1270 General Ecology | Endotherm

Endotherm
M K Labocha and J P Hayes, University of Nevada, Reno, Reno, NV, USA
ª 2008 Elsevier B.V. All rights reserved.

What Is an Endotherm? Benefits and Costs of Endothermy

Which Organisms Are Endotherms? Energetic Influence of Endotherms on Ecosystems
Source of Heat Evolution of Endothermy
Endothermy, Metabolic Rate, and Body Size Further Reading
Endothermic Response to Temperature

What Is an Endotherm? freezing point of pure water). These fishes are home-
otherms because their environmental temperature is
All organisms break complex molecules into simpler relatively constant, not because they are thermoregulat-
molecules, thereby obtaining energy to sustain life. This ing. On the other hand, some, but not most, mammals and
energy is used to transport ions, pump blood, move the birds are heterothermic. That means their body tempera-
body, and many other functions. Energy use ultimately ture varies over a wider range than is typical for
leads to the production of internal heat. Endotherms are homeotherms (but it does not necessarily track environ-
capable of producing sufficient internal heat to elevate mental temperature as in poikilotherms). This variation in
their body temperature (or part of their body) above body temperature may occur over short timescales, such
environmental temperature. In contrast, ectotherms can- as nightly torpor in bats and hummingbirds or the rather
not produce sufficient internal heat to elevate their variable body temperatures of some mammals, such as the
body temperature above environmental temperature. egg-laying echidna. Variation in body temperature of
Endotherms have higher energy use than similar-sized birds and mammals may also occur over seasonal time-
ectotherms; consequently, endotherms consume more of scales, for example, as is seen in bears and ground
the production by the ecosystem than do similar-sized squirrels. Animals that regulate their body temperature
ectotherms. The ability to be an endotherm depends on in the face of changing environmental temperature are
the rate of internal heat production, the size of the organ- called thermoregulators (Figure 1). Thermoregulation
ism, the degree of insulation, and environmental can be accomplished physiologically by altering rates of
circumstances. heat production or heat loss. Animals may also thermo-
Many endotherms, such as mammals and birds, elevate regulate behaviorally by selecting warmer or cooler
their entire body temperature above the environmental microenvironments. Animals whose body temperature
temperature. In other cases, endothermy is regional, such tracks the environmental temperature are called thermo-
that only some parts of the organism (e.g., the brain or conformers (Figure 1).
locomotor muscle of tuna or the flower of plant) are
heated above environmental temperature. In some organ-
isms, endothermy is not continuous but occurs only
40 °C
periodically or during particular activities. For example,
brooding pythons heat their body only while incubating
eggs, and insects elevate temperature of their active flight
Body temperature

Thermoregulator
muscles only prior to and during flight.
In the older literature, the term endotherm was some-
times used interchangeably with the term homeotherm,
but this usage is incorrect. Homeotherms are organisms
that maintain relatively constant body temperature. In
Thermoconformer
contrast, organisms whose body temperature varies
substantially (generally tracking the environmental tem-
perature) are called poikilotherms. Generally mammals
and birds are homeothermic while most other animals are 0 °C Environmental temperature 40 °C
poikilothermic, but this is not always the case. For exam- Figure 1 Change in body temperature of thermoregulators
ple, some Antarctic ice fish have body temperatures that and thermoconformers with environmental temperature
show extremely little variation (and are below the change.
 General Ecology | Endotherm 1271

Two other common terms in thermal physiology are temperature even if the animal is resting. Birds and mam-
cold blooded and warm blooded. These terms are no mals have much higher standard metabolic rates than
longer widely used by biologists because they convey ectotherms, and the differences in standard metabolism
little useful information beyond the temperature of an are associated with different molecular, cellular, and
animal (or its blood). For example, animals may be cold organ characteristics. Compared to ectotherms, birds
blooded at one time and warm blooded at another and mammals have higher mitochondrial volume, greater
depending on their thermal environment. Many reptiles membrane surface per tissue volume, and higher aerobic
have high body temperatures (i.e., are warm blooded enzyme activity. Their cellular membranes have greater
during the day) when the thermal environment is hot, ion fluxes, such that physiologists describe the cellular
but they may be cold blooded when the thermal environ- membranes of endotherms as being ‘leakier’ than those of
ment is cold (e.g., at night or during winter). Similarly ectotherms. Indeed, the greater leakiness of cellular mem-
heterothermic birds and mammals may be warm blooded branes has been hypothesized to be one of the major
during some parts of the diurnal or annual cycle and cold processes responsible for the higher standard metabolic
blooded at other times. rate of endotherms.
Other terms frequently used with relation to Most endothermic reptiles, fishes, and insects support
endothermy are standard metabolic rate (SMR), resting an elevated body temperature with heat produced by
metabolic rate (RMR), basal metabolic rate (BMR), and working muscle, and they do not maintain endothermy
field metabolic rate (FMR). SMR is metabolism measured at rest. So their molecular, cellular, and organ character-
at a particular environmental temperature while an ani- istics are more similar to ectotherms than they are to
mal is inactive and not digesting or absorbing food. RMR endotherms. Besides producing heat by muscle, endother-
differs from SMR in that an animal may be digesting or mic fishes developed vascular countercurrent heat
absorbing food. BMR is a special case of SMR, when exchangers to reduce heat loss and thereby promote
metabolic rate is measured within the thermoneutral endothermy. Some sea turtles also have countercurrent
zone (TNZ; see below). FMR is metabolic rate of animals heat exchangers (in their flippers). In fishes that elevate
in their natural environment. SMR, RMR, and FMR can
their brain or eye temperature, the method of heat pro-
be used to describe the metabolic rate of both endotherms
duction varies among species, and the method of heat
and ectotherms, but the term BMR applies only to
production is known not for all species. In lamnid sharks,
endotherms.
heat is transferred from muscle, whereas billfishes and
mackerel produce heat with specialized heater tissues
(evolved from ocular muscles). In plants, endothermy is
Which Organisms Are Endotherms? achieved by decoupling ATP generation from heat pro-
duction via use of an alternative electron-transport
The most familiar groups of endotherms are mammals mechanism (i.e., cyanide-insensitive respiration).
and birds. Only mammals and birds are endothermic at
rest. However, because some endothermic fish (e.g.,
tunas) must swim continuously to ventilate their gills,
these fish might also be considered to be endotherms Endothermy, Metabolic Rate, and Body
even at rest or as close to rest as their biology allows. Size
Besides birds and mammals, other endotherms include
some species of reptiles (e.g., large sea turtles, brooding The metabolic rate necessary to achieve endothermy is
pythons), ray-finned fishes (e.g., tunas and billfishes), influenced by size because heat loss to the environment is
lamnid sharks, insects (e.g., beetles, dragonflies, cicadas, roughly proportional to an animal’s surface area. Surface
moths, and bees), gymnosperm plants (i.e., cycads), and area depends on body mass (i.e., for similarly shaped
flowering plants (e.g., some species from the families organisms, surface area increases with body mass raised
Annonaceae, Aracacea, Araceae, Cyclanthaceae, and to the power 2/3). Because surface area increases more
Rafflesiaceae). slowly than body mass, larger animals generally can
achieve endothermy with lower metabolic intensities
(i.e., metabolic rates per unit mass) (Table 1). Indeed,
Source of Heat even with low metabolic intensities, animals with very
large body masses (e.g., sauropod dinosaurs) may have
Mammals and birds differ from other endotherms because been endothermic. A lower limit on size of vertebrate
they can maintain high body temperature at rest. A high endotherms also exists. The smallest vertebrate
basal metabolism is necessary for mammals and birds to endotherms are roughly 2 g in mass but the smallest
elevate their body temperature above environmental vertebrate ectotherms can be much smaller.
1272 General Ecology | Endotherm

Table 1 Basal metabolic rate (BMR) of some mammals over a large range of body mass

Body mass BMR Mass-specific BMR

Species Common name (g) (ml O2 h 1) (ml O2 g 1h 1)

Sorex minutusa Pygmy shrew 3.3 28.4 8.60

Peromyscus maniculatusb Deer mouse 14.9 44.4 3.00
Clethrionomys glareolusc Bank vole 20.9 52.5 2.50
Petrodromus tetradactylusd Four-toed elephant shrew 206.1 179.5 0.87
Erinaceus europaeuse European hedgehog 750.0 337.5 0.45
Fossa fossaf Fanaloka 2 260.0 904.0 0.40
Felis pardalisf Ocelot 10 416.0 3 229.0 0.31
Antilocapra americanag Pronghorn 37 800.0 9 318.0 0.25
Panthera oncaf Jaguar 68 900.0 12 402.0 0.18
Alces alcesh Moose 325 000.0 51 419.0 0.16
a
Sparti A (1992) Thermogenic capacity of shrews (Mammalia, Soricidae) and its relationship with basal rate of metabolism. Physiological Zoology
65: 77–96.
b
Hinds DS and Rice-Warner CN (1992) Maximum metabolism and aerobic capacity in heteromyid and other rodents. Physiological Zoology
65: 188–214.
c
Labocha MK, Sadowska ET, Baliga K, Semer AK, and Koteja P (2004) Individual variation and repeatability of basal metabolism in the bank vole,
Clethrionomys glareolus. Proceedings of the Royal Society B 271: 367–372.
d
Downs CT and Perrin MR (1995). The thermal biology of three southern African elephant-shrews. Journal of Thermal Biology 20: 445–450.
e
Shkolnik A and Schmidt-Nielsen K (1976) Temperature regulation in hedgehogs from temperate and desert environments. Physiological Zoology
49: 56–64.
f
McNab BK (1989) Basal rate of metabolism, body size and food habits in order Carnivora. In: Gittleman JL (ed.) Carnivore Behavior, Ecology, and
Evolution, pp. 335–354. Ithaca, NY: Cornell University Press.
g
Wesley DE, Knox KL, and Nagy JG (1973) Energy metabolism of pronghorn antelopes. Journal of Wildlife Management 37: 563–573.
h
Renecker LA and Hudson RJ (1986) Seasonal energy expenditures and thermoregulatory responses of moose. Canadian Journal of Zoology
64: 322–327.

Endothermic Response to Temperature

Endothermic homeotherms alter their metabolic rate

in response to environmental temperature (Figure 2).
Metabolic rate

The response to temperature depends in part on how Thermoneutral

zone (TNZ)
precisely or not endotherms maintain body tempera-
ture. Over a range of intermediate temperatures,
called the thermal neutral zone (TNZ), metabolic
rate remains constant. If body temperature and meta-
bolic rate are constant, but the thermal gradient
between the animal and the environment is changed,
then the insulation (or its reciprocal conductance) of
the animal must change. A typical endothermic home- Environmental temperature
otherm would have maximal conductance at the upper
Figure 2 Change in metabolic rate of endotherms in response
end of its TNZ and minimal conductance at the lower to environmental temperature. Other response patterns are
end of its TNZ. Above the upper end of the TNZ, possible, but this pattern is roughly typical for endotherms that
metabolic rate and body temperature increase, and the are strict thermoregulators.
animal may undertake such behaviors as panting to
increase evaporative heat loss. Below the lower end of
the TNZ, endotherms increase heat production so that or fat layers. As insulation increases, the TNZ widens and
they can offset increased heat loss in colder environ- the lower end of the TNZ occurs at lower temperatures.
ments; thereby, they balance heat production and heat Animals that are very well insulated (the arctic fox is a
loss to maintain a constant body temperature classic example) have thermal neutral zones that may
(Figure 2). extend to extremely cold temperatures (e.g., they may
Endotherms that live in cold environments (e.g., aquatic experience temperatures as low as –40  C or perhaps even
environments, where the thermal conductivity of the water lower without having to increase their basal metabolic rate).
is many times greater than that of air) tend to be well High temperatures can present problems for endotherms.
insulated, and aquatic endotherms tend to be large. When the thermal environment is hot, endotherms must
Insulation in vertebrates can take the form of fur, feathers, increase evaporation to avoid overheating. Hence, hot
 General Ecology | Endotherm 1273

environments can be challenging, particularly if ample to ectothermy is not as great as at cold environmental
water is not available. temperatures (Table 3).

Benefits and Costs of Endothermy Energetic Influence of Endotherms on

Ecosystems
The ability to control body temperature enables
endotherms to live in cold or thermally variable environ- Endothermy is more energetically costly than ectothermy.
ments yet remain active. On a geographical scale, Because endotherms use more energy than ectotherms, the
endotherms can live in regions too cold for most same amount of food can maintain a larger population of
ectotherms. On a temporal scale, endotherms can be similar-sized ectotherms than endotherms. Moreover, 90%
active year-round in regions with cold seasons, as well or more of the energy assimilated by endotherms is con-
as be active during cold parts of the day or night. verted to heat, so only a small percentage of the food
Endotherms not only have high SMR, RMR, and FMR, energy drawn from the ecosystem by endotherms is con-
they also have high aerobic capacities (maximal aerobic verted to biomass (i.e., to grow tissue or produce offspring).
metabolism). These high aerobic capacities make it pos- In other words, endotherms have lower production effi-
sible to sustain vigorous activity for extended time ciency than ectotherms.
periods. As a result, endotherms can maintain large terri- Because of the high energetic cost of endothermy,
tories and migrate for long distances. endothermic carnivores require higher prey densities
While there are diverse benefits of endothermy, than ectothermic carnivores. In systems with low primary
endothermy also has costs. In particular, endotherms productivity they will be absent or rare. Even folivorous
have higher metabolic rates than ectotherms (Figure 3), endotherms may be absent in habitats with extremely low
so they must locate and process more food to provide productivity.
the energy to sustain higher metabolism. Typically, an
endotherm requires much more energy and food
than a similar-sized ectotherm (Table 2). The cost of Evolution of Endothermy
endothermy relative to ectothermy changes with envi-
ronmental temperature. At warm environmental Because endothermy is energetically expensive and
temperatures, the energetic cost of endothermy relative evolved more than 100 million years ago (at least in

6 Mammalsa
log10 BMR = 0.614 + 0.69 log10 M
5
log10 metabolic rate (ml O2 h–1)

Birdsc
3 log10 BMR = 0.800 + 0.67 log10 M Reptilesb
log10 SMR = –0.260 + 0.76 log10 M

1
Fishb
log10 SMR = –0.657 + 0.88 log10 M
0

–1
Amphibiansb
log10 SMR = –0.469 + 0.88 log10 M
–2
–2 0 2 4 6 8
log10 body mass (g)

Figure 3 Relationship between metabolic rate and body mass in vertebrates. The data for amphibians, fish, and reptiles were
normalized to 38  C as described in White et al. Equation for birds was recalculated from Watts (used by McKechnie and Wolf) to
ml O2 h 1. aLovegrove BG (2000) The zoogeography of mammalian basal metabolic rate. American Naturalist 156: 201–219. bWhite CR,
Phillips NF, and Seymour RS (2006) The scaling and temperature dependence of vertebrate metabolism. Biology Letters 2: 125–127.
c
McKechnie AE and Wolf BO (2004) The allometry of avian basal metabolic rate: Good predictions need good data. Physiological and
Biochemical Zoology 77: 502–521.
1274 General Ecology | Endotherm

Table 2 Field metabolic rate (FMR) of endotherms and ectotherms of similar body mass

Endotherm (ENDO)/ Body mass FMR

Species Common name ectotherm (ECTO) (g) (kJ d 1)

Peromyscus maniculatusa Deer mouse ENDO 18.0 57.3

Erithacus rubeculab Robin ENDO 18.7 71.3
Mabuya striatac Striped skink ECTO 19.5 2.9
Thamnophis sirtalisd Common garter snake ECTO 22.0 5.2
Spermophilus parryie Arctic ground squirrel ENDO 630.0 817.0
Alectoris chukarf Chukar ENDO 440.0 306.2
Tiliqua scincoidesg Bluetongue lizard ECTO 574.0 45.7
Aptenodytes patagonicush King penguin ENDO 12 900.0 7410.0
Chelonia mydasi Green sea turtle ECTO 16 000.0 1867.0
a
Hayes JP (1989) Field and maximal metabolic rates of deer mice (Peromyscus maniculatus) at low and high altitudes. Physiological Zoology
62: 732–744.
b
Tatner P and Bryant DM (1993) Interspecific variation in daily energy expenditure during avian incubation. Journal of Zoology 231:
215–232.
c
Nagy KA and Knight MH (1989) Comparative field energetics of a Kalahari skink (Mabuya striata) and gecko (Pachydactylus bibroni). Copeia
1: 13–17.
d
Peterson CC, Walton BM, and Bennett AF (1998) Intrapopulation variation in ecological energetics of the garter snake Thamnophis sirtalis, with
analysis of the precision of doubly labeled water measurements. Physiological Zoology 71: 333–349.
e
Nagy KA (1994) Field bioenergetics of mammals: What determines field metabolic rates? Australian Journal of Zoology
42: 43–53.
f
Kam M, Degen AA, and Nagy KA (1987) Seasonal energy, water, and food consumption of Negev chukars and sand partridges. Ecology
68: 1029–1037.
g
Christian KA, Webb JK, and Schultz TJ (2003) Energetics of bluetongue lizard (Tiligua scincoides) in a seasonal tropical environment. Oecologia
136: 515–523.
h
Kooyman GL, Cherel Y, Le Maho Y, et al. (1992) Diving behavior and energetics during foraging cycles in king penguins. Ecological Monographs
62: 143–163.
i
Southwood AL, Reina RD, Jones VS, Speakman JR, and Jones DR (2006) Sesonal metabolism of juvenile green turtles (Chaledonia mydas) at
Heron Island, Australia. Canadian Journal of Zoology 84: 125–135.

birds and mammals), the selective forces leading to stability. Later, the aerobic capacity model posited that
the evolution of endothermy are unclear. The earliest endothermy evolved as by-product of selection for high
hypotheses to explain the evolution of endothermy pos- aerobic capacity (i.e., maximal oxygen consumption
tulated that selection for higher resting metabolism led capacity during exercise). Based on data for extant organ-
to an expanded thermal niche or increased thermal isms, aerobic capacity was argued to be inescapably

Table 3 A comparison of metabolic rates of endotherms and ectotherms of similar body mass

Endotherm (ENDO)/ Temperature Body Metabolic rate

Species Common name ectotherm (ECTO) ( C) mass (g) (ml O2 h 1)

Dipsosaurus dorsalisa Desert iguana ECTO 25 35 1.68

Dipsosaurus dorsalisa Desert iguana ECTO 30 35 2.45
Dipsosaurus dorsalisa Desert iguana ECTO 35 35 5.25
Dipsosaurus dorsalisa Desert iguana ECTO 40 35 6.30
Notoryctes caurinusb North-western marsupial ENDO 30.8 34 21.4
mole
Phyllostomus discolorc Pale spear-nosed bat ENDO 34.6 33.5 11.1
Gerbillus allenbyid Allenby’s gerbil ENDO 36.3 35.3 38.8

Acanthodactylus Fringe-toed lizard ECTO 20 9.0 1.17

erythruruse
Acanthodactylus Fringe-toed lizard ECTO 25 9.0 1.62
erythruruse
Acanthodactylus Fringe-toed lizard ECTO 30 9.0 2.25
erythruruse
Acanthodactylus Fringe-toed lizard ECTO 35 9.0 3.15
erythruruse
Crocidura crosseif Crosse’s shrew ENDO 34.3 10.2 22.4

(Continued )
 General Ecology | Endotherm 1275

Table 3 (Continued)

Endotherm (ENDO)/ Temperature Body Metabolic rate

Species Common name ectotherm (ECTO) ( C) mass (g) (ml O2 h 1)

Perognathus Little pocket mouse ENDO 34.7 8.9 9.5

longimembrisg
Pteronotus davyih Davy’s naked-backed bat ENDO 38.8 9.4 24.4
a
Bennett AF and Dawson WR (1972) Aerobic and anaerobic metabolism during activity in the lizard Dipsosaurus dorsalis. Journal of Comparative
Physiology 81: 289–299.
b
Withers PC, Thompson GG, and Seymour RS (2000) Metabolic physiology of the north-western marsupial mole, Notoryctes caurinus (Marsupialia:
Notoryctidae). Australian Journal of Zoology 48: 241–258.
c
McNab BK (1969) The economics of temperature regulation in neotropical bats. Comparative Biochemistry and Physiology 31: 227–268.
d
Haim A (1984) Adaptive variations in heat production within gerbils (genus Gerbillus) from different habitats. Oecologia 61: 49–52.
e
Pough FH and Busack SD (1978) Metabolism and activity of the Spanish fringe-toed lizard (Lacertidae: Acanthodactylus erythurus). Journal of
Thermal Biology 3: 203–205.
f
Sparti A (1990) Comparative temperature regulaton of African and European shrews. Comparative Biochemistry and Physiology A 97: 391–397.
g
Chew RM, Lindberg RG, and Hayden P (1967) Temperature regulation in the little pocket mouse, Perognathus longimembris. Comparative
Biochemistry and Physiology 21: 487–505.
h
Bonaccorso FJ, Arends A, Genoud M, Cantoni D, and Morton T (1992) Thermal ecology of moustached and ghost-faced bats (Mormoopidae) in
Venezuela. Journal of Mammalogy 73: 365–378.
For endotherms, basal metabolic rate is shown. For ectotherms, standard metabolic rate over a range of temperatures is shown.

correlated with resting metabolism. The newest models Downs CT and Perrin MR (1995) The thermal biology of three southern
African elephant-shrews. Journal of Thermal Biology 20: 445–450.
suggest that endothermy evolved as consequence of selec- Haim A (1984) Adaptive variations in heat production within gerbils
tion for intense parental care. Which of these models for (genus Gerbillus) from different habitats. Oecologia 61: 49–52.
the evolution of endothermy is best is unresolved. Hayes JP (1989) Field and maximal metabolic rates of deer mice
(Peromyscus maniculatus) at low and high altitudes. Physiological
Zoology 62: 732–744.
Hayes JP and Garland T, Jr. (1995) The evolution of endothermy:
Testing the aerobic capacity model. Evolution 49: 836–847.
Acknowledgment Heinrih B (1993) The Hot-Blooded Insects: Strategies and
Mechanisms of Thermoregulation. Cambridge, MA: Harvard
University Press.
This contribution was supported in part by US National Hinds DS and Rice-Warner CN (1992) Maximum metabolism and
Science Foundation award IOB 0344994 to J. Hayes. aerobic capacity in heteromyid and other rodents. Physiological
Zoology 65: 188–214.
Hulbert AJ and Else PL (2000) Mechanisms underlying the cost of living
See also: Homeotherms; Poikilotherms. in animals. Annual Review of Physiology 62: 207–235.
Kam M, Degen AA, and Nagy KA (1987) Seasonal energy, water, and
food consumption of Negev chukars and sand partridges. Ecology
68: 1029–1037.
Kemp TS (2006) The origin of mammalian endothermy: A paradigm for
Further Reading the evolution of complex biological structure. Zoological Journal of
the Linnean Society 147: 473–488.
Bennett AF (1991) The evolution of activity capacity. Journal of Kooyman GL, Cherel Y, Le Maho Y, et al. (1992) Diving behavior and
Experimental Biology 160: 1–23. energetics during foraging cycles in king penguins. Ecological
Bennett AF and Dawson WR (1972) Aerobic and anaerobic metabolism Monographs 62: 143–163.
during activity in the lizard Dipsosaurus dorsalis. Journal of Koteja P (2000) Energy assimilation, parental care and the evolution of
Comparative Physiology 81: 289–299. endothermy. Proceedings of the Royal Society – Biological Sciences
Bennett AF and Ruben JA (1979) Endothermy and activity in (Series B) 267: 479–484.
vertebrates. Science 206: 649–654. Labocha MK, Sadowska ET, Baliga K, Semer AK, and Koteja P (2004)
Block BA, Finnerty JR, Stewart AFR, and Kidd J (1993) Evolution of Individual variation and repeatability of basal metabolism in the bank
endothermy in fish: Mapping physiological traits on a molecular vole, Clethrionomys glareolus. Proceedings of the Royal Society B
phylogeny. Science 260: 210–214. 271: 367–372.
Bonaccorso FJ, Arends A, Genoud M, Cantoni D, and Morton T (1992) Lovegrove BG (2000) The zoogeography of mammalian basal metabolic
Thermal ecology of moustached and ghost-faced bats rate. American Naturalist 156: 201–219.
(Mormoopidae) in Venezuela. Journal of Mammalogy 73: 365–378. McKechnie AE and Wolf BO (2004) The allometry of avian basal
Chew RM, Lindberg RG, and Hayden P (1967) Temperature regulation metabolic rate: Good predictions need good data. Physiological and
in the little pocket mouse, Perognathus longimembris. Comparative Biochemical Zoology 77: 502–521.
Biochemistry and Physiology 21: 487–505. McNab BK (1969) The economics of temperature regulation in
Christian KA, Webb JK, and Schultz TJ (2003) Energetics of bluetongue neotropical bats. Comparative Biochemistry and Physiology
lizard (Tiligua scincoides) in a seasonal tropical environment. 31: 227–268.
Oecologia 136: 515–523. McNab BK (1989) Basal rate of metabolism, body size and food habits
Dickson KA and Graham JB (2004) Evolution and consequences of in order Carnivora. In: Gittleman JL (ed.) Carnivore Behavior,
endothermy in fishes. Physiological and Biochemical Zoology Ecology, and Evolution, pp. 335–354. Ithaca, NY: Cornell University
77: 998–1018. Press.
1276 Global Ecology | Energy Balance

McNab BK (2002) The Physiological Ecology of Vertebrates: A View Shkolnik A and Schmidt-Nielsen K (1976) Temperature regulation in
from Energetics, chs. 4 and 5. Ithaca, NY: Cornell University hedgehogs from temperate and desert environments. Physiological
Press. Zoology 49: 56–64.
Nagy KA (1994) Field bioenergetics of mammals: What determines field Southwood AL, Reina RD, Jones VS, Speakman JR, and Jones DR
metabolic rates? Australian Journal of Zoology 42: 43–53. (2006) Sesonal metabolism of juvenile green turtles (Chaledonia
Nagy KA and Knight MH (1989) Comparative field energetics of a mydas) at Heron Island, Australia. Canadian Journal of Zoology
Kalahari skink (Mabuya striata) and gecko (Pachydactylus bibroni). 84: 125–135.
Copeia 1: 13–17. Sparti A (1990) Comparative temperature regulaton of African and
Peterson CC, Walton BM, and Bennett AF (1998) Intrapopulation European shrews. Comparative Biochemistry and Physiology A
variation in ecological energetics of the garter snake Thamnophis 97: 391–397.
sirtalis, with analysis of the precision of doubly labeled water Sparti A (1992) Thermogenic capacity of shrews (Mammalia, Soricidae)
measurements. Physiological Zoology 71: 333–349. and its relationship with basal rate of metabolism. Physiological
Pough FH and Busack SD (1978) Metabolism and activity of the Spanish Zoology 65: 77–96.
fringe-toed lizard (Lacertidae: Acanthodactylus erythurus). Journal of Tatner P and Bryant DM (1993) Interspecific variation in daily energy
Thermal Biology 3: 203–205. expenditure during avian incubation. Journal of Zoology
Renecker LA and Hudson RJ (1986) Seasonal energy expenditures and 231: 215–232.
thermoregulatory responses of moose. Canadian Journal of Zoology Wesley DE, Knox KL, and Nagy JG (1973) Energy metabolism of
64: 322–327. pronghorn antelopes. Journal of Wildlife Management
Ruben J (1995) The evolution of endothermy in mammals and birds: 37: 563–573.
From physiology to fossils. Annual Review of Physiology White CR, Phillips NF, and Seymour RS (2006) The scaling and
57: 69–95. temperature dependence of vertebrate metabolism. Biology Letters
Schweitzer MH and Marshall CL (2001) A molecular model for the 2: 125–127.
evolution of endothermy in the theropod-bird lineage. Journal of Withers PC, Thompson GG, and Seymour RS (2000) Metabolic
Experimental Zoology 291: 317–338. physiology of the north-western marsupial mole, Notoryctes caurinus
Seymour RS and Schultze-Motel P (1997) Heat-producing flowers. (Marsupialia: Notoryctidae). Australian Journal of Zoology
Endeavour 21: 125–129. 48: 241–258.

Energy Balance
A Kleidon, Max-Planck-Institut für Biogeochemie, Jena, Germany
ª 2008 Elsevier B.V. All rights reserved.

Introduction Dynamics
Global Energy Balance Global Energy Balance and Climate
Global Entropy Budget Ecosystems and the Global Energy Balance
Radiative Exchange Further Reading

Introduction balance, how it is reflected in the seasonal and geographic

distribution of mean climatic properties, and how it inter-
All ecosystems are affected by and interact with their acts with life through ecosystem functioning.
environment. At the global scale, the Earth’s environment
is characterized by the global energy balance, the balance
of all heating and cooling terms that shape the climatic Global Energy Balance
variations in space and time, especially with respect to
At the planetary scale, the energy balance is driven by the
surface temperature, precipitation, and light. From an
absorption of sunlight and the emission of radiation to
energy balance viewpoint, the interrelationships between
space. Planetary properties and the global energy balance
ecosystems and their environment are threefold: (1) eco-
give a first impression of the relevant processes that shape
systems utilize energy sources from their environment,
the environmental conditions at the surface and how
and thereby are a part – though small – of the energy
habitable these are to life.
balance; (2) ecosystem processes are affected by environ-
mental conditions that are directly or indirectly connected
to the energy balance (e.g., precipitation affects the levels Planetary Energy Balance
of water limitation of terrestrial productivity); and (3) the The planetary energy balance is driven by the absorption of
form and functioning of ecosystems affect energy balance about 240 W m 2 of solar radiation, which is then re-emitted
terms. This article reviews the basics of the global energy into space as long-wave radiation. The planetary energy