|   | fdnacomp | 
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This program implements the compatibility method for DNA sequence data. For a four-state character without a character-state tree, as in DNA sequences, the usual clique theorems cannot be applied. The approach taken in this program is to directly evaluate each tree topology by counting how many substitutions are needed in each site, comparing this to the minimum number that might be needed (one less than the number of bases observed at that site), and then evaluating the number of sites which achieve the minimum number. This is the evaluation of the tree (the number of compatible sites), and the topology is chosen so as to maximize that number.
Compatibility methods originated with Le Quesne's (1969) suggestion that one ought to look for trees supported by the largest number of perfectly fitting (compatible) characters. Fitch (1975) showed by counterexample that one could not use the pairwise compatibility methods used in CLIQUE to discover the largest clique of jointly compatible characters.
The assumptions of this method are similar to those of CLIQUE. In a paper in the Biological Journal of the Linnean Society (1981b) I discuss this matter extensively. In effect, the assumptions are that:
That these are the assumptions of compatibility methods has been documented in a series of papers of mine: (1973a, 1978b, 1979, 1981b, 1983b, 1988b). For an opposing view arguing that arguments such as mine are invalid and that parsimony (and perhaps compatibility) methods make no substantive assumptions such as these, see the papers by Farris (1983) and Sober (1983a, 1983b, 1988), but also read the exchange between Felsenstein and Sober (1986).
There is, however, some reason to believe that the present criterion is not the proper way to correct for the presence of some sites with high rates of change in nucleotide sequence data. It can be argued that sites showing more than two nucleotide states, even if those are compatible with the other sites, are also candidates for sites with high rates of change. It might then be more proper to use DNAPARS with the Threshold option with a threshold value of 2.
Change from an occupied site to a gap is counted as one change. Reversion from a gap to an occupied site is allowed and is also counted as one change. Note that this in effect assumes that a gap N bases long is N separate events. This may be an overcorrection. When we have nonoverlapping gaps, we could instead code a gap as a single event by changing all but the first "-" in the gap into "?" characters. In this way only the first base of the gap causes the program to infer a change.
If the U (User Tree) option is used and more than one tree is supplied, the program also performs a statistical test of each of these trees against the one with highest likelihood. If there are two user trees, the test done is one which is due to Kishino and Hasegawa (1989), a version of a test originally introduced by Templeton (1983). In this implementation it uses the mean and variance of weighted compatibility differences between trees, taken across sites. If the two trees compatibilities are more than 1.96 standard deviations different then the trees are declared significantly different.
If there are more than two trees, the test done is an extension of the KHT test, due to Shimodaira and Hasegawa (1999). They pointed out that a correction for the number of trees was necessary, and they introduced a resampling method to make this correction. In the version used here the variances and covariances of the sum of weighted compatibilities of sites are computed for all pairs of trees. To test whether the difference between each tree and the best one is larger than could have been expected if they all had the same expected compatibility, compatibilities for all trees are sampled with these covariances and equal means (Shimodaira and Hasegawa's "least favorable hypothesis"), and a P value is computed from the fraction of times the difference between the tree's value and the highest compatibility exceeds that actually observed. Note that this sampling needs random numbers, and so the program will prompt the user for a random number seed if one has not already been supplied. With the two-tree KHT test no random numbers are used.
In either the KHT or the SH test the program prints out a table of the compatibility of each tree, the differences of each from the highest one, the variance of that quantity as determined by the compatibility differences at individual sites, and a conclusion as to whether that tree is or is not significantly worse than the best one.
The algorithm is a straightforward modification of DNAPARS, but with some extra machinery added to calculate, as each species is added, how many base changes are the minimum which could be required at that site. The program runs fairly quickly.
| % fdnacomp -ancseq -stepbox -printdata DNA compatibility algorithm Input (aligned) nucleotide sequence set(s): dnacomp.dat Phylip tree file (optional): Phylip weights file (optional): Phylip dnacomp program output file [dnacomp.fdnacomp]: Adding species: 1. Alpha 2. Beta 3. Gamma 4. Delta 5. Epsilon Doing global rearrangements !---------! ......... Output written to file "dnacomp.fdnacomp" Trees also written onto file "dnacomp.treefile" | 
Go to the input files for this example
Go to the output files for this example
Example 2
| % fdnacomp DNA compatibility algorithm Input (aligned) nucleotide sequence set(s): dnacomp.dat Phylip tree file (optional): dnacomptree.dat Phylip weights file (optional): Phylip dnacomp program output file [dnacomp.fdnacomp]: Output written to file "dnacomp.fdnacomp" Trees also written onto file "dnacomp.treefile" | 
Go to the input files for this example
Go to the output files for this example
| 
DNA compatibility algorithm
Version: EMBOSS:6.6.0.0
   Standard (Mandatory) qualifiers:
  [-sequence]          seqsetall  File containing one or more sequence
                                  alignments
  [-intreefile]        tree       Phylip tree file (optional)
   -weights            properties Phylip weights file (optional)
  [-outfile]           outfile    [*.fdnacomp] Phylip dnacomp program output
                                  file
   Additional (Optional) qualifiers (* if not always prompted):
   -outgrno            integer    [0] Species number to use as outgroup
                                  (Integer 0 or more)
*  -njumble            integer    [0] Number of times to randomise (Integer 0
                                  or more)
*  -seed               integer    [1] Random number seed between 1 and 32767
                                  (must be odd) (Integer from 1 to 32767)
   -[no]trout          toggle     [Y] Write out trees to tree file
*  -outtreefile        outfile    [*.fdnacomp] Phylip tree output file
                                  (optional)
   -printdata          boolean    [N] Print data at start of run
   -[no]progress       boolean    [Y] Print indications of progress of run
   -[no]treeprint      boolean    [Y] Print out tree
   -stepbox            boolean    [N] Print steps & compatibility at sites
   -ancseq             boolean    [N] Print sequences at all nodes of tree
   Advanced (Unprompted) qualifiers: (none)
   Associated qualifiers:
   "-sequence" associated qualifiers
   -sbegin1            integer    Start of each sequence to be used
   -send1              integer    End of each sequence to be used
   -sreverse1          boolean    Reverse (if DNA)
   -sask1              boolean    Ask for begin/end/reverse
   -snucleotide1       boolean    Sequence is nucleotide
   -sprotein1          boolean    Sequence is protein
   -slower1            boolean    Make lower case
   -supper1            boolean    Make upper case
   -scircular1         boolean    Sequence is circular
   -squick1            boolean    Read id and sequence only
   -sformat1           string     Input sequence format
   -iquery1            string     Input query fields or ID list
   -ioffset1           integer    Input start position offset
   -sdbname1           string     Database name
   -sid1               string     Entryname
   -ufo1               string     UFO features
   -fformat1           string     Features format
   -fopenfile1         string     Features file name
   "-outfile" associated qualifiers
   -odirectory3        string     Output directory
   "-outtreefile" associated qualifiers
   -odirectory         string     Output directory
   General qualifiers:
   -auto               boolean    Turn off prompts
   -stdout             boolean    Write first file to standard output
   -filter             boolean    Read first file from standard input, write
                                  first file to standard output
   -options            boolean    Prompt for standard and additional values
   -debug              boolean    Write debug output to program.dbg
   -verbose            boolean    Report some/full command line options
   -help               boolean    Report command line options and exit. More
                                  information on associated and general
                                  qualifiers can be found with -help -verbose
   -warning            boolean    Report warnings
   -error              boolean    Report errors
   -fatal              boolean    Report fatal errors
   -die                boolean    Report dying program messages
   -version            boolean    Report version number and exit
 | 
| Qualifier | Type | Description | Allowed values | Default | 
|---|---|---|---|---|
| Standard (Mandatory) qualifiers | ||||
| [-sequence] (Parameter 1) | seqsetall | File containing one or more sequence alignments | Readable sets of sequences | Required | 
| [-intreefile] (Parameter 2) | tree | Phylip tree file (optional) | Phylogenetic tree | |
| -weights | properties | Phylip weights file (optional) | Property value(s) | |
| [-outfile] (Parameter 3) | outfile | Phylip dnacomp program output file | Output file | <*>.fdnacomp | 
| Additional (Optional) qualifiers | ||||
| -outgrno | integer | Species number to use as outgroup | Integer 0 or more | 0 | 
| -njumble | integer | Number of times to randomise | Integer 0 or more | 0 | 
| -seed | integer | Random number seed between 1 and 32767 (must be odd) | Integer from 1 to 32767 | 1 | 
| -[no]trout | toggle | Write out trees to tree file | Toggle value Yes/No | Yes | 
| -outtreefile | outfile | Phylip tree output file (optional) | Output file | <*>.fdnacomp | 
| -printdata | boolean | Print data at start of run | Boolean value Yes/No | No | 
| -[no]progress | boolean | Print indications of progress of run | Boolean value Yes/No | Yes | 
| -[no]treeprint | boolean | Print out tree | Boolean value Yes/No | Yes | 
| -stepbox | boolean | Print steps & compatibility at sites | Boolean value Yes/No | No | 
| -ancseq | boolean | Print sequences at all nodes of tree | Boolean value Yes/No | No | 
| Advanced (Unprompted) qualifiers | ||||
| (none) | ||||
| Associated qualifiers | ||||
| "-sequence" associated seqsetall qualifiers | ||||
| -sbegin1 -sbegin_sequence | integer | Start of each sequence to be used | Any integer value | 0 | 
| -send1 -send_sequence | integer | End of each sequence to be used | Any integer value | 0 | 
| -sreverse1 -sreverse_sequence | boolean | Reverse (if DNA) | Boolean value Yes/No | N | 
| -sask1 -sask_sequence | boolean | Ask for begin/end/reverse | Boolean value Yes/No | N | 
| -snucleotide1 -snucleotide_sequence | boolean | Sequence is nucleotide | Boolean value Yes/No | N | 
| -sprotein1 -sprotein_sequence | boolean | Sequence is protein | Boolean value Yes/No | N | 
| -slower1 -slower_sequence | boolean | Make lower case | Boolean value Yes/No | N | 
| -supper1 -supper_sequence | boolean | Make upper case | Boolean value Yes/No | N | 
| -scircular1 -scircular_sequence | boolean | Sequence is circular | Boolean value Yes/No | N | 
| -squick1 -squick_sequence | boolean | Read id and sequence only | Boolean value Yes/No | N | 
| -sformat1 -sformat_sequence | string | Input sequence format | Any string | |
| -iquery1 -iquery_sequence | string | Input query fields or ID list | Any string | |
| -ioffset1 -ioffset_sequence | integer | Input start position offset | Any integer value | 0 | 
| -sdbname1 -sdbname_sequence | string | Database name | Any string | |
| -sid1 -sid_sequence | string | Entryname | Any string | |
| -ufo1 -ufo_sequence | string | UFO features | Any string | |
| -fformat1 -fformat_sequence | string | Features format | Any string | |
| -fopenfile1 -fopenfile_sequence | string | Features file name | Any string | |
| "-outfile" associated outfile qualifiers | ||||
| -odirectory3 -odirectory_outfile | string | Output directory | Any string | |
| "-outtreefile" associated outfile qualifiers | ||||
| -odirectory | string | Output directory | Any string | |
| General qualifiers | ||||
| -auto | boolean | Turn off prompts | Boolean value Yes/No | N | 
| -stdout | boolean | Write first file to standard output | Boolean value Yes/No | N | 
| -filter | boolean | Read first file from standard input, write first file to standard output | Boolean value Yes/No | N | 
| -options | boolean | Prompt for standard and additional values | Boolean value Yes/No | N | 
| -debug | boolean | Write debug output to program.dbg | Boolean value Yes/No | N | 
| -verbose | boolean | Report some/full command line options | Boolean value Yes/No | Y | 
| -help | boolean | Report command line options and exit. More information on associated and general qualifiers can be found with -help -verbose | Boolean value Yes/No | N | 
| -warning | boolean | Report warnings | Boolean value Yes/No | Y | 
| -error | boolean | Report errors | Boolean value Yes/No | Y | 
| -fatal | boolean | Report fatal errors | Boolean value Yes/No | Y | 
| -die | boolean | Report dying program messages | Boolean value Yes/No | Y | 
| -version | boolean | Report version number and exit | Boolean value Yes/No | N | 
| 
    5   13
Alpha     AACGUGGCCAAAU
Beta      AAGGUCGCCAAAC
Gamma     CAUUUCGUCACAA
Delta     GGUAUUUCGGCCU
Epsilon   GGGAUCUCGGCCC
 | 
| ((((Epsilon,Delta),Gamma),Beta),Alpha); | 
| 
DNA compatibility algorithm, version 3.69.650
 5 species,  13  sites
Name            Sequences
----            ---------
Alpha        AACGTGGCCA AAT
Beta         AAGGTCGCCA AAC
Gamma        CATTTCGTCA CAA
Delta        GGTATTTCGG CCT
Epsilon      GGGATCTCGG CCC
One most parsimonious tree found:
           +--Epsilon   
        +--4  
     +--3  +--Delta     
     !  !  
  +--2  +-----Gamma     
  !  !  
  1  +--------Beta      
  !  
  +-----------Alpha     
  remember: this is an unrooted tree!
total number of compatible sites is       11.0
steps in each site:
         0   1   2   3   4   5   6   7   8   9
     *-----------------------------------------
    0|       2   1   3   2   0   2   1   1   1
   10|   1   1   1   3                        
 compatibility (Y or N) of each site with this tree:
      0123456789
     *----------
   0 ! YYNYYYYYY
  10 !YYYN      
From    To     Any Steps?    State at upper node
                            
          1                AABGTSGCCA AAY
   1      2        maybe   AABGTCGCCA AAY
   2      3         yes    VAKDTCGCCA CAY
   3      4         yes    GGKATCTCGG CCY
   4   Epsilon     maybe   GGGATCTCGG CCC
   4   Delta        yes    GGTATTTCGG CCT
   3   Gamma        yes    CATTTCGTCA CAA
   2   Beta        maybe   AAGGTCGCCA AAC
   1   Alpha       maybe   AACGTGGCCA AAT
 | 
| ((((Epsilon,Delta),Gamma),Beta),Alpha); | 
| 
DNA compatibility algorithm, version 3.69.650
User-defined tree:
           +--Epsilon   
        +--4  
     +--3  +--Delta     
     !  !  
  +--2  +-----Gamma     
  !  !  
  1  +--------Beta      
  !  
  +-----------Alpha     
  remember: this is an unrooted tree!
total number of compatible sites is       11.0
 | 
| Program name | Description | 
|---|---|
| distmat | Create a distance matrix from a multiple sequence alignment | 
| ednacomp | DNA compatibility algorithm | 
| ednadist | Nucleic acid sequence distance matrix program | 
| ednainvar | Nucleic acid sequence invariants method | 
| ednaml | Phylogenies from nucleic acid maximum likelihood | 
| ednamlk | Phylogenies from nucleic acid maximum likelihood with clock | 
| ednapars | DNA parsimony algorithm | 
| ednapenny | Penny algorithm for DNA | 
| eprotdist | Protein distance algorithm | 
| eprotpars | Protein parsimony algorithm | 
| erestml | Restriction site maximum likelihood method | 
| eseqboot | Bootstrapped sequences algorithm | 
| fdiscboot | Bootstrapped discrete sites algorithm | 
| fdnadist | Nucleic acid sequence distance matrix program | 
| fdnainvar | Nucleic acid sequence invariants method | 
| fdnaml | Estimate nucleotide phylogeny by maximum likelihood | 
| fdnamlk | Estimates nucleotide phylogeny by maximum likelihood | 
| fdnamove | Interactive DNA parsimony | 
| fdnapars | DNA parsimony algorithm | 
| fdnapenny | Penny algorithm for DNA | 
| fdolmove | Interactive Dollo or polymorphism parsimony | 
| ffreqboot | Bootstrapped genetic frequencies algorithm | 
| fproml | Protein phylogeny by maximum likelihood | 
| fpromlk | Protein phylogeny by maximum likelihood | 
| fprotdist | Protein distance algorithm | 
| fprotpars | Protein parsimony algorithm | 
| frestboot | Bootstrapped restriction sites algorithm | 
| frestdist | Calculate distance matrix from restriction sites or fragments | 
| frestml | Restriction site maximum likelihood method | 
| fseqboot | Bootstrapped sequences algorithm | 
| fseqbootall | Bootstrapped sequences algorithm | 
Please report all bugs to the EMBOSS bug team (emboss-bug © emboss.open-bio.org) not to the original author.
Converted (August 2004) to an EMBASSY program by the EMBOSS team.
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