Bio
PhyloNetwork
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Summary
Bio::PhyloNetwork - Module to compute with Phylogenetic Networks
Package variables
Privates (from "my" definitions)
$_eN_index;
Included modules
Algorithm::Munkres
Array::Compare
Graph::Directed
IO::String
Inherit
Synopsis
use Bio::PhyloNetwork;
# Create a PhyloNetwork object from a eNewick string
my $net1=Bio::PhyloNetwork->new(
-eNewick=>'t0:((H1,(H2,l2)),H2); H1:((H3,l1)); H2:((H3,(l3,H1))); H3:(l4);'
);
# Print all available data
print $net1;
# Rebuild $net1 from its mu_data
my %mudata=$net1->mudata();
my $net2=Bio::PhyloNetwork->new(-mudata=>\%mudata,-numleaves=>4);
print $net2;
print "d=".$net1->mu_distance($net2)."\n";
# Get another one and compute distance
my $net3=Bio::PhyloNetwork->new(
-eNewick=>'(l2,((l1,(H1,l4)),H1))r; (l3)H1;'
);
print "d=".$net1->mu_distance($net3)."\n";
# ...and find an optimal alignment w.r.t. the Manhattan distance (default)
my ($weight,%alignment)=$net1->optimal_alignment($net3);
print "weight:$weight\n";
foreach my $node1 (keys %alignment) {
print "$node1 => ".$alignment{$node1}."\n";
}
# ...or the Hamming distance
my ($weightH,%alignmentH)=$net1->optimal_alignment($net3,-metric=>'Hamming');
print "weight:$weightH\n";
foreach my $node1 (keys %alignmentH) {
print "$node1 => ".$alignmentH{$node1}."\n";
}
# Test for time consistency of $net1
if ($net1->is_time_consistent) {
print "net1 is time consistent\n"
}
else {
print "net1 is not time consistent\n"
}
# create a network from the list of edges
my $net4=Bio::PhyloNetwork->new(-edges=>
[qw(r s r t s u s c t c t v u b u l3 u b v b v l4 b l2 c l1)]);
# Test for time consistency of $net3
if ($net4->is_time_consistent) {
print "net4 is time consistent\n"
}
else {
print "net4 is not time consistent\n"
}
# And print all information on net4
print $net4;
# Compute some tripartitions
my %triparts=$net1->tripartitions();
# Now these are stored
print $net1;
# And can compute the tripartition error
print "dtr=".$net1->tripartition_error($net3)."\n";
Description
This is a module to work with phylogenetic networks. Phylogenetic networks
have been studied over the last years as a richer model of the evolutionary
history of sets of organisms than phylogenetic trees, because they take not
only mutation events but also recombination and horizontal gene transfer
events into account.
The natural model for describing the evolutionary
history of a set of sequences under recombination events is a DAG, hence
this package relies on the package Graph::Directed to represent the
underlying graph of a phylogenetic network. We refer the reader to [CRV1,CRV2]
for formal definitions related to phylogenetic networks.
With this package, phylogenetic networks can be given by its eNewick
string. This description appeared in other packages related to
phylogenetic networks (see [PhyloNet] and [NetGen]); in fact, these two
packages use different descriptions. The Bio::PhyloNetwork
package allows both of them, but uses the second one in its output.
The first approach [PhyloNet] goes as follows: For each hybrid node H, say with
parents u_1,u_2,...,u_k and children v_1,v_2,...v_l: split H in k+1 different
nodes; let each of the first k copies be a child of one of the u_1,...,u_k
(one for each) and have no children (hence we will have k extra leaves);
as for the last copy, let it have no parents and have v_1,...,v_l be its
children. This way we get a forest; each of the trees will be rooted at either
one root of the phylogenetic network or a hybrid node of it; the set of leaves
(of the whole forest) will be the set of leaves of the original network
together with the set of hybrid nodes (each of them repeated as many times
as its in-degree). Then, the eNewick representation of the phylogenetic network
will be the Newick representation of all the trees in the obtained forest,
each of them with its root labeled.
The second approach [NetGen] goes as follows: For each hybrid node H, say with
parents u_1,u_2,...,u_k and children v_1,v_2,...v_l: split H in k different
nodes; let the first copy be a child of u_1 and have all v_1,v_2,...v_l as
its children; let the other copies be child of u_2,...,u_k (one for each)
and have no children. This way, we get a tree whose set of leaves is the
set of leaves of the original network together with the set of hybrid nodes
(possibly repeated). Then the Newick string of the obtained tree (note that
some internal nodes will be labeled and some leaves will be repeated) is
the eNewick string of the phylogenetic network.
For example, consider the network depicted below:
r
/ \
/ \
U V
/ \ / \
1 \ / 3
H
|
2
If the first approach is taken, we get the forest:
r
/ \
/ \
U V
/ \ / \
1 H H 3
|
H
|
2
Hence, the eNewick string is '((1,H),(H,3))r; (2)H;'.
As for the second one, one gets the tree:
r
/ \
/ \
U V
/ \ / \
1 H | 3
H
|
2
Hence, the eNewick string is '((1,H),((2)H,3))r;'.
Note: when rooting a tree, this package allows the notations
'(subtree,subtree,...)root' as well as 'root:(subtree,subtree,...)', but
the first one is used when writing eNewick strings.
Tree-child (TC) phylogenetic networks are a special class of phylogenetic
networks for which a distance, called mu-distance, is defined [CRV2]
based on certain data (mu-data) associated to every node.
Moreover, this distance extends the
Robinson-Foulds on phylogenetic trees. This package allows testing for a
phylogenetic network if it is TC and computes mu-distances between networks
over the same set of leaves.
Moreover, the mu-data allows one to define the optimal
(in some precise sense) alignment between networks
over the same set of leaves. This package also computes this optimal alignment.
Although tripartitions (see [CRV1] and the references therein) do not allow
to define distances, this package outputs tripartitions and computes a weak
form of the tripartition error.
Another useful property of Phylogenetic Networks that appears in the literature
is that of time-consistency or real-time hybrids [BSS]. Roughly speaking, a
network admits a temporal representation if it can be drawn in such a way
that tree arcs (those whose end is a tree node) are inclined downwards, while
hybridization arcs (those whose end is a hybrid node) are horizontal.
This package checks for time-consistency and, if so, a temporal representation
is provided.
Methods
Methods description
Title : new Usage : my $obj = new Bio::PhyloNetwork(); Function: Creates a new Bio::PhyloNetwork object Returns : Bio::PhyloNetwork Args : none OR -eNewick => string OR -graph => Graph::Directed object OR -edges => reference to an array OR -tree => Bio::Tree::Tree object OR -mudata => reference to a hash, -leaves => reference to an array OR -mudata => reference to a hash, -numleaves => integer
Returns a Bio::PhyloNetwork object, created according to the data given:
new() creates an empty network.
new(-eNewick => $str) creates the network whose Extended Newick representation (see description above) is the string $str.
new(-graph => $graph) creates the network with underlying graph given by the Graph::Directed object $graph
new(-tree => $tree) creates a network as a copy of the Bio::Tree::Tree object in $tree
new(-mudata =E<gt> \%mudata, -leaves => \@leaves) creates the network by reconstructing it from its mu-data stored in \%mudata and with set of leaves in \@leaves.
new(-mudata =E<gt> \%mudata, -numleaves => $numleaves) creates the network by reconstructing it from its mu-data stored in \%mudata and with set of leaves in ("l1".."l$numleaves").
|
Title : is_leaf Usage : my $b=$net->is_leaf($u) Function: tests if $u is a leaf in $net Returns : boolean Args : scalar |
Title : is_root Usage : my $b=$net->is_root($u) Function: tests if $u is the root of $net Returns : boolean Args : scalar |
Title : is_tree_node Usage : my $b=$net->is_tree_node($u) Function: tests if $u is a tree node in $net Returns : boolean Args : scalar |
Title : is_hybrid_node Usage : my $b=$net->is_hybrid_node($u) Function: tests if $u is a hybrid node in $net Returns : boolean Args : scalar |
Title : is_tree_child Usage : my $b=$net->is_tree_child() Function: tests if $net is a Tree-Child phylogenetic network Returns : boolean Args : Bio::PhyloNetwork |
Title : nodes Usage : my @nodes=$net->nodes() Function: returns the set of nodes of $net Returns : array Args : none |
Title : leaves Usage : my @leaves=$net->leaves() Function: returns the set of leaves of $net Returns : array Args : none |
Title : roots Usage : my @roots=$net->roots() Function: returns the set of roots of $net Returns : array Args : none |
Title : internal_nodes Usage : my @internal_nodes=$net->internal_nodes() Function: returns the set of internal nodes of $net Returns : array Args : none |
Title : tree_nodes Usage : my @tree_nodes=$net->tree_nodes() Function: returns the set of tree nodes of $net Returns : array Args : none |
Title : hybrid_nodes Usage : my @hybrid_nodes=$net->hybrid_nodes() Function: returns the set of hybrid nodes of $net Returns : array Args : none |
Title : graph Usage : my $graph=$net->graph() Function: returns the underlying graph of $net Returns : Graph::Directed Args : none |
Title : edges Usage : my @edges=$net->edges() Function: returns the set of edges of $net Returns : array Args : none
Each element in the array is an anonimous array whose first element is the head of the edge and the second one is the tail. |
Title : tree_edges Usage : my @tree_edges=$net->tree_edges() Function: returns the set of tree edges of $net (those whose tail is a tree node) Returns : array Args : none |
Title : hybrid_edges Usage : my @hybrid_edges=$net->hybrid_edges() Function: returns the set of hybrid edges of $net (those whose tail is a hybrid node) Returns : array Args : none |
Title : explode Usage : my @trees=$net->explode() Function: returns the representation of $net by a set of Bio::Tree:Tree objects Returns : array Args : none |
Title : mudata Usage : my %mudata=$net->mudata() Function: returns the representation of $net by its mu-data Returns : hash Args : none
$net->mudata() returns a hash with keys the nodes of $net and each value is a muVector object holding its mu-vector. |
Title : heights Usage : my %heights=$net->heights() Function: returns the heights of the nodes of $net Returns : hash Args : none
$net->heights() returns a hash with keys the nodes of $net and each value is its height. |
Title : mu_distance Usage : my $dist=$net1->mu_distance($net2) Function: Computes the mu-distance between the networks $net1 and $net2 on the same set of leaves Returns : scalar Args : Bio::PhyloNetwork |
Title : mu_distance_generalized Usage : my $dist=$net1->mu_distance($net2) Function: Computes the mu-distance between the topological restrictions of networks $net1 and $net2 on its common set of leaves Returns : scalar Args : Bio::PhyloNetwork |
Title : tripartitions Usage : my %tripartitions=$net->tripartitions() Function: returns the set of tripartitions of $net Returns : hash Args : none
$net->tripartitions() returns a hash with keys the nodes of $net and each value is a string representing the tripartition of the leaves induced by the node. A string "BCA..." associated with a node u (e.g.) means, the first leaf is in the set B(u), the second one in C(u), the third one in A(u), and so on. |
Title : is_time_consistent Usage : my $b=$net->is_time_consistent() Function: tests if $net is (strong) time-consistent Returns : boolean Args : none |
Title : temporal_representation Usage : my %time=$net->temporal_representation() Function: returns a hash containing a temporal representation of $net, or 0 if $net is not time-consistent Returns : hash Args : none |
Title : contract_elementary Usage : my ($contracted,$blocks)=$net->contract_elementary(); Function: Returns the network $contracted, obtained by contracting elementary paths of $net into edges. The reference $blocks points to a hash where, for each node of $contracted, gives the corresponding nodes of $net that have been deleted. Returns : Bio::PhyloNetwork,reference to hash Args : none |
Title : optimal_alignment Usage : my ($weight,$alignment,$wgts)=$net->optimal_alignment($net2) Function: returns the total weight of an optimal alignment, the alignment itself, and partial weights between the networks $net1 and $net2 on the same set of leaves. An optional argument allows one to use the Manhattan (default) or the Hamming distance between mu-vectors. Returns : scalar,reference to hash,reference to hash Args : Bio::PhyloNetwork, -metric => string (optional)
Supported strings for the -metric parameter are 'Manhattan' or 'Hamming'. |
Title : optimal_alignment_generalized Usage : my ($weight,%alignment)=$net->optimal_alignment_generalized($net2) Function: returns the wieght of an optimal alignment, and the alignment itself, between the topological restriction of the networks $net1 and $net2 on the set of common leaves. An optional argument allows one to use the Manhattan (default) or the Hamming distance between mu-vectors. Returns : scalar,hash Args : Bio::PhyloNetwork, -metric => string (optional)
Supported strings for the -metric parameter are 'Manhattan' or 'Hamming'. |
Title : topological_restriction Usage : my ($netr1,$netr2)=$net1->topological_restriction($net2) Function: returns the topological restriction of $net1 and $net2 on its common set of leaves Returns : Bio::PhyloNetwork, Bio::PhyloNetwork Args : Bio::PhyloNetwork |
Title : eNewick Usage : my $str=$net->eNewick() Function: returns the eNewick representation of $net without labeling internal tree nodes Returns : string Args : none |
Title : eNewick_full Usage : my $str=$net->eNewick_full() Function: returns the eNewick representation of $net labeling internal tree nodes Returns : string Args : none |
Title : display Usage : my $str=$net->display() Function: returns a string containing all the available information on $net Returns : string Args : none |
Methods code
sub new
{ my ($pkg,@args)=@_;
my $self=$pkg->SUPER::new(@args);
my ($eNewick,$edgesR,$leavesR,$numleaves,$graph,$tree,$mudataR)=
$self->_rearrange([qw(ENEWICK
EDGES
LEAVES
NUMLEAVES
GRAPH
TREE
MUDATA)],@args);
bless($self,$pkg);
$self->build_from_eNewick($eNewick) if defined $eNewick;
$self->build_from_edges(@$edgesR) if defined $edgesR;
$self->build_from_graph($graph) if defined $graph;
$self->build_from_tree($tree) if defined $tree;
if ((! defined $leavesR) && (defined $numleaves)) {
my @leaves=map {"l$_"} (1..$numleaves);
$leavesR=\@leaves;
}
$self->build_from_mudata($mudataR,$leavesR)
if ((defined $mudataR) && (defined $leavesR));
return $self;
}
} |
sub build_from_edges
{ my ($self,@edges)=@_;
my $graph=Graph::Directed->new();
$graph->add_edges(@edges);
$self->{graph}=$graph;
$self->recompute();
my $labels={};
foreach my $node ($self->nodes()) {
$labels->{$node}=$node;
}
$self->{labels}=$labels;} |
sub build_from_graph
{ my ($self,$graph)=@_;
my $graphcp=$graph->copy();
$self->{graph}=$graphcp;
$self->recompute();
my $labels={};
foreach my $node ($self->nodes()) {
$labels->{$node}=$node;
}
$self->{labels}=$labels;
}
my $_eN_index;} |
sub build_from_eNewick
{ my ($self,$string)=@_;
$_eN_index=0;
my $graph=Graph::Directed->new();
my $labels={};
my @blocks=split(/; */,$string);
foreach my $block (@blocks) {
my ($rt,$str)=get_root_and_subtree($block);
my ($rtlbl,$rttype,$rtid,$rtlng)=get_label_type_id_length($rt);
process_block($graph,$labels,$block,$rtid);
$labels->{$rtid}=$rtlbl.'';
}
$self->{graph}=$graph;
$self->{labels}=$labels;
$self->recompute();} |
sub process_block
{ my ($graph,$labels,$block,$rtid)=@_;
my ($rt,$str)=get_root_and_subtree($block);
my @substrs=my_split($str);
foreach my $substr (@substrs) {
my ($subrt,$subblock)=get_root_and_subtree($substr);
my ($subrtlbl,$subrttype,$subrtid,$subrtlng)=
get_label_type_id_length($subrt);
if (! $subrtlng eq '') {
$graph->add_weighted_edges($rtid,$subrtid,$subrtlng);
}
else {
$graph->add_edges($rtid,$subrtid);
}
if (! $subrttype eq '') {
$graph->set_edge_attribute($rtid,$subrtid,'type',$subrttype);
}
$subrtlbl.='';
if ((! defined $labels->{$subrtid})||($labels->{$subrtid} eq '')){
$labels->{$subrtid}=$subrtlbl;
} elsif (( $labels->{$subrtid} ne $subrtlbl )&&($subrtlbl ne '')) {
die("Different labels for the same node (".$labels->{$subrtid}." and $subrtlbl)");
}
if ($subblock ne "") {
process_block($graph,$labels,$subblock,$subrtid);
}
}} |
sub get_root_and_subtree
{ my ($block)=@_;
my ($rt,$str)=("","");
($rt,$str)=split(/=/,$block);
if ($rt eq $block) {
my $pos=length($rt)-1;
while ((substr($rt,$pos,1) ne ")") && ($pos >=0)) {
$pos--;
}
$rt=substr($block,$pos+1,length($block)-$pos);
$str=substr($block,0,$pos+1);
}
$rt=trim($rt);
$str=trim($str);
return ($rt,$str);} |
sub get_label_type_id_length
{ my ($string) = @_;
$string.='';
if (index($string,'#')==-1) {
my ($label,$length)=split(':',$string);
$label.='';
my $id;
if ((! defined $label) || ($label eq '')) {
$_eN_index++;
$id="T$_eN_index";
} else {
$id=$label;
}
return ($label,'',$id,$length);
}
else {
my ($label,$string2)=split('#',$string);
my ($typeid,$length)=split(':',$string2);
my $type=$typeid;
$type =~ s/\d//g;
my $id=$typeid;
$id =~ s/\D//g;
return ($label,$type,'#'.$id,$length);
} } |
sub trim
{
my ($string) = @_;
$string =~ s/^\s+//;
$string =~ s/\s+$//;
return $string; } |
sub my_split
{ my ( $string ) = @_;
my $temp="";
my @substrings;
my $level=1;
for my $i ( 1 .. length( $string ) ) {
my $char=substr($string,$i,1);
if ($char eq "(") {
$level++;
}
if ($char eq ")") {
if ($level==1) {
push @substrings, $temp;
$temp="";
}
$level--;
}
if (($char eq ",") && ($level==1)) {
push @substrings, $temp;
$temp="";
$char="";
}
$temp = $temp.$char;
}
return @substrings;} |
sub build_from_mudata
{ my ($self,$mus,$leavesR)=@_;
my $graph=Graph::Directed->new();
my @nodes=keys %{$mus};
my @leaves=@{$leavesR};
my %seen;
my @internal;
@seen{@leaves} = ();
foreach my $node (@nodes) {
push(@internal, $node) unless exists $seen{$node};
}
@internal=sort {$mus->{$b} <=> $mus->{$a} } @internal;
@nodes=(@internal,@leaves);
my $numnodes=@nodes;
for (my $i=0;$i<$numnodes;$i++) {
my $mu=$mus->{$nodes[$i]};
my $j=$i+1;
while ($mu->is_positive() && $j<$numnodes) {
if ($mu->geq_poset($mus->{$nodes[$j]})) {
$graph->add_edges(($nodes[$i],$nodes[$j]));
$mu = $mu - $mus->{$nodes[$j]};
}
$j++;
}
}
$self->build_from_graph($graph);
}
} |
sub build_from_tree
{ my ($self,$tree)=@_;
my $str;
my $io=IO::String->new($str);
my $treeio=Bio::TreeIO->new(-format => 'newick', -fh => $io);
$treeio->write_tree($tree);
$self->build_from_eNewick($str); } |
sub recompute
{ my ($self)=@_;
$self->throw("Graph is not DAG:".$self->{graph})
unless $self->{graph}->is_dag();
my @leaves=$self->{graph}->successorless_vertices();
@leaves=sort @leaves;
my $numleaves=@leaves;
my @roots=$self->{graph}->predecessorless_vertices();
my $numroots=@roots;
my @nodes=$self->{graph}->vertices();
@nodes=sort @nodes;
my $numnodes=@nodes;
foreach my $node (@nodes) {
if (! defined $self->{labels}->{$node}) {
$self->{labels}->{$node}='';
}
}
$self->{leaves}=\@leaves;
$self->{numleaves}=$numleaves;
$self->{roots}=\@roots;
$self->{numroots}=$numroots;
$self->{nodes}=\@nodes;
$self->{numnodes}=$numnodes;
$self->{mudata}={};
$self->{h}={};
$self->compute_height();
$self->compute_mu();
return $self;
}
} |
sub is_attackable
{ my ($self,$u1,$v1,$u2,$v2)=@_;
if ( $self->is_hybrid_node($v1) ||
$self->is_hybrid_node($v2) ||
$self->graph->is_reachable($v2,$u1) ||
(($u1 eq $u2)&&($v1 eq $v2)) ||
(! scalar grep {($_ ne $v2) && ($self->is_tree_node($_))}
$self->graph->successors($u2)))
{
return 0;
}
return 1;} |
sub do_attack
{ my ($self,$u1,$v1,$u2,$v2,$lbl)=@_;
my $graph=$self->{graph};
$graph->delete_edge($u1,$v1);
$graph->delete_edge($u2,$v2);
$graph->add_edge($u1,"T$lbl");
$graph->add_edge("T$lbl",$v1);
$graph->add_edge($u2,"#H$lbl");
$graph->add_edge("#H$lbl",$v2);
$graph->add_edge("T$lbl","#H$lbl");
$self->build_from_graph($graph);
}
} |
sub compute_mu
{ my ($self)=@_;
my $graph=$self->{graph};
my $mudata=$self->{mudata};
my @leaves=@{$self->{leaves}};
my $numleaves=$self->{numleaves};
for (my $i=0;$i<$numleaves;$i++) {
my $vec=Bio::PhyloNetwork::muVector->new($numleaves);
$vec->[$i]=1;
$mudata->{$leaves[$i]}=$vec;
}
my $h=1;
while (my @nodes=grep {$self->{h}->{$_} == $h} @{$self->{nodes}} )
{
foreach my $u (@nodes) {
my $vec=Bio::PhyloNetwork::muVector->new($numleaves);
foreach my $son ($graph->successors($u)) {
$vec+=$mudata->{$son};
}
$mudata->{$u}=$vec;
}
$h++;
}} |
sub compute_height
{ my ($self)=@_;
my $graph=$self->{graph};
my @leaves=@{$self->{leaves}};
foreach my $leaf (@leaves) {
$self->{h}->{$leaf}=0;
}
my $h=0;
while (my @nodes=grep {(defined $self->{h}->{$_})&&($self->{h}->{$_} == $h)}
@{$self->{nodes}} )
{
foreach my $node (@nodes) {
foreach my $parent ($graph->predecessors($node)) {
$self->{h}->{$parent}=$h+1;
}
}
$h++;
}
}
} |
sub is_leaf
{ my ($self,$node)=@_;
if ($self->{graph}->out_degree($node) == 0) {return 1;}
return 0;} |
sub is_root
{ my ($self,$node)=@_;
if ($self->{graph}->in_degree($node) == 0) {return 1;}
return 0;} |
sub is_tree_node
{ my ($self,$node)=@_;
if ($self->{graph}->in_degree($node) <= 1) {return 1;}
return 0;} |
sub is_hybrid_node
{ my ($self,$node)=@_;
if ($self->{graph}->in_degree($node) > 1) {return 1;}
return 0;} |
sub has_tree_child
{ my $g=shift(@_);
my $node=shift(@_);
my @Sons=$g->successors($node);
foreach my $son (@Sons) {
if ($g->in_degree($son)==1) {
return 1;
}
}
return 0; } |
sub is_tree_child
{ my ($self)=@_;
if (defined $self->{is_tree_child}) {
return $self->{is_tree_child};
}
$self->{is_tree_child}=0;
my $graph=$self->{graph};
foreach my $node (@{$self->{nodes}}) {
return 0 unless ($graph->out_degree($node)==0 ||
has_tree_child($graph,$node));
}
$self->{is_tree_child}=1;
return 1;
}
} |
sub nodes
{ my ($self)=@_;
return @{$self->{nodes}};} |
sub leaves
{ my ($self)=@_;
return @{$self->{leaves}};} |
sub roots
{ my ($self)=@_;
return @{$self->{roots}};} |
sub internal_nodes
{ my ($self)=@_;
return grep {! $self->is_leaf($_)} $self->nodes();} |
sub tree_nodes
{ my ($self)=@_;
return grep {$self->is_tree_node($_)} $self->nodes();} |
sub hybrid_nodes
{ my ($self)=@_;
return grep {$self->is_hybrid_node($_)} $self->nodes();} |
sub graph
{ my ($self)=@_;
return $self->{graph};} |
sub edges
{ my ($self)=@_;
return $self->{graph}->edges();} |
sub tree_edges
{ my ($self)=@_;
return grep {$self->is_tree_node($_->[1])} $self->edges();} |
sub hybrid_edges
{ my ($self)=@_;
return grep {$self->is_hybrid_node($_->[1])} $self->edges();} |
sub explode
{ my ($self)=@_;
my @trees;
$self->explode_rec(\@trees);
return @trees; } |
sub explode_rec
{ my ($self,$trees)=@_;
my @h = $self->hybrid_nodes;
if (scalar @h) {
my $v = shift @h;
for my $u ($self->{graph}->predecessors($v)) {
$self->{graph}->delete_edge($u,$v);
$self->explode_rec($trees);
$self->{graph}->add_edge($u,$v);
}
} else {
my $io = IO::String->new($self->eNewick);
my $treeio = Bio::TreeIO->new(-format => 'newick', -fh => $io);
my $tree = $treeio->next_tree;
$tree->contract_linear_paths;
push @{$trees}, $tree;
}} |
sub mudata
{ my ($self)=@_;
return %{$self->{mudata}};} |
sub mudata_node
{ my ($self,$u)=@_;
return $self->{mudata}{$u};} |
sub heights
{ my ($self)=@_;
return %{$self->{h}};} |
sub height_node
{ my ($self,$u)=@_;
return $self->{h}{$u};} |
sub mu_distance
{ my ($net1,$net2)=@_;
my @nodes1=$net1->nodes;
my @nodes2=$net2->nodes;
my $comp = Array::Compare->new;
$net1->throw("Cannot compare phylogenetic networks on different set of leaves")
unless $comp->compare($net1->{leaves},$net2->{leaves});
$net1->warn("Not a tree-child phylogenetic network")
unless $net1->is_tree_child();
$net2->warn("Not a tree-child phylogenetic network")
unless $net2->is_tree_child();
my @leaves=@{$net1->{leaves}};
my %matched1;
my %matched2;
OUTER: foreach my $node1 (@nodes1) {
foreach my $node2 (@nodes2) {
if (
(! exists $matched1{$node1}) && (! exists $matched2{$node2}) &&
($net1->{mudata}{$node1} == $net2->{mudata}{$node2})
) {
$matched1{$node1}=$node2;
$matched2{$node2}=$node1;
next OUTER;
}
}
}
return (scalar @nodes1)+(scalar @nodes2)-2*(scalar keys %matched1);} |
sub mu_distance_generalized
{ my ($net1,$net2)=@_;
my ($netr1,$netr2)=$net1->topological_restriction($net2);
return $netr1->mu_distance($netr2);
}
} |
sub mudata_string_node
{ my ($self,$u)=@_;
return $self->{mudata}->{$u}->display();} |
sub mudata_string
{ my ($self)=@_;
return $self->{mudata_string} if defined $self->{mudata_string};
my @internal=$self->internal_nodes;
my $mus=$self->{mudata};
@internal=sort {$mus->{$b} <=> $mus->{$a} } @internal;
my $str="";
foreach my $node (@internal) {
$str=$str.$self->mudata_string_node($node);
}
$self->{mudata_string}=$str;
return $str;} |
sub is_mu_isomorphic
{ my ($net1,$net2)=@_;
return ($net1->mudata_string() eq $net2->mudata_string());
}
} |
sub compute_tripartition_node
{ my ($self,$u)=@_;
$self->warn("Cannot compute tripartitions on unrooted networks. Will assume one at random")
unless ($self->{numroots} == 1);
my $root=$self->{roots}->[0];
my $graph=$self->{graph};
my $graphPruned=$graph->copy();
$graphPruned->delete_vertex($u);
my $tripartition="";
foreach my $leaf (@{$self->{leaves}}) {
my $type;
if ($graph->is_reachable($u,$leaf)) {
if ($graphPruned->is_reachable($root,$leaf)) {$type="B";}
else {$type="A";}
}
else {$type="C";}
$tripartition .= $type;
}
$self->{tripartitions}->{$u}=$tripartition;} |
sub compute_tripartitions
{ my ($self)=@_;
foreach my $node (@{$self->{nodes}}) {
$self->compute_tripartition_node($node);
}} |
sub tripartitions
{ my ($self)=@_;
$self->compute_tripartitions() unless defined $self->{tripartitions};
return %{$self->{tripartitions}};
}
} |
sub tripartition_error
{ my ($net1,$net2)=@_;
my $comp = Array::Compare->new;
$net1->throw("Cannot compare phylogenetic networks on different set of leaves")
unless $comp->compare($net1->{leaves},$net2->{leaves});
$net1->warn("Not a tree-child phylogenetic network")
unless $net1->is_tree_child();
$net2->warn("Not a tree-child phylogenetic network")
unless $net2->is_tree_child();
$net1->warn("Not a time-consistent network")
unless $net1->is_time_consistent();
$net2->warn("Not a time-consistent network")
unless $net2->is_time_consistent();
$net1->compute_tripartitions() unless defined $net1->{tripartitions};
$net2->compute_tripartitions() unless defined $net2->{tripartitions};
my @edges1=$net1->{graph}->edges();
my @edges2=$net2->{graph}->edges();
my ($FN,$FP)=(0,0);
foreach my $edge1 (@edges1) {
my $matched=0;
foreach my $edge2 (@edges2) {
if ($net1->{tripartitions}->{$edge1->[1]} eq
$net2->{tripartitions}->{$edge2->[1]}) {
$matched=1;
last;
}
}
if (! $matched) {$FN++;}
}
foreach my $edge2 (@edges2) {
my $matched=0;
foreach my $edge1 (@edges1) {
if ($net1->{tripartitions}->{$edge1->[1]} eq
$net2->{tripartitions}->{$edge2->[1]}) {
$matched=1;
last;
}
}
if (! $matched) {$FP++;}
}
return ($FN/(scalar @edges1)+$FP/(scalar @edges2))/2; }
} |
sub is_time_consistent
{ my ($self)=@_;
$self->compute_temporal_representation()
unless exists $self->{has_temporal_representation};
return $self->{has_temporal_representation};} |
sub temporal_representation
{ my ($self)=@_;
if ($self->is_time_consistent) {
return %{$self->{temporal_representation}};
}
return 0;} |
sub compute_temporal_representation
{ my ($self)=@_;
my $quotient=Graph::Directed->new();
my $classes=find_classes($self);
my %repr;
map {$repr{$_}=$classes->{$_}[0]} $self->nodes();
foreach my $e ($self->tree_edges()) {
$quotient->add_edge($repr{$e->[0]},$repr{$e->[1]});
}
my %temp;
my $depth=0;
while ($quotient->vertices()) {
if (my @svs=$quotient->predecessorless_vertices()) {
foreach my $sv (@svs) {
$temp{$sv}=$depth;
}
$quotient->delete_vertices(@svs);
} else {
return 0;
}
$depth++;
}
foreach my $node (@{$self->{nodes}}) {
$temp{$node}=$temp{$repr{$node}}
}
$self->{temporal_representation}=\%temp;
$self->{has_temporal_representation}=1;} |
sub find_classes
{ my ($self)=@_;
my $classes={};
map {$classes->{$_}=[$_]} $self->nodes();
foreach my $e ($self->hybrid_edges()) {
$classes=join_classes($classes,$e->[0],$e->[1]);
}
return $classes;} |
sub join_classes
{ my ($classes,$u,$v)=@_;
my @clu=@{$classes->{$u}};
my @clv=@{$classes->{$v}};
my @cljoin=(@clu,@clv);
map {$classes->{$_}=\@cljoin} @cljoin;
return $classes;
}
} |
sub contract_elementary
{ my ($self)=@_;
my $contracted=$self->graph->copy();
my @nodes=$self->nodes();
my $mus=$self->{mudata};
my $hs=$self->{h};
my %blocks;
foreach my $u (@nodes) {
$blocks{$u}=[$u];
}
my @elementary=grep { $contracted->out_degree($_) == 1} $self->tree_nodes();
@elementary=sort {$mus->{$b} <=> $mus->{$a} ||
$hs->{$b} <=> $hs->{$a}} @elementary;
foreach my $elem (@elementary) {
my @children=$contracted->successors($elem);
my $child=$children[0];
if ($contracted->in_degree($elem) == 1) {
my @parents=$contracted->predecessors($elem);
my $parent=$parents[0];
$contracted->add_edge($parent,$child);
}
$contracted->delete_vertex($elem);
my @blch=@{$blocks{$child}};
my @blem=@{$blocks{$elem}};
$blocks{$child}=[@blem,@blch];
delete $blocks{$elem};
}
my $contr=Bio::PhyloNetwork->new(-graph => $contracted);
return $contr,\%blocks;} |
sub optimal_alignment
{ my ($net1,$net2,%params)=@_;
my ($net1cont,$blocks1)=contract_elementary($net1);
my ($net2cont,$blocks2)=contract_elementary($net2);
my ($wc,$alignc,$weightc)=
optimal_alignment_noelementary($net1cont,$net2cont,%params);
my %alignment=();
my $totalweigth=0;
my %weigths=();
foreach my $u1 (keys %$alignc) {
my $u2=$alignc->{$u1};
my @block1=@{$blocks1->{$u1}};
my @block2=@{$blocks2->{$u2}};
while (@block1 && @block2) {
my $u1dc=pop @block1;
my $u2dc=pop @block2;
$alignment{$u1dc}=$u2dc;
$weigths{$u1dc}=$weightc->{$u1};
$totalweigth+=$weigths{$u1dc};
}
}
return $totalweigth,\%alignment,\%weigths;} |
sub optimal_alignment_noelementary
{ my ($net1,$net2,%params)=@_;
my $comp = Array::Compare->new;
$net1->throw("Cannot align phylogenetic networks on different set of leaves")
unless $comp->compare($net1->{leaves},$net2->{leaves});
my $distance;
if ((defined $params{-metric})and ($params{-metric} eq 'Hamming')) {
$distance='Hamming';
} else {
$distance='Manhattan';
}
my $numleaves=$net1->{numleaves};
my @nodes1=$net1->internal_nodes();
my @nodes2=$net2->internal_nodes();
my $numnodes1=@nodes1;
my $numnodes2=@nodes2;
my @matrix=();
for (my $i=0;$i<$numnodes1;$i++) {
my @row=();
for (my $j=0;$j<$numnodes2;$j++) {
push @row,weight($net1,$nodes1[$i],$net2,$nodes2[$j],$distance);
}
push @matrix,\@row;
}
my @alignment=();
assign(\@matrix,\@alignment);
my %alignmenthash;
my %weighthash;
my $totalw=0;
foreach my $leaf (@{$net1->{leaves}}) {
$alignmenthash{$leaf}=$leaf;
$weighthash{$leaf}=0;
}
for (my $i=0;$i<$numnodes1;$i++) {
if (defined $nodes2[$alignment[$i]]) {
$alignmenthash{$nodes1[$i]}=$nodes2[$alignment[$i]];
$weighthash{$nodes1[$i]}=$matrix[$i][$alignment[$i]];
$totalw += $matrix[$i][$alignment[$i]];
}
}
return $totalw,\%alignmenthash,\%weighthash;} |
sub optimal_alignment_generalized
{ my ($net1,$net2,%params)=@_;
my ($netr1,$netr2)=$net1->topological_restriction($net2);
return $netr1->optimal_alignment($netr2,%params); } |
sub weight
{ my ($net1,$v1,$net2,$v2,$distance)=@_;
my $w;
if (! defined $distance) {
$distance='Manhattan';
}
if ($distance eq 'Hamming') {
$w=$net1->{mudata}->{$v1}->hamming($net2->{mudata}->{$v2});
} else {
$w=$net1->{mudata}->{$v1}->manhattan($net2->{mudata}->{$v2});
}
if (($net1->is_tree_node($v1) && $net2->is_hybrid_node($v2)) ||
($net2->is_tree_node($v2) && $net1->is_hybrid_node($v1))
)
{
$w +=1/(2*$net1->{numleaves}); }
return $w;} |
sub topological_restriction
{ my ($net1,$net2)=@_;
my @leaves1=$net1->leaves();
my @leaves2=$net2->leaves();
my $numleaves1=scalar @leaves1;
my $numleaves2=scalar @leaves2;
my %position1;
for (my $i=0; $i<$numleaves1; $i++) {
$position1{$leaves1[$i]}=$i;
}
my %position2;
my @commonleaves=();
for (my $j=0; $j<$numleaves2; $j++) {
if (defined $position1{$leaves2[$j]}) {
push @commonleaves,$leaves2[$j];
$position2{$leaves2[$j]}=$j;
}
}
my $graphred1=$net1->{graph}->copy();
my $graphred2=$net2->{graph}->copy();
OUTER1:
foreach my $u ($graphred1->vertices()) {
my $mu=$net1->mudata_node($u);
foreach my $leaf (@commonleaves) {
if ($mu->[$position1{$leaf}]>0) {
next OUTER1;
}
}
$graphred1->delete_vertex($u);
}
OUTER2:
foreach my $u ($graphred2->vertices()) {
my $mu=$net2->mudata_node($u);
foreach my $leaf (@commonleaves) {
if ($mu->[$position2{$leaf}]>0) {
next OUTER2;
}
}
$graphred2->delete_vertex($u);
}
my $netr1=Bio::PhyloNetwork->new(-graph => $graphred1);
my $netr2=Bio::PhyloNetwork->new(-graph => $graphred2);
return ($netr1,$netr2);
}
} |
sub eNewick
{ my ($self)=@_;
my $str="";
my $seen={};
foreach my $root ($self->roots()) {
$str=$str.$self->eNewick_aux($root,$seen,undef)."; ";
}
return $str;} |
sub eNewick_aux
{ my ($self,$node,$seen,$parent)=@_;
my $str='';
if ($self->is_leaf($node) ||
(defined $seen->{$node}) )
{
$str=make_label($self,$parent,$node);
}
else {
$seen->{$node}=1;
my @sons=$self->{graph}->successors($node);
$str="(";
foreach my $son (@sons) {
$str=$str.$self->eNewick_aux($son,$seen,$node).",";
}
chop($str);
$str.=")".make_label($self,$parent,$node);
}
return $str;} |
sub make_label
{ my ($self,$parent,$node)=@_;
my $str='';
if ($self->is_hybrid_node($node)) {
my $lbl=$self->{labels}->{$node};
if ($lbl =~ /#/) {
$lbl='';
}
$str.=$lbl; $str.='#';
if ((defined $parent) &&
($self->graph->has_edge_attribute($parent,$node,'type'))) {
$str.=$self->graph->get_edge_attribute($parent,$node,'type');
}
$str.=substr $node,1;
} else {
$str.=$self->{labels}->{$node};
}
if ((defined $parent) &&
($self->graph->has_edge_weight($parent,$node))) {
$str.=":".$self->graph->get_edge_weight($parent,$node);
}
return $str;} |
sub eNewick_full
{ my ($self)=@_;
my $str="";
my $seen={};
foreach my $root ($self->roots()) {
$str=$str.$self->eNewick_full_aux($root,$seen,undef)."; ";
}
return $str;} |
sub eNewick_full_aux
{ my ($self,$node,$seen,$parent)=@_;
my $str='';
if ($self->is_leaf($node) ||
(defined $seen->{$node}) )
{
$str=make_label_full($self,$parent,$node);
}
else {
$seen->{$node}=1;
my @sons=$self->{graph}->successors($node);
$str="(";
foreach my $son (@sons) {
$str=$str.$self->eNewick_full_aux($son,$seen,$node).",";
}
chop($str);
$str.=")".make_label_full($self,$parent,$node);
}
return $str;} |
sub make_label_full
{ my ($self,$parent,$node)=@_;
my $str='';
if ($self->is_hybrid_node($node)) {
my $lbl=$self->{labels}->{$node};
if ($lbl =~ /#/) {
$lbl='';
}
$str.=$lbl; $str.='#';
if ((defined $parent) &&
($self->graph->has_edge_attribute($parent,$node,'type'))) {
$str.=$self->graph->get_edge_attribute($parent,$node,'type');
}
$str.=substr $node,1;
} else {
if ((defined $self->{labels}->{$node})&&($self->{labels}->{$node} ne '')) {
$str.=$self->{labels}->{$node};
}
else {
$str.=$node;
}
}
if ((defined $parent) &&
($self->graph->has_edge_weight($parent,$node))) {
$str.=":".$self->graph->get_edge_weight($parent,$node);
}
return $str;
}
use overload '""' =>\& display;} |
sub display
{ my ($self)=@_;
my $str="";
my $graph=$self->{graph};
my @leaves=$self->leaves();
my @nodes=@{$self->{nodes}};
$str.= "Leaves:\t@leaves\n";
$str.= "Nodes:\t@nodes\n";
$str.= "Graph:\t$graph\n";
$str.= "eNewick:\t".$self->eNewick()."\n";
$str.= "Full eNewick:\t".$self->eNewick_full()."\n";
$str.= "Mu-data and heights:\n";
foreach my $node (@nodes) {
$str.= "v=$node: ";
if (exists $self->{labels}->{$node}) {
$str.="\tlabel=".$self->{labels}->{$node}.",";
} else {
$str.="\tlabel=(none),";
}
$str.= "\th=".$self->{h}->{$node}.",\t mu=".$self->{mudata}->{$node}."\n";
}
if (exists $self->{has_temporal_representation}) {
$str.= "Temporal representation:\n";
if ($self->{has_temporal_representation}) {
foreach my $node (@nodes) {
$str.= "v=$node; ";
$str.= "\tt=".$self->{temporal_representation}->{$node}."\n";
}
} else {
$str.= "Does not exist.\n";
}
}
if (exists $self->{tripartitions}) {
$str.= "Tripartitions:\n";
foreach my $node (@nodes) {
$str.= "v=$node; ";
$str.= "\ttheta=".$self->{tripartitions}->{$node}."\n";
}
}
return $str;
}
1;} |
General documentation
Gabriel Cardona, gabriel(dot)cardona(at)uib(dot)es
Gabriel Valiente, valiente(at)lsi(dot)upc(dot)edu
[CRV1]
G. Cardona, F. Rossello, G. Valiente. Tripartitions do not always
discriminate phylogenetic networks. arXiv:0707.2376v1 [q-bio.PE]
[CRV2]
G. Cardona, F. Rossello, G. Valiente. A Distance Measure for
Tree-Child Phylogenetic Networks. Preprint.
[NetGen]
M.M. Morin, and B.M.E. Moret. NetGen: generating phylogenetic networks
with diploid hybrids. Bioinformatics 22 (2006), 1921-1923
[PhyloNet]
PhyloNet: "Phylogenetic Networks Toolkit".
http://bioinfo.cs.rice.edu/phylonet [BSS]
M. Baroni, C. Semple, and M. Steel. Hybrids in Real
Time. Syst. Biol. 55(1):46-56, 2006
The rest of the documentation details each of the object methods.