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forback.c
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forback.c
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/************************************************************
* HMMER - Biological sequence analysis with HMMs
* Copyright 1992-1995 Sean R. Eddy
*
* This source code is distributed under the terms of the
* GNU General Public License. See the files COPYING and
* GNULICENSE for details.
*
************************************************************/
/* forback.c
* added to 1.7.7 Tue Jan 24 13:35:42 1995
* Previous forback.c, which has been obsolete for some time, moved to Archive/.
*
* Forward-Backward algorithm for calculating likelihood of an HMM
* given a sequence. Used for full Baum-Welch expectation maximization
* and for evaulating confidence of positions in Viterbi alignments.
*
* Derived from saviterbi.c. Related to the whole family of alignment
* algorithms. Uses sahmm's, like simulated annealing does.
*/
#include <stdio.h>
#include <stdlib.h>
#include <string.h>
#include <math.h>
#include "states.h"
#include "externs.h"
#include "squid.h"
#ifdef MEMDEBUG
#include "dbmalloc.h"
#endif
/* Function: Forward()
*
* Purpose: Forward calculation.
*
* The fwd matrix is 0..L+1 rows by 0..M+1 columns for a
* sequence of length 1..L and a model of length 1..M.
* Each entry represents the scaled summed probability of getting
* into this state, starting from (0,0), inclusive of
* symbol emission.
*
* Args: hmm - model, probability-form, single precision
* s - sequence 0..L-1, upper case
* ret_fwd - RETURN: forward calculation matrix
* ret_scl - RETURN: 0..L array of scaling factors
*
* Return: (void)
* fwd and scl are alloc'ed here, and must be free'd by caller:
* Free2DArray(fwd, L+2)
* free(scl)
*/
void
Forward(struct hmm_struc *hmm, char *s, struct forback_s ***ret_fwd, float **ret_scl)
{
char *seq; /* sequence, 1..L */
int L; /* length of seq */
struct forback_s **fwd; /* the calculation grid */
int i; /* counter for sequence position: 0,1..L,L+1 */
int k; /* counter for model position: 0,1..M,M+1 */
int i_symidx; /* hint for symbol index in alphabet */
char *alphptr; /* ptr into alphabet (optimization) */
float *scl; /* scaling factors */
struct forback_s *thisrow;
struct forback_s *nextrow;
/********************************************
* Initial setup and allocations
********************************************/
/* convert sequence to 1..L for ease of indexing */
L = strlen(s);
seq = (char *) MallocOrDie((L+2) * sizeof(char));
strcpy(seq+1, s);
seq[0] = ' ';
/* allocate the calculation matrix */
fwd = (struct forback_s **) MallocOrDie (sizeof(struct forback_s *) * (L+2));
for (i = 0; i <= L+1; i++)
fwd[i] = (struct forback_s *) malloc (sizeof(struct forback_s) * (hmm->M + 2));
/* allocate the scalefactors */
scl = (float *) MallocOrDie ((L+1) * sizeof(float));
/********************************************
* Initialization
********************************************/
/* set up the 0,0 cell */
fwd[0][0].score_m = 1.0;
fwd[0][0].score_d = 0.0;
fwd[0][0].score_i = 0.0;
/* initialize the top row */
for (k = 1; k <= hmm->M; k++)
{
fwd[0][k].score_m = 0.0;
fwd[0][k].score_i = 0.0;
}
/********************************************
* Recursion: fill in the fwd matrix
********************************************/
for (i = 0; i <= L; i++)
{
/* get ptrs into current and next row. */
thisrow = fwd[i];
nextrow = fwd[i+1];
/* initialize in the next row */
nextrow[0].score_m = 0.0;
nextrow[0].score_d = 0.0;
/* optimization: get an index for this symbol
while we're in the outer loop. */
if ((alphptr = strchr(Alphabet, seq[i])) != NULL)
i_symidx = alphptr - Alphabet;
else
i_symidx = -1; /* too bad; it's a degenerate symbol */
/* First pass across all states.
Emission scores, delete states */
for (k = 0; k <= hmm->M; k++)
{
/* add emission scores to the current cell. */
if (i_symidx >= 0)
{
thisrow[k].score_m *= hmm->mat[k].p[i_symidx];
thisrow[k].score_i *= hmm->ins[k].p[i_symidx];
}
else if (i > 0) /* watch out for the "gap" at position 0 */
{
thisrow[k].score_m *= ForbackSymscore(seq[i], hmm->mat[k].p, TRUE);
thisrow[k].score_i *= ForbackSymscore(seq[i], hmm->ins[k].p, TRUE);
}
/* deal with transitions to delete state */
thisrow[k+1].score_d = thisrow[k].score_d * hmm->del[k].t[DELETE] +
thisrow[k].score_i * hmm->ins[k].t[DELETE] +
thisrow[k].score_m * hmm->mat[k].t[DELETE];
}
/* In preparation for the operations that affect
the next row, scale the current row. */
scl[i] = 0.0;
for (k = 0; k <= hmm->M; k++)
{
if (thisrow[k].score_m > scl[i]) scl[i] = thisrow[k].score_m;
if (thisrow[k].score_d > scl[i]) scl[i] = thisrow[k].score_d;
if (thisrow[k].score_i > scl[i]) scl[i] = thisrow[k].score_i;
}
scl[i] = 1.0 / scl[i];
for (k = 0; k <= hmm->M; k++)
{
thisrow[k].score_m *= scl[i];
thisrow[k].score_d *= scl[i];
thisrow[k].score_i *= scl[i];
}
/* Now deal with insert and match transitions,
affecting the next row. */
for (k = 0; k <= hmm->M; k++)
{
/* deal with transitions to insert state */
nextrow[k].score_i = thisrow[k].score_d * hmm->del[k].t[INSERT] +
thisrow[k].score_i * hmm->ins[k].t[INSERT] +
thisrow[k].score_m * hmm->mat[k].t[INSERT];
/* deal with transitions to match state */
nextrow[k+1].score_m = thisrow[k].score_d * hmm->del[k].t[MATCH] +
thisrow[k].score_i * hmm->ins[k].t[MATCH] +
thisrow[k].score_m * hmm->mat[k].t[MATCH];
}
}
/******************************************
* Debugging information
******************************************/
#ifdef EXTREME_DEBUG
DumpForbackMatrix(fwd, L, hmm->M, scl);
#endif /* EXTREME_DEBUG */
/********************************************
* Garbage collection and return: caller is responsible for free'ing fwd and scl!
********************************************/
if (ret_scl != NULL) *ret_scl = scl;
if (ret_fwd != NULL) *ret_fwd = fwd;
free(seq);
}
/* Function: Backward()
*
* Purpose: Backward calculation.
*
* The bck matrix is 0..L+1 rows by 0..M+1 columns for a
* sequence of length 1..L and a model of length 1..M.
* Each entry represents the scaled summed probability of getting
* into this state, starting from (0,0), inclusive of
* symbol emission.
*
* The scalefactors used must be the same as the ones used for
* the forward calculation. This means Backward() can only
* be called subsequent to a Forward() call.
*
* Args: hmm - model, probability form, single precision
* s - sequence 0..L-1, upper case
* scl - 0..L array of scalefactors from the forward calculation.
* ret_bck - RETURN: forward calculation matrix
*
* Return: (void)
* bck and scl are alloc'ed here, and must be free'd by caller:
* Free2DArray(bck, L+2)
* free(scl)
*/
void
Backward(struct hmm_struc *hmm, char *s, float *scl, struct forback_s ***ret_bck)
{
char *seq; /* sequence, 1..L */
int L; /* length of seq */
struct forback_s **bck; /* the calculation grid */
int i; /* counter for sequence position: 0,1..L,L+1 */
int k; /* counter for model position: 0,1..M,M+1 */
int i_symidx; /* hint for symbol index in alphabet */
char *alphptr; /* ptr into alphabet (optimization) */
struct forback_s *thisrow;
struct forback_s *nextrow;
/********************************************
* Initial setup and allocations
********************************************/
/* convert sequence to 1..L for ease of indexing */
L = strlen(s);
seq = (char *) MallocOrDie((L+3) * sizeof(char));
strcpy(seq+1, s);
seq[0] = seq[L+1] = ' ';
seq[L+2] = '\0';
/* allocate the calculation matrix */
bck = (struct forback_s **) MallocOrDie (sizeof(struct forback_s *) * (L+2));
for (i = 0; i <= L+1; i++)
bck[i] = (struct forback_s *) MallocOrDie (sizeof(struct forback_s) * (hmm->M + 2));
/********************************************
* Initialization
********************************************/
/* set up the L+1,M+1 cell */
bck[L+1][hmm->M+1].score_m = 1.0;
bck[L+1][hmm->M+1].score_d = 0.0;
bck[L+1][hmm->M+1].score_i = 0.0;
/* initialize the last row */
for (k = 0; k <= hmm->M; k++)
{
bck[L+1][k].score_m = 0.0;
bck[L+1][k].score_i = 0.0;
}
/********************************************
* Recursion: fill in the bck matrix
********************************************/
for (i = L; i >= 0; i--)
{
/* get ptrs into current and next row. */
thisrow = bck[i];
nextrow = bck[i+1];
/* initialize in this row */
thisrow[hmm->M+1].score_m = 0.0;
thisrow[hmm->M+1].score_d = 0.0;
thisrow[hmm->M+1].score_i = 0.0;
/* optimization: get an index for symbol in next row
while we're in the outer loop. */
if ((alphptr = strchr(Alphabet, seq[i])) != NULL)
i_symidx = alphptr - Alphabet;
else
i_symidx = -1; /* too bad; it's a degenerate symbol */
/* recursion, calculation of backward variable */
for (k = hmm->M; k >= 0; k--)
{
/* Transitions.
*/
thisrow[k].score_m = nextrow[k+1].score_m * hmm->mat[k].t[MATCH] +
nextrow[k].score_i * hmm->mat[k].t[INSERT] +
thisrow[k+1].score_d * hmm->mat[k].t[DELETE];
thisrow[k].score_d = nextrow[k+1].score_m * hmm->del[k].t[MATCH] +
nextrow[k].score_i * hmm->del[k].t[INSERT] +
thisrow[k+1].score_d * hmm->del[k].t[DELETE];
thisrow[k].score_i = nextrow[k+1].score_m * hmm->ins[k].t[MATCH] +
nextrow[k].score_i * hmm->ins[k].t[INSERT] +
thisrow[k+1].score_d * hmm->ins[k].t[DELETE];
/* Emissions.
*/
if (i_symidx >= 0) {
thisrow[k].score_m *= hmm->mat[k].p[i_symidx];
thisrow[k].score_i *= hmm->ins[k].p[i_symidx];
}
else if (i > 0) {
thisrow[k].score_m *= ForbackSymscore(seq[i], hmm->mat[k].p, TRUE);
thisrow[k].score_i *= ForbackSymscore(seq[i], hmm->ins[k].p, TRUE);
}
}
/* scale the current row */
for (k = 0; k <= hmm->M; k++)
{
thisrow[k].score_m *= scl[i];
thisrow[k].score_d *= scl[i];
thisrow[k].score_i *= scl[i];
}
}
/******************************************
* Debugging information
******************************************/
#ifdef EXTREME_DEBUG
DumpForbackMatrix(bck, L, hmm->M, scl);
#endif /* EXTREME_DEBUG */
/********************************************
* Garbage collection and return: caller is responsible for free'ing bck and scl!
********************************************/
if (ret_bck != NULL) *ret_bck = bck;
free(seq);
}
/* Function: AlignmentConfidence()
*
* Purpose: Determines the probability distribution for a symbol i
* being aligned to 2M+1 possible insert plus match states.
*
* These probabilities are stored in a forback_s matrix.
* The score_d entries in this matrix are unused.
*
* Args: hmm - model, probability form, single precision
* L - length of sequence
* fwd - calculated forward matrix
* bck - calculated backward matrix
* scl - scale factors used in fwd and bck
* ret_conf - RETURN: confidence probabilities for each symbol
*
* Return: void.
* conf is allocated here, must be free'd by caller.
*/
void
AlignmentConfidence(struct hmm_struc *hmm, int L,
struct forback_s **fwd, struct forback_s **bck, float *scl,
struct forback_s ***ret_conf)
{
struct forback_s **conf;
int i, k;
float norm;
/* allocate the calculation matrix */
conf = (struct forback_s **) MallocOrDie (sizeof(struct forback_s *) * (L+2));
for (i = 0; i <= L+1; i++)
conf[i] = (struct forback_s *) MallocOrDie (sizeof(struct forback_s) * (hmm->M + 2));
/* Sum.
* Note the removal of a scalefactor from the delete path, to avoid double
* counting it.
*/
for (i = 0; i <= L; i++)
for (k = 0; k <= hmm->M; k++)
{
conf[i][k].score_m = bck[i+1][k+1].score_m * hmm->mat[k].t[MATCH] +
bck[i][k+1].score_d * hmm->mat[k].t[DELETE] / scl[i] +
bck[i+1][k].score_i * hmm->mat[k].t[INSERT];
conf[i][k].score_m *= fwd[i][k].score_m;
conf[i][k].score_i = bck[i+1][k+1].score_m * hmm->ins[k].t[MATCH] +
bck[i][k+1].score_d * hmm->ins[k].t[DELETE] / scl[i] +
bck[i+1][k].score_i * hmm->ins[k].t[INSERT];
conf[i][k].score_i *= fwd[i][k].score_i;
}
/* Normalize across all match + insert states in row.
*/
for (i = 0; i <= L; i++)
{
norm = 0.0;
for (k = 0; k <= hmm->M; k++)
{
norm += conf[i][k].score_m;
norm += conf[i][k].score_i;
}
for (k = 0; k <= hmm->M; k++)
{
conf[i][k].score_m /= norm;
conf[i][k].score_i /= norm;
}
}
/* Return.
*/
*ret_conf = conf;
}
/* Function: TraceConfidence()
*
* Purpose: Find the confidence estimates on each aligned sequence symbol
* for a particular traceback, using the full matrix constructed
* by AlignmentConfidence().
*
* Args: cmx - full confidence matrix produced by AlignmentConfidence()
* L - length of seq (L+2 rows in cmx)
* tr - traceback to evaulate symbol alignment confidences on
* ret_conf - RETURN: 0..L-1 array of confidence estimates
* for each position in original sequence.
*/
void
TraceConfidence(struct forback_s **cmx, int L, struct trace_s *tr, float **ret_conf)
{
int tpos; /* position in trace */
float *conf; /* 0..L-1 array of symbol alignment confidence values */
conf = (float *) MallocOrDie (L * sizeof(float));
for (tpos = 1; tpos < tr->tlen-1; tpos++)
{
switch (tr->statetype[tpos]) {
case MATCH:
conf[tr->rpos[tpos]] = cmx[tr->rpos[tpos]+1][tr->nodeidx[tpos]].score_m;
break;
case DELETE:
break;
case INSERT:
conf[tr->rpos[tpos]] = cmx[tr->rpos[tpos]+1][tr->nodeidx[tpos]].score_i;
break;
default:
Die("Bogus statetype %d in TraceConfidence()", tr->statetype[tpos]);
}
}
*ret_conf = conf;
}
/* Function: ForbackCount()
*
* Purpose: Count probabilities of state transitions and symbol emissions from a
* forward-backward calculation on a single sequence. Results are
* stored in a counts-based HMM which will be normalized later.
* The full Baum-Welch equivalent to TraceCount() for Viterbi.
*
* Args: hmm - model. Probability form.
* prior - Used only for alphabet information.
* seq - sequence, 0..L-1
* L - length of sequence
* fwd - forward matrix produced by Forward()
* bck - backward matrix produced by Backward()
* scl - scale factors used by forward and backward
* count - counts-based HMM to accumulate counts in.
* Caller is responsible for allocating this structure
* and initializing it properly.
*
* Return: (void)
* Some counts are added to the count-based HMM.
*/
void
ForbackCount(struct hmm_struc *hmm, char *seq, int L, float weight,
struct forback_s **fwd, struct forback_s **bck,
float *scl, struct hmm_struc *count)
{
int i, k;
float tmp_m1, tmp_d1, tmp_i0;
float prob; /* overall scaled probability of the seq | model */
float wt;
int i_symidx;
char *alphptr;
/* "prob" is the total P(seq | model). We have to normalize by this.
* We want to know, how much of the total alignment probability
* flows through a particular state or state transition?
*
* Note the removal of a scale factor from the delete paths, to prevent
* double-counting it in both the fwd and bck variable.
*/
prob = fwd[L+1][hmm->M+1].score_m / weight;
if (prob == 0.0) return; /* silent! we have precision problems. */
for (i = 0; i <= L; i++)
{
/* optimization: get an index for this symbol
while we're in the outer loop. */
if (i > 0)
{
if ((alphptr = strchr(Alphabet, seq[i-1])) != NULL)
i_symidx = alphptr - Alphabet;
else
i_symidx = -1; /* too bad; it's a degenerate symbol */
}
for (k = 0; k <= hmm->M; k++)
{
tmp_m1 = fwd[i][k].score_m * hmm->mat[k].t[MATCH] * bck[i+1][k+1].score_m;
tmp_d1 = fwd[i][k].score_m * hmm->mat[k].t[DELETE] * bck[i][k+1].score_d / scl[i];
tmp_i0 = fwd[i][k].score_m * hmm->mat[k].t[INSERT] * bck[i+1][k].score_i;
count->mat[k].t[MATCH] += tmp_m1 / prob;
count->mat[k].t[DELETE] += tmp_d1 / prob;
count->mat[k].t[INSERT] += tmp_i0 / prob;
if (i > 0)
{
wt = ((tmp_m1 + tmp_d1 + tmp_i0) / prob);
if (i_symidx >= 0)
count->mat[k].p[i_symidx] += wt;
else
CountSymbol(seq[i-1], wt, count->mat[k].p);
}
count->del[k].t[MATCH] += fwd[i][k].score_d * hmm->del[k].t[MATCH] * bck[i+1][k+1].score_m / prob;
count->del[k].t[DELETE] += fwd[i][k].score_d * hmm->del[k].t[DELETE] * bck[i][k+1].score_d / (scl[i] * prob);
count->del[k].t[INSERT] += fwd[i][k].score_d * hmm->del[k].t[INSERT] * bck[i+1][k].score_i / prob;
tmp_m1 = fwd[i][k].score_i * hmm->ins[k].t[MATCH] * bck[i+1][k+1].score_m;
tmp_d1 = fwd[i][k].score_i * hmm->ins[k].t[DELETE] * bck[i][k+1].score_d / scl[i];
tmp_i0 = fwd[i][k].score_i * hmm->ins[k].t[INSERT] * bck[i+1][k].score_i;
count->ins[k].t[MATCH] += tmp_m1 / prob;
count->ins[k].t[DELETE] += tmp_d1 / prob;
count->ins[k].t[INSERT] += tmp_i0 / prob;
if (i > 0)
{
wt = ((tmp_m1 + tmp_d1 + tmp_i0) / prob);
if (i_symidx >= 0)
count->ins[k].p[i_symidx] += wt;
else
CountSymbol(seq[i-1], wt, count->ins[k].p);
}
}
}
}
/* Function: ForwardScore()
*
* Purpose: Given a forward matrix and the scalefactors,
* return the log likeihood log P(S | M) in base 2 (bits)
*/
float
ForwardScore(struct forback_s **fwd, int L, int M, float *scl)
{
int i;
float score;
score = LOG2(fwd[L+1][M+1].score_m);
for (i = 0; i <= L; i++)
score -= LOG2(scl[i]);
return score;
}
/* Function: BackwardScore()
*
* Purpose: Given a backward matrix and the scalefactors,
* return the log likeihood log P(S | M) in base 2 (bits)
*/
float
BackwardScore(struct forback_s **bck, int L, float *scl)
{
int i;
float score;
score = LOG2(bck[0][0].score_m);
for (i = 0; i <= L; i++)
score -= LOG2(scl[i]);
return score;
}
/* Function: RandomScore()
*
* Purpose: Return the log P(S | R), the log likelihood of the
* sequence given the random model.
*/
float
RandomScore(float *randomseq, char *seq)
{
float ll; /* log likelihood */
ll = 0.0;
for (; *seq != '\0'; seq++)
ll += LOG2(ForbackSymscore(*seq, randomseq, TRUE));
return ll;
}
/* Function: ForbackSymscore()
*
* Look up the score of sequence character c, given a table of values
* from the P(x | y) table for an HMM state. The table must be in
* precisely the same order as the alphabet. For nucleic acids, both
* the alphabet and the table must be in the order "ACGT".
*
* If hyperbayes is TRUE, it returns the summed probability of degenerate
* symbols. This is formally correct but produces aberrantly high scores
* on garbage degenerate sequences in database searches.
*
* Returns the looked-up score from the table.
*/
float
ForbackSymscore(char x, float *scores, int hyperbayes)
{
float result;
int count;
/* simple case: x is in the alphabet */
if (strchr(Alphabet, x) != NULL) return scores[SYMIDX(x)];
result = 0.0;
if (Alphabet_type == kAmino)
{
switch (x) {
case 'B':
result += scores[SYMIDX('N')];
result += scores[SYMIDX('D')];
count = 2;
break;
case 'Z':
result += scores[SYMIDX('Q')];
result += scores[SYMIDX('E')];
count = 2;
break;
default:
Warn("Unrecognized character %c (%d) in sequence", x, (int) x);
/* break thru to case 'X' */
case 'X':
result = 1.0;
count = 20;
break;
}
}
else if (Alphabet_type == kDNA || Alphabet_type == kRNA)
{
switch (x) {
case 'B': result = scores[1] + scores[2] + scores[3]; count = 3; break;
case 'D': result = scores[0] + scores[2] + scores[3]; count = 3; break;
case 'H': result = scores[0] + scores[1] + scores[3]; count = 3; break;
case 'K': result = scores[2] + scores[3]; count = 2; break;
case 'M': result = scores[0] + scores[1]; count = 2; break;
case 'R': result = scores[0] + scores[2]; count = 2; break;
case 'S': result = scores[1] + scores[2]; count = 2; break;
case 'T': result = scores[3]; count = 1; break;
case 'U': result = scores[3]; count = 1; break;
case 'V': result = scores[0] + scores[1] + scores[2]; count = 3; break;
case 'W': result = scores[0] + scores[3]; count = 2; break;
case 'Y': result = scores[1] + scores[3]; count = 2; break;
default:
Warn("unrecognized character %c (%d) in sequence", x, (int) x);
/* break through to case 'N' */
case 'N':
result = 1.0; count = 4; break;
}
}
else
{
Warn("unrecognized character %c (%d) in sequence\n", x, (int) x);
result = 1.0;
count = Alphabet_size;
}
/* If you want correct probabilities, set hyperbayes TRUE and
* use summed probabilities. But if you want it to work, and not
* give high probabilities on garbage poly-X sequences, set
* hyperbayes FALSE and use a simple expected probability.
*/
if (! hyperbayes) result /= (float) count;
return result;
}
/* Function: DumpForbackMatrix()
*
* Purpose: Debugging output; dump the contents of a matrix
* to stdout. Because it prints everything, it's
* only useful on small test matrices.
*
* Returns: (void)
*/
void
DumpForbackMatrix(struct forback_s **mx, int L, int M, float *scalefactors)
{
int i; /* counter for rows (sequence symbols) */
int k; /* counter for columns (states) */
for (i = 0; i <= L+1; i++)
{
/* print match states */
for (k = 0; k <= M+1; k++) printf("%6.4g ", mx[i][k].score_m);
printf(":: %.4g\n", scalefactors[i]);
/* print delete states */
for (k = 0; k <= M+1; k++) printf("%6.4g ", mx[i][k].score_d);
putchar('\n');
/* print insert states */
for (k = 0; k <= M+1; k++) printf("%6.4g ", mx[i][k].score_i);
putchar('\n');
putchar('\n');
}
}