Ethology
Overview
Ethology is the scientific study
Scientific method
Scientific method refers to a body of techniques for investigating phenomena, acquiring new knowledge, or correcting and integrating previous knowledge. To be termed scientific, a method of inquiry must be based on gathering empirical and measurable evidence subject to specific principles of...

 of animal behavior, and a sub-topic of zoology
Zoology
Zoology |zoölogy]]), is the branch of biology that relates to the animal kingdom, including the structure, embryology, evolution, classification, habits, and distribution of all animals, both living and extinct...

.

Although many naturalists have studied aspects of animal behavior throughout history, the modern discipline of ethology is generally considered to have begun during the 1930s with the work of Dutch biologist Nikolaas Tinbergen
Nikolaas Tinbergen
Nikolaas "Niko" Tinbergen was a Dutch ethologist and ornithologist who shared the 1973 Nobel Prize in Physiology or Medicine with Karl von Frisch and Konrad Lorenz for their discoveries concerning organization and elicitation of individual and social behaviour patterns in animals.In the 1960s he...

 and Austrian biologists Konrad Lorenz
Konrad Lorenz
Konrad Zacharias Lorenz was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize with Nikolaas Tinbergen and Karl von Frisch...

 and Karl von Frisch
Karl von Frisch
Karl Ritter von Frisch was an Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz....

, joint winners of the 1973 Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine administered by the Nobel Foundation, is awarded once a year for outstanding discoveries in the field of life science and medicine. It is one of five Nobel Prizes established in 1895 by Swedish chemist Alfred Nobel, the inventor of dynamite, in his will...

. Ethology is a combination of laboratory and field science, with a strong relation to certain other disciplines such as neuroanatomy
Neuroanatomy
Neuroanatomy is the study of the anatomy and organization of the nervous system. In contrast to animals with radial symmetry, whose nervous system consists of a distributed network of cells, animals with bilateral symmetry have segregated, defined nervous systems, and thus we can begin to speak of...

, ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

, and evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

.
Encyclopedia
Ethology is the scientific study
Scientific method
Scientific method refers to a body of techniques for investigating phenomena, acquiring new knowledge, or correcting and integrating previous knowledge. To be termed scientific, a method of inquiry must be based on gathering empirical and measurable evidence subject to specific principles of...

 of animal behavior, and a sub-topic of zoology
Zoology
Zoology |zoölogy]]), is the branch of biology that relates to the animal kingdom, including the structure, embryology, evolution, classification, habits, and distribution of all animals, both living and extinct...

.

Although many naturalists have studied aspects of animal behavior throughout history, the modern discipline of ethology is generally considered to have begun during the 1930s with the work of Dutch biologist Nikolaas Tinbergen
Nikolaas Tinbergen
Nikolaas "Niko" Tinbergen was a Dutch ethologist and ornithologist who shared the 1973 Nobel Prize in Physiology or Medicine with Karl von Frisch and Konrad Lorenz for their discoveries concerning organization and elicitation of individual and social behaviour patterns in animals.In the 1960s he...

 and Austrian biologists Konrad Lorenz
Konrad Lorenz
Konrad Zacharias Lorenz was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize with Nikolaas Tinbergen and Karl von Frisch...

 and Karl von Frisch
Karl von Frisch
Karl Ritter von Frisch was an Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz....

, joint winners of the 1973 Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine administered by the Nobel Foundation, is awarded once a year for outstanding discoveries in the field of life science and medicine. It is one of five Nobel Prizes established in 1895 by Swedish chemist Alfred Nobel, the inventor of dynamite, in his will...

. Ethology is a combination of laboratory and field science, with a strong relation to certain other disciplines such as neuroanatomy
Neuroanatomy
Neuroanatomy is the study of the anatomy and organization of the nervous system. In contrast to animals with radial symmetry, whose nervous system consists of a distributed network of cells, animals with bilateral symmetry have segregated, defined nervous systems, and thus we can begin to speak of...

, ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

, and evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

. Ethologists are typically interested in a behavioral process rather than in a particular animal group, and often study one type of behavior (e.g. aggression) in a number of unrelated animals.

The desire to understand animals has made ethology a rapidly growing field. Since the turn of the 21st century, many aspects of animal communication
Animal communication
Animal communication is any behavior on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, is sometimes called Zoosemiotics has played an important part in the...

, personal symbolic name use, animal emotions
Emotion in animals
There is no scientific consensus on emotion in animals, that is, what emotions certain species of animals, including humans, feel. The debate concerns primarily mammals and birds, although emotions have also been postulated for other vertebrates and even for some invertebrates.Animal lovers,...

, animal culture, learning
Learning
Learning is acquiring new or modifying existing knowledge, behaviors, skills, values, or preferences and may involve synthesizing different types of information. The ability to learn is possessed by humans, animals and some machines. Progress over time tends to follow learning curves.Human learning...

, and even sexual conduct
Animal sexuality
Animal sexual behaviour takes many different forms, even within the same species. Among animals other than humans, researchers have observed monogamy, promiscuity, sex between species, sexual arousal from objects or places, sex apparently via duress or coercion, copulation with dead animals,...

 that experts long thought they understood, have been reexamined, and new conclusions reached. New fields have developed, such as neuroethology
Neuroethology
Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system...

.

Etymology

The term ethology derives from the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

 word èthos (ήθος), meaning character. Other words that derive from ethos" include ethics and ethical. The term was first popularized by American myrmecologist William Morton Wheeler
William Morton Wheeler
William Morton Wheeler, Ph.D. was an American entomologist, myrmecologist and Harvard professor.-Early life:...

 in 1902. An earlier, slightly different sense of the term was proposed by John Stuart Mill
John Stuart Mill
John Stuart Mill was a British philosopher, economist and civil servant. An influential contributor to social theory, political theory, and political economy, his conception of liberty justified the freedom of the individual in opposition to unlimited state control. He was a proponent of...

 in his 1843 System of Logic. He recommended the development of a new science, "ethology," the purpose of which would be explanation of individual and national differences in character, on the basis of associationistic
Associationism
Associationism in philosophy refers to the idea that mental processes operate by the association of one state with its successor states.The idea is first recorded in Plato and Aristotle, especially with regard to the succession of memories....

 psychology
Psychology
Psychology is the study of the mind and behavior. Its immediate goal is to understand individuals and groups by both establishing general principles and researching specific cases. For many, the ultimate goal of psychology is to benefit society...

. This use of the word was never adopted.

Relationship with comparative psychology

Comparative psychology
Comparative psychology
Comparative psychology generally refers to the scientific study of the behavior and mental processes of non-human animals. However, scientists from different disciplines do not always agree on this definition...

 also studies animal behaviour, but, as opposed to ethology, is construed as a sub-topic of psychology
Psychology
Psychology is the study of the mind and behavior. Its immediate goal is to understand individuals and groups by both establishing general principles and researching specific cases. For many, the ultimate goal of psychology is to benefit society...

 rather than as one of biology
Biology
Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and taxonomy. Biology is a vast subject containing many subdivisions, topics, and disciplines...

. Historically, where comparative psychology researches animal behaviour in the context of what is known about human psychology, ethology researches animal behaviour in the context of what is known about animal anatomy
Anatomy
Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...

, physiology
Physiology
Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...

, neurobiology, and phylogenetic history. Furthermore, early comparative psychologists concentrated on the study of learning and tended to research behaviour in artificial situations, whereas early ethologists concentrated on behaviour in natural situations, tending to describe it as instinctive. The two approaches are complementary rather than competitive, but they do result in different perspectives and, sometimes, conflicts of opinion about matters of substance. In addition, for most of the twentieth century, comparative psychology developed most strongly in North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

, while ethology was stronger in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

. A practical difference is that early comparative psychologists concentrated on gaining extensive knowledge of the behaviour of very few species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

. Ethologists were more interested in understanding behaviour in a wide range of species to facilitate principled comparisons across taxonomic
Alpha taxonomy
Alpha taxonomy is the discipline concerned with finding, describing and naming species of living or fossil organisms. This field is supported by institutions holding collections of these organisms, with relevant data, carefully curated: such institutes include natural history museums, herbaria and...

 groups. Ethologists have made much more use of a truly comparative method
Comparative method
In linguistics, the comparative method is a technique for studying the development of languages by performing a feature-by-feature comparison of two or more languages with common descent from a shared ancestor, as opposed to the method of internal reconstruction, which analyzes the internal...

 than comparative psychologists have. Despite the historical divergence, most ethologists, at least in North America, teach in psychology departments. It is a strong belief among scientists that the mechanisms on which behavioural processes are based are the same that cause the evolution of the living species: there is therefore a strong association between these two fields.

Scala naturae and Lamarck's theories

Until the 19th century, the most common theory among scientists was still the concept of scala naturae
Great chain of being
The great chain of being , is a Christian concept detailing a strict, religious hierarchical structure of all matter and life, believed to have been decreed by the Christian God.-Divisions:...

, proposed by Aristotle
Aristotle
Aristotle was a Greek philosopher and polymath, a student of Plato and teacher of Alexander the Great. His writings cover many subjects, including physics, metaphysics, poetry, theater, music, logic, rhetoric, linguistics, politics, government, ethics, biology, and zoology...

. According to this theory, living beings were classified on an ideal pyramid that represented the simplest animals on the lower levels, with complexity increasing progressively toward the top, occupied by human beings. In the Western world of the time, people believed animal species were eternal and immutable, created with a specific purpose, as this seemed the only possible explanation for the incredible variety of living beings and their surprising adaptation to their habitats.

Jean-Baptiste Lamarck
Jean-Baptiste Lamarck
Jean-Baptiste Pierre Antoine de Monet, Chevalier de la Marck , often known simply as Lamarck, was a French naturalist...

 (1744 - 1829) was the first biologist to describe a complex theory of evolution. His theory substantially comprised two statements: first, that animal organs and behaviour can change according to the way they are used; and second, that those characteristics can transmit from one generation to the next (the example of the giraffe
Giraffe
The giraffe is an African even-toed ungulate mammal, the tallest of all extant land-living animal species, and the largest ruminant...

 whose neck becomes longer while trying to reach the upper leaves of a tree is well-known). The second statement is that every living organism, humans included, tends to reach a greater level of perfection. When Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

 went to the Galapagos Islands
Galápagos Islands
The Galápagos Islands are an archipelago of volcanic islands distributed around the equator in the Pacific Ocean, west of continental Ecuador, of which they are a part.The Galápagos Islands and its surrounding waters form an Ecuadorian province, a national park, and a...

, he was well aware of Lamarck's theories and was influenced by them.

Theory of evolution by natural selection and the beginnings of ethology

Because ethology is considered a topic of biology, ethologists have been concerned particularly with the evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 of behaviour and the understanding of behaviour in terms of the theory of natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....

. In one sense, the first modern ethologist was Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

, whose book, The Expression of the Emotions in Man and Animals, influenced many ethologists. He pursued his interest in behaviour by encouraging his protégé George Romanes
George Romanes
George John Romanes FRS was a Canadian-born English evolutionary biologist and physiologist who laid the foundation of what he called comparative psychology, postulating a similarity of cognitive processes and mechanisms between humans and other animals.He was the youngest of Charles Darwin's...

, who investigated animal learning and intelligence using an anthropomorphic method, anecdotal cognitivism
Anecdotal cognitivism
Anecdotal cognitivism is a psychological theory and animal cognition term which entails attribution of mental states to animals on the basis of anecdotes, and on the observation of particular cases, other than those observations made during controlled experiments...

, that did not gain scientific support.

Other early ethologists, such as Oskar Heinroth
Oskar Heinroth
Oskar Heinroth was a German biologist who was one of the first to apply the methods of comparative morphology to animal behaviour, and was thus one of the founders of ethology...

 and Julian Huxley
Julian Huxley
Sir Julian Sorell Huxley FRS was an English evolutionary biologist, humanist and internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentieth century evolutionary synthesis...

, instead concentrated on behaviours that can be called instinctive, or natural, in that they occur in all members of a species under specified circumstances. Their beginning for studying the behaviour of a new species was to construct an ethogram (a description of the main types of natural behaviour with their frequencies of occurrence). This provided an objective, cumulative base of data about behaviour, which subsequent researchers could check and supplement.

Fixed action patterns and animal communication

An important development, associated with the name of Konrad Lorenz
Konrad Lorenz
Konrad Zacharias Lorenz was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize with Nikolaas Tinbergen and Karl von Frisch...

 though probably due more to his teacher, Oskar Heinroth
Oskar Heinroth
Oskar Heinroth was a German biologist who was one of the first to apply the methods of comparative morphology to animal behaviour, and was thus one of the founders of ethology...

, was the identification of fixed action pattern
Fixed action pattern
In ethology, a fixed action pattern , or modal action pattern, is an instinctive behavioral sequence that is indivisible and runs to completion...

s (FAPs). Lorenz popularized FAPs as instinctive responses that would occur reliably in the presence of identifiable stimuli (called sign stimuli or releasing stimuli). These FAPs could then be compared across species, and the similarities and differences between behaviour could be easily compared with the similarities and differences in morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

. An important and much quoted study of the Anatidae
Anatidae
Anatidae is the biological family of birds that includes ducks, geese and swans. The family has a cosmopolitan distribution, occurring on all the world's continents except Antarctica and on most of the world's islands and island groups...

 (ducks and geese) by Heinroth used this technique. Ethologists noted that the stimuli that released FAPs were commonly features of the appearance or behaviour of other members of the animal's own species, and they were able to prove how important forms of animal communication
Animal communication
Animal communication is any behavior on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, is sometimes called Zoosemiotics has played an important part in the...

 could be mediated by a few simple FAPs. The most sophisticated investigation of this kind was the study by Karl von Frisch
Karl von Frisch
Karl Ritter von Frisch was an Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz....

 of the so-called "dance language" related to bee communication
Bee learning and communication
Honey bees learn and communicate in order to find food sources and for other means.-Learning:Learning is essential for efficient foraging. Honey bees are unlikely to make many repeat visits if a plant provides little in the way of reward...

. Lorenz developed an interesting theory of the evolution of animal communication based on his observations of the nature of fixed action patterns and the circumstances in which animals emit them.

Instinct

The Merriam-Webster dictionary
Webster's Dictionary
Webster's Dictionary refers to the line of dictionaries first developed by Noah Webster in the early 19th century, and also to numerous unrelated dictionaries that added Webster's name just to share his prestige. The term is a genericized trademark in the U.S.A...

 defines instinct
Instinct
Instinct or innate behavior is the inherent inclination of a living organism toward a particular behavior.The simplest example of an instinctive behavior is a fixed action pattern, in which a very short to medium length sequence of actions, without variation, are carried out in response to a...

 as a largely inheritable and unalterable tendency of an organism to make a complex and specific response to environmental stimuli without involving reason.
For ethologists, instinct
Instinct
Instinct or innate behavior is the inherent inclination of a living organism toward a particular behavior.The simplest example of an instinctive behavior is a fixed action pattern, in which a very short to medium length sequence of actions, without variation, are carried out in response to a...

 means a series of predictable behaviors for fixed action pattern
Fixed action pattern
In ethology, a fixed action pattern , or modal action pattern, is an instinctive behavioral sequence that is indivisible and runs to completion...

s. Such schemes are only acted when a precise stimulating signal is present. When such signals act as communication among members of the same species, they are known as releasers. A notable example of a releaser is the beak movements in many bird species performed by the newborns, which stimulates the mother's regurgitating process to feed her offspring. Another well-known case is the classic experiments by Tinbergen on the Graylag Goose. Like similar waterfowl
Waterfowl
Waterfowl are certain wildfowl of the order Anseriformes, especially members of the family Anatidae, which includes ducks, geese, and swans....

, the goose rolls a displaced egg
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...

 near its nest back to the others with its beak. The sight of the displaced egg triggers this mechanism. If the egg is taken away, the animal continues with the behaviour, pulling its head back as if an imaginary egg is still being maneuvered by the underside of its beak. However, it also attempts to move other egg-shaped objects, such as a giant plaster egg, door knob, or even a volleyball back into the nest. Such objects, when they exaggerate the releasers found in natural objects, can elicit a stronger version of the behavior than the natural object, so that the goose ignores its own displaced egg in favor of the giant dummy egg. These exaggerated releasers for instincts were named supernormal stimuli by Tinbergen. Tinbergen found he could produce supernormal stimuli for most instincts in animals—such as cardboard butterflies that male butterflies preferred to mate with if they had darker stripes than a real female, or dummy fish that a territorial male stickleback fish
Stickleback
The Gasterosteidae are a family of fish including the sticklebacks. FishBase currently recognises sixteen species in the family, grouped in five genera. However several of the species have a number of recognised subspecies, and the taxonomy of the family is thought to be in need of revision...

 fought more violently than a real invading male if the dummy had a brighter-colored underside. Harvard psychologist Deirdre Barrett
Deirdre Barrett
Deirdre Barrett, Ph.D. is an author and psychologist who teaches at Harvard Medical School. She is known for her research on dreams, hypnosis, and imagery and has written on evolutionary psychology. Barrett is a Past President of The International Association for the Study of Dreams and of the...

 has done research pointing out how easily humans also respond to supernormal stimuli for sexual, nurturing, feeding, and social instincts.
However, a behaviour only made of fixed action pattern
Fixed action pattern
In ethology, a fixed action pattern , or modal action pattern, is an instinctive behavioral sequence that is indivisible and runs to completion...

s would be particularly rigid and inefficient, reducing the probability of survival and reproduction, so the learning process has great importance, as does the ability to change the individual's responses based on its experience. It can be said that the more the brain
Brain
The brain is the center of the nervous system in all vertebrate and most invertebrate animals—only a few primitive invertebrates such as sponges, jellyfish, sea squirts and starfishes do not have one. It is located in the head, usually close to primary sensory apparatus such as vision, hearing,...

 is complex and the life of the individual long, the more its behaviour is "intelligent" (in the sense of being guided by experience rather than stereotyped FAPs).

Learning

Learning occurs in many ways, one of the most elementary being habituation
Habituation
Habituation can be defined as a process or as a procedure. As a process it is defined as a decrease in an elicited behavior resulting from the repeated presentation of an eliciting stimulus...

. This process consists of ignoring persistent or useless stimuli. An example of learning by habituation is the one observed in squirrels: When one of them feels threatened, the others hear its signal and go to the nearest refuge. However, if the signal comes from an individual that has caused many false alarms, the other squirrels ignore the signal.

Another common way of learning is by association
Association (psychology)
In psychology and marketing, two concepts or stimuli are associated when the experience of one leads to the effects of another, due to repeated pairing. This is sometimes called Pavlovian association for Ivan Pavlov's pioneering of classical conditioning....

, where a stimulus is, based on the experience, linked to another one that may not have anything to do with the first one. The first studies of associative learning were made by Russian physiologist Ivan Pavlov
Ivan Pavlov
Ivan Petrovich Pavlov was a famous Russian physiologist. Although he made significant contributions to psychology, he was not in fact a psychologist himself but was a mathematician and actually had strong distaste for the field....

. An example of associative behaviour is observed when a common goldfish goes close to the water surface whenever a human is going to feed it, or the excitement of a dog whenever it sees a collar
Collar (animal)
An animal collar is a device that attached to the neck of an animal to allow it to be harnessed, tied up or for various other reasons.*Pet collar. A piece of material put around the neck of certain pet animals, such as dogs or cats, for control, identification, or other purposes. Identification...

 as a prelude for a walk.

Imprinting

Being able to discriminate the members of one's own species is also of fundamental importance for reproductive success. Such discrimination can be based on a number of factors. However, this important type of learning only takes place in a very limited period of time. This kind of learning is called imprinting
Imprinting (psychology)
Imprinting is the term used in psychology and ethology to describe any kind of phase-sensitive learning that is rapid and apparently independent of the consequences of behavior...

, and was a second important finding of Lorenz. Lorenz observed that the young of birds such as geese
Goose
The word goose is the English name for a group of waterfowl, belonging to the family Anatidae. This family also includes swans, most of which are larger than true geese, and ducks, which are smaller....

 and chicken
Chicken
The chicken is a domesticated fowl, a subspecies of the Red Junglefowl. As one of the most common and widespread domestic animals, and with a population of more than 24 billion in 2003, there are more chickens in the world than any other species of bird...

s followed their mothers spontaneously from almost the first day after they were hatched, and he discovered that this response could be imitated by an arbitrary stimulus if the eggs were incubated artificially and the stimulus were presented during a critical period that continued for a few days after hatching.

Imitation

Imitation
Imitation
Imitation is an advanced behavior whereby an individual observes and replicates another's. The word can be applied in many contexts, ranging from animal training to international politics.-Anthropology and social sciences:...

 is often an important type of learning.
A well-documented example of imitative learning occurred in a group of macaque
Macaque
The macaques constitute a genus of Old World monkeys of the subfamily Cercopithecinae. - Description :Aside from humans , the macaques are the most widespread primate genus, ranging from Japan to Afghanistan and, in the case of the barbary macaque, to North Africa...

s on Hachijojima
Hachijojima
is a volcanic Japanese island in the Philippine Sea, administered by Tōkyō and located approximately south of the Special Wards of Tōkyō. It is the southernmost and most isolated of the Izu Seven Islands group of the seven northern islands of the Izu archipelago...

 Island, Japan. The macaques lived in the inland forest until the 1960s, when a group of researchers started giving them potatoes on the beach: soon, they started venturing onto the beach, picking the potatoes from the sand, and cleaning and eating them. About one year later, an individual was observed bringing a potato to the sea, putting it into the water with one hand, and cleaning it with the other. Her behaviour was soon imitated by the individuals living in contact with her; when they gave birth, they taught this practice to their young.

The National Institutes of Health reported that capuchin monkeys preferred the company of researchers who imitated them to that of researchers who did not. The monkeys not only spent more time with their imitators but also preferred to engage in a simple task with them even when provided with the option of performing the same task with a .

Mating and the fight for supremacy

Individual reproduction is the most important phase in the proliferation of individuals or genes within a species: for this reason, there exist complex mating
Mating
In biology, mating is the pairing of opposite-sex or hermaphroditic organisms for copulation. In social animals, it also includes the raising of their offspring. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization...

 ritual
Ritual
A ritual is a set of actions, performed mainly for their symbolic value. It may be prescribed by a religion or by the traditions of a community. The term usually excludes actions which are arbitrarily chosen by the performers....

s, which can be very complex even if they are often regarded as fixed action pattern
Fixed action pattern
In ethology, a fixed action pattern , or modal action pattern, is an instinctive behavioral sequence that is indivisible and runs to completion...

s (FAPs). The Stickleback
Stickleback
The Gasterosteidae are a family of fish including the sticklebacks. FishBase currently recognises sixteen species in the family, grouped in five genera. However several of the species have a number of recognised subspecies, and the taxonomy of the family is thought to be in need of revision...

's complex mating ritual was studied by Niko Tinbergen and is regarded as a notable example of a FAP.

Often in social life
Social relation
In social science, a social relation or social interaction refers to a relationship between two , three or more individuals . Social relations, derived from individual agency, form the basis of the social structure. To this extent social relations are always the basic object of analysis for social...

, animals fight for the right to reproduce, as well as social supremacy. A common example of fighting for social and sexual supremacy is the so-called pecking order
Pecking order
Pecking order or just peck order is the colloquial term for a hierarchical system of social organization in chickens. It was first described from the behaviour of poultry by Thorleif Schjelderup-Ebbe in 1921 under the German terms Hackordnung or Hackliste' ...

 among poultry
Poultry
Poultry are domesticated birds kept by humans for the purpose of producing eggs, meat, and/or feathers. These most typically are members of the superorder Galloanserae , especially the order Galliformes and the family Anatidae , commonly known as "waterfowl"...

. Every time a group of poultry cohabitate for a certain time length, they establish a pecking order. In these groups, one chicken dominates the others and can peck without being pecked. A second chicken can peck all the others except the first, and so on. Higher level chickens are easily distinguished by their well-cured aspect, as opposed to lower level chickens. While the pecking order is establishing, frequent and violent fights can happen, but once established, it is broken only when other individuals enter the group, in which case the pecking order re-establishes from scratch.

Living in groups

Several animal species, including humans, tend to live in groups. Group size
Group size measures
Many animals, including humans, tend to live in groups, herds, flocks, bands, packs, shoals, or colonies of conspecific individuals. The size of these groups, as expressed by the number of participant individuals, is an important aspect of their social environment...

 is a major aspect of their social environment. Social life
Social relation
In social science, a social relation or social interaction refers to a relationship between two , three or more individuals . Social relations, derived from individual agency, form the basis of the social structure. To this extent social relations are always the basic object of analysis for social...

 is probably a complex and effective survival strategy. It may be regarded as a sort of symbiosis
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

 among individuals of the same species: a society
Society
A society, or a human society, is a group of people related to each other through persistent relations, or a large social grouping sharing the same geographical or virtual territory, subject to the same political authority and dominant cultural expectations...

 is composed of a group of individuals belonging to the same species living within well-defined rules on food
Food
Food is any substance consumed to provide nutritional support for the body. It is usually of plant or animal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals...

 management, role assignments and reciprocal dependence.

When biologist
Biologist
A biologist is a scientist devoted to and producing results in biology through the study of life. Typically biologists study organisms and their relationship to their environment. Biologists involved in basic research attempt to discover underlying mechanisms that govern how organisms work...

s interested in evolution theory
Evolution Theory
Tin1 Yin2 Leun6 is Candy Lo's 9th studio album. It was released on 4 June 2005. For this album Candy Lo worked together with Hong Kong producer Kubert Leung with whom she worked on previous albums as well.- Track listing :...

 first started examining social behaviour, some apparently unanswerable questions arose, such as how the birth of sterile caste
Caste
Caste is an elaborate and complex social system that combines elements of endogamy, occupation, culture, social class, tribal affiliation and political power. It should not be confused with race or social class, e.g. members of different castes in one society may belong to the same race, as in India...

s, like in bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...

s, could be explained through an evolving mechanism that emphasizes the reproductive success of as many individuals as possible, or why, amongst animals living in small groups like squirrel
Squirrel
Squirrels belong to a large family of small or medium-sized rodents called the Sciuridae. The family includes tree squirrels, ground squirrels, chipmunks, marmots , flying squirrels, and prairie dogs. Squirrels are indigenous to the Americas, Eurasia, and Africa and have been introduced to Australia...

s, an individual would risk its own life to save the rest of the group. These behaviours may be examples of altruism
Altruism
Altruism is a concern for the welfare of others. It is a traditional virtue in many cultures, and a core aspect of various religious traditions, though the concept of 'others' toward whom concern should be directed can vary among cultures and religions. Altruism is the opposite of...

. Of course, not all behaviours are altruistic, as indicated by the table below. For example, revengeful behaviour was at one point claimed to have been observed exclusively in Homo sapiens. However, other species have been reported to be vengeful, including reports of vengeful camels and chimpanzees.
Classification of social behaviours
Type of behaviour Effect on the donor Effect on the receiver
Egoistic
Egotism
Egotism is "characterized by an exaggerated estimate of one's intellect, ability, importance, appearance, wit, or other valued personal characteristics" – the drive to maintain and enhance favorable views of oneself....

Increases fitness Decreases fitness
Cooperative
Cooperation
Cooperation or co-operation is the process of working or acting together. In its simplest form it involves things working in harmony, side by side, while in its more complicated forms, it can involve something as complex as the inner workings of a human being or even the social patterns of a...

Increases fitness Increases fitness
Altruistic
Altruism
Altruism is a concern for the welfare of others. It is a traditional virtue in many cultures, and a core aspect of various religious traditions, though the concept of 'others' toward whom concern should be directed can vary among cultures and religions. Altruism is the opposite of...

Decreases fitness Increases fitness
Revenge
Revenge
Revenge is a harmful action against a person or group in response to a grievance, be it real or perceived. It is also called payback, retribution, retaliation or vengeance; it may be characterized, justly or unjustly, as a form of justice.-Function in society:Some societies believe that the...

ful
Decreases fitness Decreases fitness


The existence of egoism
Egotism
Egotism is "characterized by an exaggerated estimate of one's intellect, ability, importance, appearance, wit, or other valued personal characteristics" – the drive to maintain and enhance favorable views of oneself....

 through natural selection does not pose any question to evolution theory
Evolution Theory
Tin1 Yin2 Leun6 is Candy Lo's 9th studio album. It was released on 4 June 2005. For this album Candy Lo worked together with Hong Kong producer Kubert Leung with whom she worked on previous albums as well.- Track listing :...

 and is, on the contrary, fully predicted by it, as is cooperative behaviour. It is more difficult to understand the mechanism through which altruistic behaviour initially developed.

Tinbergen's four questions for ethologists

Lorenz's collaborator, Niko Tinbergen, argued that ethology always needed to include four kinds of explanation in any instance of behaviour:
  • Function — How does the behaviour affect the animal's chances of survival and reproduction? Why does the animal respond that way instead of some other way?
  • Causation — What are the stimuli that elicit the response, and how has it been modified by recent learning?
  • Development — How does the behaviour change with age, and what early experiences are necessary for the animal to display the behaviour?
  • Evolutionary history — How does the behaviour compare with similar behaviour in related species, and how might it have begun through the process of phylogeny?


These explanations are complementary rather than mutually exclusive—all instances of behaviour require an explanation at each of these four levels. For example, the function of eating is to acquire nutrients (which ultimately aids survival and reproduction), but the immediate cause of eating is hunger (causation). Hunger and eating are evolutionarily ancient and are found in many species (evolutionary history), and develop early within an organism's lifespan (development). It is easy to confuse such questions—for example, to argue that people eat because they're hungry and not to acquire nutrients—without realizing that the reason people experience hunger is because it causes them to acquire nutrients.

Growth of the field

Due to the work of Lorenz and Tinbergen, ethology developed strongly in continental Europe during the years prior to World War II
World War II
World War II, or the Second World War , was a global conflict lasting from 1939 to 1945, involving most of the world's nations—including all of the great powers—eventually forming two opposing military alliances: the Allies and the Axis...

. After the war, Tinbergen moved to the University of Oxford
University of Oxford
The University of Oxford is a university located in Oxford, United Kingdom. It is the second-oldest surviving university in the world and the oldest in the English-speaking world. Although its exact date of foundation is unclear, there is evidence of teaching as far back as 1096...

, and ethology became stronger in the UK
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...

, with the additional influence of William Thorpe
William Homan Thorpe
William Homan Thorpe FRS was Professor of Animal Ethology at the University of Cambridge, and a significant British zoologist, ethologist and ornithologist....

, Robert Hinde
Robert Hinde
Robert Aubrey Hinde CBE FRS FBA is the Emeritus Royal Society Research Professor of Zoology at the University of Cambridge. He was formerly the master of St. John's College, Cambridge. He is the chair of British Pugwash...

, and Patrick Bateson
Patrick Bateson
Sir Patrick Bateson, FRS is an English biologist and science writer. Bateson is emeritus professor of ethology at Cambridge University and president of the Zoological Society of London since 2004....

 at the Sub-department of Animal Behaviour of the University of Cambridge
University of Cambridge
The University of Cambridge is a public research university located in Cambridge, United Kingdom. It is the second-oldest university in both the United Kingdom and the English-speaking world , and the seventh-oldest globally...

, located in the village of Madingley
Madingley
Madingley is a village near Coton and Dry Drayton on the western outskirts of Cambridge in the United Kingdom.Known as Madingelei in the Domesday Book, the village's name means "Woodland clearing of the family or followers of a man called Mada"....

. In this period, too, ethology began to develop strongly in North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

.

Lorenz, Tinbergen, and von Frisch were jointly awarded the Nobel Prize in Physiology or Medicine
Nobel Prize in Physiology or Medicine
The Nobel Prize in Physiology or Medicine administered by the Nobel Foundation, is awarded once a year for outstanding discoveries in the field of life science and medicine. It is one of five Nobel Prizes established in 1895 by Swedish chemist Alfred Nobel, the inventor of dynamite, in his will...

 in 1973 for their work of developing ethology.

Ethology is now a well-recognised scientific discipline, and has a number of journals covering developments in the subject, such as the Ethology Journal. In 1972, the International Society for Human Ethology was founded to promote exchange of knowledge and opinions concerning human behavior gained by applying ethological principles and methods and published their journal, The Human Ethology Bulletin. In 2008, in a paper published in the journal Behaviour, ethologist Peter Verbeek introduced the term "Peace Ethology" as a sub-discipline of Human Ethology that is concerned with issues of human conflict, conflict resolution, reconciliation, war, peacemaking, and peacekeeping behavior.

Today, along with actual ethologists, many biologists, zoologists, primatologists, anthropologists, veterinarians, and physicians study ethology and other related fields such as animal psychology, the study of animal social groups, and animal cognition. Some research has begun to study atypical or disordered animal behavior. Most researchers in the field have some sort of advanced degree and specialty and subspecialty training in the aforementioned fields.

Social ethology and recent developments

In 1970, the English
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

 ethologist John H. Crook published an important paper in which he distinguished comparative ethology from social ethology, and argued that much of the ethology that had existed so far was really comparative ethology—examining animals as individuals—whereas, in the future, ethologists would need to concentrate on the behaviour of social groups of animals and the social structure within them.

Also in 1970, Robert Ardrey
Robert Ardrey
Robert Ardrey was an American playwright and screenwriter who returned to his academic training in anthropology and the behavioral sciences in the 1950s....

's book The Social Contract: A Personal Inquiry into the Evolutionary Sources of Order and Disorder was published. The book and study investigated animal behaviour and then compared human behaviour to it as a similar phenomenon.

E. O. Wilson
E. O. Wilson
Edward Osborne Wilson is an American biologist, researcher , theorist , naturalist and author. His biological specialty is myrmecology, the study of ants....

's book Sociobiology: The New Synthesis
Sociobiology: The New Synthesis
Sociobiology: The New Synthesis is a book written by E. O. Wilson, which started the sociobiology debate, one of the great scientific controversies in biology of the 20th century...

 appeared in 1975, and since that time, the study of behaviour has been much more concerned with social aspects. It has also been driven by the stronger, but more sophisticated, Darwinism associated with Wilson, Robert Trivers, and William Hamilton. The related development of behavioural ecology has also helped transform ethology. Furthermore, a substantial rapprochement with comparative psychology has occurred, so the modern scientific study of behaviour offers a more or less seamless spectrum of approaches: from animal cognition
Animal cognition
Animal cognition is the title given to the study of the mental capacities of non-human animals. It has developed out of comparative psychology, but has also been strongly influenced by the approach of ethology, behavioral ecology, and evolutionary psychology...

 to more traditional comparative psychology
Comparative psychology
Comparative psychology generally refers to the scientific study of the behavior and mental processes of non-human animals. However, scientists from different disciplines do not always agree on this definition...

, ethology, sociobiology
Sociobiology
Sociobiology is a field of scientific study which is based on the assumption that social behavior has resulted from evolution and attempts to explain and examine social behavior within that context. Often considered a branch of biology and sociology, it also draws from ethology, anthropology,...

, and behavioural ecology. Sociobiology has more recently developed into evolutionary psychology
Evolutionary psychology
Evolutionary psychology is an approach in the social and natural sciences that examines psychological traits such as memory, perception, and language from a modern evolutionary perspective. It seeks to identify which human psychological traits are evolved adaptations, that is, the functional...

.

List of ethologists

People who have made notable contributions to ethology (many are comparative psychologists):
  • Robert Ardrey
    Robert Ardrey
    Robert Ardrey was an American playwright and screenwriter who returned to his academic training in anthropology and the behavioral sciences in the 1950s....

  • John C Angel
    John C Angel
    John C Angel was born in Illinois August 14, 1949. He was raised in Illinois and North Carolina and educated in England with university disciplines in physiology and ethology. He is the president of PTS Inc which specializes in fear and aggression in domesticated canines. His most recent book...

  • Adrian Simpson
    Adrian Simpson
    Adrian Simpson is a British television presenter, currently presenting on Sky News on the overnight programme. Simpson has won numerous accolades for his presenting style, which has been likened to that of Trevor McDonald in his early years....

  • Patrick Bateson
    Patrick Bateson
    Sir Patrick Bateson, FRS is an English biologist and science writer. Bateson is emeritus professor of ethology at Cambridge University and president of the Zoological Society of London since 2004....

  • John Bowlby
    John Bowlby
    Edward John Mostyn "John" Bowlby was a British psychologist, psychiatrist and psychoanalyst, notable for his interest in child development and for his pioneering work in attachment theory.- Family background :...

  • Donald Broom
    Donald Broom
    Donald Maurice Broom, is an English biologist and emeritus professor of animal welfare at Cambridge University.Donald Broom attended Whitgift School and subsequently St. Catharine's College, Cambridge. He received his BA and MA degrees in zoology and Ph.D. degree in animal behaviour from Cambridge...

  • Marian Stamp Dawkins
    Marian Stamp Dawkins
    Marian Ellina Stamp Dawkins is professor for animal behaviour at the University of Oxford, where she heads the Animal Behaviour Research Group. She has published several books, one of which has been translated into German, and many peer-reviewed papers. She is considered an expert in animal welfare...

  • Richard Dawkins
    Richard Dawkins
    Clinton Richard Dawkins, FRS, FRSL , known as Richard Dawkins, is a British ethologist, evolutionary biologist and author...

  • Irenäus Eibl-Eibesfeldt
    Irenäus Eibl-Eibesfeldt
    Irenäus Eibl-Eibesfeldt is founder of the field of human ethology. In authoring the book which bears that title, he applied ethology to humans by studying them in a perspective more common to volumes studying animal behavior....

  • John Endler
    John Endler
    Professor John A. Endler is an ethologist and evolutionary biologist noted for his work on the adaptation of vertebrates to their unique perceptual environments, and the ways in which animal sensory capacities and colour patterns co-evolve.-Life and career:...

  • Jean-Henri Fabre
  • Dian Fossey
    Dian Fossey
    Dian Fossey was an American zoologist who undertook an extensive study of gorilla groups over a period of 18 years. She studied them daily in the mountain forests of Rwanda, initially encouraged to work there by famous anthropologist Louis Leakey...

  • Karl von Frisch
    Karl von Frisch
    Karl Ritter von Frisch was an Austrian ethologist who received the Nobel Prize in Physiology or Medicine in 1973, along with Nikolaas Tinbergen and Konrad Lorenz....

  • Douglas P. Fry
    Douglas P. Fry
    Douglas P. Fry , is a docent and professor of anthropology, teacher in the Faculty of Social and Caring Sciences at Åbo Akademi University in Finland and adjunct research scientist in the Bureau of Applied Research in Anthropology at the University of Arizona.A renowned American anthropologist and...

  • Jane Goodall
    Jane Goodall
    Dame Jane Morris Goodall, DBE , is a British primatologist, ethologist, anthropologist, and UN Messenger of Peace. Considered to be the world's foremost expert on chimpanzees, Goodall is best known for her 45-year study of social and family interactions of wild chimpanzees in Gombe Stream National...

  • James L. Gould
    James L. Gould
    James L. Gould is an American ethologist, evolutionary biologist, and popular science writer. He has served as a professor of ecology and evolutionary biology at Princeton University since receiving his Ph.D in 1975...

  • Judith Hand
    Judith Hand
    Judith L. Hand is an evolutionary biologist, animal behaviorist , novelist, and pioneer in the emerging field of peace ethology. She writes on a variety of topics related to ethology, including the biological and evolutionary roots of war, gender differences in conflict resolution, empowering...

  • Clarence Ellis Harbison
    Clarence Ellis Harbison
    Clarence Ellis Harbison was an animal psychologist.-Biography:He was born in 1885 in Schenectady, New York. Previously divorced and widowed, he married Mary Louise Jorjorian in 1951 and had a son, Michael Harbison, in 1953. He died on October 1, 1960 in Preston, Connecticut.-References:...

  • Heini Hediger
    Heini Hediger
    Heini Hediger was a Swiss zoologist noted for work in proxemics in animal behavior and is known as the "father of zoo biology". Hediger was formerly the director of Tierpark Dählhölzli , Zoo Basel and Zürich Zoo ....

  • Oskar Heinroth
    Oskar Heinroth
    Oskar Heinroth was a German biologist who was one of the first to apply the methods of comparative morphology to animal behaviour, and was thus one of the founders of ethology...

  • Robert Hinde
    Robert Hinde
    Robert Aubrey Hinde CBE FRS FBA is the Emeritus Royal Society Research Professor of Zoology at the University of Cambridge. He was formerly the master of St. John's College, Cambridge. He is the chair of British Pugwash...

  • Bernard Hollander
    Bernard Hollander
    Bernard Hollander was a London psychiatrist and one of the main proponents of the new interest in phrenology in the early 20th century.-Life and work:...

  • Sarah Hrdy
  • Julian Huxley
    Julian Huxley
    Sir Julian Sorell Huxley FRS was an English evolutionary biologist, humanist and internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentieth century evolutionary synthesis...

  • Lynne Isbell
  • Julian Jaynes
    Julian Jaynes
    Julian Jaynes was an American psychologist, best known for his book The Origin of Consciousness in the Breakdown of the Bicameral Mind , in which he argued that ancient peoples were not conscious....

  • Erich Klinghammer
    Erich Klinghammer
    Erich Klinghammer was a biologist best known for his contributions to the fields of ethology and behavioural ecology, particularly that of canids...

  • John Krebs
    John Krebs
    John Richard Krebs, Baron Krebs FRS is a world leader in zoology and more specifically bird behaviour. He is currently the Principal of Jesus College, Oxford University...

  • Konrad Lorenz
    Konrad Lorenz
    Konrad Zacharias Lorenz was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize with Nikolaas Tinbergen and Karl von Frisch...

  • Aubrey Manning
    Aubrey Manning
    Professor Aubrey William George Manning OBE FRSE FIBiol is a distinguished English zoologist and broadcaster.-Life:...

  • Eugene Marais
    Eugene Marais
    Eugène Nielen Marais was a South African lawyer, naturalist, poet and writer.- His early years, before and during the Boer War :Marais was born in Pretoria, the thirteenth and last child of his parents, Jan Christiaan Nielen Marais and Catharina Helena Cornelia van Niekerk...

  • Patricia McConnell
    Patricia McConnell
    Patricia McConnell is an ethologist, author, advice columnist, and radio host. She holds a Ph.D. in zoology from the University of Wisconsin–Madison...

  • Desmond Morris
    Desmond Morris
    Desmond John Morris, born 24 January 1928 in Purton, north Wiltshire, is a British zoologist and ethologist, as well as a popular anthropologist. He is also known as a painter, television presenter and popular author.-Life:...

  • Martin Moynihan
    Martin Moynihan
    Martin Humphrey Moynihan was an influential behavioral evolutionary biologist and ornithologist who studied under Ernst Mayr and Niko Tinbergen, and was a contemporary of Desmond Morris and various other famous ethologists...

  • Caitlin O'Connell-Rodwell
  • Manny Puig
    Manny Puig
    Manny "Sharkman" Puig is an American wildlife educator and entertainer. Puig is known for his direct approach when dealing with dangerous animals such as sharks, black bears and American alligators...

  • Irene Pepperberg
    Irene Pepperberg
    Irene Maxine Pepperberg is a scientist noted for her studies in animal cognition, particularly in relation to parrots. She is an adjunct professor of psychology at Brandeis University and a lecturer at Harvard University...

  • George Romanes
    George Romanes
    George John Romanes FRS was a Canadian-born English evolutionary biologist and physiologist who laid the foundation of what he called comparative psychology, postulating a similarity of cognitive processes and mechanisms between humans and other animals.He was the youngest of Charles Darwin's...

  • Thomas A. Sebeok
  • B. F. Skinner
    B. F. Skinner
    Burrhus Frederic Skinner was an American behaviorist, author, inventor, baseball enthusiast, social philosopher and poet...

  • Barbara Smuts
    Barbara Smuts
    Barbara B. Smuts is an American anthropologist and psychologist noted for her research into baboons, dolphins, and chimpanzees.Smuts received a Bachelors Degree in Anthropology from Harvard University and a Ph.D in neurological and biological behavioral science from Stanford Medical School...

  • William Homan Thorpe
    William Homan Thorpe
    William Homan Thorpe FRS was Professor of Animal Ethology at the University of Cambridge, and a significant British zoologist, ethologist and ornithologist....

  • Niko Tinbergen
  • Jakob von Uexküll
    Jakob von Uexküll
    Jakob Johann von Uexküll was a Estonian biologist who worked in the fields of muscular physiology, animal behaviour studies, and the cybernetics of life. However, his most notable contribution is the notion of umwelt, used by semiotician Thomas Sebeok...

  • Frans de Waal
    Frans de Waal
    Fransiscus Bernardus Maria de Waal, PhD , is a Dutch primatologist and ethologist. He is the Charles Howard Candler professor of Primate Behavior in the Emory University psychology department in Atlanta, Georgia, and director of the Living Links Center at the Yerkes National Primate Research...

  • William Morton Wheeler
    William Morton Wheeler
    William Morton Wheeler, Ph.D. was an American entomologist, myrmecologist and Harvard professor.-Early life:...

  • E. O. Wilson
    E. O. Wilson
    Edward Osborne Wilson is an American biologist, researcher , theorist , naturalist and author. His biological specialty is myrmecology, the study of ants....

  • Amotz Zahavi
    Amotz Zahavi
    Amotz Zahavi is an Israeli evolutionary biologist, a Professor Emeritus at the Zoology Department of Tel Aviv University, and one of the founders of the Society for the Protection of Nature in Israel...


  • See also

    • Abnormal behaviour of birds in captivity
      Abnormal behaviour of birds in captivity
      -Abnormal behaviour :When housed under captive or commercial conditions, birds often show a range of abnormal behaviours. These are often self-injurious or harmful to other individuals, and include:*Feather pecking*Cannibalism*Vent pecking...

    • Altruism in animals
      Altruism in animals
      Altruism is a well-documented animal behaviour, which appears most obviously in kin relationships but may also be evident amongst wider social groups, in which an animal sacrifices its own well-being for the benefit of another animal.- Overview :...

    • Animal cognition
      Animal cognition
      Animal cognition is the title given to the study of the mental capacities of non-human animals. It has developed out of comparative psychology, but has also been strongly influenced by the approach of ethology, behavioral ecology, and evolutionary psychology...

    • Animal communication
      Animal communication
      Animal communication is any behavior on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, is sometimes called Zoosemiotics has played an important part in the...

    • Animal ethics
      Animal ethics
      Animal ethics can refer to:*Abolitionism *Animal law*Animal rights*Animal welfare*Intrinsic value *Wildlife managementSee also*Animal rights movement*Animal testing*Animal testing regulations*Cruelty to animals...

    • Animals in Translation
      Animals in Translation
      Animals in Translation: Using the Mysteries of Autism to Decode Animal Behavior is a 2005 book by Temple Grandin and co-written by Catherine Johnson...

       (book)
    • Animal welfare science
      Animal welfare science
      Animal Welfare Science is the scientific study of the welfare of animals as pets, in zoos, laboratories, on farms and in the wild. Although animal welfare has been of great concern for many thousands of years in religion and culture, the investigation of animal welfare using rigorous scientific...

    • Anthrozoology
      Anthrozoology
      Anthrozoology is the study of human-animal interaction. It is a modern interdisciplinary and burgeoning field that overlaps with a number of other disciplines, including anthropology, ethology, medicine, psychology, veterinary medicine and zoology...

    • Behavioral ecology
      Behavioral ecology
      Behavioral ecology, or ethoecology, is the study of the ecological and evolutionary basis for animal behavior, and the roles of behavior in enabling an animal to adapt to its environment...

    • Cognitive ethology
      Cognitive ethology
      The fusion of cognitive science and classical ethology into cognitive ethology "emphasizes observing animals under more-or-less natural conditions, with the objective of understanding the evolution, adaptation , causation, and development of the species-specific behavioral repertoire" - .- Relation...

    • Darwinian puzzle
      Darwinian puzzle
      A Darwinian puzzle is a trait that appears to reduce the fitness of individuals that possess it. Such traits attract the attention of evolutionary biologists...

    • Emotion in animals
      Emotion in animals
      There is no scientific consensus on emotion in animals, that is, what emotions certain species of animals, including humans, feel. The debate concerns primarily mammals and birds, although emotions have also been postulated for other vertebrates and even for some invertebrates.Animal lovers,...

    • Ethnic nepotism
      Ethnic nepotism
      Ethnic nepotism describes a human tendency for in-group bias or in-group favouritism applied by nepotism for people with the same ethnicity.- The theory :...

    • Etoecology
    • Intrinsic value (animal ethics)
      Intrinsic value (animal ethics)
      The intrinsic value of an animal refers to the value it possesses in its own right, as an end-in-itself, as opposed to its Instrumental value, its value to other animals . The phrase has been adopted by animal rights advocates...

    • Non-human animal sexuality
    • Phylogenetic comparative methods
      Phylogenetic comparative methods
      Phylogenetic comparative methods use information on the evolutionary relationships of organisms to compare species...

    • Sleep (non-human)
      Sleep (non-human)
      Sleep in non-human animals refers to how the behavioral and physiological state of sleep, mainly characterized by reversible unconsciousness, non-responsiveness to external stimuli, and motor passivity, appears in different categories of animals....

    • Sociophysiology
      Sociophysiology
      Sociophysiology is the "interplay between society and physical functioning" involving "collaboration of two neighboring sciences: physiology and sociology"...

    • Supernormal stimuli

    Further reading

    • Karen Shanor and Jagmeet Kanwal: Bats Sing, Mice Giggle: Revealing the Secret Lives of Animals, Icon (2009). 'Accessible to the lay reader and acceptable to the scientific community' (The Daily Telegraph
      The Daily Telegraph
      The Daily Telegraph is a daily morning broadsheet newspaper distributed throughout the United Kingdom and internationally. The newspaper was founded by Arthur B...

      ), 10 October 2009.

    External links



    Diagrams on Tinbergen's four questions
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