---
title: "Advanced Colocalization Scenarios with ColocBoost"
output: rmarkdown::html_vignette
vignette: >
  %\VignetteIndexEntry{Advanced Colocalization Scenarios with ColocBoost}
  %\VignetteEngine{knitr::rmarkdown}
  %\VignetteEncoding{UTF-8}
---

```{r, include = FALSE}
knitr::opts_chunk$set(
  collapse = TRUE,
  comment = "#>",
  dpi = 70
)
```

This vignette uses representative simulation studies to illustrate two advanced colocalization scenarios addressed by ColocBoost:

- **Multiple causal variants per trait**: Regions containing multiple causal variants, beyond the one-causal-variant-per-trait assumption.
- **Weaker effects in disease GWAS**: Shared causal signals for which the disease trait contributes weaker association evidence than the accompanying molecular traits.

# 1. Multiple causal variants within a genomic region

To reduce the combinatorial hypothesis space, Bayesian multi-trait colocalization methods commonly assume that <u>*each trait has at most one causal variant within a genomic region*</u> (**one-causal-variant-per-trait assumption**). 
This assumption becomes increasingly restrictive as the number of phenotypes increases and more distinct signals and trait-sharing patterns must be resolved. 
Collapsing these signals into a single-signal representation can obscure event-specific sharing patterns, leading to missed or incorrectly localized colocalization events. 
This concern has also been emphasized and evaluated for pairwise colocalization using COLOC (V5) (Wallace, 2021, *PLOS Genetics*).


## Scenario 1: Heterogeneous effects across traits

A common multi-signal scenario arises when multiple causal variants are shared across traits but have **heterogeneous** effects. 
Consider two traits influenced by two causal variants. Under the *one-causal-variant-per-trait* assumption, each trait is represented only by its strongest signal (Figure 2b(i)):

- **Trait 1** is represented by causal variant 1, which has the strongest association with Trait 1.
- **Trait 2** is represented by causal variant 2, which has the strongest association with Trait 2.

The resulting single-signal representations appear as two distinct trait-specific signals, 
leading to a false conclusion of no colocalization even though both causal variants are shared across the two traits. 
ColocBoost instead resolves the two shared signals as distinct colocalization events.

<img src="figures/Figure2b_i.png"
     alt="Heterogeneous effects of two causal variants across traits."
     style="max-width:100%; height:auto;">


## Scenario 2: Non-causal strongest marginal effect

Another multi-signal scenario occurs when a non-causal variant tags multiple causal variants through LD and consequently has the strongest marginal association.
Distinguishing marginal association from causal attribution motivates multi-effect fine-mapping methods such as SuSiE (Wang et al., 2020, *JRSS B*).

Consider two traits sharing the same two causal variants. Under the *one-causal-variant-per-trait* assumption (Figure 2b(ii)):

- **Trait 1 and Trait 2** are represented by the non-causal marginal lead (green dot), which has a stronger marginal association than either true causal variant (red dots).

The resulting single-signal representation incorrectly localizes the colocalized signal to a non-causal variant, whereas ColocBoost resolves the two shared causal signals as distinct colocalization events.

<img src="figures/Figure2b_ii.png"
     alt="A non-causal variant has the strongest marginal association."
     style="max-width:100%; height:auto;">

# 2. Colocalization with weaker effects in GWAS

In practice, it is often of interest to colocalize a disease GWAS with multiple molecular QTL traits to elucidate the functional basis of disease associations.
An <u>important technical aspect</u> of GWAS-xQTL colocalization is that GWAS traits often have lower per-variant contributions to heritability than molecular xQTL traits.

## Scenario 3: Weaker effects in disease GWAS

Consider a disease GWAS and an xQTL sharing the same two causal variants (Figure 2b(iii)):

- **xQTL** shows strong association evidence for both causal variants.
- **Disease GWAS** shows strong evidence for causal variant 1 but weaker evidence for causal variant 2.

COLOC (V5) identifies the event supported by the stronger GWAS signal but misses the second event with weaker GWAS evidence. 
As a two-stage approach that performs fine-mapping before colocalization, 
it may have reduced sensitivity to weaker signals with limited support in the initial single-trait analysis. 
ColocBoost identifies both shared signals using its disease-prioritized colocalization approach.

<img src="figures/Figure2b_iii.png"
     alt="Colocalization with a weaker causal effect in the disease GWAS."
     style="max-width:100%; height:auto;">

See [Mixed Data-type and Disease Prioritized Colocalization](https://statfungen.github.io/colocboost/articles/Disease_Prioritized_Colocalization.html) for practical guidance on GWAS-xQTL analysis with the ColocBoost disease-prioritized mode.
