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CCBA.ssgsea.R
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CCBA.ssgsea.R
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#-------------------------------------------------------------------------------------------------
CCBA_ssGSEA_project_dataset.v1 <- function(
#
# Project dataset into pathways or gene sets using ssGSEA
# P. Tamayo Jan 17, 2016
#
input.ds,
output.ds,
gene.set.databases,
gene.set.selection = "ALL", # "ALL" or list with names of gene sets
sample.norm.type = "rank", # "rank", "log" or "log.rank"
weight = 0.25,
statistic = "area.under.RES",
output.score.type = "ES", # "ES" or "NES"
nperm = 200, # number of random permutations for NES case
combine.mode = "combine.off", # "combine.off" do not combine *_UP and *_DN versions in
# a single score. "combine.replace" combine *_UP and
# *_DN versions in a single score that replaces the individual
# *_UP and *_DN versions. "combine.add" combine *_UP and
# *_DN versions in a single score and add it but keeping
# the individual *_UP and *_DN versions.
min.overlap = 1,
gene.names.in.desc = F, # in Protein, RNAi Ataris or hairpin gct files the gene symbols are in the descs column
correl.type = "rank") # "rank", "z.score", "symm.rank"
{
# Read input dataset
dataset <- CCBA_read_GCT_file.v1(filename = input.ds) # Read gene expression dataset (GCT format)
m <- data.matrix(dataset$ds)
if (gene.names.in.desc == T) {
gene.names <- dataset$descs
} else {
gene.names <- dataset$row.names
}
gene.descs <- dataset$descs
sample.names <- dataset$names
Ns <- length(m[1,])
Ng <- length(m[,1])
temp <- strsplit(input.ds, split="/") # Extract input file name
s <- length(temp[[1]])
input.file.name <- temp[[1]][s]
temp <- strsplit(input.file.name, split=".gct")
input.file.prefix <- temp[[1]][1]
# Sample normalization
if (sample.norm.type == "rank") {
for (j in 1:Ns) { # column rank normalization
m[,j] <- rank(m[,j], ties.method = "average")
}
m <- 10000*m/Ng
} else if (sample.norm.type == "log.rank") {
for (j in 1:Ns) { # column rank normalization
m[,j] <- rank(m[,j], ties.method = "average")
}
m <- log(10000*m/Ng + exp(1))
} else if (sample.norm.type == "log") {
m[m < 1] <- 1
m <- log(m + exp(1))
}
# Read gene set databases
max.G <- 0
max.N <- 0
for (gsdb in gene.set.databases) {
GSDB <- CCBA_Read.GeneSets.db.v1(gsdb, thres.min = 2, thres.max = 2000, gene.names = NULL)
max.G <- max(max.G, max(GSDB$size.G))
max.N <- max.N + GSDB$N.gs
}
N.gs <- 0
gs <- matrix("null", nrow=max.N, ncol=max.G)
gs.names <- vector(length=max.N, mode="character")
gs.descs <- vector(length=max.N, mode="character")
size.G <- vector(length=max.N, mode="numeric")
start <- 1
for (gsdb in gene.set.databases) {
GSDB <- CCBA_Read.GeneSets.db.v1(gsdb, thres.min = 2, thres.max = 2000, gene.names = NULL)
N.gs <- GSDB$N.gs
gs.names[start:(start + N.gs - 1)] <- GSDB$gs.names
gs.descs[start:(start + N.gs - 1)] <- GSDB$gs.desc
size.G[start:(start + N.gs - 1)] <- GSDB$size.G
gs[start:(start + N.gs - 1), 1:max(GSDB$size.G)] <- GSDB$gs[1:N.gs, 1:max(GSDB$size.G)]
start <- start + N.gs
}
N.gs <- max.N
# Select desired gene sets
if (gene.set.selection[1] == "ALL") {
gene.set.selection <- unique(gs.names)
}
locs <- match(gene.set.selection, gs.names)
# print(rbind(gene.set.selection, locs))
N.gs <- sum(!is.na(locs))
if(N.gs > 1) {
gs <- gs[locs,]
} else {
gs <- t(as.matrix(gs[locs,])) # Force vector to matrix if only one gene set specified
}
gs.names <- gs.names[locs]
gs.descs <- gs.descs[locs]
size.G <- size.G[locs]
# Check for redundant gene sets
tab <- as.data.frame(table(gs.names))
ind <- order(tab[, "Freq"], decreasing=T)
tab <- tab[ind,]
max.n <- max(10, length(gs.names))
print(tab[1:max.n,])
print(paste("Total gene sets:", length(gs.names)))
print(paste("Unique gene sets:", length(unique(gs.names))))
# Loop over gene sets
score.matrix <- score.matrix.2 <- matrix(0, nrow=N.gs, ncol=Ns)
print(paste("Size score.matrix:", dim(score.matrix)))
print(paste("Size score.matrix.2:", dim(score.matrix.2)))
for (gs.i in 1:N.gs) {
#browser()
gene.set <- gs[gs.i, 1:size.G[gs.i]]
gene.overlap <- intersect(gene.set, gene.names)
print(paste(gs.i, "gene set:", gs.names[gs.i], " overlap=", length(gene.overlap)))
if (length(gene.overlap) < min.overlap) {
score.matrix[gs.i, ] <- rep(NA, Ns)
print(paste("Size score.matrix:", dim(score.matrix)))
next
} else {
gene.set.locs <- match(gene.overlap, gene.set)
gene.names.locs <- match(gene.overlap, gene.names)
msig <- m[gene.names.locs,]
msig.names <- gene.names[gene.names.locs]
if (output.score.type == "ES") {
OPAM <- CCBA_ssGSEA.Projection.v1(data.array = m, gene.names = gene.names, n.cols = Ns,
n.rows = Ng, weight = weight, statistic = statistic,
gene.set = gene.overlap, nperm = 1, correl.type = correl.type)
score.matrix[gs.i,] <- as.matrix(t(OPAM$ES.vector))
print(paste("Size score.matrix:", dim(score.matrix)))
} else if (output.score.type == "NES") {
OPAM <- CCBA_ssGSEA.Projection.v1(data.array = m, gene.names = gene.names, n.cols = Ns,
n.rows = Ng, weight = weight, statistic = statistic,
gene.set = gene.overlap, nperm = nperm, correl.type = correl.type)
score.matrix[gs.i,] <- as.matrix(t(OPAM$NES.vector))
print(paste("Size score.matrix:", dim(score.matrix)))
}
}
}
locs <- !is.na(score.matrix[,1])
print(paste("N.gs before overlap prunning:", N.gs))
N.gs <- sum(locs)
print(paste("N.gs after overlap prunning:", N.gs))
if (nrow(score.matrix) == 1) {
score.matrix <- as.matrix(t(score.matrix[locs,]))
} else {
score.matrix <- score.matrix[locs,]
}
print(paste("Size score.matrix:", dim(score.matrix)))
gs.names <- gs.names[locs]
gs.descs <- gs.descs[locs]
initial.up.entries <- 0
final.up.entries <- 0
initial.dn.entries <- 0
final.dn.entries <- 0
combined.entries <- 0
other.entries <- 0
if (combine.mode == "combine.off") {
if (nrow(score.matrix) == 1) {
score.matrix.2 <- as.matrix(t(score.matrix))
} else {
score.matrix.2 <- score.matrix
}
print(paste("Size score.matrix.2:", dim(score.matrix.2)))
gs.names.2 <- gs.names
gs.descs.2 <- gs.descs
} else if ((combine.mode == "combine.replace") || (combine.mode == "combine.add")) {
score.matrix.2 <- NULL
gs.names.2 <- NULL
gs.descs.2 <- NULL
k <- 1
for (i in 1:N.gs) {
temp <- strsplit(gs.names[i], split="_")
body <- paste(temp[[1]][seq(1, length(temp[[1]]) -1)], collapse="_")
suffix <- tail(temp[[1]], 1)
print(paste("i:", i, "gene set:", gs.names[i], "body:", body, "suffix:", suffix))
if (suffix == "UP") { # This is an "UP" gene set
initial.up.entries <- initial.up.entries + 1
target <- paste(body, "DN", sep="_")
loc <- match(target, gs.names)
if (!is.na(loc)) { # found corresponding "DN" gene set: create combined entry
score <- score.matrix[i,] - score.matrix[loc,]
score.matrix.2 <- rbind(score.matrix.2, score)
gs.names.2 <- c(gs.names.2, body)
gs.descs.2 <- c(gs.descs.2, paste(gs.descs[i], "combined UP & DN"))
combined.entries <- combined.entries + 1
if (combine.mode == "combine.add") { # also add the "UP entry
if (nrow(score.matrix) == 1) {
score.matrix.2 <- rbind(score.matrix.2, as.matrix(t(score.matrix[i,])))
} else {
score.matrix.2 <- rbind(score.matrix.2, score.matrix[i,])
}
print(paste("Size score.matrix.2:", dim(score.matrix.2)))
gs.names.2 <- c(gs.names.2, gs.names[i])
gs.descs.2 <- c(gs.descs.2, gs.descs[i])
final.up.entries <- final.up.entries + 1
}
} else { # did not find corresponding "DN" gene set: create "UP" entry
if (nrow(score.matrix) == 1) {
score.matrix.2 <- rbind(score.matrix.2, as.matrix(t(score.matrix[i,])))
} else {
score.matrix.2 <- rbind(score.matrix.2, score.matrix[i,])
}
print(paste("Size score.matrix.2:", dim(score.matrix.2)))
gs.names.2 <- c(gs.names.2, gs.names[i])
gs.descs.2 <- c(gs.descs.2, gs.descs[i])
final.up.entries <- final.up.entries + 1
}
} else if (suffix == "DN") { # This is a "DN" gene set
initial.dn.entries <- initial.dn.entries + 1
target <- paste(body, "UP", sep="_")
loc <- match(target, gs.names)
if (is.na(loc)) { # did not find corresponding "UP" gene set: create "DN" entry
if (nrow(score.matrix) == 1) {
score.matrix.2 <- rbind(score.matrix.2, as.matrix(t(score.matrix[i,])))
} else {
score.matrix.2 <- rbind(score.matrix.2, score.matrix[i,])
}
print(paste("Size score.matrix.2:", dim(score.matrix.2)))
gs.names.2 <- c(gs.names.2, gs.names[i])
gs.descs.2 <- c(gs.descs.2, gs.descs[i])
final.dn.entries <- final.dn.entries + 1
} else { # it found corresponding "UP" gene set
if (combine.mode == "combine.add") { # create "DN" entry
if (nrow(score.matrix) == 1) {
score.matrix.2 <- rbind(score.matrix.2, as.matrix(t(score.matrix[i,])))
} else {
score.matrix.2 <- rbind(score.matrix.2, score.matrix[i,])
}
print(paste("Size score.matrix.2:", dim(score.matrix.2)))
gs.names.2 <- c(gs.names.2, gs.names[i])
gs.descs.2 <- c(gs.descs.2, gs.descs[i])
final.dn.entries <- final.dn.entries + 1
}
}
} else { # This is neither "UP nor "DN" gene set: create individual entry
if (nrow(score.matrix) == 1) {
score.matrix.2 <- rbind(score.matrix.2, as.matrix(t(score.matrix[i,])))
} else {
score.matrix.2 <- rbind(score.matrix.2, score.matrix[i,])
}
print(paste("Size score.matrix.2:", dim(score.matrix.2)))
gs.names.2 <- c(gs.names.2, gs.names[i])
gs.descs.2 <- c(gs.descs.2, gs.descs[i])
other.entries <- other.entries + 1
}
} # end for loop over gene sets
print(paste("initial.up.entries:", initial.up.entries))
print(paste("final.up.entries:", final.up.entries))
print(paste("initial.dn.entries:", initial.dn.entries))
print(paste("final.dn.entries:", final.dn.entries))
print(paste("other.entries:", other.entries))
print(paste("combined.entries:", combined.entries))
print(paste("total entries:", length(score.matrix.2[,1])))
}
# Make sure there are no duplicated gene names after adding entries
unique.gene.sets <- unique(gs.names.2)
locs <- match(unique.gene.sets, gs.names.2)
if (nrow(score.matrix) == 1) {
score.matrix.2 <- as.matrix(t(score.matrix.2[locs,]))
} else {
score.matrix.2 <- score.matrix.2[locs,]
}
gs.names.2 <- gs.names.2[locs]
gs.descs.2 <- gs.descs.2[locs]
# Final count
tab <- as.data.frame(table(gs.names.2))
ind <- order(tab[, "Freq"], decreasing=T)
tab <- tab[ind,]
print(tab[1:20,])
print(paste("Total gene sets:", length(gs.names.2)))
print(paste("Unique gene sets:", length(unique(gs.names.2))))
V.GCT <- data.frame(score.matrix.2)
colnames(V.GCT) <- sample.names
row.names(V.GCT) <- gs.names.2
CCBA_write.gct.v1(gct.data.frame = V.GCT, descs = gs.descs.2, filename = output.ds)
}
#-------------------------------------------------------------------------------------------------
CCBA_Read.GeneSets.db.v1 <- function(
#
# Read gene sets from a database (GMT file)
# P. Tamayo Jan 17, 2016
#
gs.db,
thres.min = 2,
thres.max = 2000,
gene.names = NULL)
{
temp <- readLines(gs.db)
max.Ng <- length(temp)
temp.size.G <- vector(length = max.Ng, mode = "numeric")
for (i in 1:max.Ng) {
temp.size.G[i] <- length(unlist(strsplit(temp[[i]], "\t"))) - 2
}
max.size.G <- max(temp.size.G)
gs <- matrix(rep("null", max.Ng*max.size.G), nrow=max.Ng, ncol= max.size.G)
temp.names <- vector(length = max.Ng, mode = "character")
temp.desc <- vector(length = max.Ng, mode = "character")
gs.count <- 1
for (i in 1:max.Ng) {
gene.set.size <- length(unlist(strsplit(temp[[i]], "\t"))) - 2
gs.line <- noquote(unlist(strsplit(temp[[i]], "\t")))
gene.set.name <- gs.line[1]
gene.set.desc <- gs.line[2]
gene.set.tags <- vector(length = gene.set.size, mode = "character")
for (j in 1:gene.set.size) {
gene.set.tags[j] <- gs.line[j + 2]
}
if (is.null(gene.names)) {
existing.set <- rep(TRUE, length(gene.set.tags))
} else {
existing.set <- is.element(gene.set.tags, gene.names)
}
set.size <- length(existing.set[existing.set == T])
if ((set.size < thres.min) || (set.size > thres.max)) next
temp.size.G[gs.count] <- set.size
gs[gs.count,] <- c(gene.set.tags[existing.set], rep("null", max.size.G - temp.size.G[gs.count]))
temp.names[gs.count] <- gene.set.name
temp.desc[gs.count] <- gene.set.desc
gs.count <- gs.count + 1
}
Ng <- gs.count - 1
gs.names <- vector(length = Ng, mode = "character")
gs.desc <- vector(length = Ng, mode = "character")
size.G <- vector(length = Ng, mode = "numeric")
gs.names <- temp.names[1:Ng]
gs.desc <- temp.desc[1:Ng]
size.G <- temp.size.G[1:Ng]
return(list(N.gs = Ng, gs = gs, gs.names = gs.names, gs.desc = gs.desc, size.G = size.G, max.N.gs = max.Ng))
}
#-------------------------------------------------------------------------------------------------
CCBA_ssGSEA.Projection.v1 <- function(
#
# ssGSEA projection
# P. Tamayo Jan 17, 2016
#
# Runs a 2-3x faster (2-2.5x for ES statistic and 2.5-3x faster for area.under.ES statsitic)
# version of GSEA.EnrichmentScore.5 internally that avoids overhead from the function call.
# This function use dto be OPAM.Projection.3
data.array,
gene.names,
n.cols,
n.rows,
weight = 0,
statistic = "Kolmogorov-Smirnov", # "Kolmogorov-Smirnov", # "Kolmogorov-Smirnov", "Cramer-von-Mises",
# "Anderson-Darling", "Zhang_A", "Zhang_C", "Zhang_K",
# "area.under.RES", or "Wilcoxon"
gene.set,
nperm = 200,
correl.type = "rank") # "rank", "z.score", "symm.rank"
{
ES.vector <- vector(length=n.cols)
NES.vector <- vector(length=n.cols)
p.val.vector <- vector(length=n.cols)
correl.vector <- vector(length=n.rows, mode="numeric")
# Compute ES score for signatures in each sample
# print("Computing GSEA.....")
phi <- array(0, c(n.cols, nperm))
for (sample.index in 1:n.cols) {
gene.list <- order(data.array[, sample.index], decreasing=T)
# print(paste("Computing observed enrichment for UP signature in sample:", sample.index, sep=" "))
gene.set2 <- match(gene.set, gene.names)
if (weight == 0) {
correl.vector <- rep(1, n.rows)
} else if (weight > 0) {
if (correl.type == "rank") {
correl.vector <- data.array[gene.list, sample.index]
} else if (correl.type == "symm.rank") {
correl.vector <- data.array[gene.list, sample.index]
correl.vector <- ifelse(correl.vector > correl.vector[ceiling(n.rows/2)],
correl.vector,
correl.vector + correl.vector - correl.vector[ceiling(n.rows/2)])
} else if (correl.type == "z.score") {
x <- data.array[gene.list, sample.index]
correl.vector <- (x - mean(x))/sd(x)
}
}
### Olga's Additions ###
# ptm.new = proc.time()
tag.indicator <- sign(match(gene.list, gene.set2, nomatch=0)) # notice that the sign is 0 (no tag) or 1 (tag)
no.tag.indicator <- 1 - tag.indicator
N <- length(gene.list)
Nh <- length(gene.set2)
Nm <- N - Nh
orig.correl.vector <- correl.vector
if (weight == 0) correl.vector <- rep(1, N) # unweighted case
ind = which(tag.indicator==1)
correl.vector <- abs(correl.vector[ind])^weight
sum.correl = sum(correl.vector)
up = correl.vector/sum.correl # "up" represents the peaks in the mountain plot
gaps = (c(ind-1, N) - c(0, ind)) # gaps between ranked pathway genes
down = gaps/Nm
RES = cumsum(c(up,up[Nh])-down)
valleys = RES[1:Nh]-up
max.ES = max(RES)
min.ES = min(valleys)
if( statistic == "Kolmogorov-Smirnov" ){
if( max.ES > -min.ES ){
ES <- signif(max.ES, digits=5)
arg.ES <- which.max(RES)
} else{
ES <- signif(min.ES, digits=5)
arg.ES <- which.min(RES)
}
}
if( statistic == "area.under.RES"){
if( max.ES > -min.ES ){
arg.ES <- which.max(RES)
} else{
arg.ES <- which.min(RES)
}
gaps = gaps+1
RES = c(valleys,0) * (gaps) + 0.5*( c(0,RES[1:Nh]) - c(valleys,0) ) * (gaps)
ES = sum(RES)
}
GSEA.results = list(ES = ES, arg.ES = arg.ES, RES = RES, indicator = tag.indicator)
# new.time <<- new.time + (proc.time() - ptm.new)
### End Olga's Additions ###
#GSEA.results <- GSEA.EnrichmentScore5(gene.list=gene.list, gene.set=gene.set2,
# statistic = statistic, alpha = weight, correl.vector = correl.vector)
ES.vector[sample.index] <- GSEA.results$ES
if (nperm == 0) {
NES.vector[sample.index] <- ES.vector[sample.index]
p.val.vector[sample.index] <- 1
} else {
for (r in 1:nperm) {
reshuffled.gene.labels <- sample(1:n.rows)
if (weight == 0) {
correl.vector <- rep(1, n.rows)
} else if (weight > 0) {
correl.vector <- data.array[reshuffled.gene.labels, sample.index]
}
# GSEA.results <- GSEA.EnrichmentScore5(gene.list=reshuffled.gene.labels, gene.set=gene.set2,
# statistic = statistic, alpha = weight, correl.vector = correl.vector)
### Olga's Additions ###
tag.indicator <- sign(match(reshuffled.gene.labels, gene.set2, nomatch=0)) # notice that the sign is 0 (no tag) or 1 (tag)
no.tag.indicator <- 1 - tag.indicator
N <- length(reshuffled.gene.labels)
Nh <- length(gene.set2)
Nm <- N - Nh
# orig.correl.vector <- correl.vector
if (weight == 0) correl.vector <- rep(1, N) # unweighted case
ind <- which(tag.indicator==1)
correl.vector <- abs(correl.vector[ind])^weight
sum.correl <- sum(correl.vector)
up = correl.vector/sum.correl
gaps = (c(ind-1, N) - c(0, ind))
down = gaps/Nm
RES = cumsum(c(up,up[Nh])-down)
valleys = RES[1:Nh]-up
max.ES = max(RES)
min.ES = min(valleys)
if( statistic == "Kolmogorov-Smirnov" ){
if( max.ES > -min.ES ){
ES <- signif(max.ES, digits=5)
arg.ES <- which.max(RES)
} else{
ES <- signif(min.ES, digits=5)
arg.ES <- which.min(RES)
}
}
if( statistic == "area.under.RES"){
if( max.ES > -min.ES ){
arg.ES <- which.max(RES)
} else{
arg.ES <- which.min(RES)
}
gaps = gaps+1
RES = c(valleys,0) * (gaps) + 0.5*( c(0,RES[1:Nh]) - c(valleys,0) ) * (gaps)
ES = sum(RES)
}
GSEA.results = list(ES = ES, arg.ES = arg.ES, RES = RES, indicator = tag.indicator)
### End Olga's Additions ###
phi[sample.index, r] <- GSEA.results$ES
}
if (ES.vector[sample.index] >= 0) {
pos.phi <- phi[sample.index, phi[sample.index, ] >= 0]
if (length(pos.phi) == 0) pos.phi <- 0.5
pos.m <- mean(pos.phi)
NES.vector[sample.index] <- ES.vector[sample.index]/pos.m
s <- sum(pos.phi >= ES.vector[sample.index])/length(pos.phi)
p.val.vector[sample.index] <- ifelse(s == 0, 1/nperm, s)
} else {
neg.phi <- phi[sample.index, phi[sample.index, ] < 0]
if (length(neg.phi) == 0) neg.phi <- 0.5
neg.m <- mean(neg.phi)
NES.vector[sample.index] <- ES.vector[sample.index]/abs(neg.m)
s <- sum(neg.phi <= ES.vector[sample.index])/length(neg.phi)
p.val.vector[sample.index] <- ifelse(s == 0, 1/nperm, s)
}
}
}
return(list(ES.vector = ES.vector, NES.vector = NES.vector, p.val.vector = p.val.vector))
}
#-------------------------------------------------------------------------------------------------
CCBA_read_GCT_file.v1 <- function(filename = "NULL")
#
# Reads a gene expression dataset in GCT format and converts it into an R data frame
# Pablo Tamayo Dec 30, 2015
#
{
ds <- read.delim(filename, header=T, sep="\t", skip=2, row.names=1, blank.lines.skip=T,
comment.char="", as.is=T, na.strings = "")
descs <- ds[,1]
ds <- ds[-1]
row.names <- row.names(ds)
names <- names(ds)
return(list(ds = ds, row.names = row.names, descs = descs, names = names))
}
#-------------------------------------------------------------------------------------------------
CCBA_write.gct.v1 <- function(
#
# Write data frame to a GCT file
# P. Tamayo Jan 17, 2016
#
gct.data.frame,
descs = "",
filename)
{
f <- file(filename, "w")
cat("#1.2", "\n", file = f, append = TRUE, sep = "")
cat(dim(gct.data.frame)[1], "\t", dim(gct.data.frame)[2], "\n", file = f, append = TRUE, sep = "")
cat("Name", "\t", file = f, append = TRUE, sep = "")
cat("Description", file = f, append = TRUE, sep = "")
colnames <- colnames(gct.data.frame)
cat("\t", colnames[1], file = f, append = TRUE, sep = "")
if (length(colnames) > 1) {
for (j in 2:length(colnames)) {
cat("\t", colnames[j], file = f, append = TRUE, sep = "")
}
}
cat("\n", file = f, append = TRUE, sep = "\t")
oldWarn <- options(warn = -1)
m <- matrix(nrow = dim(gct.data.frame)[1], ncol = dim(gct.data.frame)[2] + 2)
m[, 1] <- row.names(gct.data.frame)
if (length(descs) > 1) {
m[, 2] <- descs
} else {
m[, 2] <- row.names(gct.data.frame)
}
index <- 3
for (i in 1:dim(gct.data.frame)[2]) {
m[, index] <- gct.data.frame[, i]
index <- index + 1
}
write.table(m, file = f, append = TRUE, quote = FALSE, sep = "\t", eol = "\n", col.names = FALSE, row.names = FALSE)
close(f)
options(warn = 0)
}