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MICHAEL HEMANN: OK, so let's think about two linked genes.
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In this case, two very tightly linked genes.
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One of them is A, and one of them
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is B. So if we have a yeast haploid strain that is AB--
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big A, big B-- we can cross it with another strain that's
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a, little b.
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We can generate a diploid that is AB over ab.
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Here we're using, again, our linkage designation.
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So this line says they're linked together
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on the same chromosome.
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This, of course, is a diploid.
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And we can induce it to undergo meiosis,
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and we'll get a tetrad out of this.
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So let's think about two events.
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One is where we have a meiosis where
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we have basically no crossovers between A and B
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on the same chromosome, and another
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where we have a meiosis where we have one crossover.
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So with a meiosis with no crossovers, we have AB, AB,
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and we have little a, little b; little a, little b,
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no crossovers.
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So the four haploid cells that are
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going to be produced have just the unrecombined chromosomes
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from the parents, so AB, AB, little a,
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little b; little a, little b.
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These we call parental, because they
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look like these original haploid cells that went into the cross.
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So just like in the fly experiments
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we were talking about, we have parental alleles.
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So not only are they parental, but we actually
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have two kinds of haploid cells.
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One kind is big A, big B. The other kind is little a, little
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b.
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So we're going to call these PDs.
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And what PD stands for is Parental--
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because they're both parental--
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and Ditypes, so ditypes because there are two of them.
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There are two types of offspring,
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and they're both parental, so parental ditypes.
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So let's think about an event that has one crossover.
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So with one crossover, so here again we
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start with the same big A, big B; big A, big B
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to little a, little b; little a, little b.
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But we have one crossover event that's between A and B.
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And so the result of this is four, again, haploid cells.
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One is big A, big B. The other is big A, little
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b because this little b here has gone over to the top.
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The third is little a, big B, and the bottom
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is little a, little b.
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So how many different kinds of haploid cells do we have?
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Four, right.
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They're all different from one another.
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Unlike a ditype before, we have what we call a tetratype.
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And we'll indicate that with a T.
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So there are four different kinds of haploid cells.
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And so tetra, four.
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So we here have two different kinds
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of recognizable conformations of haploid cells.
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We're looking, essentially, not only not at individual cells,
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because there are individual haploid cells like big A,
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big B, and big A, big B that are common in both types,
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but we're looking at the aggregate, what
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all four of them look like together.
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So in one case, one tetrad is a parental ditype tetrad
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and the other is a tetratype tetrad.
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So if genes are really close together,
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we're only going to get single crossovers and no crossovers
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as an output.
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So genes that are tightly linked don't have a lot
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of recombination between them.
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So we really only have two possibilities.
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One is that we have a parental ditype here,
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and the other is that we have a tetratype here,
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because we're assuming that there
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aren't multiple crossovers between multiple alleles.
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So if genes are close together, we only
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have parental ditypes and tetratypes.
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All right, so let's think about distance here.
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And so our formula, our recombination formula
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for genetic distance in all organisms is centimorgans,
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or the distance and centimorgans equals 100 times the number
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of crossover gametes over the number of total gametes.
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This is true in yeast, it's true Drosophila,
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it's true in people.
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So what is the number of crossover gametes in a T
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tetrad?
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So in this kind of tetrad here at the bottom,
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how many crossover gametes do we have?
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Right, we have two of them.
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One of them is there, big A, little b, and the other
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is little a, big B. So in a T tetrad,
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we have two crossover gametes.
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So we're going to come up with a term, epsilon,
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which is essentially the number of total tetrads.
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And 4E equals the number of total gametes,
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because there are four gametes, there
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are four haploid cells per tetrad.
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So it's the number of tetrads times four.
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So here we can say that centimorgans equals
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100 times 2T, so two crossover gametes, times the number of T
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tetrads over 4E, which is the number of total gametes,
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equals 100 times T over 2E.
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So this is the genetic distance for tightly linked genes
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in yeast.
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And so again, here we're looking at genetic distance
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as a function of tetrad type.
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So above this formula, everything
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is referred to individual gametes.
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Down here, we're looking at it in terms
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of the kinds of tetrads that you're actually seeing.
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So we're now thinking about recombination
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in terms of a total tetrad, as opposed to individual gametes.
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And that will become important when we're looking now
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at double crossovers.
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