods: TG_ORA, TG_Score (target-gene-based), MiR_ORA, MiR_Score (miRNA-based).
  • Five edge-based methods: Edge_ORA, Edge_Score1D, Edge_Score2D, Edge_Topology, Edge_Network, spanning over-representation, scoring-based, topology-aware, and network-propagation approaches.

All nine methods share a common workflow: prepare method-specific input data with getAllInputData(), run the selected methods with miREA(), and visualize results with plotSummary(), plotHeatmap(), or plotHeatmapSankey().

Relation to existing packages

Several Bioconductor/CRAN packages address related but distinct questions. clusterProfiler performs over-representation and gene set enrichment analysis, but for gene enrichment analysis. It tests a list of genes against pathway gene sets, with no special focus on miRNAs. CePa incorporates pathway topology via node centrality measures but operates purely on gene-level and node-level, with no focus on miRNAs as well. multiMiR and similar miRNA-target database packages retrieve curated or predicted miRNA-gene interactions but do not provide an enrichment/significance-testing framework over them. miRNApath performs over-representation analysis by expanding each miRNA-gene pair into its own hypergeometric test unit, using a single ORA-style method. MIRit is a comprehensive, actively maintained framework for integrative miRNA-mRNA analysis, covering differential expression, correlation-based integration, and gene-set enrichment of miRNA targets. Here, we present miREA, to our knowledge the first miRNA-oriented enrichment package that adopts an MGI (miRNA-gene interaction) edge-centric enrichment. It spans over-representation, scoring, topology-aware, and network-propagation methods, and uses expression-informed edge scores rather than a plain membership test to characterize miRNA function at the pathway level. Internally, its ORA and GSEA-style methods (TG_ORA, MiR_ORA, Edge_ORA, TG_Score, MiR_Score, Edge_Score1D) reuse clusterProfiler’s enricher()/GSEA() rather than reimplementing them.

Installation

if (!requireNamespace("BiocManager", quietly = TRUE))
    install.packages("BiocManager")
BiocManager::install("miREA")

Example data

This vignette uses a small, self-contained example dataset bundled with the package (see ?miREA_example_data), derived from the real bladder urothelial carcinoma (BLCA) data used in the miREA manuscript. It covers 16 bladder-cancer-specific MSigDB gene sets and 99 TCGA samples (80 tumor and 19 normal, used for differential expression). The 80 tumor samples are also used for miRNA-gene correlation. For the full datasets used in the manuscript, see the Zenodo deposit.

library(miREA)
data(toy_pathway)
data(toy_gene_DEdata)
data(toy_mir_DEdata)
data(toy_gene_mat)
data(toy_mir_mat)
data(toy_background_MGI)
data(toy_background_GGI)

head(toy_pathway)
#>                                pathway   gene
#> 1 LINDGREN_BLADDER_CANCER_CLUSTER_1_DN  ABCF2
#> 2 LINDGREN_BLADDER_CANCER_CLUSTER_1_DN ABI3BP
#> 3 LINDGREN_BLADDER_CANCER_CLUSTER_1_DN  ACOT7
#> 4 LINDGREN_BLADDER_CANCER_CLUSTER_1_DN  ACOX3
#> 5 LINDGREN_BLADDER_CANCER_CLUSTER_1_DN  ACSF2
#> 6 LINDGREN_BLADDER_CANCER_CLUSTER_1_DN  ACSL1
dim(toy_gene_mat)
#> [1] 34721    80
dim(toy_mir_mat)
#> [1] 995  80

Step 1: Prepare input data

getAllInputData() builds the method-specific input formats (DE gene/miRNA lists, ranked score vectors, MGI dictionaries, GGI networks, etc.) from the raw expression and differential-expression tables. Here we request all nine methods at once. See ?default_method_list for the full list of supported method names.

input_data <- getAllInputData(
  methods = default_method_list,
  pathway = toy_pathway,
  mir_DEdata = toy_mir_DEdata,
  gene_DEdata = toy_gene_DEdata,
  background_MGI = toy_background_MGI,
  background_GGI = toy_background_GGI,
  GGI_source = "Omnipath",
  gene_mat = toy_gene_mat,
  mir_mat = toy_mir_mat,
  scoreFun = "rank"
)
#> Generate all input data...
#>   Start generate expression data...
#>   Start generate pathway data...
#>   Start generate pathway_GGI data...
#>   Start generate GGI data for whole network...
#>   Start generate MGI data for whole network...
#>   Start generate dict_MGI data for permutation...
#> All the input data is ready!
names(input_data)
#> [1] "raw_data"       "background_MGI" "background_GGI" "data"          
#> [5] "pathway"        "pathway_GGI"    "GGI"            "MGI"           
#> [9] "dict_MGI"

Interoperability with standard Bioconductor classes

gene_mat/mir_mat also accept a SummarizedExperiment object in place of a raw matrix/dataframe (the first assay is used), and pathway accepts a GSEABase GeneSetCollection in place of the two-column dataframe. This lets miREA plug directly into analyses that already represent expression data and gene sets in these standard classes, instead of requiring a manual conversion step.

library(SummarizedExperiment)
#> Loading required package: MatrixGenerics
#> Loading required package: matrixStats
#> 
#> Attaching package: 'MatrixGenerics'
#> The following objects are masked from 'package:matrixStats':
#> 
#>     colAlls, colAnyNAs, colAnys, colAvgsPerRowSet, colCollapse,
#>     colCounts, colCummaxs, colCummins, colCumprods, colCumsums,
#>     colDiffs, colIQRDiffs, colIQRs, colLogSumExps, colMadDiffs,
#>     colMads, colMaxs, colMeans2, colMedians, colMins, colOrderStats,
#>     colProds, colQuantiles, colRanges, colRanks, colSdDiffs, colSds,
#>     colSums2, colTabulates, colVarDiffs, colVars, colWeightedMads,
#>     colWeightedMeans, colWeightedMedians, colWeightedSds,
#>     colWeightedVars, rowAlls, rowAnyNAs, rowAnys, rowAvgsPerColSet,
#>     rowCollapse, rowCounts, rowCummaxs, rowCummins, rowCumprods,
#>     rowCumsums, rowDiffs, rowIQRDiffs, rowIQRs, rowLogSumExps,
#>     rowMadDiffs, rowMads, rowMaxs, rowMeans2, rowMedians, rowMins,
#>     rowOrderStats, rowProds, rowQuantiles, rowRanges, rowRanks,
#>     rowSdDiffs, rowSds, rowSums2, rowTabulates, rowVarDiffs, rowVars,
#>     rowWeightedMads, rowWeightedMeans, rowWeightedMedians,
#>     rowWeightedSds, rowWeightedVars
#> Loading required package: GenomicRanges
#> Loading required package: stats4
#> Loading required package: BiocGenerics
#> Loading required package: generics
#> 
#> Attaching package: 'generics'
#> The following objects are masked from 'package:base':
#> 
#>     as.difftime, as.factor, as.ordered, intersect, is.element, setdiff,
#>     setequal, union
#> 
#> Attaching package: 'BiocGenerics'
#> The following objects are masked from 'package:stats':
#> 
#>     IQR, mad, sd, var, xtabs
#> The following object is masked from 'package:utils':
#> 
#>     data
#> The following objects are masked from 'package:base':
#> 
#>     anyDuplicated, aperm, append, as.data.frame, basename, cbind,
#>     colnames, dirname, do.call, duplicated, eval, evalq, Filter, Find,
#>     get, grep, grepl, is.unsorted, lapply, Map, mapply, match, mget,
#>     order, paste, pmax, pmax.int, pmin, pmin.int, Position, rank,
#>     rbind, Reduce, rownames, sapply, saveRDS, scale, sequence, table,
#>     tapply, transform, unique, unsplit, which.max, which.min
#> Loading required package: S4Vectors
#> 
#> Attaching package: 'S4Vectors'
#> The following object is masked from 'package:utils':
#> 
#>     findMatches
#> The following objects are masked from 'package:base':
#> 
#>     expand.grid, I, unname
#> Loading required package: IRanges
#> Loading required package: Seqinfo
#> Loading required package: Biobase
#> Welcome to Bioconductor
#> 
#>     Vignettes contain introductory material; view with
#>     'browseVignettes()'. To cite Bioconductor, see
#>     'citation("Biobase")', and for packages 'citation("pkgname")'.
#> 
#> Attaching package: 'Biobase'
#> The following object is masked from 'package:MatrixGenerics':
#> 
#>     rowMedians
#> The following objects are masked from 'package:matrixStats':
#> 
#>     anyMissing, rowMedians
library(GSEABase)
#> Loading required package: annotate
#> Loading required package: AnnotationDbi
#> Loading required package: XML
#> Loading required package: graph
#> 
#> Attaching package: 'graph'
#> The following object is masked from 'package:XML':
#> 
#>     addNode

gene_se <- SummarizedExperiment(assays = list(expr = as.matrix(toy_gene_mat)))
mir_se <- SummarizedExperiment(assays = list(expr = as.matrix(toy_mir_mat)))
pathway_gsc <- GeneSetCollection(Map(function(genes, name) GeneSet(genes, setName = name),
  split(toy_pathway$gene, toy_pathway$pathway),
  names(split(toy_pathway$gene, toy_pathway$pathway))
))

input_data_bioc <- getAllInputData(
  methods = "Edge_ORA", pathway = pathway_gsc,
  mir_DEdata = toy_mir_DEdata, gene_DEdata = toy_gene_DEdata,
  background_MGI = toy_background_MGI,
  gene_mat = gene_se, mir_mat = mir_se
)
#> Generate all input data...
#>   Start generate expression data...
#>   Start generate pathway data...
#> All the input data is ready!
identical(input_data_bioc$data$Edge_ORA, input_data$data$Edge_ORA)
#> [1] TRUE

Step 2: Run enrichment analysis

miREA() requires a background for each target (TG, MiR, Edge) unless you already supplied one via getAllInputData()’s background_MGI. miREA does not ship a bundled default background. Build one from your own background_MGI/background_GGI, or download the full background used in the manuscript from the Zenodo deposit linked above.

background <- list(
  TG = unique(toy_pathway$gene),
  MiR = unique(input_data$pathway$MiR$miRNA),
  Edge = unique(paste0(toy_background_MGI$miRNA, ":", toy_background_MGI$gene))
)

result <- miREA(
  methods = default_method_list,
  input_data = input_data,
  background = background,
  minSize = NULL,
  maxSize = NULL,
  pvalueType = list(TG = "std", MiR = "std", Edge = "neg"),
  pAdjMethod = "BH",
  pvalueCutoff = 1,
  iter = 20,
  ncores = 1
)
#> Start Enrichment analysis...
#> === Start time: 2026-10-16 21:31:35 ===
#> 
#>   Start TG-ORA analysis ...
#> 
#>   TG-ORA analysis have finished!
#>     - TG_ORA finished, time cost: 0.638999999999999 s.
#> 
#>   Start TG-Score analysis ...
#> Warning in gsea(geneList = geneList, gene_sets = geneSets, weight = weight, :
#> For some pathways, in reality P-values are less than 1e-10. You can set the eps
#> argument to zero for better estimation.
#>   TG-Score analysis have finished!
#>     - TG_Score finished, time cost: 0.483000000000001 s.
#> 
#>   Start MiR-ORA analysis ...
#> 4 of 412 miRNAs are not included in the background. Those are: 
#>  hsa-miR-127-3p hsa-miR-517a-3p hsa-miR-517b-3p hsa-miR-1307-5p 
#>  Those miRNAs are deleted.
#>   MiR-ORA analysis have finished!
#>     - MiR_ORA finished, time cost: 0.0219999999999985 s.
#> 
#>   Start MiR-Score analysis ...
#> Warning in prepare_gsea_inputs(geneList, scoreType, exponent): There are ties
#> in the preranked stats (0.3% of the list). The order of those tied genes will
#> be arbitrary, which may produce unexpected results.
#>   MiR-Score analysis have finished!
#>     - MiR_Score finished, time cost: 0.776 s.
#> 
#>   Start Edge-ORA analysis ...
#>   Edge-ORA analysis have finished!
#>     - Edge_ORA finished, time cost: 0.0800000000000018 s.
#> 
#>   Start Edge-Score1D analysis ...
#> Warning in prepare_gsea_inputs(geneList, scoreType, exponent): There are ties
#> in the preranked stats (0.31% of the list). The order of those tied genes will
#> be arbitrary, which may produce unexpected results.
#> Warning in prepare_gsea_inputs(geneList, scoreType, exponent): For some
#> pathways, in reality P-values are less than 1e-10. You can set the eps argument
#> to zero for better estimation.
#>   Edge-Score1D analysis have finished!
#>     - Edge_Score1D finished, time cost: 25.22 s.
#> 
#>   Start Edge_Score2D analysis ...
#>     Generate random MGI set radius on 1 cores...
#>     Calculate pathway radius and siginificant level...
#>       1 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_1_DN : 0.937
#>       2 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_3_UP : 0.9997
#>       3 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_2B : 0
#>       4 / 16 , WU_SILENCED_BY_METHYLATION_IN_BLADDER_CANCER : 1e-04
#>       5 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_3_DN : 0
#>       6 / 16 , OSMAN_BLADDER_CANCER_UP : 0
#>       7 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_1_UP : 0
#>       8 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_2A_DN : 0
#>       9 / 16 , OSMAN_BLADDER_CANCER_DN : 0.9856
#>       10 / 16 , KEGG_BLADDER_CANCER : 0.9492
#>       11 / 16 , LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q : 0
#>       12 / 16 , LINDGREN_BLADDER_CANCER_HIGH_RECURRENCE : 0
#>       13 / 16 , WP_BLADDER_CANCER : 0
#>       14 / 16 , HP_BLADDER_NEOPLASM : 0.1978
#>       15 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_2A_UP : 0.0184
#>       16 / 16 , WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER : 0.5434
#>   Edge-Score2D analysis have finished!
#>     - Edge_Score2D finished, time cost: 0.745999999999995 s.
#> 
#>   Start Edge-Topology analysis ...
#>     Start analysis on 16 pathways...
#> Warning in Edge_Topology(MGI = input_data$data$Edge_Topology, background = background$Edge, : You are use unparallel computing. It could cause huge amount of time!!!
#>             
#> For parallel computing, please specify [ncores] parameter to be larger than 1 (We highly recommend that)!!!
#>     Calculate pathway scores and siginificant level on 1 cores...
#>       1 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_1_DN : 0.523
#>       2 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_1_UP : 0.8618
#>       3 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_2A_DN : 0.0793
#>       4 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_2A_UP : 1
#>       5 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_2B : 0.0425
#>       6 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_3_DN : 0.7068
#>       7 / 16 , LINDGREN_BLADDER_CANCER_CLUSTER_3_UP : 0.0688
#>       8 / 16 , LINDGREN_BLADDER_CANCER_HIGH_RECURRENCE : 0.8988
#>       9 / 16 , LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q : 0.2517
#>       10 / 16 , OSMAN_BLADDER_CANCER_DN : 0.6807
#>       11 / 16 , OSMAN_BLADDER_CANCER_UP : 0.5325
#>       12 / 16 , WU_SILENCED_BY_METHYLATION_IN_BLADDER_CANCER : 0.879
#>       13 / 16 , KEGG_BLADDER_CANCER : 0.8879
#>       14 / 16 , WP_BLADDER_CANCER : 0.7026
#>       15 / 16 , WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER : 0.2177
#>       16 / 16 , HP_BLADDER_NEOPLASM : 0.2336
#>   Edge-Topology analysis have finished!
#>     - Edge_Topology finished, time cost: 9.943 s.
#> 
#>   Start Edge-Network analysis ...
#>     Build restart list...
#>       Size of sparse restart_matrix: 2.58535003662109 MB.
#> Warning in Edge_Network(DEMGI = input_data$data$Edge_Network, pathway_gene = input_data$pathway$TG, : You are use unparallel computing. It could cause huge amount of time!!!
#>             
#> For parallel computing, please specify [ncores] parameter to be larger than 1 (We highly recommend that)!!!
#>     Build transition matrix...
#>       Size of transition_matrix: 8.83169555664062 MB.
#>     Execute RWR with 20 iterations on 1 cores...
#>     Summarise by pathways...
#>   Edge-Network analysis have finished!
#>     - Edge_Network finished, time cost: 2.396 s.
#> 
#> miREA analysis have finished!...
#> === End time: 2026-10-16 21:32:22 ===
result$valid_methods
#> [1] "TG_ORA"        "TG_Score"      "MiR_ORA"       "MiR_Score"    
#> [5] "Edge_ORA"      "Edge_Score1D"  "Edge_Score2D"  "Edge_Topology"
#> [9] "Edge_Network"

miREA() returns a miREAResult object, a lightweight S4 container for the per-method result tables, elapsed times, and call parameters, with $ access kept for the four fields above for convenience.

result
#> miREAResult object
#>   Methods completed: TG_ORA, TG_Score, MiR_ORA, MiR_Score, Edge_ORA, Edge_Score1D, Edge_Score2D, Edge_Topology, Edge_Network 
#>   Pathways tested: 16

result$result is a named list with one data.frame per method. For example, the TG_ORA result:

head(result$result$TG_ORA)
#>                                                                                                                         pathway
#> OSMAN_BLADDER_CANCER_DN                                                                                 OSMAN_BLADDER_CANCER_DN
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                                                       LINDGREN_BLADDER_CANCER_CLUSTER_3_UP
#> KEGG_BLADDER_CANCER                                                                                         KEGG_BLADDER_CANCER
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                                             LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                                                       LINDGREN_BLADDER_CANCER_CLUSTER_1_DN
#>                                                                                                                     Description
#> OSMAN_BLADDER_CANCER_DN                                                                                 OSMAN_BLADDER_CANCER_DN
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                                                       LINDGREN_BLADDER_CANCER_CLUSTER_3_UP
#> KEGG_BLADDER_CANCER                                                                                         KEGG_BLADDER_CANCER
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                                             LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                                                       LINDGREN_BLADDER_CANCER_CLUSTER_1_DN
#>                                                                 GeneRatio
#> OSMAN_BLADDER_CANCER_DN                                           124/575
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                               95/575
#> KEGG_BLADDER_CANCER                                                14/575
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER     6/575
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                          34/575
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                              105/575
#>                                                                  BgRatio
#> OSMAN_BLADDER_CANCER_DN                                         421/2157
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                            325/2157
#> KEGG_BLADDER_CANCER                                              42/2157
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER  16/2157
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                       115/2157
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                            374/2157
#>                                                                 RichFactor
#> OSMAN_BLADDER_CANCER_DN                                          0.2945368
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                             0.2923077
#> KEGG_BLADDER_CANCER                                              0.3333333
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER  0.3750000
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                        0.2956522
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                             0.2807487
#>                                                                 FoldEnrichment
#> OSMAN_BLADDER_CANCER_DN                                               1.104897
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                                  1.096535
#> KEGG_BLADDER_CANCER                                                   1.250435
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER       1.406739
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                             1.109081
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                                  1.053174
#>                                                                 oddsRatio
#> OSMAN_BLADDER_CANCER_DN                                          1.189575
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                             1.163406
#> KEGG_BLADDER_CANCER                                              1.385027
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER  1.657645
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                        1.164601
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                             1.090445
#>                                                                    zScore
#> OSMAN_BLADDER_CANCER_DN                                         1.4460567
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                            1.1382020
#> KEGG_BLADDER_CANCER                                             0.9879141
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER 0.9842891
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                       0.7246468
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                            0.6817292
#>                                                                    p_value
#> OSMAN_BLADDER_CANCER_DN                                         0.08379445
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                            0.14242841
#> KEGG_BLADDER_CANCER                                             0.20584038
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER 0.23507258
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                       0.26579549
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                            0.26711766
#>                                                                      padj
#> OSMAN_BLADDER_CANCER_DN                                         0.7123137
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                            0.7123137
#> KEGG_BLADDER_CANCER                                             0.7123137
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER 0.7123137
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                       0.7123137
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                            0.7123137
#>                                                                    qvalue
#> OSMAN_BLADDER_CANCER_DN                                         0.7123137
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                            0.7123137
#> KEGG_BLADDER_CANCER                                             0.7123137
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER 0.7123137
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                       0.7123137
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                            0.7123137
#>                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                             geneID
#> OSMAN_BLADDER_CANCER_DN                                         ARIH2/MRPS14/RPS6KA5/EPB41/CYTIP/SKAP1/TUBD1/ZNF852/CSTF3/VAMP1/ZNF398/RAD50/ARFGAP2/UST/MPRIP/HIF1AN/H2AC6/PHF20/SMARCA2/CDK12/MAML2/PML/ELL3/CTDSP2/ZNF763/ARID1A/UBA52/GNAS/GRIPAP1/PPP2CA/ZNF506/MRPS30/PIGL/RPLP2/TRIM38/CELF1/ASXL1/MYO1F/ZNF275/SIAH1/SKI/DIDO1/AHCYL1/NMT2/SSU72/RPL13/ZNF230/DMAC1/NSD3/SENP1/KMT5B/H2BC21/CARD11/ZNF791/ZNF780B/SAMD4B/SERINC3/ZNF493/DIABLO/BACH2/METTL4/ST13/WIPF2/CYB561D1/UBE2B/TMEM63A/TENT4A/LYRM9/WIPI2/CEP57/IDS/RPL15/MRPS25/RPS15/CAMK1D/RPL28/RPL37/ABL2/PRMT2/ZNF544/MARCHF6/ATL2/DPF2/ENOSF1/MLH3/ZNF879/RIPK1/ARHGEF2/GOSR1/RPS11/RHOH/OSBPL2/CLIP1/VWA8/LEPROTL1/TGFBR2/MDM4/FCMR/RABGAP1L/USP36/RPS6/RPS16/RPL13A/ARHGAP45/CHTOP/FLVCR1/STRBP/CELF2/NISCH/CACNA1A/HMGN4/PPM1K/SS18L2/SAFB/GNG7/ASXL2/TTF1/PRUNE1/RASGRF2/ATP8B2/NUDT5/RPL36/KCMF1/MYL12A
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                                                                                                                                                                                                                                               TM4SF1/XPOT/TRIP13/CTPS1/CLPTM1L/ARRB2/SAE1/NEDD9/CSTF3/PRPS2/BIRC5/KPNA4/NXT2/USP14/NCOA6/UCK2/PPM1D/PLK1/SCML1/ATP6V1C1/TYMS/YWHAQ/LIG1/IMPA2/WDHD1/PANK1/ZWINT/C5orf22/KIF14/OSBP/BRCA1/RBBP4/ACOT2/AKAP12/NEMP1/ZNF148/TWSG1/MELK/TCF19/CENPA/AK2/PSMC4/TK1/CKS1B/CHAF1A/CDC25A/GGCT/GMNN/SMIM29/RPA3/KIF11/RACGAP1/PXMP4/GPATCH11/POP4/SLFN13/TARS1/RBBP7/TXNRD1/PARD6B/SLC25A40/E2F1/SPATA13/DAB2/ZYG11B/OTULIN/SLC39A8/WDR53/RAD1/PAK1IP1/NAA50/NCAPG2/ZNF587/RRP1/FAM200B/EPHX1/NQO2/CENPN/TFRC/H2BC5/BRIX1/ITPR1/LNPK/DEK/MCM8/CDC6/KIF2C/TTC39A/DBF4/NEU1/GCA/MOBP/TSC22D2/RBM38/ASS1
#> KEGG_BLADDER_CANCER                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                 RPS6KA5/TP53/FGFR3/HRAS/DAPK3/EGFR/PGF/E2F1/CDKN2A/E2F3/RASSF1/MAP2K2/EGF/E2F2
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                   GCC2/MYRIP/SYTL2/SYTL5/MLPH/MADD
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                       DOCK8/SPTLC1/FUBP3/ABL1/SMARCA2/NUP214/DUSP22/MTAP/RNF38/RBM18/TOPORS/ASTN2/ANP32B/ANK2/CDK9/TOR1B/PPP6C/TGFBR1/RAD23B/PSMD5/ACO1/CREBL2/ANKH/SLC35D2/TDRP/MSANTD3/CUL5/SETX/VPS13A/KLHL9/RARA/TNFRSF10C/ASS1/APTX
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                                                                                                                                                                               ILF3/TRIP13/CTPS1/ARRB2/SAE1/TMEM108/NEDD9/FYN/CACUL1/BIRC5/RASSF5/MAN1C1/ST3GAL2/ABCF2/UCK2/ENO2/IDO1/PLK1/PRRC1/NSD2/PRSS8/SUSD6/WDHD1/ZWINT/CELSR3/VASN/KIF14/BRCA1/KLHL6/AKAP12/ZNF148/CLN3/CD80/MELK/TCF19/SNX5/SSR1/CENPA/PSMA1/PALM2AKAP2/TK1/CKS1B/CDC25A/HILPDA/VPS37C/CDK9/NCKIPSD/KIF11/RACGAP1/H1-0/RELL1/RAI14/CLIC4/SLC44A2/JPT1/MAP3K14/CERS2/E2F1/SPATA13/MACROH2A1/DAB2/CGN/ZNF703/CDC25B/NCAPG2/HMGCS1/RRP1/ABI3BP/YPEL3/HDGF/EPHX1/PEA15/ERBB3/DLGAP4/SBNO2/CENPN/HEYL/TFRC/TIMP2/CDCA7/LCOR/RBM28/POLE/NUDT1/GFOD1/MCM8/CDC6/KIF2C/TTC39A/TJP3/NEU1/SH3BP5/DYRK2/NAV2/SFMBT1/ADAM15/MOBP/GNL3L/APLNR/STAT5B/PRUNE1/EBF1/PITX2/TRIM11/SLC1A3
#>                                                                 Count
#> OSMAN_BLADDER_CANCER_DN                                           124
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP                               95
#> KEGG_BLADDER_CANCER                                                14
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER     6
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q                          34
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN                              105

All per-method results can also be flattened into a single dataframe (with a method column) via as.data.frame():

head(as.data.frame(result))
#>                                                                     method
#> OSMAN_BLADDER_CANCER_DN...1                                         TG_ORA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                            TG_ORA
#> KEGG_BLADDER_CANCER...3                                             TG_ORA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4 TG_ORA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                       TG_ORA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                            TG_ORA
#>                                                                                                                             pathway
#> OSMAN_BLADDER_CANCER_DN...1                                                                                 OSMAN_BLADDER_CANCER_DN
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                                       LINDGREN_BLADDER_CANCER_CLUSTER_3_UP
#> KEGG_BLADDER_CANCER...3                                                                                         KEGG_BLADDER_CANCER
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4 WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                             LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                                       LINDGREN_BLADDER_CANCER_CLUSTER_1_DN
#>                                                                                                                         Description
#> OSMAN_BLADDER_CANCER_DN...1                                                                                 OSMAN_BLADDER_CANCER_DN
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                                       LINDGREN_BLADDER_CANCER_CLUSTER_3_UP
#> KEGG_BLADDER_CANCER...3                                                                                         KEGG_BLADDER_CANCER
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4 WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                             LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                                       LINDGREN_BLADDER_CANCER_CLUSTER_1_DN
#>                                                                     GeneRatio
#> OSMAN_BLADDER_CANCER_DN...1                                           124/575
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                               95/575
#> KEGG_BLADDER_CANCER...3                                                14/575
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4     6/575
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                          34/575
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                              105/575
#>                                                                      BgRatio
#> OSMAN_BLADDER_CANCER_DN...1                                         421/2157
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                            325/2157
#> KEGG_BLADDER_CANCER...3                                              42/2157
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4  16/2157
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                       115/2157
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                            374/2157
#>                                                                     RichFactor
#> OSMAN_BLADDER_CANCER_DN...1                                          0.2945368
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                             0.2923077
#> KEGG_BLADDER_CANCER...3                                              0.3333333
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4  0.3750000
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                        0.2956522
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                             0.2807487
#>                                                                     FoldEnrichment
#> OSMAN_BLADDER_CANCER_DN...1                                               1.104897
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                  1.096535
#> KEGG_BLADDER_CANCER...3                                                   1.250435
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4       1.406739
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                             1.109081
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                  1.053174
#>                                                                     oddsRatio
#> OSMAN_BLADDER_CANCER_DN...1                                          1.189575
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                             1.163406
#> KEGG_BLADDER_CANCER...3                                              1.385027
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4  1.657645
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                        1.164601
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                             1.090445
#>                                                                        zScore
#> OSMAN_BLADDER_CANCER_DN...1                                         1.4460567
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                            1.1382020
#> KEGG_BLADDER_CANCER...3                                             0.9879141
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4 0.9842891
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                       0.7246468
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                            0.6817292
#>                                                                        p_value
#> OSMAN_BLADDER_CANCER_DN...1                                         0.08379445
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                            0.14242841
#> KEGG_BLADDER_CANCER...3                                             0.20584038
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4 0.23507258
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                       0.26579549
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                            0.26711766
#>                                                                          padj
#> OSMAN_BLADDER_CANCER_DN...1                                         0.7123137
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                            0.7123137
#> KEGG_BLADDER_CANCER...3                                             0.7123137
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4 0.7123137
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                       0.7123137
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                            0.7123137
#>                                                                        qvalue
#> OSMAN_BLADDER_CANCER_DN...1                                         0.7123137
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                            0.7123137
#> KEGG_BLADDER_CANCER...3                                             0.7123137
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4 0.7123137
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                       0.7123137
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                            0.7123137
#>                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                 geneID
#> OSMAN_BLADDER_CANCER_DN...1                                         ARIH2/MRPS14/RPS6KA5/EPB41/CYTIP/SKAP1/TUBD1/ZNF852/CSTF3/VAMP1/ZNF398/RAD50/ARFGAP2/UST/MPRIP/HIF1AN/H2AC6/PHF20/SMARCA2/CDK12/MAML2/PML/ELL3/CTDSP2/ZNF763/ARID1A/UBA52/GNAS/GRIPAP1/PPP2CA/ZNF506/MRPS30/PIGL/RPLP2/TRIM38/CELF1/ASXL1/MYO1F/ZNF275/SIAH1/SKI/DIDO1/AHCYL1/NMT2/SSU72/RPL13/ZNF230/DMAC1/NSD3/SENP1/KMT5B/H2BC21/CARD11/ZNF791/ZNF780B/SAMD4B/SERINC3/ZNF493/DIABLO/BACH2/METTL4/ST13/WIPF2/CYB561D1/UBE2B/TMEM63A/TENT4A/LYRM9/WIPI2/CEP57/IDS/RPL15/MRPS25/RPS15/CAMK1D/RPL28/RPL37/ABL2/PRMT2/ZNF544/MARCHF6/ATL2/DPF2/ENOSF1/MLH3/ZNF879/RIPK1/ARHGEF2/GOSR1/RPS11/RHOH/OSBPL2/CLIP1/VWA8/LEPROTL1/TGFBR2/MDM4/FCMR/RABGAP1L/USP36/RPS6/RPS16/RPL13A/ARHGAP45/CHTOP/FLVCR1/STRBP/CELF2/NISCH/CACNA1A/HMGN4/PPM1K/SS18L2/SAFB/GNG7/ASXL2/TTF1/PRUNE1/RASGRF2/ATP8B2/NUDT5/RPL36/KCMF1/MYL12A
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                                                                                                                                                                                                                               TM4SF1/XPOT/TRIP13/CTPS1/CLPTM1L/ARRB2/SAE1/NEDD9/CSTF3/PRPS2/BIRC5/KPNA4/NXT2/USP14/NCOA6/UCK2/PPM1D/PLK1/SCML1/ATP6V1C1/TYMS/YWHAQ/LIG1/IMPA2/WDHD1/PANK1/ZWINT/C5orf22/KIF14/OSBP/BRCA1/RBBP4/ACOT2/AKAP12/NEMP1/ZNF148/TWSG1/MELK/TCF19/CENPA/AK2/PSMC4/TK1/CKS1B/CHAF1A/CDC25A/GGCT/GMNN/SMIM29/RPA3/KIF11/RACGAP1/PXMP4/GPATCH11/POP4/SLFN13/TARS1/RBBP7/TXNRD1/PARD6B/SLC25A40/E2F1/SPATA13/DAB2/ZYG11B/OTULIN/SLC39A8/WDR53/RAD1/PAK1IP1/NAA50/NCAPG2/ZNF587/RRP1/FAM200B/EPHX1/NQO2/CENPN/TFRC/H2BC5/BRIX1/ITPR1/LNPK/DEK/MCM8/CDC6/KIF2C/TTC39A/DBF4/NEU1/GCA/MOBP/TSC22D2/RBM38/ASS1
#> KEGG_BLADDER_CANCER...3                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                 RPS6KA5/TP53/FGFR3/HRAS/DAPK3/EGFR/PGF/E2F1/CDKN2A/E2F3/RASSF1/MAP2K2/EGF/E2F2
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                   GCC2/MYRIP/SYTL2/SYTL5/MLPH/MADD
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                       DOCK8/SPTLC1/FUBP3/ABL1/SMARCA2/NUP214/DUSP22/MTAP/RNF38/RBM18/TOPORS/ASTN2/ANP32B/ANK2/CDK9/TOR1B/PPP6C/TGFBR1/RAD23B/PSMD5/ACO1/CREBL2/ANKH/SLC35D2/TDRP/MSANTD3/CUL5/SETX/VPS13A/KLHL9/RARA/TNFRSF10C/ASS1/APTX
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                                                                                                                                                               ILF3/TRIP13/CTPS1/ARRB2/SAE1/TMEM108/NEDD9/FYN/CACUL1/BIRC5/RASSF5/MAN1C1/ST3GAL2/ABCF2/UCK2/ENO2/IDO1/PLK1/PRRC1/NSD2/PRSS8/SUSD6/WDHD1/ZWINT/CELSR3/VASN/KIF14/BRCA1/KLHL6/AKAP12/ZNF148/CLN3/CD80/MELK/TCF19/SNX5/SSR1/CENPA/PSMA1/PALM2AKAP2/TK1/CKS1B/CDC25A/HILPDA/VPS37C/CDK9/NCKIPSD/KIF11/RACGAP1/H1-0/RELL1/RAI14/CLIC4/SLC44A2/JPT1/MAP3K14/CERS2/E2F1/SPATA13/MACROH2A1/DAB2/CGN/ZNF703/CDC25B/NCAPG2/HMGCS1/RRP1/ABI3BP/YPEL3/HDGF/EPHX1/PEA15/ERBB3/DLGAP4/SBNO2/CENPN/HEYL/TFRC/TIMP2/CDCA7/LCOR/RBM28/POLE/NUDT1/GFOD1/MCM8/CDC6/KIF2C/TTC39A/TJP3/NEU1/SH3BP5/DYRK2/NAV2/SFMBT1/ADAM15/MOBP/GNL3L/APLNR/STAT5B/PRUNE1/EBF1/PITX2/TRIM11/SLC1A3
#>                                                                     Count
#> OSMAN_BLADDER_CANCER_DN...1                                           124
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                               95
#> KEGG_BLADDER_CANCER...3                                                14
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4     6
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                          34
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                              105
#>                                                                     setSize
#> OSMAN_BLADDER_CANCER_DN...1                                              NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                 NA
#> KEGG_BLADDER_CANCER...3                                                  NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4      NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                            NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                 NA
#>                                                                     enrichmentScore
#> OSMAN_BLADDER_CANCER_DN...1                                                      NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                         NA
#> KEGG_BLADDER_CANCER...3                                                          NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4              NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                    NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                         NA
#>                                                                     NES rank
#> OSMAN_BLADDER_CANCER_DN...1                                          NA   NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                             NA   NA
#> KEGG_BLADDER_CANCER...3                                              NA   NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4  NA   NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                        NA   NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                             NA   NA
#>                                                                     leading_edge
#> OSMAN_BLADDER_CANCER_DN...1                                                 <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                    <NA>
#> KEGG_BLADDER_CANCER...3                                                     <NA>
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4         <NA>
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                               <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                    <NA>
#>                                                                     core_enrichment
#> OSMAN_BLADDER_CANCER_DN...1                                                    <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                       <NA>
#> KEGG_BLADDER_CANCER...3                                                        <NA>
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4            <NA>
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                  <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                       <NA>
#>                                                                     log2err
#> OSMAN_BLADDER_CANCER_DN...1                                              NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                 NA
#> KEGG_BLADDER_CANCER...3                                                  NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4      NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                            NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                 NA
#>                                                                     MGISize
#> OSMAN_BLADDER_CANCER_DN...1                                              NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                 NA
#> KEGG_BLADDER_CANCER...3                                                  NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4      NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                            NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                 NA
#>                                                                     score
#> OSMAN_BLADDER_CANCER_DN...1                                            NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                               NA
#> KEGG_BLADDER_CANCER...3                                                NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4    NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                          NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                               NA
#>                                                                     rand_score_dist
#> OSMAN_BLADDER_CANCER_DN...1                                                    <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                       <NA>
#> KEGG_BLADDER_CANCER...3                                                        <NA>
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4            <NA>
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                  <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                       <NA>
#>                                                                     nMoreExtreme
#> OSMAN_BLADDER_CANCER_DN...1                                                   NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                      NA
#> KEGG_BLADDER_CANCER...3                                                       NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4           NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                 NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                      NA
#>                                                                     TopMGI
#> OSMAN_BLADDER_CANCER_DN...1                                           <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                              <NA>
#> KEGG_BLADDER_CANCER...3                                               <NA>
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4   <NA>
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                         <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                              <NA>
#>                                                                     n_MGI nPerm
#> OSMAN_BLADDER_CANCER_DN...1                                            NA    NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                               NA    NA
#> KEGG_BLADDER_CANCER...3                                                NA    NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4    NA    NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                          NA    NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                               NA    NA
#>                                                                      z
#> OSMAN_BLADDER_CANCER_DN...1                                         NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                            NA
#> KEGG_BLADDER_CANCER...3                                             NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4 NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                       NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                            NA
#>                                                                     contribute_edge
#> OSMAN_BLADDER_CANCER_DN...1                                                    <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                       <NA>
#> KEGG_BLADDER_CANCER...3                                                        <NA>
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4            <NA>
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                  <NA>
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                       <NA>
#>                                                                     n_contribute_edge
#> OSMAN_BLADDER_CANCER_DN...1                                                        NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_3_UP...2                                           NA
#> KEGG_BLADDER_CANCER...3                                                            NA
#> WP_DEREGULATION_OF_RAB_AND_RAB_EFFECTOR_GENES_IN_BLADDER_CANCER...4                NA
#> LINDGREN_BLADDER_CANCER_WITH_LOH_IN_CHR9Q...5                                      NA
#> LINDGREN_BLADDER_CANCER_CLUSTER_1_DN...6                                           NA

Each method only reports pathways with pvalue and padj below pvalueCutoff. We use pvalueCutoff = 1 throughout this vignette so every pathway is returned. The exception is the ORA methods (TG_ORA, MiR_ORA, Edge_ORA). These methods only report pathways with at least one overlapping differentially expressed gene, miRNA, or edge, so some pathways can still be missing even at pvalueCutoff = 1. For example, Edge_ORA returns 9 of 16 pathways here.

Step 3: Visualize results

plotSummary() compares results across all requested methods: positive enrichment rates, a PCA of per-method pathway rankings, pairwise overlap between methods, and a p-value density plot.

plotSummary(result, penrichCutoff = 0.05)

#>   Visualization finished!
#> ***If you want to save them to pdf files directly, please specify the [plot_path] parameter!***
#> $summary

#> 
#> $pvalue

plotHeatmap() and plotHeatmapSankey() visualize a single edge-based method’s result as a miRNA-by-gene heatmap (optionally combined with a Sankey diagram showing the miRNA-gene-pathway flow). Only the five edge-based methods are supported, one at a time.

plotHeatmap(method = "Edge_Score1D", result = result, input_data = input_data,
             penrichCutoff = 0.05, n_pathway = 5, n_mir_heatmap = 10)

#>   Visualization finished!
#> ***If you want to save them to pdf files, please specify the [plot_path] parameter!***
#> [[1]]

#> 
#> [[2]]
#> [[2]][[1]]
#> A single legend
#> 
#> [[2]][[2]]
#> A single legend
#> 
#> [[2]][[3]]
#> A single legend
#> 
#> [[2]][[4]]
#> A single legend
#> 
#> [[2]][[5]]
#> A single legend
#> 
#> [[2]][[6]]
#> A single legend
plotHeatmapSankey(method = "Edge_Score1D", result = result, input_data = input_data,
                     penrichCutoff = 0.05, n_pathway = 5, n_mir_heatmap = 10,
                     n_mir_sankey = 5)

#>   Visualization finished!
#> ***If you want to save them to pdf files, please specify the [plot_path] parameter!***
#> [[1]]

#> 
#> [[2]]
#> [[2]][[1]]
#> A single legend
#> 
#> [[2]][[2]]
#> A single legend
#> 
#> [[2]][[3]]
#> A single legend
#> 
#> [[2]][[4]]
#> A single legend
#> 
#> [[2]][[5]]
#> A single legend
#> 
#> [[2]][[6]]
#> A single legend

Choosing methods for your own data

miREA supports 4 node-based and 5 edge-based methods. You do not need to run all nine. Select the method(s) most appropriate for your dataset and research question. See the method performance comparisons in the miREA manuscript (miREA’s companion bioRxiv preprint) for guidance. Edge_Topology in particular can be computationally expensive on large, real-sized pathway/background data. Consider excluding it or increasing ncores for large analyses.

Session info

sessionInfo()
#> R version 4.6.1 (2026-06-24)
#> Platform: x86_64-pc-linux-gnu
#> Running under: Ubuntu 26.04.1 LTS
#> 
#> Matrix products: default
#> BLAS:   /usr/lib/x86_64-linux-gnu/openblas-pthread/libblas.so.3 
#> LAPACK: /usr/lib/x86_64-linux-gnu/openblas-pthread/libopenblasp-r0.3.32.so;  LAPACK version 3.12.0
#> 
#> locale:
#>  [1] LC_CTYPE=en_US.UTF-8       LC_NUMERIC=C              
#>  [3] LC_TIME=en_US.UTF-8        LC_COLLATE=en_US.UTF-8    
#>  [5] LC_MONETARY=en_US.UTF-8    LC_MESSAGES=en_US.UTF-8   
#>  [7] LC_PAPER=en_US.UTF-8       LC_NAME=C                 
#>  [9] LC_ADDRESS=C               LC_TELEPHONE=C            
#> [11] LC_MEASUREMENT=en_US.UTF-8 LC_IDENTIFICATION=C       
#> 
#> time zone: Etc/UTC
#> tzcode source: system (glibc)
#> 
#> attached base packages:
#> [1] stats4    stats     graphics  grDevices utils     datasets  methods  
#> [8] base     
#> 
#> other attached packages:
#>  [1] GSEABase_1.75.0             graph_1.91.0               
#>  [3] annotate_1.91.0             XML_3.99-0.25              
#>  [5] AnnotationDbi_1.75.2        SummarizedExperiment_1.43.0
#>  [7] Biobase_2.73.2              GenomicRanges_1.65.4       
#>  [9] Seqinfo_1.3.2               IRanges_2.47.5             
#> [11] S4Vectors_0.51.10           BiocGenerics_0.59.12       
#> [13] generics_0.1.4              MatrixGenerics_1.25.0      
#> [15] matrixStats_1.5.0           miREA_0.99.2               
#> [17] BiocStyle_2.41.0           
#> 
#> loaded via a namespace (and not attached):
#>   [1] RColorBrewer_1.1-3      sys_3.4.3               jsonlite_2.0.0         
#>   [4] shape_1.4.6.1           tidydr_0.0.7            magrittr_2.0.5         
#>   [7] ggtangle_0.1.4          farver_2.1.2            rmarkdown_2.32         
#>  [10] GlobalOptions_0.1.4     fs_2.1.0                vctrs_0.7.3            
#>  [13] memoise_2.0.1           ggtree_4.3.0            htmltools_0.5.9        
#>  [16] S4Arrays_1.13.2         BiocBaseUtils_1.15.1    SparseArray_1.13.4     
#>  [19] gridGraphics_0.5-1      sass_0.4.10             bslib_0.12.0           
#>  [22] htmlwidgets_1.6.4       plyr_1.8.9              httr2_1.3.0            
#>  [25] cachem_1.1.0            buildtools_1.0.0        igraph_2.3.4           
#>  [28] lifecycle_1.0.5         iterators_1.0.14        pkgconfig_2.0.3        
#>  [31] gson_0.2.2              Matrix_1.7-6            R6_2.6.1               
#>  [34] fastmap_1.2.0           clue_0.3-68             digest_0.6.39          
#>  [37] aplot_0.3.2             enrichplot_1.99.8       colorspace_2.1-4       
#>  [40] ggnewscale_0.5.2        patchwork_1.3.2         aisdk_1.4.12           
#>  [43] ps_1.9.3                RSQLite_3.53.3          labeling_0.4.3         
#>  [46] polyclip_1.10-7         httr_1.4.9              abind_1.4-8            
#>  [49] compiler_4.6.1          withr_3.0.3             bit64_4.8.6            
#>  [52] fontquiver_0.2.1        doParallel_1.0.17       S7_0.2.2               
#>  [55] DBI_1.3.0               ggforce_0.5.0           MASS_7.3-66            
#>  [58] rappdirs_0.3.4          DelayedArray_0.39.8     rjson_0.2.23           
#>  [61] tools_4.6.1             otel_0.2.0              scatterpie_0.2.6       
#>  [64] ape_5.8-1               glue_1.8.1              callr_3.8.0            
#>  [67] nlme_3.1-171            GOSemSim_2.39.4         grid_4.6.1             
#>  [70] cluster_2.1.8.3         reshape2_1.4.5          gtable_0.3.6           
#>  [73] tidyr_1.3.2             data.table_1.18.6.1     XVector_0.53.0         
#>  [76] ggrepel_0.9.8           foreach_1.5.2           pillar_1.11.1          
#>  [79] stringr_1.6.0           yulab.utils_0.2.5       splines_4.6.1          
#>  [82] circlize_0.4.18         tweenr_2.0.3            dplyr_1.2.1            
#>  [85] treeio_1.37.1           lattice_0.23-1          bit_4.6.0              
#>  [88] tidyselect_1.2.1        fontLiberation_0.1.0    GO.db_3.23.1           
#>  [91] ComplexHeatmap_2.29.0   maketools_1.3.2         Biostrings_2.81.9      
#>  [94] knitr_1.52              fontBitstreamVera_0.1.1 RhpcBLASctl_0.23-42    
#>  [97] xfun_0.61               stringi_1.8.9           lazyeval_0.2.3         
#> [100] ggfun_0.2.2             yaml_2.3.12             evaluate_1.0.5         
#> [103] codetools_0.2-20        qvalue_2.45.0           gdtools_0.5.2          
#> [106] tibble_3.3.1            BiocManager_1.30.27     ggplotify_0.1.4        
#> [109] cli_3.6.6               systemfonts_1.3.2       xtable_1.8-8           
#> [112] processx_3.9.0          jquerylib_0.1.4         Rcpp_1.1.2             
#> [115] png_0.1-9               parallel_4.6.1          ggplot2_4.0.3          
#> [118] blob_1.3.0              clusterProfiler_4.21.3  ggalluvial_0.12.6      
#> [121] DOSE_4.7.3              tidytree_0.4.8          ggiraph_0.9.6          
#> [124] enrichit_0.2.6          scales_1.4.0            purrr_1.2.2            
#> [127] crayon_1.5.3            GetoptLong_1.1.1        rlang_1.3.0            
#> [130] KEGGREST_1.53.6