Ethology

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Honeybee workers perform the waggle dance to indicate the range and direction of food. Waggle dance photo.png
Honeybee workers perform the waggle dance to indicate the range and direction of food.
Great crested grebes perform a complex synchronised courtship display. Grebe Courtship (detail).jpg
Great crested grebes perform a complex synchronised courtship display.
Male impalas fighting during the rut Fighting impalas edit2.jpg
Male impalas fighting during the rut

Ethology is a field of zoology that investigates the behaviour of non-human animals. It has its scientific roots in the work of Charles Darwin and of American and German ornithologists of the late 19th and early 20th century, including Charles O. Whitman, Oskar Heinroth, and Wallace Craig. The modern discipline of ethology is generally considered to have begun during the 1930s with the work of the Dutch biologist Nikolaas Tinbergen and the Austrian biologists Konrad Lorenz and Karl von Frisch, the three winners of the 1973 Nobel Prize in Physiology or Medicine. Ethology combines laboratory and field science, with a strong relation to neuroanatomy, ecology, and evolutionary biology.

Contents

Etymology

The modern term ethology derives from the Greek language: ἦθος, ethos meaning "character" and -λογία , -logia meaning "the study of". The term was first popularized by the American entomologist William Morton Wheeler in 1902. [1]

History

The beginnings of ethology

Charles Darwin (1809-1882) explored the expression of emotions in animals. Charles Darwin 1880.jpg
Charles Darwin (1809–1882) explored the expression of emotions in animals.

Ethologists have been particularly interested in the evolution of behaviour and its explanation in terms of natural selection. In one sense, the first modern ethologist was Charles Darwin, whose 1872 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, who investigated animal learning and intelligence using an anthropomorphic method, anecdotal cognitivism, that did not gain scientific support. [2]

Other early ethologists, such as Eugène Marais, Charles O. Whitman, Oskar Heinroth, Wallace Craig and Julian Huxley, instead concentrated on behaviours that can be called instinctive in that they occur in all members of a species under specified circumstances. [3] [4] [1] Their starting point for studying the behaviour of a new species was to construct an ethogram, a description of the main types of behaviour with their frequencies of occurrence. This provided an objective, cumulative database of behaviour. [1]

Growth of the field

Due to the work of Konrad Lorenz and Niko Tinbergen, ethology developed strongly in continental Europe during the years prior to World War II. [1] After the war, Tinbergen moved to the University of Oxford, and ethology became stronger in the UK, with the additional influence of William Thorpe, Robert Hinde, and Patrick Bateson at the University of Cambridge. [5]

Lorenz, Tinbergen, and von Frisch were jointly awarded the Nobel Prize in Physiology or Medicine in 1973 for their work of developing ethology. [6]

Ethology is now a well-recognized scientific discipline, with its own journals such as Animal Behaviour , Applied Animal Behaviour Science , Animal Cognition , Behaviour , Behavioral Ecology and Ethology . In 1972, the International Society for Human Ethology was founded along with its journal, The Human Ethology Bulletin.[ citation needed ]

Social ethology

In 1972, the English ethologist John H. Crook 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. [7]

E. O. Wilson's book Sociobiology: The New Synthesis appeared in 1975, [8] and since that time, the study of behaviour has been much more concerned with social aspects. It has been driven by the Darwinism associated with Wilson, Robert Trivers, and W. D. Hamilton. The related development of behavioural ecology has helped transform ethology. [9] Furthermore, a substantial rapprochement with comparative psychology has occurred, so the modern scientific study of behaviour offers a spectrum of approaches. In 2020, Tobias Starzak and Albert Newen from the Institute of Philosophy II at the Ruhr University Bochum postulated that animals may have beliefs. [10]

Determinants of behaviour

Behaviour is determined by three major factors, namely inborn instincts, learning, and environmental factors. The latter include abiotic and biotic factors. Abiotic factors such as temperature or light conditions have dramatic effects on animals, especially if they are ectothermic or nocturnal. Biotic factors include members of the same species (e.g. sexual behavior), predators (fight or flight), or parasites and diseases.[ citation needed ]

Instinct

Kelp gull chicks peck at red spot on mother's beak to stimulate regurgitating reflex Larus Dominicanus with young.jpg
Kelp gull chicks peck at red spot on mother's beak to stimulate regurgitating reflex

Webster's Dictionary defines instinct as "A largely inheritable and unalterable tendency of an organism to make a complex and specific response to environmental stimuli without involving reason". [11]

Fixed action patterns

An important development, associated with the name of Konrad Lorenz though probably due more to his teacher, Oskar Heinroth, was the identification of fixed action patterns. Lorenz popularized these as instinctive responses that would occur reliably in the presence of identifiable stimuli called sign stimuli or "releasing stimuli". Fixed action patterns are now considered to be instinctive behavioural sequences that are relatively invariant within the species and that almost inevitably run to completion. [12]

One example of a releaser is the beak movements of many bird species performed by newly hatched chicks, which stimulates the mother to regurgitate food for her offspring. [13] Other examples are the classic studies by Tinbergen on the egg-retrieval behaviour and the effects of a "supernormal stimulus" on the behaviour of graylag geese. [14] [15]

One investigation of this kind was the study of the waggle dance ("dance language") in bee communication by Karl von Frisch. [16]

Learning

Habituation

Habituation is a simple form of learning and occurs in many animal taxa. It is the process whereby an animal ceases responding to a stimulus. Often, the response is an innate behavior. Essentially, the animal learns not to respond to irrelevant stimuli. For example, prairie dogs (Cynomys ludovicianus) give alarm calls when predators approach, causing all individuals in the group to quickly scramble down burrows. When prairie dog towns are located near trails used by humans, giving alarm calls every time a person walks by is expensive in terms of time and energy. Habituation to humans is therefore an important behavior in this context. [17] [18] [19]

Associative learning

Associative learning in animal behaviour is any learning process in which a new response becomes associated with a particular stimulus. [20] The first studies of associative learning were made by the Russian physiologist Ivan Pavlov, who observed that dogs trained to associate food with the ringing of a bell would salivate on hearing the bell. [21]

Imprinting

Imprinting in a moose. Moose-Imprinting-sr81-15.jpg
Imprinting in a moose.

Imprinting enables the young to discriminate the members of their own species, vital for reproductive success. This important type of learning only takes place in a very limited period of time. Konrad Lorenz observed that the young of birds such as geese and chickens 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. [22]

Cultural learning

Observational learning
Imitation

Imitation is an advanced behavior whereby an animal observes and exactly replicates the behavior of another. 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 non-imitator. [23] Imitation has been observed in recent research on chimpanzees; not only did these chimps copy the actions of another individual, when given a choice, the chimps preferred to imitate the actions of the higher-ranking elder chimpanzee as opposed to the lower-ranking young chimpanzee. [24]

Stimulus and local enhancement

Animals can learn using observational learning but without the process of imitation. One way is stimulus enhancement in which individuals become interested in an object as the result of observing others interacting with the object. [25] Increased interest in an object can result in object manipulation which allows for new object-related behaviours by trial-and-error learning. Haggerty (1909) devised an experiment in which a monkey climbed up the side of a cage, placed its arm into a wooden chute, and pulled a rope in the chute to release food. Another monkey was provided an opportunity to obtain the food after watching a monkey go through this process on four occasions. The monkey performed a different method and finally succeeded after trial-and-error. [26] Another example familiar to some cat and dog owners is the ability of their animals to open doors. The action of humans operating the handle to open the door results in the animals becoming interested in the handle and then by trial-and-error, they learn to operate the handle and open the door.

In local enhancement, a demonstrator attracts an observer's attention to a particular location. [27] Local enhancement has been observed to transmit foraging information among birds, rats and pigs. [28] The stingless bee ( Trigona corvina ) uses local enhancement to locate other members of their colony and food resources. [29]

Social transmission

A well-documented example of social transmission of a behaviour occurred in a group of macaques on Hachijojima 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. [8] 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. This behaviour was soon expressed by the individuals living in contact with her; when they gave birth, this behaviour was also expressed by their young—a form of social transmission. [30]

Teaching

Teaching is a highly specialized aspect of learning in which the "teacher" (demonstrator) adjusts their behaviour to increase the probability of the "pupil" (observer) achieving the desired end-result of the behaviour. For example, orcas are known to intentionally beach themselves to catch pinniped prey. [31] Mother orcas teach their young to catch pinnipeds by pushing them onto the shore and encouraging them to attack the prey. Because the mother orca is altering her behaviour to help her offspring learn to catch prey, this is evidence of teaching. [31] Teaching is not limited to mammals. Many insects, for example, have been observed demonstrating various forms of teaching to obtain food. Ants, for example, will guide each other to food sources through a process called "tandem running," in which an ant will guide a companion ant to a source of food. [32] It has been suggested that the pupil ant is able to learn this route to obtain food in the future or teach the route to other ants. This behaviour of teaching is also exemplified by crows, specifically New Caledonian crows. The adults (whether individual or in families) teach their young adolescent offspring how to construct and utilize tools. For example, Pandanus branches are used to extract insects and other larvae from holes within trees. [33]

Mating and the fight for supremacy

Courtship display of a sarus crane Courtship of sarus cranes.jpg
Courtship display of a sarus crane

Individual reproduction is the most important phase in the proliferation of individuals or genes within a species: for this reason, there exist complex mating rituals, which can be very complex even if they are often regarded as fixed action patterns. The stickleback's complex mating ritual, studied by Tinbergen, is regarded as a notable example. [34]

Often in social life, 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 among poultry. 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. Chickens higher in the pecking order may at times be distinguished by their healthier appearance when compared to lower level chickens.[ citation needed ] 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. [35]

Social behaviour

Several animal species, including humans, tend to live in groups. Group size is a major aspect of their social environment. Social life is probably a complex and effective survival strategy. It may be regarded as a sort of symbiosis among individuals of the same species: a society is composed of a group of individuals belonging to the same species living within well-defined rules on food management, role assignments and reciprocal dependence.[ citation needed ]

When biologists interested in evolution theory first started examining social behaviour, some apparently unanswerable questions arose, such as how the birth of sterile castes, like in bees, 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 squirrels, an individual would risk its own life to save the rest of the group. These behaviours may be examples of altruism. [36] 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 chimpanzees, [37] as well as anecdotal reports of vengeful camels. [38]

Classification of social behaviours [ citation needed ]
Type of behaviourEffect on the donorEffect on the receiver
Egoistic Neutral to Increases fitnessDecreases fitness
Cooperative Neutral to Increases fitnessNeutral to Increases fitness
Altruistic Decreases fitnessNeutral to Increases fitness
Revengeful Decreases fitnessDecreases fitness

Altruistic behaviour has been explained by the gene-centred view of evolution. [39] [40]

Benefits and costs of group living

One advantage of group living is decreased predation. If the number of predator attacks stays the same despite increasing prey group size, each prey has a reduced risk of predator attacks through the dilution effect. [9] [ page needed ] Further, according to the selfish herd theory, the fitness benefits associated with group living vary depending on the location of an individual within the group. The theory suggests that conspecifics positioned at the centre of a group will reduce the likelihood predations while those at the periphery will become more vulnerable to attack. [41] In groups, prey can also actively reduce their predation risk through more effective defence tactics, or through earlier detection of predators through increased vigilance. [9]

Another advantage of group living is an increased ability to forage for food. Group members may exchange information about food sources, facilitating the process of resource location. [9] [ page needed ] Honeybees are a notable example of this, using the waggle dance to communicate the location of flowers to the rest of their hive. [42] Predators also receive benefits from hunting in groups, through using better strategies and being able to take down larger prey. [9] [ page needed ]

Some disadvantages accompany living in groups. Living in close proximity to other animals can facilitate the transmission of parasites and disease, and groups that are too large may also experience greater competition for resources and mates. [43]

Group size

Theoretically, social animals should have optimal group sizes that maximize the benefits and minimize the costs of group living. However, in nature, most groups are stable at slightly larger than optimal sizes. [9] [ page needed ] Because it generally benefits an individual to join an optimally-sized group, despite slightly decreasing the advantage for all members, groups may continue to increase in size until it is more advantageous to remain alone than to join an overly full group. [44]

Tinbergen's four questions for ethologists

Niko Tinbergen argued that ethology needed to include four kinds of explanation in any instance of behaviour: [45] [46]

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 are hungry and not to acquire nutrients—without realizing that the reason people experience hunger is because it causes them to acquire nutrients. [47]

See also

Related Research Articles

<span class="mw-page-title-main">Konrad Lorenz</span> Austrian zoologist (1903–1989)

Konrad Zacharias Lorenz was an Austrian zoologist, ethologist, and ornithologist. He shared the 1973 Nobel Prize in Physiology or Medicine with Nikolaas Tinbergen and Karl von Frisch. He is often regarded as one of the founders of modern ethology, the study of animal behavior. He developed an approach that began with an earlier generation, including his teacher Oskar Heinroth.

<span class="mw-page-title-main">Nikolaas Tinbergen</span> Dutch zoologist and ethologist (1907–1988)

Nikolaas "Niko" Tinbergen was a Dutch biologist and ornithologist who shared the 1973 Nobel Prize in Physiology or Medicine with Karl von Frisch and Konrad Lorenz for their discoveries concerning the organization and elicitation of individual and social behavior patterns in animals. He is regarded as one of the founders of modern ethology, the study of animal behavior.

<span class="mw-page-title-main">Instinct</span> Behaviour due to innate biological factors

Instinct is the inherent inclination of a living organism towards a particular complex behaviour, containing innate (inborn) elements. The simplest example of an instinctive behaviour is a fixed action pattern (FAP), in which a very short to medium length sequence of actions, without variation, are carried out in response to a corresponding clearly defined stimulus.

Comparative psychology refers to the scientific study of the behavior and mental processes of non-human animals, especially as these relate to the phylogenetic history, adaptive significance, and development of behavior. The phrase comparative psychology may be employed in a narrow and a broad meaning. In its narrow meaning, it refers to the study of the similarities and differences in the psychology and behavior of different species. In a broader meaning, comparative psychology includes comparisons between different biological and socio-cultural groups, such as species, sexes, developmental stages, ages, and ethnicities. Research in this area addresses many different issues, uses many different methods and explores the behavior of many different species from insects to primates.

<span class="mw-page-title-main">Supernormal stimulus</span> Concept in biology and psychology

A supernormal stimulus or superstimulus is an exaggerated version of a stimulus to which there is an existing response tendency, or any stimulus that elicits a response more strongly than the stimulus for which it evolved.

<span class="mw-page-title-main">Animal cognition</span> Intelligence of non-human animals

Animal cognition encompasses the mental capacities of non-human animals including insect cognition. The study of animal conditioning and learning used in this field was developed from comparative psychology. It has also been strongly influenced by research in ethology, behavioral ecology, and evolutionary psychology; the alternative name cognitive ethology is sometimes used. Many behaviors associated with the term animal intelligence are also subsumed within animal cognition.

<span class="mw-page-title-main">Animal communication</span> Transfer of information from animal to animal

Animal communication is the transfer of information from one or a group of animals to one or more other animals that affects the current or future behavior of the receivers. Information may be sent intentionally, as in a courtship display, or unintentionally, as in the transfer of scent from predator to prey with kairomones. Information may be transferred to an "audience" of several receivers. Animal communication is a rapidly growing area of study in disciplines including animal behavior, sociology, neurology and animal cognition. Many aspects of animal behavior, such as symbolic name use, emotional expression, learning and sexual behavior, are being understood in new ways.

<i>On Aggression</i> 1963 book by Konrad Lorenz

On Aggression is a 1963 book by the ethologist Konrad Lorenz; it was translated into English in 1966. As he writes in the prologue, "the subject of this book is aggression, that is to say the fighting instinct in beast and man which is directed against members of the same species."

<span class="mw-page-title-main">Neuroethology</span> Study of animal behavior and its underlying mechanistic control by the nervous system

Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system. It is an interdisciplinary science that combines both neuroscience and ethology. A central theme of neuroethology, which differentiates it from other branches of neuroscience, is its focus on behaviors that have been favored by natural selection rather than on behaviors that are specific to a particular disease state or laboratory experiment.

<span class="mw-page-title-main">Oskar Heinroth</span> German zoologist

Oskar Heinroth was a German biologist who was one of the first to apply the methods of comparative morphology to animal behavior, and was thus one of the founders of ethology. He worked, largely isolated from most other scientists of the period, at the Berlin Aquarium where he took care of fishes, reptiles and birds, especially waterfowl.

Cognitive ethology is a branch of ethology concerned with the influence of conscious awareness and intention on the behaviour of an animal. Donald Griffin, a zoology professor in the United States, set up the foundations for researches in the cognitive awareness of animals within their habitats.

"Fixed action pattern" is an ethological term describing an instinctive behavioral sequence that is highly stereotyped and species-characteristic. Fixed action patterns are said to be produced by the innate releasing mechanism, a "hard-wired" neural network, in response to a sign/key stimulus or releaser. Once released, a fixed action pattern runs to completion.

Tinbergen's four questions, named after 20th century biologist Nikolaas Tinbergen, are complementary categories of explanations for animal behaviour. These are also commonly referred to as levels of analysis. It suggests that an integrative understanding of behaviour must include ultimate (evolutionary) explanations, in particular:

<span class="mw-page-title-main">Mobbing (animal behavior)</span> Antipredator adaptation in which individuals of prey species cooperatively attack a predator

Mobbing in animals is an antipredator adaptation in which individuals of prey species cooperatively attack or harass a predator, usually to protect their offspring. A simple definition of mobbing is an assemblage of individuals around a potentially dangerous predator. This is most frequently seen in birds, though it is also known to occur in many other animals such as the meerkat and some bovines. While mobbing has evolved independently in many species, it only tends to be present in those whose young are frequently preyed upon. This behavior may complement cryptic adaptations in the offspring themselves, such as camouflage and hiding. Mobbing calls may be used to summon nearby individuals to cooperate in the attack.

Vacuum activities are innate fixed action patterns (FAPs) of animal behaviour that are performed in the absence of a sign stimulus (releaser) that normally elicit them. This type of abnormal behaviour shows that a key stimulus is not always needed to produce an activity. Vacuum activities often take place when an animal is placed in captivity and is subjected to a lack of stimuli that would normally cause a FAP.

Socialization of animals is the process of training animals so that they can be kept in close relationship to humans.

Human ethology is the study of human behavior. Ethology as a discipline is generally thought of as a sub-category of biology, though psychological theories have been developed based on ethological ideas. The bridging between biological sciences and social sciences creates an understanding of human ethology. The International Society for Human Ethology is dedicated to advancing the study and understanding of human ethology.

Social learning refers to learning that is facilitated by observation of, or interaction with, another animal or its products. Social learning has been observed in a variety of animal taxa, such as insects, fish, birds, reptiles, amphibians and mammals.

<span class="mw-page-title-main">Hawk/goose effect</span> Behavior observed in some young birds when another bird flies above them

In ethology and cognitive ethology, the hawk/goose effect refers to a behavior observed in some young birds when another bird flies above them: if the flying bird is a goose, the young birds show no reaction, but if the flying bird is a hawk, the young birds either become more agitated or cower to reduce the danger. The observation that short-necked and long-tailed birds flying overhead caused alarm was noted by Oskar Heinroth. Friedrich Goethe conducted experiments with silhouettes to examine alarm reactions in 1937 and a more systematic study was conducted in the same year by Konrad Lorenz and Nikolaas Tinbergen which is considered one of the classic experiments of ethology.

Behavioral plasticity refers to a change in an organism's behavior that results from exposure to stimuli, such as changing environmental conditions. Behavior can change more rapidly in response to changes in internal or external stimuli than is the case for most morphological traits and many physiological traits. As a result, when organisms are confronted by new conditions, behavioral changes often occur in advance of physiological or morphological changes. For instance, larval amphibians changed their antipredator behavior within an hour after a change in cues from predators, but morphological changes in body and tail shape in response to the same cues required a week to complete.

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Further reading