Why does every mammal get 1 billion heartbeats in their life? — Transcript
Full transcript
- 0:00How much LSD should you give an elephant?
- 0:02(mellow music) Well, to a reasonable person,
- 0:04the correct answer is probably none.
- 0:06But what if you needed to do it for a scientific experiment?
- 0:10In the 1960s, the CIA was working on
- 0:12a top secret project known as MKUltra,
- 0:15and one of their main goals was to find out
- 0:16how drugs like LSD could be used to change human behavior.
- 0:21And this is where elephants come in
- 0:23because elephants are normally quite docile,
- 0:25but sometimes they just snap.
- 0:27And the hypothesis was
- 0:29that this change in behavior might be triggered
- 0:31by the release of an LSD-like substance
- 0:33that naturally occurs in their brains.
- 0:36So if that's true, then administering LSD
- 0:39to a docile elephant might reproduce that behavior.
- 0:42So the real question was how much LSD should you give
- 0:45an elephant so that the dose is large enough
- 0:47to cause a psychological reaction,
- 0:49but not so large that it causes harm?
- 0:52(mellow music)
- 0:52Well, the researchers didn't know, LSD had never been given
- 0:56to an animal that big,
- 0:57but they did know that the safe dose in cats
- 1:00was around 0.3 milligrams.
- 1:03Now, since an elephant has around
- 1:04a thousand times the mass of a cat,
- 1:06they figured we'll give it a thousand times the dose.
- 1:09They received approval to perform the experiment on Tusko,
- 1:12an Indian elephant at the Lincoln Park Zoo in Oklahoma.
- 1:16And there they injected him
- 1:17with nearly 300 milligrams of LSD.
- 1:20But within five minutes, Tusko trumpeted, collapsed,
- 1:24fell heavily onto his right side, defecated,
- 1:27and went into status epilepticus.
- 1:30They administered a few other drugs
- 1:32in an attempt to revive him,
- 1:33but Tusko died shortly thereafter.
- 1:36The mistake they made was to assume
- 1:38that safe drug dosage scales linearly with mass.
- 1:42It does not.
- 1:43And it turns out there are a lot of things like this
- 1:46that don't scale in the way you'd expect.
- 1:48(mellow music)
- 1:49For example, take the smallest mammal by mass,
- 1:52the Etruscan shrew,
- 1:53and the largest land mammal, the African bush elephant.
- 1:57Which do you think has more heartbeats
- 1:59over the course of its entire life?
- 2:01Well, an African elephant has
- 2:03around a billion heartbeats in its lifetime,
- 2:06and an Etruscan shrew also has a billion
- 2:09heartbeats in its lifetime.
- 2:11What about a wallaby?
- 2:13Also a billion.
- 2:14A two-toed sloth?
- 2:15A billion again.
- 2:16Just about every mammal, no matter
- 2:18what environment they live in, how large they are,
- 2:21or even whether they live for one year or 100 years,
- 2:24they all get around a billion heartbeats
- 2:27between the day they're born and the day they die.
- 2:29- Why a billion?
- 2:30Why not a million?
- 2:31Why not a thousand?
- 2:32It's not these other numbers.
- 2:33- That is every mammal except one.
- 2:37This video is in large part based on the book
- 2:39Scale by Geoffrey West.
- 2:41If you want to learn more,
- 2:42I'll put a link to it down in the description.
- 2:44What's even more curious is that
- 2:46just by knowing a mammal's mass,
- 2:48you can predict a staggering number
- 2:50of biological traits from its pulse rate
- 2:52and reproductive output to its total lifespan.
- 2:56The same pattern holds for cities.
- 2:58If you know the population and location,
- 3:00that allows you to forecast everything from average wages
- 3:02and patent filings to crime rates, disease prevalence,
- 3:06and even the literal speed at which pedestrians walk.
- 3:10So how is that possible?
- 3:12Well, for that, we have to go back to the case
- 3:13of Tusko the Elephant.
- 3:15Researchers assumed that safe drug dosage
- 3:17is proportional to mass.
- 3:19Double the mass, double the dose.
- 3:20But it turns out that the speed at which an animal
- 3:23can process chemical compounds
- 3:24doesn't depend directly on its mass.
- 3:27It depends more on its metabolic rate.
- 3:29That is the number of calories it uses
- 3:32in a given amount of time.
- 3:33- There's a lot going on in your body.
- 3:35Your heart is pumping, that takes energy.
- 3:38You're moving food around.
- 3:40Digestion takes energy.
- 3:42Keeping your brain going takes energy, breathing.
- 3:44Everything that you do requires energy.
- 3:47(mellow music) - [Host] For a cat, for a single day,
- 3:49they require roughly 250 kilocalories of energy,
- 3:53used by all of their trillions of cells.
- 3:56But there is nothing special about a cat's cells.
- 3:59If you take a cat's cells and an elephant's cells
- 4:01and put them under a microscope,
- 4:02you'll find they're a similar size, similar makeup,
- 4:05and they perform the same sorts of functions.
- 4:07The same is true for other animals.
- 4:10In other words, the building blocks
- 4:11of animals are always roughly the same.
- 4:14So an elephant that has a thousand times the mass
- 4:16has about a thousand times as many cells.
- 4:18So you'd expect it would need a thousand
- 4:21times as much energy.
- 4:22So 250,000 kilocalories per day.
- 4:26But that is where you run into problems.
- 4:28- So what's the issue here?
- 4:30If I'm an organism
- 4:31and I'm burning up energy all day long,
- 4:33that energy is radiated out in the form of body heat.
- 4:37So we're constantly losing heat through our surface,
- 4:40through our skin, to the environment.
- 4:43- To see why this matters, let's simplify the problem
- 4:46and do the standard physicist thing.
- 4:47Let's assume our animals are perfect spheres.
- 4:51To be clear, this isn't necessary for the argument to work,
- 4:53but it does make everything a lot easier to follow.
- 4:56Since the volume of an animal is proportional to its mass,
- 4:59our 3000 kilogram elephant has a volume a thousand times
- 5:02greater than our three kilogram cat.
- 5:04So its radius must be 10 times larger.
- 5:06That's because volume is proportional to radius cubed.
- 5:10But surface area only grows as radius squared.
- 5:13So the elephant's surface area
- 5:14only increases by a factor of 100.
- 5:17Generating a thousand times as much heat
- 5:18while only having a hundred times the surface area
- 5:20to radiate it away would end very
- 5:23poorly for the elephant.
- 5:24If this were the case, it would boil alive.
- 5:28(mellow music) So in 1838,
- 5:30French scientists proposed a different scaling law.
- 5:32Since metabolism generates heat
- 5:34and that heat is radiated through the surface,
- 5:36metabolic rate or B, should scale in proportion
- 5:39to the surface area, A, instead.
- 5:42This became known as the surface law.
- 5:44Now, we can rewrite this to see
- 5:46how metabolic rate scales as a function of mass.
- 5:48Surface area is proportional to radius squared,
- 5:51so we can swap that in,
- 5:52and if mass is proportional to radius cubed,
- 5:55then radius must be proportional to mass to the one-third.
- 5:58Plugging that in for R,
- 5:59we find that metabolic rate should scale
- 6:00with mass to the two-thirds.
- 6:02- Really familiar example would
- 6:04come from thinking about cooking.
- 6:07(mellow music)
- 6:07Maybe you want to make a big turkey for Thanksgiving.
- 6:10People commonly phrase it as how long do I need per pound?
- 6:13But that you realize is linear thinking.
- 6:15The important thing is really the thickness of the bird
- 6:18or the roast, because the heat is coming in
- 6:21by conduction from the oven
- 6:24and it's got to do thermal diffusion into the meat.
- 6:28The time for the diffusion will scale
- 6:30like the characteristic length,
- 6:32which in this case would be the thickness of the meat.
- 6:34It'll go like the dimension squared,
- 6:37whereas volume of the meat is going to be like length cubed,
- 6:41and that's proportional to the weight
- 6:43of the bird or the roast.
- 6:44So when you put those things together,
- 6:46you'll get that the characteristic amount of time needed
- 6:48to cook the roast will go like its mass
- 6:51to the two-thirds power.
- 6:52- So if I'm thinking about cooking a roast
- 6:55that weighs twice as much as another roast, how much longer?
- 6:58Is that two to the two-thirds the time?
- 7:01- Yeah, if you wanted to double the weight,
- 7:03then you only have to cook it about 60% longer,
- 7:06not 100% longer.
- 7:08- [Host] Similarly, according to this scaling,
- 7:11an elephant that's a thousand times as heavy as a cat
- 7:13should only burn a hundred times as many calories.
- 7:16So 25,000 instead of 250,000.
- 7:19And the appropriate dose of LSD for Tusko
- 7:22would've been just 30 milligrams.
- 7:25These two wildly different predictions come from different
- 7:27assumptions about how metabolic rate
- 7:29scales as a function of mass.
- 7:31But notice in both cases, it's just proportional
- 7:34to mass raised to some power.
- 7:36These kinds of relationships are called power laws
- 7:38and all power laws have a special property.
- 7:41If you take the logarithm of the X and Y values,
- 7:44you get a straight line.
- 7:45And the slope of the line is equal
- 7:47to the exponent of the power law.
- 7:49This makes it easy to identify
- 7:51different types of power laws.
- 7:52If the slope is one, that's just everyday linear scaling.
- 7:55If it's less than one, like the surface law,
- 7:58that is called sublinear scaling.
- 8:00And if it's larger than one,
- 8:01that is known as superlinear scaling.
- 8:04For nearly a hundred years, biologists generally agreed
- 8:07that the two-thirds exponent
- 8:09of the surface law was the correct one for metabolic rate.
- 8:12But then in 1932, Swiss biologist, Max Kleiber,
- 8:16decided to put it to the test.
- 8:17He took the metabolic rates
- 8:19of different animals from a small dove at 150 grams
- 8:22to a large steer at 680,000 grams,
- 8:26and then he plotted them against their mass
- 8:28on a log log plot.
- 8:29And as expected, all the data did fall on a straight line.
- 8:32But the slope wasn't two-thirds.
- 8:35Instead, it was about three quarters.
- 8:37This became known as Kleiber's Law.
- 8:40It implies that if you double an animal's mass,
- 8:42metabolic rate goes up by about 1.68,
- 8:45an increase of 68% instead of the 59%,
- 8:49which would be predicted by the two-thirds scaling law.
- 8:51So according to Kleiber's Law,
- 8:53an elephant burns roughly 178 times
- 8:56as many calories as a cat, or about 45,000 kilocalories.
- 9:00And the actual LSD dose Tusko should have received
- 9:03was 53 milligrams, which is around a sixth of the dose
- 9:06the researchers gave him.
- 9:08So that explains the dosing catastrophe
- 9:10with Tusko The Elephant.
- 9:13(mellow music)
- 9:13Kleiber's original work was based on a small data set,
- 9:16primarily made up of mammals.
- 9:18But if you plot the data for a larger range of mammals,
- 9:21as well as for other animals
- 9:22like birds, reptiles, and fish,
- 9:24you find that they all follow
- 9:25roughly the same scaling relationship.
- 9:27Now, they don't all fit perfectly onto the same line
- 9:29because the base metabolic rate
- 9:31for warm-blooded animals is higher
- 9:32than for cold-blooded ones,
- 9:34but they all scale according to the same power law,
- 9:37mass raised to the three-quarters.
- 9:39Some argue that this relationship expands all the way down
- 9:42to single cells and the molecular machines inside of them.
- 9:45If that's true, Kleiber's Law governs life
- 9:47spanning more than 25 orders of magnitude.
- 9:51But then the question is,
- 9:53if an elephant has so many more cells
- 9:56and each cell is a similar size
- 9:58to a cell in a smaller creature,
- 10:00and yet it's using proportionately less energy,
- 10:05like it's got a lower metabolic rate per pound
- 10:08or per kilo or per cell,
- 10:10that's really saying that those cells
- 10:13are functioning with much less energy.
- 10:15- It's like there's some efficiency to being big,
- 10:18and it's hard to understand why.
- 10:20What is it that the cells are providing to each other,
- 10:24somehow cooperating in some way?
- 10:26- I just put in my calculator 100 to the three quarters,
- 10:29so if you got an organism that's 100 times bigger,
- 10:32then according to this, only 31.6
- 10:35times the metabolic rate.
- 10:37So it seems like a big savings.
- 10:40- It's a massive improvement, right?
- 10:42It's a big savings.
- 10:43It's a biological fact,
- 10:44but people have been arguing now for a century
- 10:48what accounts for this three quarters.
- 10:50(mellow music)
- 10:50- In the decades following Kleiber's observation,
- 10:52the mystery only deepened.
- 10:54Researchers discovered that brain size also roughly scales
- 10:58as mass to the three quarters,
- 10:59and so does an animal's growth rate
- 11:01and the amount of blood pumped per minute.
- 11:04But not every property scales as mass to the three quarters.
- 11:07- If you ask how long a creature will live, a mammal,
- 11:11that tends to be proportional to its mass
- 11:13to the one quarter power.
- 11:15One over four, not three over four.
- 11:18- [Host] So that means if you double the mass of a mammal,
- 11:20then on average, its lifespan is around 19% longer.
- 11:24Similarly, blood circulation time also scales
- 11:26as roughly mass to the one quarter, while breathing rate
- 11:29and heart rate both scaled to the negative one quarter.
- 11:32They're not all three quarters power laws,
- 11:35but they are all multiples of a quarter.
- 11:37So the question on everyone's mind was
- 11:40where are these quarter power scaling laws coming from?
- 11:43- One popular theory emerged in the 1990s.
- 11:47(mellow music) Brian Enquist was studying
- 11:48for an undergraduate degree in biology.
- 11:50- Most of the biology classes then that you take,
- 11:53you learn about the Krebs Cycle
- 11:55and then you have to memorize all the different parts
- 11:57of a flower and then all...
- 11:59They have a different terminology for everything.
- 12:00- [Henry] But then one day he took a zoology class
- 12:03where they showed him those quarter power scaling plots.
- 12:05- [Brian] I just couldn't believe it.
- 12:07I was like, "You got to be kidding me."
- 12:09You know, this is kind of like something fundamental
- 12:11that's kind of like underlying biological diversity.
- 12:14And there I knew immediately that I wanted
- 12:17to kind of quit my specialized
- 12:19kind of plant physiology research
- 12:21and do something associated with scaling.
- 12:24(mellow music)
- 12:24- [Henry] So Enquist started studying for his PhD,
- 12:27under Professor James Brown,
- 12:29who had been thinking about scaling laws for years.
- 12:31So they knew that often when power laws appear,
- 12:34there's some form of self-similarity
- 12:35in the underlying system.
- 12:36So they wondered, what could that self-similarity be?
- 12:39And they suspected that it might have something to do
- 12:41with the way resources are transported through the body,
- 12:44specifically the networks that do this.
- 12:46They had the biological intuition,
- 12:48but to get a complete theory,
- 12:49they needed a formal mathematical framework.
- 12:52In other words, they needed a mathematician
- 12:54or a theoretical physicist.
- 12:55- And at the time, Jim was associated
- 12:58with the Santa Fe Institute,
- 13:00and he started asking around with,
- 13:02is there anyone up here that's interested in these
- 13:04biological scaling relationships?
- 13:06And then the president at the time said, "You know what?
- 13:08I know of this physicist who's up at Los Alamos
- 13:12who was talking about biological scaling relationships."
- 13:15And so we met Geoffrey and it was like immediately,
- 13:18it's like we'd been talking about the same things
- 13:21for like years.
- 13:22It's like, you know, finding,
- 13:25someone who's been like speaking your language,
- 13:27but no one could understand you.
- 13:28- [Henry] So West, Brown and Enquist teamed up
- 13:30to try and find a compelling explanation for Kleiber's Law.
- 13:33- They started by assuming three simple premises.
- 13:37The first premise is that the networks
- 13:39that distribute resources are space-filling,
- 13:41since they need to reach every cell in the body.
- 13:44The second premise is that the terminal units
- 13:46of those networks, the thinnest segments
- 13:48on the outer periphery of the delivery system,
- 13:50have the same width regardless of the size of the organism.
- 13:53That is the outermost blood vessels that carry nutrients
- 13:56to an elephant's skin cells are about as thick
- 13:58as the ones in a mouse.
- 14:00The elephant just has many more of them.
- 14:02And the third premise is that over time,
- 14:04evolution has driven these transport networks
- 14:06toward an efficient design.
- 14:08So what should such a network look like?
- 14:11(mellow music)
- 14:11Well, intuitively, to get fuel from one place to another
- 14:14as efficiently as possible,
- 14:15you want the path to be as short as possible.
- 14:17So basically a straight line.
- 14:18But because fuel needs to reach every part of the body,
- 14:21you would also need many different paths.
- 14:23And as you go to larger and larger organisms,
- 14:25the networks inside need to
- 14:27reach a larger and larger volume.
- 14:29One way to do this is to stretch all the paths
- 14:31and match the growth of the animal.
- 14:33In this case, the volume of the animal
- 14:35should scale as these internal path lengths cubed.
- 14:38Or if we rearrange that, internal path length should scale
- 14:40as volume to the one-third,
- 14:42just like the overall length of the animal does.
- 14:45But this design is incredibly wasteful.
- 14:47Take these two regions.
- 14:49The two vessels that are bringing blood here
- 14:51go through almost the same path in the body,
- 14:53and they only split up just
- 14:55before they reach their destinations.
- 14:57If instead we had just one vessel up to this point
- 14:59and split it only when the paths needed to diverge,
- 15:02we could serve the two regions
- 15:03using a lot less vessel material
- 15:05and a lot less blood to fill the vessels.
- 15:07Of course, you can extend this logic
- 15:09for all the blood vessels in the body,
- 15:10and you end up with a much more efficient design
- 15:13of branching blood vessels.
- 15:15But now we run into another issue
- 15:17because each branching point provides an opportunity
- 15:19for some of the blood to bounce back, that is reflect.
- 15:22If you have many reflections,
- 15:24that would mean it costs significantly more
- 15:25energy to pump blood around.
- 15:27So next, they argue that nature should favor structures
- 15:30that minimize reflections.
- 15:31And as it turns out, this happens
- 15:33if the cross-sectional area of the vessels
- 15:35stays the same before and after the branching.
- 15:37So if you've got a cross-sectional area
- 15:39of two centimeters squared for the main vessel,
- 15:41then each of the two branches need to have an area
- 15:43of one centimeter squared each, for large vessels at least.
- 15:47For smaller ones, daughter branches can be a bit thicker
- 15:50to allow blood to slow down
- 15:52and exchange resources with the tissue it's reaching.
- 15:55If you keep repeating this pattern across the network,
- 15:57you end up with this, a branching self-similar fractal.
- 16:01And if you look at the actual shape
- 16:03of the circulatory system, it has this geometry.
- 16:06So it seemed like they were onto something.
- 16:09- But how do you get from this
- 16:11to quarter power scaling laws?
- 16:13Well, mathematician Felix Hausdorff discovered
- 16:15that self-similar fractals have an interesting property.
- 16:18(mellow music) Take a straight line segment.
- 16:19It's completely one-dimensional and not a fractal at all.
- 16:22Hausdorff assigned this line segment a value of 1.0.
- 16:26But now imagine adding a few bends
- 16:29and more bends to those bends.
- 16:31If you keep doing this, the line becomes more
- 16:33and more fractal-like.
- 16:35Eventually, if you keep applying
- 16:36the right kind of contortions,
- 16:38that one-dimensional line segment fills up
- 16:40an entire region of the 2D plane.
- 16:42Hausdorff assigned these space-filling fractal curves
- 16:45a value of 2.0, corresponding to their dimensionality.
- 16:48The same ideas apply to a 2D surface.
- 16:51- So think about a piece of paper, right?
- 16:53Two-dimensional.
- 16:55All then the fractal network is doing is crumpling up
- 16:58that sheet of paper, and it effectively fills
- 17:01a ball of volume.
- 17:02So you can now describe that sheet of paper
- 17:05as a sphere instead of a two-dimensional sheet of paper.
- 17:09- [Henry] Repeat the right pattern of folds
- 17:10at smaller and smaller scales, and a 2D surface
- 17:13fills more and more of a 3D volume.
- 17:15In the mathematical limit, it becomes space filling
- 17:18with Hausdorff dimension of 3.0.
- 17:20- [Brian] You know, biologically, I said,
- 17:22"Well, what does that mean?"
- 17:23That enables an organism for a given size
- 17:26to pack in more of these metabolic surface areas
- 17:30than would be expected.
- 17:32And it's because of this fractal-like structure
- 17:35that enables you then to pack in and have all of these folds
- 17:38and convolutions and on top of each other
- 17:41to pack in an enormous amount of membrane surfaces.
- 17:44- As a result, the Hausdorff dimension of the surface
- 17:47of the circulatory system is roughly three,
- 17:50meaning its surface area doesn't scale
- 17:52as its length squared, but it's length cubed.
- 17:55And since the metabolic rate hinges on
- 17:57how fast resources can be exchanged across the surface area,
- 18:00well, it must also scale like length cubed.
- 18:03But remember, West, Brown and Enquist
- 18:05wanted to explain Kleiber's Law.
- 18:07So they needed to know how metabolic rate scales with mass.
- 18:12(mellow music)
- 18:12Since every cell needs to be served by the network,
- 18:14this means that the volume
- 18:15around the network should be proportional
- 18:17to the animal's mass.
- 18:18And since volume is just surface area times length
- 18:21and surface area is proportional to length cubed,
- 18:23that means both volume and mass
- 18:25must be proportional to length to the fourth,
- 18:28which can be rewritten to show that length is proportional
- 18:30to mass raised to the one quarter.
- 18:33And if you plug that into the equation
- 18:34for the metabolic rate,
- 18:35you find that the metabolic rate
- 18:36must be proportional to mass to the three-quarters,
- 18:40exactly as Max Kleiber had found.
- 18:43West, Brown and Enquist published their work in 1997,
- 18:46and it soon came to be known
- 18:48as WBE Theory, after their initials.
- 18:50- And the theory is quite rigid.
- 18:53It makes very specific predictions.
- 18:55This is good science.
- 18:57Okay?
- 18:57This is sticking your neck out,
- 18:59and it's an incredibly beautiful theory.
- 19:02(mellow music) - [Derek] Take a look at this table
- 19:03from Geoffrey West's book.
- 19:04These are the scaling exponents WBE theory predicts,
- 19:07including many that are not multiples of a quarter,
- 19:10but all follow from the same theory.
- 19:12For example, the radius of an animal's aorta
- 19:15should scale with its mass to the three-eighths or .375.
- 19:18And the area of its lungs should scale with mass
- 19:21to the 11-12ths or about 0.92.
- 19:24In total, this chart makes 26 different predictions.
- 19:27Now, these are the observed data.
- 19:30Radius of the aorta, 0.36.
- 19:32Lung area, 0.95.
- 19:35- [Steven] That's the really shocking thing
- 19:37about what they did.
- 19:37They had a table with something like, I don't know,
- 19:3920 or 30 predictions of exotic exponents,
- 19:43and that's what you really see in the data.
- 19:45So this one theory accounts not only
- 19:46for the three-quarters power of metabolism,
- 19:49but for literally dozens of other things
- 19:51that biologists have measured.
- 19:53- Some of the scaling laws are easy to explain
- 19:56once you've got the three-quarters law for metabolism.
- 19:58Take a mammal's heart rate, for instance.
- 20:01Heart rate is equal to the blood flow rate
- 20:03over the amount or volume of blood in every beat.
- 20:05(mellow music)
- 20:06The volume of blood per beat has been found
- 20:08to scale in direct proportion to an animal's mass.
- 20:10So that's just M.
- 20:11And the blood flow rate?
- 20:13Well, remember that metabolism is all about
- 20:15how nutrients get distributed around the body.
- 20:17So most biologists agree metabolic rate
- 20:19and blood flow rate are directly
- 20:20proportional to one another.
- 20:22So heart rate should scale as metabolic rate over mass.
- 20:25Swapping in the scaling law Kleiber had found,
- 20:28that gives us M to the minus one quarter,
- 20:30meaning bigger animals should have slower
- 20:32heartbeats than smaller ones.
- 20:34And this is exactly what we observe in nature.
- 20:36The world's smallest mammal, the Etruscan Shrew,
- 20:39has an extraordinary heart rate of 1200 beats per minute.
- 20:42That's 20 beats per second.
- 20:44Whereas the biggest land mammal, the African bush elephant,
- 20:47has a typical heart rate of only 30 beats per minute.
- 20:50And we can do something similar for lifespan.
- 20:53One of the leading theories is
- 20:54that an animal's lifespan is based on
- 20:56the accumulation of metabolic damage.
- 20:58That is, as each chunk of tissue in an organism
- 21:01processes nutrients over time,
- 21:03this causes damage to accumulate,
- 21:05and that over time causes the animal to die.
- 21:08So the rate at which an animal accumulates damage
- 21:10is its metabolic rate per unit of mass,
- 21:12and its lifespan should be the inverse of that rate.
- 21:15If damage accumulates faster, it dies sooner.
- 21:17If it accumulates slower, it lives longer.
- 21:20So lifespan is proportional to M over B,
- 21:23or substituting in Kleiber's Law, M to the one quarter.
- 21:27So lifespan should increase with mass.
- 21:30And you do see this in nature.
- 21:31(mellow music)
- 21:31A shrew only lives for one to two years in the wild
- 21:34while a mighty African elephant can live up to 70 years.
- 21:38So if you're a small mammal,
- 21:39you have many heartbeats per minute,
- 21:41but you live a relatively short life.
- 21:43Conversely, if you're a larger mammal,
- 21:45your heart beats much slower and you live a lot longer.
- 21:48- So it's the, you know, live fast and burnout
- 21:52and die young, right?
- 21:54Or spend it frugally and live a really long life.
- 21:57- But you might have also noticed something else.
- 22:00Heart rate scales as B over M.
- 22:02So it equals B over M times some constant.
- 22:05And lifespan scales as M over B.
- 22:08They're inverses of each other.
- 22:09They scale in equal and opposite directions.
- 22:12Now, the total number of heartbeats in an animal's life
- 22:15is just the heart rate multiplied by the lifespan.
- 22:17So when you multiply these two terms, they cancel out,
- 22:20leaving you with just a constant.
- 22:22So that suggests that no matter
- 22:24what mammal you're talking about,
- 22:26it should have roughly the same number of heartbeats.
- 22:29You can find that number by just plugging in some examples.
- 22:32Let's start with the Etruscan Shrew.
- 22:35The shrew's 1200 beats per minute multiplied by a lifespan
- 22:37of around 1.5 years gives you
- 22:39around 950 million heartbeats
- 22:41in the course of the shrew's life.
- 22:43Meanwhile, an African elephant's 30 beats per minute
- 22:45multiplied by a lifespan of around 65 years
- 22:47gives you a little over a billion heartbeats.
- 22:51And we could keep going.
- 22:52(upbeat music) But for nearly every mammal you look at,
- 22:54you keep landing at the same figure of
- 22:56around a billion heartbeats.
- 22:58This is why nearly every mammal from a tiny field mouse
- 23:01to a gazelle, from a cheetah to a hippopotamus,
- 23:05they all get around a billion heartbeats
- 23:07between the day they're born and the day they die.
- 23:12But there is one major outlier, one mammal
- 23:15that gets significantly more than a billion heartbeats.
- 23:18And that is us, humans. (mellow music)
- 23:21We are the lucky ones.
- 23:23Three centuries ago, humans were much closer
- 23:26to the standard value of one billion heartbeats.
- 23:28But around the mid 1800s, germ theory
- 23:31and better sanitation methods became widespread,
- 23:33causing a stark decrease in the number of child mortalities
- 23:36and deaths from disease.
- 23:37So life expectancy began to climb up.
- 23:39And with it, the average number of heartbeats in a lifetime.
- 23:42If you look closely, you can also see some significant drops
- 23:46like this 1918 dip from the Spanish flu pandemic
- 23:49or over here, what appears to be
- 23:50the impact of the Second World War.
- 23:52But overall, the trend is clear.
- 23:54We have systematically been increasing the number
- 23:57of heartbeats we get in our lifetime to the point
- 23:59where now the average human gets nearly three billion
- 24:02heartbeats before they die.
- 24:05I don't know if there's a better argument for science
- 24:07and technology than this.
- 24:09It has literally given the average human
- 24:11more than a full extra life.
- 24:13And it's not just humans.
- 24:14Other mammals have been observed
- 24:16to have much longer lives in captivity when they're away
- 24:18from the hazards they would naturally encounter in the wild.
- 24:22Or if you look at it purely from the number of years we get,
- 24:24we now have the lifespan of a much larger mammal,
- 24:27somewhere between an elephant and a whale.
- 24:31But there is one curious thing about this trend.
- 24:33(mellow music) Take a look at this graph.
- 24:35It looks surprisingly similar to the graph from before.
- 24:38In fact, if you overlay them, they look remarkably similar.
- 24:41Now I want you to take a guess at what this graph is.
- 24:44Have you got your answer?
- 24:46It is the number of people living in cities.
- 24:49For these two charts, we're using data from parts
- 24:52of the United Kingdom where the record keeping
- 24:54goes back several centuries,
- 24:56but the rest of the world has followed similar trends.
- 24:59Of course, that doesn't mean
- 25:00cities cause people to live longer,
- 25:03but it goes against the perception of cities
- 25:05as being full of pollution and breeding grounds for disease.
- 25:09Already in 1889, a medical doctor wrote,
- 25:12"The poisonous germs and pollutions of the city,
- 25:14it's impure air and water, bad sewage,
- 25:17and endless nuisances."
- 25:19So how do you reconcile these two views?
- 25:22Or more specifically, is there any quantitative data
- 25:25on how smaller cities compare to larger ones?
- 25:28- It turns out this is something that Geoffrey West pursued
- 25:31after his work with Brown and Enquist.
- 25:33He and collaborators like Luis Bettencourt and others
- 25:37have looked at scaling laws in cities.
- 25:39(mellow music)
- 25:39- [Host] They looked at how different properties
- 25:41like the amount of crime scale
- 25:43as the population of the city increases,
- 25:45and what they found is that if you plot
- 25:46serious crimes on a log log plot,
- 25:48the data clusters around a straight line
- 25:50with a slope of 1.15,
- 25:52meaning crime grows faster than linear or super linear.
- 25:56So for every doubling of a city's population,
- 25:59you get around 2.2 times as many criminal cases
- 26:02or around 120% more crime as opposed to the 100%
- 26:07you might naively expect.
- 26:08To make matters worse, researchers found
- 26:10that the same general pattern
- 26:11holds for the amount of wastewater
- 26:13and even the number of AIDS cases.
- 26:16The exact exponents vary a little,
- 26:18but overall, as cities grow larger,
- 26:20you systematically get more of each.
- 26:23So it seems like that medical doctor was onto something.
- 26:26And you might think life on earth would be better off
- 26:28if we all lived in small towns instead,
- 26:31but that might not be the case.
- 26:33(mellow music) In 2006, Dirk Helbing, Christian Kuhnert,
- 26:36and Geoffrey West looked at how the number
- 26:38of gas stations scales as a function of population.
- 26:41- If a city is twice as big, does it need twice
- 26:44as many gas stations?
- 26:45Because, you know, we have to supply not exactly nutrients,
- 26:48but energy, gas, for all those cars.
- 26:51- [Host] The naive expectation is that
- 26:53if you double the number of cars,
- 26:54you're going to need to double the amount of fuel,
- 26:56so double the gas stations.
- 26:58To find out whether this was true,
- 26:59they plotted the data on a log log plot
- 27:01and found a straight line.
- 27:03But the exponent wasn't one, it was about 0.8.
- 27:07This means that for every doubling, you only need around 74%
- 27:11more gas stations, which is a decent savings.
- 27:14(mellow music) The amount of roads and electrical cables
- 27:17also scale in roughly the same way.
- 27:19The rough figure that West gives in his book
- 27:21is that they all have scaling exponents of around 0.85.
- 27:23- It is interesting that some of the things
- 27:25that we can share, like you can drive
- 27:28on the road, but so can I.
- 27:29When it's shared resources, yes,
- 27:32cities can be surprisingly green.
- 27:34The argument is that cities can be even greener
- 27:36than you might think than living out
- 27:38in the middle of nowhere.
- 27:39- But cities have even bigger benefits.
- 27:42Things like total wages, GDP,
- 27:44and the number of patents all scale superlinearly,
- 27:47with exponents that cluster somewhere around 1.15,
- 27:51meaning that for every doubling in size,
- 27:53you get around 120% more of each.
- 27:57All of this becomes especially significant
- 27:59when you compare a small town of say 50,000 people
- 28:01to a city 100 times its size
- 28:04because infrastructure needs only need to go up by a factor
- 28:06of about 50 while total wages, GDP, patents and inventions,
- 28:11they all go up by a factor of 200.
- 28:13Unfortunately, disease and crime also go up
- 28:16by the same factor.
- 28:18Or look at it this way, on a per person basis,
- 28:21you'd only need about half the infrastructure,
- 28:23while you get double all the socioeconomic factors.
- 28:27So cities far from being detrimental to the world,
- 28:29they might actually be one of our best inventions
- 28:32and an indirect driver of a lot of scientific
- 28:35and technological progress.
- 28:36Perhaps this is also why people often say
- 28:38that life in the city feels faster.
- 28:41A feeling that seems justified
- 28:42because researchers looked at
- 28:44how fast people walk in cities of different sizes.
- 28:47And they found that people literally do walk
- 28:49faster in larger cities.
- 28:51- That turns out to depend on city size.
- 28:54It's not just that the sidewalks are congested or not.
- 28:57It's just like the vibe gets people amped up.
- 29:00They move faster in cities.
- 29:02- So the pace of life seems to be increasing,
- 29:05but that may come at a cost
- 29:07because as one person put it, "everything nowadays is ultra.
- 29:11Everything is being transcended continually
- 29:13in thought as well as in action.
- 29:14No one knows himself any longer.
- 29:16Young people are stirred up much too early in life
- 29:19and then carried away in the world of the times.
- 29:21Wealth and rapidity are what the world admires".
- 29:24Could it be that life is accelerating so fast
- 29:26that humans won't be able to keep up?
- 29:29Well, probably not
- 29:30because this quote was written in 1825
- 29:32by Wolfgang von Goethe.
- 29:34For the past 200 years and probably longer,
- 29:38almost every generation has felt like life was accelerating.
- 29:41And yet every generation has managed.
- 29:43So I think it's likely we can continue
- 29:45to adapt indefinitely.
- 29:47And as cities continue to grow in size,
- 29:50all of us will continue to reap the benefit
- 29:52of economies of scale in much the same way
- 29:54that mammals benefit from being larger.
- 29:57But while WBE Theory seems to predict
- 29:59where the exponents in biological scaling laws come from,
- 30:02for cities, there is no widely accepted
- 30:04explanatory theory yet.
- 30:06Trying to explain where those exponents of 0.85
- 30:09and 1.15 come from is one of the big goals for theorists.
- 30:12Although even WBE Theory is not universally accepted.
- 30:16For one, the fact it predicts the right exponents
- 30:19doesn't necessarily mean the theory itself is correct.
- 30:22There are a few other theories that predict
- 30:24some of the same scaling exponents,
- 30:26and there are also some other critiques.
- 30:29- There's a lot of discussion.
- 30:31It may look convincing.
- 30:32And personally, I tend to think it is very impressive.
- 30:35But I have very good colleagues like Peter Dodds
- 30:37at University of Vermont,
- 30:39and he thinks that a lot of the data analysis
- 30:41is either not done exactly right
- 30:43or that the data are so noisy
- 30:46that you shouldn't really take this so seriously.
- 30:49- Dodds even argues that Kleiber's Law
- 30:51itself might not be true. (mellow music)
- 30:53- In the 1960s, there's a symposium on energy metabolism,
- 30:59some name like this in animals,
- 31:02and at the end of it, they vote 29 to zero
- 31:04that it's going to be three quarters, right?
- 31:06Because, you know, you got to set some rules.
- 31:09If you go back and look at the data,
- 31:10which no one is really doing anymore, right,
- 31:12the data does not work.
- 31:13Like it doesn't work.
- 31:15(mellow music)
- 31:15- [Henry] This is a graph of metabolic rate
- 31:17as a function of mass based on a study
- 31:19that looked at 391 species of mammals,
- 31:22much larger than Kleiber's range.
- 31:23And it looks like the three-quarter slope fits quite well.
- 31:26But this is just the top part of the chart,
- 31:28the biggest mammals.
- 31:30If you zoom out, you see that the rest of the mammals appear
- 31:32to fall on a line that's closer to two-thirds.
- 31:36And in recent studies of bird metabolism,
- 31:38you also find a slope that appears closer
- 31:40to two-thirds than three-quarters.
- 31:42Could it be that those French scientists
- 31:44from centuries ago were right,
- 31:45that metabolic rate really does
- 31:47just scale with surface area?
- 31:48Well, not so fast.
- 31:51(mellow music) For one, many recent studies
- 31:52of cold-blooded animals find slopes
- 31:54that are significantly higher than two-thirds.
- 31:56The bigger problem is that measuring
- 31:58metabolic rates is difficult.
- 31:59It generally involves putting animals in a container
- 32:02and taking very precise measurements
- 32:03of their heat production or oxygen consumption,
- 32:06all while ensuring that the animal is in an unstressed,
- 32:09low activity resting state.
- 32:10Unsurprisingly, this is particularly difficult
- 32:13to pull off with large animals.
- 32:15And as a result, many studies have found scaling exponents
- 32:17where the error bars include both
- 32:19two-thirds and three-quarters.
- 32:22Today, the research community is split.
- 32:24Many uphold Kleiber's Law and the three-quarter scaling,
- 32:26while others think it's two-thirds.
- 32:28But a growing number suspect
- 32:29that there is no universal scaling exponent
- 32:32across all of life.
- 32:33In fact, it may very well be the case
- 32:35that the metabolic rate of larger mammals
- 32:37scales as their mass to the three-quarters,
- 32:39while for smaller ones, it's their mass to the two-thirds.
- 32:41- You know, it's like everything in science
- 32:43that there are people arguing, and that's good.
- 32:45- I guess the bigger exhortation
- 32:47would be someone to really measure it,
- 32:49measure things beautifully.
- 32:50You know, we're in 2026,
- 32:52has to be not just one elephant at one zoo,
- 32:55it needs to be measured again really well.
- 32:57- [Host] What everyone does agree on
- 32:58is that scaling laws are real.
- 33:00- Life is not linear all the time.
- 33:03There are deals to be had.
- 33:04There's often in real life departures from proportionality,
- 33:08and you have to be aware of it.
- 33:10Sometimes things punch above their weight,
- 33:12and as you get bigger, you get more efficient.
- 33:15Sometimes there are detriments to size.
- 33:18- So it pays to know how things scale.
- 33:21Clearly from an energy efficiency perspective,
- 33:23animals benefit from being larger.
- 33:26Similarly, all of us potentially stand
- 33:28to gain from having more people living in larger cities.
- 33:31It could lead to more discoveries and inventions,
- 33:34and in doing so, improve the standard of living
- 33:36for all of us, which might even give us more
- 33:39heartbeats in our lifetimes.
- 33:44From the surface law to Kleiber's law to WBE theory,
- 33:48how metabolic rate scales with mass
- 33:50has been one of biology's biggest debates for centuries.
- 33:52By collecting better data and analyzing it carefully,
- 33:55there's a good chance
- 33:56that the next generation of researchers could be the ones
- 33:58to finally put this debate to rest.
- 34:00It could be one of you watching or a student that you know.
- 34:04And today's sponsor, Brilliant, is helping
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