Behavioral ecology, or etoecology, is the study of the ecological Ecology is the scientific study of the distributions, abundance and relations of organisms and their interactions with the environment. Ecology includes the study of plant and animal populations, plant and animal communities and ecosystems. Ecosystems describe the web or network of relations among organisms at different scales of organization and evolutionary Evolution is the change in the inherited traits of a population of organisms through successive generations. After a population splits into smaller groups, these groups evolve independently and may eventually diversify into new species. Ultimately, life is descended from a common ancestory through a long series of these speciation events, basis for animal behavior Ethology is the scientific study of animal behavior, and a sub-topic of zoology, and the roles of behavior in enabling an animal to adapt to its environment (both intrinsic and extrinsic). Behavioral ecology emerged from ethology Ethology is the scientific study of animal behavior, and a sub-topic of zoology after Niko Tinbergen (a seminal figure in the study of animal behavior) outlined the four causes of behavior.

If an organism has a trait which provides them with a selective advantage (i.e. has an adaptive significance) in a new environment natural selection will likely favor it. Adaptive significance therefore refers to the beneficial qualities in terms of increased survival and reproduction a trait conveys.

For example, the behavior of flight Flight is the process by which an object moves either through the air, or movement beyond earth's atmosphere , by generating lift, propulsive thrust or aerostatically using buoyancy, or by simple ballistic movement has evolved numerous times in reptiles (Pterosaur Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period (220 to 65.5 million years ago). Pterosaurs are the earliest vertebrates known to have evolved powered flight. Their wings were formed by a membrane of skin, muscle, and other tissues stretching from the legs), birds, many insects and mammals (bats Bats are flying mammals in the order Chiroptera . The forelimbs of bats are webbed and developed as wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums and colugos, glide rather than fly, and only for short distances. Bats do not flap) due to its adaptive significance—for many species, flight has the potential to increase an animal's ability to escape from predators and move swiftly between habitat areas, among other things, thereby increasing the organism's chances of survival and reproduction. In all instances, the organism adapting to flight Flight is the process by which an object moves either through the air, or movement beyond earth's atmosphere , by generating lift, propulsive thrust or aerostatically using buoyancy, or by simple ballistic movement had to have "pre-adaptions" to these behavioral and anatomical changes. Feathers in birds initially evolving for thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. This process is one aspect of homeostasis: a dynamic state of stability between an animal's internal environment and its external environment . If the body is unable to maintain a normal then turned to flight due to the benefits conveyed (see Origin of avian flight); insect wings evolving from enlarged gill plates used to efficiently "sail" across the water, becoming larger until capable of flight are two good examples of this. At every stage slight improvements mean higher energy acquisition, lower energy expenditure or increased mating opportunities causing the genes that convey these traits to increase within the population. If these organisms did not have the required variation for natural selection to act upon either due to phylogenetic or genetic constraints, these behaviors would not be able to evolve.

However, it is not sufficient to apply these explanations where they seem convenient. Viewing traits and creating unsubstantiated theories or "Just So Stories" as to their adaptive nature have been deeply criticized. Stephen Jay Gould and Richard Lewontin (1979) described this as the "adaptationist programme". To be rigorous, hypotheses regarding adaptations must be theoretically or experimentally tested as with any scientific theory.

The hypothesis of the evolution of insect flight for example has been tested through wing manipulation experiments.[1] Empirical observations which adhere to the conditions prosed also provide evidence. For instance, one can suppose that when birds are not at risk of being eaten they might lose the ability to fly as the construction of functional wings are costly to produce and take away energy which could be used to increase offspring production or survival, a trend many island flightless birds such as the Kakapo The Kakapo , Strigops habroptila (Gray, 1845), also called owl parrot, is a species of flightless nocturnal parrot endemic to New Zealand. It has finely blotched yellow-green plumage, a distinct facial disc of sensory, vibrissa-like feathers, a large grey beak, short legs, large feet, and wings and a tail of relatively short length. A certain and the now extinct Dodo The dodo was a flightless bird endemic to the Indian Ocean island of Mauritius. Related to pigeons and doves, it stood about a meter (3 feet) tall, weighing about 20 kilograms (44 lb), living on fruit, and nesting on the ground demonstrate in the absence of natural predators prior to human colonization.

Contents

Proximate causation

Proximate causation is also divided into two factors which are ontogenetic Ontogeny (ontos present participle of 'to be', genesis 'creation') describes the origin and the development of an organism from the fertilized egg to its mature form. Ontogeny is studied in developmental biology, developmental psychology, developmental cognitive neuroscience, and developmental psychobiology. Ontogeny is that branch of life science and mechanistic. Ontogenetic factors are the entire sum of experience throughout the lifetime of an individual from embryo to death. Hence, factors included are learning the genetic factors giving rise to behavior in individuals. Mechanistic factors, as the name implies, are the processes of the body that give rise to behavior such as the effects of hormones A hormone is a chemical released by a cell in one part of the body, that sends out messages that affect cells in other parts of the organism. Only a small amount of hormone is required to alter cell metabolism. It is essentially a chemical messenger that transports a signal from one cell to another. All multicellular organisms produce hormones; on behavior and neuronal basis of behavior.

Optimization theory

Behavioral ecology, along with other areas of evolutionary biology Evolutionary biology is a sub-field of biology concerned with the origin of species from a common descent and descent of species, as well as their change, multiplication and diversity over time. Someone who studies evolutionary biology is known as an evolutionary biologist. To philosopher Kim Sterelny, "the development of evolutionary biology, has incorporated a number of techniques which have been borrowed from optimization theory. Optimization In mathematics and computer science, optimization, or mathematical programming, refers to choosing the best element from some set of available alternatives is a concept that stipulates strategies that offer the highest return to an animal given all the different factors and constraints facing the animal. One of the simplest ways to arrive at an optimal solution is to do a cost/benefit analysis. By considering the advantages of a behavior and the costs of a behavior, it can be seen that if the costs outweigh the benefits then a behavior will not evolve and vice versa. This is also where the concept of the trade-off becomes important. This is because it rarely pays an animal to invest maximally in any one behavior. For example, the amount of time an ectothermic Ectothermic, from the Greek εκτός "outside" and θερμός (thermós) "hot", (sometimes confusingly known as "cold-blooded") refers to organisms that control body temperature through external means. As a result, organisms are dependent on environmental heat sources and have relatively low metabolic rates. For animal such as a lizard spends foraging is constrained by its body temperature. The digestive efficiency of the lizard also increases with increases in body temperature. Lizards increase their body temperature by basking in the sun. However, the time spent basking decreases the amount of time available for foraging. Basking also increases the risk of being discovered by a predator. Therefore, the optimal basking time is the outcome of the time necessary to sufficiently warm itself to carry out its activities such as foraging Foraging theory is a branch of behavioral ecology that studies the foraging behavior of animals in response to the environment in which the animal lives. Foraging theory considers the foraging behavior of animals in reference to the payoff that an animal obtains from different foraging options. Foraging theory predicts that the foraging options. This example shows how foraging is constrained by the need to bask (intrinsic constraint) and predation In ecology, predation describes a biological interaction where a predator feeds on its prey (the organism that is attacked). Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption. The other main pressure (extrinsic constraint).

Differential reproductive success

Ultimately, behavior is subject to natural selection Natural selection is the process by which certain heritable traits—those that make it more likely for an organism to survive and successfully reproduce —become more common in a population over successive generations. It is a key mechanism of evolution just as with any other trait. Therefore animals that employ optimal behavioral strategies specific to their environment will generally leave greater numbers of offspring than their suboptimal conspecifics. Animals that leave a greater number of offspring than others of their own species are said to have greater fitness. However, environments change over time. What might be good behavior today might not be the best behavior in 10,000 years time or even 10 years time. The behavior of animals has and will continue to change in response to the environment. Behavioral ecology is one of the best ways to study these changes. As geneticist Theodosius Dobzhansky Theodosius Grygorovych Dobzhansky, also known as T. G. Dobzhansky, and sometimes Anglicized to Theodore Dobzhansky was a prominent geneticist and evolutionary biologist, and a central figure in the field of evolutionary biology for his work in shaping the unifying modern evolutionary synthesis. Dobzhansky was born in Ukraine (then part of Imperial famously wrote, "nothing in biology makes sense except in the light of evolution."

Evolutionarily stable strategies

Another driving force in the evolution of animal behavior is the concept of an evolutionarily stable strategy In game theory and behavioural ecology, an evolutionarily stable strategy is a strategy which, if adopted by a population of players, cannot be invaded by any alternative strategy that is initially rare. An ESS is an equilibrium refinement of the Nash equilibrium -- it is a Nash equilibrium which is "evolutionarily" stable meaning that (or ESS), a term derived from economic game theory Game theory is a branch of applied mathematics that is used in the social sciences, most notably in economics, as well as in biology , engineering, political science, international relations, computer science, and philosophy. Game theory attempts to mathematically capture behavior in strategic situations, or games, in which an individual's success which became prominent after John Maynard Smith John Maynard Smith, F.R.S. was a British theoretical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he then took a second degree in genetics under the well-known biologist J.B.S. Haldane. Maynard Smith was instrumental in the application of game theory to evolution and theorized on other(1982) [2] recognized the possible application of the concept of a Nash equilibrium In game theory, Nash equilibrium is a solution concept of a game involving two or more players, in which each player is assumed to know the equilibrium strategies of the other players, and no player has anything to gain by changing only his or her own strategy unilaterally. If each player has chosen a strategy and no player can benefit by changing to model the evolution of behavioral strategies.

In short, evolutionary game theory Evolutionary game theory is the application of game theory to interaction dependent strategy evolution in populations. EGT is useful in a biological context by defining a framework of strategies in which adaptive features can be modeled. It originated in 1973 with John Maynard Smith and George R. Price's formalization of evolutionarily stable asserts that only strategies In game theory, a player's strategy in a game is a complete plan of action for whatever situation might arise; this fully determines the player's behaviour. A player's strategy will determine the action the player will take at any stage of the game, for every possible history of play up to that stage that, when common in the population, cannot be "invaded" by any alternative (mutant) strategy will be an ESSs, and thus maintained in the population. In other words, at equilibrium every player should play the best strategic response to each other. When the game is two player and symmetric each player should play the strategy which is the best response to itself.

Therefore, the ESS is considered to be the evolutionary end point subsequent to the interactions. As the fitness conveyed by a strategy is influenced by what other individuals are doing (the relative frequency of each strategy in the population), behavior can be governed not only by optimality but the frequencies of strategies adopted by others and are therefore frequency dependent (frequency dependence Frequency dependent selection is the term given to an evolutionary process where the fitness of a phenotype is dependent on its frequency relative to other phenotypes in a given population. In positive frequency dependent selection, the fitness of a phenotype increases as it becomes more common. In negative frequency dependent selection, the).

Behavioral evolution is therefore influenced by both the physical environment and interactions between other individuals.

See also

References

  1. ^ Marden, J.H. & M.G. Kramer (1994) Surface-skimming stoneflies: a possible intermediate stage in insect flight evolution. Science, Vol. 266, 427-430
  2. ^ Maynard Smith, J. 1982. Evolution and the Theory of Games.

Further reading

External links

Modelling ecosystems Ecosystem models, or ecological models, are mathematical representations of ecosystems. Typically they simplify complex foodwebs down to their major components or trophic levels, and quantify these as either numbers of organisms, biomass or the inventory/concentration of some pertinent chemical element - trophic The trophic level of an organism is the position it occupies on the food chain. The word trophic derives from the Greek τροφή referring to food or feeding. A food chain represents a succession of organisms that eat another organism and are, in turn, eaten themselves. The number of steps an organism is from the start of the chain is a measure components
General Abiotic component In biology, abiotic components are non-living chemical and physical factors in the environment. Abiotic phenomena underlie all of biology. Abiotic factors, while generally downplayed, can have enormous impact on evolution · Behaviour · Biogeochemical cycle In ecology and Earth science, a biogeochemical cycle or nutrient cycle is a pathway by which a chemical element or molecule moves through both biotic and abiotic (lithosphere, atmosphere, and hydrosphere) compartments of Earth. In effect, the element is recycled, although in some cycles there may be places (called reservoirs) where the element is · Biomass Biomass, in ecology, is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms, plants or animals. The mass can be expressed as the · Biotic component Biotic factors are the factors in a biome/ecosystem/habitat that include all living things. This can later be broken down into producers, consumers , decomposers, and scavengers. Abiotic factors are the ones that are nonliving. Biotic components are the living things that shape an ecosystem. They are, any living component that affects another · Carrying capacity The carrying capacity of a biological species in an environment is the population size of the species that the environment can sustain indefinitely, given the food, habitat, water and other necessities available in the environment. For the human population, more complex variables such as sanitation and medical care are sometimes considered as part · Competition Competition is an interaction between organisms or species, in which the fitness of one is lowered by the presence of another. Limited supply of at least one resource used by both is required. Competition both within and between species is an important topic in ecology, especially community ecology. Competition is one of many interacting biotic · Ecosystem An ecosystem consists of all the organisms living in a particular area, as well as all the nonliving, physical components of the environment with which the organisms interact, such as air, soil, water, and sunlight. It is all the organisms in a given area, along with the nonliving factors with which they interact; a biological community and its · Ecosystem ecology Ecosystem ecology is the integrated study of biotic and abiotic components of ecosystems and their interactions within an ecosystem framework. This science examines how ecosystems work and relates this to their components such as chemicals, bedrock, soil, plants, and animals · Ecosystem model · Keystone species · List of feeding behaviours · Metabolic theory of ecology · Productivity
Producers Autotrophs · Chemosynthesis · Chemotrophs · Foundation species · Mixotrophs · Myco-heterotrophy · Mycotroph · Organotrophs · Photoheterotrophs · Photosynthesis · Photosynthetic efficiency · Phototrophs · Primary nutritional groups · Primary production
Consumers Apex predator · Bacterivore · Carnivores · Chemoorganotroph · Generalist and specialist species · Herbivores · Heterotroph · Heterotrophic nutrition · Mesopredator release hypothesis · Omnivores · Predation
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Species Biodiversity · Density-dependent inhibition · Ecological effects of biodiversity · Ecological extinction · Endemic species · Flagship species · Gradient analysis · Indicator species · Introduced species · Invasive species · Latitudinal gradients in species diversity · Minimum viable population · Occupancy-abundance relationship · Population viability analysis · Rapoport's rule · Relative abundance distribution · Relative species abundance · Species diversity · Species homogeneity · Species richness · Species distribution · Species-area curve · Umbrella species
Species interaction Antibiosis · Biological interaction · Commensalism · Ecological facilitation · Interspecific competition · Mutualism
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