From 5fd57d4c00ddb29265ed9e3611c3f05a4cf6365e Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Wed, 17 Jun 2020 22:23:11 +0200 Subject: [PATCH 01/28] Add draft of the qpAdm rotation method --- NAMESPACE | 2 + R/config_files.R | 11 +++ R/log.R | 21 +++--- R/qpAdm.R | 153 +++++++++++++++++++++++++++++++++++++++++ R/wrappers.R | 19 +++-- Untitled.R | 108 +++++++++++++++++++++++++++++ man/f4ratio.Rd | 18 ++++- man/qpAdm_filter.Rd | 20 ++++++ man/qpAdm_prescreen.Rd | 24 +++++++ man/qpAdm_rotation.Rd | 28 ++++++++ 10 files changed, 387 insertions(+), 17 deletions(-) create mode 100644 R/qpAdm.R create mode 100644 Untitled.R create mode 100644 man/qpAdm_filter.Rd create mode 100644 man/qpAdm_prescreen.Rd create mode 100644 man/qpAdm_rotation.Rd diff --git a/NAMESPACE b/NAMESPACE index 1f9037b..3270cd7 100644 --- a/NAMESPACE +++ b/NAMESPACE @@ -15,6 +15,8 @@ export(keep_transitions) export(loginfo) export(merge_eigenstrat) export(qpAdm) +export(qpAdm_prescreen) +export(qpAdm_rotation) export(qpWave) export(read_geno) export(read_ind) diff --git a/R/config_files.R b/R/config_files.R index 954325d..4f507d8 100644 --- a/R/config_files.R +++ b/R/config_files.R @@ -43,6 +43,17 @@ create_qpDstat_pop_file <- function(W = NULL, X = NULL, Y = NULL, Z = NULL, file } +# Generate a file with populations for a qpDstat run based on given +# list of quartets. +create_qpDstat_pop_file_quartets <- function(quartets, file) { + lines <- c() + for (q in quartets) { + lines <- c(lines, sprintf("%s %s %s %s", q[1], q[2], q[3], q[4])) + } + writeLines(lines, file) +} + + # Generate a file with populations for a qp3Pop run. create_qp3Pop_pop_file <- function(A, B, C, file) { lines <- c() diff --git a/R/log.R b/R/log.R index dda1e36..e1b0fb7 100644 --- a/R/log.R +++ b/R/log.R @@ -18,23 +18,23 @@ loginfo <- function(x, target = NA, save = FALSE, prefix = NA, dir = ".", suffix log_output <- attr(x, "log_output") targets <- names(log_output) - if (!is.na(target) && cmd != "qpAdm") + if (!is.na(target) && !cmd %in% c("qpAdm", "qpAdm_rotation")) stop(glue::glue("Specifying target does not make sense for examining the log output of {cmd}"), call. = FALSE) if (!is.na(target) && !target %in% targets) - stop(glue::glue("Target '{target}' is not present in the output (choices are: {paste(targets, collapse = ', ')})"), + stop(glue::glue("Target/model '{target}' is not present in the output (choices are: {paste(targets, collapse = ', ')})"), call. = FALSE) - # qpAdm's log output are stored as a list of character vectors but everything - # else is simply a character vector - we convert everything to a list to - # iterate over log outputs below + ## qpAdm's log output are stored as a list of character vectors but everything + ## else is simply a character vector - we convert everything to a list to + ## iterate over log outputs below if (!is.list(log_output)) log_output <- list(log_output) for (i in seq_along(log_output)) { - # write only single target log if requested by the user - if (!is.na(target) && !is.null(targets) && target != targets[i]) next + ## write only single target/model log if requested by the user + if (!is.na(target) && !is.null(targets) && target != targets[i]) next if (save) { if (is.na(prefix)) prefix <- cmd @@ -49,14 +49,17 @@ loginfo <- function(x, target = NA, save = FALSE, prefix = NA, dir = ".", suffix } else { if (cmd == "qpAdm") { title <- glue::glue("qpAdm for target '{targets[i]}'") + } else if (cmd == "qpAdm_rotation") { + title <- glue::glue("qpAdm rotation for model '{targets[i]}'") } else { title <- cmd } if (i > 1 && is.na(target)) cat("\n\n") cat(paste0("Full output log of ", title, ":\n")) - cat("==================================================\n\n") + cat("===================================================\n\n") cat(paste(log_output[[i]], collapse = "\n")) + cat("\n") } } } @@ -70,7 +73,7 @@ loginfo <- function(x, target = NA, save = FALSE, prefix = NA, dir = ".", suffix #' #' @export print.admixr_result <- function(x, ...) { - if (attr(x, "command") == "qpAdm") { + if (attr(x, "command") %in% c("qpAdm", "qpAdm_rotation")) { print.default(list( proportions = x$proportions, ranks = x$ranks, diff --git a/R/qpAdm.R b/R/qpAdm.R new file mode 100644 index 0000000..a832270 --- /dev/null +++ b/R/qpAdm.R @@ -0,0 +1,153 @@ +#' Perform a 'pre-screening' of the 'right' populations before +#' a qpAdm analysis based on Harney et al., 2020 (bioRxiv) +#' +#' @param data EIGENSTRAT dataset +#' @param candidates Character vector with potential 'outgroup' populations +#' @param left Character vector with target and source populations +#' +#' @return Data frame of all combinations of f4(L_i, L_j; R_k, R_l), +#' ordered by Zscore of the f4 statistic +#' +#' @export +qpAdm_prescreen <- function(data, candidates, left, Zcutoff = 2) { + leftcomb <- t(combn(left, 2)) + rightcomb <- t(combn(candidates, 2)) + + i <- 1 + quartets <- list() + for (l in 1:nrow(leftcomb)) { + for (r in 1:nrow(rightcomb)) { + quartets[[i]] <- c(leftcomb[l, ], rightcomb[r, ]) + i <- i + 1 + } + } + + result <- f4(data, quartets = quartets) %>% dplyr::arrange(abs(Zscore)) + + right <- dplyr::filter(result, abs(Zscore) > Zcutoff) %>% + .[, c("Y", "Z")] %>% + as.matrix %>% + as.vector %>% + unique + + list(outgroups = right, screening = result) +} + +#' Fit qpAdm models based on the rotation strategy described in +#' Harney et al. 2020 (bioRxiv) +#' +#' @param data EIGENSTRAT dataset +#' @param target Target population that is modeled as admixed +#' @param candidates Potential candidates for sources and outgroups +#' @param nsources Number of sources to pull from the candidates +#' @param ncores Number of CPU cores to utilize for model fitting +#' +#' @return qpAdm list with proportions, ranks and subsets elements (as +#' with a traditional qpAdm run) +#' +#' @export +qpAdm_rotation <- function(data, target, candidates, nsources, ncores = 1) { + ## generate combinations of possible sources and outgroups + sources <- t(combn(candidates, nsources)) + sources_outgroups <- lapply(1:nrow(sources), function(i) { + list( + sources = sources[i, ], + outgroups = setdiff(candidates, sources[i, ]) + ) + }) + + ## run qpAdm for all combinations of sources and outgroups + results_list <- parallel::mclapply(sources_outgroups, function(x) { + result <- qpAdm( + data, + target = target, sources = x$sources, outgroups = x$outgroups + ) + + names(x$sources) <- paste0("source", 1:nsources) + sources_df <- as.data.frame(t(as.matrix(x$sources))) + + ## rename sources and stderr columns (by default, proportion and + ## stderr columns are named based on the source populations - we + ## don't want that here because we want to merge all the individual + ## proportion tables, columns have to have the same name) + names(result$proportions)[2:(1 + nsources)] <- paste0("prop", 1:nsources) + names(result$proportions)[4:(3 + nsources)] <- paste0("stderr", 1:nsources) + ## add source names as two new columns + result$proportions <- cbind(result$proportions, sources_df) + ## rearrange columns + result$proportions <- dplyr::select( + result$proportions, target, names(x$sources), pvalue, + dplyr::everything() + ) + + ## reformat rank table + names(result$subsets)[7:(6 + nsources)] <- paste0("prop", 1:nsources) + ## add source names as two new columns + result$subsets <- cbind(result$subsets, sources_df) + result$subsets <- dplyr::select( + result$subsets, target, names(x$sources), pattern, + dplyr::everything() + ) + + result + }, mc.cores = ncores) + + ## extract log information before further processing of the results + log_lines <- sapply(results_list, function(i) attr(i, "log_output")) + names(log_lines) <- paste0("m", seq_along(sources_outgroups)) + + proportions <- dplyr::bind_rows(lapply(results_list, `[[`, "proportions")) + ranks <- dplyr::bind_rows(lapply(results_list, `[[`, "ranks")) + subsets <- dplyr::bind_rows(lapply(results_list, `[[`, "subsets")) + + ## add model identifier to each row in the proportions table, ... + models <- paste0("m", seq_along(sources_outgroups)) + proportions$model <- models + proportions <- dplyr::as_tibble(proportions) %>% dplyr::select(model, everything()) + ## ... ranks table, ... + ranks$model <- sort(rep(models, 2)) + ranks <- dplyr::as_tibble(ranks) %>% dplyr::select(model, everything()) + ## and subsets table + subsets$model <- sort(rep(models, 1 + 2^(nsources - 1))) + subsets <- dplyr::as_tibble(subsets) %>% dplyr::select(model, everything()) + + ## bind all tables together and add log information + results <- list(proportions = proportions, ranks = ranks, subsets = subsets) + attr(results, "command") <- "qpAdm_rotation" + attr(results, "log_output") <- log_lines + class(results) <- c("admixr_result", class(results)) + + results +} + + +#' Filter qpAdm rotation results for only 'sensible' models +#' +#' Filter for p-value larger than a specified cuttof and admixture +#' proportions between 0 and 1. +#' +#' @param x Output of a qpAdm_rotation() function +#' @param p p-value cutoff +#' +#' @return qpAdm_rotation object filtered down based on p-value +qpAdm_filter <- function(x, p) { + ## get positions of columns with estimated admixture proportions + prop_columns <- stringr::str_which(names(x$proportions), "prop") + + ## find out rows/models for which all proportions are in [0, 1] and + ## pvalue is larger than the required cutoff + pvalue <- x$proportions$pvalue > p + constr_0 <- x$proportions[, prop_columns] >= 0 + constr_1 <- x$proportions[, prop_columns] <= 1 + constr <- apply(pvalue & constr_0 & constr_1, 1, all) + + ## filter all three sub-tables to only those models that fit the criteria + x$proportions <- dplyr::arrange(x$proportions[constr, ], -pvalue) + x$ranks <- x$ranks[x$ranks$model %in% x$proportions$model, ] + x$subsets <- x$subsets[x$subsets$model %in% x$proportions$model, ] + + ## filter also only to relevant remaining log output information + attr(x, "log_output") <- attr(x, "log_output")[unique(x$proportions$model)] + + x +} diff --git a/R/wrappers.R b/R/wrappers.R index ebe066f..dc4c7d2 100644 --- a/R/wrappers.R +++ b/R/wrappers.R @@ -25,16 +25,25 @@ f4ratio <- function(data, X, A, B, C, O, outdir = NULL, params = NULL) { #' @rdname f4ratio #' #' @param f4mode Calculate the f4 statistic instead of the D statistic. +#' @param quartets List of character vectors (quartets of population/sample labels) #' #' @export -d <- function(data, W, X, Y, Z, outdir = NULL, f4mode = FALSE, params = NULL) { - check_presence(c(W, X, Y, Z), data) +d <- function(data, W, X, Y, Z, quartets = NULL, outdir = NULL, f4mode = FALSE, params = NULL) { + if (is.null(quartets)) { + check_presence(c(W, X, Y, Z, data)) + } else { + check_presence(unique(unlist(quartets)), data) + } # get the path to the population, parameter and log files config_prefix <- paste0("qpDstat__", as.integer(stats::runif(1, 0, .Machine$integer.max))) files <- get_files(outdir, config_prefix) - create_qpDstat_pop_file(W, X, Y, Z, file = files[["pop_file"]]) + if (is.null(quartets)) { + create_qpDstat_pop_file(W, X, Y, Z, file = files[["pop_file"]]) + } else { + create_qpDstat_pop_file_quartets(quartets, file = files[["pop_file"]]) + } create_par_file(files, data, params) if (f4mode) { @@ -54,8 +63,8 @@ d <- function(data, W, X, Y, Z, outdir = NULL, f4mode = FALSE, params = NULL) { #' @rdname f4ratio #' #' @export -f4 <- function(data, W, X, Y, Z, outdir = NULL, params = NULL) { - d(data, W, X, Y, Z, outdir, f4mode = TRUE) +f4 <- function(data, W, X, Y, Z, quartets, outdir = NULL, params = NULL) { + d(data, W, X, Y, Z, quartets, outdir, f4mode = TRUE) } diff --git a/Untitled.R b/Untitled.R new file mode 100644 index 0000000..26b6ec3 --- /dev/null +++ b/Untitled.R @@ -0,0 +1,108 @@ +snps <- eigenstrat(download_data()) + +result <- qpAdm( + target = c("Han", "Sardinian"), + sources = c("Vindija", "Yoruba"), + outgroups = c("Chimp", "Denisova", "Altai"), + data = snps +) + + +left <- c("Han", "Vindija", "Yoruba") +right <- setdiff(setdiff(read_ind(snps)$label, left), + c("French", "Sardinian", "Papuan", "Dinka", "Khomani_San")) + +qpAdm_prescreen(snps, right, left) + + +qpAdm_prescreen(snps, c("Chimp", "Denisova"), left) + +left <- c("Sardinian", "Dinka", "French") +candidates <- c("Chimp", "Altai", "Denisova", "Yoruba", "Mbuti") + +x = qpAdm_prescreen(snps, candidates, left) +x +arrange(x$screening, abs(Zscore)) + + + + + + +sim <- eigenstrat("/tmp/sim/snps") + +Qpadm( + target = "14", + sources = c("5", "9"), + outgroups = c("0", "7", "10", "12", "13"), + data = sim +) + +right <- c("0", "7", "10", "12", "13") +left <- c("14", "5", "9") +y <- qpAdm_prescreen(sim, right, left) + + + +right <- c("0", "1", "2", "3", "4") +left <- c("14", "5", "9") +z <- qpAdm_prescreen(sim, right, left) +z + + + + + +# all gone +right <- c("10", "12", "13") +left <- c("14", "5", "9") +z <- qpAdm_prescreen(sim, right, left, Zcutoff = 3) +z + + + +right <- c("7", "10", "12", "13") +left <- c("14", "5", "9") +z <- qpAdm_prescreen(sim, right, left, Zcutoff = 3) +z + +x <- z$screening + + + + + + + + + + + + + + + + + + + + + +library(admixr) +data <- eigenstrat("snps") +target <- "14" +candidates <- c("1", "3", "4", "5", "8", "9", "12") +nsources <- 2 +ncores = 20 + +results <- qpAdm_rotation(data, target, nsources = 2, candidates, ncores = 30) + + +props <- bind_rows(lapply(proportions, function(i) { i$src1 <- names(i)[2]; i$src2 <- names(i)[3]; names(i)[2:3] <- c("source1", "source2"); names(i)[4:5] <- c("stderr_source1", "stderr_source2"); select(i, -starts_with("stderr"))})) + +filter(props, 0 < source1, source1 < 1, 0 < source2, source2 < 1) + + + + + diff --git a/man/f4ratio.Rd b/man/f4ratio.Rd index 2dbf6d4..fe9ffd8 100644 --- a/man/f4ratio.Rd +++ b/man/f4ratio.Rd @@ -9,9 +9,19 @@ \usage{ f4ratio(data, X, A, B, C, O, outdir = NULL, params = NULL) -d(data, W, X, Y, Z, outdir = NULL, f4mode = FALSE, params = NULL) - -f4(data, W, X, Y, Z, outdir = NULL, params = NULL) +d( + data, + W, + X, + Y, + Z, + quartets = NULL, + outdir = NULL, + f4mode = FALSE, + params = NULL +) + +f4(data, W, X, Y, Z, quartets, outdir = NULL, params = NULL) f3(data, A, B, C, outdir = NULL, inbreed = FALSE, params = NULL) } @@ -25,6 +35,8 @@ f3(data, A, B, C, outdir = NULL, inbreed = FALSE, params = NULL) \item{W, X, Y, Z, A, B, C, O}{Population names according to the nomenclature used in Patterson et al., 2012.} +\item{quartets}{List of character vectors (quartets of population/sample labels)} + \item{f4mode}{Calculate the f4 statistic instead of the D statistic.} \item{inbreed}{See README.3PopTest in ADMIXTOOLS for an explanation.} diff --git a/man/qpAdm_filter.Rd b/man/qpAdm_filter.Rd new file mode 100644 index 0000000..75fc147 --- /dev/null +++ b/man/qpAdm_filter.Rd @@ -0,0 +1,20 @@ +% Generated by roxygen2: do not edit by hand +% Please edit documentation in R/qpAdm.R +\name{qpAdm_filter} +\alias{qpAdm_filter} +\title{Filter qpAdm rotation results for only 'sensible' models} +\usage{ +qpAdm_filter(x, p) +} +\arguments{ +\item{x}{Output of a qpAdm_rotation() function} + +\item{p}{p-value cutoff} +} +\value{ +qpAdm_rotation object filtered down based on p-value +} +\description{ +Filter for p-value larger than a specified cuttof and admixture +proportions between 0 and 1. +} diff --git a/man/qpAdm_prescreen.Rd b/man/qpAdm_prescreen.Rd new file mode 100644 index 0000000..b1536ec --- /dev/null +++ b/man/qpAdm_prescreen.Rd @@ -0,0 +1,24 @@ +% Generated by roxygen2: do not edit by hand +% Please edit documentation in R/qpAdm.R +\name{qpAdm_prescreen} +\alias{qpAdm_prescreen} +\title{Perform a 'pre-screening' of the 'right' populations before +a qpAdm analysis based on Harney et al., 2020 (bioRxiv)} +\usage{ +qpAdm_prescreen(data, candidates, left, Zcutoff = 2) +} +\arguments{ +\item{data}{EIGENSTRAT dataset} + +\item{candidates}{Character vector with potential 'outgroup' populations} + +\item{left}{Character vector with target and source populations} +} +\value{ +Data frame of all combinations of f4(L_i, L_j; R_k, R_l), + ordered by Zscore of the f4 statistic +} +\description{ +Perform a 'pre-screening' of the 'right' populations before +a qpAdm analysis based on Harney et al., 2020 (bioRxiv) +} diff --git a/man/qpAdm_rotation.Rd b/man/qpAdm_rotation.Rd new file mode 100644 index 0000000..39a34c2 --- /dev/null +++ b/man/qpAdm_rotation.Rd @@ -0,0 +1,28 @@ +% Generated by roxygen2: do not edit by hand +% Please edit documentation in R/qpAdm.R +\name{qpAdm_rotation} +\alias{qpAdm_rotation} +\title{Fit qpAdm models based on the rotation strategy described in +Harney et al. 2020 (bioRxiv)} +\usage{ +qpAdm_rotation(data, target, candidates, nsources, ncores = 1) +} +\arguments{ +\item{data}{EIGENSTRAT dataset} + +\item{target}{Target population that is modeled as admixed} + +\item{candidates}{Potential candidates for sources and outgroups} + +\item{nsources}{Number of sources to pull from the candidates} + +\item{ncores}{Number of CPU cores to utilize for model fitting} +} +\value{ +qpAdm list with proportions, ranks and subsets elements (as + with a traditional qpAdm run) +} +\description{ +Fit qpAdm models based on the rotation strategy described in +Harney et al. 2020 (bioRxiv) +} From cd1ed96ae6cd53d5098a67a4cc6c7d8fd9ba395e Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 13:21:36 +0200 Subject: [PATCH 02/28] Fix a surprising issue with parsing qpWave log outputs (on only some runs) --- R/output_parsers.R | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/R/output_parsers.R b/R/output_parsers.R index 7ada10c..e7854af 100644 --- a/R/output_parsers.R +++ b/R/output_parsers.R @@ -171,7 +171,7 @@ read_qpWave <- function(log_lines, details = FALSE) { a_end <- c(test_pos[-c(1, 2)], which(stringr::str_detect(log_lines, "## end of run"))) test_df <- log_lines[test_pos + 1] %>% - stringr::str_replace_all(" *[a-z0-9]+: ", "") %>% + stringr::str_replace_all("[a-z0-9]+: ", "") %>% stringr::str_replace_all(" +", "\t") %>% paste0(collapse = "\n") %>% readr::read_tsv(col_names = c("rank", "df", "chisq", "tail", "dfdiff", From 980ef620d41a0555e13cb5911999b184723ad53c Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 14:26:23 +0200 Subject: [PATCH 03/28] Update qpAdm rotation code --- R/qpAdm.R | 10 ++++++---- man/qpAdm_filter.Rd | 5 +++-- 2 files changed, 9 insertions(+), 6 deletions(-) diff --git a/R/qpAdm.R b/R/qpAdm.R index a832270..9385a87 100644 --- a/R/qpAdm.R +++ b/R/qpAdm.R @@ -73,10 +73,11 @@ qpAdm_rotation <- function(data, target, candidates, nsources, ncores = 1) { names(result$proportions)[2:(1 + nsources)] <- paste0("prop", 1:nsources) names(result$proportions)[4:(3 + nsources)] <- paste0("stderr", 1:nsources) ## add source names as two new columns - result$proportions <- cbind(result$proportions, sources_df) + result$proportions <- cbind(result$proportions, sources_df) %>% + dplyr::mutate(outgroups = paste0(x$outgroups, collapse = " & ")) ## rearrange columns result$proportions <- dplyr::select( - result$proportions, target, names(x$sources), pvalue, + result$proportions, target, names(x$sources), outgroups, pvalue, dplyr::everything() ) @@ -127,10 +128,11 @@ qpAdm_rotation <- function(data, target, candidates, nsources, ncores = 1) { #' proportions between 0 and 1. #' #' @param x Output of a qpAdm_rotation() function -#' @param p p-value cutoff +#' @param p p-value cutoff (default 0: will only filter for sensible +#' admixture proportions) #' #' @return qpAdm_rotation object filtered down based on p-value -qpAdm_filter <- function(x, p) { +qpAdm_filter <- function(x, p = 0) { ## get positions of columns with estimated admixture proportions prop_columns <- stringr::str_which(names(x$proportions), "prop") diff --git a/man/qpAdm_filter.Rd b/man/qpAdm_filter.Rd index 75fc147..94a9ffc 100644 --- a/man/qpAdm_filter.Rd +++ b/man/qpAdm_filter.Rd @@ -4,12 +4,13 @@ \alias{qpAdm_filter} \title{Filter qpAdm rotation results for only 'sensible' models} \usage{ -qpAdm_filter(x, p) +qpAdm_filter(x, p = 0) } \arguments{ \item{x}{Output of a qpAdm_rotation() function} -\item{p}{p-value cutoff} +\item{p}{p-value cutoff (default 0: will only filter for sensible +admixture proportions)} } \value{ qpAdm_rotation object filtered down based on p-value From c7c96a5737e74b93e7bbf56034859c15f6fdf560 Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 15:24:59 +0200 Subject: [PATCH 04/28] Explore sources up to a specified limit --- R/qpAdm.R | 16 ++++++++++------ man/qpAdm_rotation.Rd | 4 ++-- 2 files changed, 12 insertions(+), 8 deletions(-) diff --git a/R/qpAdm.R b/R/qpAdm.R index 9385a87..75dd453 100644 --- a/R/qpAdm.R +++ b/R/qpAdm.R @@ -39,20 +39,23 @@ qpAdm_prescreen <- function(data, candidates, left, Zcutoff = 2) { #' @param data EIGENSTRAT dataset #' @param target Target population that is modeled as admixed #' @param candidates Potential candidates for sources and outgroups -#' @param nsources Number of sources to pull from the candidates +#' @param maxsources Maximum number of sources to model (2...maxsources) #' @param ncores Number of CPU cores to utilize for model fitting #' #' @return qpAdm list with proportions, ranks and subsets elements (as #' with a traditional qpAdm run) #' #' @export -qpAdm_rotation <- function(data, target, candidates, nsources, ncores = 1) { +qpAdm_rotation <- function(data, target, candidates, maxsources = 2, ncores = 1) { ## generate combinations of possible sources and outgroups - sources <- t(combn(candidates, nsources)) - sources_outgroups <- lapply(1:nrow(sources), function(i) { + sources <- unlist(lapply(2:maxsources, function(nsrc) { + srccomb <- t(combn(candidates, nsrc)) + lapply(1:nrow(srccomb), function(j) srccomb[j, ]) + }), recursive = FALSE) + sources_outgroups <- lapply(sources, function(src) { list( - sources = sources[i, ], - outgroups = setdiff(candidates, sources[i, ]) + sources = src, + outgroups = setdiff(candidates, src) ) }) @@ -63,6 +66,7 @@ qpAdm_rotation <- function(data, target, candidates, nsources, ncores = 1) { target = target, sources = x$sources, outgroups = x$outgroups ) + nsources <- length(x$sources) names(x$sources) <- paste0("source", 1:nsources) sources_df <- as.data.frame(t(as.matrix(x$sources))) diff --git a/man/qpAdm_rotation.Rd b/man/qpAdm_rotation.Rd index 39a34c2..3c225e2 100644 --- a/man/qpAdm_rotation.Rd +++ b/man/qpAdm_rotation.Rd @@ -5,7 +5,7 @@ \title{Fit qpAdm models based on the rotation strategy described in Harney et al. 2020 (bioRxiv)} \usage{ -qpAdm_rotation(data, target, candidates, nsources, ncores = 1) +qpAdm_rotation(data, target, candidates, maxsources = 2, ncores = 1) } \arguments{ \item{data}{EIGENSTRAT dataset} @@ -14,7 +14,7 @@ qpAdm_rotation(data, target, candidates, nsources, ncores = 1) \item{candidates}{Potential candidates for sources and outgroups} -\item{nsources}{Number of sources to pull from the candidates} +\item{maxsources}{Maximum number of sources to model (2...maxsources)} \item{ncores}{Number of CPU cores to utilize for model fitting} } From aa96d57e6ad4a2a2aa5c776130058b4b1acaadb7 Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 15:54:01 +0200 Subject: [PATCH 05/28] Check for type --- R/qpAdm.R | 11 +++++++++++ 1 file changed, 11 insertions(+) diff --git a/R/qpAdm.R b/R/qpAdm.R index 75dd453..a24a952 100644 --- a/R/qpAdm.R +++ b/R/qpAdm.R @@ -33,6 +33,15 @@ qpAdm_prescreen <- function(data, candidates, left, Zcutoff = 2) { list(outgroups = right, screening = result) } + +# Check that the provided object is of the required type +check_type <- function(x, type) { + if (!inherits(x, type)) { + stop(glue::glue("Object is not of the type {type}"), call. = FALSE) + } +} + + #' Fit qpAdm models based on the rotation strategy described in #' Harney et al. 2020 (bioRxiv) #' @@ -47,6 +56,8 @@ qpAdm_prescreen <- function(data, candidates, left, Zcutoff = 2) { #' #' @export qpAdm_rotation <- function(data, target, candidates, maxsources = 2, ncores = 1) { + check_type(data, "EIGENSTRAT") + ## generate combinations of possible sources and outgroups sources <- unlist(lapply(2:maxsources, function(nsrc) { srccomb <- t(combn(candidates, nsrc)) From 687252fd471943ee5ee62affc67ab199f28e5890 Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 15:55:00 +0200 Subject: [PATCH 06/28] Check for correct size of left/right sets --- R/wrappers.R | 5 +++++ 1 file changed, 5 insertions(+) diff --git a/R/wrappers.R b/R/wrappers.R index dc4c7d2..1be29a3 100644 --- a/R/wrappers.R +++ b/R/wrappers.R @@ -108,6 +108,11 @@ f3 <- function(data, A, B, C, outdir = NULL, inbreed = FALSE, params = NULL) { #' @export qpAdm <- function(data, target, sources, outgroups, outdir = NULL, params = list(allsnps = "YES", summary = "YES", details = "YES")) { + if (length(outgroups) < length(sources) + 1) { + stop("The number of outgroup samples has to be larger or equal than the number of sources + 1", + call. = FALSE) + } + check_presence(c(target, sources, outgroups), data) results <- lapply(target, function(X) { From 4b3a38dc509103290933977ee904aaf814b36cec Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 16:26:24 +0200 Subject: [PATCH 07/28] Revert back from exploring multiple sources --- R/qpAdm.R | 27 +++++++++++++++------------ man/qpAdm_rotation.Rd | 11 +++++++++-- 2 files changed, 24 insertions(+), 14 deletions(-) diff --git a/R/qpAdm.R b/R/qpAdm.R index a24a952..8c53d28 100644 --- a/R/qpAdm.R +++ b/R/qpAdm.R @@ -48,27 +48,31 @@ check_type <- function(x, type) { #' @param data EIGENSTRAT dataset #' @param target Target population that is modeled as admixed #' @param candidates Potential candidates for sources and outgroups -#' @param maxsources Maximum number of sources to model (2...maxsources) +#' @param nsources Number of sources to pull from the candidates #' @param ncores Number of CPU cores to utilize for model fitting #' #' @return qpAdm list with proportions, ranks and subsets elements (as #' with a traditional qpAdm run) #' #' @export -qpAdm_rotation <- function(data, target, candidates, maxsources = 2, ncores = 1) { +qpAdm_rotation <- function(data, target, candidates, minimize = FALSE, nsources = 2, ncores = 1) { check_type(data, "EIGENSTRAT") ## generate combinations of possible sources and outgroups - sources <- unlist(lapply(2:maxsources, function(nsrc) { - srccomb <- t(combn(candidates, nsrc)) - lapply(1:nrow(srccomb), function(j) srccomb[j, ]) + sources <- t(combn(candidates, nsources)) + sources_outgroups <- unlist(lapply(1:nrow(sources), function(i) { + outgroups <- setdiff(candidates, sources[i, ]) + if (minimize) { + outgroups <- unlist(lapply((nsources + 1):length(outgroups), function(nout) { + outcomb <- t(combn(outgroups, nout)) + lapply(1:nrow(outcomb), function(j) outcomb[j, ]) + }), recursive = FALSE) + } else { + outgroups <- list(outgroups) + } + + lapply(outgroups, function(out) { list(sources = sources[i, ], outgroups = out) }) }), recursive = FALSE) - sources_outgroups <- lapply(sources, function(src) { - list( - sources = src, - outgroups = setdiff(candidates, src) - ) - }) ## run qpAdm for all combinations of sources and outgroups results_list <- parallel::mclapply(sources_outgroups, function(x) { @@ -77,7 +81,6 @@ qpAdm_rotation <- function(data, target, candidates, maxsources = 2, ncores = 1) target = target, sources = x$sources, outgroups = x$outgroups ) - nsources <- length(x$sources) names(x$sources) <- paste0("source", 1:nsources) sources_df <- as.data.frame(t(as.matrix(x$sources))) diff --git a/man/qpAdm_rotation.Rd b/man/qpAdm_rotation.Rd index 3c225e2..cc9d3c3 100644 --- a/man/qpAdm_rotation.Rd +++ b/man/qpAdm_rotation.Rd @@ -5,7 +5,14 @@ \title{Fit qpAdm models based on the rotation strategy described in Harney et al. 2020 (bioRxiv)} \usage{ -qpAdm_rotation(data, target, candidates, maxsources = 2, ncores = 1) +qpAdm_rotation( + data, + target, + candidates, + minimize = FALSE, + nsources = 2, + ncores = 1 +) } \arguments{ \item{data}{EIGENSTRAT dataset} @@ -14,7 +21,7 @@ qpAdm_rotation(data, target, candidates, maxsources = 2, ncores = 1) \item{candidates}{Potential candidates for sources and outgroups} -\item{maxsources}{Maximum number of sources to model (2...maxsources)} +\item{nsources}{Number of sources to pull from the candidates} \item{ncores}{Number of CPU cores to utilize for model fitting} } From ca3e96fca7f4323c137274a3093827816e2a1577 Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 18:54:36 +0200 Subject: [PATCH 08/28] Update qpAdm rotation procedure --- R/log.R | 2 +- R/qpAdm.R | 67 ++++++++++++++++++++++++++----------------- R/utils.R | 9 ++++++ man/qpAdm_filter.Rd | 2 +- man/qpAdm_rotation.Rd | 13 +++++++-- 5 files changed, 61 insertions(+), 32 deletions(-) diff --git a/R/log.R b/R/log.R index e1b0fb7..445ac9c 100644 --- a/R/log.R +++ b/R/log.R @@ -73,7 +73,7 @@ loginfo <- function(x, target = NA, save = FALSE, prefix = NA, dir = ".", suffix #' #' @export print.admixr_result <- function(x, ...) { - if (attr(x, "command") %in% c("qpAdm", "qpAdm_rotation")) { + if (attr(x, "command") %in% c("qpAdm", "qpAdm_rotation") && length(x) == 3) { print.default(list( proportions = x$proportions, ranks = x$ranks, diff --git a/R/qpAdm.R b/R/qpAdm.R index 8c53d28..22578a0 100644 --- a/R/qpAdm.R +++ b/R/qpAdm.R @@ -34,13 +34,6 @@ qpAdm_prescreen <- function(data, candidates, left, Zcutoff = 2) { } -# Check that the provided object is of the required type -check_type <- function(x, type) { - if (!inherits(x, type)) { - stop(glue::glue("Object is not of the type {type}"), call. = FALSE) - } -} - #' Fit qpAdm models based on the rotation strategy described in #' Harney et al. 2020 (bioRxiv) @@ -48,14 +41,18 @@ check_type <- function(x, type) { #' @param data EIGENSTRAT dataset #' @param target Target population that is modeled as admixed #' @param candidates Potential candidates for sources and outgroups +#' @param minimize Test also all possible subsets of outgroups? (default TRUE) #' @param nsources Number of sources to pull from the candidates #' @param ncores Number of CPU cores to utilize for model fitting +#' @param fulloutput Report also 'ranks' and 'subsets' analysis from +#' qpAdm in addition to the admixture proportions results? (default FALSE) #' #' @return qpAdm list with proportions, ranks and subsets elements (as -#' with a traditional qpAdm run) +#' with a traditional qpAdm run) or just the proportions +#' (determined by the value of the 'fulloutput' argument) #' #' @export -qpAdm_rotation <- function(data, target, candidates, minimize = FALSE, nsources = 2, ncores = 1) { +qpAdm_rotation <- function(data, target, candidates, minimize = TRUE, nsources = 2, ncores = 1, fulloutput = FALSE) { check_type(data, "EIGENSTRAT") ## generate combinations of possible sources and outgroups @@ -92,10 +89,11 @@ qpAdm_rotation <- function(data, target, candidates, minimize = FALSE, nsources names(result$proportions)[4:(3 + nsources)] <- paste0("stderr", 1:nsources) ## add source names as two new columns result$proportions <- cbind(result$proportions, sources_df) %>% - dplyr::mutate(outgroups = paste0(x$outgroups, collapse = " & ")) + dplyr::mutate(outgroups = paste0(x$outgroups, collapse = " & "), + noutgroups = length(x$outgroups)) ## rearrange columns result$proportions <- dplyr::select( - result$proportions, target, names(x$sources), outgroups, pvalue, + result$proportions, target, names(x$sources), outgroups, noutgroups, pvalue, dplyr::everything() ) @@ -130,8 +128,12 @@ qpAdm_rotation <- function(data, target, candidates, minimize = FALSE, nsources subsets$model <- sort(rep(models, 1 + 2^(nsources - 1))) subsets <- dplyr::as_tibble(subsets) %>% dplyr::select(model, everything()) - ## bind all tables together and add log information - results <- list(proportions = proportions, ranks = ranks, subsets = subsets) + ## add metadata to the results object + if (fulloutput) + results <- list(proportions = proportions, ranks = ranks, subsets = subsets) + else + results <- proportions + attr(results, "command") <- "qpAdm_rotation" attr(results, "log_output") <- log_lines class(results) <- c("admixr_result", class(results)) @@ -150,24 +152,35 @@ qpAdm_rotation <- function(data, target, candidates, minimize = FALSE, nsources #' admixture proportions) #' #' @return qpAdm_rotation object filtered down based on p-value -qpAdm_filter <- function(x, p = 0) { +qpAdm_filter <- function(x, p = 0.05) { + check_type(x, "admixr_result") + if (length(x) == 3) + proportions <- x$proportions + else + proportions <- x + ## get positions of columns with estimated admixture proportions - prop_columns <- stringr::str_which(names(x$proportions), "prop") - + prop_columns <- stringr::str_which(names(proportions), "prop") + ## find out rows/models for which all proportions are in [0, 1] and ## pvalue is larger than the required cutoff - pvalue <- x$proportions$pvalue > p - constr_0 <- x$proportions[, prop_columns] >= 0 - constr_1 <- x$proportions[, prop_columns] <= 1 + pvalue <- proportions$pvalue > p + constr_0 <- proportions[, prop_columns] >= 0 + constr_1 <- proportions[, prop_columns] <= 1 constr <- apply(pvalue & constr_0 & constr_1, 1, all) ## filter all three sub-tables to only those models that fit the criteria - x$proportions <- dplyr::arrange(x$proportions[constr, ], -pvalue) - x$ranks <- x$ranks[x$ranks$model %in% x$proportions$model, ] - x$subsets <- x$subsets[x$subsets$model %in% x$proportions$model, ] - - ## filter also only to relevant remaining log output information - attr(x, "log_output") <- attr(x, "log_output")[unique(x$proportions$model)] - - x + proportions <- dplyr::arrange(proportions[constr, ], -pvalue) + + if (length(x) == 3) { + x$proportions <- proportions + x$ranks <- x$ranks[x$ranks$model %in% proportions$model, ] + x$subsets <- x$subsets[x$subsets$model %in% proportions$model, ] + ## filter also only to relevant remaining log output information + attr(x, "log_output") <- attr(x, "log_output")[unique(proportions$model)] + return(x) + } else { + attr(proportions, "log_output") <- attr(x, "log_output")[unique(proportions$model)] + return(proportions) + } } diff --git a/R/utils.R b/R/utils.R index ed34a8f..ff828c6 100644 --- a/R/utils.R +++ b/R/utils.R @@ -58,6 +58,15 @@ get_files <- function(dir_name, prefix) { +# Check that the provided object is of the required type +check_type <- function(x, type) { + if (!inherits(x, type)) { + stop(glue::glue("Object is not of the type {type}"), call. = FALSE) + } +} + + + # Check for the presence of a given set of labels in an 'ind' file. # Fail if there a sample was not found. check_presence <- function(labels, data) { diff --git a/man/qpAdm_filter.Rd b/man/qpAdm_filter.Rd index 94a9ffc..c57071c 100644 --- a/man/qpAdm_filter.Rd +++ b/man/qpAdm_filter.Rd @@ -4,7 +4,7 @@ \alias{qpAdm_filter} \title{Filter qpAdm rotation results for only 'sensible' models} \usage{ -qpAdm_filter(x, p = 0) +qpAdm_filter(x, p = 0.05) } \arguments{ \item{x}{Output of a qpAdm_rotation() function} diff --git a/man/qpAdm_rotation.Rd b/man/qpAdm_rotation.Rd index cc9d3c3..6b82a7f 100644 --- a/man/qpAdm_rotation.Rd +++ b/man/qpAdm_rotation.Rd @@ -9,9 +9,10 @@ qpAdm_rotation( data, target, candidates, - minimize = FALSE, + minimize = TRUE, nsources = 2, - ncores = 1 + ncores = 1, + fulloutput = FALSE ) } \arguments{ @@ -21,13 +22,19 @@ qpAdm_rotation( \item{candidates}{Potential candidates for sources and outgroups} +\item{minimize}{Test also all possible subsets of outgroups? (default TRUE)} + \item{nsources}{Number of sources to pull from the candidates} \item{ncores}{Number of CPU cores to utilize for model fitting} + +\item{fulloutput}{Report also 'ranks' and 'subsets' analysis from +qpAdm in addition to the admixture proportions results? (default FALSE)} } \value{ qpAdm list with proportions, ranks and subsets elements (as - with a traditional qpAdm run) + with a traditional qpAdm run) or just the proportions + (determined by the value of the 'fulloutput' argument) } \description{ Fit qpAdm models based on the rotation strategy described in From ffae416e12bdf776b2f4b18fdba771e9d6508bb9 Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 19:07:29 +0200 Subject: [PATCH 09/28] Add type check --- R/qpAdm.R | 5 +++++ Untitled.R | 43 +++++++++++++++++++++++++++++++++++++++---- 2 files changed, 44 insertions(+), 4 deletions(-) diff --git a/R/qpAdm.R b/R/qpAdm.R index 22578a0..fee9673 100644 --- a/R/qpAdm.R +++ b/R/qpAdm.R @@ -154,6 +154,11 @@ qpAdm_rotation <- function(data, target, candidates, minimize = TRUE, nsources = #' @return qpAdm_rotation object filtered down based on p-value qpAdm_filter <- function(x, p = 0.05) { check_type(x, "admixr_result") + if (attr(x, "command") != "qpAdm_rotation") { + stop("Filtering implemented only for results of the qpAdm rotation procedure", + call. = FALSE) + } + if (length(x) == 3) proportions <- x$proportions else diff --git a/Untitled.R b/Untitled.R index 26b6ec3..f928a77 100644 --- a/Untitled.R +++ b/Untitled.R @@ -89,13 +89,13 @@ x <- z$screening library(admixr) -data <- eigenstrat("snps") +data <- eigenstrat("all") target <- "14" -candidates <- c("1", "3", "4", "5", "8", "9", "12") +candidates <- c("1", "2", "3", "4", "5", "8", "9", "11", "12", "15") nsources <- 2 -ncores = 20 +ncores = 5 -results <- qpAdm_rotation(data, target, nsources = 2, candidates, ncores = 30) +results <- qpAdm_rotation(data, target, nsources = 2, candidates, ncores = ncores) props <- bind_rows(lapply(proportions, function(i) { i$src1 <- names(i)[2]; i$src2 <- names(i)[3]; names(i)[2:3] <- c("source1", "source2"); names(i)[4:5] <- c("stderr_source1", "stderr_source2"); select(i, -starts_with("stderr"))})) @@ -106,3 +106,38 @@ filter(props, 0 < source1, source1 < 1, 0 < source2, source2 < 1) + + + + + + + + +snps <- eigenstrat(download_data()) + +read_ind(snps) + +fits <- qpAdm_rotation( + snps, + target = "French", + candidates = c("Dinka", "Mbuti", "Yoruba", "Vindija", "Altai", "Denisova", "Chimp"), + minimize = TRUE, + nsources = 2, + ncores = 30 +) + +fits + +f <- qpAdm_filter(fits) + +x <- fits$proportions + +y <- qpAdm_filter(fits)$proportions %>% dplyr::select(-model, -starts_with("stderr"), -starts_with("nsnps")) + +x = qpAdm(snps, target = "French", sources = c( "Chimp", "Denisova"), outgroups = c("Mbuti", "Yoruba", "Altai", "Vindija")) + +x = qpAdm(snps, target = "French", sources = c( "Chimp", "Vindija"), outgroups = c("Mbuti", "Yoruba", "Altai", "Denisova")) + + + qpAdm(snps, target = "French", sources = c( "Chimp", "Vindija"), outgroups = c("Mbuti", "Yoruba", "Denisova")) From f60341bcd4e1bc287ee7039910df6f5155d23c2b Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 19:33:01 +0200 Subject: [PATCH 10/28] Remove testing script --- Untitled.R | 143 ----------------------------------------------------- 1 file changed, 143 deletions(-) delete mode 100644 Untitled.R diff --git a/Untitled.R b/Untitled.R deleted file mode 100644 index f928a77..0000000 --- a/Untitled.R +++ /dev/null @@ -1,143 +0,0 @@ -snps <- eigenstrat(download_data()) - -result <- qpAdm( - target = c("Han", "Sardinian"), - sources = c("Vindija", "Yoruba"), - outgroups = c("Chimp", "Denisova", "Altai"), - data = snps -) - - -left <- c("Han", "Vindija", "Yoruba") -right <- setdiff(setdiff(read_ind(snps)$label, left), - c("French", "Sardinian", "Papuan", "Dinka", "Khomani_San")) - -qpAdm_prescreen(snps, right, left) - - -qpAdm_prescreen(snps, c("Chimp", "Denisova"), left) - -left <- c("Sardinian", "Dinka", "French") -candidates <- c("Chimp", "Altai", "Denisova", "Yoruba", "Mbuti") - -x = qpAdm_prescreen(snps, candidates, left) -x -arrange(x$screening, abs(Zscore)) - - - - - - -sim <- eigenstrat("/tmp/sim/snps") - -Qpadm( - target = "14", - sources = c("5", "9"), - outgroups = c("0", "7", "10", "12", "13"), - data = sim -) - -right <- c("0", "7", "10", "12", "13") -left <- c("14", "5", "9") -y <- qpAdm_prescreen(sim, right, left) - - - -right <- c("0", "1", "2", "3", "4") -left <- c("14", "5", "9") -z <- qpAdm_prescreen(sim, right, left) -z - - - - - -# all gone -right <- c("10", "12", "13") -left <- c("14", "5", "9") -z <- qpAdm_prescreen(sim, right, left, Zcutoff = 3) -z - - - -right <- c("7", "10", "12", "13") -left <- c("14", "5", "9") -z <- qpAdm_prescreen(sim, right, left, Zcutoff = 3) -z - -x <- z$screening - - - - - - - - - - - - - - - - - - - - - -library(admixr) -data <- eigenstrat("all") -target <- "14" -candidates <- c("1", "2", "3", "4", "5", "8", "9", "11", "12", "15") -nsources <- 2 -ncores = 5 - -results <- qpAdm_rotation(data, target, nsources = 2, candidates, ncores = ncores) - - -props <- bind_rows(lapply(proportions, function(i) { i$src1 <- names(i)[2]; i$src2 <- names(i)[3]; names(i)[2:3] <- c("source1", "source2"); names(i)[4:5] <- c("stderr_source1", "stderr_source2"); select(i, -starts_with("stderr"))})) - -filter(props, 0 < source1, source1 < 1, 0 < source2, source2 < 1) - - - - - - - - - - - - - -snps <- eigenstrat(download_data()) - -read_ind(snps) - -fits <- qpAdm_rotation( - snps, - target = "French", - candidates = c("Dinka", "Mbuti", "Yoruba", "Vindija", "Altai", "Denisova", "Chimp"), - minimize = TRUE, - nsources = 2, - ncores = 30 -) - -fits - -f <- qpAdm_filter(fits) - -x <- fits$proportions - -y <- qpAdm_filter(fits)$proportions %>% dplyr::select(-model, -starts_with("stderr"), -starts_with("nsnps")) - -x = qpAdm(snps, target = "French", sources = c( "Chimp", "Denisova"), outgroups = c("Mbuti", "Yoruba", "Altai", "Vindija")) - -x = qpAdm(snps, target = "French", sources = c( "Chimp", "Vindija"), outgroups = c("Mbuti", "Yoruba", "Altai", "Denisova")) - - - qpAdm(snps, target = "French", sources = c( "Chimp", "Vindija"), outgroups = c("Mbuti", "Yoruba", "Denisova")) From 0bf8a75b3bb9427a607ce10e68d5d8911a985d4c Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Thu, 18 Jun 2020 23:18:16 +0200 Subject: [PATCH 11/28] Add new vignette --- R/qpAdm.R | 2 + vignettes/qpAdm.Rmd | 297 ++++++++++++++++++++++++++++++++++++++++++++ 2 files changed, 299 insertions(+) create mode 100644 vignettes/qpAdm.Rmd diff --git a/R/qpAdm.R b/R/qpAdm.R index fee9673..9af2be4 100644 --- a/R/qpAdm.R +++ b/R/qpAdm.R @@ -152,6 +152,8 @@ qpAdm_rotation <- function(data, target, candidates, minimize = TRUE, nsources = #' admixture proportions) #' #' @return qpAdm_rotation object filtered down based on p-value +#' +#' @export qpAdm_filter <- function(x, p = 0.05) { check_type(x, "admixr_result") if (attr(x, "command") != "qpAdm_rotation") { diff --git a/vignettes/qpAdm.Rmd b/vignettes/qpAdm.Rmd new file mode 100644 index 0000000..05c7387 --- /dev/null +++ b/vignettes/qpAdm.Rmd @@ -0,0 +1,297 @@ +--- +title: "Fitting qpAdm models with a 'rotation method'" +author: "Martin Petr" +date: "`r Sys.Date()`" +output: rmarkdown::html_vignette +vignette: > + %\VignetteIndexEntry{Vignette Title} + %\VignetteEngine{knitr::rmarkdown} + %\VignetteEncoding{UTF-8} +--- + +```{r setup, include = FALSE} +knitr::opts_chunk$set( + collapse = TRUE, + comment = "#>" +) +``` + +## Introduction + +qpAdm model fitting is a very complex topic which relies on a deep +knowledge of the concepts behind $f$-statistics introduced by Nick +Patterson and colleagues [in +2012](https://www.genetics.org/content/192/3/1065). In our +[tutorial](https://bodkan.net/admixr/articles/tutorial.html#qpwave-and-qpadm-1) +we have looked at the very basic overview of the qpWave/qpAdm-related +functionality implemented in _admixr_, without going beyond +description of functions and function arguments. I have also stressed +how important it is to learn _how_ does qpAdm works, linking to +several important references. + +Recently, an exciting new preprint was published by Harney _et al._, +[Assessing the Performance of qpAdm: A Statistical Tool for Studying +Population +Admixture](https://www.biorxiv.org/content/10.1101/2020.04.09.032664v1). Before +we go any further, I encourage everyone to read it and also read the +superb tutorial/guide available in a [supplementary +pdf](https://www.biorxiv.org/content/10.1101/2020.04.09.032664v1.supplementary-material) +on bioRxiv. There really isn't a better source at the moment on how to +run and interpret qpAdm analyses and about the pitfals that one needs +to be aware of. + +Please, only attempt to run qpAdm if you have familiarized yoursef +with all of the above-mentioned resources. I have had many people ask +questions via email (not only about qpAdm but also other topics) to +which the only sensible answer was - "you have to read the papers and +understand the statistics". + +## _qpAdm_ "rotation scheme" + +If you have ever worked with _qpAdm_, you are well aware of the +intricacies of finding the most suitable set of models that can +explain the data. Among other things, we have need to make a decision +about the number of admixture sources we want to estimate ancestry +proportions for, which populations are the most appropriate surrogates +for those source populations because only rarely we have sampled them +directly. Furthermore, we need to carefully choose a number of so +called 'outgroup' populations (also called 'references' or 'right' +populations, depending whom you ask). This is all, of course, in +addition to other technical decisions we need to make in terms of +preparing our dataset. + +The preprint by [Harney _et +al._](https://www.biorxiv.org/content/10.1101/2020.04.09.032664v1) +described an interesting idea to find a set of the most appropriate +models (sets of source and outgroup populations) - the so called +"rotating" population, already successfully used in the past (see +references in the preprint). + +Briefly, this strategy involves defining a set of "candidate" +populations, from which we iteratively sample a defined number of +"sources" (most commonly two or three) for our "target" population of +interest. We then take as "outgroups" all candidate populations except +for these sources. Doing so, we iteratively fit qpAdm models for each +combination of target, sources and outgroups, extracting $p$-values +and other statistics of interest. After finishing the exhaustive +exploration of source-outgroup combinations, we examine all fitted +models, selecting those that seem most appropriate. + +## Implementation in _admixr_ + +In _admixr_, I have implemented a function `qpAdm_rotation()` which +does exactly what I described in the previous paragraph with one +additional feature. Given the sensitivity of _qpAdm_ to large numbers +of potential outgroups (references), we also explore, for each +combination of sources and outgroups, also models with all possible +_subsets_ of outgroups. This is to allow finding models which are as +small as possible, and also to determine which outgroups are +potentially redundant. + +In other words, if we have a target _T_ and a set of candidates +(potential sources and outgroups) _C_ = {a, b, c, d, e, f}. Then, if +we imagine an iteration of the rotation scheme in which we fix sources +_S_ = {a, b}, we have remaining candidates _C - S_ = {c, d, e, f}. The +basic implementation of the rotation procedure would simly take _C - +S_ as the set of outgroups and fitted a single model: + +- model #1: _T_, _S_ = {a, b} and outgroups = {c, d, e, f} + +In _admixr_, we would, in this situation, evaluate the following +models: + +- model #2: _T_, _S_ = {a, b} and outgroups = {c, d, e} +- model #3: _T_, _S_ = {a, b} and outgroups = {c, d, f} +- model #4: _T_, _S_ = {a, b} and outgroups = {c, e, f} +- model #5: _T_, _S_ = {a, b} and outgroups = {d, e, f}. + +This makes it possible to find the _smallest_ model (in terms of +outgroup size) that can explain our data. + +## Concrete example + +### Performing exhaustive search by rotating sources/outgroups + +As an example, let's revisit our boring example of estimating the +level of Neandertal ancestry in a French person. We use it (and not +some other more interesting example from a vast human population +history) because: + +1. It's probably the simplest possible $f$-statistics-related analysis + one could do. +2. It gives us a clear expectation of what the "truth" is. +3. It gives us a clear expectation of what models we should + _definitely_ reject. +4. It's the only thing I know how to do. + +Let's first download a small example data set: + +```{r} +library(admixr) + +snps <- eigenstrat(download_data()) +``` + +These are the individuals for which we have genotype data: + +```{r} +read_ind(snps) +``` + +The `qpAdm_rotation()` function is very simple. It accepts: + +- the name of the target population, +- the list of candidate populations, +- a logical parameter `minimize` determining whether to perform the + "minimization" of the outgroup size described in the previous + section, +- the number of sources of ancestry +- the number of CPU cores to use for an analysis (be careful with this + options as many ADMIXTOOLS analyses run in paralle often consume _a + lot_ of memory!), +- parameter `fulloutput` specifying whether we want to have all the + "ranks" and "subsets/patterns" statistics (see the tutorial for more + information) or if we just want the proportions of ancestry and + significance values for individual models (this is the default). + +Let's say we are interested in finding the proportions of archaic +human ancestry in a French individual, and also in seeing what sorts +of possible models we could find that match archaic introgression: + +```{r} +models <- qpAdm_rotation( + data = snps, + target = "French", + candidates = c("Dinka", "Mbuti", "Yoruba", "Vindija", "Altai", "Denisova", "Chimp"), + minimize = TRUE, + nsources = 2, + ncores = 30, + fulloutput = TRUE +) +``` + +On my computing cluster this takes only a couple of minutes but note +that I'm utilizing 30 CPU cores in parallel. On your computer it might +take a bit more time. If it takes _too_ much time, you might want to +restrict the set of potential candidates or turn of the minimization +option. + +Here is what the full output looks like: + +```{r} +models +``` + +We can see that we got a list with three components, as we would +expect from any other `qpAdm()` run (see the manual page and the +tutorial for description of all three elements and their meaning). The +first column is named `model` - this is just a short identifier of +each individual "rotation" run. It's values don't have any particular +meaning, but are useful for later filtering and examination. + +Let's ignore the `$ranks` and `$subsets` elements, these are not that +useful in the first step of model selection. We will focus only on the +first element, `$proportions`. + + +### Examining and filtering fitted models + +The output data.frame contains information about *all* objects, +regardless of their plausibility. We can see that by examining the +distributions of p-values (column `pvalue`) and admixture proportions +(columns `prop1` and `prop2`) of each evaluated model. + +Notice two things: + +- Many models have inferred admixture proportions _way_ outside the + [0, 1] interval - those are clearly nonsensical. +- Many models have very low p-values - knowing how qpAdm internally + operates, you will know this means these are incompatible with the + data and can be rejected. + +```{r} +library(tidyverse) + +select(models$proportions, model, pvalue, prop1, prop2) %>% + gather(parameter, value, -model) %>% + ggplot(aes(parameter, value)) + + geom_jitter() + + facet_wrap(~ parameter, scales = "free_y") +``` + +The _admixr_ package contains a simple function `qpAdm_filter()` which +accepts the object produced by the `qpAdm_rotation()` function (either +the `fulloutput` version or the simple) and removes models with any +proportion outside of the [0, 1] range and also models with p-values +lower than a specified cutoff (0.05 by default): + +```{r} +fits <- qpAdm_filter(models) +``` + +As an aside, note that you can use the function `loginfo()` to examine +the complete log output of any model by specifying the model number +(this is why each row has the model identifier). Like this (output +omitted for brevity): + +```{r, eval = FALSE} +loginfo(fits, "m81") +``` + +We can verify the filtering worked by examining the filtered set of +models. Again, note that the p-values are now nicely distributed +across the range of "insigificance" (i.e., "non-rejection") range of +[0.05, 1.0]. And, also (remember that we tried to find a set of +sources-outgroups combinations that model archaic ancestry in a French +individual?) we see two nice clusters of ancestry proportions - one +very small (which what we will see is a Neandertal component) and one +large ("modern human" component, non-Neandertal ancestry): + +```{r} +select(fits$proportions, model, pvalue, prop1, prop2) %>% + gather(parameter, value, -model) %>% + ggplot(aes(parameter, value)) + + geom_jitter() + + facet_wrap(~ parameter) + + coord_cartesian(y = c(0, 1)) +``` + +Let's now subset only to the proportions table which is most relevant +at this stage anyway. We will also ignore a couple of columns for +brevity (*note* that we are also ignoring p-values because we *cannot* +used those for model selection, they are *not* meaningful at this +stage) and order the models based on the size of the outgroup set. + +```{r} +props <- fits$proportions %>% + arrange(noutgroups) %>% + select(-c(target, noutgroups, stderr1, stderr2, nsnps_used, nsnps_target)) + +print(props, n = Inf) +``` + +Also, before you run away terrified by the huge number of "compatible" +models, please note that we have _deliberately_ added too many +candidate populations and so you might reasonably expect many of the +evaluated models are not really optimal. + +Case in point: notice that there are many models in which the +chimpanzee was fitted as a source of ancestry! Interestingly, qpAdm +used it to "sucessfully" infer archaic human ancestry (where we define +"successful model" by inferring ancestry proportions between 0 and 1, +and a high p-value). This is because Neandertal ancestry could be +considered an "ancestral component" of a modern human genome: + +```{r} +filter(props, source1 == "Chimp" | source2 == "Chimp") +``` + +## Conclusions + +At this stage of analysis you would be on your own. You would have to +decide which models are more reasonably and why, possibly based on +prior knowledge or additional statistics (such as the details +information reported in the full log output information). This is +because qpAdm on some level is as much art as it is science and +interpreting the results and finding the most appropriate models can +be quite a challenge. From ece24b3b2752aeeaca42e1ef1e5525d772aa930d Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Fri, 19 Jun 2020 15:20:35 +0200 Subject: [PATCH 12/28] Update NAMESPACE --- NAMESPACE | 1 + 1 file changed, 1 insertion(+) diff --git a/NAMESPACE b/NAMESPACE index 3270cd7..ed98458 100644 --- a/NAMESPACE +++ b/NAMESPACE @@ -15,6 +15,7 @@ export(keep_transitions) export(loginfo) export(merge_eigenstrat) export(qpAdm) +export(qpAdm_filter) export(qpAdm_prescreen) export(qpAdm_rotation) export(qpWave) From 190cfb11f6a2bdcde2ad88717afe4f4606c60695 Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Fri, 19 Jun 2020 16:04:14 +0200 Subject: [PATCH 13/28] Remove the pre-screening function for now --- NAMESPACE | 1 - R/qpAdm.R | 37 ------------------------------------- man/qpAdm_prescreen.Rd | 24 ------------------------ 3 files changed, 62 deletions(-) delete mode 100644 man/qpAdm_prescreen.Rd diff --git a/NAMESPACE b/NAMESPACE index ed98458..7869c1d 100644 --- a/NAMESPACE +++ b/NAMESPACE @@ -16,7 +16,6 @@ export(loginfo) export(merge_eigenstrat) export(qpAdm) export(qpAdm_filter) -export(qpAdm_prescreen) export(qpAdm_rotation) export(qpWave) export(read_geno) diff --git a/R/qpAdm.R b/R/qpAdm.R index 9af2be4..ba0b3c8 100644 --- a/R/qpAdm.R +++ b/R/qpAdm.R @@ -1,40 +1,3 @@ -#' Perform a 'pre-screening' of the 'right' populations before -#' a qpAdm analysis based on Harney et al., 2020 (bioRxiv) -#' -#' @param data EIGENSTRAT dataset -#' @param candidates Character vector with potential 'outgroup' populations -#' @param left Character vector with target and source populations -#' -#' @return Data frame of all combinations of f4(L_i, L_j; R_k, R_l), -#' ordered by Zscore of the f4 statistic -#' -#' @export -qpAdm_prescreen <- function(data, candidates, left, Zcutoff = 2) { - leftcomb <- t(combn(left, 2)) - rightcomb <- t(combn(candidates, 2)) - - i <- 1 - quartets <- list() - for (l in 1:nrow(leftcomb)) { - for (r in 1:nrow(rightcomb)) { - quartets[[i]] <- c(leftcomb[l, ], rightcomb[r, ]) - i <- i + 1 - } - } - - result <- f4(data, quartets = quartets) %>% dplyr::arrange(abs(Zscore)) - - right <- dplyr::filter(result, abs(Zscore) > Zcutoff) %>% - .[, c("Y", "Z")] %>% - as.matrix %>% - as.vector %>% - unique - - list(outgroups = right, screening = result) -} - - - #' Fit qpAdm models based on the rotation strategy described in #' Harney et al. 2020 (bioRxiv) #' diff --git a/man/qpAdm_prescreen.Rd b/man/qpAdm_prescreen.Rd deleted file mode 100644 index b1536ec..0000000 --- a/man/qpAdm_prescreen.Rd +++ /dev/null @@ -1,24 +0,0 @@ -% Generated by roxygen2: do not edit by hand -% Please edit documentation in R/qpAdm.R -\name{qpAdm_prescreen} -\alias{qpAdm_prescreen} -\title{Perform a 'pre-screening' of the 'right' populations before -a qpAdm analysis based on Harney et al., 2020 (bioRxiv)} -\usage{ -qpAdm_prescreen(data, candidates, left, Zcutoff = 2) -} -\arguments{ -\item{data}{EIGENSTRAT dataset} - -\item{candidates}{Character vector with potential 'outgroup' populations} - -\item{left}{Character vector with target and source populations} -} -\value{ -Data frame of all combinations of f4(L_i, L_j; R_k, R_l), - ordered by Zscore of the f4 statistic -} -\description{ -Perform a 'pre-screening' of the 'right' populations before -a qpAdm analysis based on Harney et al., 2020 (bioRxiv) -} From b76cb2e2cf193bb7cc63f165545b4ffc6fad7333 Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Fri, 19 Jun 2020 17:12:25 +0200 Subject: [PATCH 14/28] Fix d() function population checking --- R/wrappers.R | 2 +- 1 file changed, 1 insertion(+), 1 deletion(-) diff --git a/R/wrappers.R b/R/wrappers.R index 1be29a3..be4f505 100644 --- a/R/wrappers.R +++ b/R/wrappers.R @@ -30,7 +30,7 @@ f4ratio <- function(data, X, A, B, C, O, outdir = NULL, params = NULL) { #' @export d <- function(data, W, X, Y, Z, quartets = NULL, outdir = NULL, f4mode = FALSE, params = NULL) { if (is.null(quartets)) { - check_presence(c(W, X, Y, Z, data)) + check_presence(c(W, X, Y, Z), data) } else { check_presence(unique(unlist(quartets)), data) } From 69ea61979611478cf4affc7772998697afe75259 Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Fri, 19 Jun 2020 18:13:49 +0200 Subject: [PATCH 15/28] Add default "quartets" value --- R/wrappers.R | 2 +- man/f4ratio.Rd | 2 +- 2 files changed, 2 insertions(+), 2 deletions(-) diff --git a/R/wrappers.R b/R/wrappers.R index be4f505..2374a17 100644 --- a/R/wrappers.R +++ b/R/wrappers.R @@ -63,7 +63,7 @@ d <- function(data, W, X, Y, Z, quartets = NULL, outdir = NULL, f4mode = FALSE, #' @rdname f4ratio #' #' @export -f4 <- function(data, W, X, Y, Z, quartets, outdir = NULL, params = NULL) { +f4 <- function(data, W, X, Y, Z, quartets = NULL, outdir = NULL, params = NULL) { d(data, W, X, Y, Z, quartets, outdir, f4mode = TRUE) } diff --git a/man/f4ratio.Rd b/man/f4ratio.Rd index fe9ffd8..536c236 100644 --- a/man/f4ratio.Rd +++ b/man/f4ratio.Rd @@ -21,7 +21,7 @@ d( params = NULL ) -f4(data, W, X, Y, Z, quartets, outdir = NULL, params = NULL) +f4(data, W, X, Y, Z, quartets = NULL, outdir = NULL, params = NULL) f3(data, A, B, C, outdir = NULL, inbreed = FALSE, params = NULL) } From 05e843441c1cdb83265528bcd70f60b826bceb5b Mon Sep 17 00:00:00 2001 From: Martin Petr Date: Fri, 19 Jun 2020 21:26:44 +0200 Subject: [PATCH 16/28] Add new qpAdm_rotation tutorial --- docs/LICENSE-text.html | 5 +- docs/LICENSE.html | 5 +- docs/articles/index.html | 9 +- .../figure-html/unnamed-chunk-5-1.png | Bin 0 -> 104250 bytes .../figure-html/unnamed-chunk-7-1.png | Bin 0 -> 75148 bytes .../figure-html/unnamed-chunk-8-1.png | Bin 0 -> 76195 bytes .../header-attrs-2.2/header-attrs.js | 12 + docs/articles/tutorial.html | 79 ++-- docs/authors.html | 5 +- docs/index.html | 5 +- docs/news/index.html | 5 +- docs/pkgdown.yml | 3 +- docs/reference/count_snps.html | 5 +- docs/reference/download_data.html | 5 +- docs/reference/eigenstrat.html | 5 +- docs/reference/f4ratio.html | 23 +- docs/reference/filter_bed.html | 5 +- docs/reference/index.html | 5 +- docs/reference/merge_eigenstrat.html | 5 +- docs/reference/pipe.html | 5 +- docs/reference/print.EIGENSTRAT.html | 5 +- docs/reference/qpAdm.html | 5 +- docs/reference/qpWave.html | 5 +- docs/reference/read_ind.html | 5 +- docs/reference/relabel.html | 5 +- docs/reference/reset.html | 5 +- docs/reference/write_ind.html | 5 +- vignettes/qpAdm.Rmd | 363 ++++++++++-------- vignettes/tutorial.Rmd | 27 +- 29 files changed, 375 insertions(+), 236 deletions(-) create mode 100644 docs/articles/qpAdm_files/figure-html/unnamed-chunk-5-1.png create mode 100644 docs/articles/qpAdm_files/figure-html/unnamed-chunk-7-1.png create mode 100644 docs/articles/qpAdm_files/figure-html/unnamed-chunk-8-1.png create mode 100644 docs/articles/qpAdm_files/header-attrs-2.2/header-attrs.js diff --git a/docs/LICENSE-text.html b/docs/LICENSE-text.html index 5fede6e..29bfa18 100644 --- a/docs/LICENSE-text.html +++ b/docs/LICENSE-text.html @@ -94,7 +94,10 @@ diff --git a/docs/LICENSE.html b/docs/LICENSE.html index 6ad743b..0d065ab 100644 --- a/docs/LICENSE.html +++ b/docs/LICENSE.html @@ -94,7 +94,10 @@ diff --git a/docs/articles/index.html b/docs/articles/index.html index 61f71e4..a97b6fe 100644 --- a/docs/articles/index.html +++ b/docs/articles/index.html @@ -94,7 +94,10 @@ @@ -130,7 +133,9 @@

All vignettes

-
admixr - Tutorial
+
Fitting qpAdm models with a 'rotation' strategy
+
+
Tutorial and basic overview of the admixr R package
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dr-!q!zo_FRIs>yhLID5BNh?a_UN!Xpe*gguA-n(p literal 0 HcmV?d00001 diff --git a/docs/articles/qpAdm_files/header-attrs-2.2/header-attrs.js b/docs/articles/qpAdm_files/header-attrs-2.2/header-attrs.js new file mode 100644 index 0000000..dd57d92 --- /dev/null +++ b/docs/articles/qpAdm_files/header-attrs-2.2/header-attrs.js @@ -0,0 +1,12 @@ +// Pandoc 2.9 adds attributes on both header and div. We remove the former (to +// be compatible with the behavior of Pandoc < 2.8). +document.addEventListener('DOMContentLoaded', function(e) { + var hs = document.querySelectorAll("div.section[class*='level'] > :first-child"); + var i, h, a; + for (i = 0; i < hs.length; i++) { + h = hs[i]; + if (!/^h[1-6]$/i.test(h.tagName)) continue; // it should be a header h1-h6 + a = h.attributes; + while (a.length > 0) h.removeAttribute(a[0].name); + } +}); diff --git a/docs/articles/tutorial.html b/docs/articles/tutorial.html index cc2bbb4..439243c 100644 --- a/docs/articles/tutorial.html +++ b/docs/articles/tutorial.html @@ -5,13 +5,13 @@ -admixr - Tutorial • admixr +Tutorial and basic overview of the admixr R package • admixr - +