Wednesday, June 29, 2016

Why Can’t We Just Go With the Flow? – The High Cost of Heating

Biology concepts – endothermy, ectothermy, poikilothermy, thermoregulation

So you are lying in bed, cold and hungry, contemplating living like a monk just so you can have more time to live like a monk (see this post on increasing life span via reduced core temperature and caloric restriction). Your temperature is going down after bedtime and coming back up in the early morning. This implies that you can control and maintain a constant temperature - pretty impressive. But don’t get a big head, most mammals can do it, even your pet hamster.


The hypothalamus is located in the lower center of the
brain. Different parts are involved in sensing and
regulating temperature, but also for blood pressure,
circadian rhythms, and feeling full after eating.
To be able to control your temperature (thermoregulate), you must know what your temperature is in the first place. Mammals have sensors in their skin and organs which relay information about temperature to the hypothalamus of the brain. The neural sensors in skin (peripheral thermoreceptors) sense the temperature just under the surface. This can be quite different from the core temperature. Central thermoreceptors sense the temperature in the brain, bladder and muscles. Your hypothalamus sets your skin thermostat at about 72˚C, so you still feel hot when the ambient temperature is >75˚F even though your core temperature averages 98.6˚F (37˚C).

Heat is constantly being generated by your metabolism (The breaking down and building up of molecules in your cells). Burning ATP to produce work also produces heat as a byproduct, and this goes a long way to keeping our temperature around 98.6˚F. Generating internal heat to maintain a body temperature is called endothermy (endo = within, therm = heat). Mammals are endotherms, and we hold a constant temperature, so we are also homeotherms (homeo= same). However, we have seen that constant temperature is a relative term, since our circadian rhythm cycles our core temperature up and down as the day goes on.


This is an infrared image of the human body.
Temperatures increase from blue to green to
yellow to orange to red. The head is often one
of the warmer parts of the body since the brain
uses so much energy, while the testicles are
housed outside the body to keep them cooler. 
image source: http://www.medicalir.com/
In addition, holding a constant temperature doesn’t mean that all parts of the organism are the same temperature. Just like your skin is cooler than your core body temperature, parts of your internal body can be warmer than your average core temperature. During intense exercise, your muscle temperature can go to 107˚F or higher! On the other hand, sperm is damaged by high temperature, so the testicles are usually housed in an external pouch in order to keep their temperature one or two degrees below body temperature.

The higher than average temperature is O.K. for a short while or in a small part of the body, but if it involves too much volume or stays high for too long, then your core temperature can rise to dangerous levels (104˚F). On the other hand, having too low a temperature in any part of the human body can be dangerous. If ice crystals form in the cell, the jagged edges will cut the cell to ribbons and kill it; this is frostbite.

Your body thermoregulates to maintain a healthy temperature range. It finds ways to dissipate heat when the core temperature rises, such as sweating in humans, panting in dogs, or pushing more blood through the large ears of rabbits. If your core temperature is too low, you can generate heat by shivering (small muscle spasms that mean more ATP burned and more heat). Chattering teeth is just a spasm in the buccinator muscles of your jaw. These are pretty big muscles (bigger on some people I know) and can produce enough heat to keep your head warm.

Some endotherms are exquisitely adept at regulating the temperature in different parts of their body, and can save lots of energy through this differential regulation. Ground squirrels in hibernation reduce their abdominal temperature to match ambient temperature down to 0˚C, and some birds can hold a body temperature just one degree above freezing all night. These types of animals are referred to as endothermic poikilotherms (poikilo = varied).


Lizards are ectotherms, so they have temperatures near
ambient. In this image, the ambient temperature was
76.1 ˚F, so the lizard remains near that temperature.
However, the human hand is much warmer, as it has an
internal source of heat.
We are biased toward believing that all animals control their body temperature just because we do, but the vast majority of animals are ectotherms (ecto = outside). They get most of their heat from the environment, and this works for them.

Ectotherms like reptiles and insects will have low activity when it is cool, but absorbing heat by sunning themselves will speed them on their daily errands. This is because the rates of most cellular activities increase with temperature right up to the point of boiling, but low temperatures slow them down greatly. So most animals need an external source of heat to allow them to hunt, protect themselves, or seek shelter.

Some ectotherms, like moths and bees, can have their wing muscles go into spasms in order to generate enough heat for them to take off.  If they can raise their temperature in any way (sunning or spasming), they are called ectothermic poikilotherms, although some might call moths and bees partial endotherms, since the source of heat is internal. On the other hand, a few ectotherms like some fish, always have the same temperature as their environment no matter their activity or needs. These animals are referred to as ectothermic homeotherms.

There are many more ectotherms than endotherms in the world because it is a successful strategy for saving energy. It is extremely costly to maintain a high metabolic rate and a constant internal temperature, like running your furnace all winter to stay comfortable – we all know how expensive that can be. An adult human (endotherm) needs 1300-1800 kCal/day to maintain its temperature and activity, while a crocodile (ectotherm) of the same size requires only 60 kCal! Fewer calories needed means less energy expended hunting or foraging which makes surviving times easier when less food is available.

Even though they may be called cold-blooded, don’t assume that ectotherms are always cold.  Rimicaris exoculata, an ectothermic shrimp that lives next to hydrothermal vents (undersea volcanoes that spew superheated water), is happy with an internal temperature of 350˚C (662˚F). The water doesn’t boil because it is under so much pressure (for every 33 feet of water, the pressure doubles); otherwise they would be shrimp toast.



On the left is a deep-sea hydrothermal chimney called a black smoker. The temperature in the hot water column is near 700˚F. The right side image is the vent shrimp, Rimicaris exoculata. The bright spots are the dorsal eyespots and are rich in rhodopsin. They glow like cat’s eyes when light is shone on them. In the deep ocean, there is no light, so the
shrimp don’t glow normally.

Also don’t assume that ectotherms are looking for a way to warm up. Some fish are perfectly comfortable in antarctic waters at (-2˚C to -4˚C; the ocean water doesn’t freeze because the salt disrupts crystal formation). For example, Dissostichus mawsoni fish have proteins that help important molecules resist cold damage (heat shock proteins) and to stay functional at low temperatures (chaperonins). 


The left image shows Dissostichus mawsoni, the Antarctic toothfish,
swimming under an ice sheet. Up close, we can see the teeth, and
that he isn’t going to win any beauty contests.

D. mawsoni  also has an antifreeze protein in its blood that binds to ice crystals and keeps the fish from freezing solid. Now that’s cold-blooded. These notothenioid (notothen = “from the south” in Greek) fish are successful enough in this environment to make up 90% of the fish biomass in the Antarctic.

Unfortunately, the terms warm-blooded and cold-blooded have become popular for all organisms. This is wrong on so many levels. We think of snakes as cold-blooded, but on a hot sunny day, the internal temperature of a snake will be much higher than that of a mammal. And we already talked about birds, endotherms of the highest order, that can allow the temperature of their feet to come within a degree of freezing. Now, which is warm-blooded and which is cold blooded?

And where do these terms leave plants? They don’t have blood – so they can’t be cold-blooded or warm-blooded – but they are ectotherms. Some plants are even poikilotherms- they can generate some heat at certain points in their life cycle. They can’t maintain or regulate it, so they are still ectotherms, but let’s not be prejudiced against them by calling them cold-blooded.

Low and behold, there are exceptions to the rules of body temperature – wouldn’t you know it.  There is a plant that can maintain a constant temperature by producing heat – even if it is only for two days a year. And there is a mammal that seems to think ectothermy is the way to go. We’ll talk about these rule-breakers starting next time.



Shillito B, Le Bris N, Hourdez S, Ravaux J, Cottin D, Caprais JC, Jollivet D, & Gaill F (2006). Temperature resistance studies on the deep-sea vent shrimp Mirocaris fortunata. The Journal of experimental biology, 209 (Pt 5), 945-55 PMID: 16481583

Kiss AJ, Mirarefi AY, Ramakrishnan S, Zukoski CF, Devries AL, & Cheng CH (2004). Cold-stable eye lens crystallins of the Antarctic nototheniid toothfish Dissostichus mawsoni Norman. The Journal of experimental biology, 207 (Pt 26), 4633-49 PMID: 15579559



For more information, classroom activities, or laboratories on endothermy, ectothermy, or thermoregulation:

thermoregulation –

endothermy –

ectothermy –