From ad92aa358adbc7e14041392f39de141a38da57b5 Mon Sep 17 00:00:00 2001 From: Charles Plessy Date: Fri, 24 May 2024 09:18:43 +0900 Subject: [PATCH] Totally replace clusterCTSS() with distclu() and paraclu() The expression filtering is now done by `filterLowExpCTSS` because 1) it has consequences beyond tag cluster generation and 2) because it simplifies the argument list of the clustering functions. The remaining `clusterCTSS` function had a set of mutually exclusive arguments for `distclu` and `paraclu`; I find it easier to study and use the two methods when they are taken care of by different functions. --- NAMESPACE | 3 +- NEWS.md | 4 + R/AggregationMethods.R | 6 +- R/CAGEexp.R | 3 +- R/CAGEr.R | 88 +++++++++++---- R/ClusteringMethods.R | 151 ------------------------- R/Distclu.R | 57 ++++++++-- R/ExpressionProfilingMethods.R | 4 +- R/GetMethods.R | 9 +- R/Paraclu.R | 55 ++++++++- data/exampleCAGEexp.RData | Bin 164404 -> 163376 bytes man/CTSScoordinates.Rd | 4 +- man/CTSScumulativesTagClusters.Rd | 3 +- man/CTSStoGenes.Rd | 3 +- man/CustomConsensusClusters.Rd | 6 +- man/aggregateTagClusters.Rd | 8 +- man/annotateCTSS.Rd | 3 +- man/clusterCTSS.Rd | 159 --------------------------- man/consensusClusters.Rd | 3 +- man/consensusClustersDESeq2.Rd | 3 +- man/cumulativeCTSSdistribution.Rd | 6 +- man/distclu.Rd | 58 +++++++++- man/dot-ctss_summary_for_clusters.Rd | 47 ++++++++ man/exampleCAGEexp.Rd | 3 +- man/flagLowExpCTSS.Rd | 89 +++++++++++++++ man/getCTSS.Rd | 3 +- man/getExpressionProfiles.Rd | 2 +- man/normalizeTagCount.Rd | 3 +- man/paraclu.Rd | 68 +++++++++++- man/plotInterquantileWidth.Rd | 3 +- man/quantilePositions.Rd | 6 +- man/quickEnhancers.Rd | 3 +- man/resetCAGEexp.Rd | 3 +- man/summariseChrExpr.Rd | 3 +- man/tagClusters.Rd | 5 +- vignettes/CAGEexp.Rmd | 50 +++++---- 36 files changed, 516 insertions(+), 408 deletions(-) delete mode 100644 man/clusterCTSS.Rd create mode 100644 man/dot-ctss_summary_for_clusters.Rd create mode 100644 man/flagLowExpCTSS.Rd diff --git a/NAMESPACE b/NAMESPACE index e8763c4..0788b49 100644 --- a/NAMESPACE +++ b/NAMESPACE @@ -31,7 +31,6 @@ export(aggregateTagClusters) export(annotateCTSS) export(annotateConsensusClusters) export(annotateTagClusters) -export(clusterCTSS) export(consensusClustersDESeq2) export(consensusClustersGR) export(consensusClustersSE) @@ -40,8 +39,10 @@ export(cumulativeCTSSdistribution) export(distclu) export(exportToTrack) export(expressionClasses) +export(filterLowExpCTSS) export(findStrandInvaders) export(flagByUpstreamSequences) +export(flagLowExpCTSS) export(genomeName) export(getCTSS) export(getExpressionProfiles) diff --git a/NEWS.md b/NEWS.md index 29e4b4c..c827c5d 100644 --- a/NEWS.md +++ b/NEWS.md @@ -18,10 +18,14 @@ BACKWARDS-INCOMPATIBLE CHANGES standard behavior. - In cluster objects, the dominant CTSS score is now stored in the `dominantCTSS` object directly. +- The `clusterCTSS` function is replaced by the new `paraclu` and `distclu` + function. CTSS filtering is done beforehand with the new `filterLowExpCTSS` + function. BUG FIXES - The `importPublicData` function was repaired for FANTOM samples. +- CTSS filtering now works correctly with `threshold = 0, thresholdIsTpm = TRUE`. NEW FEATURES diff --git a/R/AggregationMethods.R b/R/AggregationMethods.R index 3d64e65..f3ace72 100644 --- a/R/AggregationMethods.R +++ b/R/AggregationMethods.R @@ -32,7 +32,7 @@ #' @param nrCores Number of cores to use when `useMulticore = TRUE`. Default #' (`NULL`) uses all detected cores. #' -#' @details Since the tag clusters (TCs) returned by the [`clusterCTSS`] +#' @details Since the tag clusters (TCs) returned by the CTSS clustering functions #' function are constructed separately for every CAGE sample within the CAGEr #' object, they can differ between samples in both their number, genomic #' coordinates, position of dominant TSS and overall signal. To be able to @@ -122,7 +122,7 @@ setMethod( "aggregateTagClusters", "CAGEr" .aggregateTagClustersGRL(gr.list = TC.list, CAGEexp_obj = object, maxDist = maxDist) if (excludeSignalBelowThreshold) { - filter <- .filterCtss( object + filter <- flagLowExpCTSS( object , threshold = tpmThreshold , nrPassThreshold = 1 , thresholdIsTpm = TRUE) @@ -244,7 +244,7 @@ setMethod( "CustomConsensusClusters", c("CAGEexp", "GRanges") clusters <- .ConsensusClusters(clusters) - filter <- .filterCtss( object + filter <- flagLowExpCTSS( object , threshold = threshold , nrPassThreshold = nrPassThreshold , thresholdIsTpm = thresholdIsTpm) diff --git a/R/CAGEexp.R b/R/CAGEexp.R index c9527bb..6244aab 100644 --- a/R/CAGEexp.R +++ b/R/CAGEexp.R @@ -174,7 +174,8 @@ setMethod( "initialize", "CAGEexp" #' CTSStoGenes() |> #' normalizeTagCount() |> #' getExpressionProfiles("CTSS") |> -#' clusterCTSS() |> +#' filterLowExpCTSS() |> +#' distclu() |> #' annotateTagClusters(exampleZv9_annot) |> #' cumulativeCTSSdistribution("tagClusters") |> #' quantilePositions("tagClusters") |> diff --git a/R/CAGEr.R b/R/CAGEr.R index 50c2cdd..f914572 100644 --- a/R/CAGEr.R +++ b/R/CAGEr.R @@ -159,41 +159,85 @@ setMethod("validSamples", "CAGEr", function (object, x){ }) -#' @name .filterCtss -#' @noRd -#' @param threshold,nrPassThreshold Only CTSSs with signal \code{>= threshold} in -#' \code{>= nrPassThreshold} experiments will be used for clustering and will -#' contribute towards total signal of the cluster. -#' @param thresholdIsTpm Logical, is threshold raw tag count value (FALSE) or -#' normalized signal (TRUE). -#' @title Private function -#' @details Check if a vector of strings or numbers can be used to identify a sample. +#' Flag CTSSes based on sample expression +#' +#' Flag CTSSes for that do not pass an expression threshold in at least a given +#' number of samples. This is typically used to ignore CTSSes that have been +#' seen only once in a single sample, as they can be considered to not be +#' reproduced. +#' +#' @param object An object from the _CAGEr_ package that contains expression +#' values for multiple samples. +#' +#' @param threshold Flag CTSSs with signal `< threshold`. +#' +#' @param nrPassThreshold Only flag CTSSs when signal is below threshold in at +#' least `nrPassThreshold` samples. +#' +#' @param thresholdIsTpm Logical, is threshold raw tag count value (`FALSE`) or +#' normalized signal (`TRUE`). +#' +#' @returns `flagLowExpCTSS` returns a [`Rle`] vector where `TRUE` indicates the +#' index of a CTSS that passes the filter. +#' +#' @export +#' +#' @examples +#' flagLowExpCTSS(exampleCAGEexp, threshold = 100, nrPassThreshold = 2) -setGeneric(".filterCtss", function( object - , threshold = 0 - , nrPassThreshold = 1 - , thresholdIsTpm = TRUE) { - if (threshold == 0) return(Rle(TRUE)) - standardGeneric(".filterCtss") -}) +setGeneric("flagLowExpCTSS", function( object + , threshold = 1 + , nrPassThreshold = 1 + , thresholdIsTpm = TRUE) + standardGeneric("flagLowExpCTSS") +) -setMethod(".filterCtss", "CAGEr", function (object, threshold, nrPassThreshold, thresholdIsTpm) { - .filterCtss(CTSStagCountSE(object), threshold, nrPassThreshold, thresholdIsTpm) +#' @rdname flagLowExpCTSS + +setMethod("flagLowExpCTSS", "CAGEr", function (object, threshold, nrPassThreshold, thresholdIsTpm) { + flagLowExpCTSS(CTSStagCountSE(object), threshold, nrPassThreshold, thresholdIsTpm) }) -setMethod(".filterCtss", "RangedSummarizedExperiment", function (object, threshold, nrPassThreshold, thresholdIsTpm) { +#' @rdname flagLowExpCTSS + +setMethod("flagLowExpCTSS", "RangedSummarizedExperiment", function (object, threshold, nrPassThreshold, thresholdIsTpm) { assay <- ifelse(thresholdIsTpm, "normalizedTpmMatrix", "counts") if(assay == "normalizedTpmMatrix" & is.null(assays(object)[[assay]])) stop("Normalise the CAGEr object first with ", sQuote("normalizeTagCount()"), ".") - .filterCtss(assays(object)[[assay]], threshold, nrPassThreshold, thresholdIsTpm) + flagLowExpCTSS(assays(object)[[assay]], threshold, nrPassThreshold, thresholdIsTpm) }) -setMethod(".filterCtss", "DataFrame", function (object, threshold, nrPassThreshold, thresholdIsTpm) { +#' @rdname flagLowExpCTSS + +setMethod("flagLowExpCTSS", "DataFrame", function (object, threshold, nrPassThreshold, thresholdIsTpm) { nr.pass.threshold <- rowSums.RleDataFrame(lapply(object, \(x) x > threshold) |> DataFrame()) nr.pass.threshold >= min(nrPassThreshold, ncol(object)) }) -setMethod(".filterCtss", "matrix", function (object, threshold, nrPassThreshold, thresholdIsTpm) { +#' @rdname flagLowExpCTSS + +setMethod("flagLowExpCTSS", "matrix", function (object, threshold, nrPassThreshold, thresholdIsTpm) { nr.pass.threshold <- rowSums(object > threshold) nr.pass.threshold >= min(nrPassThreshold, ncol(object)) +}) + +#' @rdname flagLowExpCTSS +#' +#' @return `filterLowExpCTSS` returns the `CAGEr` object where the output of +#' `flagLowExpCTSS` was stored internally. +#' +#' @export + +setGeneric("filterLowExpCTSS", function( object + , threshold = 1 + , nrPassThreshold = 1 + , thresholdIsTpm = TRUE) + standardGeneric("filterLowExpCTSS") +) + +#' @rdname flagLowExpCTSS + +setMethod("filterLowExpCTSS", "CAGEr", function (object, threshold, nrPassThreshold, thresholdIsTpm) { + filteredCTSSidx(object) <- flagLowExpCTSS(CTSStagCountSE(object), threshold, nrPassThreshold, thresholdIsTpm) + object }) \ No newline at end of file diff --git a/R/ClusteringMethods.R b/R/ClusteringMethods.R index ae5691b..b15e24a 100644 --- a/R/ClusteringMethods.R +++ b/R/ClusteringMethods.R @@ -1,156 +1,5 @@ #' @include CTSS.R Multicore.R -#' @name clusterCTSS -#' -#' @title Cluster CTSSs into tag clusters -#' -#' @description Clusters individual CAGE transcription start sites (CTSSs) along -#' the genome into tag clusters (TCs) using specified _ab initio_ method, or -#' assigns them to predefined genomic regions. -#' -#' @param object A [`CAGEr`] object. -#' -#' @param threshold,nrPassThreshold Ignore CTSSs with signal `< threshold` -#' in `< nrPassThreshold` experiments. -#' -#' @param thresholdIsTpm Logical indicating if `threshold` is expressed in -#' raw tag counts (`FALSE`) or normalized signal (`TRUE`). -#' -#' @param method Clustering method: `"distclu"` or `"paraclu"`. -#' -#' @param maxDist Maximal distance between two neighbouring CTSSs for them to be -#' part of the same cluster. Used only when `method = "distclu"`, -#' otherwise ignored. -#' -#' @param keepSingletonsAbove Remove "singleton" tag clusters of width 1 with -#' signal `< keepSingletonsAbove`. Default value `0` results in keeping -#' all TCs by default. Setting it to `Inf` removes all singletons. -#' -#' @param minStability Minimal stability of the cluster, where stability is -#' defined as ratio between maximal and minimal density value for which -#' this cluster is maximal scoring. For definition of stability refer to -#' Frith _et al._, Genome Research, 2007. Clusters with stability -#' `< minStability` will be discarded. Used only when `method = "paraclu"`. -#' -#' @param maxLength Maximal length of cluster in base-pairs. Clusters with length -#' `> maxLength` will be discarded. -#' -#' @param reduceToNonoverlapping Logical, should smaller clusters contained -#' within bigger cluster be removed to make a final set of tag clusters -#' non-overlapping. Used only `method = "paraclu"`. -#' -#' @param useMulticore Logical, should multicore be used. `useMulticore = TRUE` -#' has no effect on non-Unix-like platforms. -#' -#' @param nrCores Number of cores to use when `useMulticore = TRUE`. Default -#' value `NULL` uses all detected cores. -#' -#' @details The `"distclu"` method is an implementation of simple distance-based -#' clustering of data attached to sequences, where two neighbouring TSSs are -#' joined together if they are closer than some specified distance (see -#' [`GenomicRanges::reduce`] for implementation details. -#' -#' `"paraclu"` is an implementation of Paraclu algorithm for parametric -#' clustering of data attached to sequences (Frith _et al._, Genome Research, -#' 2007). Since Paraclu finds clusters within clusters (unlike distclu), -#' additional parameters (`keepSingletonsAbove`, -#' `minStability`, `maxLength` and `reduceToNonoverlapping`) can be specified to -#' simplify the output by discarding too small (singletons) or too big clusters, -#' and to reduce the clusters to a final set of non-overlapping clusters. -#' -#' Clustering is done for every CAGE dataset within the CAGEr object separately, -#' resulting in a different set of tag clusters for every CAGE dataset. TCs from -#' different datasets can further be aggregated into a single referent set of -#' consensus clusters by calling the [`aggregateTagClusters`] function. -#' -#' @return Returns the [`CAGEexp`] object, in which, the results will be stored as a `GRangesList` of -#' [`TagClusters`] objects in the metadata slot `tagClusters`. The -#' `TagClusters` objects will contain a `filteredCTSSidx` column if appropriate. -#' The clustering method name is saved in the metadata slot of the `GRangesList`. -#' -#' @references Frith _et al._ (2007) A code for transcription initiation in -#' mammalian genomes, _Genome Research_ **18**(1):1-12, -#' (\href{http://www.cbrc.jp/paraclu/}{http://www.cbrc.jp/paraclu/}). -#' -#' @author Vanja Haberle -#' -#' @seealso [`aggregateTagClusters`] -#' -#' @family CAGEr object modifiers -#' @family CAGEr clusters functions -#' -#' @examples -#' -#' # Using 'distclu', notice argument 'maxDist' -#' ce <- clusterCTSS( exampleCAGEexp, threshold = 50, thresholdIsTpm = TRUE -#' , nrPassThreshold = 1, method = "distclu", maxDist = 20 -#' , keepSingletonsAbove = 100) -#' tagClustersGR(ce, "Zf.30p.dome") -#' -#' # Using 'paraclu', notice arguments 'maxLength' and 'minStability' -#' ce <- clusterCTSS( exampleCAGEexp, threshold = 50, thresholdIsTpm = TRUE -#' , nrPassThreshold = 1, method = "paraclu" -#' , keepSingletonsAbove = 100 -#' , maxLength = 500, minStability = 1 -#' , reduceToNonoverlapping = TRUE) -#' tagClustersGR(ce, "Zf.30p.dome") -#' -#' @export - -setGeneric( "clusterCTSS" - , function( object - , threshold = 1, nrPassThreshold = 1, thresholdIsTpm = TRUE - , method = c("distclu", "paraclu"), maxDist = 20 - , keepSingletonsAbove = 0 - , minStability = 1, maxLength = 500 - , reduceToNonoverlapping = TRUE - , useMulticore = FALSE, nrCores = NULL) - standardGeneric("clusterCTSS")) - -#' @rdname clusterCTSS - -setMethod( "clusterCTSS", "CAGEexp" - , function( object, threshold, nrPassThreshold, thresholdIsTpm, method, maxDist - , keepSingletonsAbove, minStability, maxLength - , reduceToNonoverlapping, useMulticore, nrCores) { - - assay <- ifelse(isTRUE(thresholdIsTpm), "normalizedTpmMatrix", "counts") - data <- CTSStagCountSE(object) - - if (! "normalizedTpmMatrix" %in% assayNames(data)) - stop( "Could not find normalized CAGE signal values, see ?normalizeTagCount.\n" - , "clusterCTSS() needs normalized values to create its output tables, that " - , "include TPM expression columns.") - - message("\nFiltering out CTSSs below threshold...") - filteredCTSSidx(object) <- - .filterCtss(data, threshold = threshold - , nrPassThreshold = nrPassThreshold, thresholdIsTpm = thresholdIsTpm) - - message("Clustering...") - method <- match.arg(method) - - if (method == "distclu") { - ctss.cluster.list <- distclu( object = data[decode(filteredCTSSidx(object)),] - , max.dist = maxDist, keepSingletonsAbove = keepSingletonsAbove) - } else if (method == "paraclu") { - ctss.cluster.list <- paraclu( object = data[decode(filteredCTSSidx(object)),] - , minStability = minStability, maxLength = maxLength - , keepSingletonsAbove = keepSingletonsAbove - , reduceToNonoverlapping = reduceToNonoverlapping - , useMulticore = useMulticore, nrCores = nrCores) - } else if(method == "custom") { - stop("Deprecated method. See ", dQuote("CustomConsensusClusters()"), " instead.") - } - - seqlevels(ctss.cluster.list) <- seqlevels(CTSStagCountSE(object)) - seqinfo(ctss.cluster.list) <- seqinfo(CTSStagCountSE(object)) - # Changing the sequence levels may change the sort order. Re-sort - ctss.cluster.list <- sort(ctss.cluster.list) - metadata(object)$tagClusters <- ctss.cluster.list - object -}) - #' @rdname byCtss #' #' @title Apply functions to identical CTSSes. diff --git a/R/Distclu.R b/R/Distclu.R index 095abc7..33f728f 100644 --- a/R/Distclu.R +++ b/R/Distclu.R @@ -3,10 +3,20 @@ NULL #' Distance clustering #' +#' The `"distclu"` method is an implementation of simple distance-based +#' clustering of data attached to sequences, where two neighbouring TSSs are +#' joined together if they are closer than some specified distance (see +#' [`GenomicRanges::reduce`] for implementation details. +#' +#' Clustering is done for every CAGE dataset within the CAGEr object separately, +#' resulting in a different set of tag clusters for every CAGE dataset. TCs from +#' different datasets can further be aggregated into a single referent set of +#' consensus clusters by calling the [`aggregateTagClusters`] function. +#' #' @param object The [`SummarizedExperiment::RangedSummarizedExperiment`] object #' containing CTSS information, or just a [`CTSS`] object. #' -#' @param max.dist Maximal distance between two neighbouring CTSSs for them to +#' @param maxDist Maximal distance between two neighbouring CTSSs for them to #' be part of the same cluster. #' #' @param keepSingletonsAbove Remove "singleton" tag clusters of width 1 with @@ -14,27 +24,39 @@ NULL #' all TCs by default. Setting it to `Inf` removes all singletons. #' #' @family CAGEr clustering methods +#' @family CAGEr object modifiers +#' @family CAGEr clusters functions +#' +#' @seealso [`aggregateTagClusters`] +#' +#' @author Vanja Haberle +#' @author Charles Plessy #' -#' @return For `CTSS` input, a [`TagClusters`] object, and for -#' `SummarizedExperiment` in put, a [`GRangesList`] of [`TagClusters`] objects. +#' @return For `CTSS` input, a [`TagClusters`] object, for +#' `SummarizedExperiment` input, a [`GRangesList`] of [`TagClusters`] objects, +#' and for [`CAGEexp`] input, a modified object containing the tag clusters +#' stored as a `GRangesList` of [`TagClusters`] objects in its metadata slot +#' `tagClusters`. #' #' @examples #' distclu(CTSSnormalizedTpmGR(exampleCAGEexp, 1)[1:10]) #' distclu(CTSStagCountSE(exampleCAGEexp)[1:25,]) +#' ce <- distclu(exampleCAGEexp, maxDist = 20, keepSingletonsAbove = 100) +#' tagClustersGR(ce, "Zf.30p.dome") #' #' @export setGeneric("distclu", function( object, - max.dist = 20, + maxDist = 20, keepSingletonsAbove = 0 ) standardGeneric("distclu")) #' @rdname distclu setMethod("distclu", "SummarizedExperiment", - function(object, max.dist, keepSingletonsAbove) { + function(object, maxDist, keepSingletonsAbove) { ctss.cluster.list <- GRangesList() for(s in colnames(object)) { message("\t-> ", s) @@ -42,14 +64,14 @@ setMethod("distclu", "SummarizedExperiment", score(d) <- assays(object)[["normalizedTpmMatrix"]][[s]] d <- subset(d, score(d) > 0) ctss.cluster.list[[s]] <- - distclu(d, max.dist = max.dist, + distclu(d, maxDist = maxDist, keepSingletonsAbove = keepSingletonsAbove) } endoapply(ctss.cluster.list, as, "TagClusters") }) -.distclu_CTSS <- function(object, max.dist, keepSingletonsAbove) { - clusters <- reduce(GRanges(object), min = max.dist) +.distclu_CTSS <- function(object, maxDist, keepSingletonsAbove) { + clusters <- reduce(GRanges(object), min.gapwidth = maxDist) clusters <- .ctss_summary_for_clusters(object, clusters) # Remove clusters that match only one CTSS unless their expression is high enough clusters <- subset(clusters, clusters$nr_ctss > 1 | score(clusters) >= keepSingletonsAbove) @@ -60,3 +82,22 @@ setMethod("distclu", "SummarizedExperiment", #' @rdname distclu setMethod("distclu", "CTSS", .distclu_CTSS) + +#' @rdname distclu + +setMethod( "distclu", "CAGEexp", function(object, maxDist, keepSingletonsAbove) { + if (! "normalizedTpmMatrix" %in% assayNames(CTSStagCountSE(object))) + stop( "Could not find normalized CAGE signal values, see ?normalizeTagCount.\n" + , "distclu() needs normalized values to create its output tables, that " + , "include TPM expression columns.") + message("Clustering...") + ctss.cluster.list <- + distclu( object = CTSStagCountSE(object)[filteredCTSSidx(object),] + , maxDist = maxDist, keepSingletonsAbove = keepSingletonsAbove) + seqlevels(ctss.cluster.list) <- seqlevels(CTSStagCountSE(object)) + seqinfo(ctss.cluster.list) <- seqinfo(CTSStagCountSE(object)) + # Changing the sequence levels may change the sort order. Re-sort + ctss.cluster.list <- sort(ctss.cluster.list) + metadata(object)$tagClusters <- ctss.cluster.list + object +}) diff --git a/R/ExpressionProfilingMethods.R b/R/ExpressionProfilingMethods.R index d7644b3..8e04a49 100644 --- a/R/ExpressionProfilingMethods.R +++ b/R/ExpressionProfilingMethods.R @@ -32,7 +32,7 @@ #' normalized CAGE signal `>= tpmThreshold` in at least `nrPassThreshold` CAGE #' experiments are used for expression clustering. However, CTSSs along the #' genome can be spatially clustered into tag clusters for each experiment -#' separately using the [`clusterCTSS`] function, and then aggregated across +#' separately using a CTSS clustering function, and then aggregated across #' experiments into consensus clusters using [`aggregateTagClusters`] function. #' Once the consensus clusters have been created, expression clustering at the #' level of these wider genomic regions (representing entire promoters rather @@ -118,7 +118,7 @@ setMethod("getExpressionProfiles", "matrix", function (object, what, tpmThreshold, nrPassThreshold, method, xDim, yDim) { method <- match.arg(method) - idx <- .filterCtss( object, threshold = tpmThreshold + idx <- flagLowExpCTSS( object, threshold = tpmThreshold , nrPassThreshold = nrPassThreshold, thresholdIsTpm = TRUE) m <- t(scale(t(log(object + 1)), center=FALSE)) diff --git a/R/GetMethods.R b/R/GetMethods.R index 86528d3..54e5020 100644 --- a/R/GetMethods.R +++ b/R/GetMethods.R @@ -202,10 +202,7 @@ function (object){ #' #' @return `CTSScoordinatesGR` returns the coordinates as a [CTSS()] object #' wrapping genomic ranges. A `filteredCTSSidx` column metadata will be present -#' if [clusterCTSS()] was ran earlier. -#' -#' @seealso [`clusterCTSS`] -#' +#' if [`filterLowExpCTSS`] was ran earlier. #' #' @importFrom GenomeInfoDb genome genome<- #' @importFrom IRanges IRanges @@ -389,7 +386,7 @@ setMethod( "CTSSnormalizedTpmGR", "CAGEexp", function (object, samples) { #' #' @title Extract tag clusters (TCs) for individual CAGE experiments #' -#' @description Extracts tag clusters (TCs) produced by [`clusterCTSS`] function +#' @description Extracts tag clusters (TCs) #' for a specified CAGE experiment from a [`CAGEexp`] object. #' #' @param object A `CAGEexp` object. @@ -445,7 +442,7 @@ setMethod( "tagClustersGR", "CAGEexp" , function (object, sample, qLow, qUp) { tc <- metadata(object)$tagClusters[[sample]] if (is.null(tc)) - stop( "No clusters found, run ", sQuote("clusterCTSS"), " first." , call. = FALSE) + stop( "No clusters found, run ", sQuote("distclu"), "or", sQuote("paraclu"), " first." , call. = FALSE) if (! is.null(qLow) & ! is.null(qUp)) { qLowName <- paste0("q_", qLow) diff --git a/R/Paraclu.R b/R/Paraclu.R index d5fa179..5d9c332 100644 --- a/R/Paraclu.R +++ b/R/Paraclu.R @@ -1,7 +1,16 @@ #' Parametric clustering #' -#' Implementation of Paraclu - parametric clustering of data attached to -#' sequences (). +#' `"paraclu"` is an implementation of Paraclu algorithm for parametric +#' clustering of data attached to sequences (Frith _et al._, Genome Research, +#' 2007). Since Paraclu finds clusters within clusters (unlike [`distclu`]), +#' additional parameters (`minStability`, `maxLength` and `reduceToNonoverlapping`) +#' can be specified to simplify the output by discarding too big clusters, +#' and to reduce the clusters to a final set of non-overlapping clusters. +#' +#' Clustering is done for every CAGE dataset within the CAGEr object separately, +#' resulting in a different set of tag clusters for every CAGE dataset. TCs from +#' different datasets can further be aggregated into a single referent set of +#' consensus clusters by calling the [`aggregateTagClusters`] function. #' #' @param object A [`CTSS`], or a [`S4Vectors::Pairs`] object with positions #' _first_ and scores _second_. @@ -48,7 +57,17 @@ #' Running Paraclu on a [`RangedSummarizedExperiment`] object will loop on each #' sample, and return the results as a [`GRangesList`] of `TagClusters`. #' +#' Running Paraclu on a [`CAGEexp`] returnts is with the clusters stored as a +#' `GRangesList` of [`TagClusters`] objects in its metadata slot `tagClusters`. +#' #' @family CAGEr clustering methods +#' @family CAGEr object modifiers +#' @family CAGEr clusters functions +#' +#' @seealso [`aggregateTagClusters`] +#' +#' @author Vanja Haberle +#' @author Charles Plessy #' #' @importFrom utils tail #' @importFrom S4Vectors Pairs first second @@ -61,6 +80,11 @@ #' paraclu(pair[1:10]) #' paraclu(ctss[1:10]) #' paraclu(CTSStagCountSE(exampleCAGEexp)[1:25,]) +#' ce <- paraclu( exampleCAGEexp, +#' , keepSingletonsAbove = 100 +#' , maxLength = 500, minStability = 1 +#' , reduceToNonoverlapping = TRUE) +#' tagClustersGR(ce, "Zf.30p.dome") #' #' @export @@ -205,3 +229,30 @@ setMethod("paraclu", "SummarizedExperiment", } endoapply(tag.cluster.list, as, "TagClusters") }) + +#' @rdname paraclu + +setMethod( "paraclu", "CAGEexp", + function( object + , minStability = 1, maxLength = 500 + , keepSingletonsAbove = 0 + , reduceToNonoverlapping = TRUE + , useMulticore = FALSE, nrCores = NULL) { + if (! "normalizedTpmMatrix" %in% assayNames(CTSStagCountSE(object))) + stop( "Could not find normalized CAGE signal values, see ?normalizeTagCount.\n" + , "distclu() needs normalized values to create its output tables, that " + , "include TPM expression columns.") + message("Clustering...") + ctss.cluster.list <- + paraclu( object = CTSStagCountSE(object)[filteredCTSSidx(object),] + , minStability = minStability, maxLength = maxLength + , keepSingletonsAbove = keepSingletonsAbove + , reduceToNonoverlapping = reduceToNonoverlapping + , useMulticore = useMulticore, nrCores = nrCores) + seqlevels(ctss.cluster.list) <- seqlevels(CTSStagCountSE(object)) + seqinfo(ctss.cluster.list) <- seqinfo(CTSStagCountSE(object)) + # Changing the sequence levels may change the sort order. 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A \code{filteredCTSSidx} column metadata will be present -if \code{\link[=clusterCTSS]{clusterCTSS()}} was ran earlier. +if \code{\link{filterLowExpCTSS}} was ran earlier. } \description{ Extracts the genomic coordinates of all detected TSSs @@ -44,8 +44,6 @@ CTSScoordinatesGR(exampleCAGEexp) \seealso{ \code{\link{getCTSS}} -\code{\link{clusterCTSS}} - Other CAGEr accessor methods: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{CTSSnormalizedTpmDF}()}, diff --git a/man/CTSScumulativesTagClusters.Rd b/man/CTSScumulativesTagClusters.Rd index b5e36e2..f54b4c0 100644 --- a/man/CTSScumulativesTagClusters.Rd +++ b/man/CTSScumulativesTagClusters.Rd @@ -38,10 +38,11 @@ Accessor function. Other CAGEr clusters functions: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersDESeq2}()}, \code{\link{consensusClustersGR}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{plotInterquantileWidth}()}, \code{\link{quantilePositions}()}, \code{\link{tagClustersGR}()} diff --git a/man/CTSStoGenes.Rd b/man/CTSStoGenes.Rd index 6a281ad..9089bba 100644 --- a/man/CTSStoGenes.Rd +++ b/man/CTSStoGenes.Rd @@ -45,10 +45,11 @@ Other CAGEr object modifiers: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()}, diff --git a/man/CustomConsensusClusters.Rd b/man/CustomConsensusClusters.Rd index 52fb70d..c94ab47 100644 --- a/man/CustomConsensusClusters.Rd +++ b/man/CustomConsensusClusters.Rd @@ -61,10 +61,11 @@ Other CAGEr object modifiers: \code{\link{CTSStoGenes}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()}, @@ -73,10 +74,11 @@ Other CAGEr object modifiers: Other CAGEr clusters functions: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersDESeq2}()}, \code{\link{consensusClustersGR}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{plotInterquantileWidth}()}, \code{\link{quantilePositions}()}, \code{\link{tagClustersGR}()} diff --git a/man/aggregateTagClusters.Rd b/man/aggregateTagClusters.Rd index b4c7cb9..eaeeabb 100644 --- a/man/aggregateTagClusters.Rd +++ b/man/aggregateTagClusters.Rd @@ -69,7 +69,7 @@ Aggregates tag clusters (TCs) across all CAGE datasets within the CAGEr object to create a referent set of consensus clusters. } \details{ -Since the tag clusters (TCs) returned by the \code{\link{clusterCTSS}} +Since the tag clusters (TCs) returned by the CTSS clustering functions function are constructed separately for every CAGE sample within the CAGEr object, they can differ between samples in both their number, genomic coordinates, position of dominant TSS and overall signal. To be able to @@ -106,10 +106,11 @@ Other CAGEr object modifiers: \code{\link{CTSStoGenes}()}, \code{\link{CustomConsensusClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()}, @@ -118,10 +119,11 @@ Other CAGEr object modifiers: Other CAGEr clusters functions: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{CustomConsensusClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersDESeq2}()}, \code{\link{consensusClustersGR}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{plotInterquantileWidth}()}, \code{\link{quantilePositions}()}, \code{\link{tagClustersGR}()} diff --git a/man/annotateCTSS.Rd b/man/annotateCTSS.Rd index 6cf5933..3573950 100644 --- a/man/annotateCTSS.Rd +++ b/man/annotateCTSS.Rd @@ -79,10 +79,11 @@ Other CAGEr object modifiers: \code{\link{CTSStoGenes}()}, \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()}, diff --git a/man/clusterCTSS.Rd b/man/clusterCTSS.Rd deleted file mode 100644 index dccc3aa..0000000 --- a/man/clusterCTSS.Rd +++ /dev/null @@ -1,159 +0,0 @@ -% Generated by roxygen2: do not edit by hand -% Please edit documentation in R/ClusteringMethods.R -\name{clusterCTSS} -\alias{clusterCTSS} -\alias{clusterCTSS,CAGEexp-method} -\title{Cluster CTSSs into tag clusters} -\usage{ -clusterCTSS( - object, - threshold = 1, - nrPassThreshold = 1, - thresholdIsTpm = TRUE, - method = c("distclu", "paraclu"), - maxDist = 20, - keepSingletonsAbove = 0, - minStability = 1, - maxLength = 500, - reduceToNonoverlapping = TRUE, - useMulticore = FALSE, - nrCores = NULL -) - -\S4method{clusterCTSS}{CAGEexp}( - object, - threshold = 1, - nrPassThreshold = 1, - thresholdIsTpm = TRUE, - method = c("distclu", "paraclu"), - maxDist = 20, - keepSingletonsAbove = 0, - minStability = 1, - maxLength = 500, - reduceToNonoverlapping = TRUE, - useMulticore = FALSE, - nrCores = NULL -) -} -\arguments{ -\item{object}{A \code{\link{CAGEr}} object.} - -\item{threshold, nrPassThreshold}{Ignore CTSSs with signal \verb{< threshold} -in \verb{< nrPassThreshold} experiments.} - -\item{thresholdIsTpm}{Logical indicating if \code{threshold} is expressed in -raw tag counts (\code{FALSE}) or normalized signal (\code{TRUE}).} - -\item{method}{Clustering method: \code{"distclu"} or \code{"paraclu"}.} - -\item{maxDist}{Maximal distance between two neighbouring CTSSs for them to be -part of the same cluster. Used only when \code{method = "distclu"}, -otherwise ignored.} - -\item{keepSingletonsAbove}{Remove "singleton" tag clusters of width 1 with -signal \verb{< keepSingletonsAbove}. Default value \code{0} results in keeping -all TCs by default. Setting it to \code{Inf} removes all singletons.} - -\item{minStability}{Minimal stability of the cluster, where stability is -defined as ratio between maximal and minimal density value for which -this cluster is maximal scoring. For definition of stability refer to -Frith \emph{et al.}, Genome Research, 2007. Clusters with stability -\verb{< minStability} will be discarded. Used only when \code{method = "paraclu"}.} - -\item{maxLength}{Maximal length of cluster in base-pairs. Clusters with length -\verb{> maxLength} will be discarded.} - -\item{reduceToNonoverlapping}{Logical, should smaller clusters contained -within bigger cluster be removed to make a final set of tag clusters -non-overlapping. Used only \code{method = "paraclu"}.} - -\item{useMulticore}{Logical, should multicore be used. \code{useMulticore = TRUE} -has no effect on non-Unix-like platforms.} - -\item{nrCores}{Number of cores to use when \code{useMulticore = TRUE}. Default -value \code{NULL} uses all detected cores.} -} -\value{ -Returns the \code{\link{CAGEexp}} object, in which, the results will be stored as a \code{GRangesList} of -\code{\link{TagClusters}} objects in the metadata slot \code{tagClusters}. The -\code{TagClusters} objects will contain a \code{filteredCTSSidx} column if appropriate. -The clustering method name is saved in the metadata slot of the \code{GRangesList}. -} -\description{ -Clusters individual CAGE transcription start sites (CTSSs) along -the genome into tag clusters (TCs) using specified \emph{ab initio} method, or -assigns them to predefined genomic regions. -} -\details{ -The \code{"distclu"} method is an implementation of simple distance-based -clustering of data attached to sequences, where two neighbouring TSSs are -joined together if they are closer than some specified distance (see -\code{\link[GenomicRanges:inter-range-methods]{GenomicRanges::reduce}} for implementation details. - -\code{"paraclu"} is an implementation of Paraclu algorithm for parametric -clustering of data attached to sequences (Frith \emph{et al.}, Genome Research, -2007). Since Paraclu finds clusters within clusters (unlike distclu), -additional parameters (\code{keepSingletonsAbove}, -\code{minStability}, \code{maxLength} and \code{reduceToNonoverlapping}) can be specified to -simplify the output by discarding too small (singletons) or too big clusters, -and to reduce the clusters to a final set of non-overlapping clusters. - -Clustering is done for every CAGE dataset within the CAGEr object separately, -resulting in a different set of tag clusters for every CAGE dataset. TCs from -different datasets can further be aggregated into a single referent set of -consensus clusters by calling the \code{\link{aggregateTagClusters}} function. -} -\examples{ - -# Using 'distclu', notice argument 'maxDist' -ce <- clusterCTSS( exampleCAGEexp, threshold = 50, thresholdIsTpm = TRUE - , nrPassThreshold = 1, method = "distclu", maxDist = 20 - , keepSingletonsAbove = 100) -tagClustersGR(ce, "Zf.30p.dome") - -# Using 'paraclu', notice arguments 'maxLength' and 'minStability' -ce <- clusterCTSS( exampleCAGEexp, threshold = 50, thresholdIsTpm = TRUE - , nrPassThreshold = 1, method = "paraclu" - , keepSingletonsAbove = 100 - , maxLength = 500, minStability = 1 - , reduceToNonoverlapping = TRUE) -tagClustersGR(ce, "Zf.30p.dome") - -} -\references{ -Frith \emph{et al.} (2007) A code for transcription initiation in -mammalian genomes, \emph{Genome Research} \strong{18}(1):1-12, -(\href{http://www.cbrc.jp/paraclu/}{http://www.cbrc.jp/paraclu/}). -} -\seealso{ -\code{\link{aggregateTagClusters}} - -Other CAGEr object modifiers: -\code{\link{CTSStoGenes}()}, -\code{\link{CustomConsensusClusters}()}, -\code{\link{aggregateTagClusters}()}, -\code{\link{annotateCTSS}()}, -\code{\link{cumulativeCTSSdistribution}()}, -\code{\link{getCTSS}()}, -\code{\link{normalizeTagCount}()}, -\code{\link{quantilePositions}()}, -\code{\link{quickEnhancers}()}, -\code{\link{resetCAGEexp}()}, -\code{\link{summariseChrExpr}()} - -Other CAGEr clusters functions: -\code{\link{CTSScumulativesTagClusters}()}, -\code{\link{CustomConsensusClusters}()}, -\code{\link{aggregateTagClusters}()}, -\code{\link{consensusClustersDESeq2}()}, -\code{\link{consensusClustersGR}()}, -\code{\link{cumulativeCTSSdistribution}()}, -\code{\link{plotInterquantileWidth}()}, -\code{\link{quantilePositions}()}, -\code{\link{tagClustersGR}()} -} -\author{ -Vanja Haberle -} -\concept{CAGEr clusters functions} -\concept{CAGEr object modifiers} diff --git a/man/consensusClusters.Rd b/man/consensusClusters.Rd index 1a5dec7..0e920be 100644 --- a/man/consensusClusters.Rd +++ b/man/consensusClusters.Rd @@ -72,9 +72,10 @@ Other CAGEr clusters functions: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersDESeq2}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{plotInterquantileWidth}()}, \code{\link{quantilePositions}()}, \code{\link{tagClustersGR}()} diff --git a/man/consensusClustersDESeq2.Rd b/man/consensusClustersDESeq2.Rd index 71038fc..57192ba 100644 --- a/man/consensusClustersDESeq2.Rd +++ b/man/consensusClustersDESeq2.Rd @@ -32,9 +32,10 @@ Other CAGEr clusters functions: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersGR}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{plotInterquantileWidth}()}, \code{\link{quantilePositions}()}, \code{\link{tagClustersGR}()} diff --git a/man/cumulativeCTSSdistribution.Rd b/man/cumulativeCTSSdistribution.Rd index 71481de..ad2b45d 100644 --- a/man/cumulativeCTSSdistribution.Rd +++ b/man/cumulativeCTSSdistribution.Rd @@ -53,9 +53,10 @@ Other CAGEr object modifiers: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()}, @@ -65,9 +66,10 @@ Other CAGEr clusters functions: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersDESeq2}()}, \code{\link{consensusClustersGR}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{plotInterquantileWidth}()}, \code{\link{quantilePositions}()}, \code{\link{tagClustersGR}()} diff --git a/man/distclu.Rd b/man/distclu.Rd index 0597b88..5ea0837 100644 --- a/man/distclu.Rd +++ b/man/distclu.Rd @@ -4,6 +4,7 @@ \alias{distclu} \alias{distclu,SummarizedExperiment-method} \alias{distclu,CTSS-method} +\alias{distclu,CAGEexp-method} \title{Distance clustering} \usage{ distclu(object, max.dist = 20, keepSingletonsAbove = 0) @@ -11,6 +12,8 @@ distclu(object, max.dist = 20, keepSingletonsAbove = 0) \S4method{distclu}{SummarizedExperiment}(object, max.dist = 20, keepSingletonsAbove = 0) \S4method{distclu}{CTSS}(object, max.dist = 20, keepSingletonsAbove = 0) + +\S4method{distclu}{CAGEexp}(object, max.dist = 20, keepSingletonsAbove = 0) } \arguments{ \item{object}{The \code{\link[SummarizedExperiment:RangedSummarizedExperiment-class]{SummarizedExperiment::RangedSummarizedExperiment}} object @@ -24,20 +27,69 @@ signal \verb{< keepSingletonsAbove}. Default value \code{0} results in keeping all TCs by default. Setting it to \code{Inf} removes all singletons.} } \value{ -For \code{CTSS} input, a \code{\link{TagClusters}} object, and for -\code{SummarizedExperiment} in put, a \code{\link{GRangesList}} of \code{\link{TagClusters}} objects. +For \code{CTSS} input, a \code{\link{TagClusters}} object, for +\code{SummarizedExperiment} input, a \code{\link{GRangesList}} of \code{\link{TagClusters}} objects, +and for \code{\link{CAGEexp}} input, a modified object containing the tag clusters +stored as a \code{GRangesList} of \code{\link{TagClusters}} objects in its metadata slot +\code{tagClusters}. } \description{ -Distance clustering +The \code{"distclu"} method is an implementation of simple distance-based +clustering of data attached to sequences, where two neighbouring TSSs are +joined together if they are closer than some specified distance (see +\code{\link[GenomicRanges:inter-range-methods]{GenomicRanges::reduce}} for implementation details. +} +\details{ +Clustering is done for every CAGE dataset within the CAGEr object separately, +resulting in a different set of tag clusters for every CAGE dataset. TCs from +different datasets can further be aggregated into a single referent set of +consensus clusters by calling the \code{\link{aggregateTagClusters}} function. } \examples{ distclu(CTSSnormalizedTpmGR(exampleCAGEexp, 1)[1:10]) distclu(CTSStagCountSE(exampleCAGEexp)[1:25,]) +ce <- distclu(exampleCAGEexp, max.dist = 20, keepSingletonsAbove = 100) +tagClustersGR(ce, "Zf.30p.dome") } \seealso{ +\code{\link{aggregateTagClusters}} + Other CAGEr clustering methods: \code{\link{consensusClustersTpm}()}, \code{\link{paraclu}()} + +Other CAGEr object modifiers: +\code{\link{CTSStoGenes}()}, +\code{\link{CustomConsensusClusters}()}, +\code{\link{aggregateTagClusters}()}, +\code{\link{annotateCTSS}()}, +\code{\link{cumulativeCTSSdistribution}()}, +\code{\link{getCTSS}()}, +\code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, +\code{\link{quantilePositions}()}, +\code{\link{quickEnhancers}()}, +\code{\link{resetCAGEexp}()}, +\code{\link{summariseChrExpr}()} + +Other CAGEr clusters functions: +\code{\link{CTSScumulativesTagClusters}()}, +\code{\link{CustomConsensusClusters}()}, +\code{\link{aggregateTagClusters}()}, +\code{\link{consensusClustersDESeq2}()}, +\code{\link{consensusClustersGR}()}, +\code{\link{cumulativeCTSSdistribution}()}, +\code{\link{paraclu}()}, +\code{\link{plotInterquantileWidth}()}, +\code{\link{quantilePositions}()}, +\code{\link{tagClustersGR}()} +} +\author{ +Vanja Haberle + +Charles Plessy } \concept{CAGEr clustering methods} +\concept{CAGEr clusters functions} +\concept{CAGEr object modifiers} diff --git a/man/dot-ctss_summary_for_clusters.Rd b/man/dot-ctss_summary_for_clusters.Rd new file mode 100644 index 0000000..d7a374d --- /dev/null +++ b/man/dot-ctss_summary_for_clusters.Rd @@ -0,0 +1,47 @@ +% Generated by roxygen2: do not edit by hand +% Please edit documentation in R/ClusteringFunctions.R +\name{.ctss_summary_for_clusters} +\alias{.ctss_summary_for_clusters} +\title{Summarise CTSSs included in clusters} +\usage{ +.ctss_summary_for_clusters(ctss, clusters) +} +\arguments{ +\item{ctss}{A \code{\link{CTSS}} object.} + +\item{clusters}{A \code{\link{TagClusters}}, \code{\link{ConsensusClusters}} or any other +object implementing the \code{\link{GRanges}} class.} +} +\value{ +The \code{clusters} object with a new \code{dominant_CTSS} metadata in \code{CTSS} +format reporting the genomic coordinate and expression score of most +highly expressed position in each cluster, plus a \code{nr_ctss} metadata reporting +the number of expressed CTSSs in each cluster. +} +\description{ +Summarise CTSSs included in clusters +} +\examples{ +# See also benchmarks/dominant_ctss.md +(ctss <- CTSS( 'chr1', IRanges(start = 1:10, end = 1:10) + , '+', score = c(1, 0, 0, 1, 2, 0, 2, 1, 0, 1))) +(clusters <- GRanges( 'chr1', IRanges(start = c(1,9) + , end = c(8,10)), '+')) |> as("TagClusters") + +# The function assumes that all CTSSes have a score above zero +.ctss_summary_for_clusters(ctss[score(ctss)>0], clusters) +# If not the case, it will give incorrect nr_ctss and fail to remove singletons +.ctss_summary_for_clusters(ctss, clusters) + +# The function needs its output to be sorted and is not going to check it. +.ctss_summary_for_clusters(rev(ctss), clusters) +.ctss_summary_for_clusters(ctss, rev(clusters)) + +# Ties are resolved with 5' preference for both plus and minus strands. +# This may create a small bias. +ctss_minus <- ctss +strand(ctss_minus) <- '-' +clusters_minus <- clusters +strand(clusters_minus) <- '-' +.ctss_summary_for_clusters(ctss_minus, clusters_minus) +} diff --git a/man/exampleCAGEexp.Rd b/man/exampleCAGEexp.Rd index 9844a3e..8d09f2f 100644 --- a/man/exampleCAGEexp.Rd +++ b/man/exampleCAGEexp.Rd @@ -38,7 +38,8 @@ exampleCAGEexp <- CTSStoGenes() |> normalizeTagCount() |> getExpressionProfiles("CTSS") |> - clusterCTSS() |> + filterLowExpCTSS() |> + distclu() |> annotateTagClusters(exampleZv9_annot) |> cumulativeCTSSdistribution("tagClusters") |> quantilePositions("tagClusters") |> diff --git a/man/flagLowExpCTSS.Rd b/man/flagLowExpCTSS.Rd new file mode 100644 index 0000000..701a31e --- /dev/null +++ b/man/flagLowExpCTSS.Rd @@ -0,0 +1,89 @@ +% Generated by roxygen2: do not edit by hand +% Please edit documentation in R/CAGEr.R +\name{flagLowExpCTSS} +\alias{flagLowExpCTSS} +\alias{flagLowExpCTSS,CAGEr-method} +\alias{flagLowExpCTSS,RangedSummarizedExperiment-method} +\alias{flagLowExpCTSS,DataFrame-method} +\alias{flagLowExpCTSS,matrix-method} +\alias{filterLowExpCTSS} +\alias{filterLowExpCTSS,CAGEr-method} +\title{Flag CTSSes based on sample expression} +\usage{ +flagLowExpCTSS( + object, + threshold = 1, + nrPassThreshold = 1, + thresholdIsTpm = TRUE +) + +\S4method{flagLowExpCTSS}{CAGEr}( + object, + threshold = 1, + nrPassThreshold = 1, + thresholdIsTpm = TRUE +) + +\S4method{flagLowExpCTSS}{RangedSummarizedExperiment}( + object, + threshold = 1, + nrPassThreshold = 1, + thresholdIsTpm = TRUE +) + +\S4method{flagLowExpCTSS}{DataFrame}( + object, + threshold = 1, + nrPassThreshold = 1, + thresholdIsTpm = TRUE +) + +\S4method{flagLowExpCTSS}{matrix}( + object, + threshold = 1, + nrPassThreshold = 1, + thresholdIsTpm = TRUE +) + +filterLowExpCTSS( + object, + threshold = 1, + nrPassThreshold = 1, + thresholdIsTpm = TRUE +) + +\S4method{filterLowExpCTSS}{CAGEr}( + object, + threshold = 1, + nrPassThreshold = 1, + thresholdIsTpm = TRUE +) +} +\arguments{ +\item{object}{An object from the \emph{CAGEr} package that contains expression +values for multiple samples.} + +\item{threshold}{Flag CTSSs with signal \verb{< threshold}.} + +\item{nrPassThreshold}{Only flag CTSSs when signal is below threshold in at +least \code{nrPassThreshold} samples.} + +\item{thresholdIsTpm}{Logical, is threshold raw tag count value (\code{FALSE}) or +normalized signal (\code{TRUE}).} +} +\value{ +\code{flagLowExpCTSS} returns a \code{\link{Rle}} vector where \code{TRUE} indicates the +index of a CTSS that passes the filter. + +\code{filterLowExpCTSS} returns the \code{CAGEr} object where the output of +\code{flagLowExpCTSS} was stored internally. +} +\description{ +Flag CTSSes for that do not pass an expression threshold in at least a given +number of samples. This is typically used to ignore CTSSes that have been +seen only once in a single sample, as they can be considered to not be +reproduced. +} +\examples{ +flagLowExpCTSS(exampleCAGEexp, threshold = 100, nrPassThreshold = 2) +} diff --git a/man/getCTSS.Rd b/man/getCTSS.Rd index 3d21325..c68b42b 100644 --- a/man/getCTSS.Rd +++ b/man/getCTSS.Rd @@ -134,9 +134,10 @@ Other CAGEr object modifiers: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()}, diff --git a/man/getExpressionProfiles.Rd b/man/getExpressionProfiles.Rd index 2af14f9..1c250d9 100644 --- a/man/getExpressionProfiles.Rd +++ b/man/getExpressionProfiles.Rd @@ -77,7 +77,7 @@ signal at individual TSS across multiple experiments. Only TSSs with normalized CAGE signal \verb{>= tpmThreshold} in at least \code{nrPassThreshold} CAGE experiments are used for expression clustering. However, CTSSs along the genome can be spatially clustered into tag clusters for each experiment -separately using the \code{\link{clusterCTSS}} function, and then aggregated across +separately using a CTSS clustering function, and then aggregated across experiments into consensus clusters using \code{\link{aggregateTagClusters}} function. Once the consensus clusters have been created, expression clustering at the level of these wider genomic regions (representing entire promoters rather diff --git a/man/normalizeTagCount.Rd b/man/normalizeTagCount.Rd index 05bab0d..9187cd1 100644 --- a/man/normalizeTagCount.Rd +++ b/man/normalizeTagCount.Rd @@ -91,9 +91,10 @@ Other CAGEr object modifiers: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()}, diff --git a/man/paraclu.Rd b/man/paraclu.Rd index e83b545..ca19f47 100644 --- a/man/paraclu.Rd +++ b/man/paraclu.Rd @@ -6,6 +6,7 @@ \alias{paraclu,CTSS-method} \alias{paraclu,GRanges-method} \alias{paraclu,SummarizedExperiment-method} +\alias{paraclu,CAGEexp-method} \title{Parametric clustering} \usage{ paraclu( @@ -57,6 +58,16 @@ paraclu( useMulticore = FALSE, nrCores = NULL ) + +\S4method{paraclu}{CAGEexp}( + object, + minStability = 1, + maxLength = 500, + keepSingletonsAbove = 0, + reduceToNonoverlapping = TRUE, + useMulticore = FALSE, + nrCores = NULL +) } \arguments{ \item{object}{A \code{\link{CTSS}}, or a \code{\link[S4Vectors:Pairs-class]{S4Vectors::Pairs}} object with positions @@ -99,10 +110,23 @@ and \code{reduceToNonoverlapping} parameters. Running Paraclu on a \code{\link{RangedSummarizedExperiment}} object will loop on each sample, and return the results as a \code{\link{GRangesList}} of \code{TagClusters}. + +Running Paraclu on a \code{\link{CAGEexp}} returnts is with the clusters stored as a +\code{GRangesList} of \code{\link{TagClusters}} objects in its metadata slot \code{tagClusters}. } \description{ -Implementation of Paraclu - parametric clustering of data attached to -sequences (\url{http://www.cbrc.jp/paraclu/}). +\code{"paraclu"} is an implementation of Paraclu algorithm for parametric +clustering of data attached to sequences (Frith \emph{et al.}, Genome Research, +2007). Since Paraclu finds clusters within clusters (unlike \code{\link{distclu}}), +additional parameters (\code{minStability}, \code{maxLength} and \code{reduceToNonoverlapping}) +can be specified to simplify the output by discarding too big clusters, +and to reduce the clusters to a final set of non-overlapping clusters. +} +\details{ +Clustering is done for every CAGE dataset within the CAGEr object separately, +resulting in a different set of tag clusters for every CAGE dataset. TCs from +different datasets can further be aggregated into a single referent set of +consensus clusters by calling the \code{\link{aggregateTagClusters}} function. } \examples{ (ctss <- CTSSnormalizedTpmGR(exampleCAGEexp,1)) @@ -112,6 +136,11 @@ CAGEr:::.paraclu(first(pair)[1:10], second(pair)[1:10]) paraclu(pair[1:10]) paraclu(ctss[1:10]) paraclu(CTSStagCountSE(exampleCAGEexp)[1:25,]) +ce <- paraclu( exampleCAGEexp, + , keepSingletonsAbove = 100 + , maxLength = 500, minStability = 1 + , reduceToNonoverlapping = TRUE) +tagClustersGR(ce, "Zf.30p.dome") } \references{ @@ -120,8 +149,43 @@ for transcription initiation in mammalian genomes.} Genome Research 2008 18(1):1-12) } \seealso{ +\code{\link{aggregateTagClusters}} + Other CAGEr clustering methods: \code{\link{consensusClustersTpm}()}, \code{\link{distclu}()} + +Other CAGEr object modifiers: +\code{\link{CTSStoGenes}()}, +\code{\link{CustomConsensusClusters}()}, +\code{\link{aggregateTagClusters}()}, +\code{\link{annotateCTSS}()}, +\code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{getCTSS}()}, +\code{\link{normalizeTagCount}()}, +\code{\link{quantilePositions}()}, +\code{\link{quickEnhancers}()}, +\code{\link{resetCAGEexp}()}, +\code{\link{summariseChrExpr}()} + +Other CAGEr clusters functions: +\code{\link{CTSScumulativesTagClusters}()}, +\code{\link{CustomConsensusClusters}()}, +\code{\link{aggregateTagClusters}()}, +\code{\link{consensusClustersDESeq2}()}, +\code{\link{consensusClustersGR}()}, +\code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{plotInterquantileWidth}()}, +\code{\link{quantilePositions}()}, +\code{\link{tagClustersGR}()} +} +\author{ +Vanja Haberle + +Charles Plessy } \concept{CAGEr clustering methods} +\concept{CAGEr clusters functions} +\concept{CAGEr object modifiers} diff --git a/man/plotInterquantileWidth.Rd b/man/plotInterquantileWidth.Rd index 27fb37c..3b1e305 100644 --- a/man/plotInterquantileWidth.Rd +++ b/man/plotInterquantileWidth.Rd @@ -74,10 +74,11 @@ Other CAGEr clusters functions: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersDESeq2}()}, \code{\link{consensusClustersGR}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{tagClustersGR}()} } diff --git a/man/quantilePositions.Rd b/man/quantilePositions.Rd index 71565f7..9173803 100644 --- a/man/quantilePositions.Rd +++ b/man/quantilePositions.Rd @@ -71,10 +71,11 @@ Other CAGEr object modifiers: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()}, \code{\link{summariseChrExpr}()} @@ -83,10 +84,11 @@ Other CAGEr clusters functions: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersDESeq2}()}, \code{\link{consensusClustersGR}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{plotInterquantileWidth}()}, \code{\link{tagClustersGR}()} } diff --git a/man/quickEnhancers.Rd b/man/quickEnhancers.Rd index 275d49c..0963f86 100644 --- a/man/quickEnhancers.Rd +++ b/man/quickEnhancers.Rd @@ -41,10 +41,11 @@ Other CAGEr object modifiers: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{resetCAGEexp}()}, \code{\link{summariseChrExpr}()} diff --git a/man/resetCAGEexp.Rd b/man/resetCAGEexp.Rd index 68ca4f8..14508ff 100644 --- a/man/resetCAGEexp.Rd +++ b/man/resetCAGEexp.Rd @@ -30,10 +30,11 @@ Other CAGEr object modifiers: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{summariseChrExpr}()} diff --git a/man/summariseChrExpr.Rd b/man/summariseChrExpr.Rd index 3f7f059..3c2f097 100644 --- a/man/summariseChrExpr.Rd +++ b/man/summariseChrExpr.Rd @@ -33,10 +33,11 @@ Other CAGEr object modifiers: \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, \code{\link{annotateCTSS}()}, -\code{\link{clusterCTSS}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, \code{\link{getCTSS}()}, \code{\link{normalizeTagCount}()}, +\code{\link{paraclu}()}, \code{\link{quantilePositions}()}, \code{\link{quickEnhancers}()}, \code{\link{resetCAGEexp}()} diff --git a/man/tagClusters.Rd b/man/tagClusters.Rd index 3e7beb6..3046e32 100644 --- a/man/tagClusters.Rd +++ b/man/tagClusters.Rd @@ -41,7 +41,7 @@ provided, interquantile width for each TC is also calculated. The object's \emph{column data}. } \description{ -Extracts tag clusters (TCs) produced by \code{\link{clusterCTSS}} function +Extracts tag clusters (TCs) for a specified CAGE experiment from a \code{\link{CAGEexp}} object. } \examples{ @@ -71,10 +71,11 @@ Other CAGEr clusters functions: \code{\link{CTSScumulativesTagClusters}()}, \code{\link{CustomConsensusClusters}()}, \code{\link{aggregateTagClusters}()}, -\code{\link{clusterCTSS}()}, \code{\link{consensusClustersDESeq2}()}, \code{\link{consensusClustersGR}()}, \code{\link{cumulativeCTSSdistribution}()}, +\code{\link{distclu}()}, +\code{\link{paraclu}()}, \code{\link{plotInterquantileWidth}()}, \code{\link{quantilePositions}()} } diff --git a/vignettes/CAGEexp.Rmd b/vignettes/CAGEexp.Rmd index 6a1b4d4..8fb04d2 100644 --- a/vignettes/CAGEexp.Rmd +++ b/vignettes/CAGEexp.Rmd @@ -447,6 +447,29 @@ results and parameters in the further steps that would normally refer to normali (denoted as tpm), will actually be raw tag counts. +CTSS flagging +------------- + +Some CTSSes have a low expression score, and are found in only a few libraries. +In non-PCR-amplified CAGE libraries, a CTSS found in a single library with an +expression score of 1 tag represents the detection of a single mRNA molecule's +5-prime end. But it could also represent the mismapping one molecule because of +a sequencing error. To flag CTSSes that have poor reproducibility support so +that other steps of the analysis can ignore them, the `filterLowExpCTSS` +function is provided. It will add an internal flag in the `filteredCTSSidx` +column of the `CTSS` objects, set to `FALSE` where expression is lower than a +given threshold in a given number of samples. This flag is later used by CTSS +clustering and export functions. + +Let's flag low-fidelity TSSs supported by less than 1 normalized tag counts in +all of the samples. + +```{r CTSS_flagging} +ce <- filterLowExpCTSS(ce, thresholdIsTpm = TRUE, nrPassThreshold = 1, threshold = 1) +CTSSnormalizedTpmGR(ce,1) +``` + + CTSS clustering --------------- @@ -455,38 +478,23 @@ transcriptional activity of that promoter (Figure \@ref(fig:CTSSbedGraph)). TS proximity of each other give rise to a functionally equivalent set of transcripts and are likely regulated by the same promoter elements. Thus, TSSs can be spatially clustered into larger transcriptional units, called tag clusters (TCs) that correspond to individual promoters. -_CAGEr_ supports three methods for spatial clustering of TSSs along the genome, two _ab initio_ -methods driven by the data itself, as well as assigning TSSs to predefined genomic regions: +_CAGEr_ supports two methods for sample-specific spatial clustering of TSSs along the genome: - * Simple distance-based clustering in which two neighbouring TSSs are joined together if they + * `distclu()`: simple distance-based clustering in which two neighbouring TSSs are joined together if they are closer than some specified distance (greedy algorithm); - * Parametric clustering of data attached to sequences based on the density of the signal + * `paraclu()`: parametric clustering of data attached to sequences based on the density of the signal [@Frith:2007], ; - * Counting TSSs and their signal in a set of user supplied genomic regions (_e.g._ annotation - derived promoter regions or other regions of interest). - -These functionalities are provided in the function `clusterCTSS()`, which accepts additional -arguments for controlling which CTSSs will be included in the clustering as well as for -refining the final set of tag clusters. - We will perform a simple distance-based clustering using 20 bp as a maximal allowed distance -between two neighbouring TSSs. Prior to clustering we will filter out low-fidelity TSSs - the -ones supported by less than 1 normalized tag counts in all of the samples. +between two neighbouring TSSs. ```{r} -ce <- clusterCTSS( ce - , threshold = 1 - , thresholdIsTpm = TRUE - , nrPassThreshold = 1 - , method = "distclu" - , maxDist = 20 - , keepSingletonsAbove = 5) +ce <- distclu(ce, maxDist = 20, keepSingletonsAbove = 5) ``` Our final set of tag clusters will not include singletons (clusters with only one TSS), unless the -normalized signal is above 5, \emph{i.e.} it is a reasonably supported TSS. The `clusterCTSS` +normalized signal is above 5, \emph{i.e.} it is a reasonably supported TSS. The CTSS clustering functions function creates a set of clusters for each sample separately; for each cluster it returns the genomic coordinates, counts the number of TSSs within the cluster, determines the (1-based) position of the most frequently used (dominant) TSS, calculates the total CAGE signal within the cluster and CAGE