Endotherm

An endotherm (Greek: endon = "within", thermē = "heat") is an organism that maintains its body at a metabolically favourable temperature, largely by the use of heat set free by its internal bodily functions instead of relying almost purely on ambient heat. Such internally generated heat is mainly an incidental product of the animal's routine metabolism, but under conditions of excessive cold or low activity an endotherm might apply special mechanisms adapted specifically to heat production. Examples include special-function muscular exertion such as shivering, and uncoupled oxidative metabolism such as within brown adipose tissue.

Mammals and birds are the only extant universally endothermic groups of animals. Certain lamnid sharks, tuna and billfishes exhibit exothermy.

In common parlance, endotherms are characterized as "warm-blooded." The opposite of endothermy is ectothermy, although there is no absolute or clear separation between the nature of endotherms and ectotherms in general.

Mechanisms

Generating and conserving heat

Sustained energy output of an endothermic animal (mammal) and an ectothermic animal (reptile) as a function of core temperature

Many endotherms have a larger number of mitochondria per cell than ectotherms. This enables them to generate heat by increasing the rate at which they metabolize fats and sugars. Accordingly, to sustain their higher metabolism, endothermic animals typically require several times as much food as ectothermic animals do, and usually require a more sustained supply of metabolic fuel.

In many endothermic animals, a controlled temporary state of hypothermia conserves energy by permitting the body temperature to drop nearly to ambient levels. Such states may be brief, regular circadian cycles called torpor, or they might occur in much longer, even seasonal, cycles called hibernation. The body temperatures of many small birds (e.g. hummingbirds) and small mammals (e.g. tenrecs) fall dramatically during daily inactivity, such as nightly in diurnal animals or during the day in nocturnal animals, thus reducing the energy cost of maintaining body temperature. Less drastic intermittent reduction in body temperature also occurs in other, larger endotherms; for example human metabolism also slows down during sleep, causing a drop in core temperature, commonly of the order of 1 degree Celsius. At other times of the day there may be other variations in temperature, usually smaller, either endogenous or in response to external circumstances or vigorous exertion, and either an increase or a drop.[1]

The resting human body generates about two-thirds of its heat through metabolism in internal organs in the thorax and abdomen, as well as in the brain. The brain generates about 16% of the total heat produced by the body.[2]

Heat loss is a major threat to smaller creatures, as they have a larger ratio of surface area to volume. Small warm-blooded animals have insulation in the form of fur or feathers. Aquatic warm-blooded animals, such as seals, generally have deep layers of blubber under the skin and any pelage that they might have; both contribute to their insulation. Penguins have both feathers and blubber; their feathers are scale-like and serve partly for insulation and for streamlining. Endotherms that live in very cold circumstances or conditions predisposing to heat loss, such as polar waters, tend to have specialised structures of blood vessels in their extremities that act as heat exchangers. The veins are adjacent to the arteries full of warm blood. Some of the arterial heat is conducted to the cold blood and recycled back into the trunk. Birds, especially waders, often have very well-developed heat exchange mechanisms in their legs — those in the legs of emperor penguins are part of the adaptations that enable them to spend months on Antarctic winter ice.[3][4] In response to cold many warm-blooded animals also reduce blood flow to the skin by vasoconstriction to reduce heat loss. As a result, they blanch (become paler).

Avoiding overheating

In equatorial climates and during temperate summers, overheating (hyperthermia) is as great a threat as cold. In hot conditions, many warm-blooded animals increase heat loss by panting, which cools the animal by increasing water evaporation in the breath, and/or flushing, increasing the blood flow to the skin so the heat will radiate into the environment. Hairless and short-haired mammals, including humans, also sweat, since the evaporation of the water in sweat removes heat. Elephants keep cool by using their huge ears like radiators in automobiles. Their ears are thin and the blood vessels are close to the skin, and flapping their ears to increase the airflow over them causes the blood to cool, which reduces their core body temperature when the blood moves through the rest of the circulatory system.

Advantages and disadvantages of an endothermic metabolism

The major advantage of endothermy over ectothermy is decreased vulnerability to fluctuations in external temperature. Regardless of location (and hence external temperature), endothermy maintains a constant core temperature for optimum enzyme activity.

Endotherms control body temperature by internal homeostatic mechanisms. In mammals two separate homeostatic mechanisms are involved in thermoregulation - one mechanism increases body temperature, while the other decreases it. The presence of two separate mechanisms provides a very high degree of control. This is important because the core temperature of mammals can be controlled to be as close as possible to the optimum temperature for enzyme activity.

The overall rate of an animal's metabolism increases by a factor of about two for every 10 °C (18 °F) rise in temperature, limited by the need to avoid hyperthermia. Endothermy does not provide greater speed in movement than ectothermy (cold-bloodedness)—ectothermic animals can move as fast as warm-blooded animals of the same size and build when the ectotherm is near or at its optimum temperature, but often cannot maintain high metabolic activity for as long as endotherms. Endothermic/homeothermic animals can be optimally active at more times during the diurnal cycle in places of sharp temperature variations between day and night and during more of the year in places of great seasonal differences of temperature. This is accompanied by the need to expend more energy to maintain the constant internal temperature and a greater food requirement.[5] Endothermy may also provide a protection against fungal infection. While tens of thousands of fungal species infect insects, only a few hundred target mammals, and often only those with a compromised immune system. A recent study[6] suggests fungi are fundamentally ill-equipped to thrive at mammalian temperatures. The high temperatures afforded by endothermy might have provided an evolutionary advantage.

Ectotherms will increase their body temperature mostly through external heat sources such as sunlight energy, therefore they depend on the occurring environmental conditions to reach operational body temperatures. Endothermic animals mostly use internal heat production through metabolic active organs and tissues (liver, kidney, heart, brain, muscle) or specialized heat producing tissues like brown adipose tissue (BAT). In general, endotherms therefore have higher metabolic rates than ectotherms at a given body mass. As a consequence they would also need higher food intake rates, which may limit abundance of endotherms more than ectotherms.

Because ectotherms depend on environmental conditions for body temperature regulation, they typically are more sluggish at night and in the morning when they emerge from their shelters to heat up in the first sunlight. Foraging activity is therefore restricted to the day time (diurnal activity patterns) in most vertebrate ectotherms. In lizards, for instance, only a few species are known to be nocturnal (e.g. many geckos) and they mostly use 'sit and wait' foraging strategies that may not require body temperatures as high as those necessary for active foraging. Endothermic vertebrate species are therefore less dependent on the environmental conditions and have developed a high variability (both within and between species) in their diurnal activity patterns.[7]

It is thought that the evolution of endothermia was crucial in the development of mammalian species diversity in the Mesozoic period. Their endothermic capabilities provided them with a benefit over the mostly ectothermic dinosaurs that dominated the Mesozoic era. Endothermia gave the early mammals the capacity to be active during night time and avoid higher predation risk during the day. Endothermia therefore may have contributed to the fact that most mammalian taxa were already present at the end of the Mesozoic (66 million years ago) and that most extant mammalian taxa thus went through an evolutionary bottle neck (the nocturnal bottleneck hypothesis).[7]

Facultative endothermy

Many insect species are able to maintain a thoracic temperature above the ambient temperature using exercise. These are known as facultative or exercise endotherms.[8] The honey bee, for example, does so by contracting antagonistic flight muscles without moving its wings (see insect thermoregulation).[9][10][11] This form of thermogenesis is, however, only efficient above a certain temperature threshold, and below about 9–14 °C (48–57 °F), the honey bee reverts to ectothermy.[10][11][12]

Contrast between thermodynamic and biological terminology

Because of historical accident, students encounter a source of possible confusion between the terminology of physics and biology. Whereas the thermodynamic terms "exothermic" and "endothermic" respectively refer to processes that give out heat energy and processes that absorb heat energy, in biology the sense is effectively inverted. The metabolic terms "ectotherm" and "endotherm" respectively refer to organisms that rely largely on external heat to achieve a full working temperature, and to organisms that produce heat from within as a major factor in controlling their body temperatures.

References

  1. ↑ Refinetti, Roberto (2010). "The circadian rhythm of body temperature". Frontiers in Bioscience 15: 564–594. doi:10.2741/3634.
  2. ↑ http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/H/HeatTransport.html
  3. ↑ Thomas, D.B.; Fordyce, R.E. (2008). "The heterothermic loophole exploited by penguins". Australian Journal of Zoology 55: 317–321. doi:10.1071/ZO07053.
  4. ↑ Thomas, D.B., D.T. Ksepka and R.E. Fordyce. 2010. Penguin heat-retention structures evolved in a greenhouse Earth. Biology Letters (published online before print December 22, 2010, doi:10.1098/rsbl.2010.0993)
  5. ↑ Campbell, N. A.; Reece, J. B. et al. (2002). Biology (6th ed.). Benjamin/Cummings. p. 845.
  6. ↑ Robert, Vincent A. and Casadevall, Arturo (2009). "Vertebrate Endothermy Restricts Most Fungi as Potential Pathogens". The Journal of Infectious Diseases 200 (10): 1623–1626. doi:10.1086/644642. PMID 19827944.
  7. ↑ 7.0 7.1 Hut, RA; Kronfeld-Schor, N; van der Vinne, V; De la Iglesia, H (2012). "In search of a temporal niche: environmental factors". Progress in brain research. Progress in Brain Research 199: 281–304. doi:10.1016/B978-0-444-59427-3.00017-4. ISBN 9780444594273. PMID 22877672.
  8. ↑ Davenport, J. (1992). Animal life at low temperature. London: Chapman & Hall.
  9. ↑ Kammer, A. E.; Heinrich, B. (1974). "Metabolic rates related to muscle activity in bumblebees". Journal of Experimental Biology 6 (1): 219–227.
  10. ↑ 10.0 10.1 Lighton, J. R. B.; Lovegrove, B. G. (1990). "A temperature-induced switch from diffusive to convective ventilation in the honeybee". Journal of Experimental Biology 154 (1): 509–516.
  11. ↑ 11.0 11.1 Kovac, H.; Stabentheiner, A.; Hetz, S. K.; Petz, M.; Crailsheim, K. (2007). "Respiration of resting honeybees". Journal of Insect Physiology 53 (12): 1250–1261. doi:10.1016/j.jinsphys.2007.06.019. PMC 3227735. PMID 17707395.
  12. ↑ Southwick, E. E.; Heldmaier, G. (1987). "Temperature control in honey bee colonies". BioScience 37 (6): 395–399. doi:10.2307/1310562.