Last updated: 2019-07-08

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Knit directory: apaQTL/analysis/

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Unstaged changes:
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    Modified:   code/makePheno.py
    Deleted:    code/test.txt

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Rmd 75b84f4 brimittleman 2019-07-01 add code premature term 
library(reshape2)
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Many papers have started to talk about premature termination. Premature terminated isoforms may be truncated protein or may be degraded. I am going to create a measure for this and test for genetic variation associated with it in my data. The measure will be sum of the reads in intronic PAS and the sum of the UTR reads. I will use leafcutter to put the ratios onto a normal distribution. I will then test for QTLs these ratios.

mkdir ../data/PreTerm_pheno

Prepare phenotype

Total

gene start and end

genes=read.table("/project2/gilad/briana/genome_anotation_data/RefSeq_annotations/FullTranscriptByName.bed", col.names = c("chr", "Gene_start", "Gene_end", "gene", "score", "strand"),stringsAsFactors = F) %>% select(chr,Gene_start, Gene_end, gene)
totalPAS=read.table("../data/phenotype_5perc/APApeak_Phenotype_GeneLocAnno.Total.5perc.fc.gz",stringsAsFactors = F,header = T) 


totalPASPheno=totalPAS %>% melt(id.vars="chrom", variable.name="Ind", value.name = "ratio") %>% separate(ratio, into=c("count", "geneCount"), sep="/") %>% separate(chrom, into=c("chr", "start", "end", "geneID"), sep=":") %>% separate(geneID, into=c("gene", "loc","strand", "PAS"), sep="_") %>% filter(loc=="utr3" | loc=="intron") %>% inner_join(genes,by=c("chr", "gene"))%>% mutate(gene=paste(chr,Gene_start, Gene_end, gene,sep=":")) %>% group_by(Ind,gene,loc) %>% summarise(SumCount=sum(as.integer(count))) %>% ungroup() %>% group_by(Ind,gene) %>% mutate(nType=n()) %>% filter(nType==2) %>% spread(loc, SumCount) %>% mutate(total=intron+utr3,PreTermInt=paste(intron,total, sep="/"),PreTermUTR=paste(utr3,total, sep="/")) %>% select(-nType, -intron,-utr3,-total)


totalPASPheno_melt= totalPASPheno %>% melt(id.vars=c("Ind", "gene"), variable.name="Type", value.name = "Value") %>% mutate(chrom=paste(gene, Type, sep="_")) %>% spread(Ind, Value) %>% select(-gene, -Type)


#write.table(totalPASPheno_melt,"../data/PreTerm_pheno/Total_preterminationPheno.txt",quote=F, row.names=F,col.names=T, sep=" ")
#python2
gzip ../data/PreTerm_pheno/Total_preterminationPheno.txt
python prepare_phenotype_table.py ../data/PreTerm_pheno/Total_preterminationPheno.txt.gz

#activate env  

sh ../data/PreTerm_pheno/Total_preterminationPheno.txt.gz_prepare.sh

#top 2 pcs
head -n 3  ../data/PreTerm_pheno/Total_preterminationPheno.txt.gz.PCs > ../data/PreTerm_pheno/Total_preterminationPheno.txt.gz.2PCs

Nuclear

nuclearPAS=read.table("../data/phenotype_5perc/APApeak_Phenotype_GeneLocAnno.Nuclear.5perc.fc.gz",stringsAsFactors = F,header = T) 


nuclearPASPheno=nuclearPAS %>% melt(id.vars="chrom", variable.name="Ind", value.name = "ratio") %>% separate(ratio, into=c("count", "geneCount"), sep="/") %>% separate(chrom, into=c("chr", "start", "end", "geneID"), sep=":") %>% separate(geneID, into=c("gene", "loc","strand", "PAS"), sep="_") %>% filter(loc=="utr3" | loc=="intron") %>% inner_join(genes,by=c("chr", "gene"))%>% mutate(gene=paste(chr,Gene_start, Gene_end, gene,sep=":")) %>% group_by(Ind,gene,loc) %>% summarise(SumCount=sum(as.integer(count))) %>% ungroup() %>% group_by(Ind,gene) %>% mutate(nType=n()) %>% filter(nType==2) %>% spread(loc, SumCount) %>% mutate(total=intron+utr3,PreTermInt=paste(intron,total, sep="/"),PreTermUTR=paste(utr3,total, sep="/")) %>% select(-nType, -intron,-utr3,-total)


nuclearPASPheno_melt= nuclearPASPheno %>% melt(id.vars=c("Ind", "gene"), variable.name="Type", value.name = "Value") %>% mutate(chrom=paste(gene, Type, sep="_")) %>% spread(Ind, Value) %>% select(-gene, -Type)


#write.table(nuclearPASPheno_melt,"../data/PreTerm_pheno/Nuclear_preterminationPheno.txt",quote=F, row.names=F,col.names=T, sep=" ")
#python2
gzip ../data/PreTerm_pheno/Nuclear_preterminationPheno.txt
python prepare_phenotype_table.py ../data/PreTerm_pheno/Nuclear_preterminationPheno.txt.gz
#env
sh ../data/PreTerm_pheno/Nuclear_preterminationPheno.txt.gz_prepare.sh

#top 2 pcs
head -n 3  ../data/PreTerm_pheno/Nuclear_preterminationPheno.txt.gz.PCs > ../data/PreTerm_pheno/Nuclear_preterminationPheno.txt.gz.2PCs

Call QTLs

Sample list from previous work

mkdir ../data/PrematureQTLNominal
mkdir ../data/PrematureQTLPermuted
sbatch PrematureQTLNominal.sh
sbatch PrematureQTLPermuted.sh

May want to only test one number per gene but do this for now because I want to take advantage of the leafcutter normalization software.

cat ../data/PrematureQTLPermuted/Total_preterminationPheno.txt.gz.qqnorm_chr* > ../data/PrematureQTLPermuted/Total_preterminationPheno.txt.gz.qqnorm_AllChr.txt

cat ../data/PrematureQTLPermuted/Nuclear_preterminationPheno.txt.gz.qqnorm_chr* > ../data/PrematureQTLPermuted/Nuclear_preterminationPheno.txt.gz.qqnorm_AllChr.txt

Tot

totRes=read.table("../data/PrematureQTLPermuted/Total_preterminationPheno.txt.gz.qqnorm_AllChr.txt", stringsAsFactors = F,col.names = c("pid", "nvar", "shape1", "shape2", "dummy", "sid", "dist", "npval", "slope", "ppval", "bpval"))

totRes$bh=p.adjust(totRes$bpval, method="fdr")

totRes_sig=totRes %>% filter(-log10(bh)>1) 


totRes_sig_genes=totRes_sig %>% separate(pid, into=c("chr","start","end", "geneID"), sep=":") %>% separate(geneID, into=c("gene", "Frac"),sep="_") %>% select(gene) %>% unique()

write.table(totRes, file = "../data/PrematureQTLPermuted/Total_preterminationPheno.txt.gz.qqnorm_AllChrBH.txt", col.names = T, row.names = F, quote = F)
nrow(totRes_sig_genes)
[1] 40

Proportion of genes tested:

tottested_genes=totRes %>% separate(pid, into=c("chr","start","end", "geneID"), sep=":") %>% separate(geneID, into=c("gene", "Frac"),sep="_") %>% select(gene) %>% unique()
nrow(totRes_sig_genes)/nrow(tottested_genes)
[1] 0.01162453

qqplot:

qqplot(-log10(runif(nrow(totRes))), -log10(totRes$bpval),ylab="-log10 Total permuted pvalue", xlab="Uniform expectation", main="Total premature termination")
abline(0,1)

Version Author Date
dad7bd8 brimittleman 2019-07-02 
ggplot(totRes, aes(x=dist)) + geom_histogram(bins=100)
Warning: Removed 340 rows containing non-finite values (stat_bin).

Nuclear:

nucRes=read.table("../data/PrematureQTLPermuted/Nuclear_preterminationPheno.txt.gz.qqnorm_AllChr.txt", stringsAsFactors = F,col.names = c("pid", "nvar", "shape1", "shape2", "dummy", "sid", "dist", "npval", "slope", "ppval", "bpval"))

nucRes$bh=p.adjust(nucRes$bpval, method="fdr")

nucRes_sig=nucRes %>% filter(-log10(bh)>1)


nucRes_sig_genes=nucRes_sig %>% separate(pid, into=c("chr","start","end", "geneID"), sep=":") %>% separate(geneID, into=c("gene", "Frac"),sep="_") %>% select(gene) %>% unique()


write.table(nucRes, file = "../data/PrematureQTLPermuted/Nuclear_preterminationPheno.txt.gz.qqnorm_AllChrBH.txt", col.names = T, row.names = F, quote = F)
nrow(nucRes_sig_genes)
[1] 103

Proportion of genes tested:

nuctested_genes=nucRes %>% separate(pid, into=c("chr","start","end", "geneID"), sep=":") %>% separate(geneID, into=c("gene", "Frac"),sep="_") %>% select(gene) %>% unique()
nrow(nucRes_sig_genes)/nrow(nuctested_genes)
[1] 0.02003501

qqplot:

qqplot(-log10(runif(nrow(nucRes))), -log10(nucRes$bpval),ylab="-log10 Nuclear permuted pvalue", xlab="Uniform expectation", main="Nuclear premature termination")
abline(0,1)

More likely in nuclear:

prop.test(x=c(nrow(nucRes_sig_genes),nrow(totRes_sig_genes)), n=c(nrow(nuctested_genes),nrow(tottested_genes)),alternative = "greater")

    2-sample test for equality of proportions with continuity
    correction

data:  c(nrow(nucRes_sig_genes), nrow(totRes_sig_genes)) out of c(nrow(nuctested_genes), nrow(tottested_genes))
X-squared = 8.393, df = 1, p-value = 0.001883
alternative hypothesis: greater
95 percent confidence interval:
 0.003767208 1.000000000
sample estimates:
    prop 1     prop 2 
0.02003501 0.01162453

overlap with eGenes

I next want to look at the proportion of eGenes.

explainedEgenes=read.table("../data/Li_eQTLs/explainedEgenes.txt", col.names = c("gene"), stringsAsFactors = F)
unexplainedEgenes=read.table("../data/Li_eQTLs/UnexplainedEgenes.txt", col.names = c("gene"), stringsAsFactors = F)


allEgenes=bind_rows(explainedEgenes, unexplainedEgenes)

I want to test the proportion of overlap.

TotPre_uneGene=totRes_sig_genes %>% inner_join(unexplainedEgenes,by="gene")
NucPre_uneGene=nucRes_sig_genes %>% inner_join(unexplainedEgenes,by="gene")

TotPre_exeGene=totRes_sig_genes %>% inner_join(explainedEgenes,by="gene")
NucPre_exeGene=nucRes_sig_genes %>% inner_join(explainedEgenes,by="gene")

TotPre_alleGene=totRes_sig_genes %>% inner_join(allEgenes,by="gene")
NucPre_alleGene=nucRes_sig_genes %>% inner_join(allEgenes,by="gene")

Proportion of eGenes explaiend by this:

#total

nrow(TotPre_uneGene)/nrow(unexplainedEgenes)
[1] 0.007894737
nrow(TotPre_exeGene)/nrow(explainedEgenes)
[1] 0.006578947
nrow(TotPre_alleGene)/nrow(allEgenes)
[1] 0.007127193
#nuclear

nrow(NucPre_uneGene)/nrow(unexplainedEgenes)
[1] 0.01184211
nrow(NucPre_exeGene)/nrow(explainedEgenes)
[1] 0.01221805
nrow(NucPre_alleGene)/nrow(allEgenes)
[1] 0.0120614
prop.test(x=c(nrow(NucPre_uneGene),nrow(TotPre_uneGene)), n=c(nrow(unexplainedEgenes),nrow(unexplainedEgenes)))

    2-sample test for equality of proportions with continuity
    correction

data:  c(nrow(NucPre_uneGene), nrow(TotPre_uneGene)) out of c(nrow(unexplainedEgenes), nrow(unexplainedEgenes))
X-squared = 0.26932, df = 1, p-value = 0.6038
alternative hypothesis: two.sided
95 percent confidence interval:
 -0.007305071  0.015199808
sample estimates:
     prop 1      prop 2 
0.011842105 0.007894737 
prop.test(x=c(nrow(NucPre_exeGene),nrow(TotPre_exeGene)), n=c(nrow(explainedEgenes),nrow(explainedEgenes)))

    2-sample test for equality of proportions with continuity
    correction

data:  c(nrow(NucPre_exeGene), nrow(TotPre_exeGene)) out of c(nrow(explainedEgenes), nrow(explainedEgenes))
X-squared = 1.2619, df = 1, p-value = 0.2613
alternative hypothesis: two.sided
95 percent confidence interval:
 -0.003496441  0.014774636
sample estimates:
     prop 1      prop 2 
0.012218045 0.006578947

Conclusion:

Total- 13 overlaps with all eGenes, 7 ex, 6 unexplained Nuclear- 24 overlaps with all eGenes, 13 ex, 11 unexpained

All eGenes=1824 Unexplained=760 Explained=1064

Are the total in the nuclear:

totInuc=totRes_sig_genes %>% anti_join(nucRes_sig_genes,by="gene") 
nrow(totRes_sig_genes)-nrow(totInuc)
[1] 22
#did we test all of the 
totInucTESTEDnuc=totInuc %>% anti_join(nuctested_genes, by="gene") 
nrow(totInucTESTEDnuc)
[1] 2
totInucTESTEDnuc
    gene
1 IPO5P1
2 ZNF718
#all
totInuc %>% inner_join(allEgenes,by="gene")
     gene
1 ATF7IP2
2  MTHFSD
3  IPO5P1
4  ELMOD3
5 ANKRD44
#explained
totInuc %>% inner_join(explainedEgenes,by="gene")
     gene
1 ATF7IP2
2  IPO5P1
#unexplained
totInuc %>% inner_join(unexplainedEgenes,by="gene")
     gene
1  MTHFSD
2  ELMOD3
3 ANKRD44

Direction of effect size:

nucRes_sig_dir= nucRes_sig %>% separate(pid, into=c("chr","start","end", "geneID"), sep=":") %>% separate(geneID, into=c("gene", "Frac"),sep="_")
ggplot(nucRes_sig_dir, aes(x=Frac, y=slope)) + geom_boxplot() + geom_jitter(aes(col=Frac)) + labs(title="Nuclear premature APA QTL effect sizes", y="Effect size",x="") + scale_x_discrete(labels=c( "Intronic Usage","3' UTR Usage"))+ scale_color_discrete(name ="", labels=c( "Intronic Usage","3' UTR Usage")) 

totRes_sig_dir= totRes_sig %>% separate(pid, into=c("chr","start","end", "geneID"), sep=":") %>% separate(geneID, into=c("gene", "Frac"),sep="_")
ggplot(totRes_sig_dir, aes(x=Frac, y=slope)) + geom_boxplot() + geom_jitter(aes(col=Frac))+ labs(title="Total premature APA QTL effect sizes", y="Effect size",x="") + scale_x_discrete(labels=c( "Intronic Usage","3' UTR Usage"))+ scale_color_discrete(name ="", labels=c( "Intronic Usage","3' UTR Usage"))

The difference may just be due to the numbers but most of the variants are associated with decreased utr usage and increase intronic usage.


sessionInfo()
R version 3.5.1 (2018-07-02)
Platform: x86_64-pc-linux-gnu (64-bit)
Running under: Scientific Linux 7.4 (Nitrogen)

Matrix products: default
BLAS/LAPACK: /software/openblas-0.2.19-el7-x86_64/lib/libopenblas_haswellp-r0.2.19.so

locale:
 [1] LC_CTYPE=en_US.UTF-8       LC_NUMERIC=C              
 [3] LC_TIME=en_US.UTF-8        LC_COLLATE=en_US.UTF-8    
 [5] LC_MONETARY=en_US.UTF-8    LC_MESSAGES=en_US.UTF-8   
 [7] LC_PAPER=en_US.UTF-8       LC_NAME=C                 
 [9] LC_ADDRESS=C               LC_TELEPHONE=C            
[11] LC_MEASUREMENT=en_US.UTF-8 LC_IDENTIFICATION=C       

attached base packages:
[1] stats     graphics  grDevices utils     datasets  methods   base     

other attached packages:
 [1] forcats_0.3.0   stringr_1.3.1   dplyr_0.8.0.1   purrr_0.3.2    
 [5] readr_1.3.1     tidyr_0.8.3     tibble_2.1.1    ggplot2_3.1.1  
 [9] tidyverse_1.2.1 workflowr_1.4.0 reshape2_1.4.3 

loaded via a namespace (and not attached):
 [1] Rcpp_1.0.0       cellranger_1.1.0 compiler_3.5.1   pillar_1.3.1    
 [5] git2r_0.25.2     plyr_1.8.4       tools_3.5.1      digest_0.6.18   
 [9] lubridate_1.7.4  jsonlite_1.6     evaluate_0.12    nlme_3.1-137    
[13] gtable_0.2.0     lattice_0.20-38  pkgconfig_2.0.2  rlang_0.3.1     
[17] cli_1.0.1        rstudioapi_0.10  yaml_2.2.0       haven_1.1.2     
[21] withr_2.1.2      xml2_1.2.0       httr_1.3.1       knitr_1.20      
[25] hms_0.4.2        generics_0.0.2   fs_1.2.6         rprojroot_1.3-2 
[29] grid_3.5.1       tidyselect_0.2.5 glue_1.3.0       R6_2.3.0        
[33] readxl_1.1.0     rmarkdown_1.10   modelr_0.1.2     magrittr_1.5    
[37] whisker_0.3-2    backports_1.1.2  scales_1.0.0     htmltools_0.3.6 
[41] rvest_0.3.2      assertthat_0.2.0 colorspace_1.3-2 labeling_0.3    
[45] stringi_1.2.4    lazyeval_0.2.1   munsell_0.5.0    broom_0.5.1     
[49] crayon_1.3.4