
                                 fdnacomp 



Function

   DNA compatibility algorithm

Description

   Estimates phylogenies from nucleic acid sequence data using the
   compatibility criterion, which searches for the largest number of
   sites which could have all states (nucleotides) uniquely evolved on
   the same tree. Compatibility is particularly appropriate when sites
   vary greatly in their rates of evolution, but we do not know in
   advance which are the less reliable ones.

Algorithm

   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:
    1. Each character evolves independently.
    2. Different lineages evolve independently.
    3. The ancestral base at each site is unknown.
    4. The rates of change in most sites over the time spans involved in
       the the divergence of the group are very small.
    5. A few of the sites have very high rates of change.
    6. We do not know in advance which are the high and which the low
       rate sites.

   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.

Usage

   Here is a sample session with fdnacomp


% fdnacomp -ancseq -stepbox -printdata 
DNA compatibility algorithm
Input sequence: dnacomp.dat
Input tree file: 
Weights file: 
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 sequence: dnacomp.dat
Input tree file: dnacomptree.dat
Weights file: 
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

Command line arguments

   Standard (Mandatory) qualifiers:
  [-sequence]          seqsetall  File containing one or more sequence
                                  alignments
  [-intreefile]        tree       (no help text) tree value
   -weights            properties Weights file
  [-outfile]           outfile    Output file name

   Additional (Optional) qualifiers (* if not always prompted):
   -outgrno            integer    Species number to use as outgroup
*  -njumble            integer    Number of times to randomise
*  -seed               integer    Random number seed between 1 and 32767 (must
                                  be odd)
   -[no]trout          toggle     Write out trees to tree file
*  -outtreefile        outfile    Tree file name
   -printdata          boolean    Print data at start of run
   -[no]progress       boolean    Print indications of progress of run
   -[no]treeprint      boolean    Print out tree
   -stepbox            boolean    Print steps & compatibility at sites
   -ancseq             boolean    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
   -sformat1           string     Input sequence format
   -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 standard output
   -filter             boolean    Read standard input, write 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. 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 deaths


   Standard (Mandatory) qualifiers Allowed values Default
   [-sequence]
   (Parameter 1) File containing one or more sequence alignments Readable
   sets of sequences
   [-intreefile]
   (Parameter 2) (no help text) tree value Phylogenetic tree
   -weights Weights file Property value(s)
   [-outfile]
   (Parameter 3) Output file name Output file <sequence>.fdnacomp
   Additional (Optional) qualifiers Allowed values Default
   -outgrno Species number to use as outgroup Integer 0 or more 0
   -njumble Number of times to randomise Integer 0 or more 0
   -seed Random number seed between 1 and 32767 (must be odd) Integer
   from 1 to 32767 1
   -[no]trout Write out trees to tree file Toggle value Yes/No Yes
   -outtreefile Tree file name Output file
   -printdata Print data at start of run Boolean value Yes/No No
   -[no]progress Print indications of progress of run Boolean value
   Yes/No Yes
   -[no]treeprint Print out tree Boolean value Yes/No Yes
   -stepbox Print steps & compatibility at sites Boolean value Yes/No No
   -ancseq Print sequences at all nodes of tree Boolean value Yes/No No
   Advanced (Unprompted) qualifiers Allowed values Default
   (none)

Input file format

   fdnacomp reads any normal sequence USAs.

  Input files for usage example

  File: dnacomp.dat

    5   13
Alpha     AACGUGGCCAAAU
Beta      AAGGUCGCCAAAC
Gamma     CAUUUCGUCACAA
Delta     GGUAUUUCGGCCU
Epsilon   GGGAUCUCGGCCC

  Input files for usage example 2

  File: dnacomptree.dat

   ((((Epsilon,Delta),Gamma),Beta),Alpha);

Output file format

   fdnacomp output is standard: if option 1 is toggled on, the data is
   printed out, with the convention that "." means "the same as in the
   first species". Then comes a list of equally parsimonious trees, and
   (if option 2 is toggled on) a table of the number of changes of state
   required in each character. If option 5 is toggled on, a table is
   printed out after each tree, showing for each branch whether there are
   known to be changes in the branch, and what the states are inferred to
   have been at the top end of the branch. If the inferred state is a "?"
   or one of the IUB ambiguity symbols, there will be multiple
   equally-parsimonious assignments of states; the user must work these
   out for themselves by hand. A "?" in the reconstructed states means
   that in addition to one or more bases, a gap may or may not be
   present. If option 6 is left in its default state the trees found will
   be written to a tree file, so that they are available to be used in
   other programs. If the program finds multiple trees tied for best, all
   of these are written out onto the output tree file. Each is followed
   by a numerical weight in square brackets (such as [0.25000]). This is
   needed when we use the trees to make a consensus tree of the results
   of bootstrapping or jackknifing, to avoid overrepresenting replicates
   that find many tied trees.

  Output files for usage example

  File: dnacomp.fdnacomp


DNA compatibility algorithm, version 3.6b

 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


  File: dnacomp.treefile

   ((((Epsilon,Delta),Gamma),Beta),Alpha);

  Output files for usage example 2

  File: dnacomp.fdnacomp


DNA compatibility algorithm, version 3.6b

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


Data files

   None

Notes

   None.

References

   None.

Warnings

   None.

Diagnostic Error Messages

   None.

Exit status

   It always exits with status 0.

Known bugs

   None.

See also

   Program name                         Description
   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       Estimates 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 pasimony algorithm
   frestboot    Bootstrapped restriction sites algorithm
   frestdist    Distance matrix from restriction sites or fragments
   frestml      Restriction site maximum Likelihood method
   fseqboot     Bootstrapped sequences algorithm
   fseqbootall  Bootstrapped sequences algorithm

Author(s)

   This program is an EMBOSS conversion of a program written by Joe
   Felsenstein as part of his PHYLIP package.

   Although we take every care to ensure that the results of the EMBOSS
   version are identical to those from the original package, we recommend
   that you check your inputs give the same results in both versions
   before publication.

   Please report all bugs in the EMBOSS version to the EMBOSS bug team,
   not to the original author.

History

   Written (2004) - Joe Felsenstein, University of Washington.

   Converted (August 2004) to an EMBASSY program by the EMBOSS team.

Target users

   This program is intended to be used by everyone and everything, from
   naive users to embedded scripts.
