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#include "ContigID.h"
#include "ContigPath.h"
#include "ContigProperties.h"
#include "DataBase/DB.h"
#include "DataBase/Options.h"
#include "Functional.h"
#include "Graph/ContigGraph.h"
#include "Graph/ContigGraphAlgorithms.h"
#include "Graph/DirectedGraph.h"
#include "Graph/GraphIO.h"
#include "IOUtil.h"
#include "Uncompress.h"
#include "config.h"
#include <algorithm>
#include <cassert>
#include <cerrno>
#include <cstdlib>
#include <cstring> // for strerror
#include <fstream>
#include <functional>
#include <getopt.h>
#include <iostream>
#include <map>
#include <vector>
using namespace std;
#define PROGRAM "PathOverlap"
DB db;
static const char* VERSION_MESSAGE =
PROGRAM " (ABySS) " VERSION "\n"
"Written by Shaun Jackman and Tony Raymond.\n"
"\n"
"Copyright 2014 Canada's Michael Smith Genome Sciences Centre\n";
static const char* USAGE_MESSAGE =
"Usage: " PROGRAM " -k<kmer> [OPTION]... ADJ PATH\n"
"Find paths that overlap. Either output the graph of overlapping\n"
"paths, assemble overlapping paths into larger paths, or trim the\n"
"overlapping paths.\n"
"\n"
" Arguments:\n"
"\n"
" ADJ contig adjacency graph\n"
" PATH sequences of contig IDs\n"
"\n"
" Options:\n"
"\n"
" -k, --kmer=N k-mer size\n"
" -g, --graph=FILE write the contig adjacency graph to FILE\n"
" -r, --repeats=FILE write repeat contigs to FILE\n"
" --overlap find overlapping paths [default]\n"
" --assemble assemble overlapping paths\n"
" --trim trim overlapping paths\n"
" --adj output the graph in ADJ format [default]\n"
" --asqg output the graph in ASQG format\n"
" --dot output the graph in GraphViz format\n"
" --gfa output the graph in GFA1 format\n"
" --gfa1 output the graph in GFA1 format\n"
" --gfa2 output the graph in GFA2 format\n"
" --gv output the graph in GraphViz format\n"
" --sam output the graph in SAM format\n"
" --SS expect contigs to be oriented correctly\n"
" --no-SS no assumption about contig orientation [default]\n"
" -v, --verbose display verbose output\n"
" --help display this help and exit\n"
" --version output version information and exit\n"
" --db=FILE specify path of database repository in FILE\n"
" --library=NAME specify library NAME for sqlite\n"
" --strain=NAME specify strain NAME for sqlite\n"
" --species=NAME specify species NAME for sqlite\n"
"\n"
"Report bugs to <" PACKAGE_BUGREPORT ">.\n";
namespace opt {
string db;
dbVars metaVars;
unsigned k;
/** Output format. */
int format; // used by ContigProperties
/** Write the contig adjacency graph to this file. */
static string graphPath;
/** Output the IDs of contigs in overlaps to this file. */
static string repeatContigs;
/** Run a strand-specific RNA-Seq assembly. */
static int ss;
/** Mode of operation. */
enum
{
/** Find overlapping paths, do not assemble. */
OVERLAP,
/** Assemble overlapping paths. */
ASSEMBLE,
/** Trim overlapping paths. */
TRIM,
};
static int mode;
static int verbose;
}
static const char* shortopts = "g:k:r:v";
enum
{
OPT_HELP = 1,
OPT_VERSION,
OPT_DB,
OPT_LIBRARY,
OPT_STRAIN,
OPT_SPECIES
};
// enum { OPT_HELP = 1, OPT_VERSION };
static const struct option longopts[] = { { "graph", required_argument, NULL, 'g' },
{ "kmer", required_argument, NULL, 'k' },
{ "assemble", no_argument, &opt::mode, opt::ASSEMBLE },
{ "overlap", no_argument, &opt::mode, opt::OVERLAP },
{ "trim", no_argument, &opt::mode, opt::TRIM },
{ "adj", no_argument, &opt::format, ADJ },
{ "asqg", no_argument, &opt::format, ASQG },
{ "dot", no_argument, &opt::format, DOT },
{ "gfa", no_argument, &opt::format, GFA1 },
{ "gfa1", no_argument, &opt::format, GFA1 },
{ "gfa2", no_argument, &opt::format, GFA2 },
{ "gv", no_argument, &opt::format, DOT },
{ "sam", no_argument, &opt::format, SAM },
{ "SS", no_argument, &opt::ss, 1 },
{ "no-SS", no_argument, &opt::ss, 0 },
{ "repeats", required_argument, NULL, 'r' },
{ "verbose", no_argument, NULL, 'v' },
{ "help", no_argument, NULL, OPT_HELP },
{ "version", no_argument, NULL, OPT_VERSION },
{ "db", required_argument, NULL, OPT_DB },
{ "library", required_argument, NULL, OPT_LIBRARY },
{ "strain", required_argument, NULL, OPT_STRAIN },
{ "species", required_argument, NULL, OPT_SPECIES },
{ NULL, 0, NULL, 0 } };
/** A vertex of the overlap graph. */
struct Vertex
{
unsigned id;
bool sense;
/** The number of single-end contigs. */
static unsigned s_offset;
Vertex(unsigned id, bool sense)
: id(id)
, sense(sense)
{}
bool operator==(const Vertex& v) const { return id == v.id && sense == v.sense; }
ContigNode descriptor() const { return ContigNode(s_offset + id, sense); }
};
unsigned Vertex::s_offset;
/** An alignment of two overlapping contigs. */
struct Overlap
{
Vertex source;
Vertex target;
/** Overlap measured in number of contigs. */
unsigned overlap;
/** Overlap measured in bp. */
int distance;
Overlap(const Vertex& source, const Vertex& target, unsigned overlap, int distance)
: source(source)
, target(target)
, overlap(overlap)
, distance(distance)
{}
};
/** The contig IDs that have been removed from paths. */
static vector<ContigID> s_trimmedContigs;
/** The contig graph. */
typedef DirectedGraph<ContigProperties, Distance> DG;
typedef ContigGraph<DG> Graph;
typedef vector<ContigPath> Paths;
/** Return whether this vertex is a path or a contig. */
static bool
isPath(const ContigNode& u)
{
return u.id() >= Vertex::s_offset;
}
/** Return a path, complemented if necessary. */
static ContigPath
getPath(const Paths& paths, const ContigNode& u)
{
if (isPath(u)) {
unsigned i = u.id() - Vertex::s_offset;
return u.sense() ? reverseComplement(paths[i]) : paths[i];
} else
return ContigPath(1, u);
}
/** Read contig paths from the specified file.
* @param g the contig adjacency graph
* @param inPath the file of contig paths
* @param[out] pathIDs the path IDs
* @return the paths
*/
static Paths
readPaths(Graph& g, const string& inPath, vector<string>& pathIDs)
{
typedef graph_traits<Graph>::vertex_descriptor V;
assert(pathIDs.empty());
ifstream fin(inPath.c_str());
if (opt::verbose > 0)
cerr << "Reading `" << inPath << "'..." << endl;
if (inPath != "-")
assert_good(fin, inPath);
istream& in = inPath == "-" ? cin : fin;
assert_good(in, inPath);
Paths paths;
string id;
ContigPath path;
while (in >> id >> path) {
if (path.empty()) {
// Remove this contig from the graph.
V u = find_vertex(id, false, g);
clear_vertex(u, g);
remove_vertex(u, g);
} else {
pathIDs.push_back(id);
paths.push_back(path);
}
}
assert(in.eof());
return paths;
}
typedef multimap<ContigNode, Vertex> SeedMap;
/** Index the first and last contig of each path to facilitate finding
* overlaps between paths. */
static SeedMap
makeSeedMap(const Paths& paths)
{
SeedMap seedMap;
for (Paths::const_iterator it = paths.begin(); it != paths.end(); ++it) {
if (it->empty())
continue;
assert(!it->front().ambiguous());
seedMap.insert(make_pair(it->front(), Vertex(it - paths.begin(), false)));
assert(!it->back().ambiguous());
seedMap.insert(make_pair(it->back() ^ 1, Vertex(it - paths.begin(), true)));
}
return seedMap;
}
/** Check whether path starts with the sequence [first, last). */
static bool
startsWith(
ContigPath path,
bool rc,
ContigPath::const_iterator first,
ContigPath::const_iterator last)
{
if (rc)
reverseComplement(path.begin(), path.end());
assert(*first == path.front());
assert(first < last);
return unsigned(last - first) > path.size() ? false : equal(first, last, path.begin());
}
/** Check whether path starts with the sequence [first, last). */
static unsigned
findOverlap(
const Graph& g,
const Paths& paths,
ContigPath::const_iterator first,
ContigPath::const_iterator last,
const Vertex& v,
int& distance)
{
if (!startsWith(paths[v.id], v.sense, first, last))
return 0;
distance = -addProp(g, first, last).length;
return last - first;
}
typedef vector<Overlap> Overlaps;
/** Find every path that overlaps with the specified path. */
static void
findOverlaps(
const Graph& g,
const Paths& paths,
const SeedMap& seedMap,
const Vertex& v,
Overlaps& overlaps)
{
ContigPath rc;
if (v.sense) {
rc = paths[v.id];
reverseComplement(rc.begin(), rc.end());
}
const ContigPath& path = v.sense ? rc : paths[v.id];
for (ContigPath::const_iterator it = path.begin(); it != path.end(); ++it) {
if (it->ambiguous())
continue;
pair<SeedMap::const_iterator, SeedMap::const_iterator> range = seedMap.equal_range(*it);
for (SeedMap::const_iterator seed = range.first; seed != range.second; ++seed) {
if (v == seed->second)
continue;
int distance = 0;
unsigned overlap = findOverlap(g, paths, it, path.end(), seed->second, distance);
if (overlap > 0)
overlaps.push_back(Overlap(v, seed->second, overlap, distance));
}
}
}
/** Find every pair of overlapping paths. */
static Overlaps
findOverlaps(const Graph& g, const Paths& paths)
{
SeedMap seedMap = makeSeedMap(paths);
Overlaps overlaps;
for (Paths::const_iterator it = paths.begin(); it != paths.end(); ++it) {
unsigned i = it - paths.begin();
findOverlaps(g, paths, seedMap, Vertex(i, false), overlaps);
findOverlaps(g, paths, seedMap, Vertex(i, true), overlaps);
}
return overlaps;
}
/** Record the trimmed contigs. */
static void
recordTrimmedContigs(ContigPath::const_iterator first, ContigPath::const_iterator last)
{
for (ContigPath::const_iterator it = first; it != last; ++it)
if (!it->ambiguous())
s_trimmedContigs.push_back(it->contigIndex());
}
/** Remove ambiguous contigs from the ends of the path. */
static void
removeAmbiguousContigs(ContigPath& path)
{
if (!path.empty() && path.back().ambiguous())
path.erase(path.end() - 1);
if (!path.empty() && path.front().ambiguous())
path.erase(path.begin());
}
/** Remove the overlapping portion of the specified contig. */
static void
removeContigs(ContigPath& path, unsigned first, unsigned last)
{
assert(first <= path.size());
assert(last <= path.size());
if (first < last) {
recordTrimmedContigs(path.begin(), path.begin() + first);
recordTrimmedContigs(path.begin() + last, path.end());
path.erase(path.begin() + last, path.end());
path.erase(path.begin(), path.begin() + first);
} else {
recordTrimmedContigs(path.begin(), path.end());
path.clear();
}
removeAmbiguousContigs(path);
}
/** Find the largest overlap for each contig and remove it. */
static void
trimOverlaps(Paths& paths, const Overlaps& overlaps)
{
vector<unsigned> removed[2];
removed[0].resize(paths.size());
removed[1].resize(paths.size());
for (Overlaps::const_iterator it = overlaps.begin(); it != overlaps.end(); ++it) {
unsigned& a = removed[!it->source.sense][it->source.id];
unsigned& b = removed[it->target.sense][it->target.id];
a = max(a, it->overlap);
b = max(b, it->overlap);
}
for (Paths::iterator it = paths.begin(); it != paths.end(); ++it)
removeContigs(
*it, removed[0][it - paths.begin()], it->size() - removed[1][it - paths.begin()]);
}
/** Trim the ends of paths that overlap another path. */
static void
trimOverlaps(const Graph& g, Paths& paths)
{
for (Overlaps overlaps = findOverlaps(g, paths); !overlaps.empty();
overlaps = findOverlaps(g, paths)) {
cerr << "Found " << overlaps.size() / 2 << " overlaps.\n";
trimOverlaps(paths, overlaps);
}
}
static inline ContigProperties
get(vertex_bundle_t, const Graph& g, ContigNode u)
{
return u.ambiguous() ? ContigProperties(u.length() + opt::k - 1, 0) : g[u];
}
/** Add the path overlap edges to the specified graph. */
static void
addPathOverlapEdges(
Graph& g,
const Paths& paths,
const vector<string>& pathIDs,
const Overlaps& overlaps)
{
typedef graph_traits<Graph>::vertex_descriptor V;
const bool allowParallelEdge = opt::mode == opt::ASSEMBLE;
// Add the path vertices.
g_contigNames.unlock();
for (Paths::const_iterator it = paths.begin(); it != paths.end(); ++it) {
const ContigPath& path = *it;
const string& id = pathIDs[it - paths.begin()];
if (!path.empty()) {
V u = merge(g, path.begin(), path.end());
put(vertex_name, g, u, id);
}
}
g_contigNames.lock();
// Remove the single-end contigs that are in paths.
for (Paths::const_iterator it = paths.begin(); it != paths.end(); ++it)
remove_vertex_if(
g, it->begin(), it->end(), [](const ContigNode& c) { return !c.ambiguous(); });
// Add the path edges.
for (Overlaps::const_iterator it = overlaps.begin(); it != overlaps.end(); ++it) {
V u = it->source.descriptor();
V v = it->target.descriptor();
if (allowParallelEdge || !edge(u, v, g).second)
add_edge(u, v, it->distance, static_cast<DG&>(g));
else if (opt::verbose > 0)
cerr << "ambiguous overlap: " << get(vertex_name, g, u) << " -> "
<< get(vertex_name, g, v) << '\n';
}
}
typedef graph_traits<Graph>::edge_descriptor edge_descriptor;
/** A property map giving the number of contigs by which two paths
* overlap. */
typedef map<edge_descriptor, unsigned> OverlapMap;
/** Return the number of contigs by which the two paths overlap. */
static unsigned
getOverlap(
const OverlapMap& pmap,
graph_traits<Graph>::vertex_descriptor u,
graph_traits<Graph>::vertex_descriptor v)
{
if (isPath(u) && isPath(v)) {
// Both vertices are paths.
OverlapMap::const_iterator it = pmap.find(edge_descriptor(u, v));
return it == pmap.end() ? 0 : it->second;
} else {
// One of the two vertices is a contig.
return 0;
}
}
/** Merge a sequence of overlapping paths. */
static ContigPath
mergePaths(const Paths& paths, const OverlapMap& overlaps, const ContigPath& merge)
{
assert(!merge.empty());
ContigNode u = merge.front();
ContigPath path(getPath(paths, u));
for (ContigPath::const_iterator it = merge.begin() + 1; it != merge.end(); ++it) {
ContigNode v = *it;
ContigPath vpath(getPath(paths, v));
unsigned overlap = getOverlap(overlaps, u, v);
assert(path.size() > overlap);
assert(vpath.size() > overlap);
assert(equal(path.end() - overlap, path.end(), vpath.begin()));
path.insert(path.end(), vpath.begin() + overlap, vpath.end());
u = v;
}
return path;
}
/** Return true if the edge e is a path overlap. */
struct IsPathOverlap
{
IsPathOverlap(const Graph& g, const OverlapMap& pmap, const IsPositive<Graph>& pred)
: m_g(g)
, m_pmap(pmap)
, m_isPositive(pred)
{}
bool operator()(edge_descriptor e) const
{
bool stranded = true;
if (opt::ss)
stranded = m_isPositive(e);
return stranded && getOverlap(m_pmap, source(e, m_g), target(e, m_g));
}
typedef edge_descriptor argument_type;
typedef bool result_type;
private:
const Graph& m_g;
const OverlapMap& m_pmap;
const IsPositive<Graph>& m_isPositive;
};
/** Assemble overlapping paths. */
static void
assembleOverlappingPaths(Graph& g, Paths& paths, vector<string>& pathIDs)
{
if (paths.empty())
return;
// Find overlapping paths.
Overlaps overlaps = findOverlaps(g, paths);
addPathOverlapEdges(g, paths, pathIDs, overlaps);
// Create a property map of path overlaps.
OverlapMap overlapMap;
for (Overlaps::const_iterator it = overlaps.begin(); it != overlaps.end(); ++it)
overlapMap.insert(OverlapMap::value_type(
OverlapMap::key_type(it->source.descriptor(), it->target.descriptor()), it->overlap));
// Assemble unambiguously overlapping paths.
Paths merges;
assemble_if(g, back_inserter(merges), IsPathOverlap(g, overlapMap, IsPositive<Graph>(g)));
// Merge overlapping paths.
g_contigNames.unlock();
assert(!pathIDs.empty());
setNextContigName(pathIDs.back());
for (Paths::const_iterator it = merges.begin(); it != merges.end(); ++it) {
string name = createContigName();
if (opt::verbose > 0)
cerr << name << '\t' << *it << '\n';
Vertex u(paths.size(), false);
put(vertex_name, g, u.descriptor(), name);
pathIDs.push_back(name);
paths.push_back(mergePaths(paths, overlapMap, *it));
// Remove the merged paths.
for (ContigPath::const_iterator it2 = it->begin(); it2 != it->end(); ++it2) {
if (isPath(*it2))
paths[it2->id() - Vertex::s_offset].clear();
}
}
g_contigNames.lock();
}
int
main(int argc, char** argv)
{
string commandLine;
{
ostringstream ss;
char** last = argv + argc - 1;
copy(argv, last, ostream_iterator<const char*>(ss, " "));
ss << *last;
commandLine = ss.str();
}
if (!opt::db.empty())
opt::metaVars.resize(3);
bool die = false;
for (int c; (c = getopt_long(argc, argv, shortopts, longopts, NULL)) != -1;) {
istringstream arg(optarg != NULL ? optarg : "");
switch (c) {
case '?':
die = true;
break;
case 'g':
arg >> opt::graphPath;
break;
case 'k':
arg >> opt::k;
break;
case 'r':
arg >> opt::repeatContigs;
break;
case 'v':
opt::verbose++;
break;
case OPT_HELP:
cout << USAGE_MESSAGE;
exit(EXIT_SUCCESS);
case OPT_VERSION:
cout << VERSION_MESSAGE;
exit(EXIT_SUCCESS);
case OPT_DB:
arg >> opt::db;
break;
case OPT_LIBRARY:
arg >> opt::metaVars[0];
break;
case OPT_STRAIN:
arg >> opt::metaVars[1];
break;
case OPT_SPECIES:
arg >> opt::metaVars[2];
break;
}
if (optarg != NULL && !arg.eof()) {
cerr << PROGRAM ": invalid option: `-" << (char)c << optarg << "'\n";
exit(EXIT_FAILURE);
}
}
if (opt::k <= 0) {
cerr << PROGRAM ": missing -k,--kmer option\n";
die = true;
}
if (argc - optind < 2) {
cerr << PROGRAM ": missing arguments\n";
die = true;
} else if (argc - optind > 2) {
cerr << PROGRAM ": too many arguments\n";
die = true;
}
if (die) {
cerr << "Try `" << PROGRAM << " --help' for more information.\n";
exit(EXIT_FAILURE);
}
const char* adjPath = argv[optind++];
if (opt::verbose > 0)
cerr << "Reading `" << adjPath << "'..." << endl;
ifstream fin(adjPath);
assert_good(fin, adjPath);
Graph g;
fin >> g;
Vertex::s_offset = g.num_vertices() / 2;
string pathsFile(argv[optind++]);
vector<string> pathIDs;
Paths paths = readPaths(g, pathsFile, pathIDs);
switch (opt::mode) {
case opt::OVERLAP:
// Find overlapping paths, do not assemble.
addPathOverlapEdges(g, paths, pathIDs, findOverlaps(g, paths));
paths.clear();
if (opt::graphPath.empty())
opt::graphPath = "-";
break;
case opt::ASSEMBLE:
// Assemble overlapping paths.
assembleOverlappingPaths(g, paths, pathIDs);
break;
case opt::TRIM:
// Trim overlapping paths.
trimOverlaps(g, paths);
// Remove paths consisting of a single contig.
for_each_if(
paths.begin(),
paths.end(),
[](ContigPath& c) { return c.clear(); },
[](const ContigPath& c) { return c.size() == 1; });
// Add the paths to the graph.
addPathOverlapEdges(g, paths, pathIDs, Overlaps());
break;
}
// Output the paths.
for (Paths::const_iterator it = paths.begin(); it != paths.end(); ++it) {
if (it->empty())
continue;
assert(it->size() != 1);
cout << pathIDs[it - paths.begin()] << '\t' << *it << '\n';
}
assert(cout.good());
// Output the graph.
if (!opt::graphPath.empty()) {
ofstream fout;
ostream& out = opt::graphPath == "-" ? cout : (fout.open(opt::graphPath.c_str()), fout);
assert_good(out, opt::graphPath);
write_graph(out, g, PROGRAM, commandLine);
assert_good(out, opt::graphPath);
}
// Output the repeat contigs.
if (!opt::repeatContigs.empty()) {
sort(s_trimmedContigs.begin(), s_trimmedContigs.end());
s_trimmedContigs.erase(
unique(s_trimmedContigs.begin(), s_trimmedContigs.end()), s_trimmedContigs.end());
ofstream out(opt::repeatContigs.c_str());
assert_good(out, opt::repeatContigs);
for (vector<ContigID>::const_iterator it = s_trimmedContigs.begin();
it != s_trimmedContigs.end();
++it)
out << get(g_contigNames, *it) << '\n';
assert_good(out, opt::repeatContigs);
}
if (!opt::db.empty()) {
init(db, opt::db, opt::verbose, PROGRAM, opt::getCommand(argc, argv), opt::metaVars);
addToDb(db, "SS", opt::ss);
addToDb(db, "K", opt::k);
}
return 0;
}
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