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Friday, December 6, 2019

Konrad Lorenz

From Wikipedia, the free encyclopedia
https://en.wikipedia.org/wiki/Konrad_Lorenz
 
Konrad Lorenz

Konrad Lorenz.JPG
Born
Konrad Zacharias Lorenz

7 November 1903
Died27 February 1989 (aged 85)
Vienna, Austria
NationalityAustrian
Awards
Scientific career
FieldsEthology

Konrad Zacharias Lorenz (German pronunciation: [ˈkɔnʁaːt ˈloːʁɛnts]; 7 November 1903 – 27 February 1989) 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 behaviour. He developed an approach that began with an earlier generation, including his teacher Oskar Heinroth.

Lorenz studied instinctive behavior in animals, especially in greylag geese and jackdaws. Working with geese, he investigated the principle of imprinting, the process by which some nidifugous birds (i.e. birds that leave their nest early) bond instinctively with the first moving object that they see within the first hours of hatching. Although Lorenz did not discover the topic, he became widely known for his descriptions of imprinting as an instinctive bond. In 1936 he met Tinbergen, and the two collaborated in developing ethology as a separate sub-discipline of biology. A Review of General Psychology survey, published in 2002, ranked Lorenz as the 65th most cited scholar of the 20th century in the technical psychology journals, introductory psychology textbooks, and survey responses.

Lorenz's work was interrupted by the onset of World War II and in 1941 he was recruited into the German Army as a medic. In 1944 he was sent to the Eastern Front where he was captured by the Soviet Red Army and spent four years as a German prisoner of war. After the war he regretted his membership in the Nazi Party.

Lorenz wrote numerous books, some of which, such as King Solomon's Ring, On Aggression, and Man Meets Dog, became popular reading. His last work "Here I Am – Where Are You?" is a summary of his life's work and focuses on his famous studies of greylag geese.

Biography

Lorenz in 1904 with his older brother
 
In his autobiographical essay, published in 1973 in Les Prix Nobel (winners of the prizes are requested to provide such essays), Lorenz credits his career to his parents, who "were supremely tolerant of my inordinate love for animals", and to his childhood encounter with Selma Lagerlöf's The Wonderful Adventures of Nils, which filled him with a great enthusiasm about wild geese. "

At the request of his father, Adolf Lorenz, he began a premedical curriculum in 1922 at Columbia University, but he returned to Vienna in 1923 to continue his studies at the University of Vienna. He graduated as Doctor of Medicine (MD) in 1928 and became an assistant professor at the Institute of Anatomy until 1935. He finished his zoological studies in 1933 and received his second doctorate (PhD).

While still a student, Lorenz began developing what would become a large menagerie, ranging from domestic to exotic animals. In his popular book King Solomon's Ring, Lorenz recounts that while studying at the University of Vienna he kept a variety of animals at his parents' apartment, ranging from fish to a capuchin monkey named Gloria.

In 1936, at an international scientific symposium on instinct, Lorenz met his great friend and colleague Nikolaas Tinbergen. Together they studied geese—wild, domestic, and hybrid. One result of these studies was that Lorenz "realized that an overpowering increase in the drives of feeding as well as of copulation and a waning of more differentiated social instincts is characteristic of very many domestic animals". Lorenz began to suspect and fear "that analogous processes of deterioration may be at work with civilized humanity." This observation of bird hybrids caused Lorenz to believe that domestication resulting from urbanisation in humans might also cause dysgenic effects, and to argue in two papers that the Nazi eugenics policies against this were therefore scientifically justified.

Lorenz as a Soviet POW in 1944
 
In 1940 he became a professor of psychology at the University of Königsberg. He was drafted into the Wehrmacht in 1941. He sought to be a motorcycle mechanic, but instead he was assigned as a military psychologist, conducting racial studies on humans in occupied Poznań under Rudolf Hippius. The objective was to study the biological characteristics of "German-Polish half-breeds" to determine whether they 'benefitted' from the same work ethics as 'pure' Germans. The degree to which Lorenz participated in the project is unknown, but the project director Hippius referred a couple of times to Lorenz as an "examining psychologist".

Lorenz later described that he once saw transports of concentration camp inmates at Fort VII near Poznań, which made him "fully realize the complete inhumanity of the Nazis".

He was sent to the Russian front in 1944 where he quickly became a prisoner of war in the Soviet Union from 1944 to 1948. In captivity he continued to work as a medic and "got quite friendly with some Russians, mostly doctors". When he was repatriated, he was allowed to keep the manuscript of a book he had been writing, and his pet starling. He arrived back in Altenberg (his family home, near Vienna) both "with manuscript and bird intact." The manuscript became his 1973 book Behind the Mirror.

The Max Planck Society established the Lorenz Institute for Behavioral Physiology in Buldern, Germany, in 1950. In his memoirs Lorenz described the chronology of his war years differently from what historians have been able to document after his death. He himself claimed that he was captured in 1942, where in reality he was only sent to the front and captured in 1944, leaving out entirely his involvement with the Poznań project.

In 1958, Lorenz transferred to the Max Planck Institute for Behavioral Physiology in Seewiesen. He shared the 1973 Nobel Prize in Physiology or Medicine "for discoveries in individual and social behavior patterns" with two other important early ethologists, Nikolaas Tinbergen and Karl von Frisch. In 1969, he became the first recipient of the Prix mondial Cino Del Duca. He was a friend and student of renowned biologist Sir Julian Huxley (grandson of "Darwin's bulldog", Thomas Henry Huxley). Famed psychoanalyst Ralph Greenson and Sir Peter Scott were good friends. Lorenz and Karl Popper were childhood friends; many years after they met, during the celebration of Popper's 80 years, they wrote together a book entitled Die Zukunft ist offen.

He retired from the Max Planck Institute in 1973 but continued to research and publish from Altenberg and Grünau im Almtal in Austria. He died on 27 February 1989 in Altenberg. 

Ethology

Lorenz is recognized as one of the founding fathers of the field of ethology, the study of animal behavior. He is best known for his discovery of the principle of attachment, or imprinting, through which in some species a bond is formed between a new born animal and its caregiver. This principle had been discovered by Douglas Spalding in the 19th century, and Lorenz's mentor Oskar Heinroth had also worked on the topic, but Lorenz's description of Prägung, imprinting, in nidifugous birds such as greylag geese in his 1935 book Der Kumpan in der Umwelt des Vogels ("The Companion in the Environment of Birds") became the foundational description of the phenomenon.

Here, Lorenz used Jakob von Uexküll's concept of Umwelt to understand how the limited perception of animals filtered out certain phenomena with which they interacted instinctively. For example, a young goose instinctively bonds with the first moving stimulus it perceives, whether it be its mother, or a person. Lorenz showed that this behavior of imprinting is what allows the goose to learn to recognize members of its own species, enabling them to be the object of subsequent behavior patterns such as mating. He developed a theory of instinctive behavior that saw behavior patterns as largely innate but triggered through environmental stimuli, for example the hawk/goose effect. He argued that animals have an inner drive to carry out instinctive behaviors, and that if they do not encounter the right stimulus they will eventually engage in the behavior with an inappropriate stimulus.

Lorenz's approach to ethology derived from a skepticism towards the studies of animal behavior done in laboratory settings. He considered that in order to understand the mechanisms of animal behavior, it was necessary to observe their full range of behaviors in their natural context. Lorenz did not carry out much traditional fieldwork but observed animals near his home. His method involved empathizing with animals, often using anthropomorphization to imagine their mental states. He believed that animals were capable of experiencing many of the same emotions as humans.

Tinbergen, Lorenz's friend with whom he conjointly received the Nobel prize, summarized Lorenz's major contribution to ethology as making behavior a topic of biological inquiry, considering behavior a part of an animal's evolutionary equipment. Tinbergen and Lorenz contributed to making Ethology a recognized sub-discipline within Biology and founded the first specialized journal of the field "Ethology" (originally "Zeitschift für Tierpsychologie").

Politics

Lorenz joined the Nazi Party in 1938 and accepted a university chair under the Nazi regime. In his application for party membership he wrote, "I'm able to say that my whole scientific work is devoted to the ideas of the National Socialists." His publications during that time led in later years to allegations that his scientific work had been contaminated by Nazi sympathies. His published writing during the Nazi period included support for Nazi ideas of "racial hygiene" couched in pseudoscientific metaphors.

In his autobiography, Lorenz wrote:
I was frightened—as I still am—by the thought that analogous genetical processes of deterioration may be at work with civilized humanity. Moved by this fear, I did a very ill-advised thing soon after the Germans had invaded Austria: I wrote about the dangers of domestication and, in order to be understood, I couched my writing in the worst of Nazi terminology. I do not want to extenuate this action. I did, indeed, believe that some good might come of the new rulers. The precedent narrow-minded catholic regime in Austria induced better and more intelligent men than I was to cherish this naive hope. Practically all my friends and teachers did so, including my own father who certainly was a kindly and humane man. None of us as much as suspected that the word "selection", when used by these rulers, meant murder. I regret those writings not so much for the undeniable discredit they reflect on my person as for their effect of hampering the future recognition of the dangers of domestication.
After the war, Lorenz denied having been a party member, until his membership application was made public; and he denied having known the extent of the genocide, despite his position as a psychologist in the Office of Racial Policy. He was also shown to have made anti-Semitic jokes on 'Jewish characteristics' in letters to his mentor Heinroth. In 2015 the University of Salzburg posthumously rescinded an honorary doctorate awarded to Lorenz in 1983, citing his party membership and his assertions in his application that he was "always a National Socialist", and that his work "stands to serve National Socialist thought". The university also accused him of using his work to spread "basic elements of the racist ideology of National Socialism".

During the final years of his life, Lorenz supported the fledgling Austrian Green Party and in 1984 became the figurehead of the Konrad Lorenz Volksbegehren, a grass-roots movement that was formed to prevent the building of a power plant at the Danube near Hainburg an der Donau and thus the destruction of the surrounding woodland.

Contributions and legacy

With Nikolaas Tinbergen (left), 1978
 
Lorenz has been called 'The father of ethology', by Niko Tinbergen. Perhaps Lorenz's most important contribution to ethology was his idea that behavior patterns can be studied as anatomical organs. This concept forms the foundation of ethological research. However, Richard Dawkins called Lorenz a "'good of the species' man", stating that the idea of group selection was "so deeply ingrained" in Lorenz's thinking that he "evidently did not realize that his statements contravened orthodox Darwinian theory."

Together with Nikolaas Tinbergen, Lorenz developed the idea of an innate releasing mechanism to explain instinctive behaviors (fixed action patterns). They experimented with "supernormal stimuli" such as giant eggs or dummy bird beaks which they found could release the fixed action patterns more powerfully than the natural objects for which the behaviors were adapted. Influenced by the ideas of William McDougall, Lorenz developed this into a "psychohydraulic" model of the motivation of behavior, which tended towards group selectionist ideas, which were influential in the 1960s. Another of his contributions to ethology is his work on imprinting. His influence on a younger generation of ethologists; and his popular works, were important in bringing ethology to the attention of the general public. 

Lorenz claimed that there was widespread contempt for the descriptive sciences. He attributed this to the denial of perception as the source of all scientific knowledge: "a denial that has been elevated to the status of religion." He wrote that in comparative behavioral research, "it is necessary to describe various patterns of movement, record them, and above all, render them unmistakably recognizable."

There are three research institutions named after Lorenz in Austria: the Konrad Lorenz Institute for Evolution and Cognition Research (KLI) was housed in Lorenz' family mansion at Altenberg before moving to Klosterneuburg in 2013; the Konrad Lorenz Forschungsstelle (KLF) at his former field station in Grünau; and the Konrad Lorenz Institute of Ethology, an external research facility of the University of Veterinary Medicine Vienna.

Vision of the challenges facing humanity

With Nikolaas Tinbergen (right), 1978
 
Lorenz predicted the relationship between market economics and the threat of ecological catastrophe. In his 1973 book, Civilized Man's Eight Deadly Sins, Lorenz addresses the following paradox:
All the advantages that man has gained from his ever-deepening understanding of the natural world that surrounds him, his technological, chemical and medical progress, all of which should seem to alleviate human suffering... tends instead to favor humanity's destruction
Lorenz adopts an ecological model to attempt to grasp the mechanisms behind this contradiction. Thus "all species... are adapted to their environment... including not only inorganic components... but all the other living beings that inhabit the locality." p31.

Fundamental to Lorenz's theory of ecology is the function of negative feedback mechanisms, which, in hierarchical fashion, dampen impulses that occur beneath a certain threshold. The thresholds themselves are the product of the interaction of contrasting mechanisms. Thus pain and pleasure act as checks on each other:
To gain a desired prey, a dog or wolf will do things that, in other contexts, they would shy away from: run through thorn bushes, jump into cold water and expose themselves to risks which would normally frighten them. All these inhibitory mechanisms... act as a counterweight to the effects of learning mechanisms... The organism cannot allow itself to pay a price which is not worth paying. p53.
In nature, these mechanisms tend towards a 'stable state' among the living beings of an ecology:
A closer examination shows that these beings... not only do not damage each other, but often constitute a community of interests. It is obvious that the predator is strongly interested in the survival of that species, animal or vegetable, which constitutes its prey. ... It is not uncommon that the prey species derives specific benefits from its interaction with the predator species... pp31–33.
Lorenz states that humanity is the one species not bound by these mechanisms, being the only one that has defined its own environment:
[The pace of human ecology] is determined by the progress of man's technology (p35)... human ecology (economy) is governed by mechanisms of POSITIVE feedback, defined as a mechanism which tends to encourage behavior rather than to attenuate it (p43). Positive feedback always involves the danger of an 'avalanche' effect... One particular kind of positive feedback occurs when individuals OF THE SAME SPECIES enter into competition among themselves... For many animal species, environmental factors keep... intraspecies selection from [leading to] disaster... But there is no force which exercises this type of healthy regulatory effect on humanity's cultural development; unfortunately for itself, humanity has learned to overcome all those environmental forces which are external to itself p44.
Regarding aggression in human beings, Lorenz states:
Let us imagine that an absolutely unbiased investigator on another planet, perhaps on Mars, is examining human behavior on earth, with the aid of a telescope whose magnification is too small to enable him to discern individuals and follow their separate behavior, but large enough for him to observe occurrences such as migrations of peoples, wars, and similar great historical events. He would never gain the impression that human behavior was dictated by intelligence, still less by responsible morality. If we suppose our extraneous observer to be a being of pure reason, devoid of instincts himself and unaware of the way in which all instincts in general and aggression in particular can miscarry, he would be at a complete loss how to explain history at all. The ever-recurrent phenomena of history do not have reasonable causes. It is a mere commonplace to say that they are caused by what common parlance so aptly terms "human nature." Unreasoning and unreasonable human nature causes two nations to compete, though no economic necessity compels them to do so; it induces two political parties or religions with amazingly similar programs of salvation to fight each other bitterly, and it impels an Alexander or a Napoleon to sacrifice millions of lives in his attempt to unite the world under his scepter. We have been taught to regard some of the persons who have committed these and similar absurdities with respect, even as "great" men, we are wont to yield to the political wisdom of those in charge, and we are all so accustomed to these phenomena that most of us fail to realize how abjectly stupid and undesirable the historical mass behavior of humanity actually is 
Lorenz does not see human independence from natural ecological processes as necessarily bad. Indeed, he states that:
A completely new [ecology] which corresponds in every way to [humanity's] desires... could, theoretically, prove as durable as that which would have existed without his intervention (36).
However, the principle of competition, typical of Western societies, destroys any chance of this:
The competition between human beings destroys with cold and diabolic brutality... Under the pressure of this competitive fury we have not only forgotten what is useful to humanity as a whole, but even that which is good and advantageous to the individual. [...] One asks, which is more damaging to modern humanity: the thirst for money or consuming haste... in either case, fear plays a very important role: the fear of being overtaken by one's competitors, the fear of becoming poor, the fear of making wrong decisions or the fear of not being up to snuff... pp45–47.
In this book, Lorenz proposes that the best hope for mankind lies in our looking for mates based on the kindness of their hearts rather than good looks or wealth. He illustrates this with a Jewish story, explicitly described as such.

Lorenz was one of the early scientists who recognised the significance of human overpopulation. The number one deadly sin of civilized man in his book is overpopulation, which is what leads to aggression. 

Philosophical speculations

In his 1973 book Behind the Mirror: A Search for a Natural History of Human Knowledge, Lorenz considers the old philosophical question of whether our senses correctly inform us about the world as it is, or provide us only with an illusion. His answer comes from evolutionary biology. Only traits that help us survive and reproduce are transmitted. If our senses gave us wrong information about our environment, we would soon be extinct. Therefore, we can be sure that our senses give us correct information, for otherwise we would not be here to be deceived. 

Honours and awards

Works

Lorenz's best-known books are King Solomon's Ring and On Aggression, both written for a popular audience. His scientific work appeared mainly in journal articles, written in German; it became widely known to English-speaking scientists through its description in Tinbergen's 1951 book The Study of Instinct, though many of his papers were later published in English translation in the two volumes titled Studies in Animal and Human Behavior.
  • King Solomon's Ring (1949) (Er redete mit dem Vieh, den Vögeln und den Fischen, 1949)
  • Man Meets Dog (1950) (So kam der Mensch auf den Hund, 1950)
  • Evolution and Modification of Behaviour (1965)
  • On Aggression (1966) (Das sogenannte Böse. Zur Naturgeschichte der Aggression, 1963)
  • Studies in Animal and Human Behavior, Volume I (1970)
  • Studies in Animal and Human Behavior, Volume II (1971)
  • Motivation of Human and Animal Behavior: An Ethological View. With Paul Leyhausen (1973). New York: D. Van Nostrand Co. ISBN 0-442-24886-5
  • Behind the Mirror: A Search for a Natural History of Human Knowledge (1973) (Die Rückseite des Spiegels. Versuch einer Naturgeschichte menschlichen Erkennens, 1973)
  • Civilized Man's Eight Deadly Sins (1974) (Die acht Todsünden der zivilisierten Menschheit, 1973)
  • The Year of the Greylag Goose (1979) (Das Jahr der Graugans, 1979)
  • The Foundations of Ethology (1982)
  • The Waning of Humaneness (1987) (Der Abbau des Menschlichen, 1983)
  • Here I Am – Where Are You? – A Lifetime's Study of the Uncannily Human Behaviour of the Greylag Goose. (1988). Translated by Robert D. Martin from Hier bin ich – wo bist du?.
  • The Natural Science of the Human Species: An Introduction to Comparative Behavioral Research – The Russian Manuscript (1944–1948) (1995)

Emotion in animals

From Wikipedia, the free encyclopedia
https://en.wikipedia.org/wiki/Emotion_in_animals
 
A drawing of a cat by T. W. Wood in Charles Darwin's book The Expression of the Emotions in Man and Animals, described as acting "in an affectionate frame of mind".
 
Worldwide laws regarding the formal recognition of nonhuman animal sentience and suffering
  
National recognition of animal sentience
  
Partial recognition of animal sentience1
  
National recognition of animal suffering
  
Partial recognition of animal suffering2
  
No recognition of animal sentience or suffering
  
Unknown
1certain animals are excluded, only mental health is acknowledged, and/or the laws vary internally
2only includes domestic animals


Emotion is defined as any mental experience with high intensity and high hedonic content. The existence and nature of emotions in animals are believed to be correlated with those of humans and to have evolved from the same mechanisms. Charles Darwin was one of the first scientists to write about the subject, and his observational (and sometimes anecdotal) approach has since developed into a more robust, hypothesis-driven, scientific approach. Cognitive bias tests and learned helplessness models have shown feelings of optimism and pessimism in a wide range of species, including rats, dogs, cats, rhesus macaques, sheep, chicks, starlings, pigs, and honeybees. Jaak Panksepp played a large role in the study of animal emotion, basing his research on the neurological aspect. Mentioning seven core emotional feelings reflected through a variety of neuro-dynamic limbic emotional action systems, including seeking, fear, rage, lust, care, panic and play. Through brain stimulation and pharmacological challenges, such emotional responses can be effectively monitored.

Emotion has been observed and further researched through multiple different approaches including that of behaviourism, comparative, anecdotal, specifically Darwin's approach and what is most widely used today the scientific approach which has a number of subfields including functional, mechanistic, cognitive bias tests, self-medicating, spindle neurons, vocalizations and neurology.

While emotions in animals is still quite a controversial topic it has been studied in a extensive array of species both large and small including primates, rodents, elephants, horses, birds, dogs, cats, honeybees and crayfish. 

Etymology, definitions, and differentiation

The word "emotion" dates back to 1579, when it was adapted from the French word émouvoir, which means "to stir up". However, the earliest precursors of the word likely date back to the very origins of language.

Emotions have been described as discrete and consistent responses to internal or external events which have a particular significance for the organism. Emotions are brief in duration and consist of a coordinated set of responses, which may include physiological, behavioural, and neural mechanisms. Emotions have also been described as the result of evolution because they provided good solutions to ancient and recurring problems that faced ancestors.

Laterality

It has been proposed that negative, withdrawal-associated emotions are processed predominantly by the right hemisphere, whereas the left hemisphere is largely responsible for processing positive, approach-related emotions. This has been called the "laterality-valence hypothesis".

Basic and complex human emotions

In humans, a distinction is sometimes made between "basic" and "complex" emotions. Six emotions have been classified as basic: anger, disgust, fear, happiness, sadness and surprise. Complex emotions would include contempt, jealousy and sympathy. However, this distinction is difficult to maintain, and animals are often said to express even the complex emotions.

Background

Behaviourist approach

A squirrel communicating with its pup
 
Prior to the development of animal sciences such as comparative psychology and ethology, interpretation of animal behaviour tended to favour a minimalistic approach known as behaviourism. This approach refuses to ascribe to an animal a capability beyond the least demanding that would explain a behaviour; anything more than this is seen as unwarranted anthropomorphism. The behaviourist argument is, why should humans postulate consciousness and all its near-human implications in animals to explain some behaviour, if mere stimulus-response is a sufficient explanation to produce the same effects? 

Some behaviourists, such as John B. Watson, claim that stimulus–response models provide a sufficient explanation for animal behaviours that have been described as emotional, and that all behaviour, no matter how complex, can be reduced to a simple stimulus-response association. Watson described that the purpose of psychology was "to predict, given the stimulus, what reaction will take place; or given the reaction, state what the situation or stimulus is that has caused the reaction".
The cautious wording of Dixon exemplifies this viewpoint:


Moussaieff Masson and McCarthy describe a similar view (with which they disagree):


Because of the philosophical questions of consciousness and mind that are involved, many scientists have stayed away from examining animal and human emotion, and have instead studied measurable brain functions through neuroscience.

Comparative approach

In 1903, C. Lloyd Morgan published Morgan's Canon, a specialised form of Occam's razor used in ethology, in which he stated:

Darwin's approach

Charles Darwin initially planned to include a chapter on emotion in The Descent of Man but as his ideas progressed they expanded into a book, The Expression of the Emotions in Man and Animals. Darwin proposed that emotions are adaptive and serve a communicative and motivational function, and he stated three principles that are useful in understanding emotional expression: First, The Principle of Serviceable Habits takes a Lamarckian stance by suggesting that emotional expressions that are useful will be passed on to the offspring. Second, The Principle of Antithesis suggests that some expressions exist merely because they oppose an expression that is useful. Third, The Principle of the Direct Action of the Excited Nervous System on the Body suggests that emotional expression occurs when nervous energy has passed a threshold and needs to be released.

Darwin saw emotional expression as an outward communication of an inner state, and the form of that expression often carries beyond its original adaptive use. For example, Darwin remarks that humans often present their canine teeth when sneering in rage, and he suggests that this means that a human ancestor probably utilized their teeth in aggressive action. A domestic dog's simple tail wag may be used in subtly different ways to convey many meanings as illustrated in Darwin's The Expression of the Emotions in Man and Animals published in 1872.

Anecdotal approach

Evidence for emotions in animals has been primarily anecdotal, from individuals who interact with pets or captive animals on a regular basis. However, critics of animals having emotions often suggest that anthropomorphism is a motivating factor in the interpretation of the observed behaviours. Much of the debate is caused by the difficulty in defining emotions and the cognitive requirements thought necessary for animals to experience emotions in a similar way to humans. The problem is made more problematic by the difficulties in testing for emotions in animals. What is known about human emotion is almost all related or in relation to human communication.

Scientific approach

In recent years, the scientific community has become increasingly supportive of the idea of emotions in animals. Scientific research has provided insight into similarities of physiological changes between humans and animals when experiencing emotion.

Much support for animal emotion and its expression results from the notion that feeling emotions doesn't require significant cognitive processes, rather, they could be motivated by the processes to act in an adaptive way, as suggested by Darwin. Recent attempts in studying emotions in animals have led to new constructions in experimental and information gathering. Professor Marian Dawkins suggested that emotions could be studied on a functional or a mechanistic basis. Dawkins suggests that merely mechanistic or functional research will provide the answer on its own, but suggests that a mixture of the two would yield the most significant results.

Functional

Functional approaches rely on understanding what roles emotions play in humans and examining that role in animals. A widely used framework for viewing emotions in a functional context is that described by Oatley and Jenkins who see emotions as having three stages: (i) appraisal in which there is a conscious or unconscious evaluation of an event as relevant to a particular goal. An emotion is positive when that goal is advanced and negative when it is impeded (ii) action readiness where the emotion gives priority to one or a few kinds of action and may give urgency to one so that it can interrupt or compete with others and (iii) physiological changes, facial expression and then behavioural action. The structure, however, may be too broad and could be used to include all the animal kingdom as well as some plants.

Mechanistic

The second approach, mechanistic, requires an examination of the mechanisms that drive emotions and search for similarities in animals.

The mechanistic approach is utilized extensively by Paul, Harding and Mendl. Recognizing the difficulty in studying emotion in non-verbal animals, Paul et al. demonstrate possible ways to better examine this. Observing the mechanisms that function in human emotion expression, Paul et al. suggest that concentration on similar mechanisms in animals can provide clear insights into the animal experience. They noted that in humans, cognitive biases vary according to emotional state and suggested this as a possible starting point to examine animal emotion. They propose that researchers may be able to use controlled stimuli which have a particular meaning to trained animals to induce particular emotions in these animals and assess which types of basic emotions animals can experience.

Cognitive bias test

Is the glass half empty or half full?
 
A cognitive bias is a pattern of deviation in judgment, whereby inferences about other animals and situations may be drawn in an illogical fashion. Individuals create their own "subjective social reality" from their perception of the input. It refers to the question "Is the glass half empty or half full?", used as an indicator of optimism or pessimism. To test this in animals, an individual is trained to anticipate that stimulus A, e.g. a 20 Hz tone, precedes a positive event, e.g. highly desired food is delivered when a lever is pressed by the animal. The same individual is trained to anticipate that stimulus B, e.g. a 10 Hz tone, precedes a negative event, e.g. bland food is delivered when the animal presses a lever. The animal is then tested by being played an intermediate stimulus C, e.g. a 15 Hz tone, and observing whether the animal presses the lever associated with the positive or negative reward, thereby indicating whether the animal is in a positive or negative mood. This might be influenced by, for example, the type of housing the animal is kept in.

Using this approach, it has been found that rats which are subjected to either handling or playful, experimenter-administered manual stimulation (tickling) showed different responses to the intermediate stimulus: rats exposed to tickling were more optimistic. The authors stated that they had demonstrated "...for the first time a link between the directly measured positive affective state and decision making under uncertainty in an animal model." 

Cognitive biases have been shown in a wide range of species including rats, dogs, rhesus macaques, sheep, chicks, starlings and honeybees.

Self-medication with psychoactive drugs

Humans can suffer from a range of emotional or mood disorders such as depression, anxiety, fear and panic. To treat these disorders, scientists have developed a range of psychoactive drugs such as anxiolytics. Many of these drugs are developed and tested by using a range of laboratory species. It is inconsistent to argue that these drugs are effective in treating human emotions whilst denying the experience of these emotions in the laboratory animals on which they have been developed and tested.

Standard laboratory cages prevent mice from performing several natural behaviours for which they are highly motivated. As a consequence, laboratory mice sometimes develop abnormal behaviours indicative of emotional disorders such as depression and anxiety. To improve welfare, these cages are sometimes enriched with items such as nesting material, shelters and running wheels. Sherwin and Ollson tested whether such enrichment influenced the consumption of Midazolam, a drug widely used to treat anxiety in humans. Mice in standard cages, standard cages but with unpredictable husbandry, or enriched cages, were given a choice of drinking either non-drugged water or a solution of the Midazolam. Mice in the standard and unpredictable cages drank a greater proportion of the anxiolytic solution than mice from enriched cages, indicating that mice from the standard and unpredictable laboratory caging may have been experiencing greater anxiety than mice from the enriched cages. 

Spindle neurons

Spindle neurons are specialised cells found in three very restricted regions of the human brain – the anterior cingulate cortex, the frontoinsular cortex and the dorsolateral prefrontal cortex. The first two of these areas regulate emotional functions such as empathy, speech, intuition, rapid "gut reactions" and social organization in humans. Spindle neurons are also found in the brains of humpback whales, fin whales, killer whales, sperm whales, bottlenose dolphin, Risso's dolphin, beluga whales, and the African and Asian elephants.

Whales have spindle cells in greater numbers and are maintained for twice as long as humans. The exact function of spindle cells in whale brains is still not understood, but Hof and Van Der Gucht believe that they act as some sort of "high-speed connections that fast-track information to and from other parts of the cortex". They compared them to express trains that bypass unnecessary connections, enabling organisms to instantly process and act on emotional cues during complex social interactions. However, Hof and Van Der Gucht clarify that they do not know the nature of such feelings in these animals and that we cannot just apply what we see in great apes or ourselves to whales. They believe that more work is needed to know whether emotions are the same for humans and whales. 

Vocalizations

Though non-human animals cannot provide useful verbal feedback about the experiential and cognitive details of their feelings, various emotional vocalizations of other animals may be indicators of potential affective states. Beginning with Darwin and his research, it has been known that chimpanzees and other great apes perform laugh-like vocalizations, providing scientists with more symbolic self-reports of their emotional experiences.

Research with rats has revealed that under particular conditions, they emit 50-kHz ultrasonic vocalisations (USV) which have been postulated to reflect a positive affective state (emotion) analogous to primitive human joy; these calls have been termed "laughter". The 50 kHz USVs in rats are uniquely elevated by hedonic stimuli—such as tickling, rewarding electrical brain stimulation, amphetamine injections, mating, play, and aggression—and are suppressed by aversive stimuli.[6] Of all manipulations that elicit 50 kHz chirps in rats, tickling by humans elicits the highest rate of these calls.

Some vocalizations of domestic cats, such as purring, are well known to be produced in situations of positive valence, such as mother kitten interactions, contacts with familiar partner, or during tactile stimulation with inanimate objects as when rolling and rubbing. Therefore, purring can be generally considered as an indicator of "pleasure" in cats.

Low pitched bleating in sheep has been associated with some positive-valence situations, as they are produced by males as an estrus female is approaching or by lactating mothers while licking and nursing their lambs.

Neurological

Neuroscientific studies based off of the instinctual, emotional action tendencies of non-human animals accompanied by the brains neurochemical and electrical changes are deemed to best monitor relative primary process emotional/affective states. Predictions based off the research conducted on animals is what leads analysis of the neural infrastructure relevant in humans. Psycho-neuro-ethological triangulation with both humans and animals allows for further experimentation into animal emotions. Utilizing specific animals that exhibit indicators of emotional states to decode underlying neural systems aids in the discovery of critical brain variables that regulate animal emotional expressions. Comparing the results of the animals converse experiments occur predicting the affective changes that should result in humans. Specific studies where there is an increase or decrease of playfulness or separation distress vocalizations in animals, comparing humans that exhibit the predicted increases or decreases in feelings of joy or sadness, the weight of evidence constructs a concrete neural hypothesis concerning the nature of affect supporting all relevant species.

Criticism

The argument that animals experience emotions is sometimes rejected due to a lack of evidence, and those who do not believe in the idea of animal intelligence, often argue that anthropomorphism plays a role in individuals' perspectives. Those who reject that animals have the capacity to experience emotion do so mainly by referring to inconsistencies in studies that have endorsed the belief emotions exist. Having no linguistic means to communicate emotion beyond behavioral response interpretation, the difficulty of providing an account of emotion in animals relies heavily on interpretive experimentation, that relies on results from human subjects.

Some people oppose the concept of animal emotions and suggest that emotions are not universal, including in humans. If emotions are not universal, this indicates that there is not a phylogenetic relationship between human and non-human emotion. The relationship drawn by proponents of animal emotion, then, would be merely a suggestion of mechanistic features that promote adaptivity, but lack the complexity of human emotional constructs. Thus, a social life-style may play a role in the process of basic emotions developing into more complex emotions.

Darwin concluded, through a survey, that humans share universal emotive expressions and suggested that animals likely share in these to some degree. Social constructionists disregard the concept that emotions are universal. Others hold an intermediate stance, suggesting that basic emotional expressions and emotion are universal but the intricacies are developed culturally. A study by Elfenbein and Ambady indicated that individuals within a particular culture are better at recognising other cultural members' emotions.

Examples

Primates

Primates, in particular great apes, are candidates for being able to experience empathy and theory of mind. Great apes have complex social systems; young apes and their mothers have strong bonds of attachment and when a baby chimpanzee or gorilla dies, the mother will not uncommonly carry the body around for several days. Jane Goodall has described chimpanzees as exhibiting mournful behavior. Koko, a gorilla trained to use sign language, was reported to have expressed vocalizations indicating sadness after the death of her pet cat, All Ball.

Beyond such anecdotal evidence, support for empathetic reactions has come from experimental studies of rhesus macaques. Macaques refused to pull a chain that delivered food to themselves if doing so also caused a companion to receive an electric shock. This inhibition of hurting another conspecific was more pronounced between familiar than unfamiliar macaques, a finding similar to that of empathy in humans. 

Furthermore, there has been research on consolation behavior in chimpanzees. De Waal and Aureli found that third-party contacts attempt to relieve the distress of contact participants by consoling (e.g. making contact, embracing, grooming) recipients of aggression, especially those that have experienced more intense aggression. Researchers were unable to replicate these results using the same observation protocol in studies of monkeys, demonstrating a possible difference in empathy between monkeys and apes.

Other studies have examined emotional processing in the great apes. Specifically, chimpanzees were shown video clips of emotionally charged scenes, such as a detested veterinary procedure or a favorite food, and then were required to match these scenes with one of two species-specific facial expressions: "happy" (a play-face) or "sad" (a teeth-baring expression seen in frustration or after defeat). The chimpanzees correctly matched the clips to the facial expressions that shared their meaning, demonstrating that they understand the emotional significance of their facial expressions. Measures of peripheral skin temperature also indicated that the video clips emotionally affected the chimpanzees.

Rodents

In 1998, Jaak Panksepp proposed that all mammalian species are equipped with brains capable of generating emotional experiences. Subsequent work examined studies on rodents to provide foundational support for this claim. One of these studies examined whether rats would work to alleviate the distress of a conspecific. Rats were trained to press a lever to avoid the delivery of an electric shock, signaled by a visual cue, to a conspecific. They were then tested in a situation in which either a conspecific or a Styrofoam block was hoisted into the air and could be lowered by pressing a lever. Rats that had previous experience with conspecific distress demonstrated greater than ten-fold more responses to lower a distressed conspecific compared to rats in the control group, while those who had never experienced conspecific distress expressed greater than three-fold more responses to lower a distressed conspecific relative to the control group. This suggests that rats will actively work to reduce the distress of a conspecific, a phenomenon related to empathy. Comparable results have also been found in similar experiments designed for monkeys.

Langford et al. examined empathy in rodents using an approach based in neuroscience. They reported that (1) if two mice experienced pain together, they expressed greater levels of pain-related behavior than if pain was experienced individually, (2) if experiencing different levels of pain together, the behavior of each mouse was modulated by the level of pain experienced by its social partner, and (3) sensitivity to a noxious stimulus was experienced to the same degree by the mouse observing a conspecific in pain as it was by the mouse directly experiencing the painful stimulus. The authors suggest this responsiveness to the pain of others demonstrated by mice is indicative of emotional contagion, a phenomenon associated with empathy, which has also been reported in pigs. One behaviour associated with fear in rats is freezing. If female rats experience electric shocks to the feet and then witness another rat experiencing similar footshocks, they freeze more than females without any experience of the shocks. This suggests empathy in experienced rats witnessing another individual being shocked. Furthermore, the demonstrator's behaviour was changed by the behaviour of the witness; demonstrators froze more following footshocks if their witness froze more creating an empathy loop.

Several studies have also shown rodents can respond to a conditioned stimulus that has been associated with the distress of a conspecific, as if it were paired with the direct experience of an unconditioned stimulus. These studies suggest that rodents are capable of shared affect, a concept critical to empathy. 

Horses

Although not direct evidence that horses experience emotions, a 2016 study showed that domestic horses react differently to seeing photographs of positive (happy) or negative (angry) human facial expressions. When viewing angry faces, horses look more with their left eye which is associated with perceiving negative stimuli. Their heart rate also increases more quickly and they show more stress-related behaviours. One rider wrote, 'Experienced riders and trainers can learn to read the subtle moods of individual horses according to wisdom passed down from one horseman to the next, but also from years of trial-and-error. I suffered many bruised toes and nipped fingers before I could detect a curious swivel of the ears, irritated flick of the tail, or concerned crinkle above a long-lashed eye.' This suggests that horses have emotions and display them physically but is not concrete evidence.

Birds

Marc Bekoff reported accounts of animal behaviour which he believed was evidence of animals being able to experience emotions in his book The Emotional Lives of Animals. The following is an excerpt from his book: 


Bystander affiliation is believed to represent an expression of empathy in which the bystander tries to console a conflict victim and alleviate their distress. There is evidence for bystander affiliation in ravens (e.g. contact sitting, preening, or beak-to-beak or beak-to-body touching) and also for solicited bystander affiliation, in which there is post-conflict affiliation from the victim to the bystander. This indicates that ravens may be sensitive to the emotions of others, however, relationship value plays an important role in the prevalence and function of these post-conflict interactions.

The capacity of domestic hens to experience empathy has been studied. Mother hens show one of the essential underpinning attributes of empathy: the ability to be affected by, and share, the emotional state of their distressed chicks. However, evidence for empathy between familiar adult hens has not yet been found.

Dogs

A drawing by Konrad Lorenz showing facial expressions of a dog
 
Some research indicates that domestic dogs may experience negative emotions in a similar manner to humans, including the equivalent of certain chronic and acute psychological conditions. Much of this is from studies by Martin Seligman on the theory of learned helplessness as an extension of his interest in depression:
A dog that had earlier been repeatedly conditioned to associate an audible stimulus with inescapable electric shocks did not subsequently try to escape the electric shocks after the warning was presented, even though all the dog would have had to do is jump over a low divider within ten seconds. The dog didn't even try to avoid the "aversive stimulus"; it had previously "learned" that nothing it did would reduce the probability of it receiving a shock. A follow-up experiment involved three dogs affixed in harnesses, including one that received shocks of identical intensity and duration to the others, but the lever which would otherwise have allowed the dog a degree of control was left disconnected and didn't do anything. The first two dogs quickly recovered from the experience, but the third dog suffered chronic symptoms of clinical depression as a result of this perceived helplessness.
A further series of experiments showed that, similar to humans, under conditions of long-term intense psychological stress, around one third of dogs do not develop learned helplessness or long-term depression. Instead these animals somehow managed to find a way to handle the unpleasant situation in spite of their past experience. The corresponding characteristic in humans has been found to correlate highly with an explanatory style and optimistic attitude that views the situation as other than personal, pervasive, or permanent. 

Since these studies, symptoms analogous to clinical depression, neurosis, and other psychological conditions have also been accepted as being within the scope of emotion in domestic dogs. The postures of dogs may indicate their emotional state. In some instances, the recognition of specific postures and behaviors can be learned.

Psychology research has shown that when humans gaze at the face of another human, the gaze is not symmetrical; the gaze instinctively moves to the right side of the face to obtain information about their emotions and state. Research at the University of Lincoln shows that dogs share this instinct when meeting a human, and only when meeting a human (i.e. not other animals or other dogs). They are the only non-primate species known to share this instinct.

The existence and nature of personality traits in dogs have been studied (15,329 dogs of 164 different breeds). Five consistent and stable "narrow traits" were identified, described as playfulness, curiosity/fearlessness, chase-proneness, sociability and aggressiveness. A further higher order axis for shyness–boldness was also identified.

Dogs presented with images of either human or dog faces with different emotional states (happy/playful or angry/aggressive) paired with a single vocalization (voices or barks) from the same individual with either a positive or negative emotional state or brown noise. Dogs look longer at the face whose expression is congruent to the emotional state of the vocalization, for both other dogs and humans. This is an ability previously known only in humans. The behavior of a dog can not always be an indication of its friendliness. This is because when a dog wags its tail, most people interpret this as the dog expressing happiness and friendliness. Though indeed tail wagging can express these positive emotions, tail wagging is also an indication of fear, insecurity, challenging of dominance, establishing social relationships or a warning that the dog may bite.

Some researchers are beginning to investigate the question of whether dogs have emotions with the help of magnetic resonance imaging.

Elephants

Elephants are known for their empathy towards members of the same species as well as their cognitive memory. While this is true scientists continuously debate the extent to which elephants feel emotion. Observations show that elephants, like humans, are concerned with distressed or deceased individuals, and render assistance to the ailing and show a special interest in dead bodies of their own kind, however this view is interpreted as being anthropomorphic

Elephants have recently been suggested to pass mirror self-recognition tests, and such tests have been linked to the capacity for empathy. However, the experiment showing such actions did not follow the accepted protocol for tests of self-recognition, and earlier attempts to show mirror self-recognition in elephants have failed, so this remains a contentious claim. 

Elephants are also deemed to show emotion through vocal expression, specifically the rumble vocalization. Rumbles are frequency modulated, harmonically rich calls with fundamental frequencies in the infrasonic range, with clear formant structure. Elephants exhibit negative emotion and/or increased emotional intensity through their rumbles, based on specific periods of social interaction and agitation. 

Cats

Cat's response to a fear inducing stimulus.
 
It has been postulated that domestic cats can learn to manipulate their owners through vocalizations that are similar to the cries of human babies. Some cats learn to add a purr to the vocalization, which makes it less harmonious and more dissonant to humans, and therefore harder to ignore. Individual cats learn to make these vocalizations through trial-and-error; when a particular vocalization elicits a positive response from a human, the probability increases that the cat will use that vocalization in the future.

Growling can be an expression of annoyance or fear, similar to humans. When annoyed or angry, a cat wriggles and thumps its tail much more vigorously than when in a contented state. In larger felids such as lions, what appears to be irritating to them varies between individuals. A male lion may let his cubs play with his mane or tail, or he may hiss and hit them with his paws. Domestic male cats also have variable attitudes towards their family members, for example, older male siblings tend not to go near younger or new siblings and may even show hostility toward them. 

Hissing is also a vocalization associated with either offensive or defensive aggression. They are usually accompanied by a postural display intended to have a visual effect on the perceived threat. Cats hiss when they are startled, scared, angry, or in pain, and also to scare off intruders into their territory. If the hiss and growl warning does not remove the threat, an attack by the cat may follow. Kittens as young as two to three weeks will potentially hiss when first picked up by a human. 

Honeybees

Honeybees become pessimistic after being shaken
 
Honeybees ("Apis mellifera carnica") were trained to extend their proboscis to a two-component odour mixture (CS+) predicting a reward (e.g., 1.00 or 2.00 M sucrose) and to withhold their proboscis from another mixture (CS−) predicting either punishment or a less valuable reward (e.g., 0.01 M quinine solution or 0.3 M sucrose). Immediately after training, half of the honeybees were subjected to vigorous shaking for 60 s to simulate the state produced by a predatory attack on a concealed colony. This shaking reduced levels of octopamine, dopamine, and serotonin in the hemolymph of a separate group of honeybees at a time point corresponding to when the cognitive bias tests were performed. In honeybees, octopamine is the local neurotransmitter that functions during reward learning, whereas dopamine mediates the ability to learn to associate odours with quinine punishment. If flies are fed serotonin, they are more aggressive; flies depleted of serotonin still exhibit aggression, but they do so much less frequently.

Within 5 minutes of the shaking, all the trained bees began a sequence of unreinforced test trials with five odour stimuli presented in a random order for each bee: the CS+, the CS−, and three novel odours composed of ratios intermediate between the two learned mixtures. Shaken honeybees were more likely to withhold their mouthparts from the CS− and from the most similar novel odour. Therefore, agitated honeybees display an increased expectation of bad outcomes similar to a vertebrate-like emotional state. The researchers of the study stated that, "Although our results do not allow us to make any claims about the presence of negative subjective feelings in honeybees, they call into question how we identify emotions in any non-human animal. It is logically inconsistent to claim that the presence of pessimistic cognitive biases should be taken as confirmation that dogs or rats are anxious but to deny the same conclusion in the case of honeybees."

Crayfish

The freshwater crayfish Procambarus clarkii
 
Crayfish naturally explore new environments but display a general preference for dark places. A 2014 study on the freshwater crayfish Procambarus clarkii tested their responses in a fear paradigm, the elevated plus maze in which animals choose to walk on an elevated cross which offers both aversive and preferable conditions (in this case, two arms were lit and two were dark). Crayfish which experienced an electric shock displayed enhanced fearfulness or anxiety as demonstrated by their preference for the dark arms more than the light. Furthermore, shocked crayfish had relatively higher brain serotonin concentrations coupled with elevated blood glucose, which indicates a stress response. Moreover, the crayfish calmed down when they were injected with the benzodiazepine anxiolytic, chlordiazepoxide, used to treat anxiety in humans, and they entered the dark as normal. The authors of the study concluded "...stress-induced avoidance behavior in crayfish exhibits striking homologies with vertebrate anxiety."

A follow-up study using the same species confirmed the anxiolytic effect of chlordiazepoxide, but moreover, the intensity of the anxiety-like behaviour was dependent on the intensity of the electric shock until reaching a plateau. Such a quantitative relationship between stress and anxiety is also a very common feature of human and vertebrate anxiety.

Lie point symmetry

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