Temperature-size rule

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The temperature-size rule denotes the plastic response (i.e. phenotypic plasticity) of organismal body size to environmental temperature variation. [1] [2] Organisms exhibiting a plastic response are capable of allowing their body size to fluctuate with environmental temperature. First coined by David Atkinson in 1996, [3] it is considered to be a unique case of Bergmann's rule [1] that has been observed in plants, animals, birds, and a wide variety of ectotherms. [2] [4] [5] [6] [7] Although exceptions to the temperature-size rule exist, recognition of this widespread "rule" has amassed efforts to understand the physiological mechanisms (via possible tradeoffs) underlying growth and body size variation in differing environmental temperatures. [2] [8]

Contents

History

Comparative photo of a northern red fox and southern desert red fox depicting size differences with increasing latitudes (i.e. Bergmann's rule). Northern red fox & southern desert red fox.jpg
Comparative photo of a northern red fox and southern desert red fox depicting size differences with increasing latitudes (i.e. Bergmann's rule).

Relation to Bergmann's rule

In 1847, Carl Bergmann published his observations that endothermic body size (i.e. mammals) increased with increasing latitude, commonly known as Bergmann's rule. [9] His rule postulated that selection favored within species individuals with larger body sizes in cooler temperatures because the total heat loss would be diminished through lower surface area to volume ratios. [8] However, ectothermic individuals thermoregulate and allow their internal body temperature to fluctuate with environmental temperature whereas endotherms maintain a constant internal body temperature. This creates an inaccurate description of observed body size variation in ectotherms since they routinely allow evaporative heat loss and do not maintain constant internal temperatures. [8] [10] Despite this, ectotherms have largely been observed to still exhibit larger body sizes in colder environments.

Formulation of the rule

Ray (1960) originally examined body sizes in several species of ectotherms and discovered that around 80% of them exhibited larger body sizes in lower temperatures. [11] A few decades later, Atkinson (1994) performed a similar review of temperature effects on body size in ectotherms. His study, which included 92 species of ectotherms ranging from animals and plants to protists and bacteria, concluded that a reduction in temperature resulted in an increase in organism size in 83.5% of cases. [11] [12] [13] Atkinson's findings provided support for Ray's published works that ectotherms have an observable trend in body size when temperature is the primary environmental variable. The results of his study prompted him to name the increase in ectothermic body size in colder environments as the temperature-size rule.

Tradeoffs as possible underlying mechanisms

Life history model

Life history models highlighting optimal growth patterns suggest that individuals assess the environment for potential resources and other proximate factors and mature at a body size that yields the greatest reproductive success, or highest percentage of offspring surviving to reach reproductive maturity. [14]

Size at maturity

Environmental temperature is one of the most important proximate factors affecting ectotherm body size because of their need to thermoregulate. Individuals that have been observed to follow the temperature-size rule have slower growth rates in colder environments, yet they enter a period of prolonged growth that yields larger adult body sizes. [3] [15] [16] One proposed explanation for this involves a trade-off in life history traits. Ectotherms experience longer daily and seasonal activity times in warmer climates versus cooler climates, however, the increase in daily activity time is accompanied by higher infant and adult mortality rates due to predation. [16] [17] Under these environmental conditions, some individuals occupying these warmer climate environments will mature at smaller body sizes and undergo a shift in energy allocation of all acquired energy resources to reproduction. [18] [19] [20] In doing so, these individuals sacrifice growth to larger adult body sizes to ensure reproductive success, even if the trade-off results in smaller offspring that have increased mortality rates. [20]

Reproduction

Ectotherms occupying colder environments, such as mountain ranges or other areas of higher elevation, have been observed to invest in reproduction at larger adult body sizes due to a prolonged growth period. These populations of ectotherms are characterized as having smaller clutches of larger eggs, favoring a greater reproductive investment per egg and enhances offspring survival rates. [21] Individuals occupying warmer environments experience a trade-off between body size and overall reproductive success that many individuals occupying colder environments do not, hence, prolonging growth to yield greater reproductive success in colder environments could potentially be an underlying mechanism for why a large percentage of ectotherms exhibit greater body sizes in colder environments. However, a sufficient explanation for this observable pattern has yet to be produced. [14]

Investigation

Common lizard (Lacerta vivipara). Common Lizard. Lacerta vivipara (27742089869).jpg
Common lizard (Lacerta vivipara).

Supporting evidence

Eastern fence lizard (Sceloporus undulatus). Sceloporus undulatus1.JPG
Eastern fence lizard (Sceloporus undulatus).

Exceptions

Notes

The supporting evidence and the exceptions to the temperature-size rule listed above are only a few of the potential supporting/opposing evidence available for the temperature-size rule. Each was provided to support the claim that patterns of body size observed in variable environments are not 100% predictable and more research is required to identify and understand all of the mechanisms responsible.

Related Research Articles

<span class="mw-page-title-main">Centrarchidae</span> Family of fishes

Centrarchidae, better known as sunfishes, is a family of freshwater ray-finned fish belonging to the order Perciformes, native only to North America. There are eight universally included genera within the centrarchid family: Lepomis, Micropterus, Pomoxis (crappies), Enneacanthus, Centrarchus, Archoplites, Ambloplites, and Acantharchus. A genetic study in 2012 suggests that the highly distinct pygmy sunfishes of the genus Elassoma are also centrarchids.

<span class="mw-page-title-main">Endotherm</span> Organism that maintains body temperature largely by heat from internal bodily functions

An endotherm is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released 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.

<span class="mw-page-title-main">Thermoregulation</span> Ability of an organism to keep its body temperature within certain boundaries

Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature as its own body temperature, thus avoiding the need for internal thermoregulation. The internal thermoregulation process is one aspect of homeostasis: a state of dynamic stability in an organism's internal conditions, maintained far from thermal equilibrium with its environment. If the body is unable to maintain a normal temperature and it increases significantly above normal, a condition known as hyperthermia occurs. Humans may also experience lethal hyperthermia when the wet bulb temperature is sustained above 35 °C (95 °F) for six hours. Work in 2022 established by experiment that a wet-bulb temperature exceeding 30.55°C caused uncompensable heat stress in young, healthy adult humans. The opposite condition, when body temperature decreases below normal levels, is known as hypothermia. It results when the homeostatic control mechanisms of heat within the body malfunction, causing the body to lose heat faster than producing it. Normal body temperature is around 37°C(98.6°F), and hypothermia sets in when the core body temperature gets lower than 35 °C (95 °F). Usually caused by prolonged exposure to cold temperatures, hypothermia is usually treated by methods that attempt to raise the body temperature back to a normal range. It was not until the introduction of thermometers that any exact data on the temperature of animals could be obtained. It was then found that local differences were present, since heat production and heat loss vary considerably in different parts of the body, although the circulation of the blood tends to bring about a mean temperature of the internal parts. Hence it is important to identify the parts of the body that most closely reflect the temperature of the internal organs. Also, for such results to be comparable, the measurements must be conducted under comparable conditions. The rectum has traditionally been considered to reflect most accurately the temperature of internal parts, or in some cases of sex or species, the vagina, uterus or bladder.

<span class="mw-page-title-main">Ectotherm</span> Organism where internal heating sources are small or negligible

An ectotherm, more commonly referred to as a "cold-bloodedanimal", is an animal in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature. Such organisms rely on environmental heat sources, which permit them to operate at very economical metabolic rates.

<span class="mw-page-title-main">Bergmann's rule</span> Biological rule stating that larger size organisms are found in colder environments

Bergmann's rule is an ecogeographical rule that states that within a broadly distributed taxonomic clade, populations and species of larger size are found in colder environments, while populations and species of smaller size are found in warmer regions. Bergmann's rule only describes the overall size of the animals, but does not include body parts like Allen's rule does.

<span class="mw-page-title-main">Viviparous lizard</span> Species of lizard

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<span class="mw-page-title-main">Allen's rule</span> Relation of habitat temperature and limb length

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<span class="mw-page-title-main">Darwin's frog</span> Species of amphibian

Darwin’s frog, also called the Southern Darwin's frog, is a species of Chilean/Argentinian frog of the family Rhinodermatidae. It was discovered by Charles Darwin during his voyage on HMS Beagle. on a trip to Chile. In 1841, French zoologist André Marie Constant Duméril and his assistant Gabriel Bibron described and named Darwin's frog. The diet of R. darwinii consists mostly of herbivore invertebrates. R. darwinii is currently classified as an endangered species by the International Union for Conservation of Nature.

<span class="mw-page-title-main">Common side-blotched lizard</span> Species of lizard

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<i>Urosaurus ornatus</i> Species of lizard

Urosaurus ornatus, commonly known as the ornate tree lizard, is a species of lizard in the family Phrynosomatidae. The species is native to the southwestern United States and northwestern Mexico. The species, which was formerly called simply the "tree lizard", has been used to study physiological changes during the fight-or-flight response as related to stress and aggressive competition. Its life history and costs of reproduction have been documented in field populations in New Mexico and Arizona. This species has been fairly well studied because of its interesting variation in throat color in males that can correlate with different reproductive strategies,

<span class="mw-page-title-main">Sagebrush lizard</span> Species of lizard

The sagebrush lizard or sagebrush swift is a common species of phrynosomatid lizard found at mid to high altitudes in the western United States of America. It belongs to the genus Sceloporus in the Phrynosomatidae family of reptiles. Named after the sagebrush plants near which it is commonly found, the sagebrush lizard has keeled and spiny scales running along its dorsal surface.

Climatic adaptation refers to adaptations of an organism that are triggered due to the patterns of variation of abiotic factors that determine a specific climate. Annual means, seasonal variation and daily patterns of abiotic factors are properties of a climate where organisms can be adapted to. Changes in behavior, physical structure, internal mechanisms and metabolism are forms of adaptation that is caused by climate properties. Organisms of the same species that occur in different climates can be compared to determine which adaptations are due to climate and which are influenced majorly by other factors. Climatic adaptations limits to adaptations that have been established, characterizing species that live within the specific climate. It is different from climate change adaptations which refers to the ability to adapt to gradual changes of a climate. Once a climate has changed, the climate change adaptation that led to the survival of the specific organisms as a species can be seen as a climatic adaptation. Climatic adaptation is constrained by the genetic variability of the species in question.

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The eastern three-lined skink, also known commonly as the bold-striped cool-skink, is a species of skink, a lizard in the family Scincidae. The species is endemic to Australia. A. duperreyi has been extensively studied in the context of understanding the evolution of learning, viviparity in lizards, and temperature- and genetic-sex determination. A. duperreyi is classified as a species of "Least Concern" by the IUCN.

<span class="mw-page-title-main">Deep-sea gigantism</span> Tendency for deep-sea species to be larger than their shallower-water relatives

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