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“Psychobiology” redirects here. For the journal, see was distrusted by the dominant anatomists of the 18th and Cognitive, Affective, & . 19th centuries.[5] The influential work of Claude Bernard, “Biological ” redirects here. For the journal, , and helped to convince the see Biological Psychology (journal). scientific community that reliable data could be obtained For related topics, see Affective neuroscience, Behavioral from living subjects. , , , Even before the 18th and 19th century, behavioral neu- , and . roscience was beginning to take form as far back as 1700 B.C.[6] The question that seems to continually arise is Behavioral neuroscience, also known as biological psy- what is the connection between the and body. The chology,[1] biopsychology, or psychobiology[2] is the debate is formally referred to as the Mind-body problem. application of the of (in particular There are two major schools of that attempt to neurobiology), to the study of physiological, genetic, and resolve the mind–body problem; and dualism.[3] developmental mechanisms of in and and are two of several philosophers who non- animals. It typically investigates at the level participated in this debate. Plato believed that the of , , brain circuitry and the ba- was where all mental thought and processes happened.[6] sic biological processes that underlie normal and abnor- In contrast, Aristotle believed that the brain served the mal behavior. Often, in behavioral neuro- purpose of cooling down the derived from the involve non-human animal models (such as rats .[3] The Mind-body problem was a stepping stone and mice, and non-human primates) which have impli- toward attempting to understand the connection between cations for better of human pathology and the mind and body. therefore contribute to evidence-based practice. Another debate arose about was localization of function or Functional specialization (brain) versus equipotentiality which played a significant role in the 1 development in behavioral neuroscience. As a result of localization of function research, many famous people found within psychology have come to various different Behavioral neuroscience as a scientific discipline conclusions. Wilder Penfield was able to develop a map emerged from a variety of scientific and philosophical of the through studying epileptic patients traditions in the 18th and 19th centuries. In , along with Rassmussen.[3] Research on localization of people like René Descartes proposed physical models function has led behavioral to a better to explain animal and . Descartes, for understanding of which parts of the brain control example, suggested that the , a midline behavior. This is best exemplified through the unpaired structure in the brain of many organisms, was of . the point of contact between mind and body. Descartes The term “psychobiology” has been used in a variety of also elaborated on a theory in which the pneumatics contexts,emphasizing the importance of biology, which of bodily fluids could explain reflexes and other motor is the discipline that studies organic, neural and cellular behavior. This theory was inspired by moving statues in modifications in behavior, plasticity in neuroscience, and a garden in Paris.[3] biological deceases in all aspects, in addition, biology fo- Other philosophers also helped give birth to psychology. cuses and analyzes behavior and all the subjects it is con- One of the earliest textbooks in the new field, The Princi- cerned about, from a scientific point of view. In this con- ples of Psychology by (1890), argues that text, psychology helps as a complementary, but impor- the scientific study of psychology should be grounded in tant discipline in the neurobiological . The role an understanding of biology: of psychology in this questions is that of a social tool that James, like many early , had considerable backs up the main or strongest biological science. The training in . The emergence of both psychol- term “psychobiology” was first used in its modern ogy and behavioral neuroscience as legitimate sciences by in his book An Outline of Psychobiol- [7] can be traced from the emergence of physiology from ogy (1914). Dunlap also was the founder and editor-in- , particularly . Physiologists con- chief of the journal Psychobiology. In the announcement ducted experiments on living organisms, a practice that of that journal, Dunlap writes that the journal will pub-

1 2 3 RESEARCH METHODS

lish research "...bearing on the interconnection of mental • Surgical lesions - Neural tissue is destroyed by and physiological functions”, which describes the field of removing it surgically. [7] behavioral neuroscience even in its modern sense. • Electrolytic lesions - Neural tissue is destroyed through the application of electrical shock trauma. 2 Relationship to other fields of • Chemical lesions - Neural tissue is destroyed psychology and biology by the infusion of a . • Temporary lesions - Neural tissue is tem- In many cases, humans may serve as experimental porarily disabled by cooling or by the use of subjects in behavioral neuroscience experiments; how- such as tetrodotoxin. ever, a great deal of the experimental literature in be- havioral neuroscience comes from the study of non- • Transcranial magnetic stimulation - A new tech- human species, most frequently rats, mice, and mon- nique usually used with human subjects in which keys. As a result, a critical assumption in behavioral neu- a magnetic coil applied to the scalp causes unsys- roscience is that organisms share biological and behav- tematic electrical activity in nearby cortical neurons ioral similarities, enough to permit extrapolations across which can be experimentally analyzed as a func- species. This allies behavioral neuroscience closely tional lesion. with , , , and neurobiology. Behavioral neu- • Psychopharmacological manipulations - A chemical roscience also has paradigmatic and methodological sim- induces neural activity by inter- ilarities to neuropsychology, which relies heavily on the fering with . Antagonists can be study of the behavior of humans with dys- delivered systemically (such as by intravenous in- function (i.e., a non-experimentally based biological ma- jection) or locally (intracerebrally) during a surgical nipulation). procedure into the ventricles or into specific brain structures. For example, NMDA antagonist AP5 Synonyms for behavioral neuroscience, though with has been shown to inhibit the initiation of long term a slightly different emphasis and organization, in- potentiation of excitatory synaptic transmission (in clude biopsychology, biological psychology, and rodent fear conditioning) which is believed to be a psychobiology.[8] is another vital mechanism in and .[9] term often used synonymously with behavioral neuro- science, though most authors would make physiological • Optogenetic inhibition - A light activated inhibitory psychology a subfield of behavioral neuroscience, with protein is expressed in cells of interest. Power- an appropriately more narrow definition. ful millisecond timescale neuronal inhibition is in- stigated upon stimulation by the appropriate fre- quency of light delivered via fiber optics or im- 3 Research methods planted LEDs in the case of vertebrates,[10] or via external illumination for small, sufficiently translu- cent invertebrates.[11] Bacterial or The distinguishing characteristic of a behavioral neuro- Proton pumps are the two classes of proteins used science is that either the independent variable for inhibitory , achieving inhibition by of the experiment is biological, or some dependent vari- increasing cytoplasmic levels of halides (Cl-) or de- able is biological. In other words, the nervous system of creasing the cytoplasmic concentration of protons, the organism under study is permanently or temporarily respectively.[12][13] altered, or some aspect of the nervous system is measured (usually to be related to a behavioral variable). 3.2 Enhancing Neural Function

3.1 Disabling or decreasing neural func- • Electrical stimulation - A classic method in which tion neural activity is enhanced by application of a small electrical current (too small to cause significant • Lesions - A classic method in which a brain-region death). of interest is naturally or intentionally destroyed to observe any resulting changes such as degraded or • Psychopharmacological manipulations - A chemi- enhanced performance on some behavioral measure. cal receptor facilitates neural activity by en- Lesions can be placed with relatively high accuracy hancing or replacing endogenous neurotransmitters. thanks to a variety of brain 'atlases’ which provide can be delivered systemically (such as by a map of brain regions in 3-dimensional stereotactic intravenous injection) or locally (intracerebrally) coordinates. during a surgical procedure. 3.4 Genetic techniques 3

• Transcranial magnetic stimulation - In some cases be detected in an MRI apparatus and are taken to (for example, studies of ), this tech- indicate relative activity of larger scale brain regions nique can be analyzed as having a stimulatory effect (i.e., on the order of hundreds of thousands of neu- (rather than as a functional lesion). rons). • Optogenetic excitation - A light activated ex- • citatory protein is expressed in select cells. - Or EEG; and the deriva- −2 (ChR2), a light activated tive technique of event-related potentials, in which cation channel, was the first bacterial opsin shown scalp electrodes monitor the average activity of neu- to excite neurons in response to light,[14] though a rons in the cortex (again, used most frequently with number of new excitatory optogenetic tools have human subjects). now been generated by improving and imparting novel properties to ChR2[15] • Functional neuroanatomy - A more coun- terpart of . The expression of some anatomical marker is taken to reflect neural activ- 3.3 Measuring neural activity ity. For example, the expression of immediate early genes is thought to be caused by vigorous neural ac- • Optical techniques - Optical methods for recording tivity. Likewise, the injection of 2-deoxyglucose neuronal activity rely on methods that modify the prior to some behavioral task can be followed by optical properties of neurons in response to the cel- anatomical localization of that chemical; it is taken lular events associated with action potentials or neu- up by neurons that are electrically active. rotransmitter release. • Voltage sensitive dyes (VSDs) were among • MEG - shows the func- the earliest method for optically detecting ac- tioning of the through the measurement tion potentials. VSDs commonly become flu- of electromagnetic activity. Measuring the magnetic orescent in response to a ’s change fields created by the electric current flowing within in voltage, rendering action poten- the neurons identifies brain activity associated with tials detectable.[16] Genetically encoded volt- various human functions in real , with millime- age sensitive fluorescent proteins have also ter spatial accuracy. Clinicians can noninvasively been developed.[17] obtain data to help them assess neurological disor- • relies on dyes[18] or ge- ders and surgical treatments. netically encoded proteins[19] that fluoresce upon binding to the calcium that is transiently present during an action potential. 3.4 Genetic techniques • Synapto-pHluorin is a technique that relies on a fusion protein that combines a synaptic vesi- • QTL mapping - The influence of a gene in some cle membrane protein and a pH sensitive fluo- behavior can be statistically inferred by studying rescent protein. Upon synaptic vesicle release, inbred strains of some species, most commonly the chimeric protein is exposed to the higher mice. The recent sequencing of the of many pH of the synaptic cleft, causing a measurable species, most notably mice, has facilitated this tech- [20] change in fluorescence. nique. • Single-unit recording - A method whereby an elec- trode is introduced into the brain of a living animal • - Organisms, often mice, may to detect electrical activity that is generated by the be bred selectively among inbred strains to create a neurons adjacent to the electrode tip. Normally this recombinant congenic strain. This might be done to is performed with sedated animals but sometimes it isolate an experimentally interesting stretch of DNA is performed on awake animals engaged in a behav- derived from one strain on the background genome ioral event, such as a thirsty rat whisking a particu- of another strain to allow stronger inferences about lar sandpaper grade previously paired with water in the role of that stretch of DNA. order to measure the corresponding of neu- ronal firing at the decision point.[21] • - The genome may also be experimentally-manipulated; for example, knockout • Multielectrode recording - The use of a bundle of mice can be engineered to lack a particular gene, or fine electrodes to record the simultaneous activity of a gene may be expressed in a strain which does not up to hundreds of neurons. normally do so (the 'transgenic'). Advanced tech- • fMRI - Functional magnetic resonance imaging, a niques may also permit the expression or suppres- technique most frequently applied on human sub- sion of a gene to occur by injection of some regulat- jects, in which changes in cerebral blood flow can ing chemical. 4 5 NOBEL LAUREATES

3.5 Limitations and advantages • Parkinson’s Disease, a degenerative disorder of the central nervous system that often impairs the suf- Different manipulations have advantages and limitations. ferer’s motor skills and speech. Neural tissue destroyed by surgery, electric shock or neu- • rotoxcin is a permanent manipulation and therefore lim- Huntington’s Disease, a rare inherited neurological its follow-up investigation.[22] Most genetic manipulation disorder whose most obvious symptoms are abnor- techniques are also considered permanent.[22] Tempo- mal body movements and a lack of coordination. It rary lesions can be achieved with advanced in genetic also affects a number of mental abilities and some manipulations, for example, certain genes can now be aspects of personality. [22] switched on and off with diet. Pharmacological manip- • Alzheimer’s Disease, a neurodegenerative disease ulations also allow blocking of certain neurotransmitters that, in its most common form, is found in people temporarily as the function returns to its previous state over the age of 65 and is characterized by progres- [22] after the drug has been metabolized. sive cognitive deterioration, together with declin- ing activities of daily living and by neuropsychiatric symptoms or behavioral changes. 4 Topic areas in behavioral neuro- • Clinical depression, a common psychiatric disorder, science characterized by a persistent lowering of mood, loss of interest in usual activities and diminished ability In general, behavioral study similar to pleasure. themes and issues as academic psychologists, though lim- ited by the need to use nonhuman animals. As a result, the • , a psychiatric diagnosis that describes bulk of literature in behavioral neuroscience deals with a mental illness characterized by impairments in the mental processes and that are shared across dif- or expression of , most commonly ferent animal models such as: manifesting as auditory hallucinations, paranoid or bizarre delusions or disorganized speech and think- • Sensation and perception ing in the context of significant social or occupa- tional dysfunction. • Motivated behavior (hunger, thirst, sex) • , a brain development disorder that impairs • Control of movement social interaction and , and causes restricted and repetitive behavior, all starting before • Learning and memory a child is three years old. • and biological rhythms • , a physiological state characterized by cog- nitive, somatic, emotional, and behavioral compo- • nents. These components combine to create the feelings that are typically recognized as fear, appre- However, with increasing technical sophistication and hension, or worry. with the development of more precise noninvasive meth- • ods that can be applied to human subjects, behavioral Drug , including . neuroscientists are beginning to contribute to other classi- cal topic areas of psychology, philosophy, and , such as: 5 Nobel Laureates

• Language The following winners could reasonably be considered biological neuroscientists or neurobiologists. • Reasoning and decision making (This list omits winners who were almost exclusively neuroanatomists or neurophysiologists; i.e., those that did • not measure behavioral or neurobiological variables.)

Behavioral neuroscience has also had a strong history of • Charles Sherrington (1932) contributing to the understanding of medical disorders, including those that fall under the purview of clinical psy- • (1932) chology and biological (also known as • ). Although animal models do not Walter Hess (1949) exist for all mental illnesses, the field has contributed im- • Egas Moniz (1949) portant therapeutic data on a variety of conditions, in- cluding: • Georg von Békésy (1961) 5

• (1967) 7 References • (1967) [1] Breedlove, Watson, Rosenzweig, Biological Psychology: An Introduction to Behavioral and Cognitive Neuroscience, • (1973) 6/e, ISBN 978-0-87893-705-9, p. 2 • Niko Tinbergen (1973) [2] Psychobiology, Merriam-Webster’s Online Dictionary

• (1973) [3] Carlson, Neil (2007). Physiology of Behavior (9th Ed.). Allyn and Bacon. pp. 11–14. ISBN 0-205-46724-5. • Roger W. Sperry (1981) [4] James, William (1950/1890). The Principles of Psychol- • David H. Hubel (1981) ogy, Vol. One. Dover Publications, Inc. pp. 4–5. ISBN 0-486-20381-6. Check date values in: |date= (help) • Torsten N. Wiesel (1981) [5] Shepherd, Gordon M. (1991). Foundations of the Neuron • Eric R. Kandel (2000) Doctrine. Oxford University Press. ISBN 0-19-506491-7.

• (2000) [6] “”. Columbia University. Re- trieved 2014-05-04. • (2004) [7] Dewsbury, Donald (1991). “Psychobiology”. American • Linda B. Buck (2004) (46): 198–205.

• John O'Keefe (2014) [8] S. , Mark Rosenzweig and Neil V. Wat- son (2007). Biological Psychology: An Introduction to • (2014) Behavioral and Cognitive Neuroscience 6e. Sinauer As- sociates. ISBN 978-0-87893-705-9 • May-Britt Moser (2014) [9] Kim, Jeansok J.; DeCola, Joseph P.; Landeira-Fernandez, Jesus; Fanselow, Michael S. “N-methyl-D-aspartate re- ceptor antagonist APV blocks acquisition but not ex- 6 See also pression of fear conditioning.” Behavioral Neuroscience. Vol 105(1), Feb 1991, 126-133. {doi|10.1037/0735- 7044.105.1.126} • Affective neuroscience • [10] Schneider et al. “Controlling Neuronal Activity.” Biological American Journal of Psychiatry 165:562, May 2008 doi:10.1176/appi.ajp.2008.08030444 • Biology [11] Zhang, et al. “Multimodal fast optical interrogation • Cognitive neuroscience of neural circuitry.” . Vol 446. 5 April 2007. doi:10.1038/nature05744 • Developmental psychobiology [12] Chow, B. Y. et al. “High-performance genetically tar- • Epigenetics in psychology getable optical neural silencing by light-driven proton pumps.” Nature. Vol 463. 7 January 2010 • Evolutionary psychology [13] Gradinaru, Thompson, and Deisseroth. “eNpHR: a Na- • List of topics related to tronomonas enhanced for optogenetic ap- plications.” Brain cell Biology. Vol 36 (1-4). Aug 2008. • Models of abnormality doi:10.1007/s11068-008-9027-6

• Neurobiology [14] Zhang, Wang, Boyden, and Deisseroth. “Channelrhodopsin-2 and optical control of excitable • cells.” Nature Methods. VOL.3 NO.10. OCTOBER 2006 • [15] Gradinaru et al. “Molecular and Cellular Approaches for • Physical Diversifying and Extending Optogenetics.” Cell. 2010. doi:10.1016/j.cell.2010.02.037 • [16] Ebner, T. J. and Chen, G. “Use of voltage-sensitive dyes • and optical recordings in the central nervous system.” Progress in Neurobiology Volume 46, Issue 5, August • Social neuroscience 1995, 463-506. doi:10.1016/0301-0082(95)00010-S 6 8 EXTERNAL LINKS

[17] Micah S. Siegel and Ehud Y. Isacoff. “A Genetically En- coded Optical Probe of Membrane Voltage."Neuron, Vol. 19, 735–741, October, 1997

[18] O'Donovan, Hoa, Sholomenkoa, and Yeea. “Real-time imaging of neurons retrogradely and anterogradely la- belled with calcium-sensitive dyes.” Journal of Neuro- science Methods. Vol 46, Issue 2, February 1993, 91-106. doi:10.1016/0165-0270(93)90145-H

[19] Nicola Heim and Oliver Griesbeck. “Genetically Encoded Indicators of Cellular Calcium Dynamics Based on Tro- ponin C and Green Fluorescent Protein.” The Journal of Biological Chemistry, 279, 14280-14286. April 2, 2004 doi:10.1074/jbc.M312751200

[20] Gero Miesenböck, Dino A. De Angelis & James E. Roth- man1. “Visualizing secretion and synaptic transmission with pH-sensitive green fluorescent proteins.” Nature 394, 192-195 (9 July 1998) | doi:10.1038/28190

[21] von Heimendahl, M., Itskov, P., Arabzadeh, E., & Di- amond, M. (2007). Neuronal activity in rat barrel cor- tex underlying texture discrimination. PLoS Biol, 5(11), e305.

[22] T Abel, KM Lattal (2001) “Molecular mechanisms of memory acquisition, consolidation and retrieval” Current Opinion in Neurobiology

8 External links

• Biological Psychology Links

• Theory of Biological Psychology (Documents No. 9 and 10 in English) • IBRO (International Brain Research Organization) 7

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