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12 | <DIV ALIGN=RIGHT> |
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13 | version 3.6 |
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14 | </DIV> |
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15 | <P> |
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16 | <DIV ALIGN=CENTER> |
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17 | <H1>FITCH -- Fitch-Margoliash and Least-Squares Distance Methods</H1> |
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18 | </DIV> |
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19 | <P> |
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20 | © Copyright 1986-2002 by the University of |
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21 | Washington. Written by Joseph Felsenstein. Permission is granted to copy |
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22 | this document provided that no fee is charged for it and that this copyright |
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23 | notice is not removed. |
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24 | <P> |
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25 | This program carries out Fitch-Margoliash, Least Squares, |
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26 | and a number of similar methods as described in the documentation |
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27 | file for distance methods. |
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28 | <P> |
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29 | The options for FITCH are selected through the menu, which looks like |
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30 | this: |
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31 | <P> |
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32 | <TABLE><TR><TD BGCOLOR=white> |
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33 | <PRE> |
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34 | |
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35 | Fitch-Margoliash method version 3.6a3 |
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36 | |
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37 | Settings for this run: |
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38 | D Method (F-M, Minimum Evolution)? Fitch-Margoliash |
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39 | U Search for best tree? Yes |
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40 | P Power? 2.00000 |
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41 | - Negative branch lengths allowed? No |
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42 | O Outgroup root? No, use as outgroup species 1 |
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43 | L Lower-triangular data matrix? No |
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44 | R Upper-triangular data matrix? No |
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45 | S Subreplicates? No |
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46 | G Global rearrangements? No |
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47 | J Randomize input order of species? No. Use input order |
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48 | M Analyze multiple data sets? No |
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49 | 0 Terminal type (IBM PC, ANSI, none)? (none) |
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50 | 1 Print out the data at start of run No |
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51 | 2 Print indications of progress of run Yes |
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52 | 3 Print out tree Yes |
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53 | 4 Write out trees onto tree file? Yes |
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54 | |
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55 | Y to accept these or type the letter for one to change |
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56 | |
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57 | </PRE> |
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58 | <P> |
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59 | </TD></TR></TABLE> |
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60 | <P> |
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61 | Most of the input options |
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62 | (U, P, -, O, L, R, S, J, and M) are as given in |
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63 | the documentation page for distance matrix programs, and their input format is |
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64 | the same as given there. The |
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65 | U (User Tree) option has one additional feature when the N (Lengths) |
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66 | option is used. This menu option will appear only if the U (User Tree) |
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67 | option is selected. If N (Lengths) is set to "Yes" then if any branch |
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68 | in the user tree has a branch length, that branch will not have its |
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69 | length iterated. Thus you can prevent all branches from having their |
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70 | lengths changed by giving them all lengths in the user tree, or hold |
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71 | only one length unchanged by giving only that branch a length (such |
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72 | as, for example, 0.00). You may find program RETREE useful for |
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73 | adding and removing branch lengths from a tree. This option can |
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74 | also be used to compute the Average Percent Standard Deviation for a |
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75 | tree obtained from NEIGHBOR, for comparison with trees obtained by |
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76 | FITCH or KITSCH. |
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77 | <P> |
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78 | The D (methods) option allows choice between the Fitch-Margoliash |
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79 | criterion and the Minimum Evolution method (Kidd and Sgaramella-Zonta, 1971; |
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80 | Rzhetsky and Nei, 1993). Minimum Evolution (not to be confused with |
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81 | parsimony) uses the Fitch-Margoliash criterion to fit branch lengths to each |
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82 | topology, but then chooses topologies based on their total branch length |
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83 | (rather than the goodness of fit sum of squares). There is no |
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84 | constraint on negative branch lengths in the Minimum Evolution method; |
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85 | it sometimes gives rather strange results, as it can like solutions |
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86 | that have large negative branch lengths, as these reduce the total |
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87 | sum of branch lengths! |
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88 | </PRE> |
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89 | <P> |
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90 | Another input option available in FITCH that is not available in KITSCH |
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91 | or NEIGHBOR |
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92 | is the G (Global) option. G is the |
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93 | Global search option. This causes, after the last species is added to |
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94 | the tree, each possible group to be removed and re-added. This improves the |
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95 | result, since the position of every species is reconsidered. It |
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96 | approximately triples the run-time of the program. It is not an option in |
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97 | KITSCH because it is the default and is always in force there. The |
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98 | O (Outgroup) option is described in the main |
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99 | documentation file of this package. The O option has no effect if the |
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100 | tree is a user-defined tree (if the U option is in effect). The |
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101 | U (User Tree) option requires an unrooted tree; that is, it require |
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102 | that the tree have a trifurcation at its base: |
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103 | <P> |
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104 | ((A,B),C,(D,E)); |
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105 | <P> |
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106 | The output consists of an unrooted tree and the lengths of the |
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107 | interior segments. The sum of squares is printed out, and if <EM>P = 2.0</EM> |
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108 | Fitch and Margoliash's "average percent standard deviation" is also computed |
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109 | and printed out. This is the sum of squares, divided by |
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110 | N-2, and |
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111 | then square-rooted and then multiplied by 100 (n is the number of |
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112 | species on the tree): |
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113 | <P> |
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114 | APSD = ( SSQ / (N-2) )<SUP>1/2</SUP> x 100. |
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115 | <P> |
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116 | where <EM>N</EM> is the total number of off-diagonal distance measurements that |
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117 | are in the (square) distance matrix. If the S (subreplication) option |
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118 | is in force it is instead the sum of the numbers of replicates in all the |
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119 | non-diagonal |
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120 | cells of the distance matrix. But if the L or R option is also in effect, so |
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121 | that the distance matrix read in is lower- or upper-triangular, then the |
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122 | sum of replicates is only over those cells actually read in. If S is not in |
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123 | force, the number of replicates in |
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124 | each cell is assumed to be 1, so that <EM>N</EM> is <EM>n(n-1)</EM>, |
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125 | where <EM>n</EM> is the number |
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126 | of species. The <EM>APSD</EM> gives an indication of the average percentage |
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127 | error. The number of trees examined is also printed out. |
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128 | <P> |
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129 | The constants |
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130 | available for modification at the beginning of the program are: |
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131 | "smoothings", which gives the number of passes through |
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132 | the algorithm which adjusts the lengths of the segments of the tree so |
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133 | as to minimize the sum of squares, "delta", which controls the size of |
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134 | improvement in sum of squares that is used to control the number of |
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135 | iterations improving branch lengths, |
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136 | and "epsilonf", |
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137 | which defines a small quantity needed in |
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138 | some of the calculations. There is no feature saving multiply trees |
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139 | tied for best, |
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140 | partly because we do not expect exact ties except in cases where the branch |
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141 | lengths make the nature of the tie obvious, as when a branch is of zero |
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142 | length. |
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143 | <P> |
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144 | The algorithm can be slow. As the number of species |
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145 | rises, so does the number of distances from each species to the others. The |
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146 | speed of this algorithm will thus rise as the fourth power of the |
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147 | number of species, rather than as the third power as do most of the |
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148 | others. Hence it is expected to get very slow as the number of species |
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149 | is made larger. |
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150 | <P> |
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151 | <HR> |
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152 | <P> |
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153 | <H3>TEST DATA SET</H3> |
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154 | <P> |
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155 | <TABLE><TR><TD BGCOLOR=white> |
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156 | <PRE> |
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157 | 7 |
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158 | Bovine 0.0000 1.6866 1.7198 1.6606 1.5243 1.6043 1.5905 |
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159 | Mouse 1.6866 0.0000 1.5232 1.4841 1.4465 1.4389 1.4629 |
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160 | Gibbon 1.7198 1.5232 0.0000 0.7115 0.5958 0.6179 0.5583 |
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161 | Orang 1.6606 1.4841 0.7115 0.0000 0.4631 0.5061 0.4710 |
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162 | Gorilla 1.5243 1.4465 0.5958 0.4631 0.0000 0.3484 0.3083 |
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163 | Chimp 1.6043 1.4389 0.6179 0.5061 0.3484 0.0000 0.2692 |
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164 | Human 1.5905 1.4629 0.5583 0.4710 0.3083 0.2692 0.0000 |
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165 | </PRE> |
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166 | </TD></TR></TABLE> |
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167 | <P> |
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168 | <HR> |
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169 | <P> |
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170 | <H3>OUTPUT FROM TEST DATA SET (with all numerical options on)</H3> |
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171 | <P> |
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172 | <TABLE><TR><TD BGCOLOR=white> |
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173 | <PRE> |
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174 | |
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175 | 7 Populations |
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176 | |
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177 | Fitch-Margoliash method version 3.6a3 |
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178 | |
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179 | __ __ 2 |
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180 | \ \ (Obs - Exp) |
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181 | Sum of squares = /_ /_ ------------ |
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182 | 2 |
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183 | i j Obs |
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184 | |
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185 | Negative branch lengths not allowed |
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186 | |
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187 | |
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188 | Name Distances |
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189 | ---- --------- |
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190 | |
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191 | Bovine 0.00000 1.68660 1.71980 1.66060 1.52430 1.60430 |
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192 | 1.59050 |
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193 | Mouse 1.68660 0.00000 1.52320 1.48410 1.44650 1.43890 |
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194 | 1.46290 |
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195 | Gibbon 1.71980 1.52320 0.00000 0.71150 0.59580 0.61790 |
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196 | 0.55830 |
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197 | Orang 1.66060 1.48410 0.71150 0.00000 0.46310 0.50610 |
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198 | 0.47100 |
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199 | Gorilla 1.52430 1.44650 0.59580 0.46310 0.00000 0.34840 |
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200 | 0.30830 |
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201 | Chimp 1.60430 1.43890 0.61790 0.50610 0.34840 0.00000 |
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202 | 0.26920 |
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203 | Human 1.59050 1.46290 0.55830 0.47100 0.30830 0.26920 |
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204 | 0.00000 |
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205 | |
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206 | |
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207 | +---------------------------------------------Mouse |
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208 | ! |
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209 | ! +------Human |
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210 | ! +--5 |
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211 | ! +-4 +--------Chimp |
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212 | ! ! ! |
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213 | ! +--3 +---------Gorilla |
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214 | ! ! ! |
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215 | 1------------------------2 +-----------------Orang |
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216 | ! ! |
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217 | ! +---------------------Gibbon |
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218 | ! |
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219 | +------------------------------------------------------Bovine |
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220 | |
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221 | |
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222 | remember: this is an unrooted tree! |
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223 | |
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224 | Sum of squares = 0.01375 |
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225 | |
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226 | Average percent standard deviation = 1.85418 |
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227 | |
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228 | Between And Length |
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229 | ------- --- ------ |
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230 | 1 Mouse 0.76985 |
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231 | 1 2 0.41983 |
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232 | 2 3 0.04986 |
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233 | 3 4 0.02121 |
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234 | 4 5 0.03695 |
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235 | 5 Human 0.11449 |
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236 | 5 Chimp 0.15471 |
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237 | 4 Gorilla 0.15680 |
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238 | 3 Orang 0.29209 |
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239 | 2 Gibbon 0.35537 |
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240 | 1 Bovine 0.91675 |
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241 | |
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242 | |
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243 | </PRE> |
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244 | </TD></TR></TABLE> |
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