tigger/0000755000176200001440000000000013514253663011543 5ustar liggesuserstigger/NAMESPACE0000644000176200001440000000450713514251352012761 0ustar liggesusers# Generated by roxygen2: do not edit by hand export(cleanSeqs) export(findNovelAlleles) export(findUnmutatedCalls) export(generateEvidence) export(genotypeFasta) export(getMutCount) export(getMutatedPositions) export(getPopularMutationCount) export(inferGenotype) export(inferGenotypeBayesian) export(insertPolymorphisms) export(plotGenotype) export(plotNovel) export(readIgFasta) export(reassignAlleles) export(selectNovel) export(sortAlleles) export(updateAlleleNames) export(writeFasta) import(ggplot2) importFrom(alakazam,DNA_COLORS) importFrom(alakazam,checkColumns) importFrom(alakazam,getAllele) importFrom(alakazam,getFamily) importFrom(alakazam,getGene) importFrom(alakazam,translateDNA) importFrom(doParallel,registerDoParallel) importFrom(dplyr,"%>%") importFrom(dplyr,arrange) importFrom(dplyr,as_data_frame) importFrom(dplyr,bind_cols) importFrom(dplyr,bind_rows) importFrom(dplyr,combine) importFrom(dplyr,data_frame) importFrom(dplyr,desc) importFrom(dplyr,distinct) importFrom(dplyr,do) importFrom(dplyr,filter) importFrom(dplyr,glimpse) importFrom(dplyr,group_by) importFrom(dplyr,group_indices) importFrom(dplyr,inner_join) importFrom(dplyr,mutate) importFrom(dplyr,n) importFrom(dplyr,rename) importFrom(dplyr,select) importFrom(dplyr,slice) importFrom(dplyr,summarise) importFrom(dplyr,transmute) importFrom(dplyr,ungroup) importFrom(foreach,"%dopar%") importFrom(foreach,foreach) importFrom(foreach,registerDoSEQ) importFrom(graphics,plot) importFrom(gridExtra,arrangeGrob) importFrom(gtools,ddirichlet) importFrom(iterators,icount) importFrom(lazyeval,interp) importFrom(parallel,clusterEvalQ) importFrom(parallel,clusterExport) importFrom(parallel,makeCluster) importFrom(parallel,stopCluster) importFrom(rlang,":=") importFrom(rlang,.data) importFrom(rlang,sym) importFrom(rlang,syms) importFrom(shazam,calcObservedMutations) importFrom(stats,confint) importFrom(stats,cor) importFrom(stats,cov) importFrom(stats,ecdf) importFrom(stats,lm) importFrom(stats,mad) importFrom(stats,median) importFrom(stats,na.omit) importFrom(stats,sd) importFrom(stats,setNames) importFrom(stringi,"stri_sub<-") importFrom(stringi,stri_detect_fixed) importFrom(stringi,stri_length) importFrom(stringi,stri_replace_all_regex) importFrom(stringi,stri_sub) importFrom(stringi,stri_trans_toupper) importFrom(tidyr,gather) importFrom(tidyr,spread) importFrom(tidyr,unnest) tigger/README.md0000644000176200001440000000364313402556525013027 0ustar liggesusers# TIgGER # High-throughput sequencing of B cell immunoglobulin receptors is providing unprecedented insight into adaptive immunity. A key step in analyzing these data involves assignment of the germline V, D and J gene segment alleles that comprise each immunoglobulin sequence by matching them against a database of known V(D)J alleles. However, this process will fail for sequences that utilize previously undetected alleles, whose frequency in the population is unclear. TIgGER is a computational method that significantly improves V(D)J allele assignments by first determining the complete set of gene segments carried by an individual (including novel alleles) from V(D)J-rearrange sequences. TIgGER can then infer a subject's genotype from these sequences, and use this genotype to correct the initial V(D)J allele assignments. The application of TIgGER continues to identify a surprisingly high frequency of novel alleles in humans, highlighting the critical need for this approach. (TIgGER, however, can and has been used with data from other species.) ## Core Abilities ## * Detecting novel alleles * Inferring a subject's genotype * Correcting preliminary allele calls ## Required Input ## * A table of sequences from a single individual, with columns containing the following: * V(D)J-rearranged nucleotide sequence (in IMGT-gapped format) * Preliminary V allele calls * Preliminary J allele calls * Length of the junction region * Germline Ig sequences in IMGT-gapped fasta format (e.g., as those downloaded from [IMGT/GENE-DB](http://www.imgt.org/genedb)) The former can be created through the use of [IMGT/HighV-QUEST](http://www.imgt.org) and [Change-O](http://changeo.readthedocs.io). ## Contact ## For help, questions, or suggestions, please contact the [Immcantation Group](mailto:immcantation@googlegroups.com) or use the [issue tracker](https://bitbucket.org/kleinstein/tigger/issues?status=new&status=open). tigger/data/0000755000176200001440000000000013514251417012447 5ustar liggesuserstigger/data/datalist0000644000176200001440000000011313514251417014172 0ustar liggesusersGermlineIGHV SampleDb SampleGenotype SampleGermlineIGHV SampleNovel airrDb tigger/data/SampleGenotype.rda0000644000176200001440000000053313402556525016100 0ustar liggesusersBZh91AY&SY.ܱ|H.@ܠ@86zLhh~ji&LF #A"4'hĂ+[ E`44|\mkPVx@B 9RfR!>j'1h)Z'D'c،-B\I!4`ei@#Fd]5S X$Ӫ$V5a!"+F#n1pB8H$T@g\Ni'PdB鏕{D!&bkI+aL x"``,&%LGR!R bu )„vHtigger/data/SampleDb.rda0000644000176200001440000472161313402556525014650 0ustar liggesusersBZh91AY&SY~pL?c?%)K϶}E)M4>GOAlo:}_yAi }xt+^@})_w(=}x@iϭhJGϟ* fjQ+6@[Y '&Emm4w:^iFA/3|/)<})@>K)lɼ)J)E>wSPP4|ټ>)2kQxz9U6 1SԍҔi@h @h .  ^ P(@a4h >k@JoJ廊NC}﫦ylT9UhiZ|8VJw><͢lٶ5FYkHT" V((zhеK}_-}i[U6[ϢZ {TeT|}{m[{`>OOeյ{iUt֭}Um55;ϪLj5}ޯkE*>BץSM*mU 6T*Mm4%ϒfDJ}0yU*΃q+V_ MkӶժn:{S7l}=>Z\mSjlU_{H^{|Uf(JH2`5JFAFzƋ>U7c5\򍼵Uj{wU.Y7umZmmtλs{ZRלRW٘K2S-͠ mK_@Uj/elko+hI;͏UT'cv]Ϸ>=W}޽O}=7;}GeRn<"m[SٛhPSҏymX{̍Ms MTy{[ö;cR-5IZkmIQ!4}ھtR t. 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MpL%Z9s2ajڠ0lIuTY+zqƚ걪Fte:w^pHڞh6ڮcn2%u #AB q؟g;]Ҩ@.1FYnB($YHW We2+J'+D!3q#F A@N,.Z ms#-ѢPHr+_?@WB@ϯqI@HK+RZ1wRdׄmָ;ܧ?mfQQukEŪ6i6KF3N:O)Ǟ2ɒ'J|޷Կ2O&{{fVOe2C0)C&Y= s*c$|ca4mq: lV1c.0^^oY:EȦn'B/;Nȟ+0LKC$i gT+"{'1Lca\ˤda&Kj&f2ʉfR7p,C!v`/A<K>e4UTESTg> >* L=TRymfmꝑJ̙S?h$_d~7")N vԯ8ۭM=2N\: * A3$ ]pz$h\rPׁJǃʡH 7!G*35cn\j4ndHfH%!hSk6z0}xoUE43$ijnQ9dL G]T9ZLR{tk/F73j ;A> =1!8!iH81C>Y'|;)uis)R '!>hJB)D>]A֪`b)C<,HJ **'$j%ʫlV7o۳;lߩ6fYCᾫ U`D[7ky^㑃-°Lwpr閵5q#8ȹss2SZVbcMd׷3wV<^m'&ؗ{SGz23A rwB3[%G3/+jzҵ3N4ѵGY'2&bް152k/\e֫M/7//1O`3:KV'n84倚N$ j̛-iYc }>_ϨO_]1/DdǻZNGHUdj&Yʇj%N^ ( "86s:pcR7=c޼ ƺ^|lp)BGAa5Y2 )|OQ6=C=~sRK[uƤ8|Ƭoݏp ?s#(#I8WC *W>m)2 et:5&eXޫ %] A&d 3Rq\objq%;!%\&wcF{NJ({ѴӴ` \\)JPL9ȴ[ڎ˃^U:, TWHhgp KIg<;M O +dy%Hc`&eF˰OTjVV9868^V:!I;!a%d+ $ SHK={Ef(`)}2DOcRNDxqi!zph Ւ:hƠ{4`I|f1ucbd .{R#^I ' -v^8^& Bml{ ꚯU6j=\^L!:jkYxX󗇦,S00CLȹfJ.'Eӓɻ+׉8[5O[ՆosXp=1S֕NGG]7꼟k\BŃk[<$'htRD$,JIC)xoK`a)l5vn>9/s-ˢҮO돛O*եu`8CPr8*^<JWe7d:;22%\t%Z1LXNMy6z$^-wO9^xC&=#l=X)8)+ t r)+ Id<ʜ@2JR 1$k8;Hi\[g]`b7U=bh]0{Rzt4`Q.o#'Jh8^e;59Gs^ȯ=)½ SVيgH2a2+,60Ύ<̲c/hjA}) in the novel allele. \item \code{AA_DIFF}: Number of amino acids that differ between the new allele and the closest reference (\code{CLOSEST_REFERENCE}) in the \code{germline_db} database. \item \code{AA_SUBSTITUTIONS}: A comma separated list with specific amino acid differences (e.g. \code{96A>N}) in the novel allele. \item \code{SEQUENCES}: Number of sequences unambiguosly assigned to this allele. \item \code{UNMUTATED_SEQUENCES}: Number of records with the unmutated novel allele sequence. \item \code{UNMUTATED_FREQUENCY}: Proportion of records with the unmutated novel allele sequence (\code{UNMUTATED_SEQUENCES / SEQUENCE}). \item \code{ALLELIC_PERCENTAGE}: Percentage at which the (unmutated) allele is observed in the sequence dataset compared to other (unmutated) alleles. \item \code{UNIQUE_JS}: Number of unique J sequences found associated with the novel allele. The sequences are those who have been unambiguously assigned to the novel allelle (\code{POLYMORPHISM_CALL}). \item \code{UNIQUE_CDR3S}: Number of unique CDR3s associated with the inferred allele. The sequences are those who have been unambiguously assigned to the novel allelle (POLYMORPHISM_CALL). \item \code{MUT_MIN}: Minimum mutation considered by the algorithm. \item \code{MUT_MAX}: Maximum mutation considered by the algorithm. \item \code{POS_MIN}: First position of the sequence considered by the algorithm (IMGT numbering). \item \code{POS_MAX}: Last position of the sequence considered by the algorithm (IMGT numbering). \item \code{Y_INTERCEPT}: The y-intercept above which positions were considered potentially polymorphic. \item \code{ALPHA}: Significance threshold to be used when constructing the confidence interval for the y-intercept. \item \code{MIN_SEQS}: Input \code{min_seqs}. The minimum number of total sequences (within the desired mutational range and nucleotide range) required for the samples to be considered. \item \code{J_MAX}: Input \code{j_max}. The maximum fraction of sequences perfectly aligning to a potential novel allele that are allowed to utilize to a particular combination of junction length and J gene. \item \code{MIN_FRAC}: Input \code{min_frac}. The minimum fraction of sequences that must have usable nucleotides in a given position for that position to be considered. \item \code{NOTE}: Comments regarding the novel allele inferrence. } } \description{ \code{generateEvidence} builds a table of evidence metrics for the final novel V allele detection and genotyping inferrences. } \examples{ \donttest{ # Generate input data novel <- findNovelAlleles(SampleDb, SampleGermlineIGHV) genotype <- inferGenotype(SampleDb, find_unmutated=TRUE, germline_db=SampleGermlineIGHV, novel=novel) genotype_db <- genotypeFasta(genotype, SampleGermlineIGHV, novel) data_db <- reassignAlleles(SampleDb, genotype_db) # Assemble evidence table evidence <- generateEvidence(data_db, novel, genotype, genotype_db, SampleGermlineIGHV) } } \seealso{ See \link{findNovelAlleles}, \link{inferGenotype} and \link{genotypeFasta} for generating the required input. } tigger/man/updateAlleleNames.Rd0000644000176200001440000000242113445226136016167 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{updateAlleleNames} \alias{updateAlleleNames} \title{Update IGHV allele names} \usage{ updateAlleleNames(allele_calls) } \arguments{ \item{allele_calls}{a vector of strings respresenting IGHV allele names.} } \value{ Vector of strings respresenting updated IGHV allele names. } \description{ \code{updateAlleleNames} takes a set of IGHV allele calls and replaces any outdated names (e.g. IGHV1-f) with the new IMGT names. } \note{ IGMT has removed \code{IGHV2-5*10} and \code{IGHV2-5*07} as it has determined they are actually alleles \code{02} and \code{04}, respectively. The updated allele names are based on IMGT release 201408-4. } \examples{ # Create a vector that uses old gene/allele names. alleles <- c("IGHV1-c*01", "IGHV1-f*02", "IGHV2-5*07") # Update the alleles to the new names updateAlleleNames(alleles) } \references{ \enumerate{ \item Xochelli et al. (2014) Immunoglobulin heavy variable (IGHV) genes and alleles: new entities, new names and implications for research and prognostication in chronic lymphocytic leukaemia. Immunogenetics. 67(1):61-6 } } \seealso{ Like \code{updateAlleleNames}, \link{sortAlleles} can help format a list of allele names. } tigger/man/plotGenotype.Rd0000644000176200001440000000263513445226136015302 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{plotGenotype} \alias{plotGenotype} \title{Show a colorful representation of a genotype} \usage{ plotGenotype(genotype, facet_by = NULL, gene_sort = c("name", "position"), text_size = 12, silent = FALSE, ...) } \arguments{ \item{genotype}{a \code{data.frame} of alleles denoting a genotype, as returned by \link{inferGenotype}.} \item{facet_by}{a column name in \code{genotype} to facet the plot by. If \code{NULL}, then do not facet the plot.} \item{gene_sort}{a string defining the method to use when sorting alleles. If \code{"name"} then sort in lexicographic order. If \code{"position"} then sort by position in the locus, as determined by the final two numbers in the gene name.} \item{text_size}{the point size of the plotted text.} \item{silent}{if \code{TRUE} do not draw the plot and just return the ggplot object; if \code{FALSE} draw the plot.} \item{...}{additional arguments to pass to ggplot2::theme.} } \value{ A ggplot object defining the plot. } \description{ \code{plotGenotype} plots a genotype table. } \examples{ # Plot genotype plotGenotype(SampleGenotype) # Facet by subject genotype_a <- genotype_b <- SampleGenotype genotype_a$SUBJECT <- "A" genotype_b$SUBJECT <- "B" geno_sub <- rbind(genotype_a, genotype_b) plotGenotype(geno_sub, facet_by="SUBJECT", gene_sort="pos") } \seealso{ \link{inferGenotype} } tigger/man/findUnmutatedCalls.Rd0000644000176200001440000000224713514172750016375 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{findUnmutatedCalls} \alias{findUnmutatedCalls} \title{Determine which calls represent an unmutated allele} \usage{ findUnmutatedCalls(allele_calls, sample_seqs, germline_db) } \arguments{ \item{allele_calls}{a vector of strings respresenting Ig allele calls, where multiple calls are separated by a comma.} \item{sample_seqs}{V(D)J-rearranged sample sequences matching the order of the given \code{allele_calls}.} \item{germline_db}{a vector of named nucleotide germline sequences} } \value{ A vector of strings containing the members of \code{allele_calls} that represent unmutated sequences. } \description{ \code{findUnmutatedCalls} determines which allele calls would represent a perfect match with the germline sequence, given a vector of allele calls and mutation counts. In the case of multiple alleles being assigned to a sequence, only the subset that would represent a perfect match is returned. } \examples{ # Find which of the sample alleles are unmutated calls <- findUnmutatedCalls(SampleDb$V_CALL, SampleDb$SEQUENCE_IMGT, germline_db=SampleGermlineIGHV) } tigger/man/cleanSeqs.Rd0000644000176200001440000000125413445226136014523 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{cleanSeqs} \alias{cleanSeqs} \title{Clean up nucleotide sequences} \usage{ cleanSeqs(seqs) } \arguments{ \item{seqs}{a vector of nucleotide sequences.} } \value{ A modified vector of nucleotide sequences. } \description{ \code{cleanSeqs} capitalizes nucleotides and replaces all characters besides \code{c("A", "C", "G", "T", "-", ".")} with \code{"N"}. } \examples{ # Clean messy nucleotide sequences seqs <- c("AGAT.taa-GAG...ATA", "GATACAGTXXZZAGNNPPACA") cleanSeqs(seqs) } \seealso{ \link{sortAlleles} and \link{updateAlleleNames} can help format a list of allele names. } tigger/man/plotNovel.Rd0000644000176200001440000000427413514172750014573 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{plotNovel} \alias{plotNovel} \title{Visualize evidence of novel V alleles} \usage{ plotNovel(data, novel_row, v_call = "V_CALL", j_call = "J_CALL", seq = "SEQUENCE_IMGT", junction = "JUNCTION", junction_length = "JUNCTION_LENGTH", ncol = 1) } \arguments{ \item{data}{a \code{data.frame} in Change-O format. See \link{findNovelAlleles} for details.} \item{novel_row}{a single row from a data frame as output by \link{findNovelAlleles} that contains a polymorphism-containing germline allele} \item{v_call}{name of the column in \code{data} with V allele calls. Default is "V_CALL".} \item{j_call}{name of the column in \code{data} with J allele calls. Default is J_CALL.} \item{seq}{name of the column in \code{data} with the aligned, IMGT-numbered, V(D)J nucleotide sequence. Default is SEQUENCE_IMGT.} \item{junction}{Junction region nucleotide sequence, which includes the CDR3 and the two flanking conserved codons. Default is JUNCTION.} \item{junction_length}{Number of junction nucleotides in the junction sequence. Default is JUNCTION_LENGTH.} \item{ncol}{number of columns to use when laying out the plots} } \description{ \code{plotNovel} is be used to visualize the evidence of any novel V alleles found using \link{findNovelAlleles}. It can also be used to visualize the results for alleles that did } \details{ The first panel in the plot shows, for all sequences which align to a particular germline allele, the mutation frequency at each postion along the aligned sequece as a function of the sequence-wide mutation. Sequences that pass the novel allele test are colored red, while sequences that don't pass the test are colored yellow. The second panel shows the nucleotide usage at the positions as a function of sequence-wide mutation count. To avoid cases where a clonal expansion might lead to a false positive, tigger examines the combinations of J gene and junction length among sequences which perfectly match the proposed germline allele. } \examples{ # Plot the evidence for the first (and only) novel allele in the example data novel <- selectNovel(SampleNovel) plotNovel(SampleDb, novel[1, ]) } tigger/man/SampleDb.Rd0000644000176200001440000000221413514172750014270 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/data.R \docType{data} \name{SampleDb} \alias{SampleDb} \title{Example human immune repertoire data} \format{A \code{data.frame} where rows correspond to unique V(D)J sequences and columns include: \itemize{ \item \code{"SEQUENCE_IMGT"}: IMGT-gapped V(D)J nucleotide sequence. \item \code{"V_CALL"}: IMGT/HighV-QUEST V segment allele calls. \item \code{"D_CALL"}: IMGT/HighV-QUEST D segment allele calls. \item \code{"J_CALL"}: IMGT/HighV-QUEST J segment allele calls. \item \code{"JUNCTION_LENGTH"}: Junction region length. }} \description{ A \code{data.frame} of example V(D)J immunoglobulin sequences derived from a single individual (PGP1), sequenced on the Roche 454 platform, and assigned by IMGT/HighV-QUEST to IGHV1 family alleles. } \references{ \enumerate{ \item Gadala-Maria, et al. (2015) Automated analysis of high-throughput B cell sequencing data reveals a high frequency of novel immunoglobulin V gene segment alleles. PNAS. 112(8):E862-70. } } \seealso{ See \link{airrDb} for an AIRR formatted version of \code{SampleDb}. } \keyword{data} tigger/man/reassignAlleles.Rd0000644000176200001440000000454413514172750015726 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{reassignAlleles} \alias{reassignAlleles} \title{Correct allele calls based on a personalized genotype} \usage{ reassignAlleles(data, genotype_db, v_call = "V_CALL", seq = "SEQUENCE_IMGT", method = "hamming", path = NA, keep_gene = c("gene", "family", "repertoire")) } \arguments{ \item{data}{a \code{data.frame} containing V allele calls from a single subject and the sample IMGT-gapped V(D)J sequences under \code{"SEQUENCE_IMGT"}.} \item{genotype_db}{a vector of named nucleotide germline sequences matching the calls detailed in \code{allele_calls} and personalized to the subject} \item{v_call}{name of the column in \code{data} with V allele calls. Default is \code{"V_CALL"}.} \item{seq}{name of the column in \code{data} with the aligned, IMGT-numbered, V(D)J nucleotide sequence. Default is SEQUENCE_IMGT} \item{method}{the method to be used when realigning sequences to the genotype_db sequences. Currently, only \code{"hammming"} (for Hamming distance) is implemented.} \item{path}{directory containing the tool used in the realignment method, if needed. Hamming distance does not require a path to a tool.} \item{keep_gene}{a string indicating if the gene (\code{"gene"}), family (\code{"family"}) or complete repertoire (\code{"repertoire"}) assignments should be performed. Use of \code{"gene"} increases speed by minimizing required number of alignments, as gene level assignments will be maintained when possible.} } \value{ A modifed input \code{data.frame} containing the best allele call from among the sequences listed in \code{genotype_db} in the \code{V_CALL_GENOTYPED} column. } \description{ \code{reassignAlleles} uses a subject-specific genotype to correct correct preliminary allele assignments of a set of sequences derived from a single subject. } \details{ In order to save time, initial gene assignments are preserved and the allele calls are chosen from among those provided in \code{genotype_db}, based on a simple alignment to the sample sequence. } \examples{ # Extract the database sequences that correspond to the genotype genotype_db <- genotypeFasta(SampleGenotype, SampleGermlineIGHV, novel=SampleNovel) # Use the personlized genotype to determine corrected allele assignments output_db <- reassignAlleles(SampleDb, genotype_db) } tigger/man/writeFasta.Rd0000644000176200001440000000143413445226136014716 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{writeFasta} \alias{writeFasta} \title{Write to a fasta file} \usage{ writeFasta(named_sequences, file, width = 60, append = FALSE) } \arguments{ \item{named_sequences}{a vector of named string representing sequences} \item{file}{the name of the output file.} \item{width}{the number of characters to be printed per line. if not between 1 and 255, width with be infinite.} \item{append}{\code{logical} indicating if the output should be appended to \code{file} instead of overwriting it} } \value{ A named vector of strings respresenting Ig alleles. } \description{ \code{writeFasta} writes a named vector of sequences to a file in fasta format. } \seealso{ \link{readIgFasta} to do the inverse. } tigger/man/getMutCount.Rd0000644000176200001440000000306213514172750015061 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{getMutCount} \alias{getMutCount} \title{Determine the mutation counts from allele calls} \usage{ getMutCount(samples, allele_calls, germline_db) } \arguments{ \item{samples}{a vector of IMGT-gapped sample V sequences} \item{allele_calls}{a vector of strings respresenting Ig allele calls for the sequences in \code{samples}, where multiple calls are separated by a comma} \item{germline_db}{a vector of named nucleotide germline sequences matching the calls detailed in \code{allele_calls}} } \value{ A list equal in length to \code{samples}, containing the Hamming distance to each germline allele contained within each call within each element of \code{samples} } \description{ \code{getMutCount} takes a set of nucleotide sequences and their allele calls and determines the distance between that seqeunce and any germline alleles contained within the call } \examples{ # Insert a mutation into a germline sequence s2 <- s3 <- SampleGermlineIGHV[1] stringi::stri_sub(s2, 103, 103) <- "G" stringi::stri_sub(s3, 107, 107) <- "C" sample_seqs <- c(SampleGermlineIGHV[2], s2, s3) # Pretend that one sample sequence has received an ambiguous allele call sample_alleles <- c(paste(names(SampleGermlineIGHV[1:2]), collapse=","), names(SampleGermlineIGHV[2]), names(SampleGermlineIGHV[1])) # Compare each sequence to its assigned germline(s) to determine the distance getMutCount(sample_seqs, sample_alleles, SampleGermlineIGHV) } tigger/man/subsampleDb.Rd0000644000176200001440000000434713514172750015053 0ustar liggesusers% Generated by roxygen2: do not edit by hand % Please edit documentation in R/functions.R \name{subsampleDb} \alias{subsampleDb} \title{Subsample repertoire} \usage{ subsampleDb(data, gene = "V_CALL", mode = c("gene", "allele", "family"), min_n = 1, max_n = NULL, group = NULL) } \arguments{ \item{data}{a \code{data.frame} in Change-O format.} \item{gene}{name of the column in \code{data} with allele calls} \item{mode}{one of c("gene", "family", "allele") defining the degree of specificity regarding allele calls when subsetting sequences. Determines how \code{data} will be split into subsets from which the same number of sequences will be subsampled. See also \code{group}.} \item{min_n}{minimum number of observations to sample from each groupe. A group with less observations than the minimum is excluded.} \item{max_n}{maximum number of observations to sample for all \code{mode} groups. If \code{NULL}, it will be set automatically to the size of the smallest group. If \code{max_n} is larger than the availabe number of sequences for any \code{mode} group, if will be automatically adjusted and the efective \code{max_n} used will be the size of the smallest \code{mode} group.} \item{group}{columns containing additional grouping variables, e.g. sample_id. These groups will be subsampled independently. If \code{max_n} is \code{NULL}, a \code{max_n} will be automatically set for each \code{group}.} } \value{ A \code{data.frame}, subsampled from \code{data}. } \description{ \code{subsampleDb} will sample the same number of sequences for each gene, family or allele (specified with \code{mode}) in \code{data}. Samples or subjects can be subsampled indepently by setting \code{group}. } \details{ \code{data} will be split into gene, allele or family subsets (\code{mode}) from which the same number of sequences will be subsampled. If \code{mode=gene}, for each gene in the field \code{gene} from \code{data}, a maximum of \code{max_n} sequences will be subsampled. Input sequences that have multiple gene calls (ties), can be subsampled from any of their calls, but these duplicated samplings will be removed, and the final subsampled \code{data} will contain unique rows. } \examples{ # subsampleDb(SampleDb) } \seealso{ \link{selectNovel} } tigger/DESCRIPTION0000644000176200001440000000374513514253663013262 0ustar liggesusersPackage: tigger Type: Package Version: 0.4.0 Date: 2019-07-18 Authors@R: c(person("Daniel", "Gadala-Maria", role=c("aut"), email="daniel.gadala-maria@yale.edu"), person("Susanna", "Marquez", role=c("aut"), email="susanna.marquez@yale.edu"), person("Moriah", "Cohen", role=c("aut"), email="moriah.cohen@biu.ac.il"), person("Gur", "Yaari", role=c("aut"), email="gur.yaari@biu.ac.il"), person("Jason", "Vander Heiden", role=c("ctb", "cre"), email="jason.vanderheiden@yale.edu"), person("Steven", "Kleinstein", role=c("aut", "cph"), email="steven.kleinstein@yale.edu")) Title: Infers Novel Immunoglobulin Alleles from Sequencing Data Description: Infers the V genotype of an individual from immunoglobulin (Ig) repertoire sequencing data (AIRR-Seq, Rep-Seq). Includes detection of any novel alleles. This information is then used to correct existing V allele calls from among the sample sequences. Citations: Gadala-Maria, et al (2015) . License: CC BY-SA 4.0 URL: http://tigger.readthedocs.io BugReports: https://bitbucket.org/kleinstein/tigger/issues LazyData: true BuildVignettes: true VignetteBuilder: knitr Encoding: UTF-8 Depends: R (>= 3.2.5), ggplot2 (>= 3.1.1) Imports: alakazam (>= 0.3.0), dplyr (>= 0.8.1), doParallel, foreach, graphics, gridExtra, gtools, iterators, lazyeval, parallel, rlang, shazam (>= 0.2.0), stats, stringi, tidyr Suggests: knitr, testthat RoxygenNote: 6.1.1 NeedsCompilation: no Packaged: 2019-07-19 05:20:14 UTC; vandej27 Author: Daniel Gadala-Maria [aut], Susanna Marquez [aut], Moriah Cohen [aut], Gur Yaari [aut], Jason Vander Heiden [ctb, cre], Steven Kleinstein [aut, cph] Maintainer: Jason Vander Heiden Repository: CRAN Date/Publication: 2019-07-19 05:40:03 UTC tigger/build/0000755000176200001440000000000013514251416012634 5ustar liggesuserstigger/build/vignette.rds0000644000176200001440000000034413514251416015174 0ustar liggesusersQ 0 9 (]<xZlV [;旫:N04-シÔ:ħK9H@"<j/k!V܁,7l+mlZ%LDҶtk^8`4ڍ'#?#*hq5(=_C=ks0 䲄~Nnz[@d9wusrp=qrfYZwﲛtigger/vignettes/0000755000176200001440000000000013514251416013545 5ustar liggesuserstigger/vignettes/Tigger-Vignette.Rmd0000644000176200001440000003074413513673571017236 0ustar liggesusers--- title: "Tool for Immunoglobulin Genotype Elucidation via Rep-Seq (TIgGER)" author: "Daniel Gadala-Maria" date: "Last modified `r Sys.Date()`" output: pdf_document: dev: pdf fig_height: 4 fig_width: 7.5 highlight: pygments toc: yes html_document: fig_height: 4 fig_width: 7.5 highlight: pygments theme: readable toc: yes md_document: fig_height: 4 fig_width: 7.5 preserve_yaml: no toc: yes geometry: margin=1in fontsize: 11pt vignette: > %\VignetteEngine{knitr::rmarkdown} %\VignetteIndexEntry{Polymorphism detection and genotyping} %\usepackage[utf8]{inputenc} --- ## Introduction Adapative immune receptor repertoire sequencing (AIRR-Seq, Rep-Seq) data is currently the subject of much study. A key step in analyzing these data involves assigning the closest known V(D)J germline alleles to the (often somatically mutated) sample sequences using a tool such as IMGT/HighV-QUEST ([[1]][1]). However, if the sample utilizes alleles not in the germline database used for alignment, this step will fail. Additionally, this alignment has an associated error rate of ~5% ([[2]][2]), notably among sequences carrying a large number of somatic mutations. Here we provide a **T**ool for **I**mmuno**g**lobulin **G**enotype **E**lucidation via **R**ep-Seq (TIgGER). TIgGER addresses these issues by inferring the set of Immunoglobulin (Ig) alleles carried by an individual (including any novel alleles) and then using this set of alleles to correct the initial assignments given to sample sequences by existing tools. This vignette covers the following tasks: 1. Inferring the presence of novel IGHV alleles not in the germline database. 2. Inferring the personalized IGHV genotype of a sample. 3. Correcting the IGHV allele calls of a sample based on the IGHV genotype. Additional information about the methods used by TIgGER is available in: [Gadala-Maria D, Yaari G, Uduman M, Kleinstein SH (2015) Automated analysis of high-throughput B cell sequencing data reveals a high frequency of novel immunoglobulin V gene segment alleles. *PNAS* 112(8):E862-70](http://www.pnas.org/content/early/2015/02/05/1417683112). ## Input TIgGER requires two main inputs: 1. Pre-processed Ig sequence data 2. Database germline sequences AIRR-seq data is input as a data frame following the Change-O standard where each row represents a unique observation and and columns represent data about that observation. The required names of the required columns are provided below along with a description of each. Column Name | Description ----------------------|--------------------------------------------------------- `SEQUENCE_IMGT` | V(D)J sequence gapped in the IMGT gapped format ([[3]][3]) `V_CALL` | (Comma separated) name(s) of the nearest V allele(s) `J_CALL` | (Comma separated) name(s) of the nearest J allele(s) `JUNCTION_LENGTH` | Length of the junction region of the V(D)J sample An example dataset is provided with the `tigger` package as `SampleDb`. It contains unique functional sequences assigned to IGHV1 family genes isolated from individual PGP1 (referenced in Gadala-Maria *et al.* 2015). The database of germline sequences should be provided in FASTA format with sequences gapped according to the IMGT numbering scheme ([[3]][3]). IGHV alleles in the IMGT database (build 201408-4) are provided with this package as `GermlineIGHV`. You may read in your own fasta file using `readIgFasta`. ```{r, eval=TRUE, message=FALSE, warning=FALSE} # Load packages required for this example library(tigger) library(dplyr) ``` ## Novel allele detection Potential novel alleles can be detected by TIgGER. Some of these may be included in the genotype later (see below). `findNovelAlleles` will return a `data.frame` with a row for each allele tested for the presence of polymorphisms. If polymorphisms are found and the novel sequence passes all quality tests (see below), then the novel allele is named and the germline sequence is included in the `data.frame`. Additionally, the result will contain metadata on the parameters used when searching for novel alleles (which can be optionally changed in `findNovelAlleles`). ```{r, eval=TRUE, warning=FALSE} # Detect novel alleles novel <- findNovelAlleles(SampleDb, GermlineIGHV, nproc=1) ``` ```{r, eval=TRUE, warning=FALSE} # Extract and view the rows that contain successful novel allele calls novel_rows <- selectNovel(novel) novel_rows[1:3] ``` The TIgGER procedure for identifying novel alleles (see citation above) involves taking all sequences which align to a particular germline allele and, for each position along the aligned sequences, plotting the mutation frequency at that postion as a function of the sequence-wide mutation count. While mutational hot-spots and cold-spots are both expected to have a *y*-intercept around zero, polymorphic positions will have a *y*-intercept larger than zero. The required minimum *y*-intercept may be specified in `findNovelAlleles` by `y_intercept`, but defaults to `1/8`. Passing this *y*-intercept threshold is the first of three pieces of evidence that support the novel allele. The second piece of evidence supporting novel allele calls is the nucleotide usage at the polymorphic positions as a function of sequence-wide mutation count. We expect the polymorphic allele to be prevalent at all mutation counts, and we expect the mutation count equal to the number of polymorphisms in the novel sequence to be the most prevalent. Finally, to avoid cases where a clonal expansion might lead to a false positive, combinations of J gene and junction length are examined among sequences which *perfectly match* the proposed germline allele. A true novel allele is expected to utilize a wide range of J genes, and sequences with different junction length can be ruled out as not being clonally related. The maximum portion of sequences which can consist of a specific combination of J gene and junction length may be specified in `findNovelAlleles` by `j_max`. The three pieces of evidence desribed above can be viewed for any allele call made by `findNovelAlleles` using the function `plotNovel`. ```{r, eval=TRUE, warning=FALSE, fig.width=6, fig.height=8} # Plot evidence of the first (and only) novel allele from the example data plotNovel(SampleDb, novel[1, ]) ``` ## Inferring genotypes An individual's genotype can be inferred using the functions `inferGenotype` or `inferGenotypeBayesian`. Using one of this functions allows to remove from the genotype rare/erroneous allele calls which may result from mutations in allele-differentiating regions. `inferGenotype` uses a frequency method to decide which alleles belong to the subjects genotype whereas `inferGenotypeBayesian` infers an subject's genotype applying a Bayesian framework and provides a confidence estimate associated with the genotype calls. ### Frequency genotyping approach `inferGenotype` identifies the fewest alleles that account for nearly all (default is 7/8) of the allele calls made. The user may opt to only use sequences which perfectly match germline alleles, and may opt to include potential novel alleles. (The genotype output is designed to be human readable, though `plotGenotype` can be used to make a colorful visualization.) For each allele, the number of sequences which match the germline are listed in the same order as the alleles are listed. The total number of sequences that match any allele of that gene is also given. To output these alleles as a names vector of nucleotide sequences, the user may use the function `genotypeFasta`. To save this vector to a fasta file, `writeFasta` may be used. ```{r, eval=TRUE, warning=FALSE, fig.width=4, fig.height=3} # Infer the individual's genotype, using only unmutated sequences and checking # for the use of the novel alleles inferred in the earlier step. geno <- inferGenotype(SampleDb, germline_db=GermlineIGHV, novel=novel, find_unmutated=TRUE) # Save the genotype sequences to a vector genotype_db <- genotypeFasta(geno, GermlineIGHV, novel) # Visualize the genotype and sequence counts print(geno) # Make a colorful visualization. Bars indicate presence, not proportion. plotGenotype(geno, text_size = 10) ``` ### Bayesian genotyping approach The method `inferGenotypeBayesian` analyzes the posterior probabilities of possible allele distributions, considering up to four distinct alleles per V gene, corresponding to a gene duplication with both loci being heterozygous (i.e., homozygous, heterozygous with one copy of each allele, etc.). The posterior probabilities for these four possible models are compared and a Bayes factor is calculated for the two most probable models. This Bayes factor reflects the confidence in the genotyping call of the method. The bayesian method doesn't use the strict cutoff criterion `fraction_to_explain` that `inferGenotype` uses wherein only the minimum set of alleles explaining 88% (7/8) of apparently-unmutated sequences are included in the genotype. ```{r, eval=TRUE, warning=FALSE, fig.width=4, fig.height=3} # Infer the individual's genotype, using the bayesian method geno_bayesian <- inferGenotypeBayesian(SampleDb, germline_db=GermlineIGHV, novel=novel, find_unmutated=TRUE) # Visualize the genotype and sequence counts print(geno_bayesian) # Make a colorful visualization. Bars indicate presence, not proportion. plotGenotype(geno_bayesian, text_size=10) ``` ## Correcting allele calls Finally, the original V allele calls may be limited to only those within the inferred genotype. This can be done by using the function `reassignAlleles`. By correcting the calls in this manner, the user can greatly reduce the numer of ambiguous allele calls (where a single sample sequences is assigned to multiple V alleles, thus preventing the mutations analysis of allele-differentiating positions). Additionally, assignments to erroneous not-in-genotype alleles (expected to be ~5% ([[2]][2]), as mentioned above, are corrected in this manner. ```{r, eval=TRUE, warning=FALSE} # Use the personlized genotype to determine corrected allele assignments # Updated genotype will be placed in the V_CALL_GENOTYPED column sample_db <- reassignAlleles(SampleDb, genotype_db) ``` From here, one may proceed with further downstream analyses, but with the advantage of having much-improved allele calls. Besides having discovered alleles not in the IMGT database, the calls made by IMGT have been tailored to the subject's genotype, greatly reducing the number of problematic calls, as can be seen below. ```{r, eval=TRUE, warning=FALSE} # Find the set of alleles in the original calls that were not in the genotype not_in_genotype <- sample_db$V_CALL %>% strsplit(",") %>% unlist() %>% unique() %>% setdiff(names(genotype_db)) # Determine the fraction of calls that were ambigious before/after correction # and the fraction that contained original calls to non-genotype alleles. Note # that by design, only genotype alleles are allowed in "after" calls. data.frame(Ambiguous=c(mean(grepl(",", sample_db$V_CALL)), mean(grepl(",", sample_db$V_CALL_GENOTYPED))), NotInGenotype=c(mean(sample_db$V_CALL %in% not_in_genotype), mean(sample_db$V_CALL_GENOTYPED %in% not_in_genotype)), row.names=c("Before", "After")) %>% t() %>% round(3) ``` ## Evidence `generateEvidence` uses the final corrected calls, the novel alleles and genotype information, the final genotype sequences and the starting reference germlines to build a table of evidence metrics supporting the final novel V allele detection. ```{r, eval=TRUE, warning=FALSE} evidence <- generateEvidence(sample_db, novel, geno, genotype_db, GermlineIGHV, j_call = "J_CALL", junction = "JUNCTION", fields = NULL) evidence %>% select(GENE, ALLELE, POLYMORPHISM_CALL, SEQUENCES, UNMUTATED_FREQUENCY) ``` In this example, `r evidence[["SEQUENCES"]]` sequences were unambiguously assigned to allele `r evidence[["POLYMORPHISM_CALL"]]`, `r round(100*evidence[["UNMUTATED_FREQUENCY"]],2)`% of them unmutated. ## References 1. [Alamyar *et al.* (2010)](http://www.imgt.org/IMGTindex/IMGTHighV-QUEST.html) 2. [Munshaw and Kepler (2010)](http://www.ncbi.nlm.nih.gov/pubmed/20147303) 3. [Lefranc *et al.* (2003)](http://www.ncbi.nlm.nih.gov/pubmed/12477501) [1]: http://www.imgt.org/IMGTindex/IMGTHighV-QUEST.html "Alamyar et al. (2010)" [2]: http://www.ncbi.nlm.nih.gov/pubmed/20147303 "Munshaw and Kepler (2010)" [3]: http://www.ncbi.nlm.nih.gov/pubmed/12477501 "Lefranc et al. (2003)" tigger/R/0000755000176200001440000000000013514172710011735 5ustar liggesuserstigger/R/functions.R0000644000176200001440000030224113514233032014065 0ustar liggesusers # The TIgGER Trifecta ----------------------------------------------------- #' Find novel alleles from repertoire sequencing data #' #' \code{findNovelAlleles} analyzes mutation patterns in sequences thought to #' align to each germline allele in order to determine which positions #' might be polymorphic. #' #' The TIgGER allele-finding algorithm, briefly, works as follows: #' Mutations are determined through comparison to the provided germline. #' Mutation frequency at each *position* is determined as a function of #' *sequence-wide* mutation counts. Polymorphic positions exhibit a high #' mutation frequency despite sequence-wide mutation count. False positive of #' potential novel alleles resulting from clonally-related sequences are guarded #' against by ensuring that sequences perfectly matching the potential novel #' allele utilize a wide range of combinations of J gene and junction length. #' #' @param data a \code{data.frame} in Change-O format. See details. #' @param germline_db a vector of named nucleotide germline sequences #' matching the V calls in \code{data}. #' @param v_call name of the column in \code{data} with V allele calls. #' Default is V_CALL. #' @param j_call name of the column in \code{data} with J allele calls. #' Default is J_CALL. #' @param seq name of the column in \code{data} with the #' aligned, IMGT-numbered, V(D)J nucleotide sequence. #' Default is SEQUENCE_IMGT. #' @param junction Junction region nucleotide sequence, which includes #' the CDR3 and the two flanking conserved codons. Default #' is JUNCTION. #' @param junction_length Number of junction nucleotides in the junction sequence. #' Default is JUNCTION_LENGTH. #' @param germline_min the minimum number of sequences that must have a #' particular germline allele call for the allele to #' be analyzed #' @param auto_mutrange if \code{TRUE}, the algorithm will attempt to #' determine the appropriate mutation range #' automatically using the mutation count of the most #' common sequence assigned to each allele analyzed #' @param mut_range the range of mutations that samples may carry and #' be considered by the algorithm #' @param pos_range the range of IMGT-numbered positions that should be #' considered by the algorithm #' @param alpha the alpha value used for determining whether the #' fit y-intercept is greater than the \code{y_intercept} #' threshold #' @param y_intercept the y-intercept threshold above which positions should be #' considered potentially polymorphic #' @param j_max the maximum fraction of sequences perfectly aligning #' to a potential novel allele that are allowed to #' utilize to a particular combination of junction #' length and J gene #' @param min_seqs the minimum number of total sequences (within the #' desired mutational range and nucleotide range) #' required for the samples to be considered #' @param min_frac the minimum fraction of sequences that must have #' usable nucleotides in a given position for that #' position to considered #' @param nproc the number of processors to use #' #' @return #' A \code{data.frame} with a row for each known allele analyzed. #' Besides metadata on the the parameters used in the search, each row will have #' either a note as to where the polymorphism-finding algorithm exited or a #' nucleotide sequence for the predicted novel allele, along with columns providing #' additional evidence. #' #' The output contains the following columns: #' \itemize{ #' \item \code{GERMLINE_CALL}: The input (uncorrected) V call. #' \item \code{NOTE}: Comments regarding the inferrence. #' \item \code{POLYMORPHISM_CALL}: The novel allele call. #' \item \code{NT_SUBSTITUTIONS}: Mutations identified in the novel allele, relative #' to the reference germline (\code{GERMLINE_CALL}) #' \item \code{NOVEL_IMGT}: The novel allele sequence. #' \item \code{NOVEL_IMGT_COUNT}: The number of times the sequence \code{NOVEL_IMGT} #' is found in the input data. Considers the subsequence of \code{NOVEL_IMGT} #' in the \code{pos_range}. #' \item \code{NOVEL_IMGT_UNIQUE_J}: Number of distinct J calls associated to \code{NOVEL_IMGT} #' in the input data. Considers the subsequence of \code{NOVEL_IMGT} in the \code{pos_range}. #' \item \code{NOVEL_IMGT_UNIQUE_CDR3}: Number of distinct CDR3 sequences associated #' with \code{NOVEL_IMGT} in the input data. Considers the subsequence of \code{NOVEL_IMGT} #' in the \code{pos_range}. #' \item \code{PERFECT_MATCH_COUNT}: Final number of sequences retained to call the new #' allele. These are unique sequences that have V segments that perfectly match #' the predicted germline in the \code{pos_range}. #' \item \code{PERFECT_MATCH_FREQ}: \code{PERFECT_MATCH_COUNT / GERMLINE_CALL_COUNT} #' \item \code{GERMLINE_CALL_COUNT}: The number of sequences with the \code{GERMLINE_CALL} #' in the input data that were initially considered for the analysis. #' \item \code{GERMLINE_CALL_FREQ}: The fraction of sequences with the \code{GERMLINE_CALL} #' in the input data initially considered for the analysis. #' \item \code{GERMLINE_IMGT}: Germline sequence for \code{GERMLINE_CALL}. #' \item \code{GERMLINE_IMGT_COUNT}: The number of times the \code{GERMLINE_IMGT} #' sequence is found in the input data. #' \item \code{MUT_MIN}: Minimum mutation considered by the algorithm. #' \item \code{MUT_MAX}: Maximum mutation considered by the algorithm. #' \item \code{MUT_PASS_COUNT}: Number of sequences in the mutation range. #' \item \code{POS_MIN}: First position of the sequence considered by the algorithm (IMGT numbering). #' \item \code{POS_MAX}: Last position of the sequence considered by the algorithm (IMGT numbering). #' \item \code{Y_INTERCEPT}: The y-intercept above which positions were considered #' potentially polymorphic. #' \item \code{Y_INTERCEPT_PASS}: Number of positions that pass the \code{Y_INTERCEPT} threshold. #' \item \code{SNP_PASS}: Number of sequences that pass the \code{Y_INTERCEPT} threshold and are #' within the desired nucleotide range (\code{min_seqs}). #' \item \code{UNMUTATED_COUNT}: Number of unmutated sequences. #' \item \code{UNMUTATED_FREQ}: Number of unmutated sequences over \code{GERMLINE_IMGT_COUNT}. #' \item \code{UNMUTATED_SNP_J_GENE_LENGTH_COUNT}: Number of distinct combinations #' of SNP, J gene, and junction length. #' \item \code{SNP_MIN_SEQS_J_MAX_PASS}: Number of SNPs that pass both the \code{min_seqs} #' and \code{j_max} thresholds. #' \item \code{ALPHA}: Significance threshold to be used when constructing the #' confidence interval for the y-intercept. #' \item \code{MIN_SEQS}: Input \code{min_seqs}. The minimum number of total sequences #' (within the desired mutational range and nucleotide range) required #' for the samples to be considered. #' \item \code{J_MAX}: Input \code{j_max}. The maximum fraction of sequences perfectly #' aligning to a potential novel allele that are allowed to utilize to a particular #' combination of junction length and J gene. #' \item \code{MIN_FRAC}: Input \code{min_frac}. The minimum fraction of sequences that must #' have usable nucleotides in a given position for that position to be considered. #' } #' #' The following comments can appear in the \code{NOTE} column: #' #' \itemize{ #' \item \emph{Novel allele found}: A novel allele was detected. #' \item \emph{Plurality sequence too rare}: No sequence is frequent enough to pass #' the J test (\code{j_max}). #' \item \emph{A J-junction combination is too prevalent}: Not enough J diversity (\code{j_max}). #' \item \emph{No positions pass y-intercept test}: No positions above \code{y_intercept}. #' \item \emph{Insufficient sequences in desired mutational range}: #' \code{mut_range} and \code{pos_range}. #' \item \emph{Not enough sequences}: Not enough sequences in the desired mutational #' range and nucleotide range (\code{min_seqs}). #' \item \emph{No unmutated versions of novel allele found}: All observed variants of the #' allele are mutated. #' } #' #' @seealso \link{selectNovel} to filter the results to show only novel alleles. #' \link{plotNovel} to visualize the data supporting any #' novel alleles hypothesized to be present in the data and #' \link{inferGenotype} to determine if the novel alleles are frequent #' enought to be included in the subject's genotype. #' #' @examples #' \donttest{ #' # Find novel alleles and return relevant data #' novel <- findNovelAlleles(SampleDb, SampleGermlineIGHV) #' selectNovel(novel) #' # Note: In this example, with SampleGermlineIGHV, #' # which contains reference germlines retrieved on August 2018, #' # TIgGER finds the allele IGHV1-8*02_G234T. This allele #' # was added to IMGT as IGHV1-8*03 on March 28, 2018. #' } #' #' @export findNovelAlleles <- function(data, germline_db, v_call="V_CALL", j_call="J_CALL", seq="SEQUENCE_IMGT", junction="JUNCTION", junction_length="JUNCTION_LENGTH", germline_min=200, min_seqs=50, auto_mutrange=TRUE, mut_range=1:10, pos_range=1:312, y_intercept=0.125, alpha=0.05, j_max=0.15, min_frac=0.75, nproc=1) { . = idx = NULL # Keep only the db columns needed data <- data %>% dplyr::select(!!!rlang::syms(c(seq, v_call, j_call, junction_length, junction))) gc() # Keep only the columns we need and clean up the sequences missing <- c(seq, v_call, j_call, junction_length) %>% setdiff(colnames(data)) if (length(missing) != 0) { stop("Could not find required columns in the input data:\n ", paste(missing, collapse="\n ")) } empty_junctions <- sum(data[[junction_length]] == 0, na.rm=TRUE) if (empty_junctions > 0) { stop(empty_junctions, " sequences have junction ", "length of zero. ", "Please remove these sequences.") } germlines <- cleanSeqs(germline_db) names(germlines) <- getAllele(names(germlines), first=FALSE, strip_d=FALSE) data[[seq]] <- cleanSeqs(data[[seq]]) # Find which rows' calls contain which germline alleles cutoff <- ifelse(germline_min < 1, round(nrow(data)*germline_min), germline_min) allele_groups <- sapply(names(germlines), grep, data[[v_call]], fixed=TRUE, simplify=FALSE) names(allele_groups) <- names(germlines) allele_groups <- allele_groups[sapply(allele_groups, length) >= cutoff] if(length(allele_groups) == 0){ stop_message <- paste("Not enough sample sequences were assigned to any germline:\n", " (1) germline_min is too large or\n", " (2) sequences names don't match germlines.") stop(stop_message) } allele_groups <- allele_groups[sortAlleles(names(allele_groups))] # Prepare for parallel processing nproc <- ifelse(Sys.info()['sysname'] == "Windows", Sys.getenv('NUMBER_OF_PROCESSORS'), ifelse(Sys.info()['sysname'] == "Darwin", system("sysctl -n hw.ncpu", intern=TRUE), system("nproc", intern=TRUE))) %>% as.numeric() %>% min(nproc, . - 1) %>% max(1, .) if(nproc == 1) { foreach::registerDoSEQ() } else { #cluster_type = ifelse(Sys.info()['sysname'] == "Windows", "PSOCK", "FORK") cluster <- parallel::makeCluster(nproc, type="PSOCK") parallel::clusterExport(cluster, list("allele_groups", "germlines", "v_call","j_call", "junction", "junction_length", "seq", "data", "min_seqs", "auto_mutrange", "mut_range", "pos_range", "y_intercept", "alpha", "j_max", "germline_min", "min_frac", "findLowerY", "mutationRangeSubset", "positionMutations", "superSubstring"), envir=environment()) doParallel::registerDoParallel(cluster) } out_list <- foreach(idx=iterators::icount(length(allele_groups))) %dopar% { # out_list <- lapply(1:length(allele_groups), function(idx) { gc() # message(paste0("idx=",idx)) # Subset of data being analyzed allele_name <- names(allele_groups)[idx] germline <- germlines[allele_name] indicies <- allele_groups[[allele_name]] db_subset <- data[indicies, ] # If mutrange is auto, find most popular mutation count and start from there gpm <- db_subset %>% dplyr::mutate(!!v_call := allele_name ) %>% getPopularMutationCount(germline, gene_min=0, seq_min=min_seqs, seq_p_of_max=1/8, full_return=TRUE, v_call=v_call, seq=seq) # Determine the mutation range(s) to scan mut_mins <- min(mut_range) if ( auto_mutrange & sum(gpm$MUTATION_COUNT > 0) > 0 ){ mut_mins <- c(mut_mins, gpm$MUTATION_COUNT[gpm$MUTATION_COUNT > 0]) %>% unique() %>% sort() } # Create the run's return object df_run_empty <- data.frame(GERMLINE_CALL = names(germline), NOTE = "", POLYMORPHISM_CALL = NA, NT_SUBSTITUTIONS=NA, NOVEL_IMGT = NA, NOVEL_IMGT_COUNT=NA, NOVEL_IMGT_UNIQUE_J=NA, NOVEL_IMGT_UNIQUE_CDR3=NA, PERFECT_MATCH_COUNT = NA, PERFECT_MATCH_FREQ = NA, GERMLINE_CALL_COUNT = length(indicies), GERMLINE_CALL_FREQ = round(length(indicies)/nrow(data), 3), MUT_MIN = NA, MUT_MAX = NA, MUT_PASS_COUNT=NA, GERMLINE_IMGT = as.character(germline), GERMLINE_IMGT_COUNT=NA, POS_MIN = min(pos_range), POS_MAX = max(pos_range), Y_INTERCEPT = y_intercept, Y_INTERCEPT_PASS = NA, SNP_PASS=NA, UNMUTATED_COUNT=NA, UNMUTATED_FREQ=NA, UNMUTATED_SNP_J_GENE_LENGTH_COUNT=NA, SNP_MIN_SEQS_J_MAX_PASS=NA, ALPHA = alpha, MIN_SEQS = min_seqs, J_MAX = j_max, MIN_FRAC = min_frac, stringsAsFactors = FALSE) for (mut_min in rev(mut_mins)) { gc() # message(paste0("|-- mut_min=",mut_min)) if (mut_min == rev(mut_mins)[1]){ df_run <- df_run_empty } else { df_run <- dplyr::bind_rows(df_run_empty, df_run) } mut_max <- mut_min + diff(range(mut_range)) df_run$MUT_MIN[1] <- mut_min df_run$MUT_MAX[1] <- mut_max # If no sequence is frequent enough to pass the J test, give up now if(nrow(gpm) < 1) { df_run$NOTE[1] <- "Plurality sequence too rare." if(mut_mins[1] == mut_min){ return(df_run) } else { next } } # Add a mutation count column and filter out sequences not in our range db_subset_mm <- mutationRangeSubset(db_subset, germline, mut_min:mut_max, pos_range, seq=seq) df_run$MUT_PASS_COUNT[1] <- nrow(db_subset_mm) if(nrow(db_subset_mm) < min_seqs){ df_run$NOTE[1] <- paste0("Insufficient sequences (",nrow(db_subset_mm),") in desired mutational range.") if(mut_mins[1] == mut_min){ return(df_run) } else { next } } # Duplicate each sequence for all the positions to be analyzed # and find which positions are mutated pos_db <- positionMutations(db_subset_mm, germline, pos_range, seq=seq) # Find positional mut freq vs seq mut count pos_muts <- pos_db %>% dplyr::group_by(!!rlang::sym("POSITION")) %>% dplyr::mutate(PASS = mean(!! rlang::sym("OBSERVED")) >= min_frac) %>% dplyr::group_by(!!!rlang::syms(c("MUT_COUNT", "POSITION"))) %>% dplyr::summarise(POS_MUT_RATE = mean(!!rlang::sym("MUTATED"))*unique(!!rlang::sym("PASS")) ) %>% dplyr::ungroup() rm(pos_db) gc() # Calculate y intercepts, find which pass the test pass_y <- pos_muts %>% dplyr::group_by(!!rlang::sym("POSITION")) %>% dplyr::summarise(Y_INT_MIN = findLowerY(!!rlang::sym("POS_MUT_RATE"), !!rlang::sym("MUT_COUNT"), mut_min, alpha)) %>% dplyr::filter(!!rlang::sym("Y_INT_MIN") > y_intercept) df_run$Y_INTERCEPT_PASS[1] <- nrow(pass_y) if(nrow(pass_y) < 1){ df_run$NOTE[1] <- "No positions pass y-intercept test." if(mut_mins[1] == mut_min){ return(df_run) } else { next } } gl_substring <- superSubstring(germline, pass_y$POSITION) gl_minus_substring <- insertPolymorphisms(germline, pass_y$POSITION, rep("N", nrow(pass_y))) # Find the potential SNP positions and remove anything that matches # the germline at all those positions or any combo that is too rare db_y_subset_mm <- db_subset_mm %>% dplyr::group_by(1:n()) %>% dplyr::mutate(SNP_STRING = superSubstring(!! rlang::sym(seq), pass_y$POSITION)) %>% dplyr::filter(!!rlang::sym("SNP_STRING") != gl_substring) %>% dplyr::group_by(!!rlang::sym("SNP_STRING")) %>% dplyr::mutate(STRING_COUNT = n()) %>% dplyr::filter(!!rlang::sym("STRING_COUNT") >= min_seqs) df_run$SNP_PASS[1] <- nrow(db_y_subset_mm) if (nrow(db_y_subset_mm) < 1 ){ df_run$NOTE[1] <- paste("Position(s) passed y-intercept (", paste(pass_y$POSITION, collapse = ","), ") but the plurality sequence is too rare.", sep="") if(mut_mins[1] == mut_min){ return(df_run) } else { next } } # Get mutation count at all positions that are not potential SNPs pads <- paste(rep("-", min(pos_range)-1), collapse="") db_y_subset_mm$MUT_COUNT_MINUS_SUBSTRING <- db_y_subset_mm[[seq]] %>% substring(min(pos_range), max(pos_range)) %>% paste(pads, ., sep="") %>% getMutatedPositions(gl_minus_substring) %>% sapply(length) # Keep only unmutated seqences and then find the counts of J and # junction length for each of the SNP strings, and then check to # see which pass the j/junction and count requirements db_y_summary0 <- db_y_subset_mm %>% dplyr::filter(!!rlang::sym("MUT_COUNT_MINUS_SUBSTRING") == 0) df_run$UNMUTATED_COUNT[1] <- nrow(db_y_summary0) db_y_summary0 <- db_y_summary0 %>% dplyr::mutate(J_GENE = getGene(!! rlang::sym(j_call))) %>% dplyr::group_by(!!! rlang::syms(c("SNP_STRING", "J_GENE", junction_length))) %>% dplyr::summarise(COUNT = n()) df_run$UNMUTATED_SNP_J_GENE_LENGTH_COUNT[1] <- nrow(db_y_summary0) db_y_summary0 <- db_y_summary0 %>% dplyr::group_by(!!rlang::sym("SNP_STRING")) %>% dplyr::mutate(FRACTION = !! rlang::sym("COUNT")/sum(!! rlang::sym("COUNT"))) %>% dplyr::summarise(TOTAL_COUNT = sum(!! rlang::sym("COUNT")), MAX_FRAC = max(!! rlang::sym("FRACTION"))) if(nrow(db_y_summary0) < 1){ df_run$NOTE[1] <- paste("Position(s) passed y-intercept (", paste(pass_y$POSITION, collapse = ","), ") but no unmutated versions of novel allele", " found.", sep="") if(mut_mins[1] == mut_min){ return(df_run) } else { next } } # db_y_summary = db_y_summary0 %>% # filter_(~TOTAL_COUNT >= min_seqs & MAX_FRAC <= j_max) min_seqs_pass <- db_y_summary0$TOTAL_COUNT >= min_seqs j_max_pass <- db_y_summary0$MAX_FRAC <= j_max db_y_summary <- db_y_summary0[min_seqs_pass & j_max_pass, , drop=FALSE] df_run$SNP_MIN_SEQS_J_MAX_PASS[1] <- nrow(db_y_summary) if(nrow(db_y_summary) < 1){ msg <- c(NA, NA) names(msg) <- c("j_max", "min_seqs") if (sum(min_seqs_pass) == 0) { msg['min_seqs'] <- paste0("Not enough sequences (maximum total count is ", max(db_y_summary0$TOTAL_COUNT), ").") } if (sum(j_max_pass) == 0) { msg['j_max'] <- paste0("A J-junction combination is too prevalent (", round(100*max(db_y_summary0$MAX_FRAC),1),"% of sequences).") } msg <- paste(na.omit(msg), collapse=" and ") df_run$NOTE[1] <- paste("Position(s) passed y-intercept (", paste(pass_y$POSITION, collapse = ","), ") but ", msg,".", sep="") df_run$PERFECT_MATCH_COUNT[1] <- max(db_y_summary0$TOTAL_COUNT) df_run$PERFECT_MATCH_FREQ[1] <- df_run$PERFECT_MATCH_COUNT[1]/df_run$GERMLINE_CALL_COUNT[1] if(mut_mins[1] == mut_min){ return(df_run) } else { next } } germ_nts <- unlist(strsplit(gl_substring,"")) for (r in 1:nrow(db_y_summary)) { if (r > 1){ df_run <- dplyr::bind_rows(df_run[1,], df_run) } # Create the new germline snp_nts <- unlist(strsplit(db_y_summary$SNP_STRING[r],"")) remain_mut <- db_y_summary$SNP_STRING[r] %>% getMutatedPositions(gl_substring) %>% unlist() %>% unique() germ <- insertPolymorphisms(germline, pass_y$POSITION, snp_nts) is_known_allele <- germ == germlines if (sum(is_known_allele) == 0 ) { names(germ) <- mapply(paste, germ_nts[remain_mut], pass_y$POSITION[remain_mut], snp_nts[remain_mut], sep="") %>% paste(collapse="_") %>% paste(names(germline), ., sep="_") } else { # If the match is with duplicated sequences in the reference germlines, # use the first known_allele_names <- sortAlleles(names(germlines)[is_known_allele], method="position") names(germ) = known_allele_names[1] } # Save the new germline to our data frame df_run$POLYMORPHISM_CALL[1] <- names(germ) df_run$NOVEL_IMGT[1] <- as.character(germ) df_run$PERFECT_MATCH_COUNT[1] <- db_y_summary$TOTAL_COUNT[r] df_run$PERFECT_MATCH_FREQ[1] <- df_run$PERFECT_MATCH_COUNT[1]/df_run$GERMLINE_CALL_COUNT[1] df_run$NOTE[1] = "Novel allele found!" } } # end for each starting mutation counts return(df_run) } # end foreach allele if(nproc > 1) { stopCluster(cluster) } out_df <- dplyr::bind_rows(out_list) getMuSpec <- function(poly_call, germ_call) { sapply(1:length(poly_call), function(i){ p <- gsub(germ_call[i], "", poly_call[i], fixed = T) p <- strsplit(p,"_")[[1]][-1] m <- gsub("([[:alpha:]])([[:digit:]]*)([[:alpha:]])", "\\2\\1>\\3", p) paste(m, collapse=",") }) } # The number of records in the sequence dataset matching # each exact NOVEL_IMGT sequence getDbMatch <- function(novel_imgt) { novel_imgt <- stri_sub(novel_imgt, min(pos_range), max(pos_range)) novel_imgt <- stri_replace_all_regex(novel_imgt, "[-\\.]","") data_seq <- stri_sub(data[[seq]], min(pos_range), max(pos_range)) data_seq <- stri_replace_all_regex(data_seq,"[-\\.]","") sapply(novel_imgt, function(n) { # sum(grepl(n, data_seq)) sum(stri_detect_fixed(data_seq,n)) }) } # The number of distinct J in the sequence dataset associated # with the exact NOVEL_IMGT sequence getNumJ <- function(novel_imgt) { novel_imgt <- stri_sub(novel_imgt, min(pos_range), max(pos_range)) novel_imgt <- stri_replace_all_regex(novel_imgt, "[-\\.]","") data_seq <- stri_sub(data[[seq]], min(pos_range), max(pos_range)) data_seq <- stri_replace_all_regex(data_seq,"[-\\.]","") sapply(novel_imgt, function(n) { imgt_idx <- stri_detect_fixed(data_seq,n) length(unique(getGene(data[[j_call]][imgt_idx]))) }) } # The number of distinct CDR3 in the sequence dataset associated # with the exact NOVEL_IMGT sequence getNumCDR3 <- function(novel_imgt) { novel_imgt <- stri_sub(novel_imgt, min(pos_range), max(pos_range)) novel_imgt <- stri_replace_all_regex(novel_imgt, "[-\\.]","") data_seq <- stri_sub(data[[seq]], min(pos_range), max(pos_range)) data_seq <- stri_replace_all_regex(data_seq,"[-\\.]","") sapply(novel_imgt, function(n) { imgt_idx <- stri_detect_fixed(data_seq,n) seq <- data[[junction]][imgt_idx] seq <- stri_sub(seq, 4, stringi::stri_length(seq) - 3) length(unique(seq)) }) } idx <- which(!is.na(out_df$NOVEL_IMGT)) if (length(idx)>0) { out_df$NT_SUBSTITUTIONS[idx] <- getMuSpec(out_df$POLYMORPHISM_CALL[idx], out_df$GERMLINE_CALL[idx]) out_df$NOVEL_IMGT_COUNT[idx] <- getDbMatch(out_df$NOVEL_IMGT[idx]) out_df$NOVEL_IMGT_UNIQUE_J[idx] <- getNumJ(out_df$NOVEL_IMGT[idx]) if (junction %in% colnames(data)) { out_df$NOVEL_IMGT_UNIQUE_CDR3[idx] <- getNumCDR3(out_df$NOVEL_IMGT[idx]) } } out_df$GERMLINE_IMGT_COUNT <- getDbMatch(out_df$GERMLINE_IMGT) out_df$UNMUTATED_FREQ <- out_df$UNMUTATED_COUNT/out_df$GERMLINE_CALL_COUNT return(out_df) } #' Select rows containing novel alleles #' #' \code{selectNovel} takes the result from \link{findNovelAlleles} and #' selects only the rows containing unique, novel alleles. #' #' @details #' If, for instance, subject has in his genome \code{IGHV1-2*02} and a novel #' allele equally close to \code{IGHV1-2*02} and \code{IGHV1-2*05}, the novel allele may be #' detected by analyzing sequences that best align to either of these alleles. #' If \code{keep_alleles} is \code{TRUE}, both polymorphic allele calls will #' be retained. In the case that multiple mutation ranges are checked for the #' same allele, only one mutation range will be kept in the output. #' #' @param novel a \code{data.frame} of the type returned by #' \link{findNovelAlleles}. #' @param keep_alleles a \code{logical} indicating if different alleles #' leading to the same novel sequence should be kept. #' See Details. #' #' @return A \code{data.frame} containing only unique, novel alleles (if any) #' that were in the input. #' #' @examples #' novel <- selectNovel(SampleNovel) #' #' @export selectNovel <- function(novel, keep_alleles=FALSE) { # Remove non-novel rows novel <- filter(novel, !is.na(!!rlang::sym("NOVEL_IMGT"))) if (keep_alleles) { novel < novel %>% group_by(!!rlang::sym("GERMLINE_CALL")) } novel_set <- novel %>% distinct(!!rlang::sym("NOVEL_IMGT"), .keep_all=TRUE) %>% ungroup() return(novel_set) } #' Visualize evidence of novel V alleles #' #' \code{plotNovel} is be used to visualize the evidence of any novel V #' alleles found using \link{findNovelAlleles}. It can also be used to #' visualize the results for alleles that did #' #' @details #' The first panel in the plot shows, for all sequences which align to a particular #' germline allele, the mutation frequency at each postion along the aligned #' sequece as a function of the sequence-wide mutation. Sequences that pass #' the novel allele test are colored red, while sequences that don't pass #' the test are colored yellow. The second panel shows the nucleotide usage at the #' positions as a function of sequence-wide mutation count. #' #' To avoid cases where a clonal expansion might lead to a false positive, tigger examines #' the combinations of J gene and junction length among sequences which perfectly #' match the proposed germline allele. #' #' @param data a \code{data.frame} in Change-O format. See #' \link{findNovelAlleles} for details. #' @param novel_row a single row from a data frame as output by #' \link{findNovelAlleles} that contains a #' polymorphism-containing germline allele #' @param v_call name of the column in \code{data} with V allele #' calls. Default is "V_CALL". #' @param j_call name of the column in \code{data} with J allele calls. #' Default is J_CALL. #' @param seq name of the column in \code{data} with the #' aligned, IMGT-numbered, V(D)J nucleotide sequence. #' Default is SEQUENCE_IMGT. #' @param junction Junction region nucleotide sequence, which includes #' the CDR3 and the two flanking conserved codons. Default #' is JUNCTION. #' @param junction_length Number of junction nucleotides in the junction sequence. #' Default is JUNCTION_LENGTH. #' @param ncol number of columns to use when laying out the plots #' #' @examples #' # Plot the evidence for the first (and only) novel allele in the example data #' novel <- selectNovel(SampleNovel) #' plotNovel(SampleDb, novel[1, ]) #' #' @export plotNovel <- function(data, novel_row, v_call="V_CALL", j_call="J_CALL", seq="SEQUENCE_IMGT", junction="JUNCTION", junction_length="JUNCTION_LENGTH", ncol=1) { . = NULL # Use the data frame if(length(novel_row) > 0) { if(is.data.frame(novel_row) & nrow(novel_row) == 1) { pos_range <- novel_row$POS_MIN:novel_row$POS_MAX germline <- novel_row$GERMLINE_IMGT names(germline) <- novel_row$GERMLINE_CALL mut_range <- novel_row$MUT_MIN[1]:novel_row$MUT_MAX[1] novel_imgt <- novel_row$NOVEL_IMGT names(novel_imgt) <- novel_row$POLYMORPHISM_CALL min_frac <- novel_row$MIN_FRAC note <- novel_row$NOTE } else { stop("novel_row is not a data frame with only one row.") } } germline <- cleanSeqs(germline) data[[seq]] <- cleanSeqs(data[[seq]]) # Extract sequences assigned to the germline, determine which # have an appropriate range of mutations, and find the mutation # frequency of each position db_subset <- data %>% select(!!!rlang::syms(c(seq, v_call, j_call, junction_length))) %>% filter(grepl(names(germline), .data[[v_call]], fixed=TRUE)) pos_db <- db_subset %>% mutationRangeSubset(germline, mut_range, pos_range, seq=seq) if (nrow(pos_db) == 0) { warning(paste0("Insufficient sequences (",nrow(pos_db),") in desired mutational range.")) return (invisible(NULL)) } pos_db <- pos_db %>% positionMutations(germline, pos_range,seq=seq) pos_muts <- pos_db %>% group_by(!!rlang::sym("POSITION")) %>% mutate(PASS = mean(!!rlang::sym("OBSERVED")) >= min_frac) %>% group_by(!!!rlang::syms(c("MUT_COUNT", "POSITION"))) %>% summarise(POS_MUT_RATE = mean(!!rlang::sym("MUTATED"))*unique(!!rlang::sym("PASS"))) %>% ungroup() # Label the polymorphic positions as such pass_y <- unlist(strsplit(names(novel_imgt), "_"))[-1] %>% gsub("[^0-9]", "", .) %>% as.numeric() p_y_f <- unlist(strsplit(names(novel_imgt), "_"))[-1] %>% gsub("[0-9]+.", "", .) p_y_t <- unlist(strsplit(names(novel_imgt), "_"))[-1] %>% gsub(".[0-9]+", "", .) # Parse the note to find positions that passed y intercept if no novel found if (length(pass_y) == 0 & grepl("Position\\(s\\) passed y-intercept", note)) { pass_y <- note %>% gsub("Position\\(s\\) passed y-intercept \\(", "", .) %>% gsub("\\).*", "", .) %>% strsplit(",") %>% unlist %>% as.numeric p_y_f <- sapply(pass_y, function (x) substring(germline, x, x)) p_y_t <- gsub(".", "?", p_y_f) } to_from <- paste(paste("Position", pass_y), paste(paste(p_y_f, "->"), p_y_t)) names(to_from) <- pass_y pos_muts <- pos_muts %>% mutate(Polymorphic = ifelse(!!rlang::sym("POSITION") %in% pass_y, "True", "False")) pads <- paste(rep("-", min(pos_range)-1), collapse="") db_subset$MUT_COUNT_NOVEL <- db_subset[[seq]] %>% substring(min(pos_range), max(pos_range)) %>% paste(pads, ., sep="") %>% getMutatedPositions(novel_imgt) %>% sapply(length) db_subset <- db_subset %>% filter(!!rlang::sym("MUT_COUNT_NOVEL") == 0) %>% mutate(J_GENE = getGene(!!rlang::sym(j_call))) if (nrow(db_subset) == 0) { warning(paste0("Insufficient sequences (",nrow(db_subset),") with MUT_COUNT_NOVEL == 0.")) return (invisible(NULL)) } db_subset[[junction_length]] <- db_subset[[junction_length]] %>% factor(levels=min(db_subset[[junction_length]]):max(db_subset[[junction_length]])) pos_muts$Polymorphic <- pos_muts$Polymorphic %>% factor(levels = c("False", "True")) pos_db$NT <- pos_db$NT %>% factor(levels = names(DNA_COLORS)) pos_muts$GERMLINE <- names(germline) # MAKE THE FIRST PLOT if (!is.na(novel_imgt)) { POLYCOLORS <- setNames(DNA_COLORS[c(4,3)], c("False", "True")) p1 <- ggplot(pos_muts, aes_string(x="MUT_COUNT", y="POS_MUT_RATE", group="POSITION", color="Polymorphic")) + geom_line(data=filter(pos_muts, !!rlang::sym("Polymorphic") == "False"), size=0.75) + geom_line(data=filter(pos_muts, !!rlang::sym("Polymorphic") == "True"), size=0.75) + facet_grid(GERMLINE ~ .) + scale_color_manual(values = POLYCOLORS) + ylim(0,1) + xlab("Mutation Count (Sequence)") + ylab("Mutation Frequency (Position)") + theme_bw() + theme(legend.position=c(0.5,0.9), legend.justification=c(0.5,1), legend.background=element_rect(fill = "transparent")) + guides(color = guide_legend(ncol = 2, reverse = TRUE)) } else{ POLYCOLORS = setNames(DNA_COLORS[c(4,2)], c("False", "True")) p1 <- ggplot(pos_muts, aes_string(x="MUT_COUNT", y="POS_MUT_RATE", group="POSITION", color="Polymorphic")) + geom_line(size=0.75) + facet_grid(GERMLINE ~ .) + scale_color_manual(values=POLYCOLORS) + ylim(0, 1) + xlab("Mutation Count (Sequence)") + ylab("Mutation Frequency (Position)") + theme_bw() + theme(legend.position=c(0.5, 0.9), legend.justification=c(0.5, 1), legend.background=element_rect(fill="transparent")) + guides(color = guide_legend("Passed y-intercept test", ncol = 2, reverse = TRUE)) } # MAKE THE SECOND PLOT p2_data <- mutate(filter(pos_db, !!rlang::sym("POSITION") %in% pass_y), POSITION = to_from[as.character(!!rlang::sym("POSITION"))]) if (nrow(p2_data)) { p2 <- ggplot(p2_data, aes_string(x="MUT_COUNT", fill="NT")) + geom_bar(width=0.9) + guides(fill = guide_legend("Nucleotide", ncol=4)) + facet_grid(POSITION ~ .) + xlab("Mutation Count (Sequence)") + ylab("Sequence Count") + scale_fill_manual(values=DNA_COLORS, breaks=names(DNA_COLORS), drop=FALSE) + theme_bw() + theme(legend.position=c(1,1), legend.justification=c(1,1), legend.background=element_rect(fill="transparent")) } else { p2_data <- mutate(filter(pos_db, !!rlang::sym("POSITION") %in% names(which.max(table(pos_db$POSITION)))), POSITION = "No positions pass y-intercept test.") p2 <- ggplot(p2_data, aes_string(x="MUT_COUNT")) + geom_bar(width=0.9) + facet_grid(POSITION ~ .) + xlab("Mutation Count (Sequence)") + ylab("Sequence Count") + theme_bw() + theme(legend.position=c(1,1), legend.justification=c(1,1), legend.background=element_rect(fill = "transparent")) } # MAKE THE THIRD PLOT p3 <- ggplot(db_subset, aes_string(x=junction_length, fill="J_GENE")) + geom_bar(width=0.9) + guides(fill=guide_legend("J Gene", ncol=2)) + xlab("Junction Length") + ylab("Unmutated Sequence Count") + theme_bw() + theme(legend.position=c(1, 1), legend.justification=c(1, 1), legend.background=element_rect(fill="transparent")) p2_height <- length(unique(p2_data$POSITION)) if (p2_height>1) { p2_height = 0.5 * p2_height} heights <- c(1, p2_height, 1) multiplot(p1, p2, p3, cols = ncol, heights=heights) } #' Infer a subject-specific genotype using a frequency method #' #' \code{inferGenotype} infers an subject's genotype using a frequency method. #' The genotype is inferred by finding the minimum number set of alleles that #' can explain the majority of each gene's calls. The most common allele of #' each gene is included in the genotype first, and the next most common allele #' is added until the desired fraction of alleles can be explained. In this #' way, mistaken allele calls (resulting from sequences which #' by chance have been mutated to look like another allele) can be removed. #' #' @details #' Allele calls representing cases where multiple alleles have been #' assigned to a single sample sequence are rare among unmutated #' sequences but may result if nucleotides for certain positions are #' not available. Calls containing multiple alleles are treated as #' belonging to all groups. If \code{novel} is provided, all #' sequences that are assigned to the same starting allele as any #' novel germline allele will have the novel germline allele appended #' to their assignent prior to searching for unmutated sequences. #' #' @param data a \code{data.frame} containing V allele #' calls from a single subject. If #' \code{find_unmutated} is \code{TRUE}, then #' the sample IMGT-gapped V(D)J sequence should #' @param germline_db named vector of sequences containing the #' germline sequences named in #' \code{allele_calls}. Only required if #' \code{find_unmutated} is \code{TRUE}. #' @param novel an optional \code{data.frame} of the type #' novel returned by #' \link{findNovelAlleles} containing #' germline sequences that will be utilized if #' \code{find_unmutated} is \code{TRUE}. See #' Details. #' @param v_call column in \code{data} with V allele calls. #' Default is \code{"V_CALL"}. #' @param seq name of the column in \code{data} with the #' aligned, IMGT-numbered, V(D)J nucleotide sequence. #' Default is SEQUENCE_IMGT. #' @param fraction_to_explain the portion of each gene that must be #' explained by the alleles that will be included #' in the genotype. #' @param gene_cutoff either a number of sequences or a fraction of #' the length of \code{allele_calls} denoting the #' minimum number of times a gene must be #' observed in \code{allele_calls} to be included #' in the genotype. #' @param find_unmutated if \code{TRUE}, use \code{germline_db} to #' find which samples are unmutated. Not needed #' if \code{allele_calls} only represent #' unmutated samples. #' #' @return #' A \code{data.frame} of alleles denoting the genotype of the subject containing #' the following columns: #' #' \itemize{ #' \item \code{GENE}: The gene name without allele. #' \item \code{ALLELES}: Comma separated list of alleles for the given \code{GENE}. #' \item \code{COUNTS}: Comma separated list of observed sequences for each #' corresponding allele in the \code{ALLELES} list. #' \item \code{TOTAL}: The total count of observed sequences for the given \code{GENE}. #' \item \code{NOTE}: Any comments on the inferrence. #' } #' #' @note #' This method works best with data derived from blood, where a large #' portion of sequences are expected to be unmutated. Ideally, there #' should be hundreds of allele calls per gene in the input. #' #' @seealso \link{plotGenotype} for a colorful visualization and #' \link{genotypeFasta} to convert the genotype to nucleotide sequences. #' See \link{inferGenotypeBayesian} to infer a subject-specific genotype #' using a Bayesian approach. #' #' @examples #' # Infer IGHV genotype, using only unmutated sequences, including novel alleles #' inferGenotype(SampleDb, germline_db=SampleGermlineIGHV, novel=SampleNovel, #' find_unmutated=TRUE) #' #' @export inferGenotype <- function(data, germline_db=NA, novel=NA, v_call="V_CALL", seq="SEQUENCE_IMGT", fraction_to_explain=0.875, gene_cutoff=1e-4, find_unmutated=TRUE) { . = NULL allele_calls = getAllele(data[[v_call]], first=FALSE, strip_d=FALSE) # Find the unmutated subset, if requested if(find_unmutated){ if(is.na(germline_db[1])){ stop("germline_db needed if find_unmutated is TRUE") } if(!is.null(nrow(novel))){ novel <- filter(novel, !is.na(!!rlang::sym("POLYMORPHISM_CALL"))) %>% select(!!!rlang::syms(c("GERMLINE_CALL", "POLYMORPHISM_CALL", "NOVEL_IMGT"))) if(nrow(novel) > 0){ # Extract novel alleles if any and add them to germline_db novel_gl <- novel$NOVEL_IMGT names(novel_gl) <- novel$POLYMORPHISM_CALL germline_db <- c(germline_db, novel_gl) # Add the novel allele calls to allele calls of the same starting allele for(r in 1:nrow(novel)){ ind <- grep(novel$GERMLINE_CALL[r], allele_calls, fixed=TRUE) allele_calls[ind] <- allele_calls[ind] %>% sapply(paste, novel$POLYMORPHISM_CALL[r], sep=",") } } } # Find unmutated sequences allele_calls <- findUnmutatedCalls(allele_calls, as.character(data[[seq]]), germline_db) if(length(allele_calls) == 0){ stop("No unmutated sequences found! Set 'find_unmutated' to 'FALSE'.") } } # Find which rows' calls contain which genes cutoff <- ifelse(gene_cutoff < 1, length(allele_calls)*gene_cutoff, gene_cutoff) gene_regex <- allele_calls %>% strsplit(",") %>% unlist() %>% getGene(strip_d=FALSE) %>% unique() %>% paste("\\*", sep="") gene_groups <- sapply(gene_regex, grep, allele_calls, simplify=FALSE) names(gene_groups) <- gsub("\\*", "", gene_regex, fixed=TRUE) gene_groups <- gene_groups[sapply(gene_groups, length) >= cutoff] gene_groups <- gene_groups[sortAlleles(names(gene_groups))] # Make a table to store the resulting genotype GENE <- names(gene_groups) ALLELES <- COUNTS <- NOTE <- rep("", length(GENE)) TOTAL <- sapply(gene_groups, length) genotype <- cbind(GENE, ALLELES, COUNTS, TOTAL, NOTE) # For each gene, find which alleles to include for (g in GENE) { # Keep only the part of the allele calls that uses the gene being analyzed ac <- allele_calls[gene_groups[[g]]] %>% strsplit(",") %>% lapply(function(x) x[grep(paste(g, "\\*", sep=""), x)]) %>% sapply(paste, collapse=",") target <- ceiling(fraction_to_explain*length(ac)) # how many we need to explain t_ac <- table(ac) # table of allele calls potentials <- unique(unlist(strsplit(names(t_ac),","))) # potential alleles # One allele? Easy! if (length(potentials) == 1 | length(t_ac) == 1) { genotype[genotype[,"GENE"]==g,"ALLELES"] <- gsub("[^d\\*]*[d\\*]","",potentials )[1] genotype[genotype[,"GENE"]==g,"COUNTS"] <- t_ac } else { # More alleles? Let's find the fewest that can explain the needed fraction # Make a table of which alleles can explain which calls regexpotentials <- paste(gsub("\\*","\\\\*", potentials),"$",sep="") regexpotentials <- paste(regexpotentials,gsub("\\$",",",regexpotentials),sep="|") tmat = sapply(regexpotentials, function(x) grepl(x, names(t_ac),fixed=FALSE)) seqs_expl = as.data.frame(apply(tmat, 2, function(x) x*t_ac)) colnames(seqs_expl) = potentials # Cycle through the table, including alleles to explain more sequences, # until we explain enough sequences included <- counts <- character(0) tot_expl = 0 while(tot_expl < target){ allele_tot <- apply(seqs_expl, 2, sum) included <- c(included, names(which.max(allele_tot))) counts <- c(counts, max(allele_tot)) tot_expl <- max(allele_tot) + tot_expl seqs_expl <- seqs_expl[which(seqs_expl[,which.max(allele_tot)]==0),] } genotype[genotype[,"GENE"]==g,"ALLELES"] <- paste(gsub("[^d\\*]*[d\\*]","",included ),collapse=",") genotype[genotype[,"GENE"]==g,"COUNTS"] <- paste(counts,collapse=",") } } geno <- as.data.frame(genotype, stringsAsFactors = FALSE) # Check for indistinguishable calls if (find_unmutated == TRUE) { seqs <- genotypeFasta(geno, germline_db) dist_mat <- seqs %>% sapply(function(x) sapply((getMutatedPositions(seqs, x)), length)) %>% as.matrix rownames(dist_mat) <- colnames(dist_mat) for (i in 1:nrow(dist_mat)){ dist_mat[i,i] = NA } same <- which(dist_mat == 0, arr.ind=TRUE) if (nrow(same) > 0 ) { for (r in 1:nrow(same)) { inds <- as.vector(same[r,]) geno[getGene(rownames(dist_mat)[inds][1]),]$NOTE <- paste(rownames(dist_mat)[inds], collapse=" and ") %>% paste("Cannot distinguish", .) } } } rownames(geno) <- NULL return(geno) } #' Show a colorful representation of a genotype #' #' \code{plotGenotype} plots a genotype table. #' #' @param genotype a \code{data.frame} of alleles denoting a genotype, #' as returned by \link{inferGenotype}. #' @param facet_by a column name in \code{genotype} to facet the plot by. #' If \code{NULL}, then do not facet the plot. #' @param gene_sort a string defining the method to use when sorting alleles. #' If \code{"name"} then sort in lexicographic order. If #' \code{"position"} then sort by position in the locus, as #' determined by the final two numbers in the gene name. #' @param text_size the point size of the plotted text. #' @param silent if \code{TRUE} do not draw the plot and just return the ggplot #' object; if \code{FALSE} draw the plot. #' @param ... additional arguments to pass to ggplot2::theme. #' #' @return A ggplot object defining the plot. #' #' @seealso \link{inferGenotype} #' #' @examples #' # Plot genotype #' plotGenotype(SampleGenotype) #' #' # Facet by subject #' genotype_a <- genotype_b <- SampleGenotype #' genotype_a$SUBJECT <- "A" #' genotype_b$SUBJECT <- "B" #' geno_sub <- rbind(genotype_a, genotype_b) #' plotGenotype(geno_sub, facet_by="SUBJECT", gene_sort="pos") #' #' @export plotGenotype <- function(genotype, facet_by=NULL, gene_sort=c("name", "position"), text_size=12, silent=FALSE, ...) { # Check arguments gene_sort <- match.arg(gene_sort) # Split genes' alleles into their own rows alleles = strsplit(genotype$ALLELES, ",") geno2 = genotype r = 1 for (g in 1:nrow(genotype)){ for(a in 1:length(alleles[[g]])) { geno2[r, ] = genotype[g, ] geno2[r, ]$ALLELES = alleles[[g]][a] r = r + 1 } } # Set the gene order geno2$GENE = factor(geno2$GENE, levels=rev(sortAlleles(unique(geno2$GENE), method=gene_sort))) # Create the base plot p = ggplot(geno2, aes_string(x="GENE", fill="ALLELES")) + theme_bw() + theme(axis.ticks=element_blank(), axis.text.x=element_blank(), panel.grid.major=element_blank(), panel.grid.minor=element_blank(), text=element_text(size=text_size), strip.background=element_blank(), strip.text=element_text(face="bold")) + geom_bar(position="fill") + coord_flip() + xlab("Gene") + ylab("") + scale_fill_hue(name="Allele", h=c(0, 270), h.start=10) # Plot, with facets by SUBJECT if that column is present if (!is.null(facet_by)) { p = p + facet_grid(paste0(".~", facet_by)) } # Add additional theme elements p = p + do.call(theme, list(...)) # Plot if (!silent) { plot(p) } invisible(p) } #' Return the nucleotide sequences of a genotype #' #' \code{genotypeFasta} converts a genotype table into a vector of nucleotide #' sequences. #' #' @param genotype a \code{data.frame} of alleles denoting a genotype, #' as returned by \link{inferGenotype}. #' @param germline_db a vector of named nucleotide germline sequences #' matching the alleles detailed in \code{genotype}. #' @param novel an optional \code{data.frame} containing putative #' novel alleeles of the type returned by #' \link{findNovelAlleles}. #' #' @return A named vector of strings containing the germline nucleotide #' sequences of the alleles in the provided genotype. #' #' @seealso \link{inferGenotype} #' #' @examples #' # Find the sequences that correspond to the genotype #' genotype_db <- genotypeFasta(SampleGenotype, SampleGermlineIGHV, SampleNovel) #' #' @export genotypeFasta <- function(genotype, germline_db, novel=NA){ if(!is.null(nrow(novel))){ # Extract novel alleles if any and add them to germline_db novel <- filter(novel, !is.na(!!rlang::sym("POLYMORPHISM_CALL"))) %>% select(!!!rlang::syms(c("GERMLINE_CALL", "POLYMORPHISM_CALL", "NOVEL_IMGT"))) if(nrow(novel) > 0){ novel_gl <- novel$NOVEL_IMGT names(novel_gl) <- novel$POLYMORPHISM_CALL germline_db <- c(germline_db, novel_gl) } } genotype$GENE <- gsub("[Dd]\\*","*",genotype$GENE) g_names <- names(germline_db) names(g_names) <- gsub("[Dd]\\*", "*", names(germline_db)) table_calls <- mapply(paste, genotype$GENE, strsplit(genotype$ALLELES, ","), sep="*") seqs <- germline_db[as.vector(g_names[unlist(table_calls)])] if(sum(is.na(seqs)) > 0){ stop("The following genotype alleles were not found in germline_db: ", paste(unlist(table_calls)[which(is.na(seqs))], collapse = ", ")) } return(seqs) } #' Correct allele calls based on a personalized genotype #' #' \code{reassignAlleles} uses a subject-specific genotype to correct #' correct preliminary allele assignments of a set of sequences derived #' from a single subject. #' #' @details #' In order to save time, initial gene assignments are preserved and #' the allele calls are chosen from among those provided in \code{genotype_db}, #' based on a simple alignment to the sample sequence. #' #' @param data a \code{data.frame} containing V allele calls from a #' single subject and the sample IMGT-gapped V(D)J sequences under #' \code{"SEQUENCE_IMGT"}. #' @param genotype_db a vector of named nucleotide germline sequences #' matching the calls detailed in \code{allele_calls} #' and personalized to the subject #' @param v_call name of the column in \code{data} with V allele #' calls. Default is \code{"V_CALL"}. #' @param seq name of the column in \code{data} with the #' aligned, IMGT-numbered, V(D)J nucleotide sequence. #' Default is SEQUENCE_IMGT #' @param method the method to be used when realigning sequences to #' the genotype_db sequences. Currently, only \code{"hammming"} #' (for Hamming distance) is implemented. #' @param path directory containing the tool used in the #' realignment method, if needed. Hamming distance does #' not require a path to a tool. #' @param keep_gene a string indicating if the gene (\code{"gene"}), #' family (\code{"family"}) or complete repertoire #' (\code{"repertoire"}) assignments should be performed. #' Use of \code{"gene"} increases speed by minimizing required number of #' alignments, as gene level assignments will be maintained when possible. #' #' @return A modifed input \code{data.frame} containing the best allele call from #' among the sequences listed in \code{genotype_db} in the #' \code{V_CALL_GENOTYPED} column. #' #' @examples #' # Extract the database sequences that correspond to the genotype #' genotype_db <- genotypeFasta(SampleGenotype, SampleGermlineIGHV, novel=SampleNovel) #' #' # Use the personlized genotype to determine corrected allele assignments #' output_db <- reassignAlleles(SampleDb, genotype_db) #' #' @export reassignAlleles <- function(data, genotype_db, v_call="V_CALL", seq="SEQUENCE_IMGT", method="hamming", path=NA, keep_gene=c("gene", "family", "repertoire")){ # Check arguments keep_gene <- match.arg(keep_gene) # Extract data subset and prepare output vector v_sequences = as.character(data[[seq]]) v_calls = getAllele(data[[v_call]], first=FALSE, strip_d=FALSE) v_call_genotyped = rep("", length(v_calls)) if (keep_gene == "gene") { v = getGene(v_calls, first = TRUE, strip_d=FALSE) geno = getGene(names(genotype_db),strip_d=TRUE) names(geno) = names(genotype_db) } else if (keep_gene == "family") { v <- getFamily(v_calls, first = TRUE, strip_d = FALSE) geno = getFamily(names(genotype_db),strip_d=TRUE) names(geno) = names(genotype_db) } else if (keep_gene == "repertoire") { v <- rep(v_call, length(v_calls)) geno = rep(v_call, length(genotype_db)) names(geno) = names(genotype_db) } else { stop("Unknown keep_gene value: ", keep_gene) } # keep_gene == FALSE # Find which genotype genes/families are homozygous and assign those alleles first hetero = unique(geno[which(duplicated(geno))]) homo = geno[!(geno %in% hetero)] homo_alleles = names(homo) names(homo_alleles) = homo homo_calls_i = which(v %in% homo) v_call_genotyped[homo_calls_i] = homo_alleles[v[homo_calls_i]] # Now realign the heterozygote sequences to each allele of that gene for (het in hetero){ ind = which(v %in% het) if (length(ind) > 0){ het_alleles = names(geno[which(geno == het)]) het_seqs = genotype_db[het_alleles] if(method == "hamming"){ dists = lapply(het_seqs, function(x) sapply(getMutatedPositions(v_sequences[ind], x, match_instead=FALSE), length)) dist_mat = matrix(unlist(dists), ncol = length(het_seqs)) } else { stop("Only Hamming distance is currently supported as a method.") } # The sapply-apply approach could become problematic when nrow(dist_mat) # is 1 and min(best_match) has multiple values, due to the fact that R # does not always keep data structures unmutable # Explicitly specifying a list and subsequently keeping it as a list by # using lapply avoids that problem best_match = vector("list", length=nrow(dist_mat)) for (i in 1:nrow(dist_mat)) { best_match[[i]] = which(dist_mat[i, ]==min(dist_mat[i, ])) } best_alleles = lapply(best_match, function(x) het_alleles[x]) v_call_genotyped[ind] = unlist(lapply(best_alleles, paste, collapse=",")) } } # Now realign the gene-not-in-genotype calls to every genotype allele hetero_calls_i = which(v %in% hetero) not_called = setdiff(1:length(v), c(homo_calls_i, hetero_calls_i)) if(length(not_called)>1){ if(method == "hamming"){ dists = lapply(genotype_db, function(x) sapply(getMutatedPositions(v_sequences[not_called], x, match_instead=FALSE), length)) dist_mat = matrix(unlist(dists), ncol = length(genotype_db)) } else { stop("Only Hamming distance is currently supported as a method.") } # The sapply-apply approach could become problematic when nrow(dist_mat) # is 1 and min(best_match) has multiple values, due to the fact that R # does not always keep data structures unmutable # Explicitly specifying a list and subsequently keeping it as a list by # using lapply avoids that problem best_match = vector("list", length=nrow(dist_mat)) for (i in 1:nrow(dist_mat)) { best_match[[i]] = which(dist_mat[i, ]==min(dist_mat[i, ])) } best_alleles = lapply(best_match, function(x) names(genotype_db[x])) v_call_genotyped[not_called] = unlist(lapply(best_alleles, paste, collapse=",")) } if (all(v_call_genotyped == data[[v_call]])) { msg <- ("No allele assignment corrections made.") if (all(v %in% homo) & length(hetero) > 0) { keep_opt <- eval(formals(reassignAlleles)$keep_gene) i <- match(keep_gene, keep_opt) rec_opt <- paste(keep_opt[(i+1):length(keep_opt)], collapse = ", ") msg <- paste(msg, "Consider setting keep_gene to one of:", rec_opt) } warning(msg) } data$V_CALL_GENOTYPED <- v_call_genotyped return(data) } # Other Mutation-Related Functions ---------------------------------------- #' Find the location of mutations in a sequence #' #' \code{getMutatedPositions} takes two vectors of aligned sequences and #' compares pairs of sequences. It returns a list of the nucleotide positions of #' any differences. #' #' @param samples a vector of strings respresenting aligned sequences #' @param germlines a vector of strings respresenting aligned sequences #' to which \code{samples} will be compared. If only #' one string is submitted, it will be used for all #' \code{samples}. #' @param ignored_regex a regular expression indicating what characters #' should be ignored (such as gaps and N nucleotides). #' @param match_instead if \code{TRUE}, the function returns the positions #' that are the same instead of those that are #' different. #' @return A list of the nucleotide positions of any differences between the #' input vectors. #' #' @examples #' # Create strings to act as a sample sequences and a reference sequence #' seqs <- c("----GATA", "GAGAGAGA", "TANA") #' ref <- "GATAGATA" #' #' # Find the differences between the two #' getMutatedPositions(seqs, ref) #' #' @export getMutatedPositions <- function(samples, germlines, ignored_regex="[\\.N-]", match_instead=FALSE) { # If only one germline sequence is given, use it for all the sample seqs if(length(germlines) == 1){ germlines = rep(germlines, length(samples)) } if(length(samples) != length(germlines)) { stop("Number of input sequences does not match number of germlines.") } # Truncate each pair of sequences to the length of the shorter germ_mins = lapply(germlines, nchar) samp_mins = lapply(samples, nchar) min_lens = mapply(min, germ_mins, samp_mins) germ = toupper(mapply(substr, germlines, 1, min_lens, SIMPLIFY=FALSE)) samp = toupper(mapply(substr, samples, 1, min_lens, SIMPLIFY=FALSE)) # Calculate poisitions of mutations (or matches), ignoring gaps, Ns, and CDR3 samp_char = strsplit(samp,"") germ_char = strsplit(germ,"") if(!match_instead){ muts = lapply(mapply("!=", samp_char, germ_char, SIMPLIFY=FALSE), which) } else { muts = lapply(mapply("==", samp_char, germ_char, SIMPLIFY=FALSE), which) } ignore_germ = gregexpr(ignored_regex, germ) ignore_samp = gregexpr(ignored_regex, samp) ignore = mapply(c, ignore_germ, ignore_samp, SIMPLIFY=FALSE) muts = mapply(function(x, y) x[!x%in%y], muts, ignore, SIMPLIFY=FALSE) return(muts) } #' Determine the mutation counts from allele calls #' #' \code{getMutCount} takes a set of nucleotide sequences and their allele calls #' and determines the distance between that seqeunce and any germline alleles #' contained within the call #' #' @param samples a vector of IMGT-gapped sample V sequences #' @param allele_calls a vector of strings respresenting Ig allele calls for #' the sequences in \code{samples}, where multiple #' calls are separated by a comma #' @param germline_db a vector of named nucleotide germline sequences #' matching the calls detailed in \code{allele_calls} #' #' @return A list equal in length to \code{samples}, containing the Hamming #' distance to each germline allele contained within each call within #' each element of \code{samples} #' #' @examples #' # Insert a mutation into a germline sequence #' s2 <- s3 <- SampleGermlineIGHV[1] #' stringi::stri_sub(s2, 103, 103) <- "G" #' stringi::stri_sub(s3, 107, 107) <- "C" #' #' sample_seqs <- c(SampleGermlineIGHV[2], s2, s3) #' #' # Pretend that one sample sequence has received an ambiguous allele call #' sample_alleles <- c(paste(names(SampleGermlineIGHV[1:2]), collapse=","), #' names(SampleGermlineIGHV[2]), #' names(SampleGermlineIGHV[1])) #' #' # Compare each sequence to its assigned germline(s) to determine the distance #' getMutCount(sample_seqs, sample_alleles, SampleGermlineIGHV) #' #' @export getMutCount <- function(samples, allele_calls, germline_db){ call_list = strsplit(allele_calls, ",") germline_list = lapply(call_list, function(x) germline_db[x]) mut_pos_list = list() mut_count_list = list() # First, find mutations of all sequences with call count of 1 call_count = sapply(germline_list, length) cc1 = which(call_count == 1) if (length(cc1) > 0) { mut_pos_list[cc1] = getMutatedPositions(samples[cc1], unlist(germline_list[cc1])) mut_count_list[cc1] = lapply(mut_pos_list[cc1], length) } # Then find mutations of all sequences with call count > 1 ccm = which(call_count > 1) if (length(ccm) > 0){ mut_pos_list[ccm] = mapply(getMutatedPositions, germline_list[ccm], samples[ccm], SIMPLIFY=FALSE) mut_count_list[ccm] = lapply(mut_pos_list[ccm], function(x) lapply(x,length)) } return(mut_count_list) } #' Determine which calls represent an unmutated allele #' #' \code{findUnmutatedCalls} determines which allele calls would represent a #' perfect match with the germline sequence, given a vector of allele calls and #' mutation counts. In the case of multiple alleles being assigned to a #' sequence, only the subset that would represent a perfect match is returned. #' #' @param allele_calls a vector of strings respresenting Ig allele calls, #' where multiple calls are separated by a comma. #' @param germline_db a vector of named nucleotide germline sequences #' @param sample_seqs V(D)J-rearranged sample sequences matching the order #' of the given \code{allele_calls}. #' #' @return A vector of strings containing the members of \code{allele_calls} #' that represent unmutated sequences. #' #' @examples #' # Find which of the sample alleles are unmutated #' calls <- findUnmutatedCalls(SampleDb$V_CALL, SampleDb$SEQUENCE_IMGT, #' germline_db=SampleGermlineIGHV) #' #' @export findUnmutatedCalls <- function(allele_calls, sample_seqs, germline_db){ . = NULL allele_calls = getAllele(allele_calls, first = FALSE) sample_seqs = as.character(sample_seqs) # Remove calls not in germline_db not_in_db = allele_calls %>% strsplit(",") %>% unlist %>% setdiff(names(germline_db)) no_call = which(allele_calls == "") in_db = not_in_db %>% sapply(grep, allele_calls, fixed=TRUE) %>% unlist() %>% c(no_call) %>% unique() %>% setdiff(1:length(allele_calls), .) allele_calls = allele_calls[in_db] sample_seqs = sample_seqs[in_db] mut_counts = getMutCount(sample_seqs, allele_calls, germline_db) # Find which seqs are unmutated and which of the allele calls that represents unmut_i = which(sapply(mut_counts, function(x) min(unlist(x))) == 0) which_no_muts = sapply(mut_counts, function(x) grep("^0$", unlist(x)) ) unmut_alleles = rep("", length(allele_calls)) # How many alleles represent perfect matches? n_gl_unmut = sapply(which_no_muts, length) one_unmut = which(n_gl_unmut == 1) split_names = strsplit(allele_calls, ",") if (length(one_unmut) > 0){ inds = unlist(which_no_muts[one_unmut]) unmut_alleles[one_unmut] = mapply("[", split_names[one_unmut], inds) } more_unmut = which(n_gl_unmut > 1) if (length(more_unmut) > 0){ inds = which_no_muts[more_unmut] unmut_multi = mapply(function(x,y) x[unlist(y)], split_names[more_unmut], inds, SIMPLIFY = FALSE) unmut_alleles[more_unmut] = sapply(unmut_multi, paste, collapse=",") } unmut_alleles = unmut_alleles[unmut_i] return(unmut_alleles) } #' Find mutation counts for frequency sequences #' #' \code{getPopularMutationCount} determines which sequences occur frequently #' for each V gene and returns the mutation count of those sequences. #' #' @param data a \code{data.frame} in the Change-O format. See #' \link{findNovelAlleles} for a list of required #' columns. #' @param germline_db A named list of IMGT-gapped germline sequences. #' @param v_call name of the column in \code{data} with V allele calls. #' Default is V_CALL. #' @param seq name of the column in \code{data} with the #' aligned, IMGT-numbered, V(D)J nucleotide sequence. #' Default is SEQUENCE_IMG #' @param gene_min The portion of all unique sequences a gene must #' constitute to avoid exclusion. #' @param seq_min The number of copies of the V that must be present for #' to avoid exclusion. #' @param seq_p_of_max For each gene, fraction of the most common V sequence's #' count that a sequence must meet to avoid exclusion. #' @param full_return If \code{TRUE}, will return all \code{data} columns and #' will include sequences with mutation count < 1. #' #' @return A data frame of genes that have a frequent sequence mutation count #' above 1. #' #' @seealso \link{getMutatedPositions} can be used to find which positions #' of a set of sequences are mutated. #' #' @examples #' getPopularMutationCount(SampleDb, SampleGermlineIGHV) #' #' @export getPopularMutationCount <- function(data, germline_db, v_call="V_CALL", seq="SEQUENCE_IMGT", gene_min = 1e-03, seq_min = 50, seq_p_of_max = 1/8, full_return = FALSE){ modified_db <- data %>% mutate(V_GENE = getGene(!!rlang::sym(v_call))) %>% group_by(!!rlang::sym("V_GENE")) %>% mutate(V_GENE_N = n()) %>% group_by(1:n()) %>% mutate(V_SEQUENCE_IMGT = substring(!!rlang::sym(seq), 1, 312)) %>% # Count occurence of each unique IMGT-gapped V sequence group_by(!!!rlang::syms(c("V_GENE", "V_SEQUENCE_IMGT"))) %>% mutate(V_SEQUENCE_IMGT_N = n()) %>% # Determine count of most common sequence group_by(!!rlang::sym("V_GENE")) %>% mutate(V_SEQUENCE_IMGT_N_MAX = max(!!rlang::sym("V_SEQUENCE_IMGT_N"))) %>% # Remove rare V genes, rare sequences, and sequences not making up a # sufficient proportion of sequences as compared to the most common ungroup %>% distinct(!!rlang::sym("V_SEQUENCE_IMGT"), .keep_all = TRUE) %>% filter(!!rlang::sym("V_GENE_N") >= (nrow(data)*gene_min)) %>% filter(!!rlang::sym("V_SEQUENCE_IMGT_N") >= seq_min) %>% mutate(V_SEQUENCE_IMGT_P_MAX = !!rlang::sym("V_SEQUENCE_IMGT_N")/!!rlang::sym("V_SEQUENCE_IMGT_N_MAX")) %>% filter(!!rlang::sym("V_SEQUENCE_IMGT_P_MAX") >= seq_p_of_max) # Determine the mutation counts of the V sequences and append them to the db MUTATION_COUNT <- getMutCount(modified_db$V_SEQUENCE_IMGT, modified_db[[v_call]], germline_db) %>% sapply(function(x) min(unlist(x))) if (length(MUTATION_COUNT)==0){ MUTATION_COUNT <- integer(0) } merged_db <- bind_cols(modified_db, data.frame(MUTATION_COUNT)) # Strip down the data frame before returning it if (!full_return) { merged_db <- merged_db %>% filter(MUTATION_COUNT > 0) %>% select(!!!rlang::syms(c("V_GENE", "MUTATION_COUNT"))) } return(merged_db) } #' Insert polymorphisms into a nucleotide sequence #' #' \code{insertPolymorphisms} replaces nucleotides in the desired locations of a #' provided sequence. #' #' @param sequence starting nucletide sequence. #' @param positions numeric vector of positions which to be changed. #' @param nucleotides character vector of nucletides to which to change the #' positions. #' #' @return A sequence with the desired nucleotides in the provided locations. #' #' @examples #' insertPolymorphisms("HUGGED", c(1, 6, 2), c("T", "R", "I")) #' #' @export insertPolymorphisms <- function(sequence, positions, nucleotides) { if(length(positions) != length(nucleotides)){ stop("Number of nucleotides and number of positions do not match.") } names(positions) = nucleotides for (i in 1:length(positions)){ substr(sequence, positions[i], positions[i]) = names(positions[i]) } return(sequence) } # Formatting and Cleanup -------------------------------------------------- #' Read immunoglobulin sequences #' #' \code{readIgFasta} reads a fasta-formatted file of immunoglobulin (Ig) #' sequences and returns a named vector of those sequences. #' #' @param fasta_file fasta-formatted file of immunoglobuling sequences. #' @param strip_down_name if \code{TRUE}, will extract only the allele name #' from the strings fasta file's sequence names. #' @param force_caps if \code{TRUE}, will force nucleotides to #' uppercase. #' #' @return Named vector of strings respresenting Ig alleles. #' #' @seealso \link{writeFasta} to do the inverse. #' #' @export readIgFasta <- function(fasta_file, strip_down_name=TRUE, force_caps=TRUE) { all_char = readChar(fasta_file, file.info(fasta_file)$size) split_by_sequence = strsplit(all_char, "[ \t\r\n\v\f]?>") add_name_break = sapply(split_by_sequence, function(x) sub("[\r\n]",">",x)) cleaned_up = sapply(add_name_break, function(x) gsub("[ \t\r\n\v\f]", "", x)) broken_names = sapply(cleaned_up, strsplit, ">") seqs = sapply(broken_names, "[", 2) seq_names = sapply(broken_names, "[", 1) if(force_caps) { seqs = toupper(seqs) } if(strip_down_name){ seq_names = getAllele(seq_names, strip_d=FALSE) } names(seqs) = seq_names return(seqs[which(!is.na(seqs))]) } #' Write to a fasta file #' #' \code{writeFasta} writes a named vector of sequences to a file in fasta #' format. #' #' @param named_sequences a vector of named string representing sequences #' @param file the name of the output file. #' @param width the number of characters to be printed per line. #' if not between 1 and 255, width with be infinite. #' @param append \code{logical} indicating if the output should be #' appended to \code{file} instead of overwriting it #' #' @return A named vector of strings respresenting Ig alleles. #' #' @seealso \link{readIgFasta} to do the inverse. #' #' @export writeFasta <- function(named_sequences, file, width=60, append=FALSE){ . = NULL seq_names = names(named_sequences) %>% paste(">", ., "\n", sep="") seqs = as.character(named_sequences) if(is.numeric(width) & width > 0 & width < 256){ width_regex = paste("(.{", width, ",", width, "})", sep="") seqs = gsub(width_regex, "\\1\n", seqs) } seqs = seqs %>% paste("\n", sep="") %>% gsub("\n\n", "\n", .) paste(seq_names, seqs, sep="", collapse="") %>% cat(file=file, append=append) } #' Update IGHV allele names #' #' \code{updateAlleleNames} takes a set of IGHV allele calls and replaces any #' outdated names (e.g. IGHV1-f) with the new IMGT names. #' #' @param allele_calls a vector of strings respresenting IGHV allele names. #' #' @return Vector of strings respresenting updated IGHV allele names. #' #' @note #' IGMT has removed \code{IGHV2-5*10} and \code{IGHV2-5*07} as it has determined they #' are actually alleles \code{02} and \code{04}, respectively. The updated allele #' names are based on IMGT release 201408-4. #' #' @references #' \enumerate{ #' \item Xochelli et al. (2014) Immunoglobulin heavy variable (IGHV) genes #' and alleles: new entities, new names and implications for research and #' prognostication in chronic lymphocytic leukaemia. Immunogenetics. 67(1):61-6 #' } #' #' @seealso Like \code{updateAlleleNames}, \link{sortAlleles} can help #' format a list of allele names. #' #' @examples #' # Create a vector that uses old gene/allele names. #' alleles <- c("IGHV1-c*01", "IGHV1-f*02", "IGHV2-5*07") #' #' # Update the alleles to the new names #' updateAlleleNames(alleles) #' #' @export updateAlleleNames <- function(allele_calls) { . = NULL temporary_names = c("IGHV1-c*", "IGHV1-f*", "IGHV3-d*", "IGHV3-h*", "IGHV4-b*", "IGHV5-a*", "IGHV2-5*10", "IGHV2-5*07") definitive_names = c("IGHV1-38-4*", "IGHV1-69-2*", "IGHV3-38-3*", "IGHV3-69-1*", "IGHV4-38-2*", "IGHV5-10-1*", "IGHV2-5*02", "IGHV2-5*04") for (i in 1:length(temporary_names)){ allele_calls = allele_calls %>% gsub(temporary_names[i], definitive_names[i], ., fixed = TRUE) } return(allele_calls) } #' Sort allele names #' #' \code{sortAlleles} returns a sorted vector of strings respresenting Ig allele #' names. Names are first sorted by gene family, then by gene, then by allele. #' Duplicated genes have their alleles are sorted as if they were part of their #' non-duplicated counterparts (e.g. \code{IGHV1-69D*01} comes after \code{IGHV1-69*01} #' but before \code{IGHV1-69*02}), and non-localized genes (e.g. \code{IGHV1-NL1*01}) #' come last within their gene family. #' #' @param allele_calls a vector of strings respresenting Ig allele names. #' @param method a string defining the method to use when sorting alleles. #' If \code{"name"} then sort in lexicographic order. If #' \code{"position"} then sort by position in the locus, as #' determined by the final two numbers in the gene name. #' @return A sorted vector of strings respresenting Ig allele names. #' #' @seealso Like \code{sortAlleles}, \link{updateAlleleNames} can help #' format a list of allele names. #' #' @examples #' # Create a list of allele names #' alleles <- c("IGHV1-69D*01","IGHV1-69*01","IGHV1-2*01","IGHV1-69-2*01", #' "IGHV2-5*01","IGHV1-NL1*01", "IGHV1-2*01,IGHV1-2*05", #' "IGHV1-2", "IGHV1-2*02", "IGHV1-69*02") #' #' # Sort the alleles by name #' sortAlleles(alleles) #' #' # Sort the alleles by position in the locus #' sortAlleles(alleles, method="pos") #' #' @export sortAlleles <- function(allele_calls, method=c("name", "position")) { # Check arguments method <- match.arg(method) # Standardize format of submitted alleles, first SUBMITTED_CALLS <- getAllele(allele_calls, first = FALSE, strip_d= FALSE) allele_df <- data.frame( list("SUBMITTED_CALLS"=SUBMITTED_CALLS, "SUBMITTED_NAMES"=allele_calls), stringsAsFactors = FALSE) %>% arrange(SUBMITTED_CALLS) %>% # Determine the family mutate(FAMILY = getFamily(!! rlang::sym("SUBMITTED_CALLS"))) %>% # Determine the gene (exclude family); convert letters to numbers for sort mutate(GENE = getGene(!! rlang::sym("SUBMITTED_CALLS"))) %>% mutate(GENE1 = gsub("[^-]+[-S]([^-\\*D]+).*","\\1",!! rlang::sym("SUBMITTED_CALLS"))) %>% mutate(GENE1 = as.numeric(gsub("[^0-9]+", "99", !!rlang::sym("GENE1")))) %>% # If there is a second gene number, determine that, too mutate(GENE2 = gsub("[^-]+[-S][^-]+-?","",!! rlang::sym("GENE"))) %>% mutate(GENE2 = as.numeric(gsub("[^0-9]+", "99", !!rlang::sym("GENE2")))) %>% mutate(ALLELE = getAllele(!!rlang::sym("SUBMITTED_CALLS"))) %>% mutate(ALLELE = sub("[^\\*]+\\*|[^\\*]+$","", !!rlang::sym("ALLELE"))) %>% mutate(ALLELE = as.numeric(sub("_.+$", "", !!rlang::sym("ALLELE")))) # Convert missing values to 0, sort data frame allele_df[is.na(allele_df)] <- 0 if (method == "name") { sorted_df <- arrange(allele_df, !!!rlang::syms(c("FAMILY", "GENE1", "GENE2", "ALLELE"))) } else if (method == "position") { sorted_df <- arrange(allele_df, desc(!!rlang::sym("GENE1")), desc(!!rlang::sym("GENE2")), desc(!!rlang::sym("FAMILY")), desc(!!rlang::sym("ALLELE"))) } return(sorted_df$SUBMITTED_NAMES) } #' Clean up nucleotide sequences #' #' \code{cleanSeqs} capitalizes nucleotides and replaces all characters #' besides \code{c("A", "C", "G", "T", "-", ".")} with \code{"N"}. #' #' @param seqs a vector of nucleotide sequences. #' #' @return A modified vector of nucleotide sequences. #' #' @seealso \link{sortAlleles} and \link{updateAlleleNames} can #' help format a list of allele names. #' #' @examples #' # Clean messy nucleotide sequences #' seqs <- c("AGAT.taa-GAG...ATA", "GATACAGTXXZZAGNNPPACA") #' cleanSeqs(seqs) #' #' @export cleanSeqs <- function(seqs) { # . = NULL # seqs %>% # toupper %>% # gsub(".", "-", . , fixed = TRUE) %>% # gsub("[^ACGT-]", "N", .) %>% # return clean_seqs <- stri_replace_all_regex(stri_trans_toupper(seqs), "[^ACGT\\.\\-]", "N") if (!is.null(names(seqs))) { names(clean_seqs) <- names(seqs) } clean_seqs } # Private Functions ------------------------------------------------------- # Find muations-by-position compared to a germline # # \code{positionMutations} duplicates the rows of a data frame for each # position to be analyzed and determines if each sample is mutated at that # position # # @param data a Change-O db data.frame. See # \link{findNovelAlleles} for a list of required # columns. # @param germline the germline to which all the sequences should be # compared # @param pos_range the range of positions within the sequence for which # the rows should be duplicated and checked for mutation # @param seq name of the column in \code{data} with the # aligned, IMGT-numbered, V(D)J nucleotide sequence. # Default is SEQUENCE_IMGT. # @return A data frame with rows duplicated for all the positions to be # analyzed and a column indicating whether the position is mutated in # comparison to the germline # positionMutations <- function(data, germline, pos_range, seq="SEQUENCE_IMGT"){ . = NULL pos_db = pos_range %>% length() %>% rep("data", .) %>% paste(collapse=",") %>% paste("bind_rows(",., ")") %>% parse(text=.) %>% eval() pos_db$POSITION = c(sapply(pos_range, rep, nrow(data))) # Find which positions are mutated pos_db <- pos_db %>% mutate(NT = substring(!!rlang::sym(seq), !!rlang::sym("POSITION"), !!rlang::sym("POSITION"))) %>% mutate(GERM_NT = substring(germline, !!rlang::sym("POSITION"), !!rlang::sym("POSITION"))) %>% mutate(MUTATED = (!!rlang::sym("NT") != !!rlang::sym("GERM_NT") & !!rlang::sym("NT") != "N" & !!rlang::sym("NT") != "-" & !!rlang::sym("NT") != "." & !!rlang::sym("NT") != "")) %>% mutate(OBSERVED = (!!rlang::sym("NT") != "-" & !!rlang::sym("NT") != "." & !!rlang::sym("NT") != "")) return(pos_db) } # Find sequences carrying certain levels of mutation # # \code{mutationRangeSubset} determines the mutations in a \code{data.frame} of # sequences and returns the subset of sequences that meet the given mutation # count limits # # @param data a Change-O db data frame. See # \link{findNovelAlleles} for a list of required # columns. # @param germline the germline to which all the sequences should be # compared # @param pos_range the range of positions within the sequences that should # be analyzed for mutations # @param pos_range the range of mutation counts that sequences can have # and still be included # @param seq name of the column in \code{data} with the # aligned, IMGT-numbered, V(D)J nucleotide sequence. # Default is SEQUENCE_IMGT. # @return # A data.frame containing only the subset carrying the desired levels # of mutation # mutationRangeSubset <- function(data, germline, mut_range, pos_range, seq="SEQUENCE_IMGT"){ . = NULL pads = paste(rep("-", min(pos_range)-1), collapse="") data$MUT_COUNT = data[[seq]] %>% substring(min(pos_range), max(pos_range)) %>% paste(pads, ., sep="") %>% getMutatedPositions(germline) %>% sapply(length) data = data %>% filter(!!rlang::sym("MUT_COUNT") %in% mut_range) return(data) } # Find lower range of y-intercept confidence interval # # \code{findLowerY} finds the lower range of y-intercept confidence interval # # @details If mut_min is 1, a y-intercept will be searched for at 0. If # mut_min is above 1, then the "y-intercept" will be found at x = mut_min - 1. # # @param x A vector of x values # @param y A vector of y values # @param mut_min The value where the the lowest mutation count should be # found. See details. # @param alpha The alpha cutoff the be used in constructing the # confidence interval # # @return A data frame containing only the subset carrying the desired levels # of mutation # findLowerY = function(x, y, mut_min, alpha){ y = y + 1 - mut_min lowerY = suppressWarnings(confint(lm(x ~ y), level=1 - 2*alpha)[[1]]) return(lowerY) } # Enchanced substring extraction # # \code{superSubstring} is an enahnced version of \code{substring} in that # it can find disjoint positions in one call. # # @param string a single string # @param positions the positions to be extracted # # @return a substring # superSubstring = function(string, positions){ if(length(string) != 1){ stop("Please submit only one string.") } chars = sapply(positions, function(x) substring(string, x, x)) return(paste(chars, collapse="")) } # Layout multiple ggplots # # \code{multiplot} is a function provided by http://www.cookbook-r.com/ which # allows for plotting multiple ggplot objects in one panel. # # @param ... ggplot2 object(s) # @param plotlist a list alternative to ... # @param file an unused parameter, but present in the provided function # @param cols Number of columns in layout # @param layout A matrix specifying the layout. If present, 'cols' is # ignored. # @param heights A numeric vector A numeric vector or unit object # describing the heights of the rows in the layout. Will # be passed to grid.layout. Default is all plots have # the same height. multiplot <- function(..., plotlist=NULL, cols=1, layout=NULL, heights=NULL) { # Make a list from the ... arguments and plotlist plots <- c(list(...), plotlist) numPlots <- length(plots) ncol <- cols nrow <- ceiling(numPlots/cols) if (is.null(heights)) { heights = rep(1,nrow) } if (is.null(layout)) { # Make the panel # ncol: Number of columns of plots # nrow: Number of rows needed, calculated from # of cols layout <- matrix(seq(1, cols * nrow), ncol = cols, nrow = nrow) } grob <- gridExtra::arrangeGrob(grobs=plots, nrow=nrow, ncol=ncol, layout_matrix = layout, heights=heights) p <- ggplot() + layer(data = data.frame(x = NA), stat = StatIdentity, position = PositionIdentity, # geom = GeomDrawGrob, geom = GeomCustomAnn, inherit.aes = FALSE, params = list(grob = grob, xmin = 0, xmax = 1, ymin = 0, ymax = 1)) + scale_x_continuous(expand=c(0,0)) + scale_y_continuous(expand=c(0,0)) p } #' Subsample repertoire #' #' \code{subsampleDb} will sample the same number of sequences for each gene, family #' or allele (specified with \code{mode}) in \code{data}. Samples or subjects can #' be subsampled indepently by setting \code{group}. #' #' \code{data} will be split into gene, allele or family subsets (\code{mode}) from #' which the same number of sequences will be subsampled. If \code{mode=gene}, #' for each gene in the field \code{gene} from \code{data}, a maximum of #' \code{max_n} sequences will be subsampled. Input sequences #' that have multiple gene calls (ties), can be subsampled from any of their calls, #' but these duplicated samplings will be removed, and the final #' subsampled \code{data} will contain unique rows. #' #' @param data a \code{data.frame} in Change-O format. #' @param gene name of the column in \code{data} with allele calls #' @param mode one of c("gene", "family", "allele") defining the degree of #' specificity regarding allele calls when subsetting sequences. #' Determines how \code{data} will be split into subsets from #' which the same number of sequences will be subsampled. See #' also \code{group}. #' @param min_n minimum number of observations to sample from each groupe. A group with #' less observations than the minimum is excluded. #' @param max_n maximum number of observations to sample for all \code{mode} groups. #' If \code{NULL}, it will be set automatically to the size of #' the smallest group. If \code{max_n} is larger than the availabe #' number of sequences for any \code{mode} group, if will be #' automatically adjusted and the efective \code{max_n} used #' will be the size of the smallest \code{mode} group. #' @param group columns containing additional grouping variables, e.g. sample_id. #' These groups will be subsampled independently. If #' \code{max_n} is \code{NULL}, a \code{max_n} will be #' automatically set for each \code{group}. #' @return #' A \code{data.frame}, subsampled from \code{data}. #' @seealso \link{selectNovel} #' @examples #' # subsampleDb(SampleDb) subsampleDb <- function(data, gene="V_CALL", mode=c("gene", "allele", "family"), min_n=1, max_n=NULL, group=NULL) { mode <- match.arg(mode) # Check columns exist(can be NULL) check <- checkColumns(data, c(gene, group)) if (check != TRUE) { stop(check) } # If group is set, # split by group, then call this same function without grouping, # to apply the subsample within each group independently if (!is.null(group)) { group_id <- data %>% group_by(!!rlang::sym(group)) %>% group_indices() ss_data <- bind_rows( lapply(split(data, group_id), subsampleDb, gene=gene, mode=mode, min_n=min_n, max_n=max_n, group=NULL) ) return(ss_data) } # Extract gene, allele or family assignments gene_func <- switch(mode, allele=getAllele, gene=getGene, family=getFamily) # Get temporary 'gene' names (SubSet Gene) used to # create subsampling groups data[["SS_GENE"]] <- gene_func(data[[gene]], first=FALSE) # Find which rows' calls contain which germline alleles genes <- unique(unlist(strsplit(data[["SS_GENE"]], ","))) allele_groups <- sapply(genes, grep, data[["SS_GENE"]], fixed=TRUE, simplify=FALSE) allele_groups_sizes <- sapply(allele_groups, length) allele_groups <- allele_groups[ allele_groups_sizes >= min_n ] if(length(allele_groups) == 0){ stop("Not enough sample sequences (min_n) were assigned to any gene. ", "Returned `NULL`") } # Determine the number of sequences to subsample if (!is.null(max_n)) { n <- min(allele_groups_sizes, max_n) } else { n <- min(allele_groups_sizes) } # Get indices subsampled sequences ss_idx <- lapply(allele_groups, sample, size=n, replace=FALSE, prob=NULL) ss_idx <- unique(unlist(ss_idx)) data[ss_idx, ] %>% select(-!!rlang::sym("SS_GENE")) } tigger/R/data.R0000644000176200001440000001276213513673571013013 0ustar liggesusers#' Human IGHV germlines #' #' A \code{character} vector of all human IGHV germline gene segment alleles #' in IMGT/GENE-DB (2019-06-01, 372 alleles). #' See IMGT data updates: http://www.imgt.org/IMGTgenedbdoc/dataupdates.html. #' #' @name GermlineIGHV #' @docType data #' @format Values correspond to IMGT-gaped nuceltoide sequences (with #' nucleotides capitalized and gaps represented by ".") while names correspond #' to stripped-down IMGT allele names (e.g. "IGHV1-18*01"). #' #' @references #' \enumerate{ #' \item Xochelli, et al. (2014) Immunoglobulin heavy variable (IGHV) genes and #' alleles: new entities, new names and implications for research and #' prognostication in chronic lymphocytic leukaemia. Immunogenetics. 67(1):61-6. #' } #' #' @keywords data NULL #' Example Human IGHV germlines #' #' A \code{character} vector of all 344 human IGHV germline gene segment alleles #' in IMGT/GENE-DB release 201408-4. #' #' @name SampleGermlineIGHV #' @docType data #' @format Values correspond to IMGT-gaped nuceltoide sequences (with #' nucleotides capitalized and gaps represented by ".") while names correspond #' to stripped-down IMGT allele names (e.g. "IGHV1-18*01"). #' #' @references #' \enumerate{ #' \item Xochelli, et al. (2014) Immunoglobulin heavy variable (IGHV) genes and #' alleles: new entities, new names and implications for research and #' prognostication in chronic lymphocytic leukaemia. Immunogenetics. 67(1):61-6. #' } #' #' @keywords data NULL #' Example human immune repertoire data #' #' A \code{data.frame} of example V(D)J immunoglobulin sequences derived from a #' single individual (PGP1), sequenced on the Roche 454 platform, and assigned by #' IMGT/HighV-QUEST to IGHV1 family alleles. #' #' @name SampleDb #' @docType data #' @format A \code{data.frame} where rows correspond to unique V(D)J sequences and #' columns include: #' \itemize{ #' \item \code{"SEQUENCE_IMGT"}: IMGT-gapped V(D)J nucleotide sequence. #' \item \code{"V_CALL"}: IMGT/HighV-QUEST V segment allele calls. #' \item \code{"D_CALL"}: IMGT/HighV-QUEST D segment allele calls. #' \item \code{"J_CALL"}: IMGT/HighV-QUEST J segment allele calls. #' \item \code{"JUNCTION_LENGTH"}: Junction region length. #' } #' @seealso See \link{airrDb} for an AIRR formatted version of \code{SampleDb}. #' @references #' \enumerate{ #' \item Gadala-Maria, et al. (2015) Automated analysis of high-throughput B cell #' sequencing data reveals a high frequency of novel immunoglobulin V gene #' segment alleles. PNAS. 112(8):E862-70. #' } #' #' @keywords data NULL #' Example human immune repertoire data #' #' A \code{data.frame} of example V(D)J immunoglobulin sequences derived from a #' single individual (PGP1), sequenced on the Roche 454 platform, and assigned by #' IMGT/HighV-QUEST to IGHV1 family alleles. #' #' @name airrDb #' @docType data #' @format A \code{data.frame} where rows correspond to unique V(D)J sequences and #' columns include: #' \itemize{ #' \item \code{"sequence_alignment"}: IMGT-gapped V(D)J nucleotide sequence. #' \item \code{"v_call"}: IMGT/HighV-QUEST V segment allele calls. #' \item \code{"d_call"}: IMGT/HighV-QUEST D segment allele calls. #' \item \code{"j_call"}: IMGT/HighV-QUEST J segment allele calls. #' \item \code{"junction_length"}: Junction region length. #' } #' @seealso See \link{SampleDb} for Change-O formatted version of \code{airrDb}. #' @references #' \enumerate{ #' \item Gadala-Maria, et al. (2015) Automated analysis of high-throughput B cell #' sequencing data reveals a high frequency of novel immunoglobulin V gene #' segment alleles. PNAS. 112(8):E862-70. #' } #' #' @keywords data NULL #' Example novel allele detection results #' #' A \code{data.frame} of novel allele detection results from \link{findNovelAlleles}. #' Source data was a collection of V(D)J immunoglobulin sequences derived from a single #' individual (PGP1), sequenced on the Roche 454 platform, and assigned by #' IMGT/HighV-QUEST to IGHV1 family alleles. #' #' @name SampleNovel #' @docType data #' @format A \code{data.frame} where rows correspond to alleles checked for #' polymorphisms and columns give results as well as paramaters used to run #' the test. #' #' @seealso See \link{findNovelAlleles} for detailed column descriptions. #' #' @references #' \enumerate{ #' \item Gadala-Maria, et al. (2015) Automated analysis of high-throughput B cell #' sequencing data reveals a high frequency of novel immunoglobulin V gene #' segment alleles. PNAS. 112(8):E862-70. #' } #' #' @keywords data NULL #' Example genotype inferrence results #' #' A \code{data.frame} of genotype inference results from \link{inferGenotype} #' after novel allele detection via \link{findNovelAlleles}. #' Source data was a collection of V(D)J immunoglobulin sequences derived from a single #' individual (PGP1), sequenced on the Roche 454 platform, and assigned by #' IMGT/HighV-QUEST to IGHV1 family alleles. #' #' @name SampleGenotype #' @docType data #' @format A \code{data.frame} where rows correspond to genes carried by an #' individual and columns lists the alleles of those genes and their counts. #' #' @seealso See \link{inferGenotype} for detailed column descriptions. #' #' @references #' \enumerate{ #' \item Gadala-Maria, et al. (2015) Automated analysis of high-throughput B cell #' sequencing data reveals a high frequency of novel immunoglobulin V gene #' segment alleles. PNAS. 112(8):E862-70. #' } #' #' @keywords data NULLtigger/R/bayesian.R0000644000176200001440000003652513513673571013700 0ustar liggesusers#' Infer a subject-specific genotype using a Bayesian approach #' #' \code{inferGenotypeBayesian} infers an subject's genotype by applying a Bayesian framework #' with a Dirichlet prior for the multinomial distribution. Up to four distinct alleles are #' allowed in an individual’s genotype. Four likelihood distributions were generated by #' empirically fitting three high coverage genotypes from three individuals #' (Laserson and Vigneault et al, 2014). A posterior probability is calculated for the #' four most common alleles. The certainty of the highest probability model was #' calculated using a Bayes factor (the most likely model divided by second-most likely model). #' The larger the Bayes factor (K), the greater the certainty in the model. #' #' @details #' Allele calls representing cases where multiple alleles have been #' assigned to a single sample sequence are rare among unmutated #' sequences but may result if nucleotides for certain positions are #' not available. Calls containing multiple alleles are treated as #' belonging to all groups. If \code{novel} is provided, all #' sequences that are assigned to the same starting allele as any #' novel germline allele will have the novel germline allele appended #' to their assignent prior to searching for unmutated sequences. #' #' @param data a \code{data.frame} containing V allele #' calls from a single subject. If \code{find_unmutated} #' is \code{TRUE}, then the sample IMGT-gapped V(D)J sequence #' should be provided in column \code{sequence_alignment} #' @param v_call column in \code{data} with V allele calls. #' Default is \code{"V_CALL"}. #' @param seq name of the column in \code{data} with the #' aligned, IMGT-numbered, V(D)J nucleotide sequence. #' Default is SEQUENCE_IMGT. #' @param find_unmutated if \code{TRUE}, use \code{germline_db} to #' find which samples are unmutated. Not needed #' if \code{allele_calls} only represent #' unmutated samples. #' @param germline_db named vector of sequences containing the #' germline sequences named in \code{allele_calls}. #' Only required if \code{find_unmutated} is \code{TRUE}. #' @param novel an optional \code{data.frame} of the type #' novel returned by \link{findNovelAlleles} containing #' germline sequences that will be utilized if #' \code{find_unmutated} is \code{TRUE}. See Details. #' @param priors a numeric vector of priors for the multinomial distribution. #' The \code{priors} vector must be nine values that defined #' the priors for the heterozygous (two allele), #' trizygous (three allele), and quadrozygous (four allele) #' distributions. The first two values of \code{priors} define #' the prior for the heterozygous case, the next three values are for #' the trizygous case, and the final four values are for the #' quadrozygous case. Each set of priors should sum to one. #' Note, each distribution prior is actually defined internally #' by set of four numbers, with the unspecified final values #' assigned to \code{0}; e.g., the heterozygous case is #' \code{c(priors[1], priors[2], 0, 0)}. The prior for the #' homozygous distribution is fixed at \code{c(1, 0, 0, 0)}. #' #' @return #' A \code{data.frame} of alleles denoting the genotype of the subject with the log10 #' of the likelihood of each model and the log10 of the Bayes factor. The output #' contains the following columns: #' #' \itemize{ #' \item \code{GENE}: The gene name without allele. #' \item \code{ALLELES}: Comma separated list of alleles for the given \code{GENE}. #' \item \code{COUNTS}: Comma separated list of observed sequences for each #' corresponding allele in the \code{ALLELES} list. #' \item \code{TOTAL}: The total count of observed sequences for the given \code{GENE}. #' \item \code{NOTE}: Any comments on the inferrence. #' \item \code{KH}: log10 likelihood that the \code{GENE} is homozygous. #' \item \code{KD}: log10 likelihood that the \code{GENE} is heterozygous. #' \item \code{KT}: log10 likelihood that the \code{GENE} is trizygous #' \item \code{KQ}: log10 likelihood that the \code{GENE} is quadrozygous. #' \item \code{K_DIFF}: log10 ratio of the highest to second-highest zygosity likelihoods. #' } #' #' @note #' This method works best with data derived from blood, where a large #' portion of sequences are expected to be unmutated. Ideally, there #' should be hundreds of allele calls per gene in the input. #' #' @seealso \link{plotGenotype} for a colorful visualization and #' \link{genotypeFasta} to convert the genotype to nucleotide sequences. #' See \link{inferGenotype} to infer a subject-specific genotype using #' a frequency method #' #' @references #' \enumerate{ #' \item Laserson U and Vigneault F, et al. High-resolution antibody dynamics of #' vaccine-induced immune responses. PNAS. 2014 111(13):4928-33. #' } #' #' @examples #' # Infer IGHV genotype, using only unmutated sequences, including novel alleles #' inferGenotypeBayesian(SampleDb, germline_db=SampleGermlineIGHV, novel=SampleNovel, #' find_unmutated=TRUE) #' #' @export inferGenotypeBayesian <- function(data, germline_db=NA, novel=NA, v_call="V_CALL", seq="SEQUENCE_IMGT", find_unmutated=TRUE, priors=c(0.6, 0.4, 0.4, 0.35, 0.25, 0.25, 0.25, 0.25, 0.25)){ # Visibility hack . <- NULL allele_calls <- getAllele(data[[v_call]], first=FALSE, strip_d=FALSE) # Find the unmutated subset, if requested if(find_unmutated){ if(is.na(germline_db[1])){ stop("germline_db needed if find_unmutated is TRUE") } if (!is.null(nrow(novel))) { novel <- novel %>% filter(!is.na(!!rlang::sym("POLYMORPHISM_CALL"))) %>% select(!!!rlang::syms(c("GERMLINE_CALL", "POLYMORPHISM_CALL", "NOVEL_IMGT"))) if(nrow(novel) > 0){ # Extract novel alleles if any and add them to germline_db novel_gl <- novel$NOVEL_IMGT names(novel_gl) <- novel$POLYMORPHISM_CALL germline_db <- c(germline_db, novel_gl) # Add the novel allele calls to allele calls of the same starting allele for(r in 1:nrow(novel)){ ind <- grep(novel$GERMLINE_CALL[r], allele_calls, fixed=TRUE) allele_calls[ind] <- allele_calls[ind] %>% sapply(paste, novel$POLYMORPHISM_CALL[r], sep=",") } } } # Find unmutated sequences allele_calls <- findUnmutatedCalls(allele_calls, as.character(data[[seq]]), germline_db) if(length(allele_calls) == 0){ stop("No unmutated sequences found! Set 'find_unmutated' to 'FALSE'.") } } # Find which rows' calls contain which genes gene_regex <- allele_calls %>% strsplit(",") %>% unlist() %>% getGene(strip_d=FALSE) %>% unique() %>% paste("\\*", sep="") gene_groups <- sapply(gene_regex, grep, allele_calls, simplify=FALSE) names(gene_groups) <- gsub("\\*", "", gene_regex, fixed=TRUE) gene_groups <- gene_groups[sortAlleles(names(gene_groups))] # Make a table to store the resulting genotype GENE <- names(gene_groups) # ALLELES = COUNTS = NOTE = rep("", length(GENE)) # TOTAL = sapply(gene_groups, length) # genotype = cbind(GENE, ALLELES, COUNTS, TOTAL, NOTE) ALLELES <- COUNTS <- KH <- KD <- KT <- KQ <- K_DIFF <- NOTE <- rep("", length(GENE)) TOTAL <- sapply(gene_groups, length) genotype <- cbind(GENE, ALLELES, COUNTS, TOTAL, NOTE, KH, KD, KT, KQ, K_DIFF) # For each gene, find which alleles to include for (g in GENE){ # Keep only the part of the allele calls that uses the gene being analyzed ac <- allele_calls[gene_groups[[g]]] %>% strsplit(",") %>% lapply(function(x) x[grep(paste(g, "\\*", sep=""), x)]) %>% sapply(paste, collapse=",") t_ac <- table(ac) # table of allele calls potentials <- unique(unlist(strsplit(names(t_ac),","))) # potential alleles regexpotentials <- paste(gsub("\\*","\\\\*", potentials),"$",sep="") regexpotentials <- paste(regexpotentials,gsub("\\$",",",regexpotentials),sep="|") tmat <- sapply(regexpotentials, function(x) grepl(x, names(t_ac),fixed=FALSE)) if (length(potentials) == 1 | length(t_ac) == 1){ seqs_expl <- t(as.data.frame(apply(t(as.matrix(tmat)), 2, function(x) x * t_ac))) rownames(seqs_expl) <- names(t_ac)[1] }else{ seqs_expl <- as.data.frame(apply(tmat, 2, function(x) x * t_ac)) } # seqs_expl = as.data.frame(apply(tmat, 2, function(x) x*t_ac)) colnames(seqs_expl) <- potentials # Add low (fake) counts sapply(colnames(seqs_expl), function(x){if(sum(rownames(seqs_expl) %in% paste(x)) == 0){ seqs_expl <<- rbind(seqs_expl,rep(0,ncol(seqs_expl))); rownames(seqs_expl)[nrow(seqs_expl)] <<- paste(x) seqs_expl[rownames(seqs_expl) %in% paste(x),paste(x)] <<- 0.01 }}) # Build ratio dependent allele count distribution of multi assigned reads seqs_expl_single <- seqs_expl[grep(',',rownames(seqs_expl),invert = T),] seqs_expl_multi <- seqs_expl[grep(',',rownames(seqs_expl),invert = F),] if(is.null(nrow(seqs_expl_multi))){ seqs_expl_multi <- t(as.data.frame(seqs_expl_multi)) rownames(seqs_expl_multi) <- grep(',',rownames(seqs_expl),invert = F,value = T) } if(!is.null(nrow(seqs_expl_single)) && nrow(seqs_expl_single) !=0 && nrow(seqs_expl_single) != nrow(seqs_expl)){ if(nrow(seqs_expl_multi)>1){ seqs_expl_multi <- seqs_expl_multi[order(nchar(row.names(seqs_expl_multi))),] } sapply(1:nrow(seqs_expl_multi),function(x){ genes <- unlist(strsplit(row.names(seqs_expl_multi)[x],',')); counts <- seqs_expl_single[rownames(seqs_expl_single) %in% genes,genes] counts <- colSums(counts) counts_to_distribute <- seqs_expl_multi[x,genes] new_counts <- counts+((counts_to_distribute*counts)/sum(counts)) for(i in 1:length(new_counts)){ gene_tmp <- names(new_counts)[i] seqs_expl_single[rownames(seqs_expl_single) %in% gene_tmp,gene_tmp] <<- new_counts[i] } }) } # Cycle through the table, including alleles to explain more sequences, # until we explain enough sequences #included = counts = character(0) #tot_expl = 0 seqs_expl <- if(is.null(nrow(seqs_expl_single)) || nrow(seqs_expl_single) ==0 ){seqs_expl}else{seqs_expl_single} seqs_expl <- round(seqs_expl) if(sum(rowSums(seqs_expl) == 0 ) != 0){ seqs_expl <- seqs_expl[rowSums(seqs_expl)!= 0, ] } allele_tot <- sort(apply(seqs_expl, 2, sum),decreasing=TRUE) len=min(length(allele_tot),4); #print(priors) probs <-get_probabilites_with_priors(sort(c(allele_tot,rep(0,4-len)),decreasing = T)[1:4],priors = priors) probs[probs==-Inf] <- -1000 names(probs) <- c('H','D','T','Q') k <- sort(as.numeric(probs),decreasing = T); probs<-c(probs,k[1]-k[2]) names(probs)[5] <- "K_DIFF" genotype[genotype[, "GENE"] == g, "ALLELES"] <- paste(gsub("[^d\\*]*[d\\*]", "", names(allele_tot)[1:len]), collapse = ",") genotype[genotype[, "GENE"] == g, "COUNTS"] <- paste(as.numeric(allele_tot)[1:len], collapse = ",") genotype[genotype[, "GENE"] == g, "KH"] <- probs[1]; genotype[genotype[, "GENE"] == g, "KD"] <- probs[2]; genotype[genotype[, "GENE"] == g, "KT"] <- probs[3]; genotype[genotype[, "GENE"] == g, "KQ"] <- probs[4]; genotype[genotype[, "GENE"] == g, "K_DIFF"] <- probs[5]; # } } geno <- as.data.frame(genotype, stringsAsFactors = FALSE) # Check for indistinguishable calls if(find_unmutated == TRUE){ seqs <- genotypeFasta(geno, germline_db) dist_mat <- seqs %>% sapply(function(x) sapply((getMutatedPositions(seqs, x)), length)) rownames(dist_mat) <- colnames(dist_mat) for (i in 1:nrow(dist_mat)){ dist_mat[i,i] = NA } same <- which(dist_mat == 0, arr.ind=TRUE) if (nrow(same) > 0 ) { for (r in 1:nrow(same)) { inds <- as.vector(same[r,]) geno[getGene(rownames(dist_mat)[inds][1]),]$NOTE <- paste(rownames(dist_mat)[inds], collapse=" and ") %>% paste("Cannot distinguish", .) } } } rownames(geno) <- NULL return(geno) } # Calculate models likelihood # # # @param X a vector of counts # @param alpha_dirichlet alpha parameter for dirichlet distribution # @param epsilon epsilon # @param priors a vector of priors # # @return log10 of the likelihoods get_probabilites_with_priors <- function(X, alpha_dirichlet=c(0.5,0.5,0.5,0.5)*2, epsilon=0.01, priors=c(0.5,0.5,0.33,0.33,0.33,0.25,0.25,0.25,0.25)){ ## Hypotheses X<-sort(X,decreasing=TRUE) H1<-c(1,0,0,0) H2<-c(priors[1],priors[2],0,0) H3<-c(priors[3],priors[4],priors[5],0) H4<-c(priors[6],priors[7],priors[8],priors[9]) E1<-ddirichlet((H1+epsilon)/sum(H1+epsilon),alpha_dirichlet+X) E2<-ddirichlet((H2+epsilon)/sum(H2+epsilon),alpha_dirichlet+X) E3<-ddirichlet((H3+epsilon)/sum(H3+epsilon),alpha_dirichlet+X) E4<-ddirichlet((H4+epsilon)/sum(H4+epsilon),alpha_dirichlet+X) while(sort(c(E1,E2,E3,E4),decreasing=TRUE)[2] == 0 ){ X <- X/10 E1<-ddirichlet((H1+epsilon)/sum(H1+epsilon),alpha_dirichlet+X) E2<-ddirichlet((H2+epsilon)/sum(H2+epsilon),alpha_dirichlet+X) E3<-ddirichlet((H3+epsilon)/sum(H3+epsilon),alpha_dirichlet+X) E4<-ddirichlet((H4+epsilon)/sum(H4+epsilon),alpha_dirichlet+X) } return(log10(c(E1,E2,E3,E4))) } tigger/R/evidence.R0000644000176200001440000003745613513673571013673 0ustar liggesusers# Find non triplet gaps in a nucleotide sequence hasNonImgtGaps <- function (seq) { len <- ceiling(nchar(seq)/3)*3 codons <- substring(seq, seq(1, len-2, 3), seq(3, len, 3)) gaps_lengths <- nchar(gsub("[^\\.\\-]", "", codons)) if (any(gaps_lengths %% 3 != 0)) { TRUE } else { FALSE } } # Compare two IMGT gapped sequences and find AA mutations getMutatedAA <- function(ref_imgt, novel_imgt) { if (grepl("N", ref_imgt)) { stop("Unexpected N in ref_imgt") } if (grepl("N", novel_imgt)) { stop("Unexpected N in novel_imgt") } if (hasNonImgtGaps(ref_imgt)) { warning("Non IMGT gaps found in ref_imgt") } if (hasNonImgtGaps(novel_imgt)) { warning("Non IMGT gaps found in novel_imgt") } ref_imgt <- strsplit(alakazam::translateDNA(ref_imgt),"")[[1]] novel_imgt <- strsplit(alakazam::translateDNA(novel_imgt),"")[[1]] mutations <- c() diff_idx <- which(ref_imgt != novel_imgt) if (length(diff_idx)>0) { mutations <- paste0(diff_idx, ref_imgt[diff_idx],">", replace(novel_imgt[diff_idx], is.na(novel_imgt[diff_idx]),"-")) } mutations } #' Generate evidence #' #' \code{generateEvidence} builds a table of evidence metrics for the final novel V #' allele detection and genotyping inferrences. #' #' @param data a \code{data.frame} containing sequence data that has been #' passed through \link{reassignAlleles} to correct the allele #' assignments. #' @param novel the \code{data.frame} returned by \link{findNovelAlleles}. #' @param genotype the \code{data.frame} of alleles generated with \link{inferGenotype} #' denoting the genotype of the subject. #' @param genotype_db a vector of named nucleotide germline sequences in the genotype. #' Returned by \link{genotypeFasta}. #' @param germline_db the original uncorrected germline database used to by #' \link{findNovelAlleles} to identify novel alleles. #' @param j_call name of the column in \code{data} with J allele calls. #' Default is J_CALL. #' @param junction Junction region nucleotide sequence, which includes #' the CDR3 and the two flanking conserved codons. Default #' is JUNCTION #' @param fields character vector of column names used to split the data to #' identify novel alleles, if any. If \code{NULL} then the data is #' not divided by grouping variables. #' #' @return #' Returns the \code{genotype} input \code{data.frame} with the following additional columns #' providing supporting evidence for each inferred allele: #' #' \itemize{ #' \item \code{FIELD_ID}: Data subset identifier, defined with the input paramter \code{fields}. #' \item A variable number of columns, specified with the input parameter \code{fields}. #' \item \code{POLYMORPHISM_CALL}: The novel allele call. #' \item \code{NOVEL_IMGT}: The novel allele sequence. #' \item \code{CLOSEST_REFERENCE}: The closest reference gene and allele in #' the \code{germline_db} database. #' \item \code{CLOSEST_REFERENCE_IMGT}: Sequence of the closest reference gene and #' allele in the \code{germline_db} database. #' \item \code{GERMLINE_CALL}: The input (uncorrected) V call. #' \item \code{GERMLINE_IMGT}: Germline sequence for \code{GERMLINE_CALL}. #' \item \code{NT_DIFF}: Number of nucleotides that differ between the new allele and #' the closest reference (\code{CLOSEST_REFERENCE}) in the \code{germline_db} database. #' \item \code{NT_SUBSTITUTIONS}: A comma separated list of specific nucleotide #' differences (e.g. \code{112G>A}) in the novel allele. #' \item \code{AA_DIFF}: Number of amino acids that differ between the new allele and the closest #' reference (\code{CLOSEST_REFERENCE}) in the \code{germline_db} database. #' \item \code{AA_SUBSTITUTIONS}: A comma separated list with specific amino acid #' differences (e.g. \code{96A>N}) in the novel allele. #' \item \code{SEQUENCES}: Number of sequences unambiguosly assigned to this allele. #' \item \code{UNMUTATED_SEQUENCES}: Number of records with the unmutated novel allele sequence. #' \item \code{UNMUTATED_FREQUENCY}: Proportion of records with the unmutated novel allele #' sequence (\code{UNMUTATED_SEQUENCES / SEQUENCE}). #' \item \code{ALLELIC_PERCENTAGE}: Percentage at which the (unmutated) allele is observed #' in the sequence dataset compared to other (unmutated) alleles. #' \item \code{UNIQUE_JS}: Number of unique J sequences found associated with the #' novel allele. The sequences are those who have been unambiguously assigned #' to the novel allelle (\code{POLYMORPHISM_CALL}). #' \item \code{UNIQUE_CDR3S}: Number of unique CDR3s associated with the inferred allele. #' The sequences are those who have been unambiguously assigned to the #' novel allelle (POLYMORPHISM_CALL). #' \item \code{MUT_MIN}: Minimum mutation considered by the algorithm. #' \item \code{MUT_MAX}: Maximum mutation considered by the algorithm. #' \item \code{POS_MIN}: First position of the sequence considered by the algorithm (IMGT numbering). #' \item \code{POS_MAX}: Last position of the sequence considered by the algorithm (IMGT numbering). #' \item \code{Y_INTERCEPT}: The y-intercept above which positions were considered #' potentially polymorphic. #' \item \code{ALPHA}: Significance threshold to be used when constructing the #' confidence interval for the y-intercept. #' \item \code{MIN_SEQS}: Input \code{min_seqs}. The minimum number of total sequences #' (within the desired mutational range and nucleotide range) required #' for the samples to be considered. #' \item \code{J_MAX}: Input \code{j_max}. The maximum fraction of sequences perfectly #' aligning to a potential novel allele that are allowed to utilize to a particular #' combination of junction length and J gene. #' \item \code{MIN_FRAC}: Input \code{min_frac}. The minimum fraction of sequences that must #' have usable nucleotides in a given position for that position to be considered. #' \item \code{NOTE}: Comments regarding the novel allele inferrence. #' } #' #' @seealso #' See \link{findNovelAlleles}, \link{inferGenotype} and \link{genotypeFasta} #' for generating the required input. #' #' @examples #' \donttest{ #' # Generate input data #' novel <- findNovelAlleles(SampleDb, SampleGermlineIGHV) #' genotype <- inferGenotype(SampleDb, find_unmutated=TRUE, #' germline_db=SampleGermlineIGHV, #' novel=novel) #' genotype_db <- genotypeFasta(genotype, SampleGermlineIGHV, novel) #' data_db <- reassignAlleles(SampleDb, genotype_db) #' #' # Assemble evidence table #' evidence <- generateEvidence(data_db, novel, genotype, #' genotype_db, SampleGermlineIGHV) #' } #' #' @export generateEvidence <- function(data, novel, genotype, genotype_db, germline_db, j_call="J_CALL", junction="JUNCTION", fields=NULL) { # Visibility hack . <- NULL # Define set of sequences containing genotype and uncorrected calls germline_set <- c(germline_db[!names(germline_db) %in% names(genotype_db)], genotype_db) # Find closest reference .findClosestReference <- function(seq, allele_calls, ref_germ, exclude_self=F, multiple=F) { closest <- getMutCount(seq, paste(allele_calls, collapse=","), ref_germ) min_dist <- min(unlist(closest)) closest_idx <- which(unlist(closest) == min_dist) closest_names <- unique(allele_calls[closest_idx]) if (exclude_self & names(seq) %in% closest_names) { warning("Excluding self") closest_names <- closest_names[closest_names!=names(seq)] # not self } if (length(closest_names) > 1) { warning(paste0("More than one closest reference found for ", names(seq),": ", paste(closest_names, collapse=","))) # Keep the one with less mutated positions mut_pos_count <- sapply(gsub("[^_]","",closest_names), nchar) closest_names <- closest_names[mut_pos_count==min(mut_pos_count)] # Pick same length if (length(closest_names) > 1 ) { idx <- which( sapply(ref_germ[closest_names],nchar) == nchar(ref_germ[names(seq)]) ) if (length(idx) > 0 ) { closest_names <- closest_names[idx] } } # Pick same allele if (length(closest_names) > 1 ) { idx <- which( getAllele(closest_names) == gsub("_.+", "", getAllele(names(seq))) ) if (length(idx) > 0 ) { closest_names <- closest_names[idx] } } # Pick not duplicated if (length(closest_names) > 1 ) { idx <- !grepl("D\\*", closest_names) if (any(idx)) { closest_names <- closest_names[idx] } } # If still more than one, err and TODO if (length(closest_names) > 1 & multiple==FALSE) { msg <- paste0("Multiple closest reference found for ", names(seq),":\n", paste(closest_names, collapse=",")) stop(msg) } warning(paste0("Use: ", paste(closest_names, collapse=","), " (less mutated positions, not D, same length, same allele)")) } closest_names } # Subset to novel alleles final_gt <- genotype %>% dplyr::group_by(.data$GENE) %>% dplyr::filter(!duplicated(.data$ALLELES)) %>% dplyr::ungroup() %>% dplyr::mutate(ALLELES=strsplit(as.character(.data$ALLELES), ","), COUNTS=strsplit(as.character(.data$COUNTS), ",")) %>% tidyr::unnest(.data$ALLELES, .data$COUNTS) %>% dplyr::mutate(POLYMORPHISM_CALL=paste0(.data$GENE, "*" , .data$ALLELES)) %>% dplyr::filter(.data$POLYMORPHISM_CALL %in% novel$POLYMORPHISM_CALL) %>% dplyr::rename(ALLELE="ALLELES") # Add info from novel final_gt <- dplyr::inner_join(dplyr::rename(final_gt, NOTE_GT="NOTE"), novel, by=c(fields, "POLYMORPHISM_CALL")) # Add message if the same novel img sequence found from # different starting alleles, these will be novel imgt sequences # with more than one polymorphism call final_gt <- final_gt %>% dplyr::group_by(.data$NOVEL_IMGT) %>% dplyr::mutate(NUM_CALLS=length(unique(.data$POLYMORPHISM_CALL))) %>% dplyr::ungroup() idx_mult <- which(final_gt$NUM_CALLS > 1) final_gt$NUM_CALLS <- NULL if (length(idx_mult) > 0) { final_gt$NOTE_GT[idx_mult] <- paste( final_gt$NOTE_GT[idx_mult], " Found multiple polymorphism calls for the same NOVEL_IMGT.", sep="") } if (nrow(final_gt)>0) { .addEvidence <- function(df, germline_set, germline_db) { polymorphism <- df[['POLYMORPHISM_CALL']] novel_imgt <- df[["NOVEL_IMGT"]] names(novel_imgt) <- polymorphism v_call_genotyped <- data[["V_CALL_GENOTYPED"]] SEQUENCES <- sum(v_call_genotyped == polymorphism) df[["SEQUENCES"]] <- SEQUENCES closest_ref_input <- .findClosestReference(novel_imgt, names(germline_db), germline_db, exclude_self=F) closest_ref <- .findClosestReference(novel_imgt, names(germline_set), germline_set, exclude_self=F, multiple=T) if (all(getGene(closest_ref_input) != getGene(closest_ref))) { warning("closest reference gene difference") } if (all(closest_ref != polymorphism)) { warning(paste0("closest reference allele (", closest_ref ,") different from POLYMORPHISM_CALL allele (", polymorphism,")")) } df[["CLOSEST_REFERENCE"]] <- closest_ref_input nt_diff <- unlist(getMutatedPositions(novel_imgt, germline_set[[closest_ref_input]])) nt_diff_string <- "" if (nchar(novel_imgt) < nchar(germline_set)[[closest_ref_input]]) { nt_diff <- c(nt_diff, (nchar(novel_imgt)+1):nchar(germline_set[[closest_ref_input]])) } if (length(nt_diff) > 0 ) { ref_nt <- strsplit(germline_set[[closest_ref_input]],"")[[1]][nt_diff] novel_nt <- strsplit(germline_set[[polymorphism]],"")[[1]][nt_diff] nt_diff_string <- paste(paste( nt_diff, ref_nt, ">", replace(novel_nt, is.na(novel_nt), "-"), sep=""), collapse=",") } df[["NT_DIFF"]] <- length(nt_diff) df[["NT_SUBSTITUTIONS"]] <- nt_diff_string diff_aa <- getMutatedAA(germline_set[[closest_ref_input]], germline_set[[polymorphism]]) if (length(diff_aa)>0) { df[["AA_DIFF"]] <- length(diff_aa) df[["AA_SUBSTITUTIONS"]] <- paste(diff_aa,collapse=",") } else { df[["AA_DIFF"]] <- 0 df[["AA_SUBSTITUTIONS"]] <- "" } df[["COUNTS"]] <- as.numeric(df[["COUNTS"]]) df[["TOTAL"]] <- as.numeric(df[["TOTAL"]]) df[["UNMUTATED_SEQUENCES"]] <- as.numeric(df[["COUNTS"]]) df[["UNMUTATED_FREQUENCY"]] <- as.numeric(df[["COUNTS"]])/SEQUENCES df[["ALLELIC_PERCENTAGE"]] <- 100*df[["UNMUTATED_SEQUENCES"]]/as.numeric(df[["TOTAL"]]) if (SEQUENCES > 0) { df[["UNIQUE_JS"]] <- data %>% dplyr::filter(.data$V_CALL_GENOTYPED == polymorphism) %>% dplyr::distinct(.data[[j_call]]) %>% nrow() df[["UNIQUE_CDR3S"]] <- data %>% dplyr::filter(.data$V_CALL_GENOTYPED == polymorphism) %>% dplyr::distinct(translateDNA(.data[[junction]], trim=TRUE)) %>% nrow() } else { df[["UNIQUE_JS"]] <- NA df[["UNIQUE_CDR3S"]] <- NA } # Add closest germline df[["CLOSEST_REFERENCE_IMGT"]] <- cleanSeqs(germline_set[[closest_ref_input]]) data.frame(df, stringsAsFactors=FALSE) } final_gt <- final_gt %>% dplyr::rowwise() %>% do(.addEvidence(., germline_set=germline_set, germline_db=germline_db)) %>% dplyr::mutate(NOTE=trimws(paste(.data$NOTE_GT, .data$NOTE, sep=" "))) %>% dplyr::select(-c("NOTE_GT")) } return(final_gt) } tigger/R/tigger.R0000644000176200001440000001056613514233024013345 0ustar liggesusers# Project documentation for tigger # # @author Daniel Gadala-Maria # @copyright Copyright 2016 Kleinstein Lab, Yale University. All rights reserved # @license Creative Commons Attribution-NonCommercial-ShareAlike 4.0 Unported #' tigger #' #' Here we provide a \strong{T}ool for \strong{I}mmuno\strong{g}lobulin #' \strong{G}enotype \strong{E}lucidation via \strong{R}ep-Seq (TIgGER). #' TIgGER inferrs the set of Ig alleles carried by an #' individual (including any novel alleles) and then uses this set of alleles to #' correct the initial assignments given to sample sequences by existing tools. #' #' @details #' Immunoglobulin repertoire sequencing (AIRR-Seq, Rep-Seq) data is currently the #' subject of much study. A key step in analyzing these data involves assigning #' the closest known V(D)J germline alleles to the (often somatically mutated) #' sample sequences using a tool such as IMGT/HighV-QUEST. However, if the #' sample utilizes alleles not in the germline database used for alignment, this #' step will fail. Additionally, this alignment has an associated error rate of #' ~5%, notably among sequences carrying a large number of somatic #' mutations. The purpose of TIgGER is to address these issues. #' #' @section Allele detection and genotyping: #' \itemize{ #' \item \link{findNovelAlleles}: Detect novel alleles. #' \item \link{plotNovel}: Plot evidence of novel alleles. #' \item \link{inferGenotype}: Infer an Ig genotype using a frequency approach. #' \item \link{inferGenotypeBayesian}: Infer an Ig genotype using a Bayesian approach. #' \item \link{plotGenotype}: A colorful genotype visualization. #' \item \link{genotypeFasta}: Convert a genotype to sequences. #' \item \link{reassignAlleles}: Correct allele calls. #' \item \link{generateEvidence}: Generate evidence for the genotype and #' allele detection inferrence. #' } #' #' @section Mutation handling: #' \itemize{ #' \item \link{getMutatedPositions}: Find mutation locations. #' \item \link{getMutCount}: Find distance from germline. #' \item \link{findUnmutatedCalls}: Subset unmutated sequences. #' \item \link{getPopularMutationCount}: Find most common sequence's #' mutation count. #' \item \link{insertPolymorphisms}: Insert SNPs into a sequence. #' } #' #' @section Input, output and formatting: #' \itemize{ #' \item \link{readIgFasta}: Read a fasta file of Ig sequences. #' \item \link{updateAlleleNames}: Correct outdated allele names. #' \item \link{sortAlleles}: Sort allele names intelligently. #' \item \link{cleanSeqs}: Standardize sequence format. #' } #' #' @name tigger #' @docType package #' @references #' \enumerate{ #' \item Gadala-Maria, et al. (2015) Automated analysis of high-throughput B cell #' sequencing data reveals a high frequency of novel immunoglobulin V gene #' segment alleles. PNAS. 112(8):E862-70. #' } #' #' @import ggplot2 #' @importFrom alakazam getAllele getGene getFamily translateDNA DNA_COLORS checkColumns #' @importFrom doParallel registerDoParallel #' @importFrom dplyr do n desc %>% #' glimpse distinct group_indices #' as_data_frame data_frame #' bind_cols bind_rows combine inner_join #' filter select arrange #' group_by ungroup #' mutate transmute #' rename summarise #' slice #' @importFrom foreach foreach %dopar% registerDoSEQ #' @importFrom graphics plot #' @importFrom gridExtra arrangeGrob #' @importFrom gtools ddirichlet #' @importFrom iterators icount #' @importFrom lazyeval interp #' @importFrom parallel clusterEvalQ clusterExport makeCluster stopCluster #' @importFrom rlang .data := sym syms #' @importFrom shazam calcObservedMutations #' @importFrom stats na.omit setNames ecdf sd cor cov median mad #' confint lm #' @importFrom stringi stri_length stri_detect_fixed stri_replace_all_regex #' stri_sub stri_sub<- stri_trans_toupper #' @importFrom tidyr gather spread unnest NULL tigger/NEWS.md0000644000176200001440000001201513514250717012635 0ustar liggesusersVersion 0.4.0 July 18, 2019 ------------------------------------------------------------------------------- Dependencies: + Upgraded to alakazam >= 0.3.0, shazam >= 0.2.0, dplyr >= 0.8.1 and ggplot2 >= 3.1.1 Bug Fixes: + Fixed bug in `sortAlleles` that was not sorting correctly TR gene names. + Fixed bug in `positionMutations` that was counting `.` as mutated position. New Features: + Updated IGHV germline gene segment alleles in `GermlineIGHV` and moved old annotations to `SampleGermlineIGHV`. + Added the option to specify with arguments the names of the columns that contain V call (`v_call`), J call (`j_call`), sequence alignment (`seq`), junction (`junction`) and junction length (`junction_length`) in all functions that use this information. Version 0.3.1 October 19, 2018 ------------------------------------------------------------------------------- + Fixed a fatal error in `reassignAlleles` with non-existent `v_call` column. + Fixed bug in `generateEvidence` that was reporting amino acids mutations as NA instead of gaps. Version 0.3.0 October 3, 2018 ------------------------------------------------------------------------------- Bug Fixes: + Fixed a bug in `reassignAlleles` occuring with single match genotypes. + Fixed `selectNovel` improperly removing all identical novel alleles, rather than keeping a single entry. + `genotypeFasta` will now retain IMGT-numbering spacers as `.` characters instead of converting them to `-` characters. + Fixed a bug in `findNovelAlleles` causing overly aggressive minimum sequence threshold filtering. + Fixed a bug in the grouping behavior of `getPopularMutationCount`. New Features: + Added a Bayesian approach to genotype inferrence as the `inferGenotypeBayesian` function. + Added the function `generateEvidence` to build a complete evidence table from the results of `findNovelAlleles`, `inferGenotype`, `inferGenotypeBayesian`, and `reassignAlleles`. + Added multiple new evidence columns to the output of `findNovelAlleles` and adjusted the definitions/names of some existing columns. + Added behavior to the `keep_gene` argument of `reassignAlleles` to provide options for maintaining reassignments at the gene (previous `TRUE` behavior), family, or repertoire level. + Improved tie resolution in `findNovelAlleles`. Backwards Incompatible Refactors: + Renamed sample data from `germline_ighv`, `sample_db`, `genotype` and `novel_df` to `GermlineIGHV`, `SampleDb`, `SampleGenotype` and `SampleNovel`, respectively. + Renamed the `novel_df` argument to `novel` in `selectNovel`, `inferGenotype`, and `genotypeFasta`. + Renamed the `novel_df_row` argument to `novel_row` in `plotNovel`. + Argument order in `inferGenotype` was alter for clarity. + Changed the return behavior of `reassignAlleles` so that it returns the input data.frame with the `V_CALL_GENOTYPED` column appended or overwritten. + `cleanSeqs` will no longer replace `.` characters with `-`. + Renamed `clip_db` to `data` in `findNovelAlleles`, `plotNovel`, `inferGenotype` and `reassignAlleles`. Version 0.2.11 September 21, 2017 ------------------------------------------------------------------------------- + Improved memory utilization in `findNovelAlleles`. Version 0.2.10 July 1, 2017 ------------------------------------------------------------------------------- + Bugfix wherein `inferGenotype` would break when performing check for alleles that could not be distinguished. Version 0.2.9.999 May 16, 2017 ------------------------------------------------------------------------------- + Bugfix wherein `inferGenotype` would break if all sequences submitted were from a single gene and `find_unmutated` was set to `TRUE`. Version 0.2.9: March 24, 2017 ------------------------------------------------------------------------------- + License changed to Creative Commons Attribution-ShareAlike 4.0 International (CC BY-SA 4.0). Version 0.2.8: August 26, 2016 ------------------------------------------------------------------------------- + Bugfix following recent update of alakazam (0.2.5) to import selectively. + Removed unneeded dependency on shazam package (not needed as of 0.2.5.999). Version 0.2.7: July 24, 2016 ------------------------------------------------------------------------------- + More updates to work with the latest version of dplyr (0.5.0). + Bugfix in findNovelAlleles when allele passed germline_min but not min_seqs. + Fixed vignette typo and updated findUnmutatedCalls man page. Version 0.2.6: July 01, 2016 ------------------------------------------------------------------------------- + Updated code to work with the latest version of dplyr (0.5.0). Version 0.2.5.999: June 10, 2016 ------------------------------------------------------------------------------- + Fixed a bug werein `findNovelAlleles()` was not running in parallel, even when `nproc` > 1. + Changed default to `nproc=1` in `findNovelAlleles()`. Version 0.2.5: June 07, 2016 ------------------------------------------------------------------------------- + Initial CRAN release. tigger/MD50000644000176200001440000000511113514253663012051 0ustar liggesusers9255757128a68fa484b1a7f0bfca3253 *DESCRIPTION b02245b5d4979b1bcf02e31d376c6c87 *NAMESPACE 2fa63ecdbaeebdee1b4aa4e66749187a *NEWS.md fb7a259c197ff2c873fe189d69b653f6 *R/bayesian.R 43b7ce337f2ee84b94339c36417b3c37 *R/data.R dffd98cd63030fc57de6209fd61a3a08 *R/evidence.R e008566d42fe3d409e891534ab573b22 *R/functions.R f4675d20fd3e2e417091350e62bbd50d *R/tigger.R dd36e57af6a27547f887c01d7f7ea110 *README.md 59482b447d4c9b39d378e66b16adc590 *build/vignette.rds 3e3cb573afe36d5c5ca64289396c180f *data/GermlineIGHV.rda b65f789115c28c50e855e042dd8773b0 *data/SampleDb.rda 5b91397303d4a843360e11172f52fbd2 *data/SampleGenotype.rda d2c0c217f9e10329cd5e8bb2ef518013 *data/SampleGermlineIGHV.rda 996de70aa93a09176f85d473e31d143e *data/SampleNovel.rda 69480acc988a4af5f5939801d836ba34 *data/airrDb.rda 0588c898989c5fd183c025fce44656e3 *data/datalist b2ecadc9aafa513f8082c787613dc6f7 *inst/CITATION 7f75b394db0a8772042a0a0c4422e5a4 *inst/doc/Tigger-Vignette.R 80a90c307b9e1713d193440b96705aca *inst/doc/Tigger-Vignette.Rmd 13dd1603a72ada54303cb58c720d45fe *inst/doc/Tigger-Vignette.pdf af8d4005bdef68f90030cdc035157925 *man/GermlineIGHV.Rd 9583ded9b5323e91035422fa76ac49f9 *man/SampleDb.Rd ee6f174d98cdac6c8e920cffe93b8762 *man/SampleGenotype.Rd 35fccaec7da7e7a54f91ee795258b5ea *man/SampleGermlineIGHV.Rd 887b6cdecd67bce3a4b18c3b68b8ed6c *man/SampleNovel.Rd bc95a3493db0f0b3dc0cf0cea344895c *man/airrDb.Rd 65be86d49f6538199afe8a396d78e46d *man/cleanSeqs.Rd 75ae651bd9d0fa315f5e07a6d543499a *man/findNovelAlleles.Rd 26dee5609e27597515b4fcdce8b61463 *man/findUnmutatedCalls.Rd 1e8914937ac8e7410514b3cc5a07df4f *man/generateEvidence.Rd f3f4904fcfbda4d79fd9d642dc315ca4 *man/genotypeFasta.Rd 14a5c0366b9aa89652e1cb6cde02d284 *man/getMutCount.Rd b985f5ee9c7e41a4343177f1fe73116f *man/getMutatedPositions.Rd b7078e76329f01b84adceb947fd4aef9 *man/getPopularMutationCount.Rd c7cc12a8d6f536dedd171b92be547ec2 *man/inferGenotype.Rd 8e7b8f0537224e32097e8459496ba163 *man/inferGenotypeBayesian.Rd 29fc1168511d72e6c691accdba47c7fb *man/insertPolymorphisms.Rd 4888ee94f43ae2bc8b44493c102158b8 *man/plotGenotype.Rd afd8d60b0d455ddb06db699bea44f429 *man/plotNovel.Rd fe039528bfc29103382b6cfbb92c7447 *man/readIgFasta.Rd 6b9756f70961d4068c18cdb94650bbaf *man/reassignAlleles.Rd b955f370f2897613ab16833edd89397b *man/selectNovel.Rd 72ecb30aceba37af5b1bf37fc6951944 *man/sortAlleles.Rd d15ae816669e4bf50e6c032173c15934 *man/subsampleDb.Rd 96b614e74d00fad9219dcde24e456149 *man/tigger.Rd dc2623df8768f04d7f6f7d65bb51648e *man/updateAlleleNames.Rd 0ed3d8709b7dc07e58a4ac931bf905b0 *man/writeFasta.Rd 80a90c307b9e1713d193440b96705aca *vignettes/Tigger-Vignette.Rmd tigger/inst/0000755000176200001440000000000013514251416012512 5ustar liggesuserstigger/inst/doc/0000755000176200001440000000000013514251416013257 5ustar liggesuserstigger/inst/doc/Tigger-Vignette.Rmd0000644000176200001440000003074413513673571016750 0ustar liggesusers--- title: "Tool for Immunoglobulin Genotype Elucidation via Rep-Seq (TIgGER)" author: "Daniel Gadala-Maria" date: "Last modified `r Sys.Date()`" output: pdf_document: dev: pdf fig_height: 4 fig_width: 7.5 highlight: pygments toc: yes html_document: fig_height: 4 fig_width: 7.5 highlight: pygments theme: readable toc: yes md_document: fig_height: 4 fig_width: 7.5 preserve_yaml: no toc: yes geometry: margin=1in fontsize: 11pt vignette: > %\VignetteEngine{knitr::rmarkdown} %\VignetteIndexEntry{Polymorphism detection and genotyping} %\usepackage[utf8]{inputenc} --- ## Introduction Adapative immune receptor repertoire sequencing (AIRR-Seq, Rep-Seq) data is currently the subject of much study. A key step in analyzing these data involves assigning the closest known V(D)J germline alleles to the (often somatically mutated) sample sequences using a tool such as IMGT/HighV-QUEST ([[1]][1]). However, if the sample utilizes alleles not in the germline database used for alignment, this step will fail. Additionally, this alignment has an associated error rate of ~5% ([[2]][2]), notably among sequences carrying a large number of somatic mutations. Here we provide a **T**ool for **I**mmuno**g**lobulin **G**enotype **E**lucidation via **R**ep-Seq (TIgGER). TIgGER addresses these issues by inferring the set of Immunoglobulin (Ig) alleles carried by an individual (including any novel alleles) and then using this set of alleles to correct the initial assignments given to sample sequences by existing tools. This vignette covers the following tasks: 1. Inferring the presence of novel IGHV alleles not in the germline database. 2. Inferring the personalized IGHV genotype of a sample. 3. Correcting the IGHV allele calls of a sample based on the IGHV genotype. Additional information about the methods used by TIgGER is available in: [Gadala-Maria D, Yaari G, Uduman M, Kleinstein SH (2015) Automated analysis of high-throughput B cell sequencing data reveals a high frequency of novel immunoglobulin V gene segment alleles. *PNAS* 112(8):E862-70](http://www.pnas.org/content/early/2015/02/05/1417683112). ## Input TIgGER requires two main inputs: 1. Pre-processed Ig sequence data 2. Database germline sequences AIRR-seq data is input as a data frame following the Change-O standard where each row represents a unique observation and and columns represent data about that observation. The required names of the required columns are provided below along with a description of each. Column Name | Description ----------------------|--------------------------------------------------------- `SEQUENCE_IMGT` | V(D)J sequence gapped in the IMGT gapped format ([[3]][3]) `V_CALL` | (Comma separated) name(s) of the nearest V allele(s) `J_CALL` | (Comma separated) name(s) of the nearest J allele(s) `JUNCTION_LENGTH` | Length of the junction region of the V(D)J sample An example dataset is provided with the `tigger` package as `SampleDb`. It contains unique functional sequences assigned to IGHV1 family genes isolated from individual PGP1 (referenced in Gadala-Maria *et al.* 2015). The database of germline sequences should be provided in FASTA format with sequences gapped according to the IMGT numbering scheme ([[3]][3]). IGHV alleles in the IMGT database (build 201408-4) are provided with this package as `GermlineIGHV`. You may read in your own fasta file using `readIgFasta`. ```{r, eval=TRUE, message=FALSE, warning=FALSE} # Load packages required for this example library(tigger) library(dplyr) ``` ## Novel allele detection Potential novel alleles can be detected by TIgGER. Some of these may be included in the genotype later (see below). `findNovelAlleles` will return a `data.frame` with a row for each allele tested for the presence of polymorphisms. If polymorphisms are found and the novel sequence passes all quality tests (see below), then the novel allele is named and the germline sequence is included in the `data.frame`. Additionally, the result will contain metadata on the parameters used when searching for novel alleles (which can be optionally changed in `findNovelAlleles`). ```{r, eval=TRUE, warning=FALSE} # Detect novel alleles novel <- findNovelAlleles(SampleDb, GermlineIGHV, nproc=1) ``` ```{r, eval=TRUE, warning=FALSE} # Extract and view the rows that contain successful novel allele calls novel_rows <- selectNovel(novel) novel_rows[1:3] ``` The TIgGER procedure for identifying novel alleles (see citation above) involves taking all sequences which align to a particular germline allele and, for each position along the aligned sequences, plotting the mutation frequency at that postion as a function of the sequence-wide mutation count. While mutational hot-spots and cold-spots are both expected to have a *y*-intercept around zero, polymorphic positions will have a *y*-intercept larger than zero. The required minimum *y*-intercept may be specified in `findNovelAlleles` by `y_intercept`, but defaults to `1/8`. Passing this *y*-intercept threshold is the first of three pieces of evidence that support the novel allele. The second piece of evidence supporting novel allele calls is the nucleotide usage at the polymorphic positions as a function of sequence-wide mutation count. We expect the polymorphic allele to be prevalent at all mutation counts, and we expect the mutation count equal to the number of polymorphisms in the novel sequence to be the most prevalent. Finally, to avoid cases where a clonal expansion might lead to a false positive, combinations of J gene and junction length are examined among sequences which *perfectly match* the proposed germline allele. A true novel allele is expected to utilize a wide range of J genes, and sequences with different junction length can be ruled out as not being clonally related. The maximum portion of sequences which can consist of a specific combination of J gene and junction length may be specified in `findNovelAlleles` by `j_max`. The three pieces of evidence desribed above can be viewed for any allele call made by `findNovelAlleles` using the function `plotNovel`. ```{r, eval=TRUE, warning=FALSE, fig.width=6, fig.height=8} # Plot evidence of the first (and only) novel allele from the example data plotNovel(SampleDb, novel[1, ]) ``` ## Inferring genotypes An individual's genotype can be inferred using the functions `inferGenotype` or `inferGenotypeBayesian`. Using one of this functions allows to remove from the genotype rare/erroneous allele calls which may result from mutations in allele-differentiating regions. `inferGenotype` uses a frequency method to decide which alleles belong to the subjects genotype whereas `inferGenotypeBayesian` infers an subject's genotype applying a Bayesian framework and provides a confidence estimate associated with the genotype calls. ### Frequency genotyping approach `inferGenotype` identifies the fewest alleles that account for nearly all (default is 7/8) of the allele calls made. The user may opt to only use sequences which perfectly match germline alleles, and may opt to include potential novel alleles. (The genotype output is designed to be human readable, though `plotGenotype` can be used to make a colorful visualization.) For each allele, the number of sequences which match the germline are listed in the same order as the alleles are listed. The total number of sequences that match any allele of that gene is also given. To output these alleles as a names vector of nucleotide sequences, the user may use the function `genotypeFasta`. To save this vector to a fasta file, `writeFasta` may be used. ```{r, eval=TRUE, warning=FALSE, fig.width=4, fig.height=3} # Infer the individual's genotype, using only unmutated sequences and checking # for the use of the novel alleles inferred in the earlier step. geno <- inferGenotype(SampleDb, germline_db=GermlineIGHV, novel=novel, find_unmutated=TRUE) # Save the genotype sequences to a vector genotype_db <- genotypeFasta(geno, GermlineIGHV, novel) # Visualize the genotype and sequence counts print(geno) # Make a colorful visualization. Bars indicate presence, not proportion. plotGenotype(geno, text_size = 10) ``` ### Bayesian genotyping approach The method `inferGenotypeBayesian` analyzes the posterior probabilities of possible allele distributions, considering up to four distinct alleles per V gene, corresponding to a gene duplication with both loci being heterozygous (i.e., homozygous, heterozygous with one copy of each allele, etc.). The posterior probabilities for these four possible models are compared and a Bayes factor is calculated for the two most probable models. This Bayes factor reflects the confidence in the genotyping call of the method. The bayesian method doesn't use the strict cutoff criterion `fraction_to_explain` that `inferGenotype` uses wherein only the minimum set of alleles explaining 88% (7/8) of apparently-unmutated sequences are included in the genotype. ```{r, eval=TRUE, warning=FALSE, fig.width=4, fig.height=3} # Infer the individual's genotype, using the bayesian method geno_bayesian <- inferGenotypeBayesian(SampleDb, germline_db=GermlineIGHV, novel=novel, find_unmutated=TRUE) # Visualize the genotype and sequence counts print(geno_bayesian) # Make a colorful visualization. Bars indicate presence, not proportion. plotGenotype(geno_bayesian, text_size=10) ``` ## Correcting allele calls Finally, the original V allele calls may be limited to only those within the inferred genotype. This can be done by using the function `reassignAlleles`. By correcting the calls in this manner, the user can greatly reduce the numer of ambiguous allele calls (where a single sample sequences is assigned to multiple V alleles, thus preventing the mutations analysis of allele-differentiating positions). Additionally, assignments to erroneous not-in-genotype alleles (expected to be ~5% ([[2]][2]), as mentioned above, are corrected in this manner. ```{r, eval=TRUE, warning=FALSE} # Use the personlized genotype to determine corrected allele assignments # Updated genotype will be placed in the V_CALL_GENOTYPED column sample_db <- reassignAlleles(SampleDb, genotype_db) ``` From here, one may proceed with further downstream analyses, but with the advantage of having much-improved allele calls. Besides having discovered alleles not in the IMGT database, the calls made by IMGT have been tailored to the subject's genotype, greatly reducing the number of problematic calls, as can be seen below. ```{r, eval=TRUE, warning=FALSE} # Find the set of alleles in the original calls that were not in the genotype not_in_genotype <- sample_db$V_CALL %>% strsplit(",") %>% unlist() %>% unique() %>% setdiff(names(genotype_db)) # Determine the fraction of calls that were ambigious before/after correction # and the fraction that contained original calls to non-genotype alleles. Note # that by design, only genotype alleles are allowed in "after" calls. data.frame(Ambiguous=c(mean(grepl(",", sample_db$V_CALL)), mean(grepl(",", sample_db$V_CALL_GENOTYPED))), NotInGenotype=c(mean(sample_db$V_CALL %in% not_in_genotype), mean(sample_db$V_CALL_GENOTYPED %in% not_in_genotype)), row.names=c("Before", "After")) %>% t() %>% round(3) ``` ## Evidence `generateEvidence` uses the final corrected calls, the novel alleles and genotype information, the final genotype sequences and the starting reference germlines to build a table of evidence metrics supporting the final novel V allele detection. ```{r, eval=TRUE, warning=FALSE} evidence <- generateEvidence(sample_db, novel, geno, genotype_db, GermlineIGHV, j_call = "J_CALL", junction = "JUNCTION", fields = NULL) evidence %>% select(GENE, ALLELE, POLYMORPHISM_CALL, SEQUENCES, UNMUTATED_FREQUENCY) ``` In this example, `r evidence[["SEQUENCES"]]` sequences were unambiguously assigned to allele `r evidence[["POLYMORPHISM_CALL"]]`, `r round(100*evidence[["UNMUTATED_FREQUENCY"]],2)`% of them unmutated. ## References 1. [Alamyar *et al.* (2010)](http://www.imgt.org/IMGTindex/IMGTHighV-QUEST.html) 2. [Munshaw and Kepler (2010)](http://www.ncbi.nlm.nih.gov/pubmed/20147303) 3. [Lefranc *et al.* (2003)](http://www.ncbi.nlm.nih.gov/pubmed/12477501) [1]: http://www.imgt.org/IMGTindex/IMGTHighV-QUEST.html "Alamyar et al. (2010)" [2]: http://www.ncbi.nlm.nih.gov/pubmed/20147303 "Munshaw and Kepler (2010)" [3]: http://www.ncbi.nlm.nih.gov/pubmed/12477501 "Lefranc et al. 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L!HAFK>J x-j疭΄|vLnS;jHl80]ehhi9|DZ {XIBu+1$ޭ(Fd n9lV!iXыn(EDI7mװA6tJžAT o[Tdl@Ԑbk6Pzx}"vH]N9:'r endstream endobj 184 0 obj << /Author(\376\377\000D\000a\000n\000i\000e\000l\000\040\000G\000a\000d\000a\000l\000a\000-\000M\000a\000r\000i\000a)/Title(\376\377\000T\000o\000o\000l\000\040\000f\000o\000r\000\040\000I\000m\000m\000u\000n\000o\000g\000l\000o\000b\000u\000l\000i\000n\000\040\000G\000e\000n\000o\000t\000y\000p\000e\000\040\000E\000l\000u\000c\000i\000d\000a\000t\000i\000o\000n\000\040\000v\000i\000a\000\040\000R\000e\000p\000-\000S\000e\000q\000\040\000\050\000T\000I\000g\000G\000E\000R\000\051)/Subject()/Creator(LaTeX with hyperref package)/Producer(pdfTeX-1.40.19)/Keywords() /CreationDate (D:20190718222014-07'00') /ModDate (D:20190718222014-07'00') /Trapped /False /PTEX.Fullbanner (This is pdfTeX, Version 3.14159265-2.6-1.40.19 (TeX Live 2018) kpathsea version 6.3.0) >> endobj 139 0 obj << /Type /ObjStm /N 55 /First 476 /Length 3095 /Filter /FlateDecode >> stream xZYoG~Gkb`,+VlɎoE$n(MR_U=dsxDh"GuuWWHLɤ4T86(q3ii&0"4&2epLy;J0v,QDxfRd98if%X&3v"s#bv@Y0@, 8my  %uK  Vw;+edDʠAO$ aQW`q,XT #*T%@5)xe1BJBBz8Z:q  e3q9{B`jH} J}6`Q؎~Ӻ/>>a]}.:*{A-7b CʮKmZ}r[M1!.6i 9`Y;LRޅ..Xp"i>A.T9!QR#)gB4=Yd\*$!AR?%VdZnv;g@/hՂڦDm q 1<кHAHHsh% E0@B*ei 8KE33Ӥu5/+4vY'|GK-hqݐKAX%fLe)5Τc5 l%oEjbeS_k'N3xfx.4^"2LMS.9ڛ'^E@piǑ1^hE cdE0ͪ%v:QhgM R&R?Y &"U-8a|D'ۛ xzlo|/)c:_b ]nZ<"CRfK1Rb҅>S*)XMy6bfbI֍d!/h-!bXQҔ=9 WL~l1>A]L5|rɖ"[AJ0kSpy;#_r~O9-ܓKt:לMOb9ntű'%T⭡$$ ABO2$_if{Yo4Jf\ RA P0!UeE.:Sj4 Z3Wڝ@%Ϥ6Z,;# %׀fa I2-yMcT ͒V>OD1JWt8jy޻_ٛ7 `Ի1(ˡ P<ß;io^!k2yo4?_*&:Oy`qӛ #~ğ3?o;}^k~Ç|k>_IΓ!ގ>dzӅj?[ suW}?yѳoQo-NZ`\hY"|)?~/6o7# 8fp> *Ϋp^j>MzT$}CU}7%߻1`dz\9&i5#TS?\]boV}vw>f7`ʹ,{7TYV`w>w/bo`ho[[[nsP[~-e ̷9acL}0z~ç"&"[ )$4Ԃh8d@nGP 14.>~s|tȇ;u71gDmÃ鮶~ öw),16Z Wxc/绫%C- ^4B@4J-#S,yފiж*Oߜ,JCN[aWvT6z.ܰz{ԻA#icGl%$r`UM߽~~j;E( JX "$qof/M~ImjuojQ(h ̆(rw>4 Ͱj ٸ_#@3~5c_o.*#u} A fa:7z51 T'+4B!wEKF^OOH>br^69Bo; fQQˢ^ \x+0բul &+.N. rNCO>*oJ8W0d81]_@*`q8;`mDfƀ%ض`vM0Uxi۲U\![xvܼ۲ŶlFlʇm%km5mE[ Jda_mu-4mE[ WDSh!DzehKl%d8X^\a;|9xEZ/,' əb!#;ثmLe:gIـ~q|bF_lB g*Ag8_Oo#ɴ.][6 vA|.>#6 >bPn]ZgXnfBW0wGu7KأSwKwcp18A*J4kR欆f0Yi\q29K9q$[SFjMŚA[\ٖmaZxlEF,E(Id.G/.y,\/e=f̵+WE tIA:tm7\)v!xڃG}IewYZ$[BUEAϪe}5OHe߈9;# endstream endobj 185 0 obj << /Type /XRef /Index [0 186] /Size 186 /W [1 3 1] /Root 183 0 R /Info 184 0 R /ID [<5B015B57EE471F0D66E7AA1C44695CA8> <5B015B57EE471F0D66E7AA1C44695CA8>] /Length 476 /Filter /FlateDecode >> stream x%9OTas+:˰+2# " (΢BaebIlL46> _c3wyl#%s[j*q0R[̬T[aYi#퐇f\4B1;D(@#4A3 `7BtYA\Ͻ$p[976NB[<ksm{)uiuky<K 9uѬu~1%&\rCu&0W*MT7=x4Sl7ymL]_SNۻVg0 ^K~}琠AOFe *T4(fP̠AFo]m}V}G}jRӫRJM?(5{Q'hgСCA2Pq(M>(CA=% strsplit(",") %>% unlist() %>% unique() %>% setdiff(names(genotype_db)) # Determine the fraction of calls that were ambigious before/after correction # and the fraction that contained original calls to non-genotype alleles. Note # that by design, only genotype alleles are allowed in "after" calls. data.frame(Ambiguous=c(mean(grepl(",", sample_db$V_CALL)), mean(grepl(",", sample_db$V_CALL_GENOTYPED))), NotInGenotype=c(mean(sample_db$V_CALL %in% not_in_genotype), mean(sample_db$V_CALL_GENOTYPED %in% not_in_genotype)), row.names=c("Before", "After")) %>% t() %>% round(3) ## ---- eval=TRUE, warning=FALSE------------------------------------------- evidence <- generateEvidence(sample_db, novel, geno, genotype_db, GermlineIGHV, j_call = "J_CALL", junction = "JUNCTION", fields = NULL) evidence %>% select(GENE, ALLELE, POLYMORPHISM_CALL, SEQUENCES, UNMUTATED_FREQUENCY) tigger/inst/CITATION0000644000176200001440000000122413514175607013655 0ustar liggesusersbibentry(bibtype = "Article", title = "Automated analysis of high-throughput B cell sequencing data reveals a high frequency of novel immunoglobulin V gene segment alleles.", author = c(person("Daniel", "Gadala-Maria"), person("Gur", "Yaari"), person("Mohamed", "Uduman"), person("Steven H.", "Kleinstein", email="steven.kleinstein@yale.edu")), year = 2015, journal = "Proceedings of the National Academy of Sciency of the United States of America", number = 122, pages = "E862-70")