# $Id: Compatible.pm 16123 2009-09-17 12:57:27Z cjfields $ # # BioPerl module for Bio::Tree::Compatible # # Please direct questions and support issues to # # Cared for by Gabriel Valiente # # Copyright Gabriel Valiente # # You may distribute this module under the same terms as Perl itself # POD documentation - main docs before the code =head1 NAME Bio::Tree::Compatible - Testing compatibility of phylogenetic trees with nested taxa. =head1 SYNOPSIS use Bio::Tree::Compatible; use Bio::TreeIO; my $input = Bio::TreeIO->new('-format' => 'newick', '-file' => 'input.tre'); my $t1 = $input->next_tree; my $t2 = $input->next_tree; my ($incompat, $ilabels, $inodes) = Bio::Tree::Compatible::is_compatible($t1,$t2); if ($incompat) { my %cluster1 = %{ Bio::Tree::Compatible::cluster_representation($t1) }; my %cluster2 = %{ Bio::Tree::Compatible::cluster_representation($t2) }; print "incompatible trees\n"; if (scalar(@$ilabels)) { foreach my $label (@$ilabels) { my $node1 = $t1->find_node(-id => $label); my $node2 = $t2->find_node(-id => $label); my @c1 = sort @{ $cluster1{$node1} }; my @c2 = sort @{ $cluster2{$node2} }; print "label $label"; print " cluster"; map { print " ",$_ } @c1; print " cluster"; map { print " ",$_ } @c2; print "\n"; } } if (scalar(@$inodes)) { while (@$inodes) { my $node1 = shift @$inodes; my $node2 = shift @$inodes; my @c1 = sort @{ $cluster1{$node1} }; my @c2 = sort @{ $cluster2{$node2} }; print "cluster"; map { print " ",$_ } @c1; print " properly intersects cluster"; map { print " ",$_ } @c2; print "\n"; } } } else { print "compatible trees\n"; } =head1 DESCRIPTION NB: This module has exclusively class methods that work on Bio::Tree::TreeI objects. An instance of Bio::Tree::Compatible cannot itself represent a tree, and so typically there is no need to create one. Bio::Tree::Compatible is a Perl tool for testing compatibility of phylogenetic trees with nested taxa represented as Bio::Tree::Tree objects. It is based on a recent characterization of ancestral compatibility of semi-labeled trees in terms of their cluster representations. A semi-labeled tree is a phylogenetic tree with some of its internal nodes labeled, and it can represent a classification tree as well as a phylogenetic tree with nested taxa, with labeled internal nodes corresponding to taxa at a higher level of aggregation or nesting than that of their descendents. Two semi-labeled trees are compatible if their topological restrictions to the common labels are such that for each node label, the smallest clusters containing it in each of the trees coincide and, furthermore, no cluster in one of the trees properly intersects a cluster of the other tree. Future extensions of Bio::Tree::Compatible include a Bio::Tree::Supertree module for combining compatible phylogenetic trees with nested taxa into a common supertree. =head1 FEEDBACK =head2 Mailing Lists User feedback is an integral part of the evolution of this and other Bioperl modules. Send your comments and suggestions preferably to the Bioperl mailing list. Your participation is much appreciated. bioperl-l@bioperl.org - General discussion http://bioperl.org/wiki/Mailing_lists - About the mailing lists =head2 Support Please direct usage questions or support issues to the mailing list: I rather than to the module maintainer directly. Many experienced and reponsive experts will be able look at the problem and quickly address it. Please include a thorough description of the problem with code and data examples if at all possible. =head2 Reporting Bugs Report bugs to the Bioperl bug tracking system to help us keep track of the bugs and their resolution. Bug reports can be submitted via the web: http://bugzilla.open-bio.org/ =head1 SEE ALSO =over =item * Philip Daniel and Charles Semple. Supertree Algorithms for Nested Taxa. In: Olaf R. P. Bininda-Emonds (ed.) Phylogenetic Supertrees: Combining Information to Reveal the Tree of Life, I, vol. 4, chap. 7, pp. 151-171. Kluwer (2004). =item * Charles Semple, Philip Daniel, Wim Hordijk, Roderic D. M. Page, and Mike Steel: Supertree Algorithms for Ancestral Divergence Dates and Nested Taxa. Bioinformatics B<20>(15), 2355-2360 (2004). =item * Merce Llabres, Jairo Rocha, Francesc Rossello, and Gabriel Valiente: On the Ancestral Compatibility of Two Phylogenetic Trees with Nested Taxa. J. Math. Biol. B<53>(3), 340-364 (2006). =back =head1 AUTHOR - Gabriel Valiente Email valiente@lsi.upc.edu =head1 APPENDIX The rest of the documentation details each of the object methods. =cut package Bio::Tree::Compatible; use strict; # Object preamble - inherits from Bio::Root::Root use Set::Scalar; use base qw(Bio::Root::Root); =head2 postorder_traversal Title : postorder_traversal Usage : my @nodes = @{ $tree->postorder_traversal } Function: Return list of nodes in postorder Returns : reference to array of Bio::Tree::Node Args : none For example, the postorder traversal of the tree C<(((A,B)C,D),(E,F,G));> is a reference to an array of nodes with internal_id 0 through 9, because the Newick standard representation for phylogenetic trees is based on a postorder traversal. +---A +---0 | | +---+---C +---4---2 | | | | | | | | +---B | | +---1 | | | | + +-------D 9 +-------3 | | | +-----E | +-----5 | | | | +-----+-----F +-----8-----6 | | +-----G +-----7 =cut sub postorder_traversal { my($self) = @_; my @stack; my @queue; push @stack, $self->get_root_node; while (@stack) { my $node = pop @stack; push @queue, $node; foreach my $child ($node->each_Descendent(-sortby => 'internal_id')) { push @stack, $child; } } my @postorder = reverse @queue; return \@postorder; } =head2 cluster_representation Title : cluster_representation Usage : my %cluster = %{ $tree->cluster_representation } Function: Compute the cluster representation of a tree Returns : reference to hash of array of string indexed by Bio::Tree::Node Args : none For example, the cluster representation of the tree C<(((A,B)C,D),(E,F,G));> is a reference to a hash associating an array of string (descendent labels) to each node, as follows: 0 --> [A] 1 --> [B] 2 --> [A,B,C] 3 --> [D] 4 --> [A,B,C,D] 5 --> [E] 6 --> [F] 7 --> [G] 8 --> [E,F,G] 9 --> [A,B,C,D,E,F,G] =cut sub cluster_representation { my ($tree) = @_; my %cluster; my @postorder = @{ postorder_traversal($tree) }; foreach my $node ( @postorder ) { my @labeled = map { $_->id } grep { $_->id } $node->get_Descendents; push @labeled, $node->id if $node->id; $cluster{$node} = \@labeled; } return \%cluster; } =head2 common_labels Title : common_labels Usage : my $labels = $tree1->common_labels($tree2); Function: Return set of common node labels Returns : Set::Scalar Args : Bio::Tree::Tree For example, the common labels of the tree C<(((A,B)C,D),(E,F,G));> and the tree C<((A,B)H,E,(J,(K)G)I);> are: C<[A,B,E,G]>. +---A +---A | | +---+---C +-------H | | | | | | | +---B | +---B | | | + +-------D +-----------E | | | +-----E | +-------J | | | | +-----+-----F +---I | | +-----G +---G---K =cut sub common_labels { my($self,$arg) = @_; my @labels1 = map { $_->id } grep { $_->id } $self->get_nodes; my $common = Set::Scalar->new( @labels1 ); my @labels2 = map { $_->id } grep { $_->id } $arg->get_nodes; my $temp = Set::Scalar->new( @labels2 ); return $common->intersection($temp); } =head2 topological_restriction Title : topological_restriction Usage : $tree->topological_restriction($labels) Function: Compute the topological restriction of a tree to a subset of node labels Returns : Bio::Tree::Tree Args : Set::Scalar For example, the topological restrictions of each of the trees C<(((A,B)C,D),(E,F,G));> and C<((A,B)H,E,(J,(K)G)I);> to the labels C<[A,B,E,G]> are as follows: +---A +---A | | +---+---+ +---+ | | | | | +---B | +---B + | | +---E +-------E | | | +-------+ +---+---G | +---G =cut sub topological_restriction { my ($tree, $labels) = @_; for my $node ( @{ postorder_traversal($tree) } ) { unless (ref($node)) { # skip $node if already removed my @cluster = map { $_->id } grep { $_->id } $node->get_Descendents; push @cluster, $node->id if $node->id; my $cluster = Set::Scalar->new(@cluster); if ($cluster->is_disjoint($labels)) { $tree->remove_Node($node); } else { if ($node->id and not $labels->has($node->id)) { $node->{'_id'} = undef; } } } } } =head2 is_compatible Title : is_compatible Usage : $tree1->is_compatible($tree2) Function: Test compatibility of two trees Returns : boolean Args : Bio::Tree::Tree For example, the topological restrictions of the trees C<(((A,B)C,D),(E,F,G));> and C<((A,B)H,E,(J,(K)G)I);> to their common labels, C<[A,B,E,G]>, are compatible. The respective cluster representations are as follows: [A] [A] [B] [B] [E] [E] [G] [G] [A,B] [A,B] [E,G] [A,B,E,G] [A,B,E,G] As a second example, the trees C<(A,B);> and C<((B)A);> are incompatible. Their respective cluster representations are as follows: [A] [B] [B] [A,B] [A,B] The reason is, the smallest cluster containing label C is C<[A]> in the first tree but C<[A,B]> in the second tree. +---A A---B | + | +---B As a second example, the trees C<(((B,A),C),D);> and C<((A,(D,B)),C);> are also incompatible. Their respective cluster representations are as follows: [A] [A] [B] [B] [C] [C] [D] [D] [A,B] [B,D] [A,B,C] [A,B,D] [A,B,C,D] [A,B,C,D] The reason is, cluster C<[A,B]> properly intersects cluster C<[B,D]>. There are further incompatibilities between these trees: C<[A,B,C]> properly intersects both C<[B,D]> and C<[A,B,D]>. +---B +-------A | | +---+ +---+ +---D | | | | | +---+ +---A | +---+ | | + | + +-------C | +---B | | +-----------D +-----------C =cut sub is_compatible { my ($tree1, $tree2) = @_; my $common = $tree1->Bio::Tree::Compatible::common_labels($tree2); $tree1->Bio::Tree::Compatible::topological_restriction($common); $tree2->Bio::Tree::Compatible::topological_restriction($common); my @postorder1 = @{ postorder_traversal($tree1) }; my @postorder2 = @{ postorder_traversal($tree2) }; my %cluster1 = %{ cluster_representation($tree1) }; my %cluster2 = %{ cluster_representation($tree2) }; my $incompat = 0; # false my @labels; foreach my $label ( $common->elements ) { my $node1 = $tree1->find_node(-id => $label); my @labels1 = @{ $cluster1{$node1} }; my $cluster1 = Set::Scalar->new(@labels1); my $node2 = $tree2->find_node(-id => $label); my @labels2 = @{ $cluster2{$node2} }; my $cluster2 = Set::Scalar->new(@labels2); unless ( $cluster1->is_equal($cluster2) ) { $incompat = 1; # true push @labels, $label; } } my @nodes; foreach my $node1 ( @postorder1 ) { my @labels1 = @{ $cluster1{$node1} }; my $cluster1 = Set::Scalar->new(@labels1); foreach my $node2 ( @postorder2 ) { my @labels2 = @{$cluster2{$node2} }; my $cluster2 = Set::Scalar->new(@labels2); if ($cluster1->is_properly_intersecting($cluster2)) { $incompat = 1; # true push @nodes, $node1, $node2; } } } return ($incompat, \@labels, \@nodes); } 1;