Bigger animals have more cells than smaller animals. More cells means more cellular metabolism and so more heat produced. How does the amount of heat an animal produces vary with its size? We clearly expect it to go up with size, but does it increase in proportion to volume? Surface area? Something in between?
A first guess would be that metabolism (equivalently, heat produced) goes up in proportion to volume. If cells are all roughly the same size, then number of cells increases proportionately with volume. But heat is dissipated through the surface. Surface area increases in proportion to the square of length but volume increases in proportion to the cube of length. That means the ratio of surface area to volume decreases as overall size increases. The surface area to volume ratio for an elephant is much smaller than it is for a mouse. If an elephant’s metabolism per unit volume were the same as that of a mouse, the elephant’s skin would burn up.
So metabolism cannot be proportional to volume. What about surface area? Here we get into variety and controversy. Many people assume metabolism is proportional to surface area based on the argument above. This idea was first proposed by Max Rubner in 1883. Others emphasize data that supports the theory that suggests metabolism is proportional to surface area.
In the 1930’s, Max Kleiber proposed that metabolism increases according to body mass raised to the power 3/4. (I’ve been a little sloppy here using body mass and volume interchangeably. Body mass is more accurate, though to first approximation animals have uniform density.) If metabolism were proportional to volume, the exponent would be 1. If it were proportional to surface area, the exponent would be 2/3. But Kleiber’s law says it’s somewhere in between, namely 3/4. The image below comes from a paper by Kleiber from 1947.
The graph shows that on a log-log plot, the metabolism rate versus body mass for a large variety of animals has slope approximately 3/4.
So why the exponent 3/4? There is a theoretical explanation called the metabolic scaling theory proposed by Geoffrey West, Brian Enquist, and James Brown. Metabolic scaling theory says that circulatory systems and other networks are fractal-like because this is the most efficient way to serve an animal’s physiological needs. To quote Enquist:
Although living things occupy a three-dimensional space, their internal physiology and anatomy operate as if they were four-dimensional. … Fractal geometry has literally given life an added dimension.
The fractal theory would explain the power law exponent 3/4 simply: it’s the ratio of the volume dimension to the fractal dimension. However, as I suggested earlier, this theory is controversial. Some biologists dispute Kleiber’s law. Others accept Kleiber’s law as an empirical observation but dispute the theoretical explanation of West, Enquist, and Brown.
To read more about metabolism and power laws, see chapter 17 of Complexity: A Guided Tour.