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model.cpp
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model.cpp
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#include "model.h"
#include <iostream>
#include <sstream>
#include <cstdlib>
Model::Model(Sequence* seqTarget, vector<Sequence>* seqFamily, int targetIndex, Params* params)
{
_seqTarget = seqTarget;
_seqFamily = seqFamily;
_params = params;
_numrows = _seqTarget->length();
_numcols = _seqFamily->at(0).length();
_familysize = _seqFamily->size();
_targetIndex = targetIndex;
_time = _params->getNumSubst();
_bkgProbs[0] = _params->getProbA(); // A
_bkgProbs[1] = _params->getProbC(); // C
_bkgProbs[2] = _params->getProbG(); // G
_bkgProbs[3] = _params->getProbT(); // T
for (int i = 0; i < 4; i++) _logBkgProbs[i] = log(_bkgProbs[i]);
// F81 substituion model
for (int i = 0; i < 4; i++) {
for (int j = 0; j < 4; j++) {
if (i == j) _substMatrix[i][j] = exp(-_time) + (1.0 - exp(-_time)) * _bkgProbs[j];
else _substMatrix[i][j] = (1.0 - exp(-_time)) * _bkgProbs[j];
} // end of for j
} // end of for i
for (int i = 0; i < 4; i++)
for (int j = 0; j < 4; j++)
_logSubstMatrix[i][j] = log(_substMatrix[i][j]);
double val = _params->getMtoI();
_tpMI = (val != -1) ? val : DEFAULT_DELTA;
val = _params->getMtoD();
_tpMD = (val != -1) ? val : DEFAULT_DELTA;
val = _params->getItoI();
_tpII = (val != -1) ? val : DEFAULT_EPSILON;
val = _params->getDtoD();
_tpDD = (val != -1) ? val : DEFAULT_EPSILON;
val = _params->getItoD();
_tpID = (val != -1) ? val : DEFAULT_RHO;
val = _params->getDtoI();
_tpDI = (val != -1) ? val : DEFAULT_RHO;
_tpSI = _tpSD = DEFAULT_DELTA;
_tp2E = DEFAULT_TAU;
updateTransMatrix();
// Allocate memory for DP tables
for (int i = 0; i < NUM_STATES; i++) {
_forwardTable[i] = new double* [_numrows+1];
_backwardTable[i] = new double* [_numrows+1];
for (int j = 0; j < _numrows+1; j++) {
_forwardTable[i][j] = new double [_numcols+1];
_backwardTable[i][j] = new double [_numcols+1];
} // end of for j
} // end of for i
}
Model::~Model()
{
for (int i = 0; i < NUM_STATES; i++) {
for (int j = 0; j < _numrows+1; j++) {
delete [] _forwardTable[i][j];
delete [] _backwardTable[i][j];
} // end of for j
delete [] _forwardTable[i];
delete [] _backwardTable[i];
} // end of for i
}
void Model::dump()
{
cerr << "t=" << _time << endl;
cerr << "bkg prob=(" << _bkgProbs[0] << "," << _bkgProbs[1] << "," << _bkgProbs[2] << "," << _bkgProbs[3] << ")" << endl;
cerr << "subst prob:" << endl;
for (int i = 0; i < 4; i++) {
for (int j = 0; j < 4; j++) cerr << _substMatrix[i][j] << "\t";
cerr << endl;
}
cerr << "trans prob:" << endl;
for (int i = 0; i < 5; i++) {
for (int j = 0; j < 5; j++) cerr << exp(_logTransMatrix[i][j]) << "\t";
cerr << endl;
}
cerr << "forward table:M" << endl;
for (int i = 0; i <= _numrows; i++) {
for (int j = 0; j <= _numcols; j++) {
cerr << exp(_forwardTable[M][i][j]) << "\t";
}
cerr << endl;
}
cerr << "forward table:I" << endl;
for (int i = 0; i <= _numrows; i++) {
for (int j = 0; j <= _numcols; j++) {
cerr << exp(_forwardTable[I][i][j]) << "\t";
}
cerr << endl;
}
cerr << "forward table:D" << endl;
for (int i = 0; i <= _numrows; i++) {
for (int j = 0; j <= _numcols; j++) {
cerr << exp(_forwardTable[D][i][j]) << "\t";
}
cerr << endl;
}
}
void Model::updateTransMatrix()
{
double logtpMI = log(_tpMI);
double logtpMD = log(_tpMD);
double logtpII = log(_tpII);
double logtpDD = log(_tpDD);
double logtpID = log(_tpID);
double logtpDI = log(_tpDI);
double logtpSI = log(_tpSI);
double logtpSD = log(_tpSD);
double logtp2E = log(_tp2E);
for (int i = 0; i < 5; i++)
for (int j = 0; j < 5; j++)
_logTransMatrix[i][j] = NEGINF;
_logTransMatrix[M][M] = log(1.0-_tpMI-_tpMD-_tp2E);
_logTransMatrix[M][I] = logtpMI;
_logTransMatrix[M][D] = logtpMD;
_logTransMatrix[M][E] = logtp2E;
_logTransMatrix[I][M] = log(1.0-_tpII-_tpID-_tp2E);
_logTransMatrix[I][I] = logtpII;
_logTransMatrix[I][D] = logtpID;
_logTransMatrix[I][E] = logtp2E;
_logTransMatrix[D][M] = log(1.0-_tpDD-_tpDI-_tp2E);
_logTransMatrix[D][D] = logtpDD;
_logTransMatrix[D][I] = logtpDI;
_logTransMatrix[D][E] = logtp2E;
_logTransMatrix[S][M] = _logTransMatrix[M][M];
_logTransMatrix[S][I] = _logTransMatrix[M][I];
_logTransMatrix[S][D] = _logTransMatrix[M][D];
_logTransMatrix[S][E] = logtp2E;
}
void Model::estimateParameters(bool est_param, vector<double>& vecParams)
{
if (est_param == false) {
forward();
backward();
return;
}
double prevscore = NEGINF;
for (int it = 0; it < NUM_ITER; it++) {
forward();
backward();
// expected transition counts
double logTrCnts[NUM_STATES][NUM_STATES];
for (int i = 0; i < NUM_STATES; i++)
for (int j = 0; j < NUM_STATES; j++) logTrCnts[i][j] = NEGINF;
for (int i = 0; i < _numrows; i++) {
for (int j = 0; j < _numcols; j++) {
//MtoM
double logval = _forwardTable[M][i][j] + _logTransMatrix[M][M] + getLogJointProb(i+1, j+1) + _backwardTable[M][i+1][j+1];
logTrCnts[M][M] = logsum(logTrCnts[M][M], logval);
//MtoI
logval = _forwardTable[M][i][j] + _logTransMatrix[M][I] + _logBkgProbs[_seqTarget->at(i)] + _backwardTable[I][i+1][j];
logTrCnts[M][I] = logsum(logTrCnts[M][I], logval);
//MtoD
logval = _forwardTable[M][i][j] + _logTransMatrix[M][D] + getLogJointProbSingle(j+1) + _backwardTable[D][i][j+1];
logTrCnts[M][D] = logsum(logTrCnts[M][D], logval);
//ItoM
logval = _forwardTable[I][i][j] + _logTransMatrix[I][M] + getLogJointProb(i+1, j+1) + _backwardTable[M][i+1][j+1];
logTrCnts[I][M] = logsum(logTrCnts[I][M], logval);
//ItoI
logval = _forwardTable[I][i][j] + _logTransMatrix[I][I] + _logBkgProbs[_seqTarget->at(i)] + _backwardTable[I][i+1][j];
logTrCnts[I][I] = logsum(logTrCnts[I][I], logval);
//ItoD
logval = _forwardTable[I][i][j] + _logTransMatrix[I][D] + getLogJointProbSingle(j+1) + _backwardTable[D][i][j+1];
logTrCnts[I][D] = logsum(logTrCnts[I][D], logval);
//DtoM
logval = _forwardTable[D][i][j] + _logTransMatrix[D][M] + getLogJointProb(i+1, j+1) + _backwardTable[M][i+1][j+1];
logTrCnts[D][M] = logsum(logTrCnts[D][M], logval);
//DtoD
logval = _forwardTable[D][i][j] + _logTransMatrix[D][D] + getLogJointProbSingle(j+1) + _backwardTable[D][i][j+1];
logTrCnts[D][D] = logsum(logTrCnts[D][D], logval);
//DtoI
logval = _forwardTable[D][i][j] + _logTransMatrix[D][I] + _logBkgProbs[_seqTarget->at(i)] + _backwardTable[I][i+1][j];
logTrCnts[D][I] = logsum(logTrCnts[D][I], logval);
} // end of for j
} // end of for i
for (int i = 0; i < NUM_STATES; i++)
for (int j = 0; j < NUM_STATES; j++) {
logTrCnts[i][j] = exp(logTrCnts[i][j] - _logfullprob);
}
_tpMI = logTrCnts[M][I] / (logTrCnts[M][M] + logTrCnts[M][I] + logTrCnts[M][D]);
if (_tpMI < DOUBLE_MIN) _tpMI = DOUBLE_MIN;
_tpMD = logTrCnts[M][D] / (logTrCnts[M][M] + logTrCnts[M][I] + logTrCnts[M][D]);
if (_tpMD < DOUBLE_MIN) _tpMD = DOUBLE_MIN;
_tpII = logTrCnts[I][I] / (logTrCnts[I][I] + logTrCnts[I][M] + logTrCnts[I][D]);
if (_tpII < DOUBLE_MIN) _tpII = DOUBLE_MIN;
_tpDD = logTrCnts[D][D] / (logTrCnts[D][D] + logTrCnts[D][M] + logTrCnts[D][I]);
if (_tpDD < DOUBLE_MIN) _tpDD = DOUBLE_MIN;
_tpID = logTrCnts[I][D] / (logTrCnts[I][I] + logTrCnts[I][M] + logTrCnts[I][D]);
if (_tpID < DOUBLE_MIN) _tpID = DOUBLE_MIN;
_tpDI = logTrCnts[D][I] / (logTrCnts[D][D] + logTrCnts[D][M] + logTrCnts[D][I]);
if (_tpDI < DOUBLE_MIN) _tpDI = DOUBLE_MIN;
updateTransMatrix();
} // end of for
vecParams.at(0) += _tpMI;
vecParams.at(1) += _tpMD;
vecParams.at(2) += _tpII;
vecParams.at(3) += _tpDD;
vecParams.at(4) += _tpID;
vecParams.at(5) += _tpDI;
}
void Model::forward()
{
// initialize a table
_forwardTable[M][0][0] = log(1);
_forwardTable[I][0][0] = _forwardTable[D][0][0] = NEGINF;
for (int r = 0; r <= _numrows; r++) {
for (int c = 0; c <= _numcols; c++) {
if (r == 0 && c == 0) continue;
double Mr_1c_1 = (r-1 > -1 && c-1 > -1) ? _forwardTable[M][r-1][c-1] : NEGINF;
double Mr_1c = (r-1 > -1) ? _forwardTable[M][r-1][c] : NEGINF;
double Mrc_1 = (c-1 > -1) ? _forwardTable[M][r][c-1] : NEGINF;
double Ir_1c_1 = (r-1 > -1 && c-1 > -1) ? _forwardTable[I][r-1][c-1] : NEGINF;
double Ir_1c = (r-1 > -1) ? _forwardTable[I][r-1][c] : NEGINF;
double Irc_1 = (c-1 > -1) ? _forwardTable[I][r][c-1] : NEGINF;
double Dr_1c_1 = (r-1 > -1 && c-1 > -1) ? _forwardTable[D][r-1][c-1] : NEGINF;
double Dr_1c = (r-1 > -1) ? _forwardTable[D][r-1][c] : NEGINF;
double Drc_1 = (c-1 > -1) ? _forwardTable[D][r][c-1] : NEGINF;
double logdouble = getLogJointProb(r, c);
double logval = logsum(_logTransMatrix[M][M]+Mr_1c_1, _logTransMatrix[I][M]+Ir_1c_1);
logval = logsum(logval, _logTransMatrix[D][M]+Dr_1c_1);
_forwardTable[M][r][c] = logdouble + logval;
double logsingle = (r-1 > -1) ? _logBkgProbs[_seqTarget->at(r-1)] : NEGINF;
logval = logsum(_logTransMatrix[M][I]+Mr_1c, _logTransMatrix[I][I]+Ir_1c);
logval = logsum(logval, _logTransMatrix[D][I]+Dr_1c);
_forwardTable[I][r][c] = logsingle + logval;
logsingle = getLogJointProbSingle(c);
logval = logsum(_logTransMatrix[M][D]+Mrc_1, _logTransMatrix[D][D]+Drc_1);
logval = logsum(logval, _logTransMatrix[I][D]+Irc_1);
_forwardTable[D][r][c] = logsingle + logval;
} // end of for c
} // end of for r
// compute full probability
double logfullprob = _logTransMatrix[M][E]+_forwardTable[M][_numrows][_numcols];
logfullprob = logsum(logfullprob, _logTransMatrix[M][E]+_forwardTable[I][_numrows][_numcols]);
logfullprob = logsum(logfullprob, _logTransMatrix[M][E]+_forwardTable[D][_numrows][_numcols]);
_logfullprob = logfullprob;
}
void Model::backward()
{
// initialize a table
_backwardTable[M][_numrows][_numcols] = log(_tp2E);
_backwardTable[I][_numrows][_numcols] = log(_tp2E);
_backwardTable[D][_numrows][_numcols] = log(_tp2E);
for (int r = _numrows; r >= 0; r--) {
for (int c = _numcols; c >= 0; c--) {
if (r == _numrows && c == _numcols) continue;
double Mr__1c__1 = (r+1 <= _numrows && c+1 <= _numcols) ? _backwardTable[M][r+1][c+1] : NEGINF;
double Mr__1c = (r+1 <= _numrows) ? _backwardTable[M][r+1][c] : NEGINF;
double Mrc__1 = (c+1 <= _numcols) ? _backwardTable[M][r][c+1] : NEGINF;
double Ir__1c__1 = (r+1 <= _numrows && c+1 <= _numcols) ? _backwardTable[I][r+1][c+1] : NEGINF;
double Ir__1c = (r+1 <= _numrows) ? _backwardTable[I][r+1][c] : NEGINF;
double Irc__1 = (c+1 <= _numcols) ? _backwardTable[I][r][c+1] : NEGINF;
double Dr__1c__1 = (r+1 <= _numrows && c+1 <= _numcols) ? _backwardTable[D][r+1][c+1] : NEGINF;
double Dr__1c = (r+1 <= _numrows) ? _backwardTable[D][r+1][c] : NEGINF;
double Drc__1 = (c+1 <= _numcols) ? _backwardTable[D][r][c+1] : NEGINF;
double logdouble = getLogJointProb(r+1, c+1);
double logsingle_r = (r+1 <= _numrows) ? _logBkgProbs[_seqTarget->at(r)] : NEGINF;
double logsingle_c = getLogJointProbSingle(c+1);
double logval = logsum(logdouble+_logTransMatrix[M][M]+Mr__1c__1, logsingle_r+_logTransMatrix[M][I]+Ir__1c);
logval = logsum(logval, logsingle_c+_logTransMatrix[M][D]+Drc__1);
_backwardTable[M][r][c] = logval;
logval = logsum(logdouble+_logTransMatrix[I][M]+Mr__1c__1, logsingle_r+_logTransMatrix[I][I]+Ir__1c);
logval = logsum(logval, logsingle_c+_logTransMatrix[I][D]+Irc__1);
_backwardTable[I][r][c] = logval;
logval = logsum(logdouble+_logTransMatrix[D][M]+Mr__1c__1, logsingle_c+_logTransMatrix[D][D]+Drc__1);
logval = logsum(logval, logsingle_r+_logTransMatrix[D][I]+Ir__1c);
_backwardTable[D][r][c] = logval;
} // end of for c
} // end of for r
}
double Model::logsum(double logx, double logy)
{
if (logx <= NEGINF && logy <= NEGINF) return NEGINF;
if (logx <= NEGINF) return logy;
if (logy <= NEGINF) return logx;
if (logy >= logx) return (logy + log(1.0 + exp(logx - logy)));
return (logx + log(1.0 + exp(logy - logx)));
}
double Model::getLogJointProb(int r, int c)
{
// Joint probability of an alignment column based on a star topology
r--;
c--;
if (r < 0 || c < 0 || r >= _numrows || c >= _numcols) return NEGINF;
// collect non-gap characters
vector<int> bases;
bases.push_back(_seqTarget->at(r));
for (int i = 0; i < _familysize; i++) {
int ch = _seqFamily->at(i).at(c);
if (ch == GAP) continue;
bases.push_back(ch);
} // end of for i
// search cache
stringstream ss;
for (int i = 0; i < bases.size(); i++) ss << bases.at(i);
string key = ss.str();
map<string, double>::iterator iter = _emissionProbs.find(key);
if (iter != _emissionProbs.end()) {
return iter->second;
} // end of if
double logprob = NEGINF;
for (int ai = 0; ai < 4; ai++) { // for all possible ancestral characters
double val = _logBkgProbs[ai];
for (int di = 0; di < bases.size(); di++) {
int descChar = bases.at(di);
if (descChar == N) continue;
double descBkgProb = _logBkgProbs[descChar];
val += _logSubstMatrix[ai][descChar];
} // end of for di
logprob = logsum(logprob, val);
} // end of for ai
_emissionProbs[key] = logprob;
return logprob;
}
double Model::getLogJointProbSingle(int c)
{
// Joint probability of an alignment column based on a star topology
c--;
if (c < 0 || c >= _numcols) return NEGINF;
// collect non-gap characters
vector<int> bases;
for (int i = 0; i < _familysize; i++) {
int ch = _seqFamily->at(i).at(c);
if (ch == GAP) continue;
bases.push_back(ch);
} // end of for i
// search cache
stringstream ss;
for (int i = 0; i < bases.size(); i++) ss << bases.at(i);
string key = ss.str();
map<string, double>::iterator iter = _emissionProbs.find(key);
if (iter != _emissionProbs.end()) {
return iter->second;
} // end of if
double logprob = NEGINF;
for (int ai = 0; ai < 4; ai++) { // for all possible ancestral characters
double val = _logBkgProbs[ai];
for (int di = 0; di < bases.size(); di++) {
int descChar = bases.at(di);
if (descChar == N) continue;
double descBkgProb = _logBkgProbs[descChar];
val += _logSubstMatrix[ai][descChar];
} // end of for di
logprob = logsum(logprob, val);
} // end of for ai
_emissionProbs[key] = logprob;
return logprob;
}
double Model::sample_alignments(string& straln, vector<Sequence>& sequences, double threshold)
{
string aln_target;
vector<string> aln_family(_familysize);
double lls = 0; // log likelihood score
int i = _numrows;
int j = _numcols;
double MtoE = _logTransMatrix[M][E] + _forwardTable[M][i][j];
double ItoE = _logTransMatrix[M][E] + _forwardTable[I][i][j];
double DtoE = _logTransMatrix[M][E] + _forwardTable[D][i][j];
int tableindex = getNextTableIndex(MtoE, ItoE, DtoE, threshold);
lls += _logTransMatrix[tableindex][E];
while (i > 0 && j > 0) {
if (tableindex == 0) {
aln_target = ALPHS[_seqTarget->at(i-1)] + aln_target;
for (int fi=0; fi < _familysize; fi++)
aln_family[fi] = ALPHS[_seqFamily->at(fi).at(j-1)] + aln_family[fi];
double logdouble = getLogJointProb(i, j);
double mlogp = logdouble + _logTransMatrix[M][M] + _forwardTable[M][i-1][j-1];
double ilogp = logdouble + _logTransMatrix[I][M] + _forwardTable[I][i-1][j-1];
double dlogp = logdouble + _logTransMatrix[D][M] + _forwardTable[D][i-1][j-1];
tableindex = getNextTableIndex(mlogp, ilogp, dlogp, threshold);
i--;
j--;
lls += logdouble;
if (i > 0 && j > 0) lls += _logTransMatrix[tableindex][M];
else lls += _logTransMatrix[S][M];
} else if (tableindex == 1) {
aln_target = ALPHS[_seqTarget->at(i-1)] + aln_target;
for (int fi=0; fi < _familysize; fi++)
aln_family[fi] = "-" + aln_family[fi];
double logsingle = _logBkgProbs[_seqTarget->at(i-1)];
double mlogp = logsingle + _logTransMatrix[M][I] + _forwardTable[M][i-1][j];
double ilogp = logsingle + _logTransMatrix[I][I] + _forwardTable[I][i-1][j];
double dlogp = logsingle + _logTransMatrix[D][I] + _forwardTable[D][i-1][j];
tableindex = getNextTableIndex(mlogp, ilogp, dlogp, threshold);
i--;
lls += logsingle;
if (i > 0) lls += _logTransMatrix[tableindex][I];
else lls += _logTransMatrix[S][I];
} else {
aln_target = "-" + aln_target;
for (int fi=0; fi < _familysize; fi++)
aln_family[fi] = ALPHS[_seqFamily->at(fi).at(j-1)] + aln_family[fi];
double logsingle = getLogJointProbSingle(j);
double mlogp = logsingle + _logTransMatrix[M][D] + _forwardTable[M][i][j-1];
double ilogp = logsingle + _logTransMatrix[I][D] + _forwardTable[I][i][j-1];
double dlogp = logsingle + _logTransMatrix[D][D] + _forwardTable[D][i][j-1];
tableindex = getNextTableIndex(mlogp, ilogp, dlogp, threshold);
j--;
lls += logsingle;
if (j > 0) lls += _logTransMatrix[tableindex][D];
else lls += _logTransMatrix[S][D];
}
} // end of while i & j
while (i > 0) {
double logsingle = _logBkgProbs[_seqTarget->at(i-1)];
aln_target = ALPHS[_seqTarget->at(i-1)] + aln_target;
for (int fi=0; fi < _familysize; fi++)
aln_family[fi] = "-" + aln_family[fi];
i--;
lls += logsingle;
if (i > 0) lls += _logTransMatrix[I][I];
else lls += _logTransMatrix[S][I];
} // end of while i
while (j > 0) {
double logsingle = getLogJointProbSingle(j);
aln_target = "-" + aln_target;
for (int fi=0; fi < _familysize; fi++)
aln_family[fi] = ALPHS[_seqFamily->at(fi).at(j-1)] + aln_family[fi];
j--;
lls += logsingle;
if (i > 0) lls += _logTransMatrix[D][D];
else lls += _logTransMatrix[S][D];
} // end of while j
// make alignment string
straln = "";
int fi = 0;
for (int si = 0; si < _familysize+1; si++) {
string seqname = sequences.at(si).name();
straln += ">" + seqname + "\n";
if (_targetIndex == si) {
straln += aln_target + "\n";
} else {
straln += aln_family.at(fi) + "\n";
fi++;
}
} // end of for si
return (lls - _logfullprob);
}
double Model::getNextTableIndex(double mlogp, double ilogp, double dlogp, double threshold)
{
int tableindex = 0;
double sum = logsum(mlogp, ilogp);
sum = logsum(sum, dlogp);
double probM = exp(mlogp - sum);
double probI = exp(ilogp - sum);
double probX = exp(dlogp - sum);
// fine max
int maxindex = 0;
double probMax = probM;
if (probI > probM && probI > probX) {
probMax = probI;
maxindex = 1;
} else if (probX > probM && probX > probI) {
probMax = probX;
maxindex = 2;
}
if (probMax >= threshold) {
return maxindex;
}
double rndnumber = rand()/(double)RAND_MAX;
if (rndnumber < probM) tableindex = 0;
else if (rndnumber >= probM && rndnumber < (probM+probI)) tableindex = 1;
else tableindex = 2;
return tableindex;
}