goseq/DESCRIPTION0000644000175200017520000000160514136072441014424 0ustar00biocbuildbiocbuildPackage: goseq Version: 1.46.0 Date: 2019/04/26 Title: Gene Ontology analyser for RNA-seq and other length biased data Author: Matthew Young Maintainer: Matthew Young , Nadia Davidson Depends: R (>= 2.11.0), BiasedUrn, geneLenDataBase (>= 1.9.2) Imports: mgcv, graphics, stats, utils, AnnotationDbi, GO.db,BiocGenerics Suggests: edgeR, org.Hs.eg.db, rtracklayer LazyLoad: yes Description: Detects Gene Ontology and/or other user defined categories which are over/under represented in RNA-seq data License: LGPL (>= 2) Packaged: 2021-10-26 21:37:37 UTC; biocbuild biocViews: ImmunoOncology, Sequencing, GO, GeneExpression, Transcription, RNASeq git_url: https://git.bioconductor.org/packages/goseq git_branch: RELEASE_3_14 git_last_commit: 1fb5626 git_last_commit_date: 2021-10-26 Date/Publication: 2021-10-26 NeedsCompilation: no goseq/NAMESPACE0000755000175200017520000000063114136047504014140 0ustar00biocbuildbiocbuildexport(getgo,getlength,goseq,makespline,nullp,plotPWF,supportedOrganisms) importFrom(mgcv,gam,smoothCon,s,mono.con,pcls,Predict.matrix) importFrom(graphics, lines, plot) importFrom(stats, approx, binomial, dhyper, median, phyper, runif) importFrom(utils, data, relist) importFrom(AnnotationDbi, as.list, select) importFrom(GO.db,GO.db) import(BiocGenerics) import(geneLenDataBase) import(BiasedUrn) goseq/R/0000755000175200017520000000000014136047504013117 5ustar00biocbuildbiocbuildgoseq/R/getgo.R0000644000175200017520000001054314136047504014352 0ustar00biocbuildbiocbuild############################################################################# #Description: Attempts to fetch the categories specified for the given genes, genome and gene ID format #Notes: Relies on the bioconductor organism packages being installed for whatever genome you specify #Author: Matthew Young #Date Modified: 17/2/2015 getgo=function(genes,genome,id,fetch.cats=c("GO:CC","GO:BP","GO:MF")){ #Check for valid input if(any(!fetch.cats%in%c("GO:CC","GO:BP","GO:MF","KEGG"))){ stop("Invaled category specified. Categories can only be GO:CC, GO:BP, GO:MF or KEGG") } #Convert from genome ID to org.__.__.db format orgstring=as.character(.ORG_PACKAGES[match(gsub("[0-9]+",'',genome),names(.ORG_PACKAGES))]) #Multimatch or no match if(length(orgstring)!=1){ stop("Couldn't grab GO categories automatically. Please manually specify.") } #Load the library library(paste(orgstring,"db",sep='.'),character.only=TRUE) #What is the default ID that the organism package uses? coreid=strsplit(orgstring,"\\.")[[1]][3] #Now we need to convert it into the naming convention used by the organism packages userid=as.character(.ID_MAP[match(id,names(.ID_MAP))]) #Multimatch or no match if(is.na(userid) | (length(userid)!=1)){ stop("Couldn't grab GO categories automatically. Please manually specify.") } #The (now loaded) organism package contains a mapping between the internal ID and whatever #the default is (usually eg), the rest of this function is about changing that mapping to #point from categories to the ID specified #Fetch the mapping in its current format #Because GO is a directed graph, we need to get not just the genes associated with each ID, #but also those associated with its children. GO2ALLEGS does this. core2cat=NULL if(length(grep("^GO",fetch.cats))!=0){ #Get the name of the function which maps gene ids to go terms #usually this will be "GO2ALLEG" gomapFunction=.ORG_GOMAP_FUNCTION[orgstring] if(is.na(gomapFunction)) gomapFunction=.ORG_GOMAP_FUNCTION["default"] x=toTable(get(paste(orgstring,gomapFunction,sep=''))) #Keep only those ones that we specified and keep only the names # core2cat=x[x$Ontology%in%gsub("^GO:",'',fetch.cats),1:2] x[!x$Ontology%in%gsub("^GO:",'',fetch.cats),2]<-"Other" core2cat=x[,1:2] colnames(core2cat)=c("gene_id","category") } if(length(grep("^KEGG",fetch.cats))!=0){ x=toTable(get(paste(orgstring,"PATH",sep=''))) #Either add it to existing table or create a new one colnames(x)=c("gene_id","category") if(!is.null(core2cat)){ core2cat=rbind(core2cat,x) }else{ core2cat=x } } #Now we MAY have to convert the "gene_id" column to the format we are using if(coreid!=userid){ #The mapping between user id and core id, don't use the 2 object as the naming is not always consistent user2core=toTable(get(paste(orgstring,userid,sep=''))) #Throw away any user ID that doesn't appear in core2cat user2core=user2core[user2core[,1]%in%core2cat[,1],] #Make a list version of core2cat, we'll need it list_core2cat=split(core2cat[,2],core2cat[,1]) #Now we need to replicate the core IDs that need replicating list_core2cat=list_core2cat[match(user2core[,1],names(list_core2cat))] #Now we can replace the labels on this list with the user ones from user2core, #but there will be duplicates, so we have to unlist, label, then relist user2cat=split(unlist(list_core2cat,FALSE,FALSE),rep(user2core[,2],sapply(list_core2cat,length))) #Now we only want each category listed once for each entry... user2cat=sapply(user2cat,unique) ###In case you don't believe that this works as it should, here is the slow as all hell way for comparison... ###Make first list ##list_user2core=split(user2core[,1],user2core[,2]) ###Make the second ##list_core2cat=split(core2cat[,2],core2cat[,1]) ###Go through each entry in first list and expand using second... ##user2cat=sapply(list_user2core,function(u){unique(unlist(list_core2cat[u],FALSE,FALSE))}) }else{ #We don't need to convert anything (WOO!), so just make it into a list user2cat=split(core2cat[,2],core2cat[,1]) user2cat=sapply(user2cat,unique) } #remove any empty strings user2cat=lapply(user2cat,function(x){ if(length(x)>1) x=x[x!="Other"] x }) ## we don't like case sensitivity names(user2cat)<-toupper(names(user2cat)) #Now look them up return(user2cat[toupper(genes)]) } goseq/R/getlength.R0000644000175200017520000001446214136047504015232 0ustar00biocbuildbiocbuild############################################################################# #Description: Fetches gene length data for the genome and id specified. #Notes: #Author: Matthew Young, Nadia Davidson #Date Modified: 7/4/2015 getlength=function(genes,genome,id){ #geneLenDataBase contains all the gene length data, check it is installed if(!any(library()$results[,"Package"]=="geneLenDataBase")){ stop("Package geneLenDataBase is required for automatic gene length lookup.") } #Get a list of the length data files that are in the package repo=grep(".*\\..*\\.LENGTH",as.data.frame(data(package="geneLenDataBase")$results, stringsAsFactors=FALSE)$Item,ignore.case=TRUE,value=TRUE) #Is this genome/id in the local repository? if(paste(genome,id,"LENGTH",sep='.')%in%repo){ #Yes it is, so load it and group transcript lengths into genes message("Loading ",genome," length data...") data(list=paste(genome,id,"LENGTH",sep='.'),package='geneLenDataBase') data=split(get(paste(genome,id,"LENGTH",sep='.'))$Length,get(paste(genome,id,"LENGTH",sep='.'))$Gene) #Get the names of the genes used in the genome package gene_names=get(paste(genome,id,"LENGTH",sep="."))$Gene } else { ### Try to find the gene lengths from TXDB annotation packages message("Can't find ",genome,"/",id," length data in genLenDataBase...") ## Is there a TxDB package installed for the organism? installedPackages=rownames(installed.packages()) #first try looking for a match with the same type of gene IDs txdbPattern=paste("TxDb","*",genome,id,sep=".") txdbPack=installedPackages[grep(txdbPattern,installedPackages)] #if not look for the package with generic gene IDs and convert later if(length(txdbPack)==0){ orgstring=as.character(.ORG_PACKAGES[match(gsub("[0-9]+",'',genome),names(.ORG_PACKAGES))]) packageExtension=tail(strsplit(orgstring,".",fixed=T)[[1]],n=1) genericID=names(.ID_MAP[which(.ID_MAP==packageExtension)]) txdbPattern=paste("TxDb","*",genome,genericID,sep=".") txdbPack=installedPackages[grep(txdbPattern,installedPackages)] if(length(txdbPack)>0){ library(paste(orgstring,"db",sep='.'),character.only=TRUE) userid=as.character(.ID_MAP[match(id,names(.ID_MAP))]) user2core=toTable(get(paste(orgstring,userid,sep=''))) } } if(length(txdbPack)>0){ message("Found the annotation package, ",txdbPack) message("Trying to get the gene lengths from it.") library(txdbPack,character.only=TRUE) txdb<-get(txdbPack) txlens<-transcriptLengths(txdb) core2len=split(txlens$tx_len,txlens$gene_id) if(length(grep(id,txdbPattern))!=0){ data=core2len ; gene_names=names(data) } else { #We need to conver between gene IDs. Just use Matt's code from getgo.R #Now we need to replicate the core IDs that need replicating core2len=core2len[match(user2core[,1],names(core2len))] #Now we can replace the labels on this list with the user ones from user2core, #but there will be duplicates, so we have to unlist, label, then relist data=split(unlist(core2len,FALSE,FALSE),rep(user2core[,2],sapply(core2len,length))) gene_names=names(data) } } else { ## Can't use TxDB if(length(txdbPack)==0 & any(.TXDB_ORGS==genome)) { message("A TxDb annotation package exists for ",genome,". Consider installing it to get the gene lengths.",sep="") } #If it's not in geneLenDataBase, try to load the data from UCSC using rtracklayer if(!require("rtracklayer")){ stop("Can't load the rtracklayer library. Is it installed?...") } message("Trying to download from UCSC. ", "This might take a couple of minutes. ") tryCatch( { #If not, then the user will have to supply it themselves #download the relevant table from UCSC session <- browserSession() genome(session)<-genome if(id=="geneSymbol") #take the geneSymbols from refSeq genes query <- ucscTableQuery(session,"refGene") else query <- ucscTableQuery(session,id) transTable<-getTable(query) # all this below is to get the length information of each gene starts=strsplit(as.character(transTable$exonStart),",") ends=strsplit(as.character(transTable$exonEnd),",") exons=mapply(data.frame, starts, ends, SIMPLIFY=FALSE) get_length<-function(x){ sum(as.integer(as.character(x[,2]))-as.integer(as.character(x[,1])) + 1 ) } lengths=sapply(exons,get_length) # now we need to get the correct list of gene names. names2=as.character(transTable$name2) gene_names=names2 if(!any(toupper(genes)%in%toupper(names2))){ #check which column has the gene names names1=as.character(transTable$name1) if(any(toupper(genes)%in%toupper(names1))){ gene_names=names1 } } data<-split(lengths,gene_names) }, error=function(e){ stop("Length information for genome ",genome," and gene ID ",id, " is not available. You will have to specify bias.data manually.") } ) } } #In order to assign just one length to each gene, we store the Median, Min Max #and count of transcripts for each gene for later use len=data.frame(Gene=names(data),Median=sapply(data,median),Min=sapply(data,min), Max=sapply(data,max),Count=sapply(data,length)) #How many of the genes match the names the user specified with "genes" matched_frac=sum(toupper(genes)%in%toupper(gene_names))/length(genes) #None? Throw an error. if(matched_frac==0){ stop("The gene names specified do not match the gene names for genome ", genome," and ID ",id,".\n\tGene names given were: ",paste(genes[1:10],collapse=", "), "\n\tRequired gene names are: ",paste(gene_names[1:10],collapse=", ")) } #Less than 60%? Give a warning and continue if(matched_frac<.6){ warning("More than 40% of gene names specified did not match the gene names for genome ", genome," and ID ",id,". No length data will be available for these genes.\n\tGene names which failed to match were: ", paste(genes[!genes%in%gene_names][1:10],collapse=", "),"\n\tRequired gene names are: ", paste(gene_names[1:10],collapse=", ")) } #Finally, return the median transcript length for each matching gene return(len$Median[match(toupper(genes),toupper(len$Gene))]) } goseq/R/goseq.R0000644000175200017520000002335014136047504014363 0ustar00biocbuildbiocbuild############################################################################# #Description: Takes the PWF generated by nullp and uses it to generate p-values for categories specified incorporating representation bias #Notes: The automatic data fetching routines rely on the appropriate organism packages being installed. #Author: Matthew Young #Date Modified: 17/12/2010 goseq=function(pwf,genome,id,gene2cat=NULL,test.cats=c("GO:CC","GO:BP","GO:MF"),method="Wallenius",repcnt=2000,use_genes_without_cat=FALSE){ ################# Input pre-processing and validation ################### #Do some validation of input variables if(any(!test.cats%in%c("GO:CC","GO:BP","GO:MF","KEGG"))){ stop("Invalid category specified. Valid categories are GO:CC, GO:BP, GO:MF or KEGG") } if((missing(genome) | missing(id))){ if(is.null(gene2cat)){ stop("You must specify the genome and gene ID format when automatically fetching gene to GO category mappings.") } #If we're using user specified mappings, this obviously isn't a problem genome='dummy' id='dummy' } if(!any(method%in%c("Wallenius","Sampling","Hypergeometric"))){ stop("Invalid calculation method selected. Valid options are Wallenius, Sampling & Hypergeometric.") } if(!is.null(gene2cat) && (!is.data.frame(gene2cat) & !is.list(gene2cat))){ stop("Was expecting a dataframe or a list mapping categories to genes. Check gene2cat input and try again.") } #Factors are evil pwf=unfactor(pwf) gene2cat=unfactor(gene2cat) ###################### Data fetching and processing ######################## if(is.null(gene2cat)){ #When we fetch the data using getgo it will be in the list format message("Fetching GO annotations...") gene2cat=getgo(rownames(pwf),genome,id,fetch.cats=test.cats) names(gene2cat)=rownames(pwf) #Do the two rebuilds to remove any nulls cat2gene=reversemapping(gene2cat) gene2cat=reversemapping(cat2gene) }else{ #The gene2cat input accepts a number of formats, we need to check each of them in term message("Using manually entered categories.") #The options are a flat mapping (that is a data frame or matrix) or a list, where the list can be either gene->categories or category->genes if(class(gene2cat)!="list"){ #it's not a list so it must be a data.frame, work out which column contains the genes genecol_sum=as.numeric(apply(gene2cat,2,function(u){sum(u%in%rownames(pwf))})) genecol=which(genecol_sum!=0) if(length(genecol)>1){ genecol=genecol[order(-genecol_sum)[1]] warning(paste("More than one possible gene column found in gene2cat, using the one headed",colnames(gene2cat)[genecol])) } if(length(genecol)==0){ genecol=1 warning(paste("Gene column could not be identified in gene2cat conclusively, using the one headed",colnames(gene2cat)[genecol])) } othercol=1 if(genecol==1){othercol=2} #Now put it into our delicious listy format gene2cat=split(gene2cat[,othercol],gene2cat[,genecol]) #Do the appropriate builds cat2gene=reversemapping(gene2cat) gene2cat=reversemapping(cat2gene) } #!!!! #The following conditional has been flagged as a potential issue when using certain #types of input where the category names are the same as gene names (which seems like #something you should avoid anyway...). Leave it for now #!!!! #We're now garunteed to have a list (unless the user screwed up the input) but it could #be category->genes rather than the gene->categories that we want. if(sum(unique(unlist(gene2cat,use.names=FALSE))%in%rownames(pwf))>sum(unique(names(gene2cat))%in%rownames(pwf))){ gene2cat=reversemapping(gene2cat) } #Alright, we're garunteed a list going in the direction we want now. Throw out genes which we will not use gene2cat=gene2cat[names(gene2cat)%in%rownames(pwf)] #Rebuild because it's a fun thing to do cat2gene=reversemapping(gene2cat) gene2cat=reversemapping(cat2gene) ## make sure we remove duplicate entries .. e.g. see ## http://permalink.gmane.org/gmane.science.biology.informatics.conductor/46876 cat2gene=lapply(cat2gene,function(x){unique(x)}) gene2cat=lapply(gene2cat,function(x){unique(x)}) } nafrac=(sum(is.na(pwf$pwf))/nrow(pwf))*100 if(nafrac>50){ warning(paste("Missing length data for ",round(nafrac),"% of genes. Accuarcy of GO test will be reduced.",sep='')) } #Give the genes with unknown length the weight used by the median gene (not the median weighting!) pwf$pwf[is.na(pwf$pwf)]=pwf$pwf[match(sort(pwf$bias.data[!is.na(pwf$bias.data)])[ceiling(sum(!is.na(pwf$bias.data))/2)],pwf$bias.data)] ###################### Calculating the p-values ######################## # Remove all the genes with unknown GOterms unknown_go_terms=nrow(pwf)-length(gene2cat) if((!use_genes_without_cat) && unknown_go_terms>0 ){ message(paste("For",unknown_go_terms,"genes, we could not find any categories. These genes will be excluded.")) message("To force their use, please run with use_genes_without_cat=TRUE (see documentation).") message("This was the default behavior for version 1.15.1 and earlier.") pwf=pwf[rownames(pwf) %in% names(gene2cat),] } #A few variables are always useful so calculate them cats=names(cat2gene) DE=rownames(pwf)[pwf$DEgenes==1] num_de=length(DE) num_genes=nrow(pwf) pvals=data.frame(category=cats,over_represented_pvalue=NA,under_represented_pvalue=NA,stringsAsFactors=FALSE,numDEInCat=NA,numInCat=NA) if(method=="Sampling"){ #We need to know the number of DE genes in each category, make this as a mask that we can use later... num_DE_mask=rep(0,length(cats)) a=table(unlist(gene2cat[DE],FALSE,FALSE)) num_DE_mask[match(names(a),cats)]=as.numeric(a) num_DE_mask=as.integer(num_DE_mask) #We have to ensure that genes not associated with a category are included in the simulation, to do this they need an empty entry in the gene2cat list gene2cat=gene2cat[rownames(pwf)] names(gene2cat)=rownames(pwf) message("Running the simulation...") #Now do the actual simulating lookup=matrix(0,nrow=repcnt,ncol=length(cats)) for(i in 1:repcnt){ #A more efficient way of doing weighted random sampling without replacment than the built in function #The order(runif...)[1:n] bit picks n genes at random, weighting them by the PWF #The table(as.character(unlist(...))) bit then counts the number of times this random set occured in each category a=table(as.character(unlist(gene2cat[order(runif(num_genes)^(1/pwf$pwf),decreasing=TRUE)[1:num_de]],FALSE,FALSE))) lookup[i,match(names(a),cats)]=a pp(repcnt) } message("Calculating the p-values...") #The only advantage of the loop is it uses less memory... #for(i in 1:length(cats)){ # pvals[i,2:3]=c((sum(lookup[,i]>=num_DE_mask[i])+1)/(repcnt+1),(sum(lookup[,i]<=num_DE_mask[i])+1)/(repcnt+1)) # pp(length(cats)) #} pvals[,2]=(colSums(lookup>=outer(rep(1,repcnt),num_DE_mask))+1)/(repcnt+1) pvals[,3]=(colSums(lookup<=outer(rep(1,repcnt),num_DE_mask))+1)/(repcnt+1) } if(method=="Wallenius"){ message("Calculating the p-values...") #All these things are just to make stuff run faster, mostly because comparison of integers is faster than string comparison degenesnum=which(pwf$DEgenes==1) #Turn all genes into a reference to the pwf object cat2genenum=relist(match(unlist(cat2gene),rownames(pwf)),cat2gene) #This value is used in every calculation, by storing it we need only calculate it once alpha=sum(pwf$pwf) #Each category will have a different weighting so needs its own test pvals[,2:3]=t(sapply(cat2genenum,function(u){ #The number of DE genes in this category num_de_incat=sum(degenesnum%in%u) #The total number of genes in this category num_incat=length(u) #This is just a quick way of calculating weight=avg(PWF within category)/avg(PWF outside of category) avg_weight=mean(pwf$pwf[u]) weight=(avg_weight*(num_genes-num_incat))/(alpha-num_incat*avg_weight) if(num_incat==num_genes){ weight=1 } #case for the root GO terms #Now calculate the sum of the tails of the Wallenius distribution (the p-values) c(dWNCHypergeo(num_de_incat,num_incat,num_genes-num_incat,num_de,weight) +pWNCHypergeo(num_de_incat,num_incat,num_genes-num_incat,num_de,weight,lower.tail=FALSE), pWNCHypergeo(num_de_incat,num_incat,num_genes-num_incat,num_de,weight)) })) } if(method=="Hypergeometric"){ message("Calculating the p-values...") #All these things are just to make stuff run faster, mostly because comparison of integers is faster than string comparison degenesnum=which(pwf$DEgenes==1) #Turn all genes into a reference to the pwf object cat2genenum=relist(match(unlist(cat2gene),rownames(pwf)),cat2gene) #Simple hypergeometric test, one category at a time pvals[,2:3]=t(sapply(cat2genenum,function(u){ #The number of DE genes in this category num_de_incat=sum(degenesnum%in%u) #The total number of genes in this category num_incat=length(u) #Calculate the sum of the tails of the hypergeometric distribution (the p-values) c(dhyper(num_de_incat,num_incat,num_genes-num_incat,num_de)+phyper(num_de_incat,num_incat,num_genes-num_incat,num_de,lower.tail=FALSE),phyper(num_de_incat,num_incat,num_genes-num_incat,num_de)) })) } #Populate the count columns... degenesnum=which(pwf$DEgenes==1) cat2genenum=relist(match(unlist(cat2gene),rownames(pwf)),cat2gene) pvals[,4:5]=t(sapply(cat2genenum,function(u){ c(sum(degenesnum%in%u),length(u)) })) #Finally, sort by p-value pvals=pvals[order(pvals$over_represented_pvalue),] # Supplement the table with the GO term name and ontology group # but only if the enrichment categories are actually GO terms if(any(grep("^GO:",pvals$category))){ GOnames=select(GO.db,keys=pvals$category,columns=c("TERM","ONTOLOGY"))[,2:3] colnames(GOnames)<-tolower(colnames(GOnames)) pvals=cbind(pvals,GOnames) } # And return return(pvals) } goseq/R/makespline.R0000644000175200017520000000657514136047504015407 0ustar00biocbuildbiocbuild############################################################################# #Description: Fits a spline to the data (x,y) using penalized constrained least squares #Notes: #Author: Matthew Young #Date Modified: 7/4/2015 makespline=function (x, y, newX=NULL, nKnots = 6, lower_bound=10^-3){ #Should not be used outside of goseq package. Refer to the help pages for pcls in the mgcv package for more general #contstrained spline fitting. #We handle montonocily decreasing problems by reformulating them as monotonicly increasing by reflecting about "infinity" #Compare the first 10% to the last 10% ww=order(x) size=ceiling(length(y)/10) low=sum(y[ww][1:size]) hi=sum(y[ww][(length(y)-size):length(y)]) #Is the trend decreasing if(hi<=low){ #Reform as a montonicly increasing fit by reflecting about infitinity reflectionFactor=10^10 x=reflectionFactor-x } #The number of knots to use when generating the spline nKnots <- round(nKnots) if (is.null(newX)) newX <- x #We need to force the fit to go through (0,0), mono.con constructs a requirment that the LOWEST x VALUE be greater than 0 #so to force the fit through (0,0) we add a fake lowest x value of 0 by adding in a dummy point. #We use an inequality constraint rather than equality to only skew the fit enough that the spline is always >0 #This won't work if we have a monotonically decreasing function (and we don't need to it), so check that if(hi>low){ x=c(0,x) y=c(0,y) } f.ug <- gam(y ~ s(x, k = nKnots, bs = "cr"),family=binomial()) dat <- data.frame(x = x, y = y) sm <- smoothCon(s(x, k = nKnots, bs = "cr"), dat, knots = NULL)[[1]] if (length(sm$xp) < 6) warning("Few than 6 nKnots were specified!\n Possible inaccurate fitting!\n") F <- mono.con(sm$xp,TRUE) # The lower and upper parameters here don't seem to build the right constraints 100% of the time, so add them manually # ND-6/14- removing the upper limit constraint because it wasn't satisfied by initial parameters. # Hope it doesn't break the code. F$A=rbind(F$A,c(1,rep(0,ncol(F$A)-1))) #,c(rep(0,ncol(F$A)-1)),-1)) F$b=c(F$b,0) #,1) ND - should this last one be -1? G <- list(X = sm$X, C = matrix(0, 0, 0), sp = f.ug$sp, p = sm$xp, y = y, w = y * 0 + 1, Ain =F$A, bin = F$b, S = sm$S, off = 0) # do some error checking as pcls can return NaNs sometimes if this isn't true if( any((G$Ain%*%G$p - G$bin) < 0 )){ show(G$p) stop("Constraints for spline fit not satisfied by initial parameters") } #Do the actual fitting with penalized contstrained least squares p <- pcls(G) # Do some error checking for the rare case that p returns "Na" if(any(is.na(p))){ warning("The splines fit failed! Returning a flat weight distribution. This may compromise your GO analysis, please check what happened.") return(rep(mean(y),length(newX))) } #Now what we want is the value of the spline at each data point x, fv <- Predict.matrix(sm, data.frame(x = newX)) %*% p fv <- as.vector(fv) #If pcls still fails for some reason and returns negative values (or values that are so low that they'll have an effective relative weight of Inf #then we need to add a little bit to every weight to make things non-negative, the choice of minimum weight is somewhat arbitrary and serves only to #ensure that we don't have positive weights that will be infinitely prefered as a ratio. if(min(fv) gene bins.",ylab="Proportion DE",...){ #We shouldn't try and plot NAs obviously... w=!is.na(pwf$bias.data) o=order(pwf$bias.data[w]) #What is the total range in the fit? rang=max(pwf$pwf,na.rm=TRUE)-min(pwf$pwf,na.rm=TRUE) if(rang==0 & binsize=="auto") binsize=1000 if(binsize=="auto"){ #A low number of starting genes to bin, usually 100 binsize=max(1,min(100,floor(sum(w)*.08))) resid=rang #Turn off warnings till we've worked out what we're doing oldwarn=options()$warn options(warn=-1) #Keep increasing the number of genes in each bin until the scatter around the lines reaches the cutoff. #Stop if we reach only 10 bins for the entire plot while(binsize<=floor(sum(w)*.1) & resid/rang>.001){ binsize=binsize+100 #Assign each gene a "bin number" splitter=ceiling(1:length(pwf$DEgenes[w][o])/binsize) #Determine the percentage DE in each bin de=sapply(split(pwf$DEgenes[w][o],splitter),mean) #Determine the average length in each bin binlen=sapply(split(as.numeric(pwf$bias.data[w][o]),splitter),mean) #Calculate the residuals, how much the binned data deviates from the PWF resid=sum((de-approx(pwf$bias.data[w][o],pwf$pwf[w][o],binlen)$y)^2)/length(binlen) } options(warn=oldwarn) }else{ #Assign each gene a "bin number" splitter=ceiling(1:length(pwf$DEgenes[w][o])/binsize) #Determine the percentage DE in each bin de=sapply(split(pwf$DEgenes[w][o],splitter),mean) #Determine the average length in each bin binlen=sapply(split(as.numeric(pwf$bias.data[w][o]),splitter),mean) } #Now we've settled on a binsize, plot it #Did the user specify the labels? If so we can't put in the defaults or they'll be used twice and errors result. xlab=gsub("",as.character(binsize),xlab) if("xlab"%in%names(list(...))){ if("ylab"%in%names(list(...))){ plot(binlen,de,...) }else{ plot(binlen,de,ylab=ylab,...) } }else if("ylab"%in%names(list(...))){ plot(binlen,de,xlab=xlab,...) }else{ plot(binlen,de,xlab=xlab,ylab=ylab,...) } #Add the PWF lines(pwf$bias.data[w][o],pwf$pwf[w][o],col=pwf_col,lwd=pwf_lwd) } goseq/R/pp.R0000644000175200017520000000056314136047504013665 0ustar00biocbuildbiocbuild############################################################################# #Description: Prints progress through a loop #Notes: #Author: Matthew Young #Date Modified: 29/11/2012 pp=function(total,count,i=i){ if(missing(count)){count=evalq(i,envir=parent.frame())} if(missing(total)){total=evalq(stop,envir=parent.frame())} cat(round(100*(count/total)),"% \r") } goseq/R/reversemapping.R0000644000175200017520000000063014136047504016270 0ustar00biocbuildbiocbuild############################################################################# #Description: Reverses the direction of a bidirectional mapping, removes an entries pointing to NULL #Notes: #Author: Matthew Young #Date Modified: 13/12/2010 reversemapping=function(map){ tmp=unlist(map,use.names=FALSE) names(tmp)=rep(names(map),times=as.numeric(summary(map)[,1])) return(split(names(tmp),as.vector(tmp))) } goseq/R/supportedOrganisms.R0000644000175200017520000000403214136047504017151 0ustar00biocbuildbiocbuild############################################################################# #Description: Lists which genomes and gene IDs are supported by goseq (length and GO terms) #Notes: #Author: Nadia Davidson #Date Modified: 2016/10/04 supportedOrganisms=function(){ require("rtracklayer") #Start by getting the geneLenDataBase supported genomes and geneIDs geneIDs=supportedGeneIDs() #The names of the supported geneIDs is hard-coded so we can remove things like #"refGene" which actually correspond to an Entrez gene ID in geneLenDataBase uniqueIDs=c("knownGene","vegaGene","ensGene","geneSymbol") geneIDs=geneIDs[geneIDs$tablename %in% uniqueIDs,] #extract the corresponding genome IDs and rearrange genomes=strsplit(geneIDs$AvailableGenomes,",") id=rep(geneIDs$tablename,sapply(genomes,length)) description=(rep(geneIDs$GeneID,sapply(genomes,length))) length_supported=rep(TRUE,length(id)) genomes=unlist(genomes) tab_temp=data.frame(genomes,id,description,length_supported) ## Is there GO annotation package for that genome and gene ID? genome_supported_go=gsub("[0-9]+","",genomes) %in% names(.ORG_PACKAGES) id_supported_go=id %in% names(.ID_MAP) go_supported=genome_supported_go & id_supported_go tab_temp=data.frame(tab_temp,go_supported) colnames(tab_temp)<-c("Genome","Id","Id Description","Lengths in geneLeneDataBase", "GO Annotation Available") ### Add in the newer genomes which aren't supported for length, but are for GO terms base=unfactor(ucscGenomes())$db other_genomes = base[ ! base %in% genomes ] new_genomes=other_genomes[gsub("[0-9]+","",other_genomes) %in% names(.ORG_PACKAGES)] new_genomes=c(new_genomes,names(.ORG_PACKAGES)[!names(.ORG_PACKAGES) %in% gsub("[0-9]+","",c(new_genomes,genomes))]) tab_new=data.frame(new_genomes,"","",FALSE,TRUE) colnames(tab_new)<-c("Genome","Id","Id Description","Lengths in geneLeneDataBase", "GO Annotation Available") tab_supported=rbind(tab_temp,tab_new) tab_supported[order(tab_supported$Genome),] } goseq/R/zzz.R0000644000175200017520000000335314136047504014103 0ustar00biocbuildbiocbuild# ZZZ.R .onAttach <- function(libname, pkgname) { if(.Platform$OS.type=="windows" && .Platform$GUI=="Rgui" ) { winMenuAddItem("Vignettes","goseq","shell.exec(system.file(\"doc\",\"goseq.pdf\",package=\"goseq\"))") } } #These two variables are required for automatic fetching of categories to function. Their purpose is to take the UCSC genome and gene ID values given when looking up length data and convert them to the names used for the same organism and gene identifier in the organism packages. #Mappings that are primarily required by getgo, the purpose of this is to convert the UCSC genome IDs, to the bioconductor organism names, e.g. "mm"->"org.Mm." .ORG_PACKAGES=paste("org.",c("Ag.eg","At.tair","Bt.eg","Ce.eg","Cf.eg","Dm.eg","Dr.eg","EcK12.eg","EcSakai.eg","Gg.eg","Hs.eg","Mm.eg","Mmu.eg","Pf.plasmo","Pt.eg","Rn.eg","Sc.sgd","Ss.eg","Xl.eg"),sep='') names(.ORG_PACKAGES)=c("anoGam","Arabidopsis","bosTau","ce","canFam","dm","danRer","E. coli K12","E. coli Sakai","galGal","hg","mm","rheMac","Malaria","panTro","rn","sacCer","susScr","xenTro") #These are the only formats supported by getgo at the moment, the purpose is to convert the USCC gene #ID formats, to the shorthand used by the bioconductor organism packages, .e.g. "refGene"->"ENSEMBL" .ID_MAP=c("eg","eg","ENSEMBL","SYMBOL","sgd","plasmo","tair") names(.ID_MAP)=c("knownGene","refGene","ensGene","geneSymbol","sgd","plasmo","tair") #Below are the exceptions to the function name for gene to go term mappings .ORG_GOMAP_FUNCTION=c("GO2ALLEGS","GO2ALLTAIRS","GO2ALLORFS","GO2ALLORFS") names(.ORG_GOMAP_FUNCTION)=c("default","org.At.tair","org.Pf.plasmo","org.Sc.sgd") #TxDb Length databases .TXDB_ORGS=c("ce6","dm3","hg18","hg19","hg38","mm10","mm9","rn4","rn5","sacCer2","sacCer3") goseq/build/0000755000175200017520000000000014136072441014013 5ustar00biocbuildbiocbuildgoseq/build/vignette.rds0000644000175200017520000000031214136072441016346 0ustar00biocbuildbiocbuildb```b`add`b2 1# 'L/N- +GK(+fbZ&5h0X" ,LHXsS楀aM wjey~L6̜T!%ps QY_/( @hrNb1GRKҊAm&goseq/data/0000755000175200017520000000000014136047504013627 5ustar00biocbuildbiocbuildgoseq/data/genes.RData0000644000175200017520000024744514136047504015665 0ustar00biocbuildbiocbuildݻJn4HBz Rȓ!9Ǖ%:|X@>TCys?o77cPoM_?//ow25gj_اǯsw}٫wqu~b}sw5x;tO|gɵ}uOdw~yxg~MήyZ~㸪?^{w+;~Omj_ޙo_w'͝9O}vWow={;޾iO?9>>wS׼N\Kw eS^SvܝIyvo8Wz~Wz5vu>;ygoe#}]lO?}gs3c={_'e{O'u'=|bz_Wٓ{z_uՇ]Ć~u/xz]k<%C}k:UʜWξuO'W{5Ľ=-#_=o=[{yﮛw'tS }Ju?]4+[[?;wɼNyww+>.WO'O;ïdOsOͧ~:z/xg_{Oʐw?Ǟǟ>S>˧vз7mWjy>^swWVd;- gs+xʾO=Ч߰Oߵ??w+mxwtݯ,'d{;wos##>]#]o޹?]>!}Ot'쾾}5^t:~sg;kMtwΟkʵ=dO|W>͕9>ٷ?ɗ+׸vw{{ι]y.< A+6;oqЉ\^{?z}?\2ث9޹S^)յwܕk_o%WϺ^'^{Ǿ7lWsSqo|W}E~;] ]pWܕWe]{?eqsw';yGn>}Wz^j+Wl'=ﮬOO^ǻ׻{7l;2O}j\qu޵<>}޻:D64;9> Sx9~[we׿c;tO^]o?\w9W;׸S|s_Sm;~ծ;{'uѻ=|{ܑ'ǻ {ӟB=Jw^_&~>u=˯tvٓ]T_jWtŻ-꾼rtѧٓ7Oػ߲~_]߿7ɎWk]ze\:߲E>f۶;kܱ?yW#~'zNI]z|Ro~r;;w}͝5J?toOuO]}~ rWĞxB?]fm]s="W~?u\Ogx#OZO_sc}r+}<;\<ҭg{V"߫k]xw~GOm5?"ɚlWu? ;6ٕk]^:_w|wWsOuޑﬣlO˫oZyjܙ[<B^Os~}'ׯvΝ{lgcXOޟιcǾsͿʎrd;6ֻ{I}+ӵpdٻ6]9w_?7?=^ד仿~Wݽz'two}G\ɜ}zW}xڶַ_: wϲ_ώ+lο|u{Wkko5|WO|__yu}ϮqEտgxOr޳3+sܕʏ+sܱYߕٿyL;7㊏ok4߾sޫ<޵9w}O'Ow'?LD6'ƚk滻7;ʻ67~sw'''rl=/˷~{uϝl$>kw|OO٘ڍWd7']k>ww{<ή6ʏ{g꣼s<軟k='o\Z|=~rwm;uJ|~xJ~^٫D'9~OWd5|;O;;}{oW}?iscdϞ=egޕ?u\wm+{rWEWS~Uwŕ~d sOt?ʚcsݫǯz8OwIzuO9;O밫+tͫ>8Wd=}wd+ILֵ+{vޓ2sO?{OOO缳v^o;Ԟi\߹;^'wpeWd\Is~vuwpe]-oچW'j^]mtwm:|r't~{3=<;ǟ}ƿ~:Q~C]]:{*>>޽ާw|{o>uwur\OOɬOb^'O;7?mi>޾sOt':߸W箍pE_S޷ ߌ|r|_o{U~|{5'o^x9)=Y'_xW34~?ѫ:Mb#ߵ^ֺ/\mz+ӽ+󷧎O_rulo}\o_>hӰo2>z~?[0wip}x_S_uejƺqF<8;w0N`.Ibu'0>ڴxkowҦuu҆eymkL8{Y}.aݖiXﻵy·o+r}g1w;Yٷ}ۊWU}>j Ӷa>h;vqoɓ\ZڀۊI27npK >Cv1- ^ۻ 13Nmk[NQ0}%qp-}S֊mGanhNjJ_7Ӝ}2qX'-p&.a솶t}^k ffZ|o>h#vCmnMI>˷zJsNmXj>(13KI4[G| bs=W%O R^}V,r|HǞ/=~R%e~j"7,skmdZY{ޔ_% E\,]j<$|@"&ad=ag͡Q**VKмeL J_޷AVAn+WÝ)mUXeJbA׉+`gX8[a s+1 rԭU5*(9Z}l>!a[Jxǰv촑Mz>gsUÃWN۳T$'@Koz+ޘO&4Z9Y>"~ FK_Bז، کI 9˦ϰȫ$z'"9Q fSַR3Ӭ`t,S;Doԋ Sp|"}p'7t^"i :Ƌpbcݠ` jNr sbt0wE-+tI !R_~CIS&c.Q)ti#}' :l(cI.m'9!j(s S/en@nOh7e$8Is9$#rLtfix߹AINS}x&faƓtޕ!sF4>07797X>k.S(DAYR\]]~P궝m` Ë́6r'PǪrݞנhy/K[[jcD; }dqԇu5XAGk$'Pږ2ɢ+XaafW953BMyZ)D(u\'f7:}mlPRw$-Z4) r&<[]rRXr蠌khvX/r3R5#+zUI'[-䅞VP-,7^+̇A U Vm)SFeq} +Wܐ?9W]>_[1]-[8[{ҁɡ6繙wW#uQP8_~9,t "(nԊâ(A-װ#T[JrԋNs) [f9A te hQ6uӚ䤖Tj=Oп>>YFjIVMaB!{a"ab jF#01蒃N`x;kmVyjjnt>r62XEX)Um/*cX.fD0]AJ9Aj֞4I9e)*zeP5D]m-e `eXYI9DAd 2lqddַ6Iy̓kXD\;!jsmWYJ)e*thͥ S:I>MMAAu\ Om7 -q1ɤ}^vq&!ti*׉ձc|1P.sK5IdJMĬLz=# *bTt0iOD2ig\sb5/RGjlOD sAym$wYBmz7@薌́d¸/22pؚсiAtKXйGJa$tEdմ'UgT~ U2 ۮp̒jXB}.98I_㮊]-tkiJԞO7XQԿ[y ˀf;Ѻ J[nI3Pl% O+ fʐ& g'wc`nuvmm٤2JGUIΞ\.k (a醵Syg/d@ՎYt>֋|JZMVwT+SN$$[4_S  `fri-7b5?203Ƃa>q'(5em}ɤR4U. cKS! 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Each element of your vector should be named by a unique gene identifier. \item \texttt{Differentially expressed genes}: each element of your vector should be either a 1 or a 0, where 1 indicates that the gene is differentially expressed and 0 that it is not. \end{enumerate} If the organism, gene identifier or category test is currently not natively supported by \texttt{goseq}, it will also be necessary to supply additional information regarding the genes length and/or the association between categories and genes. \np Bioconductor R packages such as \texttt{Rsubread} allow for the summarization of mapped reads into a table of counts, such as reads per gene. From there, several packages exist for performing differential expression analysis on summarized data (eg. \texttt{edgeR}~\citep{Robinson07, Robinson08, Robinson10}). \texttt{goseq} will work with any method for determining differential expression and as such differential expression analysis is outside the scope of this document, but in order to facilitate ease of use, we will make use of the \texttt{edgeR} package to calculate differentially expressed (DE) genes in all the case studies in this document. \section{Reading data} We assume that the user can use appropriate in-built \texttt{R} functions (such as \texttt{read.table} or \texttt{scan}) to obtain two vectors, one containing all genes assayed in the RNA-seq experiment, the other containing all genes which are DE. If we assume that the vector of genes being assayed is named \texttt{assayed.genes} and the vector of DE genes is named \texttt{de.genes} we can construct a named vector suitable for use with \texttt{goseq} using the following: <>= gene.vector=as.integer(assayed.genes%in%de.genes) names(gene.vector)=assayed.genes head(gene.vector) @ It may be that the user can already read in a vector in this format, in which case it can then be immediately used by \texttt{goseq}. \section{GO testing of RNA-seq data} To begin the analysis, \texttt{goseq} first needs to quantify the length bias present in the dataset under consideration. This is done by calculating a Probability Weighting Function or PWF which can be thought of as a function which gives the probability that a gene will be differentially expressed (DE), based on its length alone. The PWF is calculated by fitting a monotonic spline to the binary data series of differential expression (1=DE, 0=Not DE) as a function of gene length. The PWF is used to weight the chance of selecting each gene when forming a null distribution for GO category membership. The fact that the PWF is calculated directly from the dataset under consideration makes this approach robust, only correcting for the length bias present in the data. For example, if \texttt{goseq} is run on a microarray dataset, for which no length bias exists, the calculated PWF will be nearly flat and all genes will be weighted equally, resulting in no length bias correction. \np In order to account for the length bias inherent to RNA-seq data when performing a GO analysis (or other category based tests), one cannot simply use the hypergeometric distribution as the null distribution for category membership, which is appropriate for data without DE length bias, such as microarray data. GO analysis of RNA-seq data requires the use of random sampling in order to generate a suitable null distribution for GO category membership and calculate each categories significance for over representation amongst DE genes. \np However, this random sampling is computationally expensive. In most cases, the Wallenius distribution can be used to approximate the true null distribution, without any significant loss in accuracy. The \texttt{goseq} package implements this approximation as its default option. The option to generate the null distribution using random sampling is also included as an option, but users should be aware that the default number of samples generated will not be enough to accurately call enrichment when there are a large number of go terms. \np Having established a null distribution, each GO category is then tested for over and under representation amongst the set of differentially expressed genes and the null is used to calculate a p-value for under and over representation. \section{Natively supported Gene Identifiers and category tests} \texttt{goseq} needs to know the length of each gene, as well as what GO categories (or other categories of interest) each gene is associated with. \texttt{goseq} relies on the UCSC genome browser to provide the length information for each gene. However, because the process of fetching the length of every transcript is slow and bandwidth intensive, \texttt{goseq} relies on an offline copy of this information stored in the data package \texttt{geneLenDataBase}. To see which genome/gene identifier combinations are in the local database, simply run: <>= supportedOrganisms() @ The leftmost columns in the output of this command list the genomes and gene identifiers respectively. If length data exists in the local database it is indicated in the second last column. If your genome/ID combination is not in the local database, it may be downloaded from the UCSC genome browser or taken from a \texttt{TxDb} annotation package (if installed). If your genome/ID combination is not found in any database, you will have to manually specify the gene lengths. We encourage all users to manually specify their gene lengths if provided by upstream summarization programs. e.g. \texttt{featureCounts}, as these lengths will be more accurate. \np In order to link GO categories to genes, \texttt{goseq} uses the organism packages from Bioconductor. These packages are named org...db, where is a short string identifying the genome and is a short string identifying the gene identifier. Currently, \texttt{goseq} will automatically retrieve the mapping between GO categories and genes from the relevant package (as long as it is installed) for commonly used genome/ID combinations. If GO mappings are not automatically available for your genome/ID combination, you will have to manually specify the relationship between genes and categories. Although the Genome/ID naming conventions used by the organism packages differ from the UCSC, \texttt{goseq} is able to convert between the two, so the user need only ever specify the UCSC genome/ID in most cases. The final column indicates whether the Genome/ID combination is supported for GO categories. \section{Non-native Gene Identifier or category test} If the organism, Gene Identifier or category test you wish to perform is not in the native \texttt{goseq} database, you will have to supply one or all of the following: \begin{itemize} \item \texttt{Length data}: the length of each gene in your gene identifier format. \item \texttt{Category mappings}: the mapping (usually many-to-many) between the categories you wish to test for over/under representation amongst DE genes and genes in your gene identifier format. \end{itemize} \subsection{Length data format} The length data must be formatted as a numeric vector, of the same length as the main named vector specifying gene names/DE genes. Each entry should give the length of the corresponding gene in bp. If length data is unavailable for some genes, that entry should be set to NA. \subsection{Category mapping format} The mapping between category names and genes should be given as a data frame with two columns. One column should contain the gene IDs and the other the name of an associated category. As the mapping between categories and genes is usually many-to-many, this data frame will usually have multiple rows with the same gene name and category name. \np Alternatively, mappings between genes and categories can be given as a list. The names of list entries should be gene IDs and the entries themselves should be a vector of category names to which the gene ID corresponds. \subsection{Some additional tips} Any organism for which there is an annotation on either Ensembl or the UCSC, can be easily turned into length data using the GenomicFeatures package. To do this, first create a TranscriptDb object using either makeTxDbFromBiomart or makeTxDbFromUCSC (see the help in the GenomicFeatures package on using these commands). Once you have a transcriptDb object, you can get a vector named by gene ID containing the median transcript length of each gene simply by using the command. <>= txsByGene=transcriptsBy(txdb,"gene") lengthData=median(width(txsByGene)) @ The relationship between gene identifier and GO category can usually be obtained from the Gene Ontology website (www.geneontology.org) or from the NCBI. Additionally, the bioconductor AnnotationDbi library has recently added a function "makeOrgPackageFromNCBI", which can be used to create an organism package from within R, using the NCBI data. Once created, this package can then be used to obtain the mapping between genes and gene ontology. \section{Case study: Prostate cancer data} \subsection{Introduction} This section provides an analysis of data from an RNA-seq experiment to illustrate the use of \texttt{goseq} for GO analysis. \np This experiment examined the effects of androgen stimulation on a human prostate cancer cell line, LNCaP. The data set includes more than 17 million short cDNA reads obtained for both the treated and untreated cell line and sequenced on Illumina's 1G genome analyzer. \np For each sample we were provided with the raw 35 bp RNA-seq reads from the authors. For the untreated prostate cancer cells (LNCaP cell line) there were 4 lanes totaling ~10 million, 35 bp reads. For the treated cells there were 3 lanes totaling ~7 million, 35 bp reads. All replicates were technical replicates. Reads were mapped to NCBI version 36.3 of the human genome using bowtie. Any read with which mapped to multiple locations was discarded. Using the ENSEMBL 54 annotation from biomart, each mapped read was associated with an ENSEMBL gene. This was done by associating any read that overlapped with any part of the gene (not just the exons) with that gene. Reads that did not correspond to genes were discarded. \subsection{Source of the data} The data set used in this case study is taken from \citep{Li:2008p113} and was made available from the authors upon request. \subsection{Determining the DE genes using \texttt{edgeR}} To begin with, we load in the text data and convert it the appropriate \texttt{edgeR} DGEList object. <>= library(edgeR) table.summary=read.table(system.file("extdata","Li_sum.txt",package='goseq'), sep='\t',header=TRUE,stringsAsFactors=FALSE) counts=table.summary[,-1] rownames(counts)=table.summary[,1] grp=factor(rep(c("Control","Treated"),times=c(4,3))) summarized=DGEList(counts,lib.size=colSums(counts),group=grp) @ Next, we use \texttt{edgeR} to estimate the biological dispersion and calculate differential expression using a negative binomial model. <>= disp=estimateCommonDisp(summarized) disp$common.dispersion tested=exactTest(disp) topTags(tested) @ Finally, we Format the DE genes into a vector suitable for use with \texttt{goseq} <>= genes=as.integer(p.adjust(tested$table$PValue[tested$table$logFC!=0], method="BH")<.05) names(genes)=row.names(tested$table[tested$table$logFC!=0,]) table(genes) @ \subsection{Determining Genome and Gene ID} In order to allow for automatic data retrieval, the user has to tell \texttt{goseq} what genome and gene ID format were used to summarize the data. In our case we will use the hg19 build of the human genome, we check what code this corresponds to by running: <>= head(supportedOrganisms()) @ Which lists the genome codes in the far left column, headed ``Genome''. As we are using ``hg19" and we also know that we used ENSEMBL Gene ID to summarize our read data, we check what code this corresponds to by running: <>= supportedOrganisms()[supportedOrganisms()$Genome=="hg19",] @ The gene ID codes are listed in the column second from left, titled ``Id''. We find that our gene ID code is ``ensGene". We will use these strings whenever we are asked for a genome or id. If the gene ID is missing for your Genome (for example this is the case for hg38), then the genome is not supported in the geneLengthDatabase. Gene lengths will either be automatically fetched from TxDB, UCSC or you will need to provide them manually. Supported Gene IDs to automatically fetch GO terms should usually either be Entrez (``knownGene"), Ensembl (``ensGene") or gene symbols (``geneSymbol"). \subsection{GO analysis} \subsubsection{Fitting the Probability Weighting Function (PWF)} We first need to obtain a weighting for each gene, depending on its length, given by the PWF. As you may have noticed when running supportedGenomes or supportedGeneIDs, length data is available in the local database for our gene ID, ``ensGene" and our genome, ``hg19". We will let \texttt{goseq} automatically fetch this data from its databases. <>= pwf=nullp(genes,"hg19","ensGene") head(pwf) @ \texttt{nullp} plots the resulting fit, allowing verification of the goodness of fit before continuing the analysis. Further plotting of the pwf can be performed using the \texttt{plotPWF} function. \np The output of nullp contains all the data used to create the PWF, as well as the PWF itself. It is a data frame with 3 columns, named "DEgenes", "bias.data" and "pwf" with the rownames set to the gene names. Each row corresponds to a gene with the DEgenes column specifying if the gene is DE (1 for DE, 0 for not DE), the bias.data column giving the numeric value of the DE bias being accounted for (usually the gene length or number of counts) and the pwf column giving the genes value on the probability weighting function. \subsubsection{Using the Wallenius approximation} To start with we will use the default method, to calculate the over and under expressed GO categories among DE genes. Again, we allow \texttt{goseq} to fetch data automatically, except this time the data being fetched is the relationship between ENSEMBL gene IDs and GO categories. %This hides the output of goseq <>= GO.wall=goseq(pwf,"hg19","ensGene") head(GO.wall) @ The resulting object is ordered by GO category over representation amongst DE genes. \subsubsection{Using random sampling} It may sometimes be desirable to use random sampling to generate the null distribution for category membership. For example, to check consistency against results from the Wallenius approximation. This is easily accomplished by using the \texttt{method} option to specify sampling and the \texttt{repcnt} option to specify the number of samples to generate: <>= GO.samp=goseq(pwf,"hg19","ensGene",method="Sampling",repcnt=1000) @ <>= head(GO.samp) @ You will notice that this takes far longer than the Wallenius approximation. Plotting the p-values against one another, we see that there is little difference between the two methods. However, the accuracy of the sampling method is limited by the number of samples generated, \texttt{repcnt}, such that very low p-values will not be correctly calculated. Significantly enriched GO terms may then be missed after correcting for multiple testing. <>= plot(log10(GO.wall[,2]), log10(GO.samp[match(GO.wall[,1],GO.samp[,1]),2]), xlab="log10(Wallenius p-values)",ylab="log10(Sampling p-values)", xlim=c(-3,0)) abline(0,1,col=3,lty=2) @ \subsubsection{Ignoring length bias} \texttt{goseq} also allows for one to perform a GO analysis without correcting for RNA-seq length bias. In practice, this is only useful for assessing the effect of length bias on your results. You should NEVER use this option as your final analysis. If length bias is truly not present in your data, \texttt{goseq} will produce a nearly flat PWF and no length bias correction will be applied to your data and all methods will produce the same results. \np However, if you still wish to ignore length bias in calculating GO category enrichment, this is again accomplished using the \texttt{method} option. <>= GO.nobias=goseq(pwf,"hg19","ensGene",method="Hypergeometric") head(GO.nobias) @ Ignoring length bias gives very different results from a length bias corrected analysis. <>= plot(log10(GO.wall[,2]), log10(GO.nobias[match(GO.wall[,1],GO.nobias[,1]),2]), xlab="log10(Wallenius p-values)", ylab="log10(Hypergeometric p-values)", xlim=c(-3,0), ylim=c(-3,0)) abline(0,1,col=3,lty=2) @ \subsubsection{Limiting GO categories and other category based tests} By default, \texttt{goseq} tests all three major Gene Ontology branches; Cellular Components, Biological Processes and Molecular Functions. However, it is possible to limit testing to any combination of the major branches by using the \texttt{test.cats} argument to the goseq function. This is done by specifying a vector consisting of some combination of the strings ``GO:CC", ``GO:BP" and ``GO:MF". For example, to test for only Molecular Function GO categories: <>= GO.MF=goseq(pwf,"hg19","ensGene",test.cats=c("GO:MF")) head(GO.MF) @ Native support for other category tests, such as KEGG pathway analysis are also made available via this argument. See the man \texttt{goseq} function man page for up to date information on what category tests are natively supported. \subsubsection{Making sense of the results} Having performed the GO analysis, you may now wish to interpret the results. If you wish to identify categories significantly enriched/unenriched below some p-value cutoff, it is necessary to first apply some kind of multiple hypothesis testing correction. For example, GO categories over enriched using a .05 FDR cutoff \citep{Benjamini95} are: <>= enriched.GO=GO.wall$category[p.adjust(GO.wall$over_represented_pvalue, method="BH")<.05] head(enriched.GO) @ Unless you are a machine, GO accession identifiers are probably not very meaningful to you. Information about each term can be obtained from the Gene Ontology website, \url{http://www.geneontology.org/}, or using the R package \texttt{GO.db}. <>= library(GO.db) for(go in enriched.GO[1:10]){ print(GOTERM[[go]]) cat("--------------------------------------\n") } @ \subsubsection{Understanding goseq internals} The situation may arise where it is necessary for the user to perform some of the data processing steps usually performed automatically by \texttt{goseq} themselves. With this in mind, it will be useful to step through the preprocessing steps performed automatically by \texttt{goseq} to understand what is happening. \np To start with, when \texttt{nullp} is called, \texttt{goseq} uses the genome and gene identifiers supplied to try and retrieve length information for all genes given to the \texttt{genes} argument. To do this, it retrieves the data from the database of gene lengths maintained in the package \texttt{geneLenDataBase}. This is performed by the \texttt{getlength} function in the following way: <>= len=getlength(names(genes),"hg19","ensGene") length(len) length(genes) head(len) @ After some data cleanup, the length data and the DE data is then passed to the \texttt{makespline} function to produce the PWF. The \texttt{nullp} returns a data frame which has 3 columns, the original DEgenes vector, the length bias data (in a column called bias.data) and the PWF itself (in a column named pwf). The names of the genes are also kept in this data frame as the names of the rows. If length data could not be obtained for a certain gene the corresponding entries in the "bias.data" and "pwf" columns are set to NA. \np Next we call the \texttt{goseq} function to determine over/under representation of GO categories amongst DE genes. When we do this, \texttt{goseq} looks for the appropriate organism package and tries to obtain the mapping from genes to GO categories from it. This is done using the \texttt{getgo} function as follows: <>= go=getgo(names(genes),"hg19","ensGene") length(go) length(genes) head(go) @ Note that some of the gene categories have been returned as "NULL". This means that a GO category could not be found in the database for one of the genes. In the \texttt{goseq} command, enrichment will only be calculated using genes with a GO category by default. However, in older versions of goseq (below 1.15.2), we counted all genes. i.e. genes with no categories still counted towards the total number of gene outside of any single category. It is possible to switch between these two behaviors using the \texttt{use\_genes\_without\_cat} flag in \texttt{goseq}. The first thing the getgo function does is to convert the UCSC genome/ID namings into the naming convention used by the organism packages. This is done using two hard coded conversion vectors that are included in the \texttt{goseq} package but usually hiden from the user. <>= goseq:::.ID_MAP goseq:::.ORG_PACKAGES @ It is just as valid to run the length and GO category fetching as separate steps and then pass the result to the \texttt{nullp} and \texttt{goseq} functions using the \texttt{bias.data} and \texttt{gene2cat} arguments. Thus the following two blocks of code are equivalent: <>= pwf=nullp(genes,"hg19","ensGene") go=goseq(pwf,"hg19","ensGene") @ and <>= gene_lengths=getlength(names(genes),"hg19","ensGene") pwf=nullp(genes,bias.data=gene_lengths) go_map=getgo(names(genes),"hg19","ensGene") go=goseq(pwf,"hg19","ensGene",gene2cat=go_map) @ \subsection{KEGG pathway analysis} In order to illustrate performing a category test not present in the \texttt{goseq} database, we perform a KEGG pathway analysis. For human, the mapping from KEGG pathways to genes are stored in the package org.Hs.eg.db, in the object org.Hs.egPATH. In order to test for KEGG pathway over representation amongst DE genes, we need to extract this information and put it in a format that \texttt{goseq} understands. Unfortunately, the org.Hs.eg.db package does not contain direct mappings between ENSEMBL gene ID and KEGG pathway. Therefore, we have to construct this map by combining the ENSEMBL <-> Entrez and Entrez <-> KEGG mappings. This can be done using the following code: <>= # Get the mapping from ENSEMBL 2 Entrez en2eg=as.list(org.Hs.egENSEMBL2EG) # Get the mapping from Entrez 2 KEGG eg2kegg=as.list(org.Hs.egPATH) # Define a function which gets all unique KEGG IDs # associated with a set of Entrez IDs grepKEGG=function(id,mapkeys){unique(unlist(mapkeys[id],use.names=FALSE))} # Apply this function to every entry in the mapping from # ENSEMBL 2 Entrez to combine the two maps kegg=lapply(en2eg,grepKEGG,eg2kegg) head(kegg) @ Note that this step is quite time consuming. The code written here is not the most efficient way of producing this result, but the logic is much clearer than faster algorithms. The source code for getgo contains a more efficient routine. \np We produce the PWF as before. Then, to perform a KEGG analysis, we simply make use of the \texttt{gene2cat} option in \texttt{goseq}: <>= pwf=nullp(genes,"hg19","ensGene") KEGG=goseq(pwf,gene2cat=kegg) head(KEGG) @ Note that we do not have to tell the goseq function what organism and gene ID we are using as we are manually supplying the mapping between genes and categories. \np KEGG analysis is shown as an illustration of how to supply your own mapping between gene ID and category, KEGG analysis is actually natively supported by GOseq and we could have performed it with the following code. <>= pwf=nullp(genes,'hg19','ensGene') KEGG=goseq(pwf,'hg19','ensGene',test.cats="KEGG") head(KEGG) @ Noting that this time it was necessary to tell the goseq function that we are using HG19 and ENSEMBL gene ID, as the function needs this information to automatically construct the mapping from geneid to KEGG pathway. \subsection{Extracting mappings from organism packages} If you know that the information mapping gene ID to your categories of interest is contained in the organism packages, but \texttt{goseq} fails to fetch it automatically, you may want to extract it yourself and then pass it to the \texttt{goseq} function using the \texttt{gene2cat} argument. This is done in exactly the same way as extracting the KEGG to ENSEMBL mappings in the section ``KEGG pathway analysis" above. This example is actually the worst case, where it is necessary to combine two mappings to get the desired list. If we had instead wanted the association between Entrez gene IDs and KEGG pathways, the following code would have been sufficient: <>= kegg=as.list(org.Hs.egPATH) head(kegg) @ A note on fetching GO mappings from the organism. The data structure of GO is a directed acyclic graph. This means that in addition to each GO category being associated with a set of genes, it may also have children that are associated to other genes. It is important to use the org.Hs.egGO2ALLEGS and NOT the org.Hs.egGO object to create the mapping between GO categories and gene identifiers, as the latter does not include the links to genes arising from "child" GO categories. Thank you to Christopher Fjell for pointing this out. \subsection{Correcting for other biases} It is possible that in some circumstances you will wish to correct not just for length bias, but for the total number of counts. This can make sense because power to detect DE depends on the total number of counts a gene receives, which is the product of gene length and gene expression. So correcting for read count bias will compensate for all biases, known and unknown, in power to detect DE. On the other hand, it will also remove bias resulting from differences in expression level, which may not be desirable. \np Correcting for count bias will produce a different PWF. Therefore, we need to tell \texttt{goseq} about the data on which the fraction DE depends when calculating the PWF using the \texttt{nullp} function. We then simply pass the result to \texttt{goseq} as usual. \np So, in order to tell \texttt{goseq} to correct for read count bias instead of length bias, all you need to do is supply a numeric vector, containing the number of counts for each gene to \texttt{nullp}. <>= countbias=rowSums(counts)[rowSums(counts)!=0] length(countbias) length(genes) @ To use the count bias when doing GO analysis, simply pass this vector to \texttt{nullp} using the \texttt{bias.data} option. Note that we have to supply "hg19" and "ensGene" to \texttt{goseq} as it is not used by nullp and hence not in the pwf.counts object. <>= pwf.counts=nullp(genes,bias.data=countbias) GO.counts=goseq(pwf.counts,"hg19","ensGene") head(GO.counts) @ Note that if you want to correct for length bias, but your organism/gene identifier is not natively supported, then you need to follow the same procedure as above, only the numeric vector supplied will contain each gene's length instead of its number of reads. \section{Setup} This vignette was built on: <>= sessionInfo() @ \section{Acknowledgments} Christopher Fjell for a series of bug fixes and pointing out the difference between the egGO and egGO2ALLEGS objects in the organism packages. \bibliographystyle{plainnat} \bibliography{goseq_bib} \end{document} goseq/inst/doc/goseq.pdf0000644000175200017520000514322014136072437016263 0ustar00biocbuildbiocbuild%PDF-1.5 % 3 0 obj << /Length 2465 /Filter /FlateDecode >> stream xڕYo6~_k II}(ТICAޕElo~Rvp!wzL 5e_U,\ޜ宇?g_P0jaX7~gҘZnQ*|⮡CoElO+/ۡGlViZyW/Ve(|ݰuqЀ_<|.. 8@7[u3/&h2M&3*AszeG[YԾ|X٪hJD{lwϚ~m<ޡ'_t+&3/4b+JF P{Eu4,6.= 8*AJ \b_ԍ"=[Y^t\VÚM^`Y /3bU_d=(wt3sEnP@H$VփLqHj9ԅR:_\aFQNw,InenS?]AF79::~W+e*6yD2 &.YD4:-?i(^ND ӂi>]߱f^8# $:6Cm3EXym2VizH8琁- mD(t% @܃rc⯦{ Ts|wRB|^Jn8뎓@?̶S,/*W/s*9!ɊѵRJ PYnƣ@ Vr(Qlone+m UϏLлAGe"ip.{jo_GezF[ڐ¼8ZH -RMN"U}-/f$a((6UQTXz&P&-suͻ:E168)7[֚[n2fPa# D[wy*&Ͻ,<5E-j'GnHL'9(C[bqw)Ա0ɏ$VHWxhp yo6xB )%#RXڌvT ;Rm-fm$|Ț'<^r#|d{n`w"P[saW(JH|v/ @ 7ZdhK}BJгreaW@ Os[Nv' =HR? 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As with \code{\link{getlength}} it is preferable that the same gene identifier system is used for both summarization and retrieving GO categories. Valid options for the \code{fetch.cats} arguement are any combination of "GO:CC", "GO:BP", "GO:MF" & "KEGG". The three GO terms refer to the Cellular Component, Biological Process and Molecular Function respectively. "KEGG" refers to KEGG pathways. Note that \code{getgo} is a convenience function, designed to make extracting mappings between GO categories and Gene ID easy. For less common organisms and/or gene ID \code{getgo} may fail to return a mapping even when a legitimate mapping exists in the revelant organism package. If \code{getgo} fails, you should always try to build the mapping yourself from the organism package (if one exists) before deciding that the information is unavailable. Further information and examples of this can be found in the package Vignette. } \value{ A list where each entry is named by a gene and contains a vector of all the associated GO categories. This can be used directly with the \code{gene2cat} option in \code{\link{goseq}}. } \author{ Matthew D. Young \email{myoung@wehi.edu.au} } %\note{ %% ~~further notes~~ %} %% ~Make other sections like Warning with \section{Warning }{....} ~ \seealso{ \code{\link{supportedGenomes}}, \code{\link{supportedGeneIDs}}, \code{\link{goseq}} } \examples{ genes <- c("ENSG00000124208", "ENSG00000182463", "ENSG00000124201", "ENSG00000124205", "ENSG00000124207") getgo(genes,'hg19','ensGene') } % Add one or more standard keywords, see file 'KEYWORDS' in the % R documentation directory. % \keyword{ ~kwd1 } % \keyword{ ~kwd2 }% __ONLY ONE__ keyword per line goseq/man/getlength.Rd0000644000175200017520000000520614136047504015744 0ustar00biocbuildbiocbuild\name{getlength} \Rdversion{1.1} \alias{getlength} %- Also NEED an '\alias' for EACH other topic documented here. \title{ Retrieves Gene length data } \description{ Gets the length of each gene in a vector. } \usage{ getlength(genes, genome, id) } %- maybe also 'usage' for other objects documented here. \arguments{ \item{genes}{ A vector or list of the genes for which length information is required. } \item{genome}{ A string identifying the genome that \code{genes} refer to. For a list of supported organisms run \code{\link{supportedGenomes}}. } \item{id}{ A string identifying the gene identifier used by \code{genes}. For a list of supported gene IDs run \code{\link{supportedGeneIDs}}. } } \details{ Length data is obtained from data obtained from the UCSC genome browser for each combination of \code{genome} and \code{id}. As fetching this data at runtime is time consuming, a local copy of the length information for common genomes and gene ID are included in the \pkg{geneLenDataBase} package. This function uses this package to fetch the required data. The length of a gene is taken to be the median length of all its mature, mRNA, transcripts. It is always preferable to obtain length information directly for the gene ID used to summarize your count data, rather than converting IDs and then using the supplied databases. Even when two genes have a one-to-one mapping between different identifier conventions (which is often not the case), they frequently refer to slightly different regions of the genome with different lengths. It is therefore recommended that the user perform the full analysis in terms of only one gene ID, or manually obtain their own length data for the identifier used to bin reads by gene. } \value{ Returns a vector of the gene lengths, in the same order as \code{genes}. If length data is unavailable for a particular gene NA is returned in that position. The returned vector is intended for use with the \code{bias.data} option of the \code{\link{nullp}} function. } %\references{ %% ~put references to the literature/web site here ~ %} \author{ Matthew D. Young \email{myoung@wehi.edu.au} } %\note{ %% ~~further notes~~ %} %% ~Make other sections like Warning with \section{Warning }{....} ~ \seealso{ \code{\link{supportedGenomes}}, \code{\link{supportedGeneIDs}}, \code{\link{nullp}}, \pkg{geneLenDataBase} } \examples{ genes <- c("ENSG00000124208", "ENSG00000182463", "ENSG00000124201", "ENSG00000124205", "ENSG00000124207") getlength(genes,'hg19','ensGene') } %% Add one or more standard keywords, see file 'KEYWORDS' in the %% R documentation directory. %\keyword{ ~kwd1 } %\keyword{ ~kwd2 }% __ONLY ONE__ keyword per line goseq/man/goseq.Rd0000644000175200017520000001650614136047504015106 0ustar00biocbuildbiocbuild\name{goseq} \Rdversion{1.1} \alias{goseq} %- Also NEED an '\alias' for EACH other topic documented here. \title{ goseq Gene Ontology analyser } \description{ Does selection-unbiased testing for category enrichment amongst differentially expressed (DE) genes for RNA-seq data. By default, tests gene ontology (GO) categories, but any categories may be tested. } \usage{ goseq(pwf, genome, id, gene2cat = NULL, test.cats=c("GO:CC", "GO:BP", "GO:MF"), method = "Wallenius", repcnt = 2000, use_genes_without_cat=FALSE) } %- maybe also 'usage' for other objects documented here. \arguments{ \item{pwf}{ An object containing gene names, DE calls, the probability weighting function. Usually generated by \code{\link{nullp}}. } \item{genome}{ A string identifying the genome that \code{genes} refer to. For a list of supported organisms run \code{\link{supportedGenomes}}. } \item{id}{ A string identifying the gene identifier used by \code{genes}. For a list of supported gene IDs run \code{\link{supportedGeneIDs}}. } \item{gene2cat}{ A data frame with two columns containing the mapping between genes and the categories of interest. Alternatively, a list where the names are genes and each entry is a vector containing GO categories associated with that gene (this is the output produced by \code{\link{getgo}}). If set to \code{NULL} goseq attempts to fetch GO categories automatically using \code{\link{getgo}}. } \item{test.cats}{ A vector specifying which categories to test for over representation amongst DE genes. See details for allowed options. } \item{method}{ The method to use to calculate the unbiased category enrichment scores. Valid options are "Wallenius", "Sampling" & "Hypergeometric". "Hypergeometric" and "Sampling" should almost never be used (see details). } \item{repcnt}{ Number of random samples to be calculated when random sampling is used. Ignored unless \code{method="Sampling"}. } \item{use_genes_without_cat}{ A boolean to indicate whether genes without a categorie should still be used. For example, a large number of gene may have no GO term annotated. If this option is set to FALSE, those genes will be ignored in the calculation of p-values (default behaviour). If this option is set to TRUE, then these genes will count towards the total number of genes outside the category being tested (default behaviour prior to version 1.15.2). } } \details{ The \code{pwf} argument is almost always the output of the function \code{\link{nullp}}. This is a data frame with 3 columns, named "DEgenes", "bias.data" and "pwf" with the rownames set to the gene names. Each row corresponds to a gene with the DEgenes column specifying if the gene is DE (1 for DE, 0 for not DE), the bias.data column giving the numeric value of the DE bias being accounted for (usually the gene length or number of counts) and the pwf column giving the genes value on the probability weighting function. \code{goseq} obtains length data from UCSC and GO mappings from the organim packages (see \code{link{getgo}} and \code{\link{getlength}} for details). If your data is in an unsupported format you will need to obtain the GO category mapping and supply them to the \code{goseq} function using the \code{gene2cat} arguement. To use your own gene to category mapping with \code{goseq}, use the \code{gene2cat} arguement. This arguement takes a data.frame, with one column containing gene IDs and the other containing the associated categories. As the mapping from gene <-> category is in general many to many there will be multiple rows containing the same gene identifier. Alternatively, \code{gene2cat} can take a list, where the names are the genes and the entries are the GO categories associated with the genes. This is the format produced by the \code{\link{getgo}} function and is more space efficient than the data.frame representation. If \code{gene2cat} is left as \code{NULL}, \code{goseq} attempts to use \code{\link{getgo}} to fetch GO catgeory to gene identifier mappings. The PWF is usually calculated using the \code{\link{nullp}} function to correct for length bias. However, \code{goseq} will work with any vector of weights. Any bias can be accounted for so long as a weight for each gene is supplied using this arguement. \code{NA}s are allowed in the "pwf" and "bias.data" columns of the PWF data frame (these usually occur as a result of missing length data for some genes). Any entry which is \code{NA} is set to the weighting of the median gene. Valid options for the \code{test.cats} arguement are any combination of "GO:CC", "GO:BP", "GO:MF" & "KEGG". The three GO terms refer to the Cellular Component, Biological Process and Molecular Function respectively. "KEGG" refers to KEGG pathways. The three methods, "Wallenius", "Sampling" & "Hypergeometric", calculate the p-values as follows. "Wallenius" approximates the true distribution of numbers of members of a category amongst DE genes by the Wallenius non-central hypergeometric distribution. This distribution assumes that within a category all genes have the same probability of being chosen. Therefore, this approximation works best when the range in probabilities obtained by the probability weighting function is small. "Wallenius" is the recommended method for calculating p-values. "Sampling" uses random sampling to approximate the true distribution and uses it to calculate the p-values for over (and under) representation of categories. In practice, its use quickly becomes computationally prohibitive because \code{repcnt} would need to be set very high for most applications. CAUTION: "Hypergeometric" should NEVER be used for producing results for biological interpretation. If there is genuinly no bias in power to detect DE in your experiment, the PWF will reflect this and the other methods will produce accuracte results. "Hypergeometric" assumes there is no bias in power to detect differential expression at all and calculates the p-values using a standard hypergeometric distribution. Useful if you wish to test the effect of selection bias on your results. } \value{ goseq returns a data frame with several columns. The first column gives the name of the category, the second gives the p-value for the associated category being over represented amongst DE genes. The third column gives the p-value for the associated category being under represented amongst DE genes. The p-values have not been corrected for multiple hypothesis testing. The fourth and fifth columns give the number of differentially expressed genes in the category and total genes in the category respectively. If any of the categories was a GO term, there will be two additional columns for the GO term and its ontology. } \references{ Young, M. D., Wakefield, M. J., Smyth, G. K., Oshlack, A. (2010) \emph{Gene ontology analysis for RNA-seq: accounting for selection bias} Genome Biology Date: Feb 2010 Vol: 11 Issue: 2 Pages: R14 } \author{ Matthew D. Young \email{myoung@wehi.edu.au} } %\note{ %% ~~further notes~~ %} %% ~Make other sections like Warning with \section{Warning }{....} ~ \seealso{ \code{\link{nullp}}, \code{\link{getgo}}, \code{\link{getlength}} } \examples{ data(genes) pwf <- nullp(genes,'hg19','ensGene') pvals <- goseq(pwf,'hg19','ensGene') head(pvals) } % Add one or more standard keywords, see file 'KEYWORDS' in the % R documentation directory. %\keyword{ ~kwd1 } %\keyword{ ~kwd2 }% __ONLY ONE__ keyword per line goseq/man/makespline.Rd0000644000175200017520000000370714136047504016117 0ustar00biocbuildbiocbuild\name{makespline} \Rdversion{1.1} \alias{makespline} %- Also NEED an '\alias' for EACH other topic documented here. \title{ Monotonic Spline } \description{ Fits a monotonic cubic spline to the data provided, using the penalized constrained least squares method from the \code{mgcv} package. } \usage{ makespline(x, y, newX=NULL, nKnots = 6, lower_bound = 10^-3) } %- maybe also 'usage' for other objects documented here. \arguments{ \item{x}{ The predictor variable. } \item{y}{ The response variable. Must be the same length as \code{x}. } \item{newX}{ The points at which to return the value on the fitted spline. If not specified \code{x} is used. } \item{nKnots}{ The number of knots to use in fitting the spline. } \item{lower_bound}{ The spline cannot drop below this value. } } \details{ This uses the \code{pcls} function from the \pkg{mgcv} package to produce the fit. The montonicity constraint is enforced using \code{mono.con} from the same package. The \code{lower_bound} argument is only used on the rare occasions when the fitting function becomes negative or arbitrarily close to zero. If this does occur \code{lower_bound} is added everywhere to ensure that no one length is given essentially infinite weighting. } \value{ Returns a vector of values containing the value of the fit at each point \code{newX}. } \references{ Package \pkg{mgcv}. In particular this function is a modification of an example given in the man page for \code{pcls}. } \author{ Matthew D. Young \email{myoung@wehi.edu.au}. } %\note{ %% ~~further notes~~ %} %% ~Make other sections like Warning with \section{Warning }{....} ~ %\seealso{ %% ~~objects to See Also as \code{\link{help}}, ~~~ %} \examples{ y <- c( rbinom(50,p=0.4,size=1), rbinom(50,p=0.6,size=1) ) x <- 1:100 plot(x,y) p <- makespline(x,y) lines(x,p) } %% Add one or more standard keywords, see file 'KEYWORDS' in the %% R documentation directory. %\keyword{ ~kwd1 } %\keyword{ ~kwd2 }% __ONLY ONE__ keyword per line goseq/man/nullp.Rd0000644000175200017520000000657314136047504015125 0ustar00biocbuildbiocbuild\name{nullp} \Rdversion{1.1} \alias{nullp} %- Also NEED an '\alias' for EACH other topic documented here. \title{ Probability Weighting Function } \description{ Calculates a Probability Weighting Function for a set of genes based on a given set of biased data (usually gene length) and each genes status as differentially expressed or not. } \usage{ nullp(DEgenes, genome, id, bias.data=NULL,plot.fit=TRUE) } %- maybe also 'usage' for other objects documented here. \arguments{ \item{DEgenes}{ A named binary vector where 1 represents DE, 0 not DE and the names are gene IDs. } \item{genome}{ A string identifying the genome that \code{genes} refer to. For a list of supported organisms run \code{\link{supportedGenomes}}. } \item{id}{ A string identifying the gene identifier used by \code{genes}. For a list of supported gene IDs run \code{\link{supportedGeneIDs}}. } \item{bias.data}{ A numeric vector containing the data on which the DE may depend. Usually this is the median transcript length of each gene in bp. If set to \code{NULL} \code{nullp} will attempt to fetch length using \code{\link{getlength}}. } \item{plot.fit}{ Plot the PWF or not? Calls \code{\link{plotPWF}} with default values if \code{TRUE}. } } \details{ It is essential that the entire analysis pipeline, from summarizing raw reads through to using \code{goseq} be done in just one gene identifier format. If your data is in a different format you will need to obtain the gene lengths and supply them to the \code{nullp} function using the \code{bias.data} arguement. Converting to a supported format from another format should be avoided whenever possible as this will almost always result in data loss. \code{NA}s are allowed in the bias.data vector if you do not have information about a certain gene. Setting a gene to \code{NA} is preferable to removing it from the analysis. If \code{bias.data} is left as \code{NULL}, \code{nullp} attempts to use \code{\link{getlength}} to fetch GO catgeory to gene identifier mappings. It is recommended you review the fit produced by the \code{nullp} function before proceeding by leaving \code{plot.fit} as \code{TRUE}. } \value{ A data frame with 3 columns, named "DEgenes", "bias.data" and "pwf" with the rownames set to the gene names. Each row corresponds to a gene with the DEgenes column specifying if the gene is DE (1 for DE, 0 for not DE), the bias.data column giving the numeric value of the DE bias being accounted for (usually the gene length or number of counts) and the pwf column giving the genes value on the probability weighting function. This object is usually passed to \code{goseq} to calculate enriched categories or \code{plotPWF} for further plotting. } \references{ Young, M. D., Wakefield, M. J., Smyth, G. K., Oshlack, A. (2010) \emph{Gene ontology analysis for RNA-seq: accounting for selection bias} Genome Biology Date: Feb 2010 Vol: 11 Issue: 2 Pages: R14 } \author{ Matthew D. Young \email{myoung@wehi.edu.au} } %\note{ %% ~~further notes~~ %} %% ~Make other sections like Warning with \section{Warning }{....} ~ \seealso{ \code{\link{supportedGenomes}}, \code{\link{supportedGeneIDs}}, \code{\link{goseq}}, \code{\link{getlength}} } \examples{ data(genes) pwf <- nullp(genes, 'hg19', 'ensGene') } % Add one or more standard keywords, see file 'KEYWORDS' in the % R documentation directory. %\keyword{ ~kwd1 } %\keyword{ ~kwd2 }% __ONLY ONE__ keyword per line goseq/man/plotPWF.Rd0000644000175200017520000000531014136047504015312 0ustar00biocbuildbiocbuild\name{plotPWF} \Rdversion{1.1} \alias{plotPWF} %- Also NEED an '\alias' for EACH other topic documented here. \title{ Plot the Probability Weighting Function } \description{ Plots the Probability Weighting Function created by \code{\link{nullp}} by binning together genes. } \usage{ plotPWF(pwf,binsize="auto",pwf_col=3,pwf_lwd=2,xlab="Biased Data in gene bins.",ylab="Proportion DE",...) } %- maybe also 'usage' for other objects documented here. \arguments{ \item{pwf}{ A data frame with 3 columns named DEgenes, bias.data & pwf and row names giving the gene names. Usually generated by \code{\link{nullp}}. } \item{binsize}{ Calculate and plot the fraction of genes that are DE in bins of this size. If set to "auto" the best binsize for visualization is attempted to be found automatically. } \item{pwf_col}{ The colour of the probability weighting function } \item{pwf_lwd}{ The width of the probability weighting function } \item{xlab}{ The x-axis label. \code{} is replaced by the binsize used. } \item{ylab}{ The y-axis label. } \item{...}{ Extra arguments that are passed to plot. } } \details{ This function is almost always called using the output from the \code{\link{nullp}} function. However, it can be used to visualize the length (or any other type of quantifyable) bias in ability to detect DE in a data set. The \code{pwf} arguement needs to be a data frame with 3 columns each containing numeric entries (although NAs are permitted in the bias.data and pwf columns), which must be named "DEgenes", "bias.data" and "pwf", although they can appear in any order. The row names are taken to be the gene names. The DEgenes column should be 0s or 1s where 1 represents a DE gene, 0 a gene which is not DE. The bias.data column is a quantification of the quantity for which there is a bias in detecting DE for the associated gene, this is usually gene length or the number of counts associated with a gene. Finally, the pwf column gives the probability weighting to be applied for a given gene. } \value{ Nothing is returned. } \references{ Young, M. D., Wakefield, M. J., Smyth, G. K., Oshlack, A. (2010) \emph{Gene ontology analysis for RNA-seq: accounting for selection bias} Genome Biology Date: Feb 2010 Vol: 11 Issue: 2 Pages: R14 } \author{ Matthew D. Young \email{myoung@wehi.edu.au} } %\note{ %% ~~further notes~~ %} %% ~Make other sections like Warning with \section{Warning }{....} ~ \seealso{ \code{\link{nullp}} } \examples{ data(genes) pwf <- nullp(genes, 'hg19', 'ensGene',plot.fit=FALSE) plotPWF(pwf,binsize=200) } % Add one or more standard keywords, see file 'KEYWORDS' in the % R documentation directory. %\keyword{ ~kwd1 } %\keyword{ ~kwd2 }% __ONLY ONE__ keyword per line goseq/man/supportedOrganisms.Rd0000644000175200017520000000314514136047504017673 0ustar00biocbuildbiocbuild\name{supportedOrganisms} \Rdversion{1.1} \alias{supportedOrganisms} %- Also NEED an '\alias' for EACH other topic documented here. \title{ Supported Organisms } \description{ Lists which genomes and gene ids are supported for gene lengths and for gene ontology } \usage{ supportedOrganisms() } \details{ Goseq allows a user to provide their own bias data (usually gene lengths) and/or gene categories (usually gene ontologies), but goseq also provides this data automatically for many commonly used species. This function lists which genome and gene ids are automatically supported by goseq. The first to third columns list the genomes, gene ids, and gene id descriptions respectively. The fourth column indicates whether this combination of genome and id are available in the geneLengthDataBase. If a particular combination is absent, goseq may still automatically fetch the gene lengths from either a TxDB annotation package (must be installed) or download the data from UCSC. For example gene lengths for \code{hg38} are not supported in \code{geneLengthDataBase} but may be fetched by these other means. However, this is not always the case. The final column indicates if GO terms will be automatically fetched for the geneome and id combination. Goseq relies on an \code{org} annotation package (e.g. \code{org.Hs.eg.db}) existing for the organism. In general, if either the lengths or GO terms are not supported, the user must enter this information manually. } \value{ A data.frame containing supported genomes and gene ids } \author{ Nadia Davidson \email{nadia.davidson@mcri.edu.au} } \examples{ supportedOrganisms() } goseq/vignettes/0000755000175200017520000000000014136072441014724 5ustar00biocbuildbiocbuildgoseq/vignettes/goseq.Rnw0000644000175200017520000007126614136047504016550 0ustar00biocbuildbiocbuild\documentclass[12pt]{article} \topmargin 0in \headheight 0in \headsep 0in \oddsidemargin 0in \evensidemargin 0in \textwidth 176mm \textheight 215mm \usepackage{natbib} \usepackage{Sweave} \usepackage{url} \DeclareGraphicsRule{.tif}{png}{.png}{`convert #1 `dirname #1`/`basename #1 .tif`.png} \newcommand{\np}{\vspace{5mm}\par} \begin{document} %\VignetteIndexEntry{goseq User's Guide} \title{\texttt{goseq}: Gene Ontology testing for RNA-seq datasets} \author{Matthew D. Young \and Nadia Davidson % \\ \texttt{nadia.davidson@mcri.edu.au} \and Matthew J. Wakefield \and Gordon K.\ Smyth \and Alicia Oshlack} % Please increment date when working on this document, so that % date shows genuine change date, not merely date of compile. \date{8 September 2017} \maketitle \section{Introduction} This document gives an introduction to the use of the \texttt{goseq} R Bioconductor package~\citep{Young10}. This package provides methods for performing Gene Ontology analysis of RNA-seq data, taking length bias into account~\citep{Oshlack:2009p149}. The methods and software used by \texttt{goseq} are equally applicable to other category based test of RNA-seq data, such as KEGG pathway analysis. \np Once installed, the \texttt{goseq} package can be easily loaded into R using: <>= library(goseq) @ <>= options(width=84) @ \np In order to perform a GO analysis of your RNA-seq data, \texttt{goseq} only requires a simple named vector, which contains two pieces of information. \begin{enumerate} \item \texttt{Measured genes}: all genes for which RNA-seq data was gathered for your experiment. Each element of your vector should be named by a unique gene identifier. \item \texttt{Differentially expressed genes}: each element of your vector should be either a 1 or a 0, where 1 indicates that the gene is differentially expressed and 0 that it is not. \end{enumerate} If the organism, gene identifier or category test is currently not natively supported by \texttt{goseq}, it will also be necessary to supply additional information regarding the genes length and/or the association between categories and genes. \np Bioconductor R packages such as \texttt{Rsubread} allow for the summarization of mapped reads into a table of counts, such as reads per gene. From there, several packages exist for performing differential expression analysis on summarized data (eg. \texttt{edgeR}~\citep{Robinson07, Robinson08, Robinson10}). \texttt{goseq} will work with any method for determining differential expression and as such differential expression analysis is outside the scope of this document, but in order to facilitate ease of use, we will make use of the \texttt{edgeR} package to calculate differentially expressed (DE) genes in all the case studies in this document. \section{Reading data} We assume that the user can use appropriate in-built \texttt{R} functions (such as \texttt{read.table} or \texttt{scan}) to obtain two vectors, one containing all genes assayed in the RNA-seq experiment, the other containing all genes which are DE. If we assume that the vector of genes being assayed is named \texttt{assayed.genes} and the vector of DE genes is named \texttt{de.genes} we can construct a named vector suitable for use with \texttt{goseq} using the following: <>= gene.vector=as.integer(assayed.genes%in%de.genes) names(gene.vector)=assayed.genes head(gene.vector) @ It may be that the user can already read in a vector in this format, in which case it can then be immediately used by \texttt{goseq}. \section{GO testing of RNA-seq data} To begin the analysis, \texttt{goseq} first needs to quantify the length bias present in the dataset under consideration. This is done by calculating a Probability Weighting Function or PWF which can be thought of as a function which gives the probability that a gene will be differentially expressed (DE), based on its length alone. The PWF is calculated by fitting a monotonic spline to the binary data series of differential expression (1=DE, 0=Not DE) as a function of gene length. The PWF is used to weight the chance of selecting each gene when forming a null distribution for GO category membership. The fact that the PWF is calculated directly from the dataset under consideration makes this approach robust, only correcting for the length bias present in the data. For example, if \texttt{goseq} is run on a microarray dataset, for which no length bias exists, the calculated PWF will be nearly flat and all genes will be weighted equally, resulting in no length bias correction. \np In order to account for the length bias inherent to RNA-seq data when performing a GO analysis (or other category based tests), one cannot simply use the hypergeometric distribution as the null distribution for category membership, which is appropriate for data without DE length bias, such as microarray data. GO analysis of RNA-seq data requires the use of random sampling in order to generate a suitable null distribution for GO category membership and calculate each categories significance for over representation amongst DE genes. \np However, this random sampling is computationally expensive. In most cases, the Wallenius distribution can be used to approximate the true null distribution, without any significant loss in accuracy. The \texttt{goseq} package implements this approximation as its default option. The option to generate the null distribution using random sampling is also included as an option, but users should be aware that the default number of samples generated will not be enough to accurately call enrichment when there are a large number of go terms. \np Having established a null distribution, each GO category is then tested for over and under representation amongst the set of differentially expressed genes and the null is used to calculate a p-value for under and over representation. \section{Natively supported Gene Identifiers and category tests} \texttt{goseq} needs to know the length of each gene, as well as what GO categories (or other categories of interest) each gene is associated with. \texttt{goseq} relies on the UCSC genome browser to provide the length information for each gene. However, because the process of fetching the length of every transcript is slow and bandwidth intensive, \texttt{goseq} relies on an offline copy of this information stored in the data package \texttt{geneLenDataBase}. To see which genome/gene identifier combinations are in the local database, simply run: <>= supportedOrganisms() @ The leftmost columns in the output of this command list the genomes and gene identifiers respectively. If length data exists in the local database it is indicated in the second last column. If your genome/ID combination is not in the local database, it may be downloaded from the UCSC genome browser or taken from a \texttt{TxDb} annotation package (if installed). If your genome/ID combination is not found in any database, you will have to manually specify the gene lengths. We encourage all users to manually specify their gene lengths if provided by upstream summarization programs. e.g. \texttt{featureCounts}, as these lengths will be more accurate. \np In order to link GO categories to genes, \texttt{goseq} uses the organism packages from Bioconductor. These packages are named org...db, where is a short string identifying the genome and is a short string identifying the gene identifier. Currently, \texttt{goseq} will automatically retrieve the mapping between GO categories and genes from the relevant package (as long as it is installed) for commonly used genome/ID combinations. If GO mappings are not automatically available for your genome/ID combination, you will have to manually specify the relationship between genes and categories. Although the Genome/ID naming conventions used by the organism packages differ from the UCSC, \texttt{goseq} is able to convert between the two, so the user need only ever specify the UCSC genome/ID in most cases. The final column indicates whether the Genome/ID combination is supported for GO categories. \section{Non-native Gene Identifier or category test} If the organism, Gene Identifier or category test you wish to perform is not in the native \texttt{goseq} database, you will have to supply one or all of the following: \begin{itemize} \item \texttt{Length data}: the length of each gene in your gene identifier format. \item \texttt{Category mappings}: the mapping (usually many-to-many) between the categories you wish to test for over/under representation amongst DE genes and genes in your gene identifier format. \end{itemize} \subsection{Length data format} The length data must be formatted as a numeric vector, of the same length as the main named vector specifying gene names/DE genes. Each entry should give the length of the corresponding gene in bp. If length data is unavailable for some genes, that entry should be set to NA. \subsection{Category mapping format} The mapping between category names and genes should be given as a data frame with two columns. One column should contain the gene IDs and the other the name of an associated category. As the mapping between categories and genes is usually many-to-many, this data frame will usually have multiple rows with the same gene name and category name. \np Alternatively, mappings between genes and categories can be given as a list. The names of list entries should be gene IDs and the entries themselves should be a vector of category names to which the gene ID corresponds. \subsection{Some additional tips} Any organism for which there is an annotation on either Ensembl or the UCSC, can be easily turned into length data using the GenomicFeatures package. To do this, first create a TranscriptDb object using either makeTxDbFromBiomart or makeTxDbFromUCSC (see the help in the GenomicFeatures package on using these commands). Once you have a transcriptDb object, you can get a vector named by gene ID containing the median transcript length of each gene simply by using the command. <>= txsByGene=transcriptsBy(txdb,"gene") lengthData=median(width(txsByGene)) @ The relationship between gene identifier and GO category can usually be obtained from the Gene Ontology website (www.geneontology.org) or from the NCBI. Additionally, the bioconductor AnnotationDbi library has recently added a function "makeOrgPackageFromNCBI", which can be used to create an organism package from within R, using the NCBI data. Once created, this package can then be used to obtain the mapping between genes and gene ontology. \section{Case study: Prostate cancer data} \subsection{Introduction} This section provides an analysis of data from an RNA-seq experiment to illustrate the use of \texttt{goseq} for GO analysis. \np This experiment examined the effects of androgen stimulation on a human prostate cancer cell line, LNCaP. The data set includes more than 17 million short cDNA reads obtained for both the treated and untreated cell line and sequenced on Illumina's 1G genome analyzer. \np For each sample we were provided with the raw 35 bp RNA-seq reads from the authors. For the untreated prostate cancer cells (LNCaP cell line) there were 4 lanes totaling ~10 million, 35 bp reads. For the treated cells there were 3 lanes totaling ~7 million, 35 bp reads. All replicates were technical replicates. Reads were mapped to NCBI version 36.3 of the human genome using bowtie. Any read with which mapped to multiple locations was discarded. Using the ENSEMBL 54 annotation from biomart, each mapped read was associated with an ENSEMBL gene. This was done by associating any read that overlapped with any part of the gene (not just the exons) with that gene. Reads that did not correspond to genes were discarded. \subsection{Source of the data} The data set used in this case study is taken from \citep{Li:2008p113} and was made available from the authors upon request. \subsection{Determining the DE genes using \texttt{edgeR}} To begin with, we load in the text data and convert it the appropriate \texttt{edgeR} DGEList object. <>= library(edgeR) table.summary=read.table(system.file("extdata","Li_sum.txt",package='goseq'), sep='\t',header=TRUE,stringsAsFactors=FALSE) counts=table.summary[,-1] rownames(counts)=table.summary[,1] grp=factor(rep(c("Control","Treated"),times=c(4,3))) summarized=DGEList(counts,lib.size=colSums(counts),group=grp) @ Next, we use \texttt{edgeR} to estimate the biological dispersion and calculate differential expression using a negative binomial model. <>= disp=estimateCommonDisp(summarized) disp$common.dispersion tested=exactTest(disp) topTags(tested) @ Finally, we Format the DE genes into a vector suitable for use with \texttt{goseq} <>= genes=as.integer(p.adjust(tested$table$PValue[tested$table$logFC!=0], method="BH")<.05) names(genes)=row.names(tested$table[tested$table$logFC!=0,]) table(genes) @ \subsection{Determining Genome and Gene ID} In order to allow for automatic data retrieval, the user has to tell \texttt{goseq} what genome and gene ID format were used to summarize the data. In our case we will use the hg19 build of the human genome, we check what code this corresponds to by running: <>= head(supportedOrganisms()) @ Which lists the genome codes in the far left column, headed ``Genome''. As we are using ``hg19" and we also know that we used ENSEMBL Gene ID to summarize our read data, we check what code this corresponds to by running: <>= supportedOrganisms()[supportedOrganisms()$Genome=="hg19",] @ The gene ID codes are listed in the column second from left, titled ``Id''. We find that our gene ID code is ``ensGene". We will use these strings whenever we are asked for a genome or id. If the gene ID is missing for your Genome (for example this is the case for hg38), then the genome is not supported in the geneLengthDatabase. Gene lengths will either be automatically fetched from TxDB, UCSC or you will need to provide them manually. Supported Gene IDs to automatically fetch GO terms should usually either be Entrez (``knownGene"), Ensembl (``ensGene") or gene symbols (``geneSymbol"). \subsection{GO analysis} \subsubsection{Fitting the Probability Weighting Function (PWF)} We first need to obtain a weighting for each gene, depending on its length, given by the PWF. As you may have noticed when running supportedGenomes or supportedGeneIDs, length data is available in the local database for our gene ID, ``ensGene" and our genome, ``hg19". We will let \texttt{goseq} automatically fetch this data from its databases. <>= pwf=nullp(genes,"hg19","ensGene") head(pwf) @ \texttt{nullp} plots the resulting fit, allowing verification of the goodness of fit before continuing the analysis. Further plotting of the pwf can be performed using the \texttt{plotPWF} function. \np The output of nullp contains all the data used to create the PWF, as well as the PWF itself. It is a data frame with 3 columns, named "DEgenes", "bias.data" and "pwf" with the rownames set to the gene names. Each row corresponds to a gene with the DEgenes column specifying if the gene is DE (1 for DE, 0 for not DE), the bias.data column giving the numeric value of the DE bias being accounted for (usually the gene length or number of counts) and the pwf column giving the genes value on the probability weighting function. \subsubsection{Using the Wallenius approximation} To start with we will use the default method, to calculate the over and under expressed GO categories among DE genes. Again, we allow \texttt{goseq} to fetch data automatically, except this time the data being fetched is the relationship between ENSEMBL gene IDs and GO categories. %This hides the output of goseq <>= GO.wall=goseq(pwf,"hg19","ensGene") head(GO.wall) @ The resulting object is ordered by GO category over representation amongst DE genes. \subsubsection{Using random sampling} It may sometimes be desirable to use random sampling to generate the null distribution for category membership. For example, to check consistency against results from the Wallenius approximation. This is easily accomplished by using the \texttt{method} option to specify sampling and the \texttt{repcnt} option to specify the number of samples to generate: <>= GO.samp=goseq(pwf,"hg19","ensGene",method="Sampling",repcnt=1000) @ <>= head(GO.samp) @ You will notice that this takes far longer than the Wallenius approximation. Plotting the p-values against one another, we see that there is little difference between the two methods. However, the accuracy of the sampling method is limited by the number of samples generated, \texttt{repcnt}, such that very low p-values will not be correctly calculated. Significantly enriched GO terms may then be missed after correcting for multiple testing. <>= plot(log10(GO.wall[,2]), log10(GO.samp[match(GO.wall[,1],GO.samp[,1]),2]), xlab="log10(Wallenius p-values)",ylab="log10(Sampling p-values)", xlim=c(-3,0)) abline(0,1,col=3,lty=2) @ \subsubsection{Ignoring length bias} \texttt{goseq} also allows for one to perform a GO analysis without correcting for RNA-seq length bias. In practice, this is only useful for assessing the effect of length bias on your results. You should NEVER use this option as your final analysis. If length bias is truly not present in your data, \texttt{goseq} will produce a nearly flat PWF and no length bias correction will be applied to your data and all methods will produce the same results. \np However, if you still wish to ignore length bias in calculating GO category enrichment, this is again accomplished using the \texttt{method} option. <>= GO.nobias=goseq(pwf,"hg19","ensGene",method="Hypergeometric") head(GO.nobias) @ Ignoring length bias gives very different results from a length bias corrected analysis. <>= plot(log10(GO.wall[,2]), log10(GO.nobias[match(GO.wall[,1],GO.nobias[,1]),2]), xlab="log10(Wallenius p-values)", ylab="log10(Hypergeometric p-values)", xlim=c(-3,0), ylim=c(-3,0)) abline(0,1,col=3,lty=2) @ \subsubsection{Limiting GO categories and other category based tests} By default, \texttt{goseq} tests all three major Gene Ontology branches; Cellular Components, Biological Processes and Molecular Functions. However, it is possible to limit testing to any combination of the major branches by using the \texttt{test.cats} argument to the goseq function. This is done by specifying a vector consisting of some combination of the strings ``GO:CC", ``GO:BP" and ``GO:MF". For example, to test for only Molecular Function GO categories: <>= GO.MF=goseq(pwf,"hg19","ensGene",test.cats=c("GO:MF")) head(GO.MF) @ Native support for other category tests, such as KEGG pathway analysis are also made available via this argument. See the man \texttt{goseq} function man page for up to date information on what category tests are natively supported. \subsubsection{Making sense of the results} Having performed the GO analysis, you may now wish to interpret the results. If you wish to identify categories significantly enriched/unenriched below some p-value cutoff, it is necessary to first apply some kind of multiple hypothesis testing correction. For example, GO categories over enriched using a .05 FDR cutoff \citep{Benjamini95} are: <>= enriched.GO=GO.wall$category[p.adjust(GO.wall$over_represented_pvalue, method="BH")<.05] head(enriched.GO) @ Unless you are a machine, GO accession identifiers are probably not very meaningful to you. Information about each term can be obtained from the Gene Ontology website, \url{http://www.geneontology.org/}, or using the R package \texttt{GO.db}. <>= library(GO.db) for(go in enriched.GO[1:10]){ print(GOTERM[[go]]) cat("--------------------------------------\n") } @ \subsubsection{Understanding goseq internals} The situation may arise where it is necessary for the user to perform some of the data processing steps usually performed automatically by \texttt{goseq} themselves. With this in mind, it will be useful to step through the preprocessing steps performed automatically by \texttt{goseq} to understand what is happening. \np To start with, when \texttt{nullp} is called, \texttt{goseq} uses the genome and gene identifiers supplied to try and retrieve length information for all genes given to the \texttt{genes} argument. To do this, it retrieves the data from the database of gene lengths maintained in the package \texttt{geneLenDataBase}. This is performed by the \texttt{getlength} function in the following way: <>= len=getlength(names(genes),"hg19","ensGene") length(len) length(genes) head(len) @ After some data cleanup, the length data and the DE data is then passed to the \texttt{makespline} function to produce the PWF. The \texttt{nullp} returns a data frame which has 3 columns, the original DEgenes vector, the length bias data (in a column called bias.data) and the PWF itself (in a column named pwf). The names of the genes are also kept in this data frame as the names of the rows. If length data could not be obtained for a certain gene the corresponding entries in the "bias.data" and "pwf" columns are set to NA. \np Next we call the \texttt{goseq} function to determine over/under representation of GO categories amongst DE genes. When we do this, \texttt{goseq} looks for the appropriate organism package and tries to obtain the mapping from genes to GO categories from it. This is done using the \texttt{getgo} function as follows: <>= go=getgo(names(genes),"hg19","ensGene") length(go) length(genes) head(go) @ Note that some of the gene categories have been returned as "NULL". This means that a GO category could not be found in the database for one of the genes. In the \texttt{goseq} command, enrichment will only be calculated using genes with a GO category by default. However, in older versions of goseq (below 1.15.2), we counted all genes. i.e. genes with no categories still counted towards the total number of gene outside of any single category. It is possible to switch between these two behaviors using the \texttt{use\_genes\_without\_cat} flag in \texttt{goseq}. The first thing the getgo function does is to convert the UCSC genome/ID namings into the naming convention used by the organism packages. This is done using two hard coded conversion vectors that are included in the \texttt{goseq} package but usually hiden from the user. <>= goseq:::.ID_MAP goseq:::.ORG_PACKAGES @ It is just as valid to run the length and GO category fetching as separate steps and then pass the result to the \texttt{nullp} and \texttt{goseq} functions using the \texttt{bias.data} and \texttt{gene2cat} arguments. Thus the following two blocks of code are equivalent: <>= pwf=nullp(genes,"hg19","ensGene") go=goseq(pwf,"hg19","ensGene") @ and <>= gene_lengths=getlength(names(genes),"hg19","ensGene") pwf=nullp(genes,bias.data=gene_lengths) go_map=getgo(names(genes),"hg19","ensGene") go=goseq(pwf,"hg19","ensGene",gene2cat=go_map) @ \subsection{KEGG pathway analysis} In order to illustrate performing a category test not present in the \texttt{goseq} database, we perform a KEGG pathway analysis. For human, the mapping from KEGG pathways to genes are stored in the package org.Hs.eg.db, in the object org.Hs.egPATH. In order to test for KEGG pathway over representation amongst DE genes, we need to extract this information and put it in a format that \texttt{goseq} understands. Unfortunately, the org.Hs.eg.db package does not contain direct mappings between ENSEMBL gene ID and KEGG pathway. Therefore, we have to construct this map by combining the ENSEMBL <-> Entrez and Entrez <-> KEGG mappings. This can be done using the following code: <>= # Get the mapping from ENSEMBL 2 Entrez en2eg=as.list(org.Hs.egENSEMBL2EG) # Get the mapping from Entrez 2 KEGG eg2kegg=as.list(org.Hs.egPATH) # Define a function which gets all unique KEGG IDs # associated with a set of Entrez IDs grepKEGG=function(id,mapkeys){unique(unlist(mapkeys[id],use.names=FALSE))} # Apply this function to every entry in the mapping from # ENSEMBL 2 Entrez to combine the two maps kegg=lapply(en2eg,grepKEGG,eg2kegg) head(kegg) @ Note that this step is quite time consuming. The code written here is not the most efficient way of producing this result, but the logic is much clearer than faster algorithms. The source code for getgo contains a more efficient routine. \np We produce the PWF as before. Then, to perform a KEGG analysis, we simply make use of the \texttt{gene2cat} option in \texttt{goseq}: <>= pwf=nullp(genes,"hg19","ensGene") KEGG=goseq(pwf,gene2cat=kegg) head(KEGG) @ Note that we do not have to tell the goseq function what organism and gene ID we are using as we are manually supplying the mapping between genes and categories. \np KEGG analysis is shown as an illustration of how to supply your own mapping between gene ID and category, KEGG analysis is actually natively supported by GOseq and we could have performed it with the following code. <>= pwf=nullp(genes,'hg19','ensGene') KEGG=goseq(pwf,'hg19','ensGene',test.cats="KEGG") head(KEGG) @ Noting that this time it was necessary to tell the goseq function that we are using HG19 and ENSEMBL gene ID, as the function needs this information to automatically construct the mapping from geneid to KEGG pathway. \subsection{Extracting mappings from organism packages} If you know that the information mapping gene ID to your categories of interest is contained in the organism packages, but \texttt{goseq} fails to fetch it automatically, you may want to extract it yourself and then pass it to the \texttt{goseq} function using the \texttt{gene2cat} argument. This is done in exactly the same way as extracting the KEGG to ENSEMBL mappings in the section ``KEGG pathway analysis" above. This example is actually the worst case, where it is necessary to combine two mappings to get the desired list. If we had instead wanted the association between Entrez gene IDs and KEGG pathways, the following code would have been sufficient: <>= kegg=as.list(org.Hs.egPATH) head(kegg) @ A note on fetching GO mappings from the organism. The data structure of GO is a directed acyclic graph. This means that in addition to each GO category being associated with a set of genes, it may also have children that are associated to other genes. It is important to use the org.Hs.egGO2ALLEGS and NOT the org.Hs.egGO object to create the mapping between GO categories and gene identifiers, as the latter does not include the links to genes arising from "child" GO categories. Thank you to Christopher Fjell for pointing this out. \subsection{Correcting for other biases} It is possible that in some circumstances you will wish to correct not just for length bias, but for the total number of counts. This can make sense because power to detect DE depends on the total number of counts a gene receives, which is the product of gene length and gene expression. So correcting for read count bias will compensate for all biases, known and unknown, in power to detect DE. On the other hand, it will also remove bias resulting from differences in expression level, which may not be desirable. \np Correcting for count bias will produce a different PWF. Therefore, we need to tell \texttt{goseq} about the data on which the fraction DE depends when calculating the PWF using the \texttt{nullp} function. We then simply pass the result to \texttt{goseq} as usual. \np So, in order to tell \texttt{goseq} to correct for read count bias instead of length bias, all you need to do is supply a numeric vector, containing the number of counts for each gene to \texttt{nullp}. <>= countbias=rowSums(counts)[rowSums(counts)!=0] length(countbias) length(genes) @ To use the count bias when doing GO analysis, simply pass this vector to \texttt{nullp} using the \texttt{bias.data} option. Note that we have to supply "hg19" and "ensGene" to \texttt{goseq} as it is not used by nullp and hence not in the pwf.counts object. <>= pwf.counts=nullp(genes,bias.data=countbias) GO.counts=goseq(pwf.counts,"hg19","ensGene") head(GO.counts) @ Note that if you want to correct for length bias, but your organism/gene identifier is not natively supported, then you need to follow the same procedure as above, only the numeric vector supplied will contain each gene's length instead of its number of reads. \section{Setup} This vignette was built on: <>= sessionInfo() @ \section{Acknowledgments} Christopher Fjell for a series of bug fixes and pointing out the difference between the egGO and egGO2ALLEGS objects in the organism packages. \bibliographystyle{plainnat} \bibliography{goseq_bib} \end{document} goseq/vignettes/goseq_bib.bib0000644000175200017520000001700614136047504017342 0ustar00biocbuildbiocbuild%% This BibTeX bibliography file in UTF-8 format was created using Papers. %% http://mekentosj.com/papers/ @article{Young10, author = {M. D. Young and M. J. Wakefield and G. K. Smyth and A. Oshlack}, journal = {Genome Biology}, title = {Gene ontology analysis for {RNA}-seq: accounting for selection bias}, number = {}, pages = {R14}, volume = {11}, year = {2010}, month = {Feb}, } @article{Benjamini95, author = {Y. Benjamini and Y. Hochberg}, journal = {Journal of the Royal Statistical Society: Series B}, title = {Controlling the false discovery rate: a practical and powerful approach to multiple testing}, number = {}, pages = {289--300}, volume = {57}, year = {1995}, month = {}, date-added = {2009-05-19 23:56:23 +1000}, date-modified = {2009-07-03 11:54:22 +1000}, doi = {}, local-url = {f}, uri = {}, read = {Yes}, rating = {5} } @article{Robinson07, author = {M. D. Robinson and G. K. Smyth}, journal = {Bioinformatics}, title = {Moderated statistical tests for assessing differences in tag abundance}, number = {21}, pages = {2881--2887}, volume = {23}, year = {2007}, month = {}, date-added = {2009-05-19 23:56:23 +1000}, date-modified = {2009-07-03 11:54:22 +1000}, doi = {}, local-url = {f}, uri = {}, read = {Yes}, rating = {5} } @article{Robinson08, author = {M. D. Robinson and G. K. Smyth}, journal = {Biostatistics}, title = {Small-sample estimation of negative binomial dispersion, with applications to SAGE data}, number = {2}, pages = {321--332}, volume = {9}, year = {2008}, month = {}, date-added = {2009-05-19 23:56:23 +1000}, date-modified = {2009-07-03 11:54:22 +1000}, doi = {}, local-url = {f}, uri = {}, read = {Yes}, rating = {5} } @article{Robinson10, author = {M. D. Robinson and D. J. McCarthy and G. K. Smyth}, journal = {Bioinformatics}, title = {edge{R}: a Bioconductor package for differential expression analysis of digital gene expression data}, abstract = {SUMMARY: It is expected that emerging digital gene expression (DGE) technologies will overtake microarray technologies in the near future for many functional genomics applications. One of the fundamental data analysis tasks, especially for gene expression studies, involves determining whether there is evidence that counts for a transcript or exon are significantly different across experimental conditions. edgeR is a Bioconductor software package for examining differential expression of replicated count data. An overdispersed Poisson model is used to account for both biological and technical variability. Empirical Bayes methods are used to moderate the degree of overdispersion across transcripts, improving the reliability of inference. The methodology can be used even with the most minimal levels of replication, provided at least one phenotype or experimental condition is replicated. The software may have other applications beyond sequencing data, such as proteome peptide count data. AVAILABILITY: The package is freely available under the LGPL licence from the Bioconductor web site (http://bioconductor.org).}, affiliation = {Cancer Program, Garvan Institute of Medical Research, 384 Victoria Street, Darlinghurst, NSW 2010, Australia. mrobinson@wehi.edu.au}, number = {1}, pages = {139--40}, volume = {26}, year = {2010}, month = {Jan}, language = {eng}, date-added = {2010-02-02 14:01:09 +1100}, date-modified = {2010-02-02 14:01:09 +1100}, doi = {10.1093/bioinformatics/btp616}, pii = {btp616}, pmid = {19910308}, uri = {papers://C20AA413-5252-4F2C-BFAB-68084A7E2228/Paper/p567}, rating = {0} } @article{Li:2008p113, author = {Hairi Li and Michael T Lovci and Young-Soo Kwon and Michael G Rosenfeld and Xiang-Dong Fu and Gene W Yeo}, journal = {Proc Natl Acad Sci USA}, title = {Determination of tag density required for digital transcriptome analysis: application to an androgen-sensitive prostate cancer model}, abstract = {High-throughput sequencing has rapidly gained popularity for transcriptome analysis in mammalian cells because of its ability to generate digital and quantitative information on annotated genes and to detect transcripts and mRNA isoforms. Here, we described a double-random priming method for deep sequencing to profile double poly(A)-selected RNA from LNCaP cells before and after androgen stimulation. From approximately 20 million sequence tags, we uncovered 71% of annotated genes and identified hormone-regulated gene expression events that are highly correlated with quantitative real time PCR measurement. A fraction of the sequence tags were mapped to constitutive and alternative splicing events to detect known and new mRNA isoforms expressed in the cell. Finally, curve fitting was used to estimate the number of tags necessary to reach a "saturating" discovery rate among individual applications. This study provides a general guide for analysis of gene expression and alternative splicing by deep sequencing.}, affiliation = {Department of Cellular and Molecular Medicine, University of California, San Diego, 9500 Gilman Drive, La Jolla, CA 92093-0651, USA.}, number = {51}, pages = {20179--84}, volume = {105}, year = {2008}, month = {Dec}, language = {eng}, keywords = {Automation, Male, Alternative Splicing, Expressed Sequence Tags, Polymerase Chain Reaction, Cell Line: Tumor, Prostatic Neoplasms, Gene Expression Profiling, Animals, RNA: Messenger, Androgens}, date-added = {2009-11-26 12:49:28 +1100}, date-modified = {2009-11-26 12:49:28 +1100}, doi = {10.1073/pnas.0807121105}, pii = {0807121105}, pmid = {19088194}, uri = {papers://C20AA413-5252-4F2C-BFAB-68084A7E2228/Paper/p113}, read = {Yes}, rating = {0} } @article{Oshlack:2009p149, author = {Alicia Oshlack and Matthew J Wakefield}, journal = {Biol Direct}, title = {Transcript length bias in {RNA}-seq data confounds systems biology}, abstract = {BACKGROUND: Several recent studies have demonstrated the effectiveness of deep sequencing for transcriptome analysis (RNA-seq) in mammals. As RNA-seq becomes more affordable, whole genome transcriptional profiling is likely to become the platform of choice for species with good genomic sequences. As yet, a rigorous analysis methodology has not been developed and we are still in the stages of exploring the features of the data. RESULTS: We investigated the effect of transcript length bias in RNA-seq data using three different published data sets. For standard analyses using aggregated tag counts for each gene, the ability to call differentially expressed genes between samples is strongly associated with the length of the transcript. CONCLUSION: Transcript length bias for calling differentially expressed genes is a general feature of current protocols for RNA-seq technology. This has implications for the ranking of differentially expressed genes, and in particular may introduce bias in gene set testing for pathway analysis and other multi-gene systems biology analyses. REVIEWERS: This article was reviewed by Rohan Williams (nominated by Gavin Huttley), Nicole Cloonan (nominated by Mark Ragan) and James Bullard (nominated by Sandrine Dudoit).}, affiliation = {Walter and Eliza Hall Institute of Medical Research, Parkville, Vic, Australia. oshlack@wehi.edu.au}, pages = {14}, volume = {4}, year = {2009}, month = {Jan}, language = {eng}, keywords = {Transcription: Genetic, Base Sequence, Humans, Gene Expression Profiling, Oligonucleotide Array Sequence Analysis, Bias (Epidemiology), Systems Biology, RNA: Messenger}, date-added = {2009-11-26 13:00:07 +1100}, date-modified = {2009-11-26 13:00:07 +1100}, doi = {10.1186/1745-6150-4-14}, pii = {1745-6150-4-14}, pmid = {19371405}, uri = {papers://C20AA413-5252-4F2C-BFAB-68084A7E2228/Paper/p149}, read = {Yes}, rating = {0} }