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The Evolutionary history of related organisms
Students: Please DOWNLOAD this PowerPoint and then view it in presentation mode to be able to view the animations. For best learning, do not rush through the slides. PHYLOGENY… The Evolutionary history of related organisms Photographic images within this PowerPoint were obtained from:
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Distinguishing Phylogeny vs. Taxonomy
Taxonomy – a hierarchical structure of grouping & naming or classifying organisms based on similarities and differences Phylogeny – Depicts the evolutionary relationships of a group of organisms. Phylogenetic trees indicate when/where species diverged from a common ancestor.
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Phylogenetic trees show lines of descent
The branching patterns that make up a phylogenetic tree are not usually observed directly, rather they are inferred from data Red circle – extinction. What does this dead-end branch depict? Try to answer the question before you click again. Make sure you do this each time a question is asked. An extinction Image adapted from: Baum, D.A. and Smith S.D. Tree thinking: an introduction to phylogenetic biology. Roberts, 2013.
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Before we can create phylogenetic trees, we need to ask…
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How are organisms classified?
Let’s start with a set of species named as follows… A B C D E F We have evidence that: B and C are more closely related to each other than to any other species A is related to B and C, but not as closely as B and C are related to each other D, E and F are more closely related to each other than to A, B or C Let’s start here and learn a simple way to organize the species based on relatedness.
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How are organisms classified?
We know: B and C are closely related Start by grouping the most closely related organisms together. Use parentheses to put them together in a set… …and a comma to show that they share a common ancestor. B C (B C) (B,C)
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How are organisms classified?
We know: B and C are closely related A is related to both B and C, but not as closely as B and C are related to each other Now add the next most closely related organism Place this organism next to the first set, add parentheses to group them into a larger set, and a comma to show the common ancestor. A (B,C) (B,C) (A (B,C)) (A,(B,C))
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How are organisms classified?
What about the other organisms? D E F D, E and F are more closely related to each other than to A, B or C. If D, E and F are equally related to each other (or you’re not sure!) put them together in a set using parentheses. And separate them with commas to show they share a common ancestry. D E F (D E F) (D,E,F)
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How are organisms classified?
Let’s bring this all together. Because the DEF set is more distantly related from the ABC set, we keep the 2 sets separate but put them next to each other. Then we add a final set of parentheses to enclose all of the species as one larger set. Don’t forget the comma, to show the common ancestor between the two sets that make up the larger set. ((A,(B,C)),(D,E,F)) (A,(B,C)) (D,E,F)
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How are organisms classified?
One last thing! To show that you have completed the tree, you add a semi colon to the end of the set. ((A,(B,C)),(D,E,F));
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How are organisms classified?
((A,(B,C)),(D,E,F)); What you have just made is a phylogenetic tree! Click to see how this can be converted into a phylogenetic tree…
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(B,C) Present Don’t worry about the terms for right now… Past
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(A,(B,C)) Present Past
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(D,E,F) Present Past
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((A,(B,C)),(D,E,F)); Present Past
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((A,(B,C)),(D,E,F)); Present Past
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Some common terminology
Branches indicate evolutionary path Descendants (Current taxa) at tips Nodes indicate common ancestor Other terms shown here will be addressed later. Past Present
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Now that we’ve seen how trees are put together let’s look at how to interpret them.
Past Present
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Unless stated otherwise, assume that absolute time is NOT represented
C D α δ Trees ONLY depict relatedness and not time unless specifically stated otherwise. Alpha happened after beta (alpha occurred closer to the present than beta). We don’t know if alpha came before or after delta. β Was α before or after δ? Was α before or after β? You can’t tell
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More terminology A cladogram indicates relatedness only!
Sister taxa are most closely related because the MRCA (most recent common ancestor) that they share is more recent than the ones they share with another taxon on this tree A cladogram shows relatedness only! Branch length is not always meaningful – Taxon B and C are not more or less evolved than taxon D, E, and F. Sister taxa are NOT closely related because they are next to one another at the tips. A and B are NOT sister taxa, whereas B and C are, because they share an MRCA. Branch length is not always meaningful
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Same order – different tree?
Let’s look at these two trees. The taxa are in the same order at the tips, but the branching appears to be different. Tree on the left: human and mouse are sister taxa - most closely related, share a common ancestor. Tree on the right: lizard and mouse are most closely related. These circles show just one difference. There are more. You should identify them. Use the Tree Thinking Article. What’s different? Images via Wikimedia Commons used under Creative Commons Attribution 2.0 Generic License.
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Don’t be distracted by morphological similarity
It can be tempting to think about what organisms look like when interpreting these trees, but you really only want to focus on branching patterns. Lizards and frogs have many similar morphological features –four legs, they might go in water and live on land, they both eat insects, etc. But, this tree shows us that lizards are more closely related to mice and humans. While they do share an MRCA with frogs, that ancestor lived farther in the past that the MRCA circled in red and it is the SAME common ancestor shared with humans. Use the fish in this tree as the outgroup – to help root the tree; we know fish are distantly related, as they are not tetrapods. Most Recent Common Ancestor (MRCA) Is a lizard more closely related to a frog or a human? Images via Wikimedia Commons used under Creative Commons Attribution 2.0 Generic License
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Assembling a Tree: Practice #2
Let’s practice putting that tree together ourselves, using the set method from earlier in this tutorial Five organisms – How are they related? We know that the human and the mouse are most closely related. So let’s group them together in a set. We’ll do this one with pictures! ( , )
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Assembling a Tree: Practice #2
Let’s practice putting that tree together ourselves, using the set method from earlier in this tutorial Five organisms – How are they related? Next, we know that lizards are related most closely to the human/mouse set… ( ,( , ))
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Assembling a Tree: Practice #2
Let’s practice putting that tree together ourselves, using the set method from earlier in this tutorial Five organisms – How are they related? And frogs are related most closely to the human/mouse/lizard set… ( ,( ,( , ))))
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Assembling a Tree: Practice #2
Let’s practice putting that tree together ourselves, using the set method from earlier in this tutorial Five organisms – How are they related? Finally, we add the fish in. ( ,( ,( ,( , ))))
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Assembling a Tree: Practice #2
What’s missing from this tree? ( ,( ,( ,( , )))) ; The semi-colon at the end. This indicates that the tree is complete. If we wanted to write this out using names, so that we could enter it into a phylogeny-making computer program, it would look like this: (Fish,(Frog,(Lizard,(Human,Mouse))));
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Trees can be drawn in various ways yet contain the same phylogenetic information
The taxa at the tips of all 4 trees are in the same order – are the evolutionary relationships the same? All of these trees depict the exact same evolutionary relationships among taxa A through F. For example, all trees show that F and C are sister taxa, along with E and D. B is equally closely related to C and F. The BCF clade shares an MRCA with the ED clade. Phylogenetically, these trees are identical. Gregory Evo Edu Outreach 1:
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Rotation around the nodes does not change the information in a tree.
These three trees are identical 4 3 2 If you rotate tree 1 at node 1 only, you get tree 2! If you rotate node 1, 2, and 3 you get tree 3. If you rotate tree 1 at nodes 1, 2, and 4, you get tree 3! 1 Tree 1 Tree 3 Tree 2 If we rotate around the first node… …we get this …we get this If we rotate around the 1st, 2rd 4th nodes…
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There are no “main lines” in evolution
Just because the human has a “straight shot” does not mean it is more related to the common ancestor than the others. The “straight shot” also does not imply that the human (or the shark) is more advanced than the other taxa. Images via Wikimedia Commons used under Creative Commons Attribution 2.0 Generic License
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Clades can be expanded or collapsed
If we collapse these two regions… …we get this tree …we get this tree We call this “pruning” the tree. If you prune tree (a) by collapsing taxa A, B, C and D into M and collapsing E and F into K, you wind up with tree (b). If you prune (a) by collapsing EFGH into N, collapsing C and D into J, and rotating at each node, you get tree (c).
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A tree can depict polytomy
plant animal fungi If we are unsure which of these is most likely: plant fungi animal Rearranging them to meet at a common point shows uncertainty about how lineages relate to each other. This arrangement is a called a polytomy.
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Monophyletic groups (clades)
Monophyletic group (clade) A clade is a grouping that includes a common ancestor and all the descendants (living and extinct) of that ancestor Scientists try and make sure that each taxon (a named group of organisms) corresponds to a clade Clades = branch-systems have a special importance in understanding diversity. They are monophyletic. Monophyletic groups include an ancestor and all of their descendants. (all members are most-closely related).
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How can we reconstruct phylogenies?
Many methods, some quite sophisticated Simplest is to apply the principle of parsimony: Occam’s Razor - All other things being equal, the simplest solution is the best. Can be done on Morphological or Molecular data The rest of this activity will focus on parsimony analysis with molecular data
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THE PARSIMONY CRITERION
Select the tree that implies the minimum number of character changes summed across all characters of interest
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Let’s use molecular data
RW A A C G A T T C T A A A G G A T T BW A G C G C T T C T A A A G G A T T MW A G C G C T T A T A A A G G A T T LBW A G C G C T T A T A T A G G G T T RW A A C A A BW G C C A A MW G C A A A LBW G C A T G RW = Rock Wren (outgroup) BW = Banded Wren MW = Moustached Wren LBW = Long-Billed Wren First we need to align the DNA (or RNA) sequence data. For each species, we have nucleotide data at 5 positions in the DNA sequence. First, we align the gene sequences that we want to compare and look for places where they differ
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Three possible trees Tree 1 Tree 2 Tree 3
We already know that RW is the outgroup. There are three phylogenies (trees) that we would like to test. Only one of these can be correct. There are three possible trees shown here. For each tree, RW is acting as the outgroup – this is the organism we think is most distantly related to the group of interest – we are trying to figure out the phylogenetic relationship between the moustached wren, the banded wren, and the long-billed wren. Tree 3 Image via Wikimedia Commons used under Creative Commons Attribution 2.0 Generic License.
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Three possible trees ( ,( ,( , ))); ( ,( ,( , ))); Tree 1 Tree 2
( ,( ,( , ))); ( ,( ,( , ))); Tree 1 Tree 2 Here we’ve included the parentheses and commas we would have used to develop the top two trees. Tree 3 Image via Wikimedia Commons used under Creative Commons Attribution 2.0 Generic License.
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Map the characters onto tree 1
These sequences from four different wren species are aligned. Each column (1, 2, 3…) can be considered a character trait. We will therefore analyze each sequence position (“character”) separately. 1 2 3 4 5 LBW MW BW RW A A C A A RW 4 5 3 BW G C C A A T G MW 1 2 LBW Note the first two character states could be mapped onto two different branches (either branch leading to RW or the branch leading to (LBW, MW,BW). For example, RW could have evolved an A from a G (1 step) or A could have evolved to a C on the branch to (LBW, MW, BW) (Also 1 step). Because these first two characters are uninformative as all they do is define the outrgroup, regardless of which branch it is placed on. It does not change the length of the tree or the topology. We prefer to map characters that define the outgroup on that branch to address the misconception that outgroup lineages do not change and evolve. The number mapped above on the tree depicts the “character” change to the left. For 1, the rock wren “character” changed from a G (guanine) to an A (adenine). For 2, the rock wren “character” changed from a C (cytosine) to an A (adenine). Let’s map the remaining character changes Total cost (length) = 6 steps
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Actually there is more than one way to map character 3
1 2 3 4 5 LBW MW BW RW 3 A A C A A RW 3 3 BW G C C A A T G MW LBW Either way the character contributes 2 steps to the overall cost i.e. – you can map this character change in two different ways, each for the same cost (two separate changes needed).
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Map the characters onto tree 2
1 2 3 4 5 MW BW LBW RW A A C A A RW 3 4 5 BW G C C A A T G 3 MW 1 2 LBW Length = 6 steps
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Map the characters onto tree 3
1 2 3 4 5 BW MW LBW RW A A C A A RW 4 5 BW G C C A A T G 3 MW 1 2 LBW Length = 5
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What was the cost of each tree?
Tree 1: length = 6 Tree 2: length = 6 It is most likely that of the 3 possible trees we looked at, for this sequence data, tree 3 is the most likely tree. So, long-billed and moustached wrens are sister taxa (share a common ancestor). Note that these are only 3 of the many possible trees; thankfully, we can use a computer program to do this work for us when we have more taxa and longer sequences. Tree 3 has the lowest cost, is therefore the most parsimonious tree and the one that we believe to be the most likely tree based on Occam’s Razor. Tree 3: length = 5
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This is just the beginning . .
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