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//! Arena-based partial order alignment (POA) graph types.
//!
//! This module ports `spoa`'s pointer-based graph (`unique_ptr<Node>` / raw `Node*`) to an
//! index-based arena: nodes and edges live in flat `Vec`s on [`Graph`], and are referenced by
//! [`NodeId`] / [`EdgeId`] indices rather than pointers. This avoids `unsafe` entirely while
//! preserving the original's semantics.
use std::collections::HashSet;
use std::fmt;
/// Errors raised by [`Graph::add_alignment`] and its convenience wrappers.
///
/// Mirrors the three `std::invalid_argument` throws in `spoa::Graph::AddAlignment`
/// (`graph.cpp:162-166,184-188,191-194`).
#[derive(Debug, Clone, Copy, PartialEq, Eq)]
pub enum GraphError {
/// `seq` and `weights` have different lengths (`graph.cpp:162-166`).
UnequalWeights,
/// An alignment entry's sequence index is out of bounds for `seq` (`graph.cpp:184-188`).
InvalidAlignment,
/// No alignment entry references any position in `seq` (`graph.cpp:191-194`).
MissingSequence,
}
impl fmt::Display for GraphError {
fn fmt(&self, f: &mut fmt::Formatter<'_>) -> fmt::Result {
let message = match self {
GraphError::UnequalWeights => "sequence and weights are of unequal size",
GraphError::InvalidAlignment => "invalid alignment",
GraphError::MissingSequence => "missing sequence in alignment",
};
write!(f, "[spoars::Graph::add_alignment] error: {message}")
}
}
impl std::error::Error for GraphError {}
/// Index of a [`Node`] within [`Graph::nodes`].
#[derive(Debug, Clone, Copy, PartialEq, Eq, Hash, PartialOrd, Ord)]
pub struct NodeId(pub u32);
/// Index of an [`Edge`] within [`Graph::edges`].
#[derive(Debug, Clone, Copy, PartialEq, Eq, Hash, PartialOrd, Ord)]
pub struct EdgeId(pub u32);
/// A single POA graph node: one aligned "column" position holding one coded symbol.
///
/// Mirrors `spoa::Graph::Node` (`graph.hpp:40-71`), minus the `id` field (an arena `Node`'s
/// identity is its index in [`Graph::nodes`], carried externally as a [`NodeId`]).
#[derive(Debug, Clone)]
pub struct Node {
/// The coded symbol (see [`Graph::coder`] / [`Graph::decoder`]) this node represents.
pub code: u32,
/// Edges whose head is this node.
pub inedges: Vec<EdgeId>,
/// Edges whose tail is this node.
pub outedges: Vec<EdgeId>,
/// Other nodes aligned to this one (same column, different symbol).
pub aligned_nodes: Vec<NodeId>,
}
impl Node {
/// The distinct input-sequence labels passing through this node — the set-union of the
/// sequence-index labels on all of its in-edges and out-edges — sorted ascending.
///
/// This is the set [`Node::coverage`] counts; exposing it lets callers derive per-node strand
/// or sequence membership without re-walking edges. See [`Graph::sequence_starts`] /
/// [`Graph::sequence_path`] for what a label indexes.
pub fn labels(&self, graph: &Graph) -> Vec<u32> {
let mut labels: HashSet<u32> = HashSet::new();
for &edge_id in self.inedges.iter().chain(self.outedges.iter()) {
labels.extend(graph.edges[edge_id.0 as usize].labels.iter().copied());
}
let mut labels: Vec<u32> = labels.into_iter().collect();
labels.sort_unstable();
labels
}
/// Number of distinct input sequences passing through this node.
///
/// Mirrors `spoa::Graph::Node::Coverage` (`graph.cpp:32-47`): the size of the set-union of
/// the sequence-index labels on all of this node's in-edges and out-edges (i.e.
/// `self.labels(graph).len()`).
pub fn coverage(&self, graph: &Graph) -> u32 {
self.labels(graph).len() as u32
}
/// Returns the head of this node's first out-edge (in insertion order) whose `labels`
/// contains `label` — i.e. the next node visited by sequence `label` after this one — or
/// `None` if no out-edge carries that label (this node is `label`'s last).
///
/// Mirrors `spoa::Graph::Node::Successor` (`graph.cpp:22-30`).
pub fn successor(&self, graph: &Graph, label: u32) -> Option<NodeId> {
for &edge_id in &self.outedges {
let edge = &graph.edges[edge_id.0 as usize];
if edge.labels.contains(&label) {
return Some(edge.head);
}
}
None
}
/// The raw input byte this node represents, decoded via `graph` (`graph.decode(self.code)`),
/// or `None` if the code is unknown to `graph` (which cannot happen for a node obtained from
/// that same graph). A convenience over reading [`Node::code`] and calling [`Graph::decode`].
pub fn base(&self, graph: &Graph) -> Option<u8> {
graph.decode(self.code)
}
}
/// A directed edge between two POA graph nodes, tagged with the sequences that traverse it.
///
/// Mirrors `spoa::Graph::Edge` (`graph.hpp:72-100`).
#[derive(Debug, Clone)]
pub struct Edge {
/// Source node of this edge.
pub tail: NodeId,
/// Destination node of this edge.
pub head: NodeId,
/// Sequence-index labels of every input sequence that traverses this edge. Load-bearing for
/// `Successor`-style MSA-row and GFA P-line reconstruction in later tasks.
pub labels: Vec<u32>,
/// Sum of per-sequence weights of all sequences traversing this edge.
pub weight: i64,
}
/// An arena-based partial order alignment graph.
///
/// Mirrors `spoa::Graph` (`graph.hpp:25-320`), replacing its `unique_ptr<Node>` / raw `Node*`
/// pointer graph with flat `Vec<Node>` / `Vec<Edge>` arenas indexed by [`NodeId`] / [`EdgeId`].
#[derive(Debug, Clone)]
pub struct Graph {
/// Arena of all nodes ever added to the graph, indexed by [`NodeId`].
pub(crate) nodes: Vec<Node>,
/// Arena of all edges ever added to the graph, indexed by [`EdgeId`].
pub(crate) edges: Vec<Edge>,
/// Maps a raw input byte (0-255) to its assigned code, or -1 if not yet seen.
pub(crate) coder: [i32; 256],
/// Maps a code back to its raw input byte, or -1 if the code is unused.
pub(crate) decoder: Vec<i32>,
/// Number of distinct symbol codes assigned so far.
pub(crate) num_codes: u32,
/// For each added sequence (in order), the [`NodeId`] of its first node.
pub(crate) sequences: Vec<NodeId>,
/// Nodes in topological-sort order, recomputed by [`Graph::topological_sort`] after every
/// `add_alignment` call.
pub(crate) rank_to_node: Vec<NodeId>,
/// Nodes forming the generated consensus sequence, in traversal order, populated by
/// [`Graph::traverse_heaviest_bundle`].
pub(crate) consensus: Vec<NodeId>,
}
impl Default for Graph {
fn default() -> Graph {
Graph::new()
}
}
impl Graph {
/// Creates an empty graph, with the byte-to-code table cleared to "unassigned" (`-1`).
///
/// Mirrors `spoa::Graph::Graph` (`graph.cpp:65-74`).
pub fn new() -> Graph {
Graph {
nodes: Vec::new(),
edges: Vec::new(),
coder: [-1; 256],
decoder: vec![-1; 256],
num_codes: 0,
sequences: Vec::new(),
rank_to_node: Vec::new(),
consensus: Vec::new(),
}
}
// ---- Public read-only accessors -----------------------------------------------------------
//
// The graph is *built* through `add_alignment*` and *summarized* through `generate_consensus*`/
// `generate_msa`/`to_gfa`/`to_dot`. These accessors expose the underlying arena so downstream
// crates can also *inspect* the DAG directly — enumerate nodes/edges, follow the topological
// order, read the consensus path as node ids, and decode a node's coded symbol back to its raw
// input byte. All are immutable borrows: the arena is only ever mutated through the crate's own
// graph-construction methods, preserving its invariants.
/// All nodes ever added, indexed by [`NodeId`]'s inner `u32` (`self.nodes()[id.0 as usize]`,
/// or use [`Graph::node`]). Nodes are never removed, so a [`NodeId`] stays valid for the
/// graph's lifetime.
pub fn nodes(&self) -> &[Node] {
&self.nodes
}
/// All edges ever added, indexed by [`EdgeId`]'s inner `u32` (or use [`Graph::edge`]).
pub fn edges(&self) -> &[Edge] {
&self.edges
}
/// The [`Node`] for `id`.
///
/// # Panics
/// Panics if `id` is out of range (mirrors slice indexing); every id handed out by this crate
/// is always in range.
pub fn node(&self, id: NodeId) -> &Node {
&self.nodes[id.0 as usize]
}
/// The [`Edge`] for `id`.
///
/// # Panics
/// Panics if `id` is out of range (mirrors slice indexing).
pub fn edge(&self, id: EdgeId) -> &Edge {
&self.edges[id.0 as usize]
}
/// Number of nodes in the graph.
pub fn num_nodes(&self) -> usize {
self.nodes.len()
}
/// Number of edges in the graph.
pub fn num_edges(&self) -> usize {
self.edges.len()
}
/// Number of distinct symbol codes assigned so far (the alphabet size seen across all added
/// sequences).
pub fn num_codes(&self) -> u32 {
self.num_codes
}
/// The raw input byte that a node's [`Node::code`] represents, or `None` if `code` was never
/// assigned. Inverse of [`Graph::encode`]; use it to turn a node back into its base, e.g.
/// `graph.decode(graph.node(id).code)`.
pub fn decode(&self, code: u32) -> Option<u8> {
match self.decoder.get(code as usize).copied() {
Some(byte) if byte >= 0 => Some(byte as u8),
_ => None,
}
}
/// The code assigned to a raw input `base`, or `None` if that byte has not been seen in any
/// added sequence. Inverse of [`Graph::decode`].
pub fn encode(&self, base: u8) -> Option<u32> {
match self.coder[base as usize] {
code if code >= 0 => Some(code as u32),
_ => None,
}
}
/// Nodes in topological-sort order, recomputed after every `add_alignment*` call. Iterating
/// this yields a valid processing order (every node appears after all of its predecessors);
/// it is the same order the alignment fill and consensus traversal use.
pub fn rank_order(&self) -> &[NodeId] {
&self.rank_to_node
}
/// For each sequence added (in insertion order), the [`NodeId`] of its first node — the entry
/// points for per-sequence traversal via [`Node::successor`].
pub fn sequence_starts(&self) -> &[NodeId] {
&self.sequences
}
/// The nodes of the most recently generated consensus path, in traversal order, or an empty
/// slice if no consensus has been generated yet. Populated by [`Graph::generate_consensus`] /
/// [`Graph::generate_consensus_min_coverage`]; `graph.consensus_nodes().iter().filter_map(|&n|
/// graph.decode(graph.node(n).code))` reconstructs the same bytes those methods return as a
/// `String`.
pub fn consensus_nodes(&self) -> &[NodeId] {
&self.consensus
}
/// The nodes sequence `seq_index` traverses, in order, from its start
/// ([`Graph::sequence_starts`]`()[seq_index]`) following that sequence's labeled edges via
/// [`Node::successor`], as a borrowing iterator (no allocation).
///
/// `seq_index` is the 0-based index assigned in `add_alignment*` order (see the ordering
/// guarantee on [`Graph::add_alignment`]); it is also the row index in [`Graph::generate_msa`].
/// Decoding the visited nodes (`graph.decode(graph.node(id).code)`) reconstructs the sequence's
/// bases in order.
///
/// # Panics
/// Panics if `seq_index >= self.sequence_starts().len()`.
pub fn sequence_path_iter(&self, seq_index: usize) -> impl Iterator<Item = NodeId> + '_ {
let label = seq_index as u32;
let mut next = Some(self.sequences[seq_index]);
std::iter::from_fn(move || {
let current = next?;
next = self.nodes[current.0 as usize].successor(self, label);
Some(current)
})
}
/// The nodes sequence `seq_index` traverses, in order, collected into a `Vec`. Allocating form
/// of [`Graph::sequence_path_iter`]; see it for the index semantics.
///
/// # Panics
/// Panics if `seq_index >= self.sequence_starts().len()`.
pub fn sequence_path(&self, seq_index: usize) -> Vec<NodeId> {
self.sequence_path_iter(seq_index).collect()
}
/// For each [`NodeId`], the MSA column it occupies (aligned peer nodes share a column), plus the
/// total column count. This is the exact mapping [`Graph::generate_msa`] uses internally to
/// place each node's symbol; a downstream caller can use it to map bases to MSA columns without
/// materializing the character-grid MSA.
///
/// The returned `Vec` is indexed by `NodeId`'s inner `u32`; the `u32` is the number of columns.
pub fn msa_columns(&self) -> (Vec<u32>, u32) {
self.initialize_msa()
}
/// One entry per MSA column, listing the `(sequence_index, NodeId)` pairs present in that
/// column — the column-major inverse of [`Graph::msa_columns`], saving callers from inverting
/// the node→column map themselves.
///
/// A `(sequence_index, node)` pair is emitted for column `c` when `sequence_index` traverses
/// `node` (via [`Graph::sequence_path_iter`]) and `node` occupies column `c`. Within each
/// column, pairs are ordered by `sequence_index` then by traversal order.
pub fn column_members(&self) -> Vec<Vec<(u32, NodeId)>> {
let (node_id_to_column, row_size) = self.initialize_msa();
let mut columns: Vec<Vec<(u32, NodeId)>> = vec![Vec::new(); row_size as usize];
for seq_index in 0..self.sequences.len() {
for node in self.sequence_path_iter(seq_index) {
let column = node_id_to_column[node.0 as usize] as usize;
columns[column].push((seq_index as u32, node));
}
}
columns
}
/// Appends a new node with the given code and returns its id.
///
/// Mirrors `spoa::Graph::AddNode` (`graph.cpp:76-79`): the new node's id is simply the
/// arena's length before the push.
pub(crate) fn add_node(&mut self, code: u32) -> NodeId {
let id = NodeId(self.nodes.len() as u32);
self.nodes.push(Node {
code,
inedges: Vec::new(),
outedges: Vec::new(),
aligned_nodes: Vec::new(),
});
id
}
/// Adds a directed edge from `tail` to `head`, or if one already exists, folds the new
/// sequence into it.
///
/// Mirrors `spoa::Graph::AddEdge` (`graph.cpp:81-91`). `label` is the in-progress sequence's
/// index (`sequences.len()` at the caller's call time); it is load-bearing for
/// `Successor`-style MSA-row and GFA P-line reconstruction, so both the dedup path and the
/// new-edge path record it.
pub(crate) fn add_edge(&mut self, tail: NodeId, head: NodeId, weight: i64, label: u32) {
for &edge_id in &self.nodes[tail.0 as usize].outedges {
let edge = &mut self.edges[edge_id.0 as usize];
if edge.head == head {
edge.labels.push(label);
edge.weight += weight;
return;
}
}
let edge_id = EdgeId(self.edges.len() as u32);
self.edges.push(Edge {
tail,
head,
labels: vec![label],
weight,
});
self.nodes[tail.0 as usize].outedges.push(edge_id);
self.nodes[head.0 as usize].inedges.push(edge_id);
}
/// Appends one node per base of `seq[begin..end]`, chained by edges weighted with the sum of
/// each pair of adjacent bases' weights. Returns the id of the first node appended, or `None`
/// if `begin == end`.
///
/// Mirrors `spoa::Graph::AddSequence` (`graph.cpp:93-110`). Assumes every byte in
/// `seq[begin..end]` has already been registered in [`Graph::coder`].
fn add_sequence(
&mut self,
seq: &[u8],
weights: &[u32],
begin: u32,
end: u32,
) -> Option<NodeId> {
if begin == end {
return None;
}
let label = self.sequences.len() as u32;
let first = NodeId(self.nodes.len() as u32);
let mut prev: Option<NodeId> = None;
for i in begin..end {
let code = self.coder[seq[i as usize] as usize] as u32;
let curr = self.add_node(code);
if let Some(p) = prev {
let weight = weights[(i - 1) as usize] as i64 + weights[i as usize] as i64;
self.add_edge(p, curr, weight, label);
}
prev = Some(curr);
}
Some(first)
}
/// Threads `seq` into the graph along `alignment`, its alignment to the graph's existing
/// nodes, weighting each base-to-base edge by `weights`.
///
/// `alignment` is a list of `(node_index, seq_index)` pairs mirroring `spoa::Alignment`
/// (`vector<pair<int32_t, int32_t>>`): `-1` in either slot is the "no match" sentinel — a
/// `-1` node index means the sequence base has no counterpart yet in the graph (insertion),
/// and a `-1` sequence index means an existing graph node has no counterpart in this sequence
/// (deletion).
///
/// Mirrors `spoa::Graph::AddAlignment` (`graph.cpp:155-247`).
///
/// # Sequence-index ordering (guaranteed)
/// The Nth call to any `add_alignment*` method (`N` counting from 1, over non-empty sequences)
/// assigns that sequence the label / index `N - 1`. That same 0-based index is:
/// - the sequence's position in [`Graph::sequence_starts`],
/// - the argument to [`Graph::sequence_path`] / [`Graph::sequence_path_iter`], and
/// - the row index of that sequence in [`Graph::generate_msa`]'s output.
///
/// This correspondence is a stable part of the API, not an implementation accident.
pub fn add_alignment(
&mut self,
alignment: &[(i32, i32)],
seq: &[u8],
weights: &[u32],
) -> Result<(), GraphError> {
if seq.is_empty() {
return Ok(());
}
if seq.len() != weights.len() {
return Err(GraphError::UnequalWeights);
}
// Coder registration (graph.cpp:168-173): assign every not-yet-seen byte a fresh code.
for &base in seq {
let byte = base as usize;
if self.coder[byte] == -1 {
self.coder[byte] = self.num_codes as i32;
self.decoder[self.num_codes as usize] = base as i32;
self.num_codes += 1;
}
}
if alignment.is_empty() {
// Empty-alignment fast path (graph.cpp:175-179): the whole sequence is new.
if let Some(start) = self.add_sequence(seq, weights, 0, seq.len() as u32) {
self.sequences.push(start);
}
self.topological_sort();
return Ok(());
}
let mut valid: Vec<u32> = Vec::new();
for &(node_idx, seq_idx) in alignment {
if seq_idx != -1 {
if seq_idx < 0 || seq_idx as usize >= seq.len() {
return Err(GraphError::InvalidAlignment);
}
// `node_idx` flows straight into `self.nodes[..]` in the aligned-bases loop
// below; `-1` is the "new node" sentinel, so reject any other out-of-range
// value here rather than let it panic on the index.
if node_idx != -1 && (node_idx < 0 || node_idx as usize >= self.nodes.len()) {
return Err(GraphError::InvalidAlignment);
}
valid.push(seq_idx as u32);
}
}
if valid.is_empty() {
return Err(GraphError::MissingSequence);
}
let valid_front = valid[0];
let valid_back = valid[valid.len() - 1];
// Add unaligned prefix bases (positions before the first aligned base).
let mut begin = self.add_sequence(seq, weights, 0, valid_front);
let mut prev: Option<NodeId> = begin.map(|_| NodeId(self.nodes.len() as u32 - 1));
// Add unaligned suffix bases (positions after the last aligned base) up front, matching
// the C++ call order; `last` is the first node of that suffix chain, chained in below.
let last = self.add_sequence(seq, weights, valid_back + 1, seq.len() as u32);
// Add aligned bases (graph.cpp:202-240).
for &(node_idx, seq_idx) in alignment {
if seq_idx == -1 {
continue;
}
let seq_idx = seq_idx as usize;
let code = self.coder[seq[seq_idx] as usize] as u32;
let curr = if node_idx == -1 {
self.add_node(code)
} else {
let existing = NodeId(node_idx as u32);
if self.nodes[existing.0 as usize].code == code {
existing
} else {
let mut matched: Option<NodeId> = None;
for &aligned in &self.nodes[existing.0 as usize].aligned_nodes {
if self.nodes[aligned.0 as usize].code == code {
matched = Some(aligned);
break;
}
}
match matched {
Some(node) => node,
None => {
// Cross-link the new node into `existing`'s aligned-node set
// (graph.cpp:216-231).
let new_node = self.add_node(code);
let existing_aligned =
self.nodes[existing.0 as usize].aligned_nodes.clone();
for aligned in existing_aligned {
self.nodes[aligned.0 as usize].aligned_nodes.push(new_node);
self.nodes[new_node.0 as usize].aligned_nodes.push(aligned);
}
self.nodes[existing.0 as usize].aligned_nodes.push(new_node);
self.nodes[new_node.0 as usize].aligned_nodes.push(existing);
new_node
}
}
}
};
if begin.is_none() {
begin = Some(curr);
}
if let Some(p) = prev {
let label = self.sequences.len() as u32;
let weight = weights[seq_idx - 1] as i64 + weights[seq_idx] as i64;
self.add_edge(p, curr, weight, label);
}
prev = Some(curr);
}
if let Some(last_node) = last {
let label = self.sequences.len() as u32;
let weight =
weights[valid_back as usize] as i64 + weights[(valid_back + 1) as usize] as i64;
let p = prev.expect("aligned-bases loop always sets prev when `valid` is non-empty");
self.add_edge(p, last_node, weight, label);
}
if let Some(start) = begin {
self.sequences.push(start);
}
self.topological_sort();
Ok(())
}
/// Convenience wrapper: builds a uniform per-base `weights` vector from a single `weight` and
/// delegates to [`Graph::add_alignment`].
///
/// Mirrors `spoa::Graph::AddAlignment(alignment, sequence, sequence_len, weight)`
/// (`graph.cpp:119-125`).
pub fn add_alignment_weight(
&mut self,
alignment: &[(i32, i32)],
seq: &[u8],
weight: u32,
) -> Result<(), GraphError> {
let weights = vec![weight; seq.len()];
self.add_alignment(alignment, seq, &weights)
}
/// Convenience wrapper: derives per-base weights from Phred-scaled `quality` bytes
/// (`quality[i] - 33`) and delegates to [`Graph::add_alignment`].
///
/// Mirrors `spoa::Graph::AddAlignment(alignment, sequence, sequence_len, quality, quality_len)`
/// (`graph.cpp:137-146`).
pub fn add_alignment_quality(
&mut self,
alignment: &[(i32, i32)],
seq: &[u8],
quality: &[u8],
) -> Result<(), GraphError> {
// Treats each quality byte as unsigned (0-255). C++'s `char` is signed on our targets, so
// a byte >= 128 would produce a different (negative) `q - 33` before its narrowing to
// uint32_t and thus diverge from the oracle. This is unreachable for valid Phred+33
// (always < 127); a future differential-fuzz test feeding garbage quality bytes would need
// to account for the sign difference here.
let weights: Vec<u32> = quality.iter().map(|&q| (q as i32 - 33) as u32).collect();
self.add_alignment(alignment, seq, &weights)
}
/// Computes a topological order of [`Graph::nodes`] into [`Graph::rank_to_node`].
///
/// Mirrors `spoa::Graph::TopologicalSort` (`graph.cpp:249-303`): an iterative (explicit-stack)
/// depth-first search over the graph, keyed by a per-node three-state `marks` array
/// (0 = unvisited, 1 = on-stack/in-progress, 2 = done/ranked) and a per-node `ignored` flag.
/// `ignored[n]` is set the moment `n` is discovered as another node's `aligned_nodes` entry;
/// an ignored node never emits its own rank, since its rank is instead emitted immediately
/// after the node that discovered it (see the `is_valid` branch below), keeping every
/// aligned-node group contiguous in `rank_to_node`.
///
/// For each node visited, unfinished (`marks != 2`) in-edge tails are pushed first, then —
/// only if the node itself is not ignored — unfinished aligned nodes are pushed (and marked
/// ignored). This exact push order is load-bearing: it determines the resulting rank order,
/// which later tasks' DP indexing and consensus tie-breaking depend on matching byte-for-byte.
fn topological_sort(&mut self) {
self.rank_to_node.clear();
let mut marks = vec![0u8; self.nodes.len()];
let mut ignored = vec![false; self.nodes.len()];
let mut stack: Vec<NodeId> = Vec::new();
for start in 0..self.nodes.len() {
if marks[start] != 0 {
continue;
}
stack.push(NodeId(start as u32));
while let Some(&curr) = stack.last() {
let curr_idx = curr.0 as usize;
let mut is_valid = true;
if marks[curr_idx] != 2 {
for &edge_id in &self.nodes[curr_idx].inedges {
let tail = self.edges[edge_id.0 as usize].tail;
if marks[tail.0 as usize] != 2 {
stack.push(tail);
is_valid = false;
}
}
if !ignored[curr_idx] {
for &aligned in &self.nodes[curr_idx].aligned_nodes {
if marks[aligned.0 as usize] != 2 {
stack.push(aligned);
ignored[aligned.0 as usize] = true;
is_valid = false;
}
}
}
debug_assert!(is_valid || marks[curr_idx] != 1, "Graph is not a DAG");
if is_valid {
marks[curr_idx] = 2;
if !ignored[curr_idx] {
self.rank_to_node.push(curr);
for &aligned in &self.nodes[curr_idx].aligned_nodes {
self.rank_to_node.push(aligned);
}
}
} else {
marks[curr_idx] = 1;
}
}
if is_valid {
stack.pop();
}
}
}
debug_assert!(
self.is_topologically_sorted(),
"Graph is not topologically sorted"
);
}
/// Generates the consensus sequence, keeping every node regardless of coverage.
///
/// Equivalent to `min_coverage = -1`: since [`Node::coverage`] is always `>= 0`, no node is
/// ever excluded. Mirrors `spoa::Graph::GenerateConsensus()` (`graph.cpp:368-374`), which
/// itself never filters by coverage.
///
/// # Note on run-length-sensitive data
/// The consensus is the graph's *heaviest bundle* — the path maximizing summed edge weight — so
/// across long homopolymer or low-complexity runs the consensus length can be inflated relative
/// to a per-column majority vote. This is correct spoa behavior. If you need run-length-faithful
/// output, prefer the column view ([`Graph::msa_columns`] / [`Graph::column_members`] /
/// [`Graph::generate_msa`]) and reduce per column yourself.
pub fn generate_consensus(&mut self) -> String {
self.generate_consensus_min_coverage(-1)
}
/// Generates the consensus sequence, dropping any consensus node whose [`Node::coverage`] is
/// below `min_coverage`.
///
/// Mirrors `spoa::Graph::GenerateConsensus(std::int32_t min_coverage)` (`graph.cpp:377-386`).
pub fn generate_consensus_min_coverage(&mut self, min_coverage: i32) -> String {
self.traverse_heaviest_bundle();
let mut dst = String::new();
for i in 0..self.consensus.len() {
let node_id = self.consensus[i];
let node = &self.nodes[node_id.0 as usize];
if node.coverage(self) as i32 >= min_coverage {
let code = node.code;
let byte = self.decoder[code as usize];
dst.push(byte as u8 as char);
}
}
dst
}
/// Maps every node id to its multiple-sequence-alignment column, folding each node's
/// `aligned_nodes` peers into the same column as their representative. Returns
/// `(node_id_to_column, row_size)`, where `row_size` is the total number of distinct MSA
/// columns (the number of aligned-node *groups* in topological order, not [`Graph::nodes`]'s
/// length).
///
/// Mirrors `spoa::Graph::InitializeMultipleSequenceAlignment` (`graph.cpp:321-337`). That
/// loop's C-style `for (i = 0; i < rank_to_node_.size(); ++i, ++j)` advances `j` once per
/// outer iteration, but its body *also* does `++i` once per aligned peer
/// (`graph.cpp:325-331`) to skip over the peers that `topological_sort` already placed
/// immediately after their representative in `rank_to_node`. A plain `for (i, node) in
/// rank_to_node.iter().enumerate()` cannot replicate that mid-loop `++i`, so this is ported
/// with an explicit `while` loop and a manually advanced `i`, matching both increments
/// exactly (see the module-level task notes on this double-increment).
fn initialize_msa(&self) -> (Vec<u32>, u32) {
let mut node_id_to_column = vec![0u32; self.nodes.len()];
let mut j: u32 = 0;
let mut i: usize = 0;
while i < self.rank_to_node.len() {
let node_id = self.rank_to_node[i];
node_id_to_column[node_id.0 as usize] = j;
for &aligned in &self.nodes[node_id.0 as usize].aligned_nodes {
node_id_to_column[aligned.0 as usize] = j;
i += 1;
}
i += 1;
j += 1;
}
(node_id_to_column, j)
}
/// Generates a multiple sequence alignment: one row per input sequence (in `add_alignment`
/// order), each padded to the same width with `'-'` gap characters, plus an optional
/// trailing consensus row.
///
/// Mirrors `spoa::Graph::GenerateMultipleSequenceAlignment` (`graph.cpp:339-366`). Each
/// sequence's row is built by walking its nodes via [`Node::successor`] (labeled by that
/// sequence's index) from its first node to its last, writing each visited node's decoded
/// symbol into that node's MSA column (from [`Graph::initialize_msa`]); columns the sequence
/// never visits stay `'-'`. When `include_consensus` is `true`,
/// [`Graph::traverse_heaviest_bundle`] is (re-)run and one more row is appended the same way,
/// from [`Graph::consensus`].
pub fn generate_msa(&mut self, include_consensus: bool) -> Vec<String> {
let (node_id_to_column, row_size) = self.initialize_msa();
let mut dst: Vec<String> = Vec::with_capacity(self.sequences.len() + 1);
for i in 0..self.sequences.len() {
let mut row = vec!['-'; row_size as usize];
for node_id in self.sequence_path_iter(i) {
let code = self.nodes[node_id.0 as usize].code;
let byte = self.decoder[code as usize];
row[node_id_to_column[node_id.0 as usize] as usize] = byte as u8 as char;
}
dst.push(row.into_iter().collect());
}
if include_consensus {
self.traverse_heaviest_bundle();
let mut row = vec!['-'; row_size as usize];
for i in 0..self.consensus.len() {
let node_id = self.consensus[i];
let code = self.nodes[node_id.0 as usize].code;
let byte = self.decoder[code as usize];
row[node_id_to_column[node_id.0 as usize] as usize] = byte as u8 as char;
}
dst.push(row.into_iter().collect());
}
dst
}
/// Finds the heaviest-weighted path through the graph (the "heaviest bundle") and stores it,
/// in traversal order, into [`Graph::consensus`].
///
/// Mirrors `spoa::Graph::TraverseHeaviestBundle` (`graph.cpp:465-509`). Runs a single
/// forward DP pass over [`Graph::rank_to_node`], scoring each node by the heaviest-weighted
/// path reaching it (`scores`) and recording that path's predecessor (`predecessors`); ties
/// are broken by preferring the inedge whose *tail's own* predecessor chain scored highest
/// (see the tie-break note on the inedge comparison below). If the best-scoring node found
/// (`max`) still has outgoing edges once the forward pass completes, the bundle has run into
/// a competing branch partway through, so [`Graph::branch_completion`] is called repeatedly
/// (invalidating and rescoring the losing branch) until `max` is a true sink. The consensus is
/// then read off by walking `predecessors` back from `max` to the bundle's start and
/// reversing.
fn traverse_heaviest_bundle(&mut self) {
if self.rank_to_node.is_empty() {
return;
}
let mut predecessors: Vec<Option<NodeId>> = vec![None; self.nodes.len()];
let mut scores: Vec<i64> = vec![-1; self.nodes.len()];
let mut max: Option<NodeId> = None;
for i in 0..self.rank_to_node.len() {
let node_id = self.rank_to_node[i];
let node_idx = node_id.0 as usize;
for &edge_id in &self.nodes[node_idx].inedges {
let edge = &self.edges[edge_id.0 as usize];
// Tie-break short-circuit (graph.cpp:477-478): mirrors C++'s
// `(scores[it->id] < jt->weight) || (scores[it->id] == jt->weight &&
// scores[predecessors[it->id]->id] <= scores[jt->tail->id])` exactly. The `&&`
// only evaluates its right side when the first `scores[it] < weight` comparison
// is false; C++ relies on this to dereference `predecessors[it->id]` safely
// (weights are positive and scores start at -1, so on a node's first inedge the
// first comparison is always true and the null predecessor is never touched).
// With `Option<NodeId>` we replicate that: only read `predecessors[node_idx]`
// (and `.expect` it to be `Some`) inside the branch that C++'s short-circuit
// guarantees is only reached once a predecessor already exists.
let take = if scores[node_idx] < edge.weight {
true
} else if scores[node_idx] == edge.weight {
let current_pred = predecessors[node_idx].expect(
"scores[node_idx] == edge.weight (both != -1's initial trivial case \
would have taken the first branch) implies a predecessor was already \
recorded",
);
scores[current_pred.0 as usize] <= scores[edge.tail.0 as usize]
} else {
false
};
if take {
scores[node_idx] = edge.weight;
predecessors[node_idx] = Some(edge.tail);
}
}
if let Some(pred) = predecessors[node_idx] {
scores[node_idx] += scores[pred.0 as usize];
}
if max.is_none() || scores[max.unwrap().0 as usize] < scores[node_idx] {
max = Some(node_id);
}
}
let mut max = max.expect("rank_to_node is non-empty, so max is always assigned");
if !self.nodes[max.0 as usize].outedges.is_empty() {
let mut node_id_to_rank = vec![0u32; self.nodes.len()];
for (rank, &node_id) in self.rank_to_node.iter().enumerate() {
node_id_to_rank[node_id.0 as usize] = rank as u32;
}
while !self.nodes[max.0 as usize].outedges.is_empty() {
let rank = node_id_to_rank[max.0 as usize];
max = self.branch_completion(rank, &mut scores, &mut predecessors);
}
}
// Traceback (graph.cpp:502-508).
self.consensus.clear();
let mut node = max;
while let Some(pred) = predecessors[node.0 as usize] {
self.consensus.push(node);
node = pred;
}
self.consensus.push(node);
self.consensus.reverse();
}
/// Resolves a heaviest-bundle traversal that ran into a competing branch at `rank`: the
/// branch not taken by `rank`'s node is invalidated (scores reset to `-1`), then every node
/// from `rank + 1` onward is rescored, skipping any inedge whose tail was invalidated (either
/// just now, or earlier in this same rescan). Returns the best-scoring node found in the
/// rescanned range.
///
/// Mirrors `spoa::Graph::BranchCompletion` (`graph.cpp:511-549`). The inedge tie-break
/// comparison is the same short-circuit as [`Graph::traverse_heaviest_bundle`]'s forward
/// pass; see that method's doc comment for why the predecessor dereference is safe.
fn branch_completion(
&self,
rank: u32,
scores: &mut [i64],
predecessors: &mut [Option<NodeId>],
) -> NodeId {
let start = self.rank_to_node[rank as usize];
for &out_edge_id in &self.nodes[start.0 as usize].outedges {
let head = self.edges[out_edge_id.0 as usize].head;
for &in_edge_id in &self.nodes[head.0 as usize].inedges {
let tail = self.edges[in_edge_id.0 as usize].tail;
if tail != start {
scores[tail.0 as usize] = -1;
}
}
}
let mut max: Option<NodeId> = None;
for i in (rank as usize + 1)..self.rank_to_node.len() {
let node_id = self.rank_to_node[i];
let node_idx = node_id.0 as usize;
scores[node_idx] = -1;
predecessors[node_idx] = None;
for &edge_id in &self.nodes[node_idx].inedges {
let edge = &self.edges[edge_id.0 as usize];
if scores[edge.tail.0 as usize] == -1 {
continue;
}
// Same tie-break short-circuit as traverse_heaviest_bundle (graph.cpp:534-535).
let take = if scores[node_idx] < edge.weight {
true
} else if scores[node_idx] == edge.weight {
let current_pred = predecessors[node_idx].expect(
"scores[node_idx] == edge.weight implies a predecessor was already \
recorded on an earlier (non-skipped) inedge",
);
scores[current_pred.0 as usize] <= scores[edge.tail.0 as usize]
} else {
false
};
if take {
scores[node_idx] = edge.weight;
predecessors[node_idx] = Some(edge.tail);
}
}
if let Some(pred) = predecessors[node_idx] {
scores[node_idx] += scores[pred.0 as usize];
}
if max.is_none() || scores[max.unwrap().0 as usize] < scores[node_idx] {
max = Some(node_id);
}
}
max.expect("rank + 1 < rank_to_node.len() whenever BranchCompletion is called, since it's only called while `max` still has outedges")
}
/// Emits the graph in GFA (Graphical Fragment Assembly) format: one `H` header line, one
/// `S`/`L` segment/link line per node/out-edge, and one `P` path line per input sequence
/// (plus, when `include_consensus` is `true`, one final `P\tConsensus\t...` line).
///
/// Mirrors `spoa::PrintGfa` (`third_party/spoa/src/main.cpp:123-203`), returning a `String`
/// instead of writing to `std::cout`. `headers[i]` becomes sequence `i`'s `P`-line name;
/// `is_reversed[i]`, if non-empty, reverses that same `P`-line's node path and flips every
/// node's orientation suffix from `+` to `-`. Node and edge ids are emitted 1-based
/// (`id + 1`), matching upstream's GFA convention (GFA disallows id `0`). A node (or a `L`
/// line whose *both* endpoints) that appears in [`Graph::consensus`] gets a trailing
/// `\tic:Z:true` tag; [`Graph::consensus`] must already be populated by a prior
/// [`Graph::generate_consensus`] (or a min-coverage variant) call, since this method only
/// reads it and never recomputes it.
///
/// # Panics (debug only)
///
/// Debug-asserts that `headers` has at least one entry per sequence, and that
/// `is_reversed` (if non-empty) does too — mirroring upstream's
/// `[spoa::PrintGfa] error: missing header(s)` early return, but as a precondition rather
/// than a silent empty-string result, since this library's callers (the oracle, and later
/// the CLI) always supply enough of both.
pub fn to_gfa(
&self,
headers: &[String],
is_reversed: &[bool],
include_consensus: bool,
) -> String {
debug_assert!(
headers.len() >= self.sequences.len(),
"to_gfa: missing header(s): {} header(s) for {} sequence(s)",
headers.len(),
self.sequences.len()
);
debug_assert!(
is_reversed.is_empty() || is_reversed.len() >= self.sequences.len(),
"to_gfa: missing reversion flag(s): {} flag(s) for {} sequence(s)",
is_reversed.len(),
self.sequences.len()
);
let mut is_consensus_node = vec![false; self.nodes.len()];
for &node_id in &self.consensus {
is_consensus_node[node_id.0 as usize] = true;
}
let mut out = String::new();
out.push_str("H\tVN:Z:1.0\n");
for (id, node) in self.nodes.iter().enumerate() {
let symbol = self.decoder[node.code as usize] as u8 as char;
out.push_str(&format!("S\t{}\t{symbol}", id + 1));
if is_consensus_node[id] {
out.push_str("\tic:Z:true");
}
out.push('\n');
for &edge_id in &node.outedges {
let edge = &self.edges[edge_id.0 as usize];
let head_id = edge.head.0 as usize;
out.push_str(&format!(
"L\t{}\t+\t{}\t+\tOM\tew:f:{}",
id + 1,
head_id + 1,
edge.weight
));
if is_consensus_node[id] && is_consensus_node[head_id] {
out.push_str("\tic:Z:true");
}
out.push('\n');
}
}
for i in 0..self.sequences.len() {
out.push_str(&format!("P\t{}\t", headers[i]));
// 1-based node ids along this sequence's path (see `sequence_path_iter`).
let mut path: Vec<u32> = self.sequence_path_iter(i).map(|node| node.0 + 1).collect();
let reversed = !is_reversed.is_empty() && is_reversed[i];
if reversed {
path.reverse();
}
let sign = if reversed { '-' } else { '+' };
for (j, node_id) in path.iter().enumerate() {
if j != 0 {
out.push(',');
}
out.push_str(&format!("{node_id}{sign}"));
}
out.push_str("\t*\n");
}
if include_consensus {
out.push_str("P\tConsensus\t");
for (i, &node_id) in self.consensus.iter().enumerate() {
if i != 0 {
out.push(',');
}
out.push_str(&format!("{}+", node_id.0 + 1));
}
out.push_str("\t*\n");
}
out
}
/// Emits the graph in Graphviz DOT format.
///
/// Mirrors `spoa::Graph::PrintDot` (`third_party/spoa/src/graph.cpp:640-680`), returning a
/// `String` instead of writing to a file. Node and edge ids are emitted 0-based (unlike
/// [`Graph::to_gfa`]'s 1-based ids). A node in [`Graph::consensus`] is filled `goldenrod1`;
/// an edge between two CONSECUTIVE consensus nodes (`consensus_rank[tail] + 1 ==
/// consensus_rank[head]`, `-1` for non-consensus nodes — reproduced verbatim, including its
/// edge case where a non-consensus tail's `-1 + 1 == 0` can coincidentally match a
/// consensus head at rank 0) is colored `goldenrod1`; and each aligned-node pair (same MSA
/// column, different symbol) is linked once by a dotted, arrowhead-less edge, guarded by
/// `jt.id > it.id` so each pair is only emitted from its lower-id member. [`Graph::consensus`]
/// must already be populated by a prior [`Graph::generate_consensus`] (or a min-coverage
/// variant) call, since this method only reads it and never recomputes it.
pub fn to_dot(&self) -> String {
let mut consensus_rank: Vec<i32> = vec![-1; self.nodes.len()];
for (rank, &node_id) in self.consensus.iter().enumerate() {
consensus_rank[node_id.0 as usize] = rank as i32;
}
let mut out = String::new();
out.push_str(&format!(
"digraph {} {{\n graph [rankdir = LR]\n",
self.sequences.len()
));
for (id, node) in self.nodes.iter().enumerate() {
let symbol = self.decoder[node.code as usize] as u8 as char;
out.push_str(&format!(" {id}[label = \"{id} - {symbol}\""));
if consensus_rank[id] != -1 {
out.push_str(", style = filled, fillcolor = goldenrod1");
}
out.push_str("]\n");
for &edge_id in &node.outedges {
let edge = &self.edges[edge_id.0 as usize];
let head_id = edge.head.0 as usize;
out.push_str(&format!(" {id} -> {head_id} [label = \"{}\"", edge.weight));
if consensus_rank[id] + 1 == consensus_rank[head_id] {
out.push_str(", color = goldenrod1");
}
out.push_str("]\n");
}
for &aligned in &node.aligned_nodes {
let aligned_id = aligned.0 as usize;
if aligned_id > id {
out.push_str(&format!(
" {id} -> {aligned_id} [style = dotted, arrowhead = none]\n"
));
}
}
}
out.push_str("}\n");
out
}
/// Returns whether [`Graph::rank_to_node`] is a valid topological order of [`Graph::nodes`]:
/// every node's in-edge tails appear at an earlier rank than the node itself.
///
/// Mirrors `spoa::Graph::IsTopologicallySorted` (`graph.cpp:305-319`).
pub fn is_topologically_sorted(&self) -> bool {
debug_assert!(
self.nodes.len() == self.rank_to_node.len(),
"Topological sort not called"
);
let mut visited = vec![false; self.nodes.len()];
for &node_id in &self.rank_to_node {
let node = &self.nodes[node_id.0 as usize];
for &edge_id in &node.inedges {
let tail = self.edges[edge_id.0 as usize].tail;
if !visited[tail.0 as usize] {
return false;
}
}
visited[node_id.0 as usize] = true;
}
true
}
}
#[cfg(test)]
mod tests {
use super::*;
#[test]
fn add_node_assigns_sequential_ids() {
let mut g = Graph::new();
let a = g.add_node(0);
let b = g.add_node(1);
assert_eq!(a.0, 0);
assert_eq!(b.0, 1);
assert_eq!(g.nodes.len(), 2);
}
/// The public read-only accessors expose a coherent view of a built graph: the arena slices
/// and their id-indexed lookups agree, codes round-trip through `encode`/`decode`, and the
/// rank/sequence/consensus views are populated and decodable.
#[test]
fn public_accessors_expose_a_coherent_view_of_a_built_graph() {
let mut g = Graph::new();
// First sequence seeds the linear chain (nodes 0,1,2); the second is aligned onto it
// position-for-position so the two merge into one coverage-2 chain rather than a disjoint
// second chain.
g.add_alignment(&[], b"ACG", &[1, 1, 1]).unwrap();
g.add_alignment(&[(0, 0), (1, 1), (2, 2)], b"ACG", &[1, 1, 1])
.unwrap();
// Arena slices and id lookups agree.
assert_eq!(g.num_nodes(), g.nodes().len());
assert_eq!(g.num_edges(), g.edges().len());
assert_eq!(g.num_nodes(), 3);
for (i, node) in g.nodes().iter().enumerate() {
let id = NodeId(i as u32);
assert_eq!(g.node(id).code, node.code);
}
for (i, edge) in g.edges().iter().enumerate() {
let id = EdgeId(i as u32);
assert_eq!(g.edge(id).tail, edge.tail);
}
// Codes round-trip; the alphabet is exactly {A, C, G}.
assert_eq!(g.num_codes(), 3);
for &base in b"ACG" {
let code = g.encode(base).expect("base was added");
assert_eq!(g.decode(code), Some(base));
}
assert_eq!(g.encode(b'T'), None); // never seen
assert_eq!(g.decode(999), None); // never assigned
// Node::base decodes each node back to its input byte, in rank order spelling "ACG".
let spelled: Vec<u8> = g
.rank_order()
.iter()
.map(|&id| g.node(id).base(&g).expect("node code is decodable"))
.collect();
assert_eq!(spelled, b"ACG");
assert_eq!(g.rank_order().len(), 3);
// One start per added sequence; every node has coverage 2 (both sequences traverse it).
assert_eq!(g.sequence_starts().len(), 2);
for node in g.nodes() {
assert_eq!(node.coverage(&g), 2);
}
// Consensus is empty until generated, then decodes to the same String the API returns.
assert!(g.consensus_nodes().is_empty());
let consensus = g.generate_consensus();
let from_nodes: String = g
.consensus_nodes()
.iter()
.map(|&id| g.node(id).base(&g).expect("node code is decodable") as char)
.collect();
assert_eq!(from_nodes, consensus);
assert_eq!(consensus, "ACG");
}
/// Builds a small graph with one aligned (substituted) column so the MSA has a folded column,
/// and exercises the RELAY accessors: `sequence_path` decodes back to each input;
/// `msa_columns`/`column_members` agree and fold aligned peers; `Node::labels` matches coverage.
#[test]
fn relay_accessors_map_sequences_to_msa_columns() {
// "ACGT" then "ACTT" aligned onto it: position 2 diverges (G vs T) into an aligned column.
let mut g = Graph::new();
g.add_alignment(&[], b"ACGT", &[1, 1, 1, 1]).unwrap();
g.add_alignment(&[(0, 0), (1, 1), (2, 2), (3, 3)], b"ACTT", &[1, 1, 1, 1])
.unwrap();
assert_eq!(g.sequence_starts().len(), 2);
// sequence_path (and its iterator) decode back to the exact input bytes, indexed by the
// add order (seq 0 = "ACGT", seq 1 = "ACTT").
let decode_path = |g: &Graph, i: usize| -> Vec<u8> {
g.sequence_path(i)
.iter()
.map(|&n| g.node(n).base(g).unwrap())
.collect()
};
assert_eq!(decode_path(&g, 0), b"ACGT");
assert_eq!(decode_path(&g, 1), b"ACTT");
// The iterator form yields the same node sequence.
assert_eq!(
g.sequence_path_iter(0).collect::<Vec<_>>(),
g.sequence_path(0)
);
// msa_columns: 4 columns; the divergent G and T share their column (aligned peers folded).
let (node_to_col, row_size) = g.msa_columns();
assert_eq!(row_size, 4);
let path0 = g.sequence_path(0); // A C G T
let path1 = g.sequence_path(1); // A C T T
assert_eq!(
node_to_col[path0[0].0 as usize],
node_to_col[path1[0].0 as usize]
); // shared A
assert_eq!(
node_to_col[path0[2].0 as usize],
node_to_col[path1[2].0 as usize]
); // G/T folded
assert_ne!(path0[2], path1[2]); // ...but they are distinct nodes
// column_members inverts the map: the divergent column lists both sequences; column 0 too.
let columns = g.column_members();
assert_eq!(columns.len(), row_size as usize);
let divergent = node_to_col[path0[2].0 as usize] as usize;
let seqs_in_divergent: Vec<u32> = columns[divergent].iter().map(|&(s, _)| s).collect();
assert_eq!(seqs_in_divergent, vec![0, 1]);
// Every (seq, node) entry actually sits in the column it is filed under.
for (col, members) in columns.iter().enumerate() {
for &(_seq, node) in members {
assert_eq!(node_to_col[node.0 as usize] as usize, col);
}
}
// Node::labels == the set coverage counts, sorted; the shared "A" node carries both labels.
let a_node = path0[0];
assert_eq!(g.node(a_node).labels(&g), vec![0, 1]);
for node in g.nodes() {
assert_eq!(node.labels(&g).len() as u32, node.coverage(&g));
}
}
/// The MSA row order matches the `sequence_path`/`sequence_starts` index order (RELAY #3 doc
/// guarantee), and each MSA row's non-gap characters decode that sequence.
#[test]
fn generate_msa_row_order_matches_sequence_index() {
let mut g = Graph::new();
g.add_alignment(&[], b"ACGT", &[1, 1, 1, 1]).unwrap();
g.add_alignment(&[(0, 0), (1, 1), (2, 2), (3, 3)], b"ACTT", &[1, 1, 1, 1])
.unwrap();
let msa = g.generate_msa(false);
assert_eq!(msa.len(), 2);
for (i, row) in msa.iter().enumerate() {
let ungapped: String = row.chars().filter(|&c| c != '-').collect();
let expected: String = g
.sequence_path(i)
.iter()
.map(|&n| g.node(n).base(&g).unwrap() as char)
.collect();
assert_eq!(ungapped, expected, "MSA row {i} must spell sequence {i}");
}
}
#[test]
fn add_edge_dedups_repeated_tail_head_pairs() {
let mut g = Graph::new();
let a = g.add_node(0);
let b = g.add_node(1);
g.add_edge(a, b, 1, 0);
g.add_edge(a, b, 1, 1);
assert_eq!(
g.edges.len(),
1,
"second add_edge should reuse the existing edge"
);
let edge = &g.edges[0];
assert_eq!(edge.labels, vec![0, 1]);
assert_eq!(edge.weight, 2);
assert_eq!(g.nodes[a.0 as usize].outedges, vec![EdgeId(0)]);
assert_eq!(g.nodes[b.0 as usize].inedges, vec![EdgeId(0)]);
}
/// Locks the tie-break short-circuit in `traverse_heaviest_bundle`'s inedge comparison
/// (mirroring `graph.cpp:477-478`) independent of the oracle. Builds a diamond
/// `S -> A -> M` / `S -> B -> M` where `A->M` and `B->M` have EQUAL weight but `A`'s
/// cumulative score (via `S->A`) is far heavier than `B`'s (via `S->B`), so the correct
/// winner at the tie is decided by comparing `scores[predecessors[M]]` against
/// `scores[jt.tail]` — not by the local edge weight (which is tied) and not by insertion
/// order. Exercised with the two inedges added in BOTH orders to confirm the outcome (and
/// the absence of any panic from an eager predecessor `.unwrap()`) is order-independent.
#[test]
fn traverse_heaviest_bundle_breaks_equal_weight_inedge_ties_by_predecessor_score() {
for reversed in [false, true] {
let mut g = Graph::new();
let s = g.add_node(0);
let a = g.add_node(1);
let b = g.add_node(2);
let m = g.add_node(3);
// Decoder entries so `generate_consensus` produces readable output.
g.decoder[0] = b'S' as i32;
g.decoder[1] = b'A' as i32;
g.decoder[2] = b'B' as i32;
g.decoder[3] = b'M' as i32;
g.add_edge(s, a, 100, 0); // heavy branch into A
g.add_edge(s, b, 1, 1); // light branch into B
if reversed {
g.add_edge(b, m, 5, 1);
g.add_edge(a, m, 5, 0); // equal weight to B->M, added second
} else {
g.add_edge(a, m, 5, 0);
g.add_edge(b, m, 5, 1); // equal weight to A->M, added second
}
g.topological_sort();
let consensus = g.generate_consensus();
// The heavier-scoring predecessor (A, via the 100-weight S->A edge) must win the
// tie at M regardless of which equal-weight inedge (A->M or B->M) was added first;
// B (the lighter branch) must NOT appear in the consensus.
assert_eq!(
consensus, "SAM",
"reversed={reversed}: tie must resolve to the heavier predecessor (A), not \
insertion order"
);
}
}
#[test]
fn coverage_counts_distinct_labels_across_in_and_out_edges() {
// a -[label 0]-> b -[label 0,1]-> c
let mut g = Graph::new();
let a = g.add_node(0);
let b = g.add_node(1);
let c = g.add_node(2);
g.add_edge(a, b, 1, 0);
g.add_edge(b, c, 1, 0);
g.add_edge(b, c, 1, 1);
assert_eq!(g.nodes[a.0 as usize].coverage(&g), 1);
assert_eq!(g.nodes[b.0 as usize].coverage(&g), 2);
assert_eq!(g.nodes[c.0 as usize].coverage(&g), 2);
}
#[test]
fn add_alignment_weight_with_empty_alignment_builds_linear_chain() {
// First sequence into an empty graph always takes the empty-alignment fast path
// (graph.cpp:175-179): a straight-line chain of one node per base.
let mut g = Graph::new();
g.add_alignment_weight(&[], b"ACGT", 1).unwrap();
assert_eq!(g.nodes.len(), 4, "one node per base");
assert_eq!(g.edges.len(), 3, "one edge between consecutive bases");
assert_eq!(g.num_codes, 4, "four distinct symbols registered");
assert_eq!(g.sequences.len(), 1, "one sequence recorded");
assert_eq!(g.sequences[0], NodeId(0));
// Interior edges sum the weights of both endpoints, matching AddSequence
// (graph.cpp:104-106).
for edge in &g.edges {
assert_eq!(edge.weight, 2);
assert_eq!(edge.labels, vec![0]);
}
}
#[test]
fn add_alignment_weight_aligns_mismatch_via_aligned_nodes_cross_link() {
// Second sequence "AG" aligned against first sequence "AC": position 0 (A) matches an
// existing node, position 1 (G) mismatches the existing C node and must fork into a new,
// cross-linked aligned node (graph.cpp:207-231).
let mut g = Graph::new();
g.add_alignment_weight(&[], b"AC", 1).unwrap();
let node_a = NodeId(0);
let node_c = NodeId(1);
g.add_alignment_weight(&[(0, 0), (1, 1)], b"AG", 1).unwrap();
assert_eq!(g.nodes.len(), 3, "A, C, and a new G node");
let node_g = NodeId(2);
assert_eq!(
g.nodes[node_g.0 as usize].code,
g.coder[b'G' as usize] as u32
);
// Aligned-node cross-linking: C and G recorded as aligned to each other.
assert_eq!(g.nodes[node_c.0 as usize].aligned_nodes, vec![node_g]);
assert_eq!(g.nodes[node_g.0 as usize].aligned_nodes, vec![node_c]);
// Two sequences recorded, both starting at node A.
assert_eq!(g.sequences, vec![node_a, node_a]);
// A->C edge (from the first sequence) is untouched; a new A->G edge carries the second
// sequence's label and summed weight.
let edge_ac = g
.edges
.iter()
.find(|e| e.tail == node_a && e.head == node_c)
.expect("A->C edge exists");
assert_eq!(edge_ac.labels, vec![0]);
assert_eq!(edge_ac.weight, 2);
let edge_ag = g
.edges
.iter()
.find(|e| e.tail == node_a && e.head == node_g)
.expect("A->G edge exists");
assert_eq!(edge_ag.labels, vec![1]);
assert_eq!(edge_ag.weight, 2);
}
#[test]
fn add_alignment_weight_threads_unaligned_prefix_and_suffix() {
// Seed: "ACGT" -> nodes 0=A,1=C,2=G,3=T; codes A=0,C=1,G=2,T=3; sequences=[0].
let mut g = Graph::new();
g.add_alignment_weight(&[], b"ACGT", 1).unwrap();
// Second sequence "TACGTA" maps ONLY its middle 'C' (seq index 2) to the existing C node
// (node 1), forcing BOTH an unaligned prefix ("TA", indices 0-1) and an unaligned suffix
// ("GTA", indices 3-5) through `add_sequence` (graph.cpp:196-243).
g.add_alignment_weight(&[(1, 2)], b"TACGTA", 1).unwrap();
// 4 seed + 2 prefix + 3 suffix = 9 nodes; the 'C' column reuses the existing node.
assert_eq!(g.nodes.len(), 9);
let node_c = NodeId(1);
let (prefix_t, prefix_a) = (NodeId(4), NodeId(5));
let (suffix_g, suffix_t, suffix_a) = (NodeId(6), NodeId(7), NodeId(8));
// Returned start node is the prefix's FIRST node (graph.cpp:244, `begin`).
assert_eq!(g.sequences, vec![NodeId(0), prefix_t]);
// Node codes of the freshly created prefix/suffix nodes.
assert_eq!(
g.nodes[prefix_t.0 as usize].code,
g.coder[b'T' as usize] as u32
);
assert_eq!(
g.nodes[prefix_a.0 as usize].code,
g.coder[b'A' as usize] as u32
);
assert_eq!(
g.nodes[suffix_g.0 as usize].code,
g.coder[b'G' as usize] as u32
);
assert_eq!(
g.nodes[suffix_t.0 as usize].code,
g.coder[b'T' as usize] as u32
);
assert_eq!(
g.nodes[suffix_a.0 as usize].code,
g.coder[b'A' as usize] as u32
);
// Every new edge belongs to sequence 1 with the summed adjacent-base weight (1+1=2).
let find_edge = |tail: NodeId, head: NodeId| {
g.edges
.iter()
.find(|e| e.tail == tail && e.head == head)
.unwrap_or_else(|| panic!("edge {tail:?}->{head:?} exists"))
};
// Prefix chain, then the seam edge into the reused 'C' node.
assert_eq!(find_edge(prefix_t, prefix_a).labels, vec![1]);
assert_eq!(find_edge(prefix_t, prefix_a).weight, 2);
assert_eq!(find_edge(prefix_a, node_c).labels, vec![1]);
assert_eq!(find_edge(prefix_a, node_c).weight, 2);
// Seam edge out of the reused 'C' node into the suffix chain, then the suffix chain.
assert_eq!(find_edge(node_c, suffix_g).labels, vec![1]);
assert_eq!(find_edge(node_c, suffix_g).weight, 2);
assert_eq!(find_edge(suffix_g, suffix_t).labels, vec![1]);
assert_eq!(find_edge(suffix_g, suffix_t).weight, 2);
assert_eq!(find_edge(suffix_t, suffix_a).labels, vec![1]);
assert_eq!(find_edge(suffix_t, suffix_a).weight, 2);
// The reused 'C' node now sits on both sequences: original in/out plus the new seams.
assert_eq!(g.nodes[node_c.0 as usize].inedges.len(), 2);
assert_eq!(g.nodes[node_c.0 as usize].outedges.len(), 2);
}
#[test]
fn add_alignment_rejects_unequal_weights() {
let mut g = Graph::new();
let err = g.add_alignment(&[], b"ACGT", &[1, 2, 3]).unwrap_err();
assert!(matches!(err, GraphError::UnequalWeights));
}
#[test]
fn add_alignment_rejects_alignment_missing_sequence_positions() {
let mut g = Graph::new();
g.add_alignment_weight(&[], b"AC", 1).unwrap();
let err = g
.add_alignment_weight(&[(0, -1), (1, -1)], b"AC", 1)
.unwrap_err();
assert!(matches!(err, GraphError::MissingSequence));
}
#[test]
fn add_alignment_rejects_out_of_range_sequence_index() {
let mut g = Graph::new();
let err = g.add_alignment_weight(&[(-1, 5)], b"AC", 1).unwrap_err();
assert!(matches!(err, GraphError::InvalidAlignment));
}
#[test]
fn add_alignment_rejects_out_of_range_node_index() {
// `node_idx` flows straight into `self.nodes[..]`; out-of-range values (too large for
// the current node count, or below the `-1` "new node" sentinel) must return
// InvalidAlignment rather than panic on the index.
let mut g = Graph::new();
assert!(matches!(
g.add_alignment_weight(&[(5, 0)], b"AC", 1).unwrap_err(),
GraphError::InvalidAlignment
));
assert!(matches!(
g.add_alignment_weight(&[(-2, 0)], b"AC", 1).unwrap_err(),
GraphError::InvalidAlignment
));
}
#[test]
fn initialize_msa_folds_aligned_peers_into_one_column() {
// Seed "AC" (A->C), then align "AG" (forks C's mismatch into a new aligned node G,
// graph.cpp:207-231), then align "AT" (extends the SAME aligned group by matching
// against C's existing aligned_nodes, cross-linking a third node T to both C and G).
// C, G, and T end up topo-adjacent in `rank_to_node` (aligned peers are emitted
// immediately after their representative by `topological_sort`), which is exactly the
// shape that exercises `InitializeMultipleSequenceAlignment`'s double-increment
// (graph.cpp:325-331: the outer loop's `++i` PLUS the inner aligned_nodes loop's own
// `++i` per peer) — a naive `enumerate()` port would instead give C, G, and T three
// separate columns.
let mut g = Graph::new();
g.add_alignment_weight(&[], b"AC", 1).unwrap();
g.add_alignment_weight(&[(0, 0), (1, 1)], b"AG", 1).unwrap();
g.add_alignment_weight(&[(0, 0), (1, 1)], b"AT", 1).unwrap();
assert_eq!(g.nodes.len(), 4, "sanity: A, C, G, T");
let node_a = NodeId(0);
let node_c = NodeId(1);
let node_g = NodeId(2);
let node_t = NodeId(3);
assert_eq!(
g.nodes[node_c.0 as usize].aligned_nodes.len(),
2,
"sanity: C's aligned group grew to include both G and T"
);
let (node_id_to_column, row_size) = g.initialize_msa();
assert_eq!(
row_size, 2,
"row_size must count distinct columns (A's, and the aligned C/G/T group's), not \
the 4 individual nodes"
);
assert_eq!(node_id_to_column[node_a.0 as usize], 0);
let aligned_column = node_id_to_column[node_c.0 as usize];
assert_ne!(aligned_column, node_id_to_column[node_a.0 as usize]);
assert_eq!(
node_id_to_column[node_g.0 as usize], aligned_column,
"G must share C's column"
);
assert_eq!(
node_id_to_column[node_t.0 as usize], aligned_column,
"T must share C's column"
);
}
/// Builds a small, deterministic (tie-free) graph: three sequences "AC", "AC", "AG" fed
/// through `add_alignment_weight` in that order. The two "AC" sequences fold their C's
/// into one shared node and double A->C's weight (4) over A->G's (2), so
/// `traverse_heaviest_bundle`'s heaviest-bundle consensus is unambiguously "AC" (node G is
/// never a tie candidate) — letting the GFA/DOT unit tests below assert an exact, fully
/// predictable expected string without depending on any tie-break rule.
fn small_gfa_dot_graph() -> Graph {
let mut g = Graph::new();
g.add_alignment_weight(&[], b"AC", 1).unwrap();
g.add_alignment_weight(&[(0, 0), (1, 1)], b"AC", 1).unwrap();
g.add_alignment_weight(&[(0, 0), (1, 1)], b"AG", 1).unwrap();
g
}
#[test]
fn to_gfa_emits_s_l_p_lines_with_one_based_ids_and_consensus_tags() {
let mut g = small_gfa_dot_graph();
assert_eq!(
g.generate_consensus(),
"AC",
"sanity: deterministic consensus"
);
let headers = vec!["s0".to_string(), "s1".to_string(), "s2".to_string()];
let gfa = g.to_gfa(&headers, &[], true);
let expected = concat!(
"H\tVN:Z:1.0\n",
"S\t1\tA\tic:Z:true\n",
"L\t1\t+\t2\t+\tOM\tew:f:4\tic:Z:true\n",
"L\t1\t+\t3\t+\tOM\tew:f:2\n",
"S\t2\tC\tic:Z:true\n",
"S\t3\tG\n",
"P\ts0\t1+,2+\t*\n",
"P\ts1\t1+,2+\t*\n",
"P\ts2\t1+,3+\t*\n",
"P\tConsensus\t1+,2+\t*\n",
);
assert_eq!(gfa, expected);
}
#[test]
fn to_gfa_reverses_path_and_flips_sign_when_is_reversed_is_set() {
// Single sequence "ACG" -> nodes 1,2,3 (1-based). With `is_reversed[0] = true`, the P-line
// path must be walked, then REVERSED, with every node's orientation suffix flipped from
// `+` to `-` — mirroring `PrintGfa`'s `std::reverse(path)` + `(ir ? "-" : "+")`
// (main.cpp:180-189). This exercises the reversal/sign-flip branch the corpus/`&[]` tests
// never hit.
let mut g = Graph::new();
g.add_alignment_weight(&[], b"ACG", 1).unwrap();
// No consensus generated: `consensus` stays empty, so no `ic:Z:true` tags appear and the
// P-line reversal is the only thing under test.
let headers = vec!["only".to_string()];
let gfa = g.to_gfa(&headers, &[true], false);
let expected = concat!(
"H\tVN:Z:1.0\n",
"S\t1\tA\n",
"L\t1\t+\t2\t+\tOM\tew:f:2\n",
"S\t2\tC\n",
"L\t2\t+\t3\t+\tOM\tew:f:2\n",
"S\t3\tG\n",
"P\tonly\t3-,2-,1-\t*\n",
);
assert_eq!(gfa, expected);
}
#[test]
fn to_dot_fills_consensus_nodes_colors_consensus_edges_and_dots_aligned_pairs() {
let mut g = small_gfa_dot_graph();
assert_eq!(
g.generate_consensus(),
"AC",
"sanity: deterministic consensus"
);
let dot = g.to_dot();
let expected = concat!(
"digraph 3 {\n",
" graph [rankdir = LR]\n",
" 0[label = \"0 - A\", style = filled, fillcolor = goldenrod1]\n",
" 0 -> 1 [label = \"4\", color = goldenrod1]\n",
" 0 -> 2 [label = \"2\"]\n",
" 1[label = \"1 - C\", style = filled, fillcolor = goldenrod1]\n",
" 1 -> 2 [style = dotted, arrowhead = none]\n",
" 2[label = \"2 - G\"]\n",
"}\n",
);
assert_eq!(dot, expected);
}
#[test]
fn topological_sort_ranks_every_node_including_aligned_groups() {
// Build a branching graph: seed "AC" (A->C), then align "AG" so its 'A' reuses the
// existing A node but its 'G' mismatches the existing C node and forks into a new,
// cross-linked aligned node (graph.cpp:207-231). This gives real in-edges (A->C, A->G)
// plus a genuine aligned-node group ({C, G}), which is exactly the shape TopologicalSort
// must handle: `add_alignment` already invokes `topological_sort` internally, so
// `rank_to_node` should come out fully populated and valid without any direct call here.
let mut g = Graph::new();
g.add_alignment_weight(&[], b"AC", 1).unwrap();
g.add_alignment_weight(&[(0, 0), (1, 1)], b"AG", 1).unwrap();
assert_eq!(
g.rank_to_node.len(),
g.nodes.len(),
"every node must be assigned a rank"
);
assert!(
g.is_topologically_sorted(),
"rank_to_node must be a valid topological order"
);
}
}