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Week 10: Consensus trees, tree distances, tests of tree shape Genome 570 March, 2012 Week 10: Consensus trees, tree distances, tests of tree shape – p.1/42

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Page 1: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Week 10: Consensus trees, tree distances, tests of treeshape

Genome 570

March, 2012

Week 10: Consensus trees, tree distances, tests of tree shape – p.1/42

Page 2: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Trees we will use for consensus trees

DA C B E D FG A CG F B E D A CG F B E

Week 10: Consensus trees, tree distances, tests of tree shape – p.2/42

Page 3: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Trees we will use for consensus trees

A C B E D FG A CG F B E D A CG F B E D

(for unrooted trees we would use partitions induced by branches insteadof clades)

Week 10: Consensus trees, tree distances, tests of tree shape – p.3/42

Page 4: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Trees we will use for consensus trees

A C B E D FG A CG F B E D A CG F B E D

Week 10: Consensus trees, tree distances, tests of tree shape – p.4/42

Page 5: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Trees we will use for consensus trees

A C B E D FG A CG F B E D A CG F B E D

(Do we count this one if the trees are considered rooted? unrooted?)

Week 10: Consensus trees, tree distances, tests of tree shape – p.5/42

Page 6: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Their strict consensus tree

A CG F B E D

Week 10: Consensus trees, tree distances, tests of tree shape – p.6/42

Page 7: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A distressing case for the strict consensus tree

A B C D E F G B C D E F G A

Only one species moves ...

Week 10: Consensus trees, tree distances, tests of tree shape – p.7/42

Page 8: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A distressing case for the strict consensus tree

A B C D E F G

... but the strict consensus tree becomes totally unresolved.

Week 10: Consensus trees, tree distances, tests of tree shape – p.8/42

Page 9: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Majority-rule consensus tree

A CG F B E D

100

100

67

67100

Week 10: Consensus trees, tree distances, tests of tree shape – p.9/42

Page 10: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

The Adams consensus tree

For rooted trees, Adams (1972, 1986) suggested:

1. Take all rooted triples on each tree.

2. Retain those that are not contradicted, where lack of resolution doesnot count as contradiction.

3. Construct a tree of these.

Week 10: Consensus trees, tree distances, tests of tree shape – p.10/42

Page 11: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Two of the possible triples to examine

DA C B E D FG A CG F B E D A CG F B E

The green triple shows the same rooted topology on all three trees. Thered triple is contradicted and does not get used in the Adams ConsensusTree.

Week 10: Consensus trees, tree distances, tests of tree shape – p.11/42

Page 12: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

The Adams consensus tree

A CG F B E D

Week 10: Consensus trees, tree distances, tests of tree shape – p.12/42

Page 13: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Steel, Böcker, and Dress’s shocking disproof

Steel, M., S. Böcker, and A. W. M. Dress. 2000. Simple but fundamentallimits for supertree and consensus tree methods. Systematic Biology 49(2):363-368.

They put forward three minimal requirements for an unrooted Adams-likeconsensus tree based on observations of quartets, rather than triples.Note that a quartet, like a triple, has three possible topologies, butunrooted ones: ((A,B),(C,D)) and ((A,C),(B,D)) and ((A,D),(B,C)).

The result shouldn’t be altered by relabelling all the species in aconsistent way.

The result should not depend on the order in which the trees areinput.

If a quartet appears in all trees, it should appear in the consensus.

Alas, they then show there is no consensus tree method for unrootedtrees that can satisfy all of these!

Week 10: Consensus trees, tree distances, tests of tree shape – p.13/42

Page 14: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A consensus subtree

F C A B G DFCA BDE F AB D E

Week 10: Consensus trees, tree distances, tests of tree shape – p.14/42

Page 15: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A consensus subtree

F C A B G DFCA BDE F AB D E

Week 10: Consensus trees, tree distances, tests of tree shape – p.15/42

Page 16: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A consensus subtree

F C A B G DFCA BDE F AB D E

B DFA

Week 10: Consensus trees, tree distances, tests of tree shape – p.16/42

Page 17: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

The symmetric difference metric

AB

C

D

F

E G E

F

AD

B

C

G

Partitions

{ADF | BCEG}

{BC | ADEFG}

Partitions

{ADF | BCEG}

{EG | ABCDF}

{DF | ABCEG} {AD | BCEFG}

{BC | ADEFG}

The symmetric difference is the number of partitions that are in one butnot both of these lists, in this case 3.

Week 10: Consensus trees, tree distances, tests of tree shape – p.17/42

Page 18: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

The Robinson-Foulds and Branch Length distances

D

E0.1 0.1

G

B

C0.2

0.2 0.15

A0.1

0.2F

0.30.05

0.4

0.2 E

F0.1

0.2

0.3

B

C0.2

0.150.1 0.3

D0.2

0.20.1

A

G

0.4

0.2

0.3

0.1

0.05

0.2

none

0.15

0.1

0.1

0.2

0.2

{ADF | BCEG}

{AD | BCEFG}

{A | BCDEFG}

{B | ACDEFG}

{C | ABDEFG}

{D | ABCEFG}

{E | ABCDFG}

{F | ABCDEG}

{G | ABCDEF}

{DF | ABCEG}

{EG | ABCDF}

Partitions Branch lengths

{BC | ADEFG}

0.15

0.1

0.1

0.2

0.2

0.3

none

none

0.2

0.2

0.1

0.3

difference

−0.2

0.1

−0.1

0.3

0.1

−0.15

0.0

0.0

0.0

0.0

0.1

0.0

RF: Σ | difference | = 1.05

BLD: Σ (difference)2

= 0.1925Week 10: Consensus trees, tree distances, tests of tree shape – p.18/42

Page 19: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Asymmetry at base of a tree

A tree with n tips can, at the bottom fork, have clades whose ultimatesizes are 1 : n − 1, 2 : n − 2, 3 : n − 3, . . . n − 1 : 1.

Under a random branching model, these possibilities are all equiprobable(this is true when there is random death of lineages too). As we can’t tellwhich clade is on the left, we can symmetrize this, always putting thesmaller clade on the left.

So with 7 tips we have equal probabilities of 1 : 6, 2 : 5, and 3 : 4. With aneven number of tips (say 8) we have to be more careful, as 4 : 4 is thenonly half as probable as as each of the others 1 : 7, 2 : 6, and 3 : 5..

Week 10: Consensus trees, tree distances, tests of tree shape – p.19/42

Page 20: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Harding’s probability of a tree shape

2:2

1:1 1:1 1:11:11:2

1:3

2:4

4:629

25

23

1 1 111

31

Week 10: Consensus trees, tree distances, tests of tree shape – p.20/42

Page 21: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Birth-death process tree

ab

c

d

efghij

Week 10: Consensus trees, tree distances, tests of tree shape – p.21/42

Page 22: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

If we remove the extinct lineages

ab

c

d

efghij

Week 10: Consensus trees, tree distances, tests of tree shape – p.22/42

Page 23: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

We get ...

ab

c

d

efghij

Week 10: Consensus trees, tree distances, tests of tree shape – p.23/42

Page 24: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Reconstructed lineages from that tree

abcdefghij

Week 10: Consensus trees, tree distances, tests of tree shape – p.24/42

Page 25: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Birth-death processes and tree shapes

With branching rate λ and death rate µ,

probability of survival of a lineage for t is

s(t) = Prob (n > 0 | t) =(λ − µ)e(λ−µ)t

λ e(λ−µ)t − µ

Week 10: Consensus trees, tree distances, tests of tree shape – p.25/42

Page 26: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Number of reconstructed and actual lineages

Nee, May, and Harvey (1994) argue that the number of lineages at time t

ago who have descendants in the present, given that they started T timeago, is expected to be

E[NR(t, T)] =e(λ−µ)t

s(T − t)

s(T)

and the expected number that actually were there then, given that at leastone is still alive now, is

E[N(t, T)] =e(λ−µ)t

s(T)−

s(t) − s(T)

s(t)s(T − t)

Week 10: Consensus trees, tree distances, tests of tree shape – p.26/42

Page 27: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Reconstructed and actual lineages

0 101

10

100

20

2

5

50

155

Actual

Reconstructed

Time

Spe

cies

Week 10: Consensus trees, tree distances, tests of tree shape – p.27/42

Page 28: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Rooted tree with ancestors in “intermediate” position

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.28/42

Page 29: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Rooted tree with ancestors in “centered” position

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.29/42

Page 30: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Rooted tree with ancestors in “weighted” position

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.30/42

Page 31: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Rooted tree with ancestors in “inner” position

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.31/42

Page 32: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Drawn with diagonal lines

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.32/42

Page 33: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Drawn so as to be V-shaped

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.33/42

Page 34: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A curvogram

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.34/42

Page 35: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A swoopogram

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.35/42

Page 36: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A eurogram

AB

CD

EFG

HIJ

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.36/42

Page 37: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

A circular drawing of a rooted tree

A

B

CD E

F

G

H

I

JK

L

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.37/42

Page 38: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

The equal-angle algorithm

A

B

I

J

C D

E

F

GH

o72

o72

o144

o72

o216

o36

o108 o

72

o36

Week 10: Consensus trees, tree distances, tests of tree shape – p.38/42

Page 39: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

The equal-angle algorithm tree for the example

AB

C

D

E

F

G

H

I

J

KL

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.39/42

Page 40: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

The n-body algorithm for the example

A

B

C

D

E

F

G

H

I

J

K

L

M

Week 10: Consensus trees, tree distances, tests of tree shape – p.40/42

Page 41: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

The equal-daylight algorithm

F

H

AB

I

J

C D

E

G

Week 10: Consensus trees, tree distances, tests of tree shape – p.41/42

Page 42: Week 10: Consensus trees, tree distances, tests of tree shapeevolution.gs.washington.edu/gs570/2012/week10.pdf · Week 10: Consensus trees, tree distances, tests of tree shape –

Equal-daylight tree for the example

ABCD

E

F

G

HI

J

K

L

MWeek 10: Consensus trees, tree distances, tests of tree shape – p.42/42