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Why does every mammal get 1 billion heartbeats in their life? — Transcript

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  1. 0:00How much LSD should you give an elephant?
  2. 0:02(mellow music) Well, to a reasonable person,
  3. 0:04the correct answer is probably none.
  4. 0:06But what if you needed to do it for a scientific experiment?
  5. 0:10In the 1960s, the CIA was working on
  6. 0:12a top secret project known as MKUltra,
  7. 0:15and one of their main goals was to find out
  8. 0:16how drugs like LSD could be used to change human behavior.
  9. 0:21And this is where elephants come in
  10. 0:23because elephants are normally quite docile,
  11. 0:25but sometimes they just snap.
  12. 0:27And the hypothesis was
  13. 0:29that this change in behavior might be triggered
  14. 0:31by the release of an LSD-like substance
  15. 0:33that naturally occurs in their brains.
  16. 0:36So if that's true, then administering LSD
  17. 0:39to a docile elephant might reproduce that behavior.
  18. 0:42So the real question was how much LSD should you give
  19. 0:45an elephant so that the dose is large enough
  20. 0:47to cause a psychological reaction,
  21. 0:49but not so large that it causes harm?
  22. 0:52(mellow music)
  23. 0:52Well, the researchers didn't know, LSD had never been given
  24. 0:56to an animal that big,
  25. 0:57but they did know that the safe dose in cats
  26. 1:00was around 0.3 milligrams.
  27. 1:03Now, since an elephant has around
  28. 1:04a thousand times the mass of a cat,
  29. 1:06they figured we'll give it a thousand times the dose.
  30. 1:09They received approval to perform the experiment on Tusko,
  31. 1:12an Indian elephant at the Lincoln Park Zoo in Oklahoma.
  32. 1:16And there they injected him
  33. 1:17with nearly 300 milligrams of LSD.
  34. 1:20But within five minutes, Tusko trumpeted, collapsed,
  35. 1:24fell heavily onto his right side, defecated,
  36. 1:27and went into status epilepticus.
  37. 1:30They administered a few other drugs
  38. 1:32in an attempt to revive him,
  39. 1:33but Tusko died shortly thereafter.
  40. 1:36The mistake they made was to assume
  41. 1:38that safe drug dosage scales linearly with mass.
  42. 1:42It does not.
  43. 1:43And it turns out there are a lot of things like this
  44. 1:46that don't scale in the way you'd expect.
  45. 1:48(mellow music)
  46. 1:49For example, take the smallest mammal by mass,
  47. 1:52the Etruscan shrew,
  48. 1:53and the largest land mammal, the African bush elephant.
  49. 1:57Which do you think has more heartbeats
  50. 1:59over the course of its entire life?
  51. 2:01Well, an African elephant has
  52. 2:03around a billion heartbeats in its lifetime,
  53. 2:06and an Etruscan shrew also has a billion
  54. 2:09heartbeats in its lifetime.
  55. 2:11What about a wallaby?
  56. 2:13Also a billion.
  57. 2:14A two-toed sloth?
  58. 2:15A billion again.
  59. 2:16Just about every mammal, no matter
  60. 2:18what environment they live in, how large they are,
  61. 2:21or even whether they live for one year or 100 years,
  62. 2:24they all get around a billion heartbeats
  63. 2:27between the day they're born and the day they die.
  64. 2:29- Why a billion?
  65. 2:30Why not a million?
  66. 2:31Why not a thousand?
  67. 2:32It's not these other numbers.
  68. 2:33- That is every mammal except one.
  69. 2:37This video is in large part based on the book
  70. 2:39Scale by Geoffrey West.
  71. 2:41If you want to learn more,
  72. 2:42I'll put a link to it down in the description.
  73. 2:44What's even more curious is that
  74. 2:46just by knowing a mammal's mass,
  75. 2:48you can predict a staggering number
  76. 2:50of biological traits from its pulse rate
  77. 2:52and reproductive output to its total lifespan.
  78. 2:56The same pattern holds for cities.
  79. 2:58If you know the population and location,
  80. 3:00that allows you to forecast everything from average wages
  81. 3:02and patent filings to crime rates, disease prevalence,
  82. 3:06and even the literal speed at which pedestrians walk.
  83. 3:10So how is that possible?
  84. 3:12Well, for that, we have to go back to the case
  85. 3:13of Tusko the Elephant.
  86. 3:15Researchers assumed that safe drug dosage
  87. 3:17is proportional to mass.
  88. 3:19Double the mass, double the dose.
  89. 3:20But it turns out that the speed at which an animal
  90. 3:23can process chemical compounds
  91. 3:24doesn't depend directly on its mass.
  92. 3:27It depends more on its metabolic rate.
  93. 3:29That is the number of calories it uses
  94. 3:32in a given amount of time.
  95. 3:33- There's a lot going on in your body.
  96. 3:35Your heart is pumping, that takes energy.
  97. 3:38You're moving food around.
  98. 3:40Digestion takes energy.
  99. 3:42Keeping your brain going takes energy, breathing.
  100. 3:44Everything that you do requires energy.
  101. 3:47(mellow music) - [Host] For a cat, for a single day,
  102. 3:49they require roughly 250 kilocalories of energy,
  103. 3:53used by all of their trillions of cells.
  104. 3:56But there is nothing special about a cat's cells.
  105. 3:59If you take a cat's cells and an elephant's cells
  106. 4:01and put them under a microscope,
  107. 4:02you'll find they're a similar size, similar makeup,
  108. 4:05and they perform the same sorts of functions.
  109. 4:07The same is true for other animals.
  110. 4:10In other words, the building blocks
  111. 4:11of animals are always roughly the same.
  112. 4:14So an elephant that has a thousand times the mass
  113. 4:16has about a thousand times as many cells.
  114. 4:18So you'd expect it would need a thousand
  115. 4:21times as much energy.
  116. 4:22So 250,000 kilocalories per day.
  117. 4:26But that is where you run into problems.
  118. 4:28- So what's the issue here?
  119. 4:30If I'm an organism
  120. 4:31and I'm burning up energy all day long,
  121. 4:33that energy is radiated out in the form of body heat.
  122. 4:37So we're constantly losing heat through our surface,
  123. 4:40through our skin, to the environment.
  124. 4:43- To see why this matters, let's simplify the problem
  125. 4:46and do the standard physicist thing.
  126. 4:47Let's assume our animals are perfect spheres.
  127. 4:51To be clear, this isn't necessary for the argument to work,
  128. 4:53but it does make everything a lot easier to follow.
  129. 4:56Since the volume of an animal is proportional to its mass,
  130. 4:59our 3000 kilogram elephant has a volume a thousand times
  131. 5:02greater than our three kilogram cat.
  132. 5:04So its radius must be 10 times larger.
  133. 5:06That's because volume is proportional to radius cubed.
  134. 5:10But surface area only grows as radius squared.
  135. 5:13So the elephant's surface area
  136. 5:14only increases by a factor of 100.
  137. 5:17Generating a thousand times as much heat
  138. 5:18while only having a hundred times the surface area
  139. 5:20to radiate it away would end very
  140. 5:23poorly for the elephant.
  141. 5:24If this were the case, it would boil alive.
  142. 5:28(mellow music) So in 1838,
  143. 5:30French scientists proposed a different scaling law.
  144. 5:32Since metabolism generates heat
  145. 5:34and that heat is radiated through the surface,
  146. 5:36metabolic rate or B, should scale in proportion
  147. 5:39to the surface area, A, instead.
  148. 5:42This became known as the surface law.
  149. 5:44Now, we can rewrite this to see
  150. 5:46how metabolic rate scales as a function of mass.
  151. 5:48Surface area is proportional to radius squared,
  152. 5:51so we can swap that in,
  153. 5:52and if mass is proportional to radius cubed,
  154. 5:55then radius must be proportional to mass to the one-third.
  155. 5:58Plugging that in for R,
  156. 5:59we find that metabolic rate should scale
  157. 6:00with mass to the two-thirds.
  158. 6:02- Really familiar example would
  159. 6:04come from thinking about cooking.
  160. 6:07(mellow music)
  161. 6:07Maybe you want to make a big turkey for Thanksgiving.
  162. 6:10People commonly phrase it as how long do I need per pound?
  163. 6:13But that you realize is linear thinking.
  164. 6:15The important thing is really the thickness of the bird
  165. 6:18or the roast, because the heat is coming in
  166. 6:21by conduction from the oven
  167. 6:24and it's got to do thermal diffusion into the meat.
  168. 6:28The time for the diffusion will scale
  169. 6:30like the characteristic length,
  170. 6:32which in this case would be the thickness of the meat.
  171. 6:34It'll go like the dimension squared,
  172. 6:37whereas volume of the meat is going to be like length cubed,
  173. 6:41and that's proportional to the weight
  174. 6:43of the bird or the roast.
  175. 6:44So when you put those things together,
  176. 6:46you'll get that the characteristic amount of time needed
  177. 6:48to cook the roast will go like its mass
  178. 6:51to the two-thirds power.
  179. 6:52- So if I'm thinking about cooking a roast
  180. 6:55that weighs twice as much as another roast, how much longer?
  181. 6:58Is that two to the two-thirds the time?
  182. 7:01- Yeah, if you wanted to double the weight,
  183. 7:03then you only have to cook it about 60% longer,
  184. 7:06not 100% longer.
  185. 7:08- [Host] Similarly, according to this scaling,
  186. 7:11an elephant that's a thousand times as heavy as a cat
  187. 7:13should only burn a hundred times as many calories.
  188. 7:16So 25,000 instead of 250,000.
  189. 7:19And the appropriate dose of LSD for Tusko
  190. 7:22would've been just 30 milligrams.
  191. 7:25These two wildly different predictions come from different
  192. 7:27assumptions about how metabolic rate
  193. 7:29scales as a function of mass.
  194. 7:31But notice in both cases, it's just proportional
  195. 7:34to mass raised to some power.
  196. 7:36These kinds of relationships are called power laws
  197. 7:38and all power laws have a special property.
  198. 7:41If you take the logarithm of the X and Y values,
  199. 7:44you get a straight line.
  200. 7:45And the slope of the line is equal
  201. 7:47to the exponent of the power law.
  202. 7:49This makes it easy to identify
  203. 7:51different types of power laws.
  204. 7:52If the slope is one, that's just everyday linear scaling.
  205. 7:55If it's less than one, like the surface law,
  206. 7:58that is called sublinear scaling.
  207. 8:00And if it's larger than one,
  208. 8:01that is known as superlinear scaling.
  209. 8:04For nearly a hundred years, biologists generally agreed
  210. 8:07that the two-thirds exponent
  211. 8:09of the surface law was the correct one for metabolic rate.
  212. 8:12But then in 1932, Swiss biologist, Max Kleiber,
  213. 8:16decided to put it to the test.
  214. 8:17He took the metabolic rates
  215. 8:19of different animals from a small dove at 150 grams
  216. 8:22to a large steer at 680,000 grams,
  217. 8:26and then he plotted them against their mass
  218. 8:28on a log log plot.
  219. 8:29And as expected, all the data did fall on a straight line.
  220. 8:32But the slope wasn't two-thirds.
  221. 8:35Instead, it was about three quarters.
  222. 8:37This became known as Kleiber's Law.
  223. 8:40It implies that if you double an animal's mass,
  224. 8:42metabolic rate goes up by about 1.68,
  225. 8:45an increase of 68% instead of the 59%,
  226. 8:49which would be predicted by the two-thirds scaling law.
  227. 8:51So according to Kleiber's Law,
  228. 8:53an elephant burns roughly 178 times
  229. 8:56as many calories as a cat, or about 45,000 kilocalories.
  230. 9:00And the actual LSD dose Tusko should have received
  231. 9:03was 53 milligrams, which is around a sixth of the dose
  232. 9:06the researchers gave him.
  233. 9:08So that explains the dosing catastrophe
  234. 9:10with Tusko The Elephant.
  235. 9:13(mellow music)
  236. 9:13Kleiber's original work was based on a small data set,
  237. 9:16primarily made up of mammals.
  238. 9:18But if you plot the data for a larger range of mammals,
  239. 9:21as well as for other animals
  240. 9:22like birds, reptiles, and fish,
  241. 9:24you find that they all follow
  242. 9:25roughly the same scaling relationship.
  243. 9:27Now, they don't all fit perfectly onto the same line
  244. 9:29because the base metabolic rate
  245. 9:31for warm-blooded animals is higher
  246. 9:32than for cold-blooded ones,
  247. 9:34but they all scale according to the same power law,
  248. 9:37mass raised to the three-quarters.
  249. 9:39Some argue that this relationship expands all the way down
  250. 9:42to single cells and the molecular machines inside of them.
  251. 9:45If that's true, Kleiber's Law governs life
  252. 9:47spanning more than 25 orders of magnitude.
  253. 9:51But then the question is,
  254. 9:53if an elephant has so many more cells
  255. 9:56and each cell is a similar size
  256. 9:58to a cell in a smaller creature,
  257. 10:00and yet it's using proportionately less energy,
  258. 10:05like it's got a lower metabolic rate per pound
  259. 10:08or per kilo or per cell,
  260. 10:10that's really saying that those cells
  261. 10:13are functioning with much less energy.
  262. 10:15- It's like there's some efficiency to being big,
  263. 10:18and it's hard to understand why.
  264. 10:20What is it that the cells are providing to each other,
  265. 10:24somehow cooperating in some way?
  266. 10:26- I just put in my calculator 100 to the three quarters,
  267. 10:29so if you got an organism that's 100 times bigger,
  268. 10:32then according to this, only 31.6
  269. 10:35times the metabolic rate.
  270. 10:37So it seems like a big savings.
  271. 10:40- It's a massive improvement, right?
  272. 10:42It's a big savings.
  273. 10:43It's a biological fact,
  274. 10:44but people have been arguing now for a century
  275. 10:48what accounts for this three quarters.
  276. 10:50(mellow music)
  277. 10:50- In the decades following Kleiber's observation,
  278. 10:52the mystery only deepened.
  279. 10:54Researchers discovered that brain size also roughly scales
  280. 10:58as mass to the three quarters,
  281. 10:59and so does an animal's growth rate
  282. 11:01and the amount of blood pumped per minute.
  283. 11:04But not every property scales as mass to the three quarters.
  284. 11:07- If you ask how long a creature will live, a mammal,
  285. 11:11that tends to be proportional to its mass
  286. 11:13to the one quarter power.
  287. 11:15One over four, not three over four.
  288. 11:18- [Host] So that means if you double the mass of a mammal,
  289. 11:20then on average, its lifespan is around 19% longer.
  290. 11:24Similarly, blood circulation time also scales
  291. 11:26as roughly mass to the one quarter, while breathing rate
  292. 11:29and heart rate both scaled to the negative one quarter.
  293. 11:32They're not all three quarters power laws,
  294. 11:35but they are all multiples of a quarter.
  295. 11:37So the question on everyone's mind was
  296. 11:40where are these quarter power scaling laws coming from?
  297. 11:43- One popular theory emerged in the 1990s.
  298. 11:47(mellow music) Brian Enquist was studying
  299. 11:48for an undergraduate degree in biology.
  300. 11:50- Most of the biology classes then that you take,
  301. 11:53you learn about the Krebs Cycle
  302. 11:55and then you have to memorize all the different parts
  303. 11:57of a flower and then all...
  304. 11:59They have a different terminology for everything.
  305. 12:00- [Henry] But then one day he took a zoology class
  306. 12:03where they showed him those quarter power scaling plots.
  307. 12:05- [Brian] I just couldn't believe it.
  308. 12:07I was like, "You got to be kidding me."
  309. 12:09You know, this is kind of like something fundamental
  310. 12:11that's kind of like underlying biological diversity.
  311. 12:14And there I knew immediately that I wanted
  312. 12:17to kind of quit my specialized
  313. 12:19kind of plant physiology research
  314. 12:21and do something associated with scaling.
  315. 12:24(mellow music)
  316. 12:24- [Henry] So Enquist started studying for his PhD,
  317. 12:27under Professor James Brown,
  318. 12:29who had been thinking about scaling laws for years.
  319. 12:31So they knew that often when power laws appear,
  320. 12:34there's some form of self-similarity
  321. 12:35in the underlying system.
  322. 12:36So they wondered, what could that self-similarity be?
  323. 12:39And they suspected that it might have something to do
  324. 12:41with the way resources are transported through the body,
  325. 12:44specifically the networks that do this.
  326. 12:46They had the biological intuition,
  327. 12:48but to get a complete theory,
  328. 12:49they needed a formal mathematical framework.
  329. 12:52In other words, they needed a mathematician
  330. 12:54or a theoretical physicist.
  331. 12:55- And at the time, Jim was associated
  332. 12:58with the Santa Fe Institute,
  333. 13:00and he started asking around with,
  334. 13:02is there anyone up here that's interested in these
  335. 13:04biological scaling relationships?
  336. 13:06And then the president at the time said, "You know what?
  337. 13:08I know of this physicist who's up at Los Alamos
  338. 13:12who was talking about biological scaling relationships."
  339. 13:15And so we met Geoffrey and it was like immediately,
  340. 13:18it's like we'd been talking about the same things
  341. 13:21for like years.
  342. 13:22It's like, you know, finding,
  343. 13:25someone who's been like speaking your language,
  344. 13:27but no one could understand you.
  345. 13:28- [Henry] So West, Brown and Enquist teamed up
  346. 13:30to try and find a compelling explanation for Kleiber's Law.
  347. 13:33- They started by assuming three simple premises.
  348. 13:37The first premise is that the networks
  349. 13:39that distribute resources are space-filling,
  350. 13:41since they need to reach every cell in the body.
  351. 13:44The second premise is that the terminal units
  352. 13:46of those networks, the thinnest segments
  353. 13:48on the outer periphery of the delivery system,
  354. 13:50have the same width regardless of the size of the organism.
  355. 13:53That is the outermost blood vessels that carry nutrients
  356. 13:56to an elephant's skin cells are about as thick
  357. 13:58as the ones in a mouse.
  358. 14:00The elephant just has many more of them.
  359. 14:02And the third premise is that over time,
  360. 14:04evolution has driven these transport networks
  361. 14:06toward an efficient design.
  362. 14:08So what should such a network look like?
  363. 14:11(mellow music)
  364. 14:11Well, intuitively, to get fuel from one place to another
  365. 14:14as efficiently as possible,
  366. 14:15you want the path to be as short as possible.
  367. 14:17So basically a straight line.
  368. 14:18But because fuel needs to reach every part of the body,
  369. 14:21you would also need many different paths.
  370. 14:23And as you go to larger and larger organisms,
  371. 14:25the networks inside need to
  372. 14:27reach a larger and larger volume.
  373. 14:29One way to do this is to stretch all the paths
  374. 14:31and match the growth of the animal.
  375. 14:33In this case, the volume of the animal
  376. 14:35should scale as these internal path lengths cubed.
  377. 14:38Or if we rearrange that, internal path length should scale
  378. 14:40as volume to the one-third,
  379. 14:42just like the overall length of the animal does.
  380. 14:45But this design is incredibly wasteful.
  381. 14:47Take these two regions.
  382. 14:49The two vessels that are bringing blood here
  383. 14:51go through almost the same path in the body,
  384. 14:53and they only split up just
  385. 14:55before they reach their destinations.
  386. 14:57If instead we had just one vessel up to this point
  387. 14:59and split it only when the paths needed to diverge,
  388. 15:02we could serve the two regions
  389. 15:03using a lot less vessel material
  390. 15:05and a lot less blood to fill the vessels.
  391. 15:07Of course, you can extend this logic
  392. 15:09for all the blood vessels in the body,
  393. 15:10and you end up with a much more efficient design
  394. 15:13of branching blood vessels.
  395. 15:15But now we run into another issue
  396. 15:17because each branching point provides an opportunity
  397. 15:19for some of the blood to bounce back, that is reflect.
  398. 15:22If you have many reflections,
  399. 15:24that would mean it costs significantly more
  400. 15:25energy to pump blood around.
  401. 15:27So next, they argue that nature should favor structures
  402. 15:30that minimize reflections.
  403. 15:31And as it turns out, this happens
  404. 15:33if the cross-sectional area of the vessels
  405. 15:35stays the same before and after the branching.
  406. 15:37So if you've got a cross-sectional area
  407. 15:39of two centimeters squared for the main vessel,
  408. 15:41then each of the two branches need to have an area
  409. 15:43of one centimeter squared each, for large vessels at least.
  410. 15:47For smaller ones, daughter branches can be a bit thicker
  411. 15:50to allow blood to slow down
  412. 15:52and exchange resources with the tissue it's reaching.
  413. 15:55If you keep repeating this pattern across the network,
  414. 15:57you end up with this, a branching self-similar fractal.
  415. 16:01And if you look at the actual shape
  416. 16:03of the circulatory system, it has this geometry.
  417. 16:06So it seemed like they were onto something.
  418. 16:09- But how do you get from this
  419. 16:11to quarter power scaling laws?
  420. 16:13Well, mathematician Felix Hausdorff discovered
  421. 16:15that self-similar fractals have an interesting property.
  422. 16:18(mellow music) Take a straight line segment.
  423. 16:19It's completely one-dimensional and not a fractal at all.
  424. 16:22Hausdorff assigned this line segment a value of 1.0.
  425. 16:26But now imagine adding a few bends
  426. 16:29and more bends to those bends.
  427. 16:31If you keep doing this, the line becomes more
  428. 16:33and more fractal-like.
  429. 16:35Eventually, if you keep applying
  430. 16:36the right kind of contortions,
  431. 16:38that one-dimensional line segment fills up
  432. 16:40an entire region of the 2D plane.
  433. 16:42Hausdorff assigned these space-filling fractal curves
  434. 16:45a value of 2.0, corresponding to their dimensionality.
  435. 16:48The same ideas apply to a 2D surface.
  436. 16:51- So think about a piece of paper, right?
  437. 16:53Two-dimensional.
  438. 16:55All then the fractal network is doing is crumpling up
  439. 16:58that sheet of paper, and it effectively fills
  440. 17:01a ball of volume.
  441. 17:02So you can now describe that sheet of paper
  442. 17:05as a sphere instead of a two-dimensional sheet of paper.
  443. 17:09- [Henry] Repeat the right pattern of folds
  444. 17:10at smaller and smaller scales, and a 2D surface
  445. 17:13fills more and more of a 3D volume.
  446. 17:15In the mathematical limit, it becomes space filling
  447. 17:18with Hausdorff dimension of 3.0.
  448. 17:20- [Brian] You know, biologically, I said,
  449. 17:22"Well, what does that mean?"
  450. 17:23That enables an organism for a given size
  451. 17:26to pack in more of these metabolic surface areas
  452. 17:30than would be expected.
  453. 17:32And it's because of this fractal-like structure
  454. 17:35that enables you then to pack in and have all of these folds
  455. 17:38and convolutions and on top of each other
  456. 17:41to pack in an enormous amount of membrane surfaces.
  457. 17:44- As a result, the Hausdorff dimension of the surface
  458. 17:47of the circulatory system is roughly three,
  459. 17:50meaning its surface area doesn't scale
  460. 17:52as its length squared, but it's length cubed.
  461. 17:55And since the metabolic rate hinges on
  462. 17:57how fast resources can be exchanged across the surface area,
  463. 18:00well, it must also scale like length cubed.
  464. 18:03But remember, West, Brown and Enquist
  465. 18:05wanted to explain Kleiber's Law.
  466. 18:07So they needed to know how metabolic rate scales with mass.
  467. 18:12(mellow music)
  468. 18:12Since every cell needs to be served by the network,
  469. 18:14this means that the volume
  470. 18:15around the network should be proportional
  471. 18:17to the animal's mass.
  472. 18:18And since volume is just surface area times length
  473. 18:21and surface area is proportional to length cubed,
  474. 18:23that means both volume and mass
  475. 18:25must be proportional to length to the fourth,
  476. 18:28which can be rewritten to show that length is proportional
  477. 18:30to mass raised to the one quarter.
  478. 18:33And if you plug that into the equation
  479. 18:34for the metabolic rate,
  480. 18:35you find that the metabolic rate
  481. 18:36must be proportional to mass to the three-quarters,
  482. 18:40exactly as Max Kleiber had found.
  483. 18:43West, Brown and Enquist published their work in 1997,
  484. 18:46and it soon came to be known
  485. 18:48as WBE Theory, after their initials.
  486. 18:50- And the theory is quite rigid.
  487. 18:53It makes very specific predictions.
  488. 18:55This is good science.
  489. 18:57Okay?
  490. 18:57This is sticking your neck out,
  491. 18:59and it's an incredibly beautiful theory.
  492. 19:02(mellow music) - [Derek] Take a look at this table
  493. 19:03from Geoffrey West's book.
  494. 19:04These are the scaling exponents WBE theory predicts,
  495. 19:07including many that are not multiples of a quarter,
  496. 19:10but all follow from the same theory.
  497. 19:12For example, the radius of an animal's aorta
  498. 19:15should scale with its mass to the three-eighths or .375.
  499. 19:18And the area of its lungs should scale with mass
  500. 19:21to the 11-12ths or about 0.92.
  501. 19:24In total, this chart makes 26 different predictions.
  502. 19:27Now, these are the observed data.
  503. 19:30Radius of the aorta, 0.36.
  504. 19:32Lung area, 0.95.
  505. 19:35- [Steven] That's the really shocking thing
  506. 19:37about what they did.
  507. 19:37They had a table with something like, I don't know,
  508. 19:3920 or 30 predictions of exotic exponents,
  509. 19:43and that's what you really see in the data.
  510. 19:45So this one theory accounts not only
  511. 19:46for the three-quarters power of metabolism,
  512. 19:49but for literally dozens of other things
  513. 19:51that biologists have measured.
  514. 19:53- Some of the scaling laws are easy to explain
  515. 19:56once you've got the three-quarters law for metabolism.
  516. 19:58Take a mammal's heart rate, for instance.
  517. 20:01Heart rate is equal to the blood flow rate
  518. 20:03over the amount or volume of blood in every beat.
  519. 20:05(mellow music)
  520. 20:06The volume of blood per beat has been found
  521. 20:08to scale in direct proportion to an animal's mass.
  522. 20:10So that's just M.
  523. 20:11And the blood flow rate?
  524. 20:13Well, remember that metabolism is all about
  525. 20:15how nutrients get distributed around the body.
  526. 20:17So most biologists agree metabolic rate
  527. 20:19and blood flow rate are directly
  528. 20:20proportional to one another.
  529. 20:22So heart rate should scale as metabolic rate over mass.
  530. 20:25Swapping in the scaling law Kleiber had found,
  531. 20:28that gives us M to the minus one quarter,
  532. 20:30meaning bigger animals should have slower
  533. 20:32heartbeats than smaller ones.
  534. 20:34And this is exactly what we observe in nature.
  535. 20:36The world's smallest mammal, the Etruscan Shrew,
  536. 20:39has an extraordinary heart rate of 1200 beats per minute.
  537. 20:42That's 20 beats per second.
  538. 20:44Whereas the biggest land mammal, the African bush elephant,
  539. 20:47has a typical heart rate of only 30 beats per minute.
  540. 20:50And we can do something similar for lifespan.
  541. 20:53One of the leading theories is
  542. 20:54that an animal's lifespan is based on
  543. 20:56the accumulation of metabolic damage.
  544. 20:58That is, as each chunk of tissue in an organism
  545. 21:01processes nutrients over time,
  546. 21:03this causes damage to accumulate,
  547. 21:05and that over time causes the animal to die.
  548. 21:08So the rate at which an animal accumulates damage
  549. 21:10is its metabolic rate per unit of mass,
  550. 21:12and its lifespan should be the inverse of that rate.
  551. 21:15If damage accumulates faster, it dies sooner.
  552. 21:17If it accumulates slower, it lives longer.
  553. 21:20So lifespan is proportional to M over B,
  554. 21:23or substituting in Kleiber's Law, M to the one quarter.
  555. 21:27So lifespan should increase with mass.
  556. 21:30And you do see this in nature.
  557. 21:31(mellow music)
  558. 21:31A shrew only lives for one to two years in the wild
  559. 21:34while a mighty African elephant can live up to 70 years.
  560. 21:38So if you're a small mammal,
  561. 21:39you have many heartbeats per minute,
  562. 21:41but you live a relatively short life.
  563. 21:43Conversely, if you're a larger mammal,
  564. 21:45your heart beats much slower and you live a lot longer.
  565. 21:48- So it's the, you know, live fast and burnout
  566. 21:52and die young, right?
  567. 21:54Or spend it frugally and live a really long life.
  568. 21:57- But you might have also noticed something else.
  569. 22:00Heart rate scales as B over M.
  570. 22:02So it equals B over M times some constant.
  571. 22:05And lifespan scales as M over B.
  572. 22:08They're inverses of each other.
  573. 22:09They scale in equal and opposite directions.
  574. 22:12Now, the total number of heartbeats in an animal's life
  575. 22:15is just the heart rate multiplied by the lifespan.
  576. 22:17So when you multiply these two terms, they cancel out,
  577. 22:20leaving you with just a constant.
  578. 22:22So that suggests that no matter
  579. 22:24what mammal you're talking about,
  580. 22:26it should have roughly the same number of heartbeats.
  581. 22:29You can find that number by just plugging in some examples.
  582. 22:32Let's start with the Etruscan Shrew.
  583. 22:35The shrew's 1200 beats per minute multiplied by a lifespan
  584. 22:37of around 1.5 years gives you
  585. 22:39around 950 million heartbeats
  586. 22:41in the course of the shrew's life.
  587. 22:43Meanwhile, an African elephant's 30 beats per minute
  588. 22:45multiplied by a lifespan of around 65 years
  589. 22:47gives you a little over a billion heartbeats.
  590. 22:51And we could keep going.
  591. 22:52(upbeat music) But for nearly every mammal you look at,
  592. 22:54you keep landing at the same figure of
  593. 22:56around a billion heartbeats.
  594. 22:58This is why nearly every mammal from a tiny field mouse
  595. 23:01to a gazelle, from a cheetah to a hippopotamus,
  596. 23:05they all get around a billion heartbeats
  597. 23:07between the day they're born and the day they die.
  598. 23:12But there is one major outlier, one mammal
  599. 23:15that gets significantly more than a billion heartbeats.
  600. 23:18And that is us, humans. (mellow music)
  601. 23:21We are the lucky ones.
  602. 23:23Three centuries ago, humans were much closer
  603. 23:26to the standard value of one billion heartbeats.
  604. 23:28But around the mid 1800s, germ theory
  605. 23:31and better sanitation methods became widespread,
  606. 23:33causing a stark decrease in the number of child mortalities
  607. 23:36and deaths from disease.
  608. 23:37So life expectancy began to climb up.
  609. 23:39And with it, the average number of heartbeats in a lifetime.
  610. 23:42If you look closely, you can also see some significant drops
  611. 23:46like this 1918 dip from the Spanish flu pandemic
  612. 23:49or over here, what appears to be
  613. 23:50the impact of the Second World War.
  614. 23:52But overall, the trend is clear.
  615. 23:54We have systematically been increasing the number
  616. 23:57of heartbeats we get in our lifetime to the point
  617. 23:59where now the average human gets nearly three billion
  618. 24:02heartbeats before they die.
  619. 24:05I don't know if there's a better argument for science
  620. 24:07and technology than this.
  621. 24:09It has literally given the average human
  622. 24:11more than a full extra life.
  623. 24:13And it's not just humans.
  624. 24:14Other mammals have been observed
  625. 24:16to have much longer lives in captivity when they're away
  626. 24:18from the hazards they would naturally encounter in the wild.
  627. 24:22Or if you look at it purely from the number of years we get,
  628. 24:24we now have the lifespan of a much larger mammal,
  629. 24:27somewhere between an elephant and a whale.
  630. 24:31But there is one curious thing about this trend.
  631. 24:33(mellow music) Take a look at this graph.
  632. 24:35It looks surprisingly similar to the graph from before.
  633. 24:38In fact, if you overlay them, they look remarkably similar.
  634. 24:41Now I want you to take a guess at what this graph is.
  635. 24:44Have you got your answer?
  636. 24:46It is the number of people living in cities.
  637. 24:49For these two charts, we're using data from parts
  638. 24:52of the United Kingdom where the record keeping
  639. 24:54goes back several centuries,
  640. 24:56but the rest of the world has followed similar trends.
  641. 24:59Of course, that doesn't mean
  642. 25:00cities cause people to live longer,
  643. 25:03but it goes against the perception of cities
  644. 25:05as being full of pollution and breeding grounds for disease.
  645. 25:09Already in 1889, a medical doctor wrote,
  646. 25:12"The poisonous germs and pollutions of the city,
  647. 25:14it's impure air and water, bad sewage,
  648. 25:17and endless nuisances."
  649. 25:19So how do you reconcile these two views?
  650. 25:22Or more specifically, is there any quantitative data
  651. 25:25on how smaller cities compare to larger ones?
  652. 25:28- It turns out this is something that Geoffrey West pursued
  653. 25:31after his work with Brown and Enquist.
  654. 25:33He and collaborators like Luis Bettencourt and others
  655. 25:37have looked at scaling laws in cities.
  656. 25:39(mellow music)
  657. 25:39- [Host] They looked at how different properties
  658. 25:41like the amount of crime scale
  659. 25:43as the population of the city increases,
  660. 25:45and what they found is that if you plot
  661. 25:46serious crimes on a log log plot,
  662. 25:48the data clusters around a straight line
  663. 25:50with a slope of 1.15,
  664. 25:52meaning crime grows faster than linear or super linear.
  665. 25:56So for every doubling of a city's population,
  666. 25:59you get around 2.2 times as many criminal cases
  667. 26:02or around 120% more crime as opposed to the 100%
  668. 26:07you might naively expect.
  669. 26:08To make matters worse, researchers found
  670. 26:10that the same general pattern
  671. 26:11holds for the amount of wastewater
  672. 26:13and even the number of AIDS cases.
  673. 26:16The exact exponents vary a little,
  674. 26:18but overall, as cities grow larger,
  675. 26:20you systematically get more of each.
  676. 26:23So it seems like that medical doctor was onto something.
  677. 26:26And you might think life on earth would be better off
  678. 26:28if we all lived in small towns instead,
  679. 26:31but that might not be the case.
  680. 26:33(mellow music) In 2006, Dirk Helbing, Christian Kuhnert,
  681. 26:36and Geoffrey West looked at how the number
  682. 26:38of gas stations scales as a function of population.
  683. 26:41- If a city is twice as big, does it need twice
  684. 26:44as many gas stations?
  685. 26:45Because, you know, we have to supply not exactly nutrients,
  686. 26:48but energy, gas, for all those cars.
  687. 26:51- [Host] The naive expectation is that
  688. 26:53if you double the number of cars,
  689. 26:54you're going to need to double the amount of fuel,
  690. 26:56so double the gas stations.
  691. 26:58To find out whether this was true,
  692. 26:59they plotted the data on a log log plot
  693. 27:01and found a straight line.
  694. 27:03But the exponent wasn't one, it was about 0.8.
  695. 27:07This means that for every doubling, you only need around 74%
  696. 27:11more gas stations, which is a decent savings.
  697. 27:14(mellow music) The amount of roads and electrical cables
  698. 27:17also scale in roughly the same way.
  699. 27:19The rough figure that West gives in his book
  700. 27:21is that they all have scaling exponents of around 0.85.
  701. 27:23- It is interesting that some of the things
  702. 27:25that we can share, like you can drive
  703. 27:28on the road, but so can I.
  704. 27:29When it's shared resources, yes,
  705. 27:32cities can be surprisingly green.
  706. 27:34The argument is that cities can be even greener
  707. 27:36than you might think than living out
  708. 27:38in the middle of nowhere.
  709. 27:39- But cities have even bigger benefits.
  710. 27:42Things like total wages, GDP,
  711. 27:44and the number of patents all scale superlinearly,
  712. 27:47with exponents that cluster somewhere around 1.15,
  713. 27:51meaning that for every doubling in size,
  714. 27:53you get around 120% more of each.
  715. 27:57All of this becomes especially significant
  716. 27:59when you compare a small town of say 50,000 people
  717. 28:01to a city 100 times its size
  718. 28:04because infrastructure needs only need to go up by a factor
  719. 28:06of about 50 while total wages, GDP, patents and inventions,
  720. 28:11they all go up by a factor of 200.
  721. 28:13Unfortunately, disease and crime also go up
  722. 28:16by the same factor.
  723. 28:18Or look at it this way, on a per person basis,
  724. 28:21you'd only need about half the infrastructure,
  725. 28:23while you get double all the socioeconomic factors.
  726. 28:27So cities far from being detrimental to the world,
  727. 28:29they might actually be one of our best inventions
  728. 28:32and an indirect driver of a lot of scientific
  729. 28:35and technological progress.
  730. 28:36Perhaps this is also why people often say
  731. 28:38that life in the city feels faster.
  732. 28:41A feeling that seems justified
  733. 28:42because researchers looked at
  734. 28:44how fast people walk in cities of different sizes.
  735. 28:47And they found that people literally do walk
  736. 28:49faster in larger cities.
  737. 28:51- That turns out to depend on city size.
  738. 28:54It's not just that the sidewalks are congested or not.
  739. 28:57It's just like the vibe gets people amped up.
  740. 29:00They move faster in cities.
  741. 29:02- So the pace of life seems to be increasing,
  742. 29:05but that may come at a cost
  743. 29:07because as one person put it, "everything nowadays is ultra.
  744. 29:11Everything is being transcended continually
  745. 29:13in thought as well as in action.
  746. 29:14No one knows himself any longer.
  747. 29:16Young people are stirred up much too early in life
  748. 29:19and then carried away in the world of the times.
  749. 29:21Wealth and rapidity are what the world admires".
  750. 29:24Could it be that life is accelerating so fast
  751. 29:26that humans won't be able to keep up?
  752. 29:29Well, probably not
  753. 29:30because this quote was written in 1825
  754. 29:32by Wolfgang von Goethe.
  755. 29:34For the past 200 years and probably longer,
  756. 29:38almost every generation has felt like life was accelerating.
  757. 29:41And yet every generation has managed.
  758. 29:43So I think it's likely we can continue
  759. 29:45to adapt indefinitely.
  760. 29:47And as cities continue to grow in size,
  761. 29:50all of us will continue to reap the benefit
  762. 29:52of economies of scale in much the same way
  763. 29:54that mammals benefit from being larger.
  764. 29:57But while WBE Theory seems to predict
  765. 29:59where the exponents in biological scaling laws come from,
  766. 30:02for cities, there is no widely accepted
  767. 30:04explanatory theory yet.
  768. 30:06Trying to explain where those exponents of 0.85
  769. 30:09and 1.15 come from is one of the big goals for theorists.
  770. 30:12Although even WBE Theory is not universally accepted.
  771. 30:16For one, the fact it predicts the right exponents
  772. 30:19doesn't necessarily mean the theory itself is correct.
  773. 30:22There are a few other theories that predict
  774. 30:24some of the same scaling exponents,
  775. 30:26and there are also some other critiques.
  776. 30:29- There's a lot of discussion.
  777. 30:31It may look convincing.
  778. 30:32And personally, I tend to think it is very impressive.
  779. 30:35But I have very good colleagues like Peter Dodds
  780. 30:37at University of Vermont,
  781. 30:39and he thinks that a lot of the data analysis
  782. 30:41is either not done exactly right
  783. 30:43or that the data are so noisy
  784. 30:46that you shouldn't really take this so seriously.
  785. 30:49- Dodds even argues that Kleiber's Law
  786. 30:51itself might not be true. (mellow music)
  787. 30:53- In the 1960s, there's a symposium on energy metabolism,
  788. 30:59some name like this in animals,
  789. 31:02and at the end of it, they vote 29 to zero
  790. 31:04that it's going to be three quarters, right?
  791. 31:06Because, you know, you got to set some rules.
  792. 31:09If you go back and look at the data,
  793. 31:10which no one is really doing anymore, right,
  794. 31:12the data does not work.
  795. 31:13Like it doesn't work.
  796. 31:15(mellow music)
  797. 31:15- [Henry] This is a graph of metabolic rate
  798. 31:17as a function of mass based on a study
  799. 31:19that looked at 391 species of mammals,
  800. 31:22much larger than Kleiber's range.
  801. 31:23And it looks like the three-quarter slope fits quite well.
  802. 31:26But this is just the top part of the chart,
  803. 31:28the biggest mammals.
  804. 31:30If you zoom out, you see that the rest of the mammals appear
  805. 31:32to fall on a line that's closer to two-thirds.
  806. 31:36And in recent studies of bird metabolism,
  807. 31:38you also find a slope that appears closer
  808. 31:40to two-thirds than three-quarters.
  809. 31:42Could it be that those French scientists
  810. 31:44from centuries ago were right,
  811. 31:45that metabolic rate really does
  812. 31:47just scale with surface area?
  813. 31:48Well, not so fast.
  814. 31:51(mellow music) For one, many recent studies
  815. 31:52of cold-blooded animals find slopes
  816. 31:54that are significantly higher than two-thirds.
  817. 31:56The bigger problem is that measuring
  818. 31:58metabolic rates is difficult.
  819. 31:59It generally involves putting animals in a container
  820. 32:02and taking very precise measurements
  821. 32:03of their heat production or oxygen consumption,
  822. 32:06all while ensuring that the animal is in an unstressed,
  823. 32:09low activity resting state.
  824. 32:10Unsurprisingly, this is particularly difficult
  825. 32:13to pull off with large animals.
  826. 32:15And as a result, many studies have found scaling exponents
  827. 32:17where the error bars include both
  828. 32:19two-thirds and three-quarters.
  829. 32:22Today, the research community is split.
  830. 32:24Many uphold Kleiber's Law and the three-quarter scaling,
  831. 32:26while others think it's two-thirds.
  832. 32:28But a growing number suspect
  833. 32:29that there is no universal scaling exponent
  834. 32:32across all of life.
  835. 32:33In fact, it may very well be the case
  836. 32:35that the metabolic rate of larger mammals
  837. 32:37scales as their mass to the three-quarters,
  838. 32:39while for smaller ones, it's their mass to the two-thirds.
  839. 32:41- You know, it's like everything in science
  840. 32:43that there are people arguing, and that's good.
  841. 32:45- I guess the bigger exhortation
  842. 32:47would be someone to really measure it,
  843. 32:49measure things beautifully.
  844. 32:50You know, we're in 2026,
  845. 32:52has to be not just one elephant at one zoo,
  846. 32:55it needs to be measured again really well.
  847. 32:57- [Host] What everyone does agree on
  848. 32:58is that scaling laws are real.
  849. 33:00- Life is not linear all the time.
  850. 33:03There are deals to be had.
  851. 33:04There's often in real life departures from proportionality,
  852. 33:08and you have to be aware of it.
  853. 33:10Sometimes things punch above their weight,
  854. 33:12and as you get bigger, you get more efficient.
  855. 33:15Sometimes there are detriments to size.
  856. 33:18- So it pays to know how things scale.
  857. 33:21Clearly from an energy efficiency perspective,
  858. 33:23animals benefit from being larger.
  859. 33:26Similarly, all of us potentially stand
  860. 33:28to gain from having more people living in larger cities.
  861. 33:31It could lead to more discoveries and inventions,
  862. 33:34and in doing so, improve the standard of living
  863. 33:36for all of us, which might even give us more
  864. 33:39heartbeats in our lifetimes.
  865. 33:44From the surface law to Kleiber's law to WBE theory,
  866. 33:48how metabolic rate scales with mass
  867. 33:50has been one of biology's biggest debates for centuries.
  868. 33:52By collecting better data and analyzing it carefully,
  869. 33:55there's a good chance
  870. 33:56that the next generation of researchers could be the ones
  871. 33:58to finally put this debate to rest.
  872. 34:00It could be one of you watching or a student that you know.
  873. 34:04And today's sponsor, Brilliant, is helping
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