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Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system. It is an interdisciplinary science that combines both
neuroscience Neuroscience is the science, scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a Multidisciplinary approach, multidisciplinary science that combines physiology, an ...
(study of the nervous system) and
ethology Ethology is the scientific study of animal behaviour, usually with a focus on behaviour under natural conditions, and viewing behaviour as an evolutionarily adaptive trait. Behaviourism as a term also describes the scientific and objecti ...
(study of animal behavior in natural conditions). A central theme of neuroethology, which differentiates it from other branches of neuroscience, is its focus on behaviors that have been favored by
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
(e.g., finding mates, navigation, locomotion, and predator avoidance) rather than on behaviors that are specific to a particular disease state or laboratory experiment. Neuroethologists hope to uncover general principles of the nervous system from the study of animals with exaggerated or specialized behaviors. They endeavor to understand how the nervous system translates biologically relevant stimuli into natural behavior. For example, many bats are capable of echolocation which is used for prey capture and navigation. The auditory system of bats is often cited as an example for how acoustic properties of sounds can be converted into a sensory map of behaviorally relevant features of sounds.


Philosophy

Neuroethology is an integrative approach to the study of animal behavior that draws upon several disciplines. Its approach stems from the
theory A theory is a rational type of abstract thinking about a phenomenon, or the results of such thinking. The process of contemplative and rational thinking is often associated with such processes as observational study or research. Theories may ...
that animals' nervous systems have evolved to address problems of sensing and acting in certain environmental niches and that their nervous systems are best understood in the context of the problems they have evolved to solve. In accordance with Krogh's principle, neuroethologists often study animals that are "specialists" in the behavior the researcher wishes to study e.g. honeybees and social behavior, bat echolocation, owl sound localization, etc. The scope of neuroethological inquiry might be summarized by Jörg-Peter Ewert, a pioneer of neuroethology, when he considers the types of questions central to neuroethology in his 1980 introductory text to the field: # How are stimuli detected by an organism? # How are environmental stimuli in the external world represented in the nervous system? # How is information about a stimulus acquired, stored and recalled by the nervous system? # How is a behavioral pattern encoded by neural networks? # How is behavior coordinated and controlled by the nervous system? # How can the ontogenetic development of behavior be related to neural mechanisms? Often central to addressing questions in neuroethology are comparative methodologies, drawing upon knowledge about related organisms' nervous systems, anatomies, life histories, behaviors and environmental niches. While it is not unusual for many types of neurobiology experiments to give rise to behavioral questions, many neuroethologists often begin their research programs by observing a species' behavior in its natural environment. Other approaches to understanding nervous systems include the systems identification approach, popular in
engineering Engineering is the use of scientific principles to design and build machines, structures, and other items, including bridges, tunnels, roads, vehicles, and buildings. The discipline of engineering encompasses a broad range of more speciali ...
. The idea is to stimulate the system using a non-natural stimulus with certain properties. The system's response to the stimulus may be used to analyze the operation of the system. Such an approach is useful for
linear Linearity is the property of a mathematical relationship ('' function'') that can be graphically represented as a straight line. Linearity is closely related to '' proportionality''. Examples in physics include rectilinear motion, the linear ...
systems, but the nervous system is notoriously
nonlinear In mathematics and science, a nonlinear system is a system in which the change of the output is not proportional to the change of the input. Nonlinear problems are of interest to engineers, biologists, physicists, mathematicians, and many oth ...
, and neuroethologists argue that such an approach is limited. This argument is supported by experiments in the auditory system, which show that neural responses to complex sounds, like social calls, can not be predicted by the knowledge gained from studying the responses due to pure tones (one of the non-natural stimuli favored by auditory neurophysiologists). This is because of the non-linearity of the system. Modern neuroethology is largely influenced by the research techniques used. Neural approaches are necessarily very diverse, as is evident through the variety of questions asked, measuring techniques used, relationships explored, and model systems employed. Techniques utilized since 1984 include the use of intracellular dyes, which make maps of identified neurons possible, and the use of brain slices, which bring vertebrate brains into better observation through intracellular electrodes (Hoyle 1984). Currently, other fields toward which neuroethology may be headed include
computational neuroscience Computational neuroscience (also known as theoretical neuroscience or mathematical neuroscience) is a branch of neuroscience which employs mathematical models, computer simulations, theoretical analysis and abstractions of the brain to u ...
,
molecular genetics Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
, neuroendocrinology and
epigenetics In biology, epigenetics is the study of stable phenotypic changes (known as ''marks'') that do not involve alterations in the DNA sequence. The Greek prefix '' epi-'' ( "over, outside of, around") in ''epigenetics'' implies features that are ...
. The existing field of neural modeling may also expand into neuroethological terrain, due to its practical uses in
robotics Robotics is an interdisciplinary branch of computer science and engineering. Robotics involves design, construction, operation, and use of robots. The goal of robotics is to design machines that can help and assist humans. Robotics integrat ...
. In all this, neuroethologists must use the right level of simplicity to effectively guide research towards accomplishing the goals of neuroethology. Critics of neuroethology might consider it a branch of neuroscience concerned with 'animal trivia'. Though neuroethological subjects tend not to be traditional neurobiological model systems (i.e. ''
Drosophila ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many speci ...
'', ''
C. elegans ''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek ''caeno-'' (recent), ''rhabditis'' (r ...
'', or ''
Danio rerio The zebrafish (''Danio rerio'') is a freshwater fish belonging to the minnow family (Cyprinidae) of the order Cypriniformes. Native to South Asia, it is a popular aquarium fish, frequently sold under the trade name zebra danio (and thus often ca ...
''), neuroethological approaches emphasizing comparative methods have uncovered many concepts central to neuroscience as a whole, such as lateral inhibition, coincidence detection, and sensory maps. The discipline of neuroethology has also discovered and explained the only vertebrate behavior for which the entire neural circuit has been described: the
electric fish An electric fish is any fish that can generate electric fields. Most electric fish are also electroreceptive, meaning that they can sense electric fields. The only exception is the stargazer family. Electric fish, although a small minority, inc ...
jamming avoidance response. Beyond its conceptual contributions, neuroethology makes indirect contributions to advancing human health. By understanding simpler nervous systems, many clinicians have used concepts uncovered by neuroethology and other branches of
neuroscience Neuroscience is the science, scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a Multidisciplinary approach, multidisciplinary science that combines physiology, an ...
to develop treatments for devastating human diseases.


History

Neuroethology owes part of its existence to the establishment of ethology as a unique discipline within
zoology Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, an ...
. Although animal behavior had been studied since the time of
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
(384–342 BC), it was not until the early twentieth century that ethology finally became distinguished from natural science (a strictly descriptive field) and ecology. The main catalysts behind this new distinction were the research and writings of
Konrad Lorenz Konrad Zacharias Lorenz (; 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 regarde ...
and Niko Tinbergen. Konrad Lorenz was born in Austria in 1903, and is widely known for his contribution of the theory of
fixed action pattern A fixed action pattern is an ethological term describing an instinctive behavioral sequence that is highly stereotyped and species-characteristic. Fixed action patterns are said to be produced by the innate releasing mechanism, a "hard-wired" neura ...
s (FAPs): endogenous, instinctive behaviors involving a complex sequence of movements that are triggered ("released") by a certain kind of stimulus. This sequence always proceeds to completion, even if the original stimulus is removed. It is also species-specific and performed by nearly all members. Lorenz constructed his famous "hydraulic model" to help illustrate this concept, as well as the concept of action specific energy, or drives. Niko Tinbergen was born in the Netherlands in 1907 and worked closely with Lorenz in the development of the FAP theory; their studies focused on the egg retrieval response of nesting geese. Tinbergen performed extensive research on the releasing mechanisms of particular FAPs, and used the bill-pecking behavior of baby herring gulls as his model system. This led to the concept of the
supernormal stimulus A supernormal stimulus or superstimulus is an exaggerated version of a stimulus to which there is an existing response tendency, or any stimulus that elicits a response more strongly than the stimulus for which it evolved. For example, it is p ...
. Tinbergen is also well known for his
four questions Ma Nishtana ( he, מה נשתנה) are the first two words in a phrase meaning "Why is tonight different from all other nights?" The phrase appears at the beginning of each line of The Four Questions, traditionally asked via song by the youngest ...
that he believed ethologists should be asking about any given animal behavior; among these is that of the mechanism of the behavior, on a physiological, neural and
molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bio ...
level, and this question can be thought of in many regards as the keystone question in neuroethology. Tinbergen also emphasized the need for ethologists and neurophysiologists to work together in their studies, a unity that has become a reality in the field of neuroethology. Unlike
behaviorism Behaviorism is a systematic approach to understanding the behavior of humans and animals. It assumes that behavior is either a reflex evoked by the pairing of certain antecedent stimuli in the environment, or a consequence of that individual ...
, which studies animals' reactions to non-natural stimuli in artificial,
laboratory A laboratory (; ; colloquially lab) is a facility that provides controlled conditions in which scientific or technological research, experiments, and measurement may be performed. Laboratory services are provided in a variety of settings: physi ...
conditions, ethology sought to categorize and analyze the natural behaviors of animals in a field setting. Similarly, neuroethology asks questions about the neural bases of ''naturally occurring'' behaviors, and seeks to mimic the natural context as much as possible in the laboratory. Although the development of ethology as a distinct discipline was crucial to the advent of neuroethology, equally important was the development of a more comprehensive understanding of
neuroscience Neuroscience is the science, scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a Multidisciplinary approach, multidisciplinary science that combines physiology, an ...
. Contributors to this new understanding were the Spanish Neuroanatomist, Ramon y Cajal (born in 1852), and physiologists Charles Sherrington, Edgar Adrian,
Alan Hodgkin Sir Alan Lloyd Hodgkin (5 February 1914 – 20 December 1998) was an English physiologist and biophysicist who shared the 1963 Nobel Prize in Physiology or Medicine with Andrew Huxley and John Eccles. Early life and education Hodgkin was bo ...
, and
Andrew Huxley Sir Andrew Fielding Huxley (22 November 191730 May 2012) was an English physiologist and biophysicist. He was born into the prominent Huxley family. After leaving Westminster School in central London, he went to Trinity College, Cambridge ...
. Charles Sherrington, who was born in Great Britain in 1857, is famous for his work on the nerve synapse as the site of transmission of nerve impulses, and for his work on reflexes in the spinal cord. His research also led him to hypothesize that every muscular activation is coupled to an inhibition of the opposing muscle. He was awarded a Nobel Prize for his work in 1932 along with Lord Edgar Adrian who made the first physiological recordings of neural activity from single nerve fibers. Alan Hodgkin and Andrew Huxley (born 1914 and 1917, respectively, in Great Britain), are known for their collaborative effort to understand the production of action potentials in the giant axons of squid. The pair also proposed the existence of ion channels to facilitate action potential initiation, and were awarded the Nobel Prize in 1963 for their efforts. As a result of this pioneering research, many scientists then sought to connect the physiological aspects of the nervous and sensory systems to specific behaviors. These scientists – Karl von Frisch,
Erich von Holst Erich Walther von Holst (28 November 1908 – 26 May 1962) was a German behavioral physiologist who was a Baltic German native of Riga, Livonia and was related to historian Hermann Eduard von Holst (1841–1904). In the 1950s he found ...
, and Theodore Bullock – are frequently referred to as the "fathers" of neuroethology. Neuroethology did not really come into its own, though, until the 1970s and 1980s, when new, sophisticated experimental methods allowed researchers such as
Masakazu Konishi was a Japanese neurobiologist, known for his research on the neuroscience underlying the behavior of owls and songbirds. Early life and education Konishi was born on 17 February 1933 in Kyoto, Japan, the only child of poor "Nishijin" weavers. ...
, Walter Heiligenberg, Jörg-Peter Ewert, and others to study the neural circuits underlying verifiable behavior.


Modern neuroethology

The International Society for Neuroethology represents the present discipline of neuroethology, which was founded on the occasion of the NATO-Advanced Study Institute "Advances in Vertebrate Neuroethology" (August 13–24, 1981) organized by J.-P. Ewert, D.J. Ingle and R.R. Capranica, held at the University of Kassel in Hofgeismar, Germany (cf. report Trends in Neurosci. 5:141-143,1982). Its first president was Theodore H. Bullock. The society has met every three years since its first meeting in Tokyo in 1986. Its membership draws from many research programs around the world; many of its members are students and faculty members from medical schools and neurobiology departments from various universities. Modern advances in
neurophysiology Neurophysiology is a branch of physiology and neuroscience that studies nervous system function rather than nervous system architecture. This area aids in the diagnosis and monitoring of neurological diseases. Historically, it has been dominated ...
techniques have enabled more exacting approaches in an ever-increasing number of animal systems, as size limitations are being dramatically overcome. Survey of the most recent (2007) congress of the ISN meeting symposia topics gives some idea of the field's breadth: *Comparative aspects of spatial memory (rodents, birds, humans, bats) *Influences of higher processing centers in active sensing (primates, owls, electric fish, rodents, frogs) *Animal signaling plasticity over many time scales (electric fish, frogs, birds) *Song production and learning in passerine birds *Primate sociality *Optimal function of sensory systems (flies, moths, frogs, fish) *Neuronal complexity in behavior (insects, computational) *Contributions of genes to behavior (Drosophila, honeybees, zebrafish) *Eye and head movement (crustaceans, humans, robots) *Hormonal actions in brain and behavior (rodents, primates, fish, frogs, and birds) *Cognition in insects (honeybee)


Application to technology

Neuroethology can help create advancements in
technology Technology is the application of knowledge to reach practical goals in a specifiable and reproducible way. The word ''technology'' may also mean the product of such an endeavor. The use of technology is widely prevalent in medicine, scien ...
through an advanced understanding of animal behavior. Model systems were generalized from the study of simple and related animals to humans. For example, the neuronal cortical space map discovered in bats, a specialized champion of hearing and navigating, elucidated the concept of a computational space map. In addition, the discovery of the space map in the barn owl led to the first neuronal example of the Jeffress model. This understanding is translatable to understanding spatial localization in humans, a mammalian relative of the bat. Today, knowledge learned from neuroethology are being applied in new technologies. For example, Randall Beer and his colleagues used algorithms learned from insect walking behavior to create robots designed to walk on uneven surfaces (Beer et al.). Neuroethology and technology contribute to one another bidirectionally. Neuroethologists seek to understand the neural basis of a behavior as it would occur in an animal's natural environment but the techniques for neurophysiological analysis are lab-based, and cannot be performed in the field setting. This dichotomy between field and lab studies poses a challenge for neuroethology. From the neurophysiology perspective, experiments must be designed for controls and objective rigor, which contrasts with the ethology perspective – that the experiment be applicable to the animal's natural condition, which is uncontrolled, or subject to the dynamics of the environment. An early example of this is when Walter Rudolf Hess developed focal brain stimulation technique to examine a cat's brain controls of vegetative functions in addition to other behaviors. Even though this was a breakthrough in technological abilities and technique, it was not used by many neuroethologists originally because it compromised a cat's natural state, and, therefore, in their minds, devalued the experiments' relevance to real situations. When intellectual obstacles like this were overcome, it led to a golden age of neuroethology, by focusing on simple and robust forms of behavior, and by applying modern neurobiological methods to explore the entire chain of sensory and neural mechanisms underlying these behaviors (Zupanc 2004). New technology allows neuroethologists to attach electrodes to even very sensitive parts of an animal such as its brain while it interacts with its environment. The founders of neuroethology ushered this understanding and incorporated technology and creative experimental design. Since then even indirect technological advancements such as battery-powered and waterproofed instruments have allowed neuroethologists to mimic natural conditions in the lab while they study behaviors objectively. In addition, the electronics required for amplifying neural signals and for transmitting them over a certain distance have enabled neuroscientists to record from behaving animals performing activities in naturalistic environments. Emerging technologies can complement neuroethology, augmenting the feasibility of this valuable perspective of natural neurophysiology. Another challenge, and perhaps part of the beauty of neuroethology, is experimental design. The value of neuroethological criteria speak to the reliability of these experiments, because these discoveries represent behavior in the environments in which they evolved. Neuroethologists foresee future advancements through using new technologies and techniques, such as computational neuroscience, neuroendocrinology, and molecular genetics that mimic natural environments.


Case studies


Jamming avoidance response

In 1963, Akira Watanabe and Kimihisa Takeda discovered the behavior of the jamming avoidance response in the knifefish ''Eigenmannia'' sp. In collaboration with T.H. Bullock and colleagues, the behavior was further developed. Finally, the work of W. Heiligenberg expanded it into a full neuroethology study by examining the series of neural connections that led to the behavior. ''Eigenmannia'' is a weakly electric fish that can generate electric discharges through electrocytes in its tail. Furthermore, it has the ability to electrolocate by analyzing the perturbations in its electric field. However, when the frequency of a neighboring fish's current is very close (less than 20 Hz difference) to that of its own, the fish will avoid having their signals interfere through a behavior known as Jamming Avoidance Response. If the neighbor's frequency is higher than the fish's discharge frequency, the fish will lower its frequency, and vice versa. The sign of the frequency difference is determined by analyzing the "beat" pattern of the incoming interference which consists of the combination of the two fish's discharge patterns. Neuroethologists performed several experiments under ''Eigenmannia''s natural conditions to study how it determined the sign of the frequency difference. They manipulated the fish's discharge by injecting it with curare which prevented its natural electric organ from discharging. Then, an electrode was placed in its mouth and another was placed at the tip of its tail. Likewise, the neighboring fish's electric field was mimicked using another set of electrodes. This experiment allowed neuroethologists to manipulate different discharge frequencies and observe the fish's behavior. From the results, they were able to conclude that the electric field frequency, rather than an internal frequency measure, was used as a reference. This experiment is significant in that not only does it reveal a crucial neural mechanism underlying the behavior but also demonstrates the value neuroethologists place on studying animals in their natural habitats.


Feature analysis in toad vision

The recognition of prey and predators in the toad was first studied in depth by Jörg-Peter Ewert (Ewert 1974; see also 2004). He began by observing the natural prey-catching behavior of the common toad (''Bufo bufo'') and concluded that the animal followed a sequence that consisted of stalking, binocular fixation, snapping, swallowing and mouth-wiping. However, initially, the toad's actions were dependent on specific features of the sensory stimulus: whether it demonstrated worm or anti-worm configurations. It was observed that the worm configuration, which signaled prey, was initiated by movement along the object's long axis, whereas anti-worm configuration, which signaled predator, was due to movement along the short axis. (Zupanc 2004). Ewert and coworkers adopted a variety of methods to study the predator versus prey behavior response. They conducted recording experiments where they inserted electrodes into the brain, while the toad was presented with worm or anti-worm stimuli. This technique was repeated at different levels of the visual system and also allowed feature detectors to be identified. In focus was the discovery of prey-selective neurons in the optic tectum, whose axons could be traced towards the snapping pattern generating cells in the hypoglossal nucleus. The discharge patterns of prey-selective tectal neurons in response to prey objects – in freely moving toads – "predicted" prey-catching reactions such as snapping. Another approach, called stimulation experiment, was carried out in freely moving toads. Focal electrical stimuli were applied to different regions of the brain, and the toad's response was observed. When the thalamic-pretectal region was stimulated, the toad exhibited escape responses, but when the tectum was stimulated in an area close to prey-selective neurons, the toad engaged in prey catching behavior (Carew 2000). Furthermore, neuroanatomical experiments were carried out where the toad's thalamic-pretectal/tectal connection was lesioned and the resulting deficit noted: the prey-selective properties were abolished both in the responses of prey-selective neurons and in the prey catching behavior. These and other experiments suggest that prey selectivity results from pretecto-tectal influences. Ewert and coworkers showed in toads that there are stimulus-response mediating pathways that translate perception (of visual sign stimuli) into action (adequate behavioral responses). In addition there are modulatory loops that initiate, modify or specify this mediation (Ewert 2004). Regarding the latter, for example, the telencephalic caudal ventral striatum is involved in a loop gating the stimulus-response mediation in a manner of directed attention. The telencephalic ventral medial pallium („primordium hippocampi"), however, is involved in loops that either modify prey-selection due to associative learning or specify prey-selection due to non-associative learning, respectively.


Computational neuroethology

Computational neuroethology (CN or CNE) is concerned with the computer modelling of the neural mechanisms underlying animal behaviors. Together with the term "artificial ethology," the term "computational neuroethology" was first published in literature by Achacoso and Yamamoto in the Spring of 1990, based on their pioneering work on the connectome of C. elegans in 1989, with further publications in 1992. Computational neuroethology was argued for in depth later in 1990 by Randall Beer and by
Dave Cliff Dave Cliff (born 25 June 1944) is a British jazz guitarist. Career Cliff was born in Hexham, Northumberland. In 1967, he moved to Leeds and gained a diploma in jazz studies from Leeds College of Music while studying with bassist Peter Ind an ...
both of whom acknowledged the strong influence of Michael Arbib's ''Rana Computatrix'' computational model of neural mechanisms for visual guidance in frogs and toads. CNE systems work within a closed-loop environment; that is, they perceive their (perhaps artificial) environment directly, rather than through human input, as is typical in AI systems. For example, Barlow et al. developed a time-dependent model for the retina of the horseshoe crab ''Limulus polyphemus'' on a Connection Machine (Model CM-2). Instead of feeding the model retina with idealized input signals, they exposed the simulation to digitized video sequences made underwater, and compared its response with those of real animals.


Model systems

* Bat echolocation – nocturnal flight navigation and prey capture; location of objects using echo returns of its own call * Oscine
bird song Bird vocalization includes both bird calls and bird songs. In non-technical use, bird songs are the bird sounds that are melodious to the human ear. In ornithology and birding, songs (relatively complex vocalizations) are distinguished by fun ...
– zebra finch (''
Taeniopygia guttata The Sunda zebra finch (''Taeniopygia guttata'') is a species of bird in the family Estrildidae. It is found in the Lesser Sundas The Lesser Sunda Islands or nowadays known as Nusa Tenggara Islands ( id, Kepulauan Nusa Tenggara, formerly ) ...
''), canary ('' Serinus canaria'') and white-crowned sparrow ('' Zonotrichia leucophrys''); song learning as a model for human speech development *
Electric fish An electric fish is any fish that can generate electric fields. Most electric fish are also electroreceptive, meaning that they can sense electric fields. The only exception is the stargazer family. Electric fish, although a small minority, inc ...
– navigation, communication, Jamming Avoidance Response (JAR),
corollary discharge In physiology, an efference copy or efferent copy is an internal copy of an outflowing ('' efferent''), movement-producing signal generated by an organism's motor system.Jeannerod, Marc (2003): "Action Monitoring and Forward Control of Movements ...
, expectation generators, and spike timing dependent plasticity *
Barn owl The barn owl (''Tyto alba'') is the most widely distributed species of owl in the world and one of the most widespread of all species of birds, being found almost everywhere except for the polar and desert regions, Asia north of the Himala ...
auditory spatial map – nocturnal prey location and capture * Toad vision – discrimination of prey versus predator �
Video
"Image processing in the toad's visual system: behavior, brain function

*
Circadian rhythm A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., endogenous) and responds to ...
– influence of various circadian controlled behaviors by the
suprachiasmatic nucleus The suprachiasmatic nucleus or nuclei (SCN) is a tiny region of the brain in the hypothalamus, situated directly above the optic chiasm. It is responsible for controlling circadian rhythms. The neuronal and hormonal activities it generates regu ...
* Cricket song – mate attraction and corollary discharge * Fish
Mauthner cells The Mauthner cells are a pair of big and easily identifiable neurons (one for each half of the body) located in the rhombomere 4 of the hindbrain in fish and amphibians that are responsible for a very fast escape reflex (in the majority of animals � ...
– C-start escape response and underwater directional hearing * Fly – Microscale directional hearing in '' Ormia ochracea'', sex differences of the visual system in
Bibionidae Bibionidae (March flies) is a family of flies (Diptera) containing approximately 650–700 species worldwide. Adults are nectar feeders and emerge in numbers in spring. Because of the likelihood of adults flies being found ''in copula'', they h ...
, and spatial navigation in chasing behavior of Fannia canicularis *
Noctuid moth The Noctuidae, commonly known as owlet moths, cutworms or armyworms, are a family of moths. They are considered the most controversial family in the superfamily Noctuoidea because many of the clades are constantly changing, along with the other ...
s – ultrasound avoidance response to bat calls * '' Aplysia'' sea hares – learning and memory in startle response * Rat –
spatial memory In cognitive psychology and neuroscience, spatial memory is a form of memory responsible for the recording and recovery of information needed to plan a course to a location and to recall the location of an object or the occurrence of an event. Sp ...
and navigation * Salmon homing – olfactory imprinting and
thyroid hormone File:Thyroid_system.svg, upright=1.5, The thyroid system of the thyroid hormones T3 and T4 rect 376 268 820 433 Thyroid-stimulating hormone rect 411 200 849 266 Thyrotropin-releasing hormone rect 297 168 502 200 Hypothalamus rect 66 216 386 25 ...
s *
Crayfish Crayfish are freshwater crustaceans belonging to the clade Astacidea, which also contains lobsters. In some locations, they are also known as crawfish, craydids, crawdaddies, crawdads, freshwater lobsters, mountain lobsters, rock lobsters, m ...
– escape and startle behaviors, aggression and formation of social hierarchies *
Cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted th ...
fish – aggression and attack behaviors *
Honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...
– learning, navigation, vision, olfaction, flight, aggression, foraging *
Monarch butterfly The monarch butterfly or simply monarch (''Danaus plexippus'') is a milkweed butterfly (subfamily Danainae) in the family Nymphalidae. Other common names, depending on region, include milkweed, common tiger, wanderer, and black-veined brown. ...
– navigational mechanisms
More Model Systems and Information


See also

* Niko Tinbergen * Karl von Frisch *
Konrad Lorenz Konrad Zacharias Lorenz (; 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 regarde ...
*
Erich von Holst Erich Walther von Holst (28 November 1908 – 26 May 1962) was a German behavioral physiologist who was a Baltic German native of Riga, Livonia and was related to historian Hermann Eduard von Holst (1841–1904). In the 1950s he found ...
* Theodore H. Bullock * Jörg-Peter Ewert * Eric Knudsen *
Masakazu Konishi was a Japanese neurobiologist, known for his research on the neuroscience underlying the behavior of owls and songbirds. Early life and education Konishi was born on 17 February 1933 in Kyoto, Japan, the only child of poor "Nishijin" weavers. ...
* Martin Giurfa


References


Sources

* Beer D., Randall, Roy E. Ritzmann, Thomas McKenna (1993) Biological neural networks in invertebrate neuroethology and robotics. Boston : Academic Press. * Camhi, J.M. (1984) Neuroethology: Nerve cells and the Natural behavior of Animals, Sinauer Associates. * Carew, T.J. (2000) Feature analysis in Toads. Behavioral Neurobiology, Sunderland, MA: Sinauer, pp. 95–119. * Carew, T.J. (2000) Behavioral neurobiology: The Cellular Organization of Natural Behavior, Sinauer Associates. * Ewert, J.-P. (1974) The neural basis of visually guided behavior. Scientific American 230(3):34-42 * Ewert J.-P. (2004) Motion perception shapes the visual world of amphibians. In: Prete F.R. (Ed.) Complex Worlds from Simpler Nervous Systems. Cambridge, MA, MIT Press, pp. 117–160 * Hoyle, G. (1984) The scope of Neuroethology. Behavioural Brain Science 7:367-412. Graham Hoyle put forth a rather narrow definition of the goals and subject matter of neuroethology and links the field to the field of ethology. This is followed by commentaries from many prominent neuroethologists. It makes for fascinating reading. * Metzner, W. (1993) The Jamming avoidance response in ''Eigenmannia'' is controlled by two separate motor pathways. The Journal of Neuroscience. 13(5):1862-1878 * Pfluger, H.-J. and R. Menzel (1999) Neuroethology, its roots and future. J Comp Physiol A 185:389-392. * Zupanc, G.K.H. (2004) Behavioral Neurobiology: An Integrative Approach. Oxford University Press: Oxford, UK.


Further reading


Textbooks

* Sillar, K.T., Picton, L.P., Heitler, W.J. (2016) The Neuroethology of Predation and Escape. John Wiley & Sons Inc., New York. * Zupanc, G.K.H. (2004) Behavioral Neurobiology an Integrative Approach. Oxford University Press, New York. * Carew, T.J. (2000) Behavioral Neurobiology: The Cellular Organization of Natural Behavior. Sinauer, Sunderland Mass. * Simmons, P., Young, D. (1999) Nerve Cells and Animal Behaviour. Second Edition. Cambridge University Press, New York. * Simmons, P., Young, D. (2010) Nerve Cells and Animal Behaviour. Third Edition. Cambridge University Press, New York. * Camhi J. (1984) Neuroethology: Nerve Cells and the Natural Behavior of Animals. Sinauer Associates, Sunderland Mass. * Guthrie, D.M. (1980) Neuroethology: An Introduction. Wiley, New York. * Ewert, J.-P. (1980) Neuroethology: An Introduction to the Neurophysiological Fundamentals of Behaviour. Springer-Verlag, New York. * Ewert, J.-P. (1976) Neuroethologie: Einführung in die neurophysiologischen Grundlagen des Verhaltens. HT 181. Springer-Verlag Heidelberg, Berlin, New York. * Kandel, E.R. (1976) Cellular Basis of Behavior: An Introduction to Behavioral Neurobiology. W.H. Freeman * Roeder, K.D. (1967) Nerve Cells and Insect Behavior. Harvard University Press, Cambridge Mass. * Marler, P., Hamilton, W.J. (1966) Mechanisms of Animal Behavior. John Wiley & Sons Inc., New York.


Articles

*
Günther K. H. Zupanc (2010), Neuroethology, Scholarpedia, 5(10):5306.


External links


International Society for Neuroethology
*https://web.archive.org/web/20071006201121/http://www.tamie.org/insect.png
Collected Neuroethology articles in Scholarpedia
{{Zoology Ethology Neurophysiology