Animal Behavior/History

Early Influences

Prehistoric Period

Cave painting of a horse at Lascaux

Our early ancestors critically depended on a detailed understanding of animal behavior. Specifically, knowledge of habitat preference, movement patterns, and sensory biology of prey as well as predators were essential. Rock paintings, figurines, and carvings, depicting scenes from daily life and religious rituals, commonly feature detailed likenesses of many kinds of animals. Early forms of rock art emerged during the Paleolithic (200,000 BCE), however, the advent of modern man brought about extraordinary cave wall paintings (40,000 BCE) of animals, hunting scenes and other graphic illustrations of prehistoric life.

Holes in Neolithic skulls illustrate that patients dating back to such early times have been subjected to head surgery, presumably with the aim of curing headaches or mental disorders. Sumerian descriptions show the use of Opium, derived from poppy plants, to produce euphoriant effects.

Native American

A common theme contends that the world was populated by many peoples who were subsequently transformed into different kinds of animals. Thus all animals and humans share the presence of a soul. The close bond between them is shown in frequent rituals where the behavior of different animals is simulated. Imagery of animals is common. The Great Serpent Mound represents an effigy mound located on a plateau along Ohio Brush Creek in Adams County, Ohio. Conforming to the topography, the snake's head approaches a cliff point, the snake then winds back and forth for more than 200 meters and ends with a triple coiled tail.

Egyptian Mythology

Horus, an ancient Egyptian falcon headed-deity.

Ancient Egyptian societies showed a keen interest in the temperaments and sensory abilities of different animals and ascribed them to individual deities.

Ancient Egyptians had a highly developed view of the afterlife with elaborate rituals for preparing the body and soul for a peaceful existence after death. Beliefs about the soul and its needs led to numerous ceremonies for the preservation of the body. The brain apparently was not considered an essential element for eternity, as it was treated with a distinct lack of dignity. It was often spooned into small pieces and removed through the nose, then unceremoniously discarded.

Classical Antiquity

Ancient Greek Period

Aristotle
Aristotle, 340 BC
Born	384 BC Stageira, Chalcidice, Greece
Died	322 BC Euboea, Greece
Nationality	Greek
Fields	Philosophy, Psychology, Animal Behavior

Plato held the notion that the human mind as an entity pre-existed somewhere in the heavens, before being sent down to join a body on earth. Hippocrates believed that the brain was not only involved with sensation but was also the seat of intelligence. Like many before and after him, Aristotle was intrigued by the natural world around him. He, however, stands out as a seminal thinker on a wide range of subjects as he was able to move beyond his substantial abilities at observation by forwarding daring hypotheses in order to categorize and make sense of what he saw. Aristotle provided the first written records of mutualistic ties between individuals, animal tool use, and brood parasitism. His views on human nature considered our behavior as constituting formed habits and a desire for happiness. With his proposed concept of a tabula rasa he forwarded the thesis that individuals are born with no innate or built-in mental content, in a word "blank", and that their entire resource of knowledge is built up gradually from their experiences and sensory perceptions of the outside world. Herophilus studied the nervous system and distinguished between sensory nerves and motor nerves. Erasistratus studied the brain and distinguished between the cerebrum and cerebellum.

Roman Empire

As a physician in a gladiator school, Galen gained a unique view of the behavioral deficits that resulted from acute head trauma. Not surprisingly he wrote extensively on the connection between paralysis and severance of the spinal cord. His many audacious operations, including brain and eye surgery, were not again used for almost two millennia.

Middle Ages

Naturalists and philosophers focused on classical texts and, in particular, on the Bible in Latin. Accepted were Aristotle's views on all matters of science, including psychology. As knowledge of Greek declined, the West found itself cut off from its Greek philosophical and scientific roots. General levels of literacy further decreased as learning became largely restricted to monastic and cathedral schools that were primarily aimed at training clergy. The Church exerted an overpowering role in philosophical thought and nontheological scholarship was rare. Despite this many influences on the study of behavior in later periods began to emerge.

Arabists

Arabian scholars describe cranial and spinal nerves, along with surgical procedures for neurological disorders.

Renaissance

The principal features of the Renaissance (14th-16th century) were the revival of learning based on classical sources and the broad advancements of science. It pioneered scientific study through deductive reasoning. René Descartes supplied a foundation for the pursuit of knowledge.

• Unity of all sciences through deductive reasoning
• Never accept anything as true that is not known clearly to be such
• Divide problems into as many parts as possible, and proceed from the simplest to the hardest

He argues that he perceives his body through the use of the senses, which often proven unreliable. So he concludes that thinking is his essence as it is the only thing about him that cannot be doubted. Descartes defines "thought" as every activity of which a person is immediately conscious. In his Mind/body dualism he viewed the body as a machine that follows the laws of physics. In contrast, he regarded the mind (or soul) as a nonmaterial entity that is not subject to such physical constraints. The Cartesians (i.e., the followers of Descartes) aimed to explain behavior using simple, natural, and entirely mechanistic principles. In their view only humans were endowed with an ability to reason.

With the advent of the scientific revolution, medieval worldviews steadily gave way to increasingly analytical methodologies and a weakening of vitalist traditions. Although medical thought continued to be guided by the occult, critical thinking began to chip away at the long-held notions based on Galen. Hippocrates' views are revived and Renaissance artists such as Leonardo da Vinci conducted detailed studies of human anatomy in order to better portray the human form. As Andreas Vesalius corrects Greek medical errors, he revolutionizes medicine. William Harvey applies scientific methodology to human physiology and solves the human circulatory system. Paracelsus rejects occultism and pioneers the use of chemicals and minerals in medicine.

Enlightenment

Thinkers of the Age of Reason (18th century) perceive their efforts as bringing to an end previous periods of irrationality, superstition, and tyranny. John Locke's philosophy revives Aristiotles theory of a tabula rasa, where the (human) mind is at birth a "blank slate" without rules for processing data, and where data is added and rules for processing are formed solely by one's sensory experiences. The concept of a blank mind at birth emphasized the individual's freedom to author one's own human character. Moreover, as members of ones society will become what society make of them, it becomes essential to assure universal access to learning opportunities, medical care, and possibilities for personal development.

Evolution

In the second half of the 19th century the simple taxonomic classification of all life forms was supplanted by a growing focus on parsimonious and scientifically testable (phylegenetic) hypotheses that could explain the relatedness among groups of organisms. Studies of the fossil record, the geographic distribution of faunas, and the characteristics that linked the diversity of live, convinced a growing number of scientists that species indeed changed over time. Evolution became viewed as a branching process, whereby populations are altered over time and may specialize into separate branches, hybridize together, or terminate by extinction. However, the mechanism driving these changes remained unclear until the 1859 publication of Charles Darwin's On the Origin of Species, which provided the theory of evolution with the likely mechanism of natural selection.

• Charles Darwin (12 February 1809 – 19 April 1882) represents arguably among the most significant scientists of all time. He realized and presented compelling evidence for natural selection as the main mechanism by which all species of life have evolved over time from common ancestors. Evolution, as an explanation for the diversity of life, had emerged earlier and was generally accepted by most in his lifetime. His 1859 book On the Origin of Species established evolutionary descent with modification as the dominant scientific explanation of diversification in nature and this view emerged as the primary explanation in modern evolutionary theory. He also examined human evolution and sexual selection in The Descent of Man, and Selection in Relation to Sex, followed by The Expression of the Emotions in Man and Animals. Darwin suggested that all human characteristics are fundamentally inherited and all have been subject to evolution. Emotions may best be understood as consequences of adaptive reactions.
• George John Romanes, a Canadian-born English evolutionary biologist and physiologist, laid the foundation for comparative psychology, postulating a similarity of cognitive processes and mechanisms between humans and animals.

Psychophysics

Psychophysicists, such as Gustav Theodor Fechner and Wilhelm Wundt, aimed to relate matter to mind via a study of inner experiences or feelings towards defined physical stimuli. Psychophysicists explicitly rejected the concepts of vitalists, i.e. that the functions of a living organism are due to a vital principle distinct from physicochemical forces. The main focus lay on the way subjects perceive experimental stimuli that can be objectively measured, such as pure tones varying in intensity, or lights varying in luminance. A broad range of senses has been studied, including vision, hearing, touch (including skin and enteric perception), taste, smell, or the sense of time.

Psychoanalysis

Sigismund Schlomo Freud
Sigmund Freud, 1920
Born	May 6, 1856 Freiberg in Mähren, Moravia, now in the Czech Republic
Died	September 23, 1939 London, England, United Kingdom
Nationality	Austrian
Fields	Neurology, Philosophy, Psychiatry, Psychology, Psychotherapy, Psychoanalysis, Literature
Institutions	University of Vienna, Austria
Known for	Psychoanalysis

• Psychoanalysis is a family of psychological theories and methods based on the pioneering work of Sigmund Freud. As a result of talking with his patients, he came to believe that psychopathologies stemmed from culturally unacceptable, thus repressed and unconscious, desires and fantasies of a sexual nature. Moreover he stressed the presumed value of dreams as sources of insight into unconscious desires. The analyst's goal is thus aimed to discover connections among the unconscious components of patients' mental processes. This ought to help liberate the patient from unexamined or unconscious barriers of transference and resistance, that is, past patterns of relating that are no longer serviceable or that inhibit freedom.
• Carl G. Jung's unique and broadly influential approach to psychology emphasized understanding the psyche through exploring the worlds of dreams, art, mythology, world religion and philosophy. Jung emphasized the importance of balance and harmony. He cautioned that modern humans rely too heavily on science and logic and would benefit from integrating spirituality and appreciation of the unconscious realm. His Collective unconscious refers to a human experience reservoir, a kind of knowledge, we are all born with but can never be directly conscious of. His Archetypes represent generic, idealized models of personalities with unlearned tendencies to experience things in a certain way.

Comparative Psychology and Behaviorism

This school of thought represents a movement in psychology that advocates the use of strict experimental procedures to study observable behaviors (or responses) in relation to environment (or stimuli). Behaviorists regard learning and environmental conditions as dominant over the effects of genetics and heredity. Concepts, such as learning or memory are explored using artificial tasks in strictly controlled environments (e.g., Skinner Box, Morris' water maze, Radial arm maze).

Comparative psychology viewed science less as a framework for explaining events but rather for predicting and controlling them. Adoption of the observational techniques that had proven useful in the study of the behavior of animals was viewed as the best way to advance the scientific study of human beings. The external environment represented the main determining influence on behavior and its manipulation held the most promise for improving the condition of human beings. The concept of a tabula rasa was revived.

A common tenet held that the complex phenomena of behavior could be explained by and reduced to clear, distinct, reflexive units of behavior.

William James described psychology as "the science of mental life." As a rigid application of the scientific method is only possible for learned behaviors, he felt that our studies should foremost focus on studying those.

Ivan P. Pavlov realized that behavior changes when an organism comes to associate one stimulus with another. Specifically, a reflexive or automatic response transfers from one stimulus to another - Classical Conditioning.

Edward L. Thorndike Using a Puzzle Box, he demonstrated that behaviors that are followed by a positive outcome are often repeated, while those followed by a negative outcome or none at all are extinguished (i.e., Operant Conditioning). He postulated the Law of effect where behavior changes when the results of an action proved of consequence (e.g. rewarding, noxious).

John B. Watson stimulus-response theory of psychology, emotional reactions are learned in much the same way as other skills

B.F. Skinner contends that nearly all behavior is shaped by complex patterns of reinforcement in a person's environment. Skinner Box: An animal placed inside the box is rewarded with a small bit of food each time it makes the desired response, such as pressing a lever or pecking a key. A device outside the box records the animal's responses.

Ethology

"When, as by a miracle, the lovely butterfly bursts from the chrysalis full-winged and perfect, ... it has, for the most part, nothing to learn, because its little life flows from its organization like melody from a music box"—Douglas A. Spalding, 1873

Ethology, the biological study of behavior under naturalistic conditions, emerged out of diverse roots by individuals largely trained in a zoological tradition. It aimed to provide a more comprehensive study of animal behavior than previously available. Where comparative psychology studies the behavior of animals in the context of what is known about human psychology, ethology explores animal behavior in the context of animal anatomy and physiology. Ethologists specifically considered the significance of inherited elements of behavior alongside learned components. Influenced by evolutionary biologists such as Thomas Hunt Morgan and others, many aspects of behavior may indeed be accessible to selection. Charles Otis Whitman examined the importance of phylogeny to the origins of behavior. Oskar Heinroth rather focused on how the adaptive value of behavior may explain behavioral differences among taxa. Experiments, in which he raised young birds separate from other individuals of their own species, demonstrated that many species-specific behaviors were nonetheless present. Interest in genetically-programmed elements (instincts) were contrasted to those subject to learning (e.g., imprinting). Using a comparative approach ethologists are thus interested to search for the biological basis of behavior (including human behavior) by comparing specific activities to those of close relatives.

Karl von Frisch
Born	November 20, 1886
Died	June 12, 1982
Citizenship	Austrian
Fields	ethology
Known for	honey bee language
Notable awards	Nobel Prize in Physiology or Medicine in 1973

Karl von Frisch studied the senses of honey bees, identified their mechanisms of communication, and showed their sensitivity to ultraviolet and polarized light. He unlocked the basic secrets of the forager's waggle dance, research which continues to yield new insights to this day. Techniques developed by behaviorists, such as classical and operant conditioning, have proven valuable to ethologists. Animals trained to respond to a particular stimulus in the appropriate behavioral context can demonstrate their ability to discriminate between different types of stimulus. For example, von Frisch used classical conditioning to show that fish could hear, and bees were able to perceive colors.

Konrad Lorenz' work included studies of imprinting (i.e., the capacity to learn specific types of information at certain critical periods in development) and instinct (i.e., the capacity of an animal to complete a complex behavior in response to a first-time encounter with a given situation). Filial imprinting refers to the case where young of a species need to acquire the image of what their parents look like (e.g. ducks). Instinctive behavior can usually be elicited by a limited combination of complex environmental cues. Such a sign stimulus refers to a particular subset of features or qualities of a natural stimulus or from complex environmental cues that is sufficient to elicit a specific behavioral response. They noted that different individuals produce identical behavioral response to specific sign stimuli - Motor patterns (or Fixed Action Pattern, FAP). Once initiated, FAPs continue until completion. A releasing mechanism is a functionally organized, neural circuit that recognizes a specific releaser and produces the appropriate response. Due to their stereotype, behaviors can be cataloged in form of an Ethogram (i.e., a comprehensive compilation of the behavioral repertory of a species, a catalog of actions as complete and precise as possible). An internal factor that determines how likely an animal is to perform a behavior (i.e., Action-Specific Energy).

The causation of behavior is viewed in the context of a metaphor - the Psychohydraulic model. Action specific energy is akin to a liquid building up in a vessel. A high level of action specific energy results when a behavior has not been performed in some time. Pressure from the water pushes onto a piston which is also pulled by sign stimuli in the environment, represented by weights. As pressure mounts the piston finally opens a valve that allows water to flow out (representing the performance of the behavior) Example: An animal that has had a good meal will have a low drive to eat more (i.e., little water in the apparatus). As it goes for a while without eating, the drive (i.e., hunger) builds up. If food is then encountered, the animal will try to consume it. If no good food is encountered, less preferred items will be accepted. This model has faced much criticism as performance of many behaviors appear to increase the tendency to perform it again (i.e. aggression), rather than bring about a decrease.

Three Ethologists shared the 1973 Nobel Prize for Physiology or Medicine in recognition of their work over the previous 30 years as founders of the field of ethology.

• Karl von Frisch social behavior of bees
• Konrad Z. Lorenz studied the behavior of geese, ravens, and ducks
• Niko Tinbergen behavioral studies of gulls and stickle backs. "Man likes to consider himself a rationale being; he not only fails to recognize his instincts, he even scoffs at instincts and considers them inferior. This attitude makes him fall victim to similarity between man and animals; he recognizes himself only too often in an animal. Also, he has rather a higher opinion of instincts; for instance, he recognizes that they are at the bottom of his subjective experiences. And he has doubts about the all-controlling power of reason, for he sees its limits too often."

Behavioral Genetics

John Paul Scott
Born	December 17, 1909 Kansas City, KS, United States
Died	March 26, 2000, Bowling Green, OH, United States
Nationality	American
Institutions	Jackson Laboratory, Bar Harbor, ME and Bowling Green State University, Bowling Green, OH, USA
Known for	Genetics of Dog "Temperament"
Influences	Sewall Wright, WC Allee

Behavioral Genetics explores the hereditability of behavioral traits. More recently, this has given rise to a largely quantitative emphasis on using molecular genetic techniques in order to isolate individual genes that play a role in behavior, neural and behavioral pathologies, or individual human characteristics.

Each inherited feature of an organism is determined by a section of genetic code - a gene which represents the smallest hereditary unit. The genome refers to one individual's entire set of hereditary information and includes genes as well as non-coding sequences of DNA. The genomes for any two individuals, except for those of identical twins or cloned animals, are unique. Such differences (i.e., polymorphism) produce different versions of the same genes, called alleles. Despite these differences in genetic makeup, we also find major commonalities across much of the code shared by all forms of life. The specific genetic makeup of an individual is called its genotype. A trait refers to a specific characteristic or feature of an organism (although in psychology the term may be used to represent a component of personality). The term phenotype represents the specific manifestation of a trait or its physical appearance and constitution and refers to the actual state of that trait (e.g., the trait eye color has the phenotypes blue, brown and hazel). The individual's phenotype emerges as interaction of genetic information with environmental conditions.

Sir Francis Galton (February 16, 1822 – January 17, 1911), a younger relative of Charles Darwin, was an English explorer, geographer, and statistician. Intrigued by Darwin's concept of natural selection and with a keen interest in the breeding of domesticated animals, Galton explored variation in human populations and its implications. He became interested at first in the question of whether human ability was hereditary. He presented evidence for the inheritance of a range of abilities - "a man's natural abilities are derived by inheritance, under exactly the same limitations as are the form and physical features of the whole organic world". He also proposed adoption studies, including trans-racial adoption studies, to separate out the effects of heredity and environment. He was instrumental in the development of quantitative methods to test human intelligence, statistical correlation, fingerprint analysis. self-report questionnaires, and word-association studies. In a darker side, he became a vociferous proponent of eugenics where he proposed to improve the human race via selective breeding of those individuals who managed to score high marks on intelligence tests.

• John Paul Scott - genetics of behavior by studying personality traits in breeds of dogs

Behavioral Ecology

"Nothing in biology makes sense except in the light of evolution ..."—Theodosius Dobzhansky

Behavioral ecology is the study of the ecological and evolutionary basis for animal behavior. Primarily concerned with ultimate (evolutionary) consequences, researchers aim to identify phylogenetic contingencies within behavior (i.e., to what degree behavior is inherited from its ancestors) and to examine its functional significance (i.e., how the behavior impacts the bearers reproductive success). Behavioral ecologists contend that in order to minimize ad-hoc explanations arising from anecdotal observations a rigorous examination of the adaptive consequences of a behavior are essential. Specifically, it explores what a trait is good for in an evolutionary context.

Optimization theory is concerned with assessing the success of behavior. It attempts to identify those behavioral strategies that offer the highest return under a given set of conditions using a cost/benefit analysis. In this context success or fitness is judged by considering the number of offspring that the bearer of the behavior would contribute to the next generation relative to its suboptimal conspecifics. Optimization theory, such as optimal foraging theory, states that individuals ought to adopt strategies that allow it to consume the most energy while expending the least amount of energy. Evolutionary game theory asserts that the success of strategies should not only consider its inherent benefits, as the actual success depends on what other strategies exist in the population. An evolutionarily stable strategy is a strategy which when adopted by all members cannot be beaten out by any invading, mutant strategy.

• John R. Krebs - acoustic communication in birds and crickets, optimal foraging theory
• N. B. Davies - the economics of behavioral strategies

Sociobiology

Edward O Wilson
Edward O. Wilson, 2007
Born	June 10, 1929 Birmingham, AL, United States
Nationality	American
Institutions	Harvard University, USA
Known for	Sociobiology
Influences	Ronald Fisher
Notable awards	Pulitzer Prize (1979), Crafoord Prize (1990), Pulitzer Prize (1991), Nierenberg Prize (2001)

Sociobiology aims to explain behavior in all species by considering the evolutionary advantages of social behaviours. Sociobiology is based on the idea that all animals, including humans, will tend to act in ways to improve their own reproductive success, and that this will result in social processes conducive to genetic fitness. For example, cultural practices and values related to sexuality are viewed as shaped by natural selection (e.g., societal rules for incest avoidance). The concept has been criticized for its reliance on innate predispositions over learning, and by the view that presents situations are not representative of historical conditions.

The selfish genes hypothesis states that the individual is not as important to the population as its genes. Kin selection refers to changes in gene frequency across generations when biological relatives influence the reproductive success of each other. Thus, a gene that prompts behavior which enhances the fitness of relatives but lowers that of the individual displaying the behavior, may nonetheless increase in frequency, as relatives often share the same genes. The enhanced fitness of relatives can at times more than compensate for the fitness loss incurred by the individuals displaying the behavior.

• E.O. Wilson American entomologist, naturalist, and environmentalist.
• William D. Hamilton British evolutionary biologist, considered one of the greatest evolutionary theorists of the 20th century. Hamilton became famous for his theoretical work on kin selection - Hamilton's rule.

Evolutionary Psychology

Evolutionary Psychology attempts to explain "useful" mental and psychological traits—such as memory, perception, or language—as adaptations, i.e., as the functional products of natural selection.

Neuroscience

The last frontier of the biological sciences--their ultimate challenge--is to understand the biological basis of consciousness and the mental processes by which we perceive, act, learn, and remember.—Eric Kandel, 2000

Luigi Galvani
Luigi Galvani, Italian physician famous for pioneering bioelectricity
Born	September 9, 1737 Bologna, Papal States
Died	December 4, 1798 (aged 61) Bologna, Papal States
Institutions	University of Bologna
Known for	Bioelectricity

In 1791, Luigi Galvani discovered that the muscles of dead frog's legs twitched when struck by an electric spark. According to popular version of the story, Galvani had been conducting experiments with static electricity by rubbing frog skin. Galvani's assistant touched an exposed sciatic nerve of the skinned frog carcass with a metal scalpel that had picked up a charge, they saw sparks and the dead frog's leg kicked as if in life. The observation made Galvani appreciate the relationship between electricity and muscle movement. Galvani's cousin. who engaged in a public demonstration of electrical stimulation of muscles on the body of an executed criminal, reputedly provided the inspiration for Mary Shelley's Frankenstein. With stimulation of the body's face, the jaws began to quiver, facial muscles were horribly contorted, an eye opened, the right hand was raised and clenched, and muscles in legs and thighs moved.

The late 19th century witnessed the rise of modern neuroscience. Hypotheses that distinct brain functions were contained within specific regions of the brain emerged with evidence from patients with epileptic seizures, head trauma, and stroke. Extensive characterizations of neurons throughout the brain led Camillo Golgi and Santiago Ramón y Cajal to propose the neuron doctrine, i.e., that the function of the brain is a product of the electrical and chemical activity of individual neurons. Further support came from the discovery that muscles and neurons showed electrical excitability, which could be transferred onto neighboring cells. Electrical properties are assessed either by recording from an electrode placed into the lumen of a cell (intracellular) or from its surrounding vicinity (extracellular). Intracellular recording techniques aim to either measure the current needed to maintain a constant voltage (voltage clamp) or measure the voltage needed to keep the current constant (current clamp) across the cell membrane. Using such techniques, Alan Hodgkin and Andrew Huxley identified the ionic mechanisms that underly the electrical properties of neurons in giant axons of squid. John Carew Eccles identified synapses as the sites of communication between neurons.

Generations of neuroscientists had perceived the brain as a rather passive, reactive organ whose functional capacities were maintained by a sustained flow of sensory input.

Neuroethology and Behavioral Neuroscience

"It is an urgent task of ethologists and neurophysiologists to join efforts in the training of ‘etho-physiologists'"—Tinbergen, 1951. The Study of Instinct

Neuroethology is a synthesis of neuroscience and the study of behavior that explores neural mechanisms of natural behavior. This is in contrast to other approaches to neuroscience that study the nervous system in isolation, or in the context of artificial conditions. The neuroethological approach stems from the idea that nervous systems have evolved to address problems of sensing and acting in certain environmental conditions. The functions of nervous systems, therefore, are best understood in the context of the problems they have evolved to solve.

Using physiological experiments, Erich von Holst showed that behaviors can be elicited experimentally even in the absence of any sensory input. In contrast to Sherrington's reflex theory, van Holst postulated that basic central nervous function required permanent production of endogenous inhibitory stimulation. Behavior emerged when inhibitory control was diminished. He also demonstrated that internal, autonomous, rhythmic behavior can be independent of environmental factors. By slicing a worm into separate segments, and attaching each segment to a sensitive voltmeter, he noticed distinct, consecutive deflections on the meter which demonstrated as a wave moving through the severed parts from the front to the end of the entire cut-up specimen at approximately the speed of a contraction wave of a wriggling earthworm.

Eric Kandel
Eric Kandel, 2006
Born	November 7, 1929 Vienna, Austria
Fields	Neuroscience
Institutions	Columbia University College of Physicians and Surgeons
Known for	Physiology of memory
Notable awards	Nobel Prize in Physiology or Medicine 2000

Behavioral Neuroscience, a subdicipline of neuroscience, is largely concerned with the function of neural systems in generating behavior.

Mechanisms of sensory processing have utilized a variety of model systems, including the moth's acoustic startle response to bat sounds, jamming avoidance in electric fish, or sound localization in owls. A detailed understanding of the control and coordination of movement patters has emerged from investigations of food processing in lobsters, prey capture in frogs, or swimming in lampreys. Advances in our knowledge of the hormonal control of behavior have profited from work on the division of labor in bee colonies, behavioral changes due to molting in moths, and aggression in lizzards.

Sensory receptors and neurons exhibit functional tuning to a highly specific subset of stimulus conditions. For instance mechanoreceptors with sensory hairs detect air currents in crickets by transducing both direction and velocity of relevant stimulus energies. Directional tuning is determined by the preferred orientation of the hair's socket. Velocity tuning occurs when longer hairs respond to low wind velocities while shorter hairs need higher wind velocity to activate them. Each mechanoreceptor is innervated by a single sensory neuron and its directional tuning curve represents the cell's sensitivity to wind direction. Most substrates for sensory processing show a strict spatial organization - its topographic mapping. A system for extracting temporal information can be constructed via delay lines, a system which utilizes the fact that electrical conduction in neurons is not instantaneous; it is delayed depending on the length of the axon and other properties of the neuron (e.g. myelination). The resulting delays can be used for time sensitive calculations, such as calculation of inter-aural time differences in sound localization. A central pattern generator (CPG) is a network of neurons (or even a single neuron) which is able to exhibit rhythmic activity in the absence of sensory input.

• Erich von Holst demonstrated that behaviors can be elicited experimentally even in the absence of any sensory input.
• Eric Kandel observed synaptic changes in the Aplysia siphon-withdrawal reflex as a function of Hebbian learning.
• Walter Heiligenberg developed one of the best neurologically understood behavioral patterns in vertebrate, the jamming avoidance response in the weakly electric fish Eigenmannia.

Modern Psychology and Linguistics

Incidental Learning: Children readily imitated behavior exhibited by other children and adults who are significant to them, especially when such behavior is reinforced. A primary rationale for Special Education Inclusion holds that children with special needs will model their behavior after that of nondisabled peers and thereby improve communication, motor, social and play skills.

• Albert Bandura

Language Learning: Humans have an inborn capacity to extract word meanings, sentence structure, and grammatical rules from the complex stream of sounds they hear.

• Noam Chomski

Human Universals: Cultural universals are elements, patterns, traits, or institutions that are common to all human cultures worldwide.

• Steven Pinker

TED talks and more

• Steven Pinker: Human nature and the blank slate (Feb 2003)

<<Return to Contents Page | <<Chapter 1