/* * MultiSpeciesCoalescent.java * * Copyright (c) 2002-2015 Alexei Drummond, Andrew Rambaut and Marc Suchard * * This file is part of BEAST. * See the NOTICE file distributed with this work for additional * information regarding copyright ownership and licensing. * * BEAST is free software; you can redistribute it and/or modify * it under the terms of the GNU Lesser General Public License as * published by the Free Software Foundation; either version 2 * of the License, or (at your option) any later version. * * BEAST is distributed in the hope that it will be useful, * but WITHOUT ANY WARRANTY; without even the implied warranty of * MERCHANTABILITY or FITNESS FOR A PARTICULAR PURPOSE. See the * GNU Lesser General Public License for more details. * * You should have received a copy of the GNU Lesser General Public * License along with BEAST; if not, write to the * Free Software Foundation, Inc., 51 Franklin St, Fifth Floor, * Boston, MA 02110-1301 USA */ package dr.evomodel.speciation; import dr.evolution.coalescent.DemographicFunction; import dr.evolution.coalescent.ScaledDemographic; import dr.evolution.tree.NodeRef; import dr.evolution.tree.TreeUtils; import dr.evolution.util.Units; import dr.inference.model.Likelihood; import dr.inference.model.Model; import jebl.util.FixedBitSet; import java.util.Arrays; /** * Compute coalecent log-liklihood of a set of gene trees embedded inside one species tree. * * @author Joseph Heled, Graham Jones * Date: 26/05/2008 */ public class MultiSpeciesCoalescent extends Likelihood.Abstract implements Units { private final SpeciesTreeModel spTree; private final SpeciesBindings species; private boolean checkCompatibility; private final boolean[] compatibleCheckRequited; public MultiSpeciesCoalescent(SpeciesBindings species, SpeciesTreeModel tree) { super(tree); spTree = tree; this.species = species; spTree.addModelRestoreListener(this); // recompute on any change in geneTree - // possible optimization: keep track which tree changed. final SpeciesBindings.GeneTreeInfo[] trees = species.getGeneTrees(); for(SpeciesBindings.GeneTreeInfo geneTree : trees) { geneTree.tree.addModelListener(this); } compatibleCheckRequited = new boolean[trees.length]; Arrays.fill(compatibleCheckRequited, false); checkCompatibility = false; } // override this for efficiency, otherwise the overridden makeDirty, which results in additional overhead is called public void modelRestored(Model model) { super.makeDirty(); } // Upon a direct "make dirty" enable all compatibility checks, since the last call to calculateLogLikelihood may have // found a non compatible tree and returned -inf. This case is not explicitly saved. public void makeDirty() { super.makeDirty(); checkCompatibility = true; for(int i = 0; i < species.getGeneTrees().length; i++) { compatibleCheckRequited[i] = true; } } protected double calculateLogLikelihood() { if( checkCompatibility ) { boolean compatibility = true; for(int i = 0; i < compatibleCheckRequited.length; ++i) { if( compatibleCheckRequited[i] ) { if( !spTree.isCompatible(species.getGeneTrees()[i]) ) { compatibility = false; } compatibleCheckRequited[i] = false; //System.out.println("check compatibility:" + species.getGeneTrees()[i].tree.getId() + " - " + compatibility ); } } if( !compatibility ) { return Double.NEGATIVE_INFINITY; } checkCompatibility = false; } double logl = 0; int[] info = {0, 0}; for( SpeciesBindings.GeneTreeInfo geneTree : species.getGeneTrees() ) { final double v = treeLogLikelihood(geneTree, spTree.getRoot(), info, geneTree.popFactor()); assert ! Double.isNaN(v); // if( Double.isNaN(v) ) { // double x = 0; // } logl += v; } ccc += 1; return logl; } int ccc = 0; private final boolean verbose = false; private double treeLogLikelihood(SpeciesBindings.GeneTreeInfo geneTree, NodeRef node, int[] info, double popFactor) { // number of lineages remaining at node int nLineages; // location in coalescent list (optimization) int indexInClist = 0; // accumulated log-likelihood inBranchh from node to it's parent double like = 0; final double t0 = spTree.getNodeHeight(node); final SpeciesBindings.CoalInfo[] cList = geneTree.getCoalInfo(); if( verbose ) { if( spTree.isRoot(node) ) { System.err.println("gtree:" + geneTree.tree.getId()); System.err.println("t0 " + t0); for(int k = 0; k < cList.length; ++k) { System.err.println(k + " " + cList[k].ctime + " " + cList[k].sinfo[0] + " " + cList[k].sinfo[1]); } } } if( spTree.isExternal(node) ) { nLineages = geneTree.nLineages(spTree.speciesIndex(node)); indexInClist = 0; } else { //assert spTree.getChildCount(node) == 2; nLineages = 0; for(int nc = 0; nc < 2; ++nc) { final NodeRef child = spTree.getChild(node, nc); like += treeLogLikelihood(geneTree, child, info, popFactor); nLineages += info[0]; indexInClist = Math.max(indexInClist, info[1]); } // The root of every gene tree (last coalescent point) should be always above // root of species tree if( indexInClist >= cList.length) { int k = 1; } assert indexInClist < cList.length; // Skip over (presumably, not tested by assert) non interesting coalescent // events to the first event before speciation point while( cList[indexInClist].ctime < t0 ) { ++indexInClist; } } final boolean isRoot = spTree.isRoot(node); // Upper limit // use of (t0 + spTree.getBranchLength(node)) caused problem since there was a tiny difference // between those (supposedly equal) values. we should track where the discrepancy comes from. final double stopTime = isRoot ? Double.MAX_VALUE : spTree.getNodeHeight(spTree.getParent(node)); // demographic function is 0 based (relative to node height) // time away from node double lastTime = 0.0; // demographic function across branch DemographicFunction demog = spTree.getNodeDemographic(node); if( popFactor > 0 ) { demog = new ScaledDemographic(demog, popFactor); } // if(false) { // final double duration = isRoot ? cList[cList.length - 1].ctime - t0 : stopTime; // double demographicAtCoalPoint = demog.getDemographic(duration); // double intervalArea = demog.getIntegral(0, duration); // if( demographicAtCoalPoint < 1e-12 * (duration/intervalArea) ) { // return Double.NEGATIVE_INFINITY; // } // } // Species sharing this branch FixedBitSet subspeciesSet = spTree.spSet(node); if( verbose ) { System.err.println(TreeUtils.uniqueNewick(spTree, node) + " nl " + nLineages + " " + subspeciesSet + " t0 - st " + t0 + " - " + stopTime); } while( nLineages > 1 ) { // if ( !( indexInClist < cList.length ) ) { // System.err.println( indexInClist ) ; // } assert ( indexInClist < cList.length ); final double nextT = cList[indexInClist].ctime; // while rare they can be equal if( nextT >= stopTime ) { break; } if( nonEmptyIntersection(cList[indexInClist].sinfo, subspeciesSet) ) { final double time = nextT - t0; if( time > 0 ) { final double interval = demog.getIntegral(lastTime, time); lastTime = time; final int nLineageOver2 = (nLineages * (nLineages - 1)) / 2; like -= nLineageOver2 * interval; final double pop = demog.getDemographic(time); assert( pop > 0 ); like -= Math.log(pop); } --nLineages; } ++indexInClist; } if( nLineages > 1 ) { // add term for No coalescent until root final double interval = demog.getIntegral(lastTime, stopTime - t0); final int nLineageOver2 = (nLineages * (nLineages - 1)) / 2; like -= nLineageOver2 * interval; } info[0] = nLineages; info[1] = indexInClist; if( verbose ) { System.err.println(TreeUtils.uniqueNewick(spTree, node) + " stopTime " + stopTime + " nl " + nLineages + " icl " + indexInClist); } return like; } public void modelChangedEvent(Model model, Object object, int index) { //super.modelChangedEvent(model, object, index); // not the above for efficiency, otherwise the overridden makeDirty, which results in additional overhead is called. super.makeDirty(); if( model == spTree ) { if( object == spTree && index != -1 ) { // Species tree scaling checkCompatibility = true; Arrays.fill(compatibleCheckRequited, true); } } else { final SpeciesBindings.GeneTreeInfo[] trees = species.getGeneTrees(); for(int i = 0; i < species.getGeneTrees().length; i++) { if( trees[i].tree == model ) { checkCompatibility = true; compatibleCheckRequited[i] = true; break; } } } } private boolean nonEmptyIntersection(FixedBitSet[] sinfo, FixedBitSet subspeciesSet) { for( FixedBitSet nodeSpSet : sinfo ) { if( nodeSpSet.intersectCardinality(subspeciesSet) == 0 ) { return false; } } return true; } public Type getUnits() { return spTree.getUnits(); } public void setUnits(Type units) { assert false; } }