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

, adaptive behavior is any

behavior Behavior (American English) or behaviour (British English) is the range of actions of Individual, individuals, organisms, systems or Artificial intelligence, artificial entities in some environment. These systems can include other systems or or ...

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 Heredity, heritable traits characteristic of a population over generation ...

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

sexual selection Sexual selection is a mechanism of evolution in which members of one 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 the opposite sex ...

of the most fit mate, and defending a

territory A territory is an area of land, sea, or space, belonging or connected to a particular country, person, or animal. In international politics, a territory is usually a geographic area which has not been granted the powers of self-government, ...

harem A harem is a domestic space that is reserved for the women of the house in a Muslim family. A harem may house a man's wife or wives, their pre-pubescent male children, unmarried daughters, female domestic Domestic worker, servants, and other un ...

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, fro ...

of unrelated young, and being a subordinate in a

dominance hierarchy In the zoological field of ethology, 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. Dif ...

. 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 Phenotypic trait, traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cell (biology), cells or orga ...

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

evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...

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

Non-heritable

Population Population is a set of humans or other organisms in a given region or area. Governments conduct a census to quantify the resident population size within a given jurisdiction. The term is also applied to non-human animals, microorganisms, and pl ...

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 Resource (biology), resources an ...

, 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, Attitude (psychology), attitudes, and preferences. The ability to learn is possessed by humans, non-human animals, and ...

Organisms An organism is any living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have been pr ...

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

classical conditioning Classical conditioning (also respondent conditioning and Pavlovian conditioning) is a behavioral procedure in which a biologically potent Stimulus (physiology), stimulus (e.g. food, a puff of air on the eye, a potential rival) is paired with a n ...

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. A learned adaptive behavior must have a

psychological Psychology is the scientific study of mind and behavior. Its subject matter includes the behavior of humans and nonhumans, both consciousness, conscious and Unconscious mind, unconscious phenomena, and mental processes such as thoughts, feel ...

, as well as a

biological Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, origin, evolution, and distribution of ...

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

Kin selection

Kin selection Kin selection is a process whereby natural selection favours a trait due to its positive effects on the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin selection can lead ...

(commonly referred to as

altruism Altruism is the concern for the well-being of others, independently of personal benefit or reciprocity. The word ''altruism'' was popularised (and possibly coined) by the French philosopher Auguste Comte in French, as , for an antonym of egoi ...

) 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. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...

.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. Frequency is an important parameter used in science and engineering to specify the rate of oscillatory and vibratory phenomena, such as mechanical vibrations, audio ...

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

cost 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 case the amount of money expended to acquire it i ...

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 (''Agelaius phoeniceus''). Therefore, correct learning of the birdsong early in life is necessary for territory defense in birds. European beavers (''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 has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...

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, recombination, and/or

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

. 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, 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 sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...

. 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''.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. Male ''L. calliptreus'' 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. Species that possess positive adaptive behaviors will inherently acquire evolutionary advantages. For example, adaptive behavior is a mechanism of population stabilization.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 and predator-prey dynamics. 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 disturbances and a changing environment. In

ecology Ecology () is the natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...

, the coexistence of

organisms An organism is any living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have been pr ...

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 Anthropogenic may also refer to things that have been generated by humans, as follows: * Human impact on the enviro ...

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

ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...

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

Measurement

An organism's behavioral strategies and ability to adapt 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 adaptation.

References

{{Ethology Behavioral ecology Evolutionary biology