Bringing the World's Best Biology to You
00:00:23;24 I’m interested in how living organisms move.
00:00:26;27 It’s a fundamental problem of life.
00:00:29;14 Life is endowed with movement.
00:00:31;28 So, what are some examples of movement
00:00:35;20 that occurs in living systems?
00:00:37;21 Yes?
00:00:39;15 Student: Heart beating. Heart beating — good.
00:00:41;20 What else Come on, fire. Yes?
00:00:44;22 Student: Blood flow. What?
00:00:47;12 Blood flow is a consequence of the heart,
00:00:51;26 so it’s kind of a consequence rather than a fundamental action itself.
00:00:58;00 In the very back? Student: Muscle contraction.
00:01:00;09 Muscle contraction, okay, that’s…
00:01:02;05 these are obvious ones.
00:01:03;25 Now, let’s think outside of the whole animal
00:01:05;24 to smaller things. Yes?
00:01:08;07 Student: Peristalsis. Peristalsis?
00:01:12;01 Okay, so that’s another kind of muscle.
00:01:14;01 It’s called smooth muscle.
00:01:15;27 It’s kind of what makes your GI system work.
00:01:19;22 So, now we’ve covered all muscles:
00:01:21;14 cardiac muscle, skeletal muscle, smooth muscle.
00:01:25;09 Let’s start thinking at a cellular level. Yes? Student: Vesicle transport.
00:01:29;18 Ooo, yes, that’s my favorite. Okay.
00:01:32;06 Haha, because that’s what I work on.
00:01:33;29 So, vesicle transport in a cell.
00:01:35;22 Student: Cells moving with either cilia or...
00:01:37;25 Okay, so cilia and flagella.
00:01:40;00 So, we can think of sperm swimming.
00:01:42;13 We can think of a paramecium in pond water swimming.
00:01:47;17 We can think of the cilia in your bronchial tracts
00:01:52;02 that are involved in pushing things out of your lungs.
00:01:56;06 So, that’s pretty good. I think we’ve covered a lot.
00:01:58;27 So, let’s just have a look at some of that
00:02:02;00 and just see what they look like.
00:02:03;21 I know you’ve seen them in textbooks, but…
00:02:08;04 okay, so, here’s one thing, which is just a cell moving.
00:02:11;24 This is a white blood cell in your body — really amazing.
00:02:14;21 So, it’s moving.
00:02:16;17 Second thing is it’s chasing that little thing in front of it,
00:02:19;08 which is a bacteria.
00:02:21;05 And look how smart the cell is.
00:02:23;06 The bacteria is trying to, like, get away,
00:02:25;22 but the cell actually not only can move;
00:02:27;21 it can sense its location until eventually… boom.
00:02:31;17 It engulfed it.
00:02:33;29 So, that just shows you how smart cells are
00:02:36;22 but also illustrates the phenomena
00:02:39;27 of cell movement.
00:02:41;15 Here is cell division.
00:02:43;03 So, you learned all the stages of mitosis.
00:02:45;02 What are these threads here?
00:02:47;12 Chromosomes. Okay.
00:02:48;28 So, here, the cell is going to metaphase.
00:02:50;11 You learned about that. So, on the right side, there’s some…
00:02:53;18 see all that jiggling going on?
00:02:55;13 The chromosomes are moving back and forth.
00:02:57;03 They’re actually trying to be moved, physically moved,
00:03:00;05 to the metaphase plate.
00:03:01;26 It doesn’t just happen.
00:03:03;12 There’s actually physical motion that’s required
00:03:05;27 to move chromosomes.
00:03:07;09 And now, actually — another amazing thing —
00:03:08;20 the cell senses when all the chromosomes are there
00:03:11;04 and then another physical event happens.
00:03:13;18 The e… the sister chromatids separate,
00:03:17;07 and there’s a physical motion of them apart,
00:03:19;20 followed by another physical motion,
00:03:21;10 which is a constriction of the cell in two.
00:03:24;06 So, there’s all kinds of movement that’s going on there
00:03:27;03 during cell division.
00:03:28;18 So, that’s happening in all of the dividing cells of your body
00:03:31;14 right now.
00:03:33;08 It’s obviously… too much of that is what goes wrong in cancer.
00:03:38;11 Now let’s look at vesicle motion,
00:03:42;14 which one of the students mentioned.
00:03:44;13 So, this is actually looking inside of a nerve axon.
00:03:47;01 Just look at it. I mean, there’s…
00:03:49;02 these little dots are membrane organelles, vesicles.
00:03:51;26 You can see all kinds of movement going up and down.
00:03:56;16 Do you know what these big, elongated things are?
00:04:00;29 You want to have a guess?
00:04:04;04 It’s kind of choking on this now.
00:04:06;06 What do you think those big, elongated things are?
00:04:08;13 Nah, that doesn’t count. No teachers allowed.
00:04:12;24 What might be a big, elongated thing inside of a cell?
00:04:19;15 Take a guess.
00:04:22;08 You should not be afraid of uttering the wrong answer.
00:04:25;11 I love talking to elementary school kids
00:04:27;11 because they just will always raise their hand for any question,
00:04:30;19 and anything is fair game to come out of their mouth.
00:04:35;12 So, you should try to maintain that same kind of spirit.
00:04:39;18 Yeah? Student: Mitochondria. Mito… did you say mitochondria?
00:04:42;25 Okay. Those are mitochondria.
00:04:47;03 So, those are mitochondria moving inside of the cell.
00:04:50;18 Okay. So, you know, why is there all that movement in the cell?
00:04:53;19 I mean, maybe this is a bit of a corny video,
00:04:55;29 but… you know, we make things in certain parts of the city
00:04:59;19 and then we have to distribute them so that…
00:05:02;12 you know, things are made in some places.
00:05:04;24 They’re distributed in cargo containers
00:05:07;27 that are transported by engines
00:05:11;21 that move things around in a city,
00:05:14;08 and the cell is pretty much the same way.
00:05:17;09 There are places where things are made
00:05:19;04 inside of cells,
00:05:20;28 and then they have to be transported on roadways.
00:05:22;11 So, that’s what I’ll tell you about.
00:05:23;29 But… this just shows a blowup on…
00:05:26;28 of one of those roadways.
00:05:28;09 It’s called a microtubule.
00:05:30;13 It shows some cargo being carried,
00:05:33;15 and it shows one of these motor proteins,
00:05:35;26 called kinesin, moving along that microtubule.
00:05:38;08 Now, let me just tell you how I started this problem.
00:05:42;00 I was a graduate student.
00:05:43;28 I was actually a neuroscience student,
00:05:46;01 so initially I wasn’t interested in this.
00:05:48;15 I was, like, interested in how the nervous system develops,
00:05:50;26 how neurodegeneradiv…
00:05:54;02 neurodegenerative diseases are produced.
00:05:57;20 And the reason why it’s so interesting…
00:05:59;15 the reason I got connected into transport is that neurons have…
00:06:05;15 for all these problems, for neurodegeneration or development…
00:06:10;06 they’re extremely unusual cells
00:06:12;15 and rely very heavily on intracellular transport.
00:06:17;03 The reason is that the neurons are the longest cells in your body
00:06:20;22 by far.
00:06:22;12 Okay? So, for example,
00:06:24;19 if you are using a muscle in your toe,
00:06:28;05 that muscle in your toe is actually being controlled
00:06:30;10 by a neuron that starts in your spinal cord.
00:06:33;19 Okay?
00:06:35;13 Now, the part that’s in your spinal cord is the part that has the nucleus, the DNA.
00:06:38;16 It has all the synthetic machinery,
00:06:40;15 like ribosomes and all of that.
00:06:42;16 And then… so, that’s the part…
00:06:45;14 that cell body is in your spinal cord.
00:06:47;02 But then it has this very, very long tube,
00:06:49;08 which is also called a… axon. Okay?
00:06:55;21 Now, the axon is sending a…
00:06:57;16 what does the axon do, that you probably know about?
00:06:59;16 Action potential.
00:07:01;07 So, it’s firing down an electrical signal
00:07:03;07 to stimulate your muscle.
00:07:05;20 But think about it… all the things that are at the very end of that nerve cell,
00:07:09;25 like all your neurotransmitters and all of that,
00:07:12;22 they’re not made there.
00:07:14;07 They’re not made by your muscle.
00:07:15;18 They’re actually made near your spinal cord,
00:07:17;28 so they have to be shipped
00:07:20;26 from your spinal cord to the very end.
00:07:23;14 Now, I’m gonna give you an idea of the scale of that transport.
00:07:26;11 I’m gonna now blow up a 20-micron cell biol…
00:07:30;14 cell body that’s normally in your spinal cord,
00:07:32;21 and I’m gonna create a monster cell body
00:07:34;19 that is about six feet.
00:07:38;28 So, a little smaller than this table, okay?
00:07:42;11 And I’m gonna scale everything up proportionately,
00:07:44;21 and I’m gonna put this six-foot cell body in Newark, okay?
00:07:49;01 The axon, that tube that comes out of that six-foot diameter cell body,
00:07:55;28 would go all the way from Newark halfway down Long Island.
00:07:59;09 Okay?
00:08:02;16 So, you can imagine the transport that has to go on to get material
00:08:07;29 that’s delivered in Newark from this six-foot, you know, factory
00:08:11;22 all the way down to Long Island.
00:08:13;21 There has to be a pretty good transport system.
00:08:16;05 So, that’s… and when I started as a student,
00:08:18;28 we knew there was a transport system.
00:08:21;02 We didn’t know how it worked.
00:08:23;25 So, by analogy, there have to be roadways —
00:08:25;28 we didn’t know what the roadways were.
00:08:27;15 There have to be motors —
00:08:29;03 we didn’t know what the motors were.
00:08:31;07 We kind of knew what the cargos were.
00:08:32;21 Someone mentioned vesicle transport. Okay.
00:08:35;14 So, we knew there had to be vesicles going down there.
00:08:39;10 And there have to be signals controlling all this,
00:08:41;06 because you can’t have this chaotic transport system.
00:08:45;06 So, we didn’t know any of these questions,
00:08:46;27 so I’m gonna just focus on the motor.
00:08:49;03 I want to just really relate in your head
00:08:50;29 the size of these things we’re talking about.
00:08:52;27 Because, you know, everyone talks about nanotechnology.
00:08:55;16 It’s all cool now.
00:08:57;09 But everything we talk about with nanotechnology
00:08:59;20 is mostly micron-scale technology.
00:09:02;15 Nano means what? What is a nanometer?
00:09:06;23 Students: 10 to the negative 9th. 10 to the negative 9th, okay?
00:09:10;06 So, you know... that iPod is not,
00:09:14;17 you know, 10 to the negative 9th.
00:09:16;23 Okay.
00:09:18;18 These motors are pretty darn close to 10 to the negative 9th.
00:09:20;28 They’re about… the motor part is about 10 to the minus 8th meters.
00:09:24;23 That’s about a millionth of an inch in size.
00:09:29;00 So, we have to study things that are that small.
00:09:32;21 The other amazing thing is…
00:09:35;00 also, think about engineering.
00:09:37;06 Those of you who are engineers…
00:09:39;22 our cars that we put out on the road burn hydrocarbons,
00:09:43;00 convert them into motion.
00:09:45;14 These biological motors use ATP, adenosine triphosphate.
00:09:48;12 Look how much better they are than your car.
00:09:51;14 So, your car is pretty terrible in converting…
00:09:54;27 like, the theoretical energy derived from breaking down hydrocarbons
00:09:59;14 into useful work.
00:10:01;05 And the motors work at,
00:10:03;05 you know, maybe six-fold better efficiency.
00:10:05;26 So, in fact, engineers are actually quite interested
00:10:09;13 in understanding how biological systems
00:10:11;22 that are created through evolution…
00:10:13;15 how they do the things that they’re doing.
00:10:15;13 The things we’re talking about are proteins
00:10:17;16 made by the genome.
00:10:19;03 The genome basically doesn’t do much in biology,
00:10:22;09 in terms of the actual work of biology.
00:10:24;15 It’s just the instruction book
00:10:27;22 that is churning out and making these proteins,
00:10:29;14 which are acting as machines.
00:10:31;27 So, you know, a typical protein
00:10:34;06 may have about 500 amino acids strung together.
00:10:37;13 My favorite protein, the motor protein kinesin,
00:10:39;27 has about 960 amino acids.
00:10:42;22 And those 960 amino acids come together
00:10:46;07 to create some kind of machine
00:10:48;28 that’s able to act as an engine.
00:10:50;22 Okay? So, just to get that…
00:10:53;05 so, just one minute…
00:10:55;28 think of some other protein machines.
00:10:57;17 Just talk to your neighbor for one minute.
00:10:59;04 Just try… and if you can’t name another protein,
00:11:01;22 think of a specific activity, a specific function,
00:11:05;16 that a protein is doing in biology, okay?
00:11:08;09 So, just talk to your neighbor, see what you can come up with.
00:11:19;17 Okay. Maybe time’s up.
00:11:21;29 Really quick… I can’t go through everyone.
00:11:23;21 Let’s just name a few quick examples.
00:11:25;23 Student: Hemoglobin. Shout it out. Student: Hemoglobin.
00:11:28;10 What does hemoglobin do? Student: It stores oxygen in the blood.
00:11:31;23 Carries oxygen in red blood cells.
00:11:34;07 What… did you guys come up with something?
00:11:36;28 Student: I think we did enzymes.
00:11:38;18 Enzymes are a big case.
00:11:41;21 So, can… a motor protein is an enzyme.
00:11:43;22 Can you think of a specific enzyme, other than a motor protein?
00:11:46;24 Student: DNA helicase. DNA helicase?
00:11:49;15 Is that what you said?
00:11:50;24 Sure, that’s a great thing.
00:11:52;04 What does… do you know what DNA helicase does?
00:11:57;05 Yeah, okay, it unwinds DNA,
00:12:00;07 for example during replication.
00:12:02;23 Okay, one more example. Anyone, anyone in the back?
00:12:05;08 Example of a protein. Yes.
00:12:06;29 Student: Amylase? Amylase. What does amylase do?
00:12:09;16 Student: It breaks down carbohydrates.
00:12:12;22 It breaks down carbohydrates, so that’s an enzyme
00:12:14;23 that’s found in your intestinal tract to break down food.
00:12:16;21 So, I’m gonna tell you a little bit about how I discovered
00:12:19;17 a new kind of protein, okay?,
00:12:21;01 because this also gives you an idea of how discoveries are made.
00:12:23;06 If you want to, like, really understand something,
00:12:24;29 you have to build it. Right?
00:12:27;09 That’s like… an engineer, you know, would approach things that way —
00:12:30;05 or an auto mechanic, right?
00:12:31;29 If you want to become a really good auto mechanic,
00:12:35;03 which I’m not…
00:12:36;25 if you can take the car engine apart, you know, all the pieces,
00:12:39;24 lie ‘em out on a big table, and put ‘em all back together again,
00:12:42;19 then you can actually understand how that car works.
00:12:46;12 So, that’s the general philosophy.
00:12:48;04 So, here’s the general experiment.
00:12:50;03 We started with this system.
00:12:51;26 We’re looking at a nerve cell
00:12:54;12 with a fancy microscope that has computers, video cameras, all of that.
00:12:57;22 And we can see all kinds of stuff happening,
00:13:00;06 but we don’t know what the roadways are,
00:13:02;08 we don’t know what the engines are,
00:13:04;05 so how do you take that apart?
00:13:06;12 Okay.
00:13:09;17 We used… to study this, instead of studying it in people,
00:13:11;23 because most people don’t want to donate their nerves…
00:13:15;25 and actually, studying it in mice,
00:13:17;25 you could do it,
00:13:19;16 but it turns out that squid were a much better system
00:13:21;19 for studying this problem.
00:13:23;24 The reason is the squid has an axon
00:13:25;28 that’s 50 to 100 times bigger
00:13:28;13 than a human axon. Okay?
00:13:31;07 So, you can easily dissect it out of the squid,
00:13:33;18 and believe it or not,
00:13:35;10 you can actually squeeze out the inside of the axon of a squid
00:13:38;24 just like toothpaste. It’s really amazing.
00:13:40;25 And out comes this little line of the inside of a cell
00:13:45;18 that, you know, you could put on a glass slide.
00:13:48;06 And that’s… and then…
00:13:51;05 this experiment was squeezing it out,
00:13:53;07 putting it on a glass slide,
00:13:55;26 and literally mushing it up with some buffer, some solution,
00:14:00;05 and a pipette tip.
00:14:01;22 You know, like one of these yellow tips. You probably use them in class?
00:14:04;15 Anyway, we mushed it up
00:14:07;06 and then watched it under the microscope.
00:14:09;05 And we learned some things,
00:14:11;04 because now we can see… in this mushed-up thing,
00:14:13;02 what can you see there that you couldn’t see before?
00:14:16;01 Huh? Student: Roads. Like what?
00:14:19;04 Student: The roads, like…
00:14:21;10 So, you can see something that looks like roadways,
00:14:24;04 some really discrete things, now,
00:14:25;06 that look like roadways.
00:14:27;17 And this alone got rid of a lot of other theories
00:14:29;18 about how this transport system worked.
00:14:31;15 There were clear roadways.
00:14:33;27 We actually discovered what the roadways were
00:14:35;25 using an electron microscope.
00:14:37;13 They were… turned out to be microtubules,
00:14:38;25 but we didn’t know what the motor was.
00:14:40;26 So, here’s the experiment to find out
00:14:43;02 what the motor was.
00:14:44;25 And that was this idea of taking everything apart
00:14:47;27 and trying to put it back together.
00:14:49;15 So, we knew what the roadways were.
00:14:51;05 They were these things called microtubules.
00:14:52;20 We could purify them, add ‘em into a test tube.
00:14:55;21 We needed cargo.
00:14:57;15 So, we would thought…
00:14:59;11 we thought, okay, we can get mitochondria, stuff like that,
00:15:02;02 and it should move along the roadways,
00:15:05;19 and the motors should be there.
00:15:07;06 As long as we gave it some energy source, like ATP,
00:15:10;23 which we could fortunately buy.
00:15:13;02 But… bummer. It didn’t work.
00:15:15;27 You know, so… we thought for sure this experiment
00:15:19;04 was gonna work.
00:15:20;21 But two things about science:
00:15:22;05 you fail a lot of times,
00:15:24;26 and second of all, when you fail
00:15:26;29 you can score bigger than if you succeeded.
00:15:29;09 Because sometimes when you fail,
00:15:31;12 it means that some of your ideas were wrong going into it,
00:15:33;27 and it means that if you figure out where you’re wrong
00:15:36;12 you can make, actually, a bigger discovery
00:15:38;13 at the end of the day.
00:15:39;24 So, what to do next, besides desperation?
00:15:43;25 We kind of figured out maybe something is missing.
00:15:46;23 So, we added back… I’ll call it “magic juice”,
00:15:48;25 but it’s basically kind of soluble proteins from the cells,
00:15:53;17 proteins that are just kind of floating around
00:15:56;01 in what’s called the cytoplasm.
00:15:57;19 We added that back to see if it would work.
00:15:59;25 And the answer is, it worked.
00:16:02;17 The cargo now started moving.
00:16:04;13 Very cool!
00:16:05;28 But the next experiment was even cooler,
00:16:09;13 and this was a total surprise.
00:16:12;19 But I need to set this up for you
00:16:14;12 so you can understand it.
00:16:16;05 Before I show you the experiment,
00:16:18;10 I’m gonna show you another kind of transportation system, okay?,
00:16:22;09 that you can probably relate to as kids your age.
00:16:29;19 Okay. This is The Boss, here.
00:16:35;27 Now, I hope I can do that when I’m 62 too.
00:16:43;12 But how is he being transported?
00:16:45;18 This is not a dumb video. How is he being transported?
00:16:49;04 How is he being transported?
00:16:51;20 Yes? Student: It’s by people’s hands who are moving him along.
00:16:54;28 Right. People are stationary, right?
00:16:56;27 And their hands are moving him, okay?
00:16:59;23 Now, I’m gonna show you that I discovered crowd surfing
00:17:03;26 well before pop culture did —
00:17:06;12 at a molecular level.
00:17:08;06 So, this was the experiment.
00:17:14;09 That magic juice kind of…
00:17:17;11 one time, almost by accident,
00:17:20;24 I just spread this magic soluble protein solution on a glass slide.
00:17:25;18 Okay?
00:17:27;18 And what happened was that the magic juice
00:17:30;21 contained motor proteins in it.
00:17:33;12 The motor proteins would stick to the surface of the glass slide
00:17:37;09 just like the people in that rock concert.
00:17:40;08 But if you now have this situation
00:17:42;02 where your walkers are stationary
00:17:44;17 and they grab hold of this roadway,
00:17:47;20 these microtubules,
00:17:49;23 what will happen is that these motor proteins,
00:17:52;10 as they’re trying to move along the roadway,
00:17:54;25 are gonna be transporting the microtubule across the surface,
00:17:59;24 this lawn of motor proteins.
00:18:01;28 And can you see those,
00:18:03;16 those little snakes now moving along the glass?
00:18:07;29 Now, this was a solution of maybe 10,000 proteins,
00:18:11;04 but it provided a way to find out,
00:18:13;28 what was the one protein
00:18:16;28 that was causing this motion?
00:18:20;13 And that… you can use this technique called biochemistry, basically.
00:18:24;02 And biochemical purification
00:18:26;19 is kind of simple conceptually.
00:18:28;06 You have 10,000 proteins.
00:18:30;17 Let’s separate them by some various methods,
00:18:33;10 chromatography for example,
00:18:35;13 and put them into ten different bins, okay?
00:18:38;00 And now, you look at those ten different, like, bowls of proteins,
00:18:42;10 where proteins are separated
00:18:43;27 and ask, which of the ten bowls
00:18:46;20 will do that when you coat a glass surface?
00:18:48;08 Ah, this bowl does, the other [nine] don’t.
00:18:50;24 You throw away the other nine.
00:18:52;21 You keep that bowl.
00:18:54;06 You then separate the proteins
00:18:56;01 into ten more bowls,
00:18:57;28 so you now separate ‘em finer and finer.
00:18:59;28 You look on the microscope.
00:19:01;19 You ask, which one does that?
00:19:03;11 You find the one bowl, until you eventually get down to one protein.
00:19:07;08 And that one protein turned out to be
00:19:09;14 a completely new protein,
00:19:11;02 a new molecular motor
00:19:13;07 that was driving this kind of transport.
00:19:15;21 So, that was pretty cool.
00:19:18;03 My mom and I got to name it.
00:19:19;29 You know, you could… you should call your mom on the phone
00:19:21;29 and ask them for advice sometimes.
00:19:24;05 So, I called my mom, and she…
00:19:26;02 you know, the usual weekend call.
00:19:27;12 You know, I said, how are you doin’?
00:19:29;00 Fine, you know.
00:19:30;03 I said, well, mom, I actually got some really cool dis…
00:19:32;14 you know, a really cool thing.
00:19:34;06 I think I found this completely new,
00:19:35;23 you know, molecular motor protein.
00:19:38;00 And you know, she would say, oh, that’s nice dear.
00:19:39;15 Are you taking your vitamins?
00:19:41;02 And I go, mom!
00:19:42;23 This is actually really important.
00:19:44;15 And I said, I actually now need to name it.
00:19:46;05 I need to come up with a name.
00:19:47;22 And she actually said, oh, you know,
00:19:49;09 I’ll call up my friend who’s a Greek scholar.
00:19:51;20 So, she actually came up with the name kinesin,
00:19:54;18 from kinein — kinetic energy, motion.
00:19:57;18 So, umm… and this is what this…
00:20:01;11 after another 15 years,
00:20:03;16 how we think this motor protein actually works.
00:20:06;13 It binds ATP — that’s adenosine triphosphate —
00:20:10;27 and it… the ATP binding causes these two engines, in blue,
00:20:16;03 to basically walk along this track in this hand-over-hand manner.
00:20:20;22 And the ATP actually causes
00:20:22;20 a change in part of this motor.
00:20:25;11 See that red part over there?
00:20:27;13 When the ATP comes in,
00:20:29;04 it causes that red piece to zipper up
00:20:31;16 along the side of the main body of the enzyme.
00:20:36;05 It’s very much like… an analogy is like
00:20:38;25 a judo expert, you know, with the throw of the arm.
00:20:42;04 If the judo expert throws their arm forward,
00:20:43;29 they can throw an opponent from a rear side to a forward side.
00:20:47;08 That’s what this motor protein does.
00:20:49;05 It plays leapfrog between these two kinds of engines
00:20:53;09 so that one engine goes in front,
00:20:55;00 and then the other engine,
00:20:56;28 and then the other engine.
00:20:58;06 And this way, it can walk along a track.
Lasker Award winner Vale visits a high school class and explains how he became fascinated with molecular motors and how they work. He also describes the experiments that led to his discovery of the motor protein kinesin.
Professor of the Department of Cellular and Molecular Pharmacology; Investigator in the Howard Hughes Medical Institute
University of California, San Francisco Continue Reading
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