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====Insects==== Serotonin is evolutionarily conserved and appears across the animal kingdom. It is seen in insect processes in roles similar to in the human central nervous system, such as memory, appetite, sleep, and behavior.<ref>{{cite journal |doi=10.14800/nt.314 |title=Serotonin, serotonin receptors and their actions in insects |journal=Neurotransmitter |year=2015 |volume=2 |pages=1β14 |doi-access=free }}</ref><ref name="Huser_2012" /> Some circuits in [[mushroom bodies]] are serotonergic.<ref name="Schoofs-et-al-2017">{{cite journal | vauthors = Schoofs L, De Loof A, Van Hiel MB | title = Neuropeptides as Regulators of Behavior in Insects | journal = Annual Review of Entomology | volume = 62 | issue = 1 | pages = 35β52 | date = January 2017 | pmid = 27813667 | doi = 10.1146/annurev-ento-031616-035500 | publisher = [[Annual Reviews (publisher)|Annual Reviews]] | doi-access = free }}</ref> (See specific ''Drosophila'' example below, [[#Dipterans|Β§Dipterans]].) =====Acrididae===== Locust swarming is initiated ''but not maintained'' by serotonin,<ref name="Wang-Kang-2014">{{cite journal | vauthors = Wang X, Kang L | title = Molecular mechanisms of phase change in locusts | journal = Annual Review of Entomology | volume = 59 | issue = 1 | pages = 225β244 | date = 2014-01-07 | pmid = 24160426 | doi = 10.1146/annurev-ento-011613-162019 | publisher = [[Annual Reviews (publisher)|Annual Reviews]] | quote = <br />p.{{spaces}}231,<br />The change in the number of several potential neurotransmitters ... such as serotonin... may play an important role in remodeling the CNS during phase change (26, 56, 80).<br />p.{{spaces}}233,<br />In the locust ''S. gregaria'', the amount of serotonin in the thoracic ganglia was positively correlated with the extent of gregarious behavior induced by different periods of crowding. A series of pharmacological and behavioral experiments demonstrated that serotonin plays a key role in inducing initial behavioral gregarization (2, 80). However, serotonin is not responsible for maintaining gregarious behavior because its amount in long-term gregarious locusts is less than half that in long-term solitarious locusts (80). In ''L. migratoria'', the injection of serotonin can also slightly initiate gregarious behavior, but serotonin when accompanying crowding treatment induced more solitarious-like behavior than did serotonin injection alone (48). Significant differences in serotonin levels were not found in brain tissues between the two phases of ''L. migratoria''. A recent report by Tanaka & Nishide (97) measured attraction/avoidance behavior in ''S. gregaria'' after single and multiple injections of serotonin at different concentrations. Serotonin had only a short-term effect on the level of some locomotor activities and was not involved in the control of gregarious behavior (97). In addition, it is not clear how the neurotransmitter influences this unique behavior, because a binary logistic regression model used in these studies for the behavioral assay focused mostly on only one behavioral parameter representing an overall phase state. Obviously, behavioral phase change might involve alternative regulatory mechanisms in different locust species. Therefore, these studies demonstrate that CNS regulatory mechanisms governing initiation and maintenance of phase change are species specific and involve the interactions between these neurotransmitters.<br />Given the key roles of aminergic signaling, what are the downstream pathways involved in the establishment of long-term memory? Ott et al. (63) investigated the role of [] protein kinase[] in the phase change in ''S. gregaria'': ... cAMP-dependent protein kinase A (PKA). Through use of pharmacological and RNAi intervention, these authors have demonstrated that PKA... has a critical role in modulating the propensity of locusts to acquire and express gregarious behavior. ... Unfortunately, although a correlation between serotonin and PKA was hypothesized, direct evidence was not provided. | doi-access = free }}</ref> with release being triggered by tactile contact between individuals.<ref name="Zhang-et-al-2019">{{cite journal | vauthors = Zhang L, Lecoq M, Latchininsky A, Hunter D | title = Locust and Grasshopper Management | journal = Annual Review of Entomology | volume = 64 | issue = 1 | pages = 15β34 | date = January 2019 | pmid = 30256665 | doi = 10.1146/annurev-ento-011118-112500 | publisher = [[Annual Reviews (publisher)|Annual Reviews]] | quote-page = 20 | s2cid = 52843907 | doi-access = free | quote = ...gregarization is evoked by... tactile stimulation... Tactile stimuli trigger the increase of biogenic amines, particularly serotonin, in the locust nervous system (1, 116); these amines play critical roles in the neurophysiology of locust behavioral phase change. }}</ref> This transforms social preference from aversion to a gregarious state that enables coherent groups.<ref name="Anstey">{{cite journal | vauthors = Anstey ML, Rogers SM, Ott SR, Burrows M, Simpson SJ | title = Serotonin mediates behavioral gregarization underlying swarm formation in desert locusts | journal = Science | volume = 323 | issue = 5914 | pages = 627β630 | date = January 2009 | pmid = 19179529 | doi = 10.1126/science.1165939 | s2cid = 5448884 | bibcode = 2009Sci...323..627A }} * {{cite news | vauthors = Morgan J |date=29 January 2009 |title=Locust swarms 'high' on serotonin |work=BBC News |url=http://news.bbc.co.uk/2/hi/science/nature/7858996.stm}}</ref><ref name="Zhang-et-al-2019" /><ref name="Wang-Kang-2014" /> Learning in flies and honeybees is affected by the presence of serotonin.<ref>{{cite journal | vauthors = Sitaraman D, LaFerriere H, Birman S, Zars T | title = Serotonin is critical for rewarded olfactory short-term memory in Drosophila | journal = Journal of Neurogenetics | volume = 26 | issue = 2 | pages = 238β244 | date = June 2012 | pmid = 22436011 | doi = 10.3109/01677063.2012.666298 | s2cid = 23639918 }}</ref><ref>{{cite journal | vauthors = Bicker G, Menzel R | title = Chemical codes for the control of behaviour in arthropods | journal = Nature | volume = 337 | issue = 6202 | pages = 33β39 | date = January 1989 | pmid = 2562906 | doi = 10.1038/337033a0 | s2cid = 223750 | bibcode = 1989Natur.337...33B }}</ref> {{anchor|Insecticide|Insecticides}} =====Role in insecticides===== Insect 5-HT receptors have similar sequences to the vertebrate versions, but pharmacological differences have been seen. Invertebrate drug response has been far less characterized than mammalian pharmacology and the potential for species selective insecticides has been discussed.<ref>{{cite journal | vauthors = Cai M, Li Z, Fan F, Huang Q, Shao X, Song G | title = Design and synthesis of novel insecticides based on the serotonergic ligand 1-[(4-aminophenyl)ethyl]-4-[3-(trifluoromethyl)phenyl]piperazine (PAPP) | journal = Journal of Agricultural and Food Chemistry | volume = 58 | issue = 5 | pages = 2624β2629 | date = March 2010 | pmid = 20000410 | doi = 10.1021/jf902640u | bibcode = 2010JAFC...58.2624C }}</ref> {{anchor|Hymenoptera}} =====Hymenopterans===== [[Wasp]]s and [[hornets]] have serotonin in their venom,<ref>{{cite book | vauthors = Manahan SE |title=Toxicological Chemistry and Biochemistry |edition=3rd |publisher=CRC Press |year=2002 |isbn=978-1-4200-3212-3 |page=393 }}</ref> which causes pain and inflammation<ref name="Chen_2010" >{{cite journal | vauthors = Chen J, Lariviere WR | title = The nociceptive and anti-nociceptive effects of bee venom injection and therapy: a double-edged sword | journal = Progress in Neurobiology | volume = 92 | issue = 2 | pages = 151β183 | date = October 2010 | pmid = 20558236 | pmc = 2946189 | doi = 10.1016/j.pneurobio.2010.06.006 }}</ref><ref name="Erspamer-1966" /> as do [[scorpion]]s.<ref>{{cite book | vauthors = Postma TL |chapter=Neurotoxic Animal Poisons and Venoms |chapter-url=http://www.sciencedirect.com/science/article/pii/B9780323052603500496 |pages=463β489 | veditors = Dobbs MR |year=2009 |title=Clinical Neurotoxicology |publisher=W.B. Saunders |doi=10.1016/B978-032305260-3.50049-6 |isbn=978-0-323-05260-3 }}</ref><ref name="Erspamer-1966" /> ''[[Pheidole dentata]]'' takes on more and more tasks in [[ant colony|the colony]] as it gets older, which requires it to respond to more and more [[olfaction|olfactory]] cues in the course of performing them. This olfactory response broadening was demonstrated to go along with increased serotonin and [[dopamine]], but not [[octopamine]] in 2006.<ref name="Gadenne-et-al-2016">{{cite journal | vauthors = Gadenne C, Barrozo RB, Anton S | title = Plasticity in Insect Olfaction: To Smell or Not to Smell? | journal = Annual Review of Entomology | volume = 61 | issue = 1 | pages = 317β333 | date = 2016-03-11 | pmid = 26982441 | doi = 10.1146/annurev-ento-010715-023523 | publisher = [[Annual Reviews (publisher)|Annual Reviews]] | hdl-access = free | s2cid = 207568844 | hdl = 11336/19586 }}</ref> =====Dipterans===== If flies are fed serotonin, they are more aggressive; flies depleted of serotonin still exhibit aggression, but they do so much less frequently.<ref name="Dierick">{{cite journal | vauthors = Dierick HA, Greenspan RJ | title = Serotonin and neuropeptide F have opposite modulatory effects on fly aggression | journal = Nature Genetics | volume = 39 | issue = 5 | pages = 678β682 | date = May 2007 | pmid = 17450142 | doi = 10.1038/ng2029 | s2cid = 33768246 }}</ref> In [[Dipteran crop|their crops]] it plays a vital role in digestive motility produced by contraction. Serotonin that acts on the crop is exogenous to the crop itself and 2012 research suggested that it probably originated in the serotonin neural plexus in the thoracic-abdominal synganglion.<ref name="Stoffolano-Haselton-2013">{{cite journal | vauthors = Stoffolano JG, Haselton AT | title = The adult Dipteran crop: a unique and overlooked organ | journal = Annual Review of Entomology | volume = 58 | issue = 1 | pages = 205β225 | date = 2013-01-07 | pmid = 23317042 | doi = 10.1146/annurev-ento-120811-153653 | publisher = [[Annual Reviews (publisher)|Annual Reviews]] | author-link = John Stoffolano }}</ref> In 2011 a ''[[Drosophila]]'' serotonergic mushroom body was found to work in concert with ''[[Amnesiac gene|Amnesiac]]'' to form memories.<ref name="Schoofs-et-al-2017" /> In 2007 serotonin was found to promote aggression in ''[[Diptera]]'', which was counteracted by [[neuropeptide F]] β a surprising find given that they both promote [[insect courtship|courtship]], which is usually similar to aggression in most respects.<ref name="Schoofs-et-al-2017" />
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