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Integrative Analysis of Differential Gene Expression, Transcript Usage, and RNA Secondary Structure in Early- and Late-Onset Colorectal Cancer

by Salomon Marquez

11/02/2025

This site contains a summary of my bioinformatics master's thesis developed at CIMA University of Navarra under the supervision of Fernando Pastor Rodriguez and Igor Ruiz de los Mozos.

For a detailed overview of the project, you can check the following resources:


Abstract

Colorectal cancer is increasingly diagnosed in individuals younger than 50 years, a trend referred to as early-onset colorectal cancer. This study applied an integrative transcriptomic workflow to characterize molecular differences between early-onset and late-onset colorectal cancer. A total of 86 matched tumor and normal samples were analyzed using a combination of differential gene expression, differential transcript usage, and RNA secondary structure prediction.

Differential gene expression analysis identified a 40-gene signature uniquely deregulated in early-onset colorectal cancer, enriched in immune-related, inflammatory, and cell-matrix adhesion processes. Differential transcript usage analysis revealed substantial differences in transcript-level complexity between cohorts, with late-onset colorectal cancer showing extensive isoform remodeling, whereas early-onset tumors displayed a more restricted pattern of transcript usage. Despite these global differences, six conserved isoform switches were consistently detected in both cohorts, including TPM2, LMNB2, MAP3K20, and the long non-coding RNA HOXB-AS3. RNA secondary structure prediction further characterized significant isoforms, revealing a predominance of stem motifs.

These results indicate that gene-level and isoform-level analyses capture complementary aspects of colorectal cancer transcriptomic variation. The integrative framework presented here provides a methodological basis for future large-scale studies and supports the exploration of RNA structural features as potential biomarkers and therapeutic targets in colorectal cancer.

keywords: colorectal-cancer, deseq2-analysisdtu-analysisrna-structure-prediction, nf-core-rnaseqisoformSwitchAnalyzeR, hpc-clusternextflow


Objective

The primary goal of this project is to identify genes and transcripts that are differentially regulated in early-onset and late-onset colorectal cancer, and to evaluate whether changes in isoform usage are associated with differences in their RNA secondary structure through computational prediction.


Proposed solution

We adopted an integrative transcriptomic strategy to investigate molecular differences between early-onset and late-onset colorectal cancer (EOCRC and LOCRC). Building upon a previously conducted gene-level study from Marx et al., we extend the analysis by incorporating isoform-level regulation and RNA secondary structure prediction, thereby moving beyond expression changes alone to explore post-transcriptional and structural layers of regulation. The overall integrative analysis workflow is summarized below:

tfm-workflow Schematic overview of the integrative transcriptomic analysis workflow implemented in this study.


From concept to implementation

The main milestones of this project were tracked using a dedicated 🏁 roadmap on GitHub projects, which facilitated version control and documentation of analytical decisions. Among these decisions, the main technical and methodological challenges encountered during the development of this project and the mitigation strategies adopted are listed below:


Summary of products obtained

This project produced the following deliverables.

  • A master’s thesis manuscript documenting the analytical workflow, results, and their biological interpretation.
  • A configured and validated nf-core/rnaseq Nextflow pipeline for reproducible RNA-seq analysis on high-performance computing (HPC) clusters.
  • A Zenodo repository hosting primary and processed data generated in this study.
  • A GitHub repository containing primary scripts and documentation.

Results

This is a summary of results after the implementation of the integrative transcriptomic workflow for the analysis of EOCRC and LOCRC samples.

1. nf-core/rnaseq pipeline — EOCRC

SRA study SRP357925
21 EOCRC patient pairs

2. nf-core/rnaseq pipeline — LOCRC

SRA study SRP479528
22 LOCRC patient pairs

3. Differential Gene Expression (DGE)

4. Differential Transcript Usage (DTU)

  • IsoformSwitchAnalyzeR Analysis
Show R script
# Install and load modules and libraries 
.libPaths("~/R/x86_64-pc-linux-gnu-library/4.4")

# Check if package is installed, if not then install
if (!require("IsoformSwitchAnalyzeR", quietly = TRUE)) {
    if (!require("BiocManager", quietly = TRUE))
        install.packages("BiocManager")
    BiocManager::install("IsoformSwitchAnalyzeR")
}

library(scales)
library(ggplot2)
library(tidyr)
library(dplyr)
library(S4Vectors)
library(reshape2)
library(corrplot)
library(factoextra)
library(SummarizedExperiment)
library(DESeq2)
library(IsoformSwitchAnalyzeR)
library(tidyverse)
library(dbplyr)
library(BiocParallel)
library(Biostrings)

## ------------------------------------------------------------------------------------------------
# Adjust workers according to cluster cores
ncores <- 16
register(MulticoreParam(workers = ncores))

# Import quantifications
salmonQuant <- importIsoformExpression(
    parentDir = "dtu_data/salmon"
)

head(salmonQuant$abundance, 2)
head(salmonQuant$counts, 2)

saveRDS(salmonQuant, file = "dtu_data/salmonQuant_42_44.rds")

## ------------------------------------------------------------------------------------------------
# Read 'se' object 
se_42 <- readRDS("dtu_data/gene.SummarizedExperiment.metadata_42.rds")
se_44 <- readRDS("dtu_data/gene.SummarizedExperiment.metadata_44.rds")

# Display 'se' object
se_42
se_44

# Display colNames 
colnames(se_42)
colnames(se_44)

# Display condition
colData(se_42)$condition
colData(se_44)$condition

# Display condition in salmonQuant
colnames(salmonQuant$abundance)

## ------------------------------------------------------------------------------------------------
# Define design matrix
myDesign <- data.frame(
    sampleID = colnames(salmonQuant$abundance)[-1],    # Exclude isoform_id column and include all sample names
    condition = c(colData(se_42)$condition, colData(se_44)$condition)  # Combine conditions from both se_42 and se_44
)
myDesign

# Define comparisonsToMake
mycomparisonsToMake <- data.frame(
    condition_1 = c("normal_eocrc", "normal_locrc"),
    condition_2 = c("tumor_eocrc", "tumor_locrc") 
)
mycomparisonsToMake

## ------------------------------------------------------------------------------------------------
cat("1. Create switchAnalyzeRlist...\n")

# Create switchAnalyzeRlist
aSwitchList <- importRdata(
    isoformCountMatrix   = salmonQuant$counts,
    isoformRepExpression = salmonQuant$abundance,
    designMatrix         = myDesign,
    comparisonsToMake    = mycomparisonsToMake,
    isoformExonAnnoation = "dtu_data/gencode.v49.primary_assembly.annotation.gtf.gz",
    isoformNtFasta       = "dtu_data/gencode.v49.transcripts.fa.gz",
    fixStringTieAnnotationProblem = TRUE,
    showProgress = TRUE
)

cat("View aSwitchList summary...\n")

# View summary report
summary(aSwitchList)
saveRDS(aSwitchList, file = "dtu_outputs/aSwitchList_gencode_42_44.rds")

## ------------------------------------------------------------------------------------------------
cat("2. Prefiltering...\n")
cat("Prefiltering summary with geneExpressionCutoff = 1; isoformExpressionCutoff = 0; Isoform fraction = 0...\n")

# Prefilter
mySwitchList <- preFilter(aSwitchList)
summary(mySwitchList)
saveRDS(mySwitchList, file = "dtu_outputs/mySwitchList_gencode_42_44_filstandard.rds")

## ------------------------------------------------------------------------------------------------
cat("3. Differential Expression Analysis using TestSatuRn...\n")

# Identify differentially used isoforms with SatuRn
mySwitchList <- isoformSwitchTestSatuRn(
    mySwitchList,
    reduceToSwitchingGenes = TRUE,  # focus on switching genes relevant to CRC
    showProgress = TRUE
)

# Summarize results
extractSwitchSummary(mySwitchList)

## ------------------------------------------------------------------------------------------------
cat("4. Obtain Open Reading Frames (ORFs)...\n")

# Obtain Open Reading Frames (ORFs)
mySwitchList <- analyzeORF(
    mySwitchList,
    showProgress = TRUE 
)

## ------------------------------------------------------------------------------------------------
cat("5. Extracting sequences...\n")

# Extract Sequences
mySwitchList <- extractSequence(
    mySwitchList,
    outputPrefix = "dtu_42_44"
)
saveRDS(mySwitchList, file = "dtu_outputs/mySwitchList_42_44_nt.rds")

## ------------------------------------------------------------------------------------------------
cat("6. Alternative and intron retention analysis...\n")

# Analyze alternative splicing
mySwitchList <- analyzeAlternativeSplicing(
    switchAnalyzeRlist = mySwitchList,
    quiet = TRUE
)

## ------------------------------------------------------------------------------------------------
# Analyze Intron Retention
mySwitchList <- analyzeIntronRetention(
    switchAnalyzeRlist = mySwitchList,
    onlySwitchingGenes = TRUE,
    alpha = 0.05,
    dIFcutoff = 0.1,
    showProgress = TRUE,
    quiet = TRUE
)

saveRDS(mySwitchList, file = "dtu_outputs/mySwitchList_Asplicing_Iretention_42_44.rds")

# Extract 10 top switching genes (by q-value)
extractTopSwitches(
    mySwitchList, 
    filterForConsequences = FALSE, 
    n = 10, 
    sortByQvals = TRUE
)

## ------------------------------------------------------------------------------------------------
cat("7. Analyze Consequences of Isoform Switches...\n")

# AnalyzeSwitchConsequences
# tss -> alternative 5'UTR
# tts -> alternative 3'UTR

consequencesOfInterest <- c(
    "tss",
    "tts",
    "exon_number",
    "intron_structure",
    "intron_retention"
)

bioMechanismeAnalysis <- analyzeSwitchConsequences(
    mySwitchList,
    consequencesToAnalyze = consequencesOfInterest, 
    showProgress = TRUE
)

extractSwitchSummary(bioMechanismeAnalysis, filterForConsequences = FALSE)
extractSwitchSummary(bioMechanismeAnalysis, filterForConsequences = TRUE)

saveRDS(bioMechanismeAnalysis, file = "dtu_outputs/mySwitchList_consequences_42_44.rds")
class: SummarizedExperiment 
dim: 78428 42 
metadata(8): abstract Accession ... Submission Relevance
assays(5): counts counts_length_scaled counts_scaled lengths tpm
rownames(78428): ENSG00000000003.17 ENSG00000000005.6 ...
  ENSG00000310592.1 ENSG00000310593.1
rowData names(3): transcript_id gene_id gene_name
colnames(42): srr17866817 srr17866818 ... srr17866857 srr17866858
colData names(32): age_at_surgery tissue ... source_name condition
class: SummarizedExperiment 
dim: 78428 44 
metadata(9): abstract Accession ... Submission Relevance
assays(5): counts counts_length_scaled counts_scaled lengths tpm
rownames(78428): ENSG00000000003.17 ENSG00000000005.6 ...
  ENSG00000310592.1 ENSG00000310593.1
rowData names(3): transcript_id gene_id gene_name
colnames(44): srr27320655 srr27320656 ... srr27320697 srr27320698
colData names(33): age_at_surgery tissue ... source_name condition
[1] "srr17866817" "srr17866818" "srr17866819" "srr17866820" "srr17866821"
[6] "srr17866822" "srr17866823" "srr17866824" "srr17866825" "srr17866826"
[11] "srr17866827" "srr17866828" "srr17866829" "srr17866830" "srr17866831"
[16] "srr17866832" "srr17866833" "srr17866834" "srr17866835" "srr17866836"
[21] "srr17866837" "srr17866838" "srr17866839" "srr17866840" "srr17866841"
[26] "srr17866842" "srr17866843" "srr17866844" "srr17866845" "srr17866846"
[31] "srr17866847" "srr17866848" "srr17866849" "srr17866850" "srr17866851"
[36] "srr17866852" "srr17866853" "srr17866854" "srr17866855" "srr17866856"
[41] "srr17866857" "srr17866858"
[1] "srr27320655" "srr27320656" "srr27320657" "srr27320658" "srr27320659"
[6] "srr27320660" "srr27320661" "srr27320662" "srr27320663" "srr27320664"
[11] "srr27320665" "srr27320666" "srr27320667" "srr27320668" "srr27320669"
[16] "srr27320670" "srr27320671" "srr27320672" "srr27320673" "srr27320674"
[21] "srr27320675" "srr27320676" "srr27320677" "srr27320678" "srr27320679"
[26] "srr27320680" "srr27320681" "srr27320682" "srr27320683" "srr27320684"
[31] "srr27320685" "srr27320686" "srr27320687" "srr27320688" "srr27320689"
[36] "srr27320690" "srr27320691" "srr27320692" "srr27320693" "srr27320694"
[41] "srr27320695" "srr27320696" "srr27320697" "srr27320698"
[1] tumor_eocrc  normal_eocrc tumor_eocrc  normal_eocrc tumor_eocrc 
[6] normal_eocrc tumor_eocrc  normal_eocrc tumor_eocrc  normal_eocrc
[11] tumor_eocrc  normal_eocrc tumor_eocrc  normal_eocrc tumor_eocrc 
[16] normal_eocrc tumor_eocrc  normal_eocrc tumor_eocrc  normal_eocrc
[21] tumor_eocrc  normal_eocrc tumor_eocrc  normal_eocrc tumor_eocrc 
[26] normal_eocrc tumor_eocrc  normal_eocrc tumor_eocrc  normal_eocrc
[31] tumor_eocrc  normal_eocrc tumor_eocrc  normal_eocrc tumor_eocrc 
[36] normal_eocrc tumor_eocrc  normal_eocrc tumor_eocrc  normal_eocrc
[41] tumor_eocrc  normal_eocrc
Levels: normal_eocrc tumor_eocrc
[1] normal_locrc tumor_locrc  normal_locrc tumor_locrc  normal_locrc
[6] tumor_locrc  normal_locrc tumor_locrc  normal_locrc tumor_locrc 
[11] normal_locrc tumor_locrc  normal_locrc tumor_locrc  normal_locrc
[16] tumor_locrc  normal_locrc tumor_locrc  normal_locrc tumor_locrc 
[21] normal_locrc tumor_locrc  normal_locrc tumor_locrc  normal_locrc
[26] tumor_locrc  normal_locrc tumor_locrc  normal_locrc tumor_locrc 
[31] normal_locrc tumor_locrc  normal_locrc tumor_locrc  normal_locrc
[36] tumor_locrc  normal_locrc tumor_locrc  normal_locrc tumor_locrc 
[41] normal_locrc tumor_locrc  normal_locrc tumor_locrc 
Levels: normal_locrc tumor_locrc
[1] "isoform_id"  "srr17866817" "srr17866818" "srr17866819" "srr17866820"
[6] "srr17866821" "srr17866822" "srr17866823" "srr17866824" "srr17866825"
[11] "srr17866826" "srr17866827" "srr17866828" "srr17866829" "srr17866830"
[16] "srr17866831" "srr17866832" "srr17866833" "srr17866834" "srr17866835"
[21] "srr17866836" "srr17866837" "srr17866838" "srr17866839" "srr17866840"
[26] "srr17866841" "srr17866842" "srr17866843" "srr17866844" "srr17866845"
[31] "srr17866846" "srr17866847" "srr17866848" "srr17866849" "srr17866850"
[36] "srr17866851" "srr17866852" "srr17866853" "srr17866854" "srr17866855"
[41] "srr17866856" "srr17866857" "srr17866858" "srr27320655" "srr27320656"
[46] "srr27320657" "srr27320658" "srr27320659" "srr27320660" "srr27320661"
[51] "srr27320662" "srr27320663" "srr27320664" "srr27320665" "srr27320666"
[56] "srr27320667" "srr27320668" "srr27320669" "srr27320670" "srr27320671"
[61] "srr27320672" "srr27320673" "srr27320674" "srr27320675" "srr27320676"
[66] "srr27320677" "srr27320678" "srr27320679" "srr27320680" "srr27320681"
[71] "srr27320682" "srr27320683" "srr27320684" "srr27320685" "srr27320686"
[76] "srr27320687" "srr27320688" "srr27320689" "srr27320690" "srr27320691"
[81] "srr27320692" "srr27320693" "srr27320694" "srr27320695" "srr27320696"
[86] "srr27320697" "srr27320698"
      sampleID    condition
1  srr17866817  tumor_eocrc
2  srr17866818 normal_eocrc
3  srr17866819  tumor_eocrc
4  srr17866820 normal_eocrc
5  srr17866821  tumor_eocrc
6  srr17866822 normal_eocrc
7  srr17866823  tumor_eocrc
8  srr17866824 normal_eocrc
9  srr17866825  tumor_eocrc
10 srr17866826 normal_eocrc
11 srr17866827  tumor_eocrc
12 srr17866828 normal_eocrc
13 srr17866829  tumor_eocrc
14 srr17866830 normal_eocrc
15 srr17866831  tumor_eocrc
16 srr17866832 normal_eocrc
17 srr17866833  tumor_eocrc
18 srr17866834 normal_eocrc
19 srr17866835  tumor_eocrc
20 srr17866836 normal_eocrc
21 srr17866837  tumor_eocrc
22 srr17866838 normal_eocrc
23 srr17866839  tumor_eocrc
24 srr17866840 normal_eocrc
25 srr17866841  tumor_eocrc
26 srr17866842 normal_eocrc
27 srr17866843  tumor_eocrc
28 srr17866844 normal_eocrc
29 srr17866845  tumor_eocrc
30 srr17866846 normal_eocrc
31 srr17866847  tumor_eocrc
32 srr17866848 normal_eocrc
33 srr17866849  tumor_eocrc
34 srr17866850 normal_eocrc
35 srr17866851  tumor_eocrc
36 srr17866852 normal_eocrc
37 srr17866853  tumor_eocrc
38 srr17866854 normal_eocrc
39 srr17866855  tumor_eocrc
40 srr17866856 normal_eocrc
41 srr17866857  tumor_eocrc
42 srr17866858 normal_eocrc
43 srr27320655 normal_locrc
44 srr27320656  tumor_locrc
45 srr27320657 normal_locrc
46 srr27320658  tumor_locrc
47 srr27320659 normal_locrc
48 srr27320660  tumor_locrc
49 srr27320661 normal_locrc
50 srr27320662  tumor_locrc
51 srr27320663 normal_locrc
52 srr27320664  tumor_locrc
53 srr27320665 normal_locrc
54 srr27320666  tumor_locrc
55 srr27320667 normal_locrc
56 srr27320668  tumor_locrc
57 srr27320669 normal_locrc
58 srr27320670  tumor_locrc
59 srr27320671 normal_locrc
60 srr27320672  tumor_locrc
61 srr27320673 normal_locrc
62 srr27320674  tumor_locrc
63 srr27320675 normal_locrc
64 srr27320676  tumor_locrc
65 srr27320677 normal_locrc
66 srr27320678  tumor_locrc
67 srr27320679 normal_locrc
68 srr27320680  tumor_locrc
69 srr27320681 normal_locrc
70 srr27320682  tumor_locrc
71 srr27320683 normal_locrc
72 srr27320684  tumor_locrc
73 srr27320685 normal_locrc
74 srr27320686  tumor_locrc
75 srr27320687 normal_locrc
76 srr27320688  tumor_locrc
77 srr27320689 normal_locrc
78 srr27320690  tumor_locrc
79 srr27320691 normal_locrc
80 srr27320692  tumor_locrc
81 srr27320693 normal_locrc
82 srr27320694  tumor_locrc
83 srr27320695 normal_locrc
84 srr27320696  tumor_locrc
85 srr27320697 normal_locrc
86 srr27320698  tumor_locrc
1. Create switchAnalyzeRlist...

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                    comparison estimated_genes_with_dtu
1 normal_eocrc vs normal_locrc              1027 - 1712
2  normal_eocrc vs tumor_eocrc                214 - 357
3   tumor_eocrc vs tumor_locrc               858 - 1429
View aSwitchList summary...
This switchAnalyzeRlist list contains:
398518 isoforms from 58088 genes
6 comparison from 4 conditions (in total 86 samples)

Feature analyzed:
[1] "ORFs, ntSequence"
2. Prefiltering...
Prefiltering summary with geneExpressionCutoff = 1; isoformExpressionCutoff = 0; Isoform fraction = 0...
This switchAnalyzeRlist list contains:
119137 isoforms from 14067 genes
6 comparison from 4 conditions (in total 86 samples)

Feature analyzed:
[1] "ORFs, ntSequence"

[1] "SatuRn results with standard prefiltering...\n"
                  Comparison nrIsoforms nrSwitches nrGenes
1 normal_eocrc vs tumor_eocrc         14         10      10
2 normal_locrc vs tumor_locrc        133        122     104
3                    Combined        140        128     109

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            gene_ref gene_id gene_name  condition_1 condition_2
1  geneComp_00055095    TPM2      TPM2 normal_eocrc tumor_eocrc
2  geneComp_00110879  SORBS2    SORBS2 normal_locrc tumor_locrc
3  geneComp_00113199    TPM2      TPM2 normal_locrc tumor_locrc
4  geneComp_00091701   FSIP1     FSIP1 normal_locrc tumor_locrc
5  geneComp_00102515 OSBPL1A   OSBPL1A normal_locrc tumor_locrc
6  geneComp_00098800 MAP3K20   MAP3K20 normal_locrc tumor_locrc
7  geneComp_00061664    CD44      CD44 normal_locrc tumor_locrc
8  geneComp_00063393  CXCL12    CXCL12 normal_locrc tumor_locrc
9  geneComp_00101397     NLN       NLN normal_locrc tumor_locrc
10 geneComp_00093642   HNF4A     HNF4A normal_locrc tumor_locrc
  gene_switch_q_value Rank
1         0.0001116680    1
2         0.0001544585    2
3         0.0001544585    3
4         0.0001622092    4
5         0.0002382206    5
6         0.0003502090    6
7         0.0004922797    7
8         0.0006191435    8
9         0.0006191435    9
10        0.0009308128   10

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                  Comparison nrIsoforms nrSwitches nrGenes
1 normal_eocrc vs tumor_eocrc         14         10      10
2 normal_locrc vs tumor_locrc        133        122     104
3                    Combined        140        128     109
                  Comparison nrIsoforms nrSwitches nrGenes
1 normal_eocrc vs tumor_eocrc         14         10      10
2 normal_locrc vs tumor_locrc        132        122     103
3                    Combined        140        128     109

5. RNA Secondary Structure Analysis (RSS)


Contributing

This is a thesis repository, but suggestions for improvements are welcome via GitHub issues.