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Would you like to inspect the original subtitles? These are the user uploaded subtitles that are being translated: 0 00:00:00,000 --> 00:00:01,500 1 00:00:01,500 --> 00:00:06,450 MICHAEL HEMANN: OK, so let's think about two linked genes. 2 00:00:06,450 --> 00:00:10,080 In this case, two very tightly linked genes. 3 00:00:10,080 --> 00:00:16,740 4 00:00:16,740 --> 00:00:19,440 One of them is A, and one of them 5 00:00:19,440 --> 00:00:26,685 is B. So if we have a yeast haploid strain that is AB-- 6 00:00:26,685 --> 00:00:30,060 big A, big B-- we can cross it with another strain that's 7 00:00:30,060 --> 00:00:31,440 a, little b. 8 00:00:31,440 --> 00:00:38,660 We can generate a diploid that is AB over ab. 9 00:00:38,660 --> 00:00:41,220 Here we're using, again, our linkage designation. 10 00:00:41,220 --> 00:00:44,240 So this line says they're linked together 11 00:00:44,240 --> 00:00:45,890 on the same chromosome. 12 00:00:45,890 --> 00:00:49,670 This, of course, is a diploid. 13 00:00:49,670 --> 00:00:53,510 And we can induce it to undergo meiosis, 14 00:00:53,510 --> 00:00:58,410 and we'll get a tetrad out of this. 15 00:00:58,410 --> 00:01:00,030 So let's think about two events. 16 00:01:00,030 --> 00:01:02,210 One is where we have a meiosis where 17 00:01:02,210 --> 00:01:06,230 we have basically no crossovers between A and B 18 00:01:06,230 --> 00:01:08,090 on the same chromosome, and another 19 00:01:08,090 --> 00:01:12,650 where we have a meiosis where we have one crossover. 20 00:01:12,650 --> 00:01:26,290 So with a meiosis with no crossovers, we have AB, AB, 21 00:01:26,290 --> 00:01:31,660 and we have little a, little b; little a, little b, 22 00:01:31,660 --> 00:01:33,110 no crossovers. 23 00:01:33,110 --> 00:01:35,890 So the four haploid cells that are 24 00:01:35,890 --> 00:01:39,640 going to be produced have just the unrecombined chromosomes 25 00:01:39,640 --> 00:01:47,290 from the parents, so AB, AB, little a, 26 00:01:47,290 --> 00:01:51,040 little b; little a, little b. 27 00:01:51,040 --> 00:01:55,630 These we call parental, because they 28 00:01:55,630 --> 00:02:01,430 look like these original haploid cells that went into the cross. 29 00:02:01,430 --> 00:02:05,990 So just like in the fly experiments 30 00:02:05,990 --> 00:02:08,389 we were talking about, we have parental alleles. 31 00:02:08,389 --> 00:02:11,030 So not only are they parental, but we actually 32 00:02:11,030 --> 00:02:15,770 have two kinds of haploid cells. 33 00:02:15,770 --> 00:02:20,180 One kind is big A, big B. The other kind is little a, little 34 00:02:20,180 --> 00:02:21,080 b. 35 00:02:21,080 --> 00:02:23,900 So we're going to call these PDs. 36 00:02:23,900 --> 00:02:28,410 And what PD stands for is Parental-- 37 00:02:28,410 --> 00:02:30,180 because they're both parental-- 38 00:02:30,180 --> 00:02:37,255 and Ditypes, so ditypes because there are two of them. 39 00:02:37,255 --> 00:02:38,630 There are two types of offspring, 40 00:02:38,630 --> 00:02:41,750 and they're both parental, so parental ditypes. 41 00:02:41,750 --> 00:02:44,390 So let's think about an event that has one crossover. 42 00:02:44,390 --> 00:02:55,070 So with one crossover, so here again we 43 00:02:55,070 --> 00:03:03,280 start with the same big A, big B; big A, big B 44 00:03:03,280 --> 00:03:09,720 to little a, little b; little a, little b. 45 00:03:09,720 --> 00:03:14,850 But we have one crossover event that's between A and B. 46 00:03:14,850 --> 00:03:20,350 And so the result of this is four, again, haploid cells. 47 00:03:20,350 --> 00:03:26,030 One is big A, big B. The other is big A, little 48 00:03:26,030 --> 00:03:30,810 b because this little b here has gone over to the top. 49 00:03:30,810 --> 00:03:39,000 The third is little a, big B, and the bottom 50 00:03:39,000 --> 00:03:41,880 is little a, little b. 51 00:03:41,880 --> 00:03:52,830 So how many different kinds of haploid cells do we have? 52 00:03:52,830 --> 00:03:57,650 53 00:03:57,650 --> 00:03:58,770 Four, right. 54 00:03:58,770 --> 00:04:02,000 They're all different from one another. 55 00:04:02,000 --> 00:04:14,440 Unlike a ditype before, we have what we call a tetratype. 56 00:04:14,440 --> 00:04:16,700 And we'll indicate that with a T. 57 00:04:16,700 --> 00:04:21,360 So there are four different kinds of haploid cells. 58 00:04:21,360 --> 00:04:22,970 And so tetra, four. 59 00:04:22,970 --> 00:04:26,660 So we here have two different kinds 60 00:04:26,660 --> 00:04:31,790 of recognizable conformations of haploid cells. 61 00:04:31,790 --> 00:04:36,680 We're looking, essentially, not only not at individual cells, 62 00:04:36,680 --> 00:04:39,680 because there are individual haploid cells like big A, 63 00:04:39,680 --> 00:04:43,983 big B, and big A, big B that are common in both types, 64 00:04:43,983 --> 00:04:45,650 but we're looking at the aggregate, what 65 00:04:45,650 --> 00:04:47,550 all four of them look like together. 66 00:04:47,550 --> 00:04:51,230 So in one case, one tetrad is a parental ditype tetrad 67 00:04:51,230 --> 00:04:54,830 and the other is a tetratype tetrad. 68 00:04:54,830 --> 00:05:04,150 69 00:05:04,150 --> 00:05:06,910 So if genes are really close together, 70 00:05:06,910 --> 00:05:15,180 we're only going to get single crossovers and no crossovers 71 00:05:15,180 --> 00:05:17,460 as an output. 72 00:05:17,460 --> 00:05:19,680 So genes that are tightly linked don't have a lot 73 00:05:19,680 --> 00:05:21,250 of recombination between them. 74 00:05:21,250 --> 00:05:23,650 So we really only have two possibilities. 75 00:05:23,650 --> 00:05:26,730 One is that we have a parental ditype here, 76 00:05:26,730 --> 00:05:29,735 and the other is that we have a tetratype here, 77 00:05:29,735 --> 00:05:31,110 because we're assuming that there 78 00:05:31,110 --> 00:05:34,410 aren't multiple crossovers between multiple alleles. 79 00:05:34,410 --> 00:05:45,820 So if genes are close together, we only 80 00:05:45,820 --> 00:05:53,800 have parental ditypes and tetratypes. 81 00:05:53,800 --> 00:05:55,940 All right, so let's think about distance here. 82 00:05:55,940 --> 00:05:59,770 And so our formula, our recombination formula 83 00:05:59,770 --> 00:06:04,360 for genetic distance in all organisms is centimorgans, 84 00:06:04,360 --> 00:06:11,200 or the distance and centimorgans equals 100 times the number 85 00:06:11,200 --> 00:06:24,560 of crossover gametes over the number of total gametes. 86 00:06:24,560 --> 00:06:26,640 This is true in yeast, it's true Drosophila, 87 00:06:26,640 --> 00:06:28,700 it's true in people. 88 00:06:28,700 --> 00:06:45,790 So what is the number of crossover gametes in a T 89 00:06:45,790 --> 00:06:48,190 tetrad? 90 00:06:48,190 --> 00:06:53,880 So in this kind of tetrad here at the bottom, 91 00:06:53,880 --> 00:06:55,860 how many crossover gametes do we have? 92 00:06:55,860 --> 00:06:59,630 93 00:06:59,630 --> 00:07:01,380 Right, we have two of them. 94 00:07:01,380 --> 00:07:04,460 One of them is there, big A, little b, and the other 95 00:07:04,460 --> 00:07:12,870 is little a, big B. So in a T tetrad, 96 00:07:12,870 --> 00:07:17,580 we have two crossover gametes. 97 00:07:17,580 --> 00:07:21,120 So we're going to come up with a term, epsilon, 98 00:07:21,120 --> 00:07:29,770 which is essentially the number of total tetrads. 99 00:07:29,770 --> 00:07:39,920 And 4E equals the number of total gametes, 100 00:07:39,920 --> 00:07:42,080 because there are four gametes, there 101 00:07:42,080 --> 00:07:45,000 are four haploid cells per tetrad. 102 00:07:45,000 --> 00:07:48,380 So it's the number of tetrads times four. 103 00:07:48,380 --> 00:07:53,390 So here we can say that centimorgans equals 104 00:07:53,390 --> 00:08:03,110 100 times 2T, so two crossover gametes, times the number of T 105 00:08:03,110 --> 00:08:09,020 tetrads over 4E, which is the number of total gametes, 106 00:08:09,020 --> 00:08:16,480 equals 100 times T over 2E. 107 00:08:16,480 --> 00:08:21,310 So this is the genetic distance for tightly linked genes 108 00:08:21,310 --> 00:08:22,490 in yeast. 109 00:08:22,490 --> 00:08:29,180 And so again, here we're looking at genetic distance 110 00:08:29,180 --> 00:08:32,059 as a function of tetrad type. 111 00:08:32,059 --> 00:08:40,919 112 00:08:40,919 --> 00:08:43,200 So above this formula, everything 113 00:08:43,200 --> 00:08:45,480 is referred to individual gametes. 114 00:08:45,480 --> 00:08:47,580 Down here, we're looking at it in terms 115 00:08:47,580 --> 00:08:50,740 of the kinds of tetrads that you're actually seeing. 116 00:08:50,740 --> 00:08:53,310 So we're now thinking about recombination 117 00:08:53,310 --> 00:08:58,410 in terms of a total tetrad, as opposed to individual gametes. 118 00:08:58,410 --> 00:09:00,720 And that will become important when we're looking now 119 00:09:00,720 --> 00:09:02,540 at double crossovers. 8915

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