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These are the user uploaded subtitles that are being translated: 0 00:00:01,060 --> 00:00:02,850 PROFESSOR: What's an enzyme? 1 00:00:02,850 --> 00:00:04,000 How does it work? 2 00:00:04,000 --> 00:00:07,450 Well, it's time to really talk about what an enzyme does. 3 00:00:07,450 --> 00:00:11,700 And so we're going to meet one particular enzyme. 4 00:00:11,700 --> 00:00:12,950 And that enzyme-- 5 00:00:15,200 --> 00:00:16,450 section two-- 6 00:00:19,570 --> 00:00:37,640 this enzyme is called triose phosphate isomerase. 7 00:00:44,610 --> 00:00:49,880 Triose phosphate isomerase, TIM amongst friends. 8 00:00:49,880 --> 00:00:50,100 OK? 9 00:00:50,100 --> 00:00:51,730 So we call this enzyme TIM. 10 00:00:51,730 --> 00:00:54,860 Many people do who work on it just refer to it as TIM. 11 00:00:54,860 --> 00:00:58,340 Now, we talked about oses, we talked about sugars. 12 00:00:58,340 --> 00:01:02,020 We said that six carbon sugars were hexoses. 13 00:01:02,020 --> 00:01:05,050 Five carbon sugars were pentoses. 14 00:01:05,050 --> 00:01:08,500 Three carbon sugars are trioses. 15 00:01:08,500 --> 00:01:12,130 So this is going to be a three carbon sugar that has a 16 00:01:12,130 --> 00:01:13,180 phosphate group on it. 17 00:01:13,180 --> 00:01:15,570 That's what a triose phosphate is. 18 00:01:15,570 --> 00:01:16,640 Pretty straightforward. 19 00:01:16,640 --> 00:01:20,030 Actually, most of this stuff makes sense when you get down to it. 20 00:01:20,030 --> 00:01:26,990 But this enzyme is going to carry out a transformation on one triose 21 00:01:26,990 --> 00:01:31,760 phosphate and turn it into another triose phosphate. 22 00:01:31,760 --> 00:01:34,380 It's going to turn it into an isomer. 23 00:01:34,380 --> 00:01:36,640 Same atoms, but rearranged differently. 24 00:01:36,640 --> 00:01:38,150 That's what an isomer is. 25 00:01:38,150 --> 00:01:41,960 So it's going to take one triose with a phosphate on it and turn it into 26 00:01:41,960 --> 00:01:44,600 another triose with a phosphate on it. 27 00:01:44,600 --> 00:01:46,210 Sounds easy. 28 00:01:46,210 --> 00:01:47,460 Let's take a look. 29 00:01:50,380 --> 00:01:51,930 So triose-- 30 00:01:51,930 --> 00:01:55,940 here is one triose. 31 00:01:55,940 --> 00:02:01,550 Remember that I need to have three carbons and-- 32 00:02:01,550 --> 00:02:03,250 here we go. 33 00:02:03,250 --> 00:02:05,720 I've got a hydroxyl here. 34 00:02:05,720 --> 00:02:08,075 I've got there. 35 00:02:11,680 --> 00:02:16,400 And instead of having another hydroxyl here-- remember, sugar had one 36 00:02:16,400 --> 00:02:18,510 carbonyl, hydroxyl, hydroxyl. 37 00:02:18,510 --> 00:02:22,130 But here I've now, instead, got my phosphate. 38 00:02:27,060 --> 00:02:31,640 That is a particular triose with a phosphate on it. 39 00:02:31,640 --> 00:02:42,220 And this guy goes by the name glyceraldehyde 3-phosphate. 40 00:02:49,070 --> 00:02:52,130 Because it's on the third carbon that that phosphate is. 41 00:02:52,130 --> 00:02:56,000 It's glyceraldehyde, and 'cause that's a lot to say, it's just G3P. 42 00:02:58,520 --> 00:03:00,420 So we've got G3P here. 43 00:03:00,420 --> 00:03:01,470 All right? 44 00:03:01,470 --> 00:03:04,060 Now I'm going to give you another triose. 45 00:03:04,060 --> 00:03:06,050 Let's take this triose here. 46 00:03:06,050 --> 00:03:08,380 Got to have three carbons. 47 00:03:08,380 --> 00:03:11,120 And we know we're going to have a phosphate on it, so I put that 48 00:03:11,120 --> 00:03:12,370 phosphate on. 49 00:03:16,370 --> 00:03:17,650 How is it different? 50 00:03:17,650 --> 00:03:32,360 Well, it's different here because I've got my double bond here, and I've got 51 00:03:32,360 --> 00:03:33,400 a hydroxyl there. 52 00:03:33,400 --> 00:03:37,720 Whereas here I had my double bond here and hydroxyl here. 53 00:03:37,720 --> 00:03:39,690 Not a big difference, right? 54 00:03:39,690 --> 00:03:40,610 All right. 55 00:03:40,610 --> 00:03:51,530 This guy is called dihydroxyacetone phosphate. 56 00:03:51,530 --> 00:03:56,050 And I don't care if you know what all those words mean, but 57 00:03:56,050 --> 00:03:57,190 that's what it's called. 58 00:03:57,190 --> 00:04:01,090 And I'm not expecting you to know these chemical structures here. 59 00:04:01,090 --> 00:04:03,680 You're going to know how to name chemical structures, and we're just 60 00:04:03,680 --> 00:04:07,380 going to call DHAP, dihydroxyacetone phosphate. 61 00:04:07,380 --> 00:04:10,620 So I have a triose phosphate, and I have another triose phosphate. 62 00:04:10,620 --> 00:04:16,310 What do I need to do to convert this triose phosphate into this one? 63 00:04:16,310 --> 00:04:24,380 Oh, by the way, this reaction is going to have to go on in fermentation. 64 00:04:24,380 --> 00:04:27,590 And when we learn about how fermentation works, this is one of 65 00:04:27,590 --> 00:04:28,510 those reactions. 66 00:04:28,510 --> 00:04:34,430 So this is one of the things that Buchner squeezed out of yeast, is an 67 00:04:34,430 --> 00:04:38,160 enzyme that can carry out this particular transformation. 68 00:04:38,160 --> 00:04:39,740 So what do I got to do? 69 00:04:45,120 --> 00:04:45,490 Let's see. 70 00:04:45,490 --> 00:04:46,990 I'm going to-- 71 00:04:46,990 --> 00:04:49,440 STUDENT: Move the H? 72 00:04:49,440 --> 00:04:50,740 PROFESSOR: Proton up here. 73 00:04:50,740 --> 00:04:51,570 The H up here. 74 00:04:51,570 --> 00:04:54,510 So when I'm moving a hydrogen, I'm just moving a proton, because that's 75 00:04:54,510 --> 00:04:54,920 all the nucleus-- 76 00:04:54,920 --> 00:04:56,610 I'm going to move the H up here. 77 00:04:56,610 --> 00:04:58,060 STUDENT: And take an electron down there. 78 00:04:58,060 --> 00:04:59,740 PROFESSOR: Well, I'm going to actually take an H up here. 79 00:04:59,740 --> 00:05:02,500 So basically, I'm just going to put the H here and the H here, and then 80 00:05:02,500 --> 00:05:05,970 I'm going to get the C double bond-- somebody said you've 81 00:05:05,970 --> 00:05:06,820 got to move two Hs. 82 00:05:06,820 --> 00:05:07,610 This can't be that hard. 83 00:05:07,610 --> 00:05:09,380 H's don't weigh very much, right? 84 00:05:09,380 --> 00:05:10,660 So I'm just going to move these Hs. 85 00:05:10,660 --> 00:05:12,700 Sounds pretty easy. 86 00:05:12,700 --> 00:05:14,530 Well. 87 00:05:14,530 --> 00:05:17,250 And it actually turns out to be chemically pretty 88 00:05:17,250 --> 00:05:20,900 favorable to do this. 89 00:05:20,900 --> 00:05:23,790 Chemists can measure that with something called the free energy. 90 00:05:26,990 --> 00:05:30,810 Whether one structure is energetically more or less favorable 91 00:05:30,810 --> 00:05:32,590 than another structure. 92 00:05:32,590 --> 00:05:43,360 So the free energy of G3P and the free energy here of DHAP-- 93 00:05:43,360 --> 00:05:47,250 DHAP is actually more energetically favorable. 94 00:05:47,250 --> 00:05:49,000 It's a lower energy state. 95 00:05:49,000 --> 00:05:50,810 It's a more favorable state. 96 00:05:50,810 --> 00:05:53,140 It's actually lower by-- 97 00:05:53,140 --> 00:05:58,630 we write delta G zero prime, which is a measure of free energy. 98 00:05:58,630 --> 00:06:00,880 And it's better. 99 00:06:00,880 --> 00:06:07,090 On its own, if you ask these two molecules, this is an energetically 100 00:06:07,090 --> 00:06:08,620 more favorable state. 101 00:06:08,620 --> 00:06:13,220 So why doesn't G3P just become DHAP spontaneously? 102 00:06:13,220 --> 00:06:14,470 STUDENT: [INAUDIBLE]. 103 00:06:17,090 --> 00:06:17,560 PROFESSOR: Sorry? 104 00:06:17,560 --> 00:06:18,370 STUDENT: Activation energy. 105 00:06:18,370 --> 00:06:19,310 PROFESSOR: Activation-- 106 00:06:19,310 --> 00:06:21,900 what's activation energy? 107 00:06:21,900 --> 00:06:24,170 I mean, we say, because there's an activation barrier. 108 00:06:24,170 --> 00:06:25,840 There's an activation energy. 109 00:06:25,840 --> 00:06:29,422 What does it mean to say there's an activation energy? 110 00:06:29,422 --> 00:06:30,730 STUDENT: You need energy to start the reaction. 111 00:06:30,730 --> 00:06:32,670 PROFESSOR: I need energy, so I just like light a match? 112 00:06:36,030 --> 00:06:38,910 [LAUGHTER] 113 00:06:38,910 --> 00:06:45,380 STUDENT: Wouldn't you need to invest some energy into breaking the bonds in 114 00:06:45,380 --> 00:06:48,270 G3P so that you can move the hydrogen [INAUDIBLE]? 115 00:06:48,270 --> 00:06:50,110 PROFESSOR: So why do I have to invest energy? 116 00:06:50,110 --> 00:06:53,080 People always tell you to start investing, but you know, you're young. 117 00:06:53,080 --> 00:06:55,840 Why you need to start investing here, all this energy? 118 00:06:55,840 --> 00:06:57,870 What's the point? 119 00:06:57,870 --> 00:06:58,745 Yeah? 120 00:06:58,745 --> 00:07:01,720 STUDENT: You'll get more energy back when the bonds reform another one. 121 00:07:01,720 --> 00:07:03,010 PROFESSOR: I'll get more energy back. 122 00:07:03,010 --> 00:07:04,760 That's what they always tell you about why you should invest, because you can 123 00:07:04,760 --> 00:07:05,570 get more back. 124 00:07:05,570 --> 00:07:05,890 [LAUGHTER] 125 00:07:05,890 --> 00:07:09,100 And then the molecule pays some interest rate. 126 00:07:09,100 --> 00:07:11,600 But what's going on? 127 00:07:11,600 --> 00:07:11,740 Yeah? 128 00:07:11,740 --> 00:07:16,190 STUDENT: You need to invest energy so the reaction favors the product. 129 00:07:16,190 --> 00:07:17,840 PROFESSOR: That actually favors the product right now. 130 00:07:17,840 --> 00:07:20,050 I gave you a reaction that favors that product. 131 00:07:20,050 --> 00:07:22,710 It actually would rather be DHAP than G3P. 132 00:07:22,710 --> 00:07:25,080 It's going in the right direction. 133 00:07:25,080 --> 00:07:26,030 Better free energy. 134 00:07:26,030 --> 00:07:28,005 So it's already doing that. 135 00:07:28,005 --> 00:07:29,280 I don't need any energy. 136 00:07:29,280 --> 00:07:30,322 Yep? 137 00:07:30,322 --> 00:07:34,706 STUDENT: So globally, the product has a lower free energy 138 00:07:34,706 --> 00:07:35,890 state than the reactant-- 139 00:07:35,890 --> 00:07:36,876 PROFESSOR: It does. 140 00:07:36,876 --> 00:07:44,822 STUDENT: --but locally, the reactant is in a lower free energy state than 141 00:07:44,822 --> 00:07:45,730 in transition. 142 00:07:45,730 --> 00:07:46,600 PROFESSOR: Transition. 143 00:07:46,600 --> 00:07:49,000 Tell me about this transition. 144 00:07:49,000 --> 00:07:53,860 You're saying that to get from here to there, from one molecule to another 145 00:07:53,860 --> 00:07:57,180 molecule, how do I get there? 146 00:07:57,180 --> 00:08:00,960 I have to pass through some intermediate state, 147 00:08:00,960 --> 00:08:03,370 some transition state. 148 00:08:03,370 --> 00:08:11,770 In fact, to get from here to there, from here to here, it turns out I have 149 00:08:11,770 --> 00:08:15,450 to pass through this state. 150 00:08:19,810 --> 00:08:27,020 Chemically, I'm not going to go into the details of why, but it turns out 151 00:08:27,020 --> 00:08:39,380 that in order to do this, I need to transition through this guy. 152 00:08:44,210 --> 00:08:45,570 Let's take a look at what's going on. 153 00:08:45,570 --> 00:08:49,200 Everything below this line is the same. 154 00:08:49,200 --> 00:08:55,210 But above this line, while I'm doing some moving around here, this is a 155 00:08:55,210 --> 00:08:56,880 very different molecule. 156 00:08:56,880 --> 00:08:58,970 I have this double bond here. 157 00:08:58,970 --> 00:09:01,710 And this guy goes by the funky name cis-enediol. 158 00:09:06,110 --> 00:09:11,410 And what it is is this is what's called the transition state. 159 00:09:11,410 --> 00:09:24,270 So cis-enediol is the transition state, or the intermediate. 160 00:09:24,270 --> 00:09:28,250 And it is a very unhappy molecule. 161 00:09:28,250 --> 00:09:31,460 It has very high free energy. 162 00:09:31,460 --> 00:09:33,740 It is not a happy state. 163 00:09:33,740 --> 00:09:39,785 In fact, it is there. 164 00:09:43,770 --> 00:09:49,210 So it turns out that the reason that this reaction is not going forward is 165 00:09:49,210 --> 00:09:51,320 not that it wouldn't like to be DHAP. 166 00:09:51,320 --> 00:09:52,600 It would love to be DHAP. 167 00:09:52,600 --> 00:09:55,940 You appreciate, I'm anthropomorphizing when I speak about the desires of 168 00:09:55,940 --> 00:09:58,100 relatively small molecules, right? 169 00:09:58,100 --> 00:10:02,970 But nonetheless, it is energetically more favorable to be DHAP. 170 00:10:02,970 --> 00:10:06,340 But to get from here to there, you have to climb a mountain. 171 00:10:06,340 --> 00:10:09,480 You have to go through an intermediate molecular state, which is 172 00:10:09,480 --> 00:10:12,730 extraordinarily unfavorable. 173 00:10:12,730 --> 00:10:17,050 The chance of this happening on its own is roughly nil. 174 00:10:17,050 --> 00:10:24,020 Because this is something like 26 kilocalories per mole, which is a lot 175 00:10:24,020 --> 00:10:26,260 of energy needed. 176 00:10:26,260 --> 00:10:29,700 Very rare that by chance the molecules are just going to 177 00:10:29,700 --> 00:10:31,010 happen to go do that. 178 00:10:31,010 --> 00:10:34,540 And the truth is, it's actually worse than that. 179 00:10:34,540 --> 00:10:40,030 In this state, not only is this a high energy state, but rather than going 180 00:10:40,030 --> 00:10:45,280 from here to there, this molecule is so unhappy that its phosphate will 181 00:10:45,280 --> 00:10:46,840 usually break off. 182 00:10:46,840 --> 00:10:49,150 Especially in the presence of water. 183 00:10:49,150 --> 00:10:55,700 So the chance of going from here to here, very, very low. 184 00:10:55,700 --> 00:10:58,900 And then the chance of managing to keep going without losing the 185 00:10:58,900 --> 00:11:01,190 phosphate, forget it. 186 00:11:01,190 --> 00:11:04,780 This is just not going to happen on its own. 187 00:11:04,780 --> 00:11:07,000 You need to help it. 188 00:11:07,000 --> 00:11:09,180 That is what an enzyme does. 189 00:11:09,180 --> 00:11:11,830 Turns out you don't have to invest any energy at all. 190 00:11:11,830 --> 00:11:14,490 No energy is needed to make this happen. 191 00:11:14,490 --> 00:11:16,200 I didn't have to spend any energy. 192 00:11:16,200 --> 00:11:19,450 I don't have to invest any energy, and I don't have to get back any energy. 193 00:11:19,450 --> 00:11:21,180 None of that is needed. 194 00:11:21,180 --> 00:11:25,120 Because in fact, it's an energetically favorable reaction. 195 00:11:25,120 --> 00:11:28,840 It just has to go through some intermediate state. 196 00:11:28,840 --> 00:11:33,770 And we have to gently help it over the mountain. 197 00:11:33,770 --> 00:11:36,520 Now that we've finished this segment, test yourself with this question. 14957

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