behavioral ecology Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for animal behavior due to ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined four questions to address when ...

, adaptive behavior is any

behavior Behavior (American English) or behaviour (British English) is the range of actions and mannerisms made by individuals, organisms, systems or artificial entities in some environment. These systems can include other systems or organisms as wel ...

that contributes directly or indirectly to an individual's

reproductive success Reproductive success is an individual's production of offspring per breeding event or lifetime. This is not limited by the number of offspring produced by one individual, but also the reproductive success of these offspring themselves. Reproduct ...

, and is thus subject to the forces of

natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charle ...

.Ecology and Behavior ''from'' Biology: The Unity and Diversity of Life 10th edition. Starr and Taggart 2004. Thompson publishers Examples include favoring

kin __NOTOC__ Kin usually refers to kinship and family. Kin or KIN may also refer to: Culture and religion *Otherkin, people who identify as not entirely human *Kinism, a white supremacist religious movement * Kinh, the majority ethnic group of V ...

altruistic behavior Altruism is the principle and moral practice of concern for the welfare and/or happiness of other human beings or animals, resulting in a quality of life both material and spiritual. It is a traditional virtue in many cultures and a core asp ...

sexual selection Sexual selection is a mode of natural selection in which members of one biological sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of t ...

of the most fit mate, and defending a

territory A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...

harem Harem (Persian: حرمسرا ''haramsarā'', ar, حَرِيمٌ ''ḥarīm'', "a sacred inviolable place; harem; female members of the family") refers to domestic spaces that are reserved for the women of the house in a Muslim family. A hare ...

from rivals. Conversely, non-adaptive behavior is any behavior that is counterproductive to an individual's survival or reproductive success. Examples might include altruistic behaviors which do not favor kin,

adoption Adoption is a process whereby a person assumes the parenting of another, usually a child, from that person's biological or legal parent or parents. Legal adoptions permanently transfer all rights and responsibilities, along with filiation, from ...

of unrelated young, and being a subordinate in a

dominance hierarchy In biology, a dominance hierarchy (formerly and colloquially called a pecking order) is a type of social hierarchy that arises when members of animal social animal , social groups interact, creating a ranking system. A dominant higher-ranking i ...

. Adaptations are commonly defined as evolved solutions to recurrent environmental problems of survival and reproduction.Buss, D.M. and Greiling, H. 1999. Adaptive individual differences. Journal of Personality, 67: 209-243. Individual differences commonly arise through both

heritable Heredity, also called inheritance or biological inheritance, is the passing on of traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cells or organisms acquire the genetic informa ...

and non-heritable adaptive behavior. Both have been proven to be influential in the

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

of species' adaptive behaviors, although non-heritable adaptation remains a controversial subject.

Non-heritable

Population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...

s change through the process of

. Each individual in a population has a unique role in their particular environment. This role, commonly known as an

ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...

, is simply how an organism lives in an environment in relation to others.Staddon, J. E. R. (1983). ''Adaptive Behavior and Learning''. Cambridge University Press. Over successive generations, the organism must adapt to their surrounding conditions in order to develop their niche. An organism's niche will evolve as changes in the external environment occur. The most successful species in nature are those that are able to use adaptive behaviors to build on prior knowledge, thereby increasing their overall knowledge bank. In turn, this will improve their overall survival and reproductive success.

Learning

Many species have the ability to adapt through

learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, value (personal and cultural), values, attitudes, and preferences. The ability to learn is possessed by humans, animals, and some machine learning, machines ...

Organisms In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fungi; ...

will often learn through various psychological and cognitive processes, such as

operant Operant conditioning, also called instrumental conditioning, is a learning process where behaviors are modified through the association of stimuli with reinforcement or punishment. In it, operants—behaviors that affect one's environment—are c ...

and

classical conditioning Classical conditioning (also known as Pavlovian or respondent conditioning) is a behavioral procedure in which a biologically potent stimulus (e.g. food) is paired with a previously neutral stimulus (e.g. a triangle). It also refers to the learni ...

and discrimination memory. This learning process allows organisms to modify their behavior to survive in unpredictable environments. Organisms begin as naive individuals and learning allows them to obtain the knowledge they need to adapt and survive. It is important to remember that a learned adaptive behavior must have a

psychological Psychology is the scientific study of mind and behavior. Psychology includes the study of conscious and unconscious phenomena, including feelings and thoughts. It is an academic discipline of immense scope, crossing the boundaries between t ...

, as well as a

biological Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary in ...

, component; without the integration of these two disciplines, behavioral adaptation cannot occur.

Kin selection

Kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like Altruism in animals, altruistic behavio ...

(commonly referred to as

altruism Altruism is the principle and moral practice of concern for the welfare and/or happiness of other human beings or animals, resulting in a quality of life both material and spiritual. It is a traditional virtue in many cultures and a core as ...

) is an example of an adaptive behavior that directly influences the genetic composition of a population. It involves evolutionary strategies that favor the persistence of an organism's relatives, often at the cost of the organism's own survival and

reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual or ...

.Eberhard, M.J.W. 1975. The evolution of social behavior by kin selection. The Quarterly Review of Biology, 50: 1-33. This will result in population gene

frequency Frequency is the number of occurrences of a repeating event per unit of time. It is also occasionally referred to as ''temporal frequency'' for clarity, and is distinct from ''angular frequency''. Frequency is measured in hertz (Hz) which is eq ...

variation over successive generations, based on the interactions between related individuals. The probability of altruism increases when the

cost In production, research, retail, and accounting, a cost is the value of money that has been used up to produce something or deliver a service, and hence is not available for use anymore. In business, the cost may be one of acquisition, in which ...

is low for the donor, or when there is a high level of gain for the beneficiary. In addition, individuals often display altruistic behaviors when the relative is genetically similar to them. This means offspring or siblings are more likely to benefit from altruism than someone more distantly related, such as a cousin, aunt, or uncle. Kin selection has played a large role in the evolution of social and adaptive behaviors in chimpanzees. Closely related chimpanzees will form a kin group that cooperates to protect a territory, thereby increasing their access to females and resources.Morin, P.A., Moore, J.J., Chakraborty, R., Jin, L., Goodall, J. and Woodruff, D.S. Kin selection, social structure, gene flow, and the evolution of chimpanzees. Science, 265: 1193-1201. By working together with close relatives, they can ensure that their genes will persist in the next generation, even if circumstances make them unable to reproduce themselves. This behavioral adaptation coincides with the chimpanzee's ability to distinguish kin from non-kin (referred to as visual kin selection) allowing chimps to formulate large, complex societies, where they use altruistic methods to ensure their genes persist in future generations. A wide variety of species, including lions, honeybees, and other insects have displayed kin selection as an adaptive behavior.

Territorial defense

As mentioned above, chimpanzees work together with closely related kin to protect their

. Defending territory from rivals (known as territoriality) is a learnt adaptive behavior performed by several ecological species. The advantage of being territorial varies depending on the species of interest, but the underlying principle is always to increase overall fitness. Many species will display territoriality in order to acquire food, compete for mates, or have the safest lair. Bird song is an example of learned territorial defense. Studies show that birds with high-quality songs will use them as a stimulus to deter predators from their territorial range. Higher quality songs have been proven to act as the best defense mechanism in a variety of bird species, such as the

red-winged blackbird The red-winged blackbird (''Agelaius phoeniceus'') is a passerine bird of the family Icteridae found in most of North America and much of Central America. It breeds from Alaska and Newfoundland south to Florida, the Gulf of Mexico, Mexico, and ...

(''Agelaius phoeniceus''). Therefore, correct learning of the birdsong early in life is necessary for territory defense in birds.

European beavers The Eurasian beaver (''Castor fiber'') or European beaver is a beaver species that was once widespread in Eurasia, but was hunted to near-extinction for both its fur and castoreum. At the turn of the 20th century, only about 1,200 beavers surviv ...

(''Castor fiber'') are another species that use territory defense. They are very protective of their territory because they spend a great deal of time and energy familiarizing themselves with the location and building a habitat. Beavers have developed the technique of scent-marking as a way to intimidate intruders. This scent acts as a "psychological fence", thereby decreasing the possibility of injury or death by predation.

Controversy

There is debate on whether or not there exists a biological component associated with the learning process in ecological adaptive behavior. Many researchers suggest that the biological and psychological disciplines are integrated, while others believe that the non-heritable component is strictly psychological. They argue that non-heritable traits cannot evolve over successive generations.

Heritable

Organisms can also express

adaptive behaviors. These behaviors are encoded in their

genes In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...

and inherited from their parents. This gives the organisms the ability to respond to situations with their innate response mechanisms. Using these mechanisms, they can respond appropriately to their internal and external environment without having to learn.

Natural selection

Heritable adaptive behavior evolves through

. In this case, some genes better equip individuals to respond to environmental or physiological cues, thereby increasing reproductive success and causing these genes to persist in future generations. Non-adaptive behaviors cause a decrease in reproductive success so the genes are lost from future generations. These adaptive and non-adaptive characteristics arise through

random mutation Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...

, recombination, and/or

genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...

. Essentially, natural selection is a mechanism that contributes to directional gene selection in individuals that reproduce. Traits that cause greater reproductive success of an organism are favored, whereas those that reduce reproductive success are selected against. In contrast to learning, natural selection is solely a biological concept. It is the biological and genetic component that allows an adaptive behavior to be inherited with no connection to the environment. This form of adaptive behavior is most commonly considered in

ecological studies In epidemiology, ecological studies are used to understand the relationship between outcome and exposure at a population level, where 'population' represents a group of individuals with a shared characteristic such as geography, ethnicity, socio-ec ...

, and therefore natural selection is often used to explain ecological adaptive behavior in organisms.

Sexual selection

While kin selection is non-heritable and a direct result of the environment, sexual selection is a heritable adaptive behavior, and can therefore can be acted upon by natural selection. Sexual selection refers specifically to competition for mates.Andersson, M. (1995). Sexual Selection. Princeton, New Jersey: Princeton University Press. Many traits or features that are characteristic of a certain species can be explained by sexual selection as an adaptive behavior; this is because competition for mates results in specific traits being inherited. Only the species that are able to successfully compete and obtain a mate will reproduce and pass their genes on to the next generation. Therefore, species-specific genetic traits must be inherited, allowing individuals to be successful in their designated environments. There are many environmental examples of sexual selection as an adaptive behavior; one popular example is

sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...

. Sexual dimorphism is a morphological, phenotypic difference between males and females of the same species.Barlow, G.W. 2005. How do we decide that a species is sex-role reversed?. The Quarterly Review of Biology, 80: 28-35. A common example of sexual dimorphism is difference in body size. Sexual dimorphism can specifically be seen in the fish species,

Lamprologus callipterus ''Lamprologus callipterus'' is a species of cichlid endemic to Lake Tanganyika where it very actively moves about in search of crustaceans and other invertebrates. Males of this species can reach a length of TL while the females only grow to T ...

.Kazutaka, O., Masanori, K. and Tetsu, S. 2010. Unusual allometry for sexual size dimorphism in a cichlid where males are extremely larger than females. Journal of Biosciences, 35: 257-265. These male fish are substantially larger (sometimes up to 60 times) than their female counterparts. The male's increased size is advantageous because the larger individuals are able to compete for females, and subsequently defend their offspring, which grow inside empty snail shells until birth. Basically, the larger the male fish, the greater the adaptive benefit. The advantage of being larger and more powerful is demonstrated in the male's ability to compete and protect. In contrast to the males, the female fish must remain small in order to lay eggs inside the snail shells. It is evident that size plays a significant role in the reproductive success of this particular fish species. Large size is a common adaptive behavioral trait that is inherited through sexual selection and reproduction, as demonstrated in Lamprologus callipterus and other sexually dimorphic species.

Importance

It has been proven that adaptive behavior is crucial in the process of

, and thus is important in the

evolutionary process Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...

. Species that possess positive adaptive behaviors will inherently acquire evolutionary advantages. For example, adaptive behavior is a mechanism of

population stabilization Zero population growth, sometimes abbreviated ZPG, is a condition of demographic balance where the number of people in a specified population neither grows nor declines; that is, the number of births plus in-migrants equals the number of death ...

.Valdovinos, F.S., Ramos-Jiliberto, R., Garay-Narvaez, L., Urbani, P. and Dunne, J.A. 2010. Consequences of adaptive behavior for the structure and dynamic of food webs. Ecology Letters, 13: 1546-1559. In natural communities, organisms are able to interact with each other creating complex

food webs A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one ...

and

predator-prey dynamics Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation (wh ...

. Adaptive behavior helps modulate the dynamics of feeding relationships by having a direct effect on their feeding traits and strategies. These adaptive behaviors allow for resilience and resistance in the face of

disturbance Disturbance and its variants may refer to: Math and science * Disturbance (ecology), a temporary change in average environmental conditions that causes a pronounced change in an ecosystem * Disturbance (geology), linear zone of faults and folds ...

s and a changing environment. In

ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...

, the coexistence of

organisms In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fungi; ...

in natural ecosystems and the stability of populations are central topics. Currently, we live in a world experiencing great changes at a fast rate, mostly due to

anthropogenic Anthropogenic ("human" + "generating") is an adjective that may refer to: * Anthropogeny, the study of the origins of humanity Counterintuitively, anthropogenic may also refer to things that have been generated by humans, as follows: * Human im ...

impacts on organisms and the environment. By studying adaptive behavior one can understand

ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...

complexity – how it emerges, how it evolves, and how it can be maintained.

Measurement

An organism's behavioral strategies and ability to

adapt ADAPT (formerly American Disabled for Attendant Programs Today) is a United States grassroots disability rights organization with chapters in 30 states and Washington, D.C. They use nonviolent direct action in order to bring about disability just ...

will determine how they respond under different environmental conditions. Fitness is a common measure of adaptive success, and can be defined as the number of descendants left over after a group exhibits a particular behavioral strategy.Houston, A. and McNamara, J.M. (1999) Models of Adaptive Behavior. Cambridge University Press. Successful strategies will result in increased survival and

, which in turn can be characterized as a profitable behavioral adaption.

References

{{Ethology Behavioral ecology Ethology Evolutionary biology