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PROFESSOR: Section 3, Linkage Maps, where our hero Alfred Sturtevant, the
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19-year-old junior at Columbia, takes home the data and figures out why the
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Chromosome Theory really truly is right.
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So he starts out saying black and vestigial, right up there in
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the cross we did.
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How far apart were they?
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What was the recombination frequency?
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STUDENT: 17%.
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PROFESSOR: 17%, so let's put them down on the map and write--
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I'll move it over.
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We'll just put it there.
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17%.
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Now he goes to some other piece of data in the lab and he says, oh, yeah,
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someone did a cross with vestigial and that eye color thing I showed you over
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there, cinnabar, an eye color mutant.
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And what they found was that vestigial and cinnabar had 8%
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recombination frequency.
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He said, where should I put it on the map?
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Cinnabar is 8% away from vestigial.
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Where should it go on the map?
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STUDENT: Depends how far it is from it.
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PROFESSOR: Depends on which side it's on.
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So maybe cinnabar is over here, or maybe cinnabar is over here, but
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either way it's 8% to the left or 8% to the right of vestigial, if this
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Chromosome Theory is right.
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Which is it?
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How do we know whether it's to the right or to the left?
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[INTERPOSING VOICES]
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PROFESSOR: Ah, testable predictions.
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For the first time in all this stuff, we're hearing some testable
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predictions.
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Testable prediction--
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if it's over here on the left and it's 8% away, 8% over here, what should the
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distance be--
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we'll call that cinnabar, because that's what it was--
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what should the distance be between black and cinnabar?
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STUDENT: 9%.
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PROFESSOR: It should be 9%.
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But what if instead cinnabar is over here?
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How far will black be away?
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STUDENT: 25.
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PROFESSOR: It'll be the 17 plus 8.
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It'll be 17 plus 8.
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He checks the data.
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Black and vestigial are 17, vestigial to cinnabar is 8, black
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to cinnabar is 9.
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He had two choices, and it turned out that this looks like it's the map--
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9, 8, 17.
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Now he goes further.
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He gets another cross.
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There's a mutant called lobe.
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Lobe has 5% recombination away from vestigial.
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Where do you want to stick it?
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One side or the other.
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If it's over here at 5%, the lobe mutant--
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give me a testable prediction.
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Cinnabar to lobe will be how much?
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STUDENT: 13.
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PROFESSOR: 13.
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It is.
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How far will it be from black to lobe?
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STUDENT: 22.
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PROFESSOR: How much?
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STUDENT: 22.
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PROFESSOR: 22.
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It is.
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This is a very good all-nighter.
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He's sitting there drawing things.
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Try another one over here, curved wing.
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It's 3 away from lobed.
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If it's on that side, then this will be 8.
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That will be 16, et cetera, et cetera.
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Every new gene that I drop into that map ends up making lots of testable
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predictions.
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And just the opposite of all the other stuff that has been going on, those
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testable predictions are being tested and they're turning out to be right.
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That is so cool.
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Alfred Sturtevant is saying the idea that these things lie on a line makes
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these testable predictions of all those distances are going to be, and
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they're all checking out.
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He does one other thing.
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He says this involves pair wise.
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These are two factor crosses, crosses that just involve two genes
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simultaneously.
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They're like we've been drawing over there.
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It's just pairwise testing of particular pairs of genes.
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What about three factor crosses?
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What do I mean by a three factor cross?
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Well, I'm going to forget about black and cinnabar and vestigial and all
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those sort of things for the moment.
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And I'm just going to make it very simple and say, suppose I had a cross
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where in the F1 I had a, b, c, plus, plus, plus.
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I'm going to cross it to a, b, c, a, b, c, and I'm going to look at the
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possible gametes that could come out in a three factor cross.
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How many possible gametes can come out in a three factor cross?
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I've got two choices at the first locus--
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at the first gene, I've got two choices at the next gene and I've got
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two choices and the next gene.
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That makes eight possible choices, right.
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So let's actually write those down.
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Let's see.
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If this theory is right, this Chromosome Theory is right, I've got
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my a, b, c, I've got my plus, plus, plus, and crossovers could occur.
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Maybe there's no crossovers or maybe there's a crossover here or a
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crossover there.
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Oh, I don't know.
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Let's for the sake of argument, say this is 10% recombination here and 10%
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recombination here just in my simple example.
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Well, what are the possibilities?
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I could have no recombinations at all.
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If no recombination occurs, no crossing over occurs, then what kind
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of gametes can I get out of that?
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I could get a, b, c, or I could get plus plus plus.
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Now I could have one recombination that separates a away.
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So what could I get there?
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For a recombination occurs that separates from b and c, what kind of
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gametes could there be?
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a plus plus or plus b c will be the kinds of gametes that I could get. a
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plus plus or plus b c, and one recombination can do it.
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What's the chance of one recombination occurring, a recombination occurring
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here but not here?
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What's the chance of recombination here?
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We said 10%.
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What's the chance of no recombination here?
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STUDENT: 90.
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PROFESSOR: 90%.
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So that's going to be 10% times 90%, so that means 9% of the time
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we might see that.
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Let's just check out one over here.
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What's the chance there's no recombination here and no
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recombination here?
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STUDENT: 90 times 90.
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PROFESSOR: 90 times 90, 90% times 90% is 81%.
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Let's try this recombination now over here that separates away c.
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What are the possible gametes?
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We'll have recombination separating c.
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What do we get, a b plus or plus b c--
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I'm sorry, or plus plus c.
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There we go.
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And what's the probability of that occurring?
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STUDENT: 90%.
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PROFESSOR: 90% times 10% equals 9%.
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But can I ever get a gamete where b has been separated away from little a
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and little c?
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STUDENT: Two recombinations.
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PROFESSOR: It takes two recombinations, not one recombination.
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Two won't do, so one won't do it.
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You need two recombinations.
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You need this event and that event.
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Two recombinations, two recombination events, and then you
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can separate b away.
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But because b is in the middle, it takes two, and you're going to get
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plus b plus, a plus c.
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And what's the chance of two recombinations?
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10% times 10% equals 1%.
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One of the three is very hard to separate away from the others because
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it takes two crossovers if it's in the middle.
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That is a pretty cool testable prediction that based on the two
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factor distances and the map of who's in the middle, you can predict that
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one of them is going to be very much harder to separate from the others
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because it takes two crossovers.
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And when you look at the data, that's what you see.
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The sun rises over Manhattan.
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Bleary Alfred Sturtevant realizes that the Chromosome Theory checks out.
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It makes amazing testable predictions of this whole thing.
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Sturtevant writes it up and it becomes one of the most
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famous papers in genetics.
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Why is it such a famous paper in genetics?
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Let's just flip to a fly map here.
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This is an example of the fruit fly genetic map.
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This has only got a teeny fraction of what's on the fruit fly genetic map.
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You've got vermilion and scalloped wings and here's cinnabar and
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vestigial and lobed curved and black body.
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Right over here is where we've been working on the map.
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And all of these different traits are there.
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This is, like, way cool, this map that he's produced and that scientists
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throughout the rest and rest of the century produce.
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We've got a question for you now about recombination frequency.
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Try answering this one to be sure that you understand how
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recombination works.
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