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The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial cells, known as hair cells, which respond to displacement caused by motion and transduce these signals into electrical impulses via excitatory synapses. Lateral lines play an important role in schooling behavior, predation, and orientation.

Early in the evolution of fish, some of the sensory organs of the lateral line were modified to function as the electroreceptors called ampullae of Lorenzini. The lateral line system is ancient and basal to the vertebrate clade, as it is found in fishes that diverged over 400 million years ago.

Function

The lateral line system allows the detection of movement, vibration, and pressure gradients in the water surrounding an animal. It plays an essential role in orientation, predation, and fish schooling by providing spatial awareness and the ability to navigate in the environment.^[1] Analysis has shown that the lateral line system should be an effective passive sensing system able to discriminate between submerged obstacles by their shape.^[2]^[3] The lateral line allows fish to navigate and hunt in water with poor visibility.^[4]

The lateral line system enables predatory fishes to detect vibrations made by their prey, and to orient towards the source to begin predatory action.^[5] Blinded predatory fishes remain able to hunt, but not when lateral line function is inhibited by cobalt ions.^[6]

The lateral line plays a role in fish schooling. Blinded Pollachius virens were able to integrate into a school, whereas fish with severed lateral lines could not.^[7] It may have evolved further to allow fish to forage in dark caves. In Mexican blind cave fish, Astyanax mexicanus, neuromasts in and around the orbit of the eye are bigger and around twice as sensitive as those of surface-living fish.^[8]

One function of schooling may be to confuse the lateral line of predatory fishes. A single prey fish creates a simple particle velocity pattern, whereas the pressure gradients of many closely swimming (schooling) prey fish overlap, creating a complex pattern. This makes it difficult for predatory fishes to identify individual prey through lateral line perception.^[9]

Anatomy

Lateral lines are usually visible as faint lines of pores running along each side of a fish's body.^[1] The functional units of the lateral line are the neuromasts, discrete mechanoreceptive organs that sense movement in water. There are two main varieties: canal neuromasts and superficial neuromasts. Superficial neuromasts are on the surface of the body, while canal neuromasts are along the lateral lines in subdermal, fluid-filled canals. Each neuromast consists of receptive hair cells whose tips are covered by a flexible jellylike cupula. Hair cells typically possess both glutamatergic afferent connections and cholinergic efferent connections.^[11]^[12] The receptive hair cells are modified epithelial cells; they typically possess bundles of 40–50 microvilli "hairs" which function as the mechanoreceptors.^[12] Within each bundle, the hairs are organized in a rough "staircase" from shortest to longest.^[13]

Signal transduction

The hair cells are stimulated by the deflection of their hair bundles in the direction of the tallest "hairs" or stereocilia. The deflection allows cations to enter through a mechanically gated channel, causing depolarization or hyperpolarization of the hair cell. Depolarization opens Ca_v1.3 calcium channels in the basolateral membrane.^[14]

Hair cells use a system of transduction with rate coding to transmit the directionality of a stimulus. The hair cells produce a constant, tonic rate of firing. As mechanical motion is transmitted through water to the neuromast, the cupula bends and is displaced according to the strength of the stimulus. This results in a shift in the cell's ionic permeability. Deflection towards the longest hair results in depolarization of the hair cell, increased neurotransmitter release at the excitatory afferent synapse, and a higher rate of signal transduction. Deflection towards the shorter hair has the opposite effect, hyperpolarizing the hair cell and producing a decreased rate of neurotransmitter release. These electrical impulses are then transmitted along afferent lateral neurons to the brain.^[12]

While both varieties of neuromasts utilize this method of transduction, their specialized organization gives them different mechanoreceptive capacities. Superficial organs are exposed more directly to the external environment. The organization of the bundles within their organs is seemingly haphazard, incorporating various shapes and sizes of microvilli within bundles. This suggests coarse but wide-ranging detection.^[13] In contrast, the structure of canal organs allow canal neuromasts more sophisticated mechanoreception, such as of pressure differentials. As current moves across the pores, a pressure differential is created, inducing a flow in the canal fluid. This moves the cupulae of the neuromasts in the canal, resulting in a deflection of the hairs in the direction of the flow.^[15]

Electrophysiology

The mechanoreceptive hair cells of the lateral line structure are integrated into more complex circuits through their afferent and efferent connections. The synapses that directly participate in the transduction of mechanical information are excitatory afferent connections that utilize glutamate.^[16] Species vary in their neuromast and afferent connections, providing differing mechanoreceptive properties. For instance, the superficial neuromasts of the midshipman fish, Porichthys notatus, are sensitive to specific stimulation frequencies.^[17] One variety is attuned to collect information about acceleration, at stimulation frequencies between 30 and 200 Hz. The other type obtains information about velocity, and is most receptive to stimulation below 30 Hz.^[17]

The efferent synapses to hair cells are inhibitory and use acetylcholine as a transmitter. They are crucial participants in a corollary discharge system designed to limit self-generated interference. When a fish moves, it creates disturbances in the water that could be detected by the lateral line system, potentially interfering with the detection of other biologically relevant signals. To prevent this, an efferent signal is sent to the hair cell upon motor action, resulting in inhibition which counteracts the excitation resulting from reception of the self-generated stimulation. This allows the fish to detect external stimuli without interference from its own movements.^[18]

Signals from the hair cells are transmitted along lateral neurons to the brain. The area where these signals most often terminate is the medial octavolateralis nucleus (MON), which probably processes and integrates mechanoreceptive information.^[19] The deep MON contains distinct layers of basilar and non-basilar crest cells, suggesting computational pathways analogous to the electrosensory lateral line lobe of electric fish. The MON is likely involved in the integration of excitatory and inhibitory parallel circuits to interpret mechanoreceptive information.^[20]

Evolution

The use of mechanosensitive hairs is homologous to the functioning of hair cells in the auditory and vestibular systems, indicating a close link between these systems.^[12] Due to many overlapping functions and their great similarity in ultrastructure and development, the lateral line system and the inner ear of fish are often grouped together as the octavolateralis system (OLS).^[23] Here, the lateral line system detects particle velocities and accelerations with frequencies below 100 Hz. These low frequencies create large wavelengths, which induce strong particle accelerations in the near field of swimming fish that do not radiate into the far field as acoustic waves due to an acoustic short circuit. The auditory system detects pressure fluctuations with frequencies above 100 Hz that propagate to the far field as waves.^[24]

The lateral line system is ancient and basal to the vertebrate clade; it is found in groups of fishes that diverged over 400 million years ago, including the lampreys, cartilaginous fishes, and bony fishes.^[25]^[26] Most amphibian larvae and some fully aquatic adult amphibians possess mechanosensitive systems comparable to the lateral line.^[27] The terrestrial tetrapods have secondarily lost their lateral line organs, which are ineffective when not submerged.^[26]

The electroreceptive organs, called ampullae of Lorenzini, appearing as pits in the skin of sharks and some other fishes, evolved from the lateral line organ.^[21] Passive electroreception using ampullae is an ancestral trait in the vertebrates, meaning that it was present in their last common ancestor.^[22]

>440 mya

430 mya

Amphibians
(as larvae; some as adults)
	Other tetrapods
(no lateral line)

Ray-finned fishes

425 mya

Ampullae of Lorenzini

Lateral line

References

^ ^a ^b Bleckmann, Horst; Zelick, Randy (2009-03-01). "Lateral line system of fish". Integrative Zoology. 4 (1): 13–25. doi:10.1111/j.1749-4877.2008.00131.x. PMID 21392273.
^ Bouffanais, Roland; Weymouth, Gabriel D.; Yue, Dick K. P. (2 June 2010). "Hydrodynamic object recognition using pressure sensing" (PDF). Proceedings of the Royal Society A: Mathematical, Physical and Engineering Sciences. 467 (2125): 19–38. doi:10.1098/rspa.2010.0095. S2CID 233793.
^ Lakkam, Sreetej; Balamurali, B. T.; Bouffanais, Roland (2 August 2019). "Hydrodynamic object identification with artificial neural models". Scientific Reports. 9 (1): 11242. arXiv:1903.00828. Bibcode:2019NatSR...911242L. doi:10.1038/s41598-019-47747-8. PMC 6677828. PMID 31375742.
^ Coombs, Sheryl; Braun, Christopher B. (2003). "Information Processing by the Lateral Line System". In Collin, Shaun P.; Marshall, N. Justin (eds.). Sensory Processing in Aquatic Environments. Springer Science & Business Media. pp. 122–138. doi:10.1007/978-0-387-22628-6_7. ISBN 978-0-387-95527-8.
^ Coombs, S. I.; Braun, C. B.; Donovan, B. (2001). "The orienting response of Lake Michigan mottled sculpin is mediated by canal neuromasts". Journal of Experimental Biology. 204 (2): 337–348. doi:10.1242/jeb.204.2.337. PMID 11136619.
^ Karlsen, H. E.; Sand, O. (1987). "Selective and Reversible Blocking of the Lateral Line in Freshwater Fish". The Journal of Experimental Biology. 133 (1): 249–262. doi:10.1242/jeb.133.1.249.
^ Pitcher, T.; Partridge, B.; Wardle, C. (1976). "A blind fish can school". Science. 194 (4268): 963–965. Bibcode:1976Sci...194..963P. doi:10.1126/science.982056. PMID 982056.
^ Yoshizawa, Masato; Jeffery, William; Van Netten, Sietse; McHenry, Matthew (2014). "The sensitivity of lateral line receptors and their role in the behavior of Mexican blind cavefish, Astyanax mexicanus". The Journal of Experimental Biology. 217 (6): 886–895. doi:10.1242/jeb.094599. PMC 3951362. PMID 24265419.
^ Larsson, M. (2009). "Possible functions of the octavolateralis system in fish schooling". Fish and Fisheries. 10 (3): 344–355. Bibcode:2009AqFF...10..344L. doi:10.1111/j.1467-2979.2009.00330.x.
^ Wark, Abigail R.; Mills, Margaret G.; Dang, Lam-Ha; et al. (2012). "Genetic Architecture of Variation in the Lateral Line Sensory System of Threespine Sticklebacks". G3: Genes, Genomes, Genetics. 2 (9): 1047–1056. doi:10.1534/g3.112.003079. PMC 3429919. PMID 22973542.
^ Russell, I. J. (1971). "The Pharmacology of Efferent Synapses in the Lateral-Line System of Xenopus Laevis". Journal of Experimental Biology. 54 (3): 643–659. doi:10.1242/jeb.54.3.643. PMID 4326222.
^ ^a ^b ^c ^d Flock, Å. (1967). "Ultrastructure and function in the lateral line organs". In P. Cahn (ed.). Lateral Line Detectors; proceedings of a conference held at Yeshiva University, New York, April 16-18, 1966. Indiana University Press. pp. 163–197.
^ ^a ^b Peach, M. B.; Rouse, G. W. (2000). "The Morphology of the Pit Organs and Lateral Line Canal Neuromasts of Mustelus Antarcticus (Chondrichthyes: Triakidae)". Journal of the Marine Biological Association of the United Kingdom. 80 (1): 155–162. Bibcode:2000JMBUK..80..155P. doi:10.1017/s0025315499001678. S2CID 85963954.
^ Baker, Clare (June 11, 2018). "Insights into Electroreceptor Development and Evolution from Molecular Comparisons with Hair Cells". Integrative and Comparative Biology. 58 (2): 329–340. doi:10.1093/icb/icy037. PMC 6927855. PMID 29846597.
^ Kuiper, J. W. (1967). "Frequency Characteristics and Functional Significance of the Lateral Line Organ". In P. Cahn (ed.). Lateral Line Detectors; proceedings of a conference held at Yeshiva University, New York, April 16-18, 1966. Indiana University Press. pp. 105–121.
^ Flock, A.; Lam, D. M. K. (1974). "Neurotransmitter synthesis in inner ear and lateral line sense organs". Nature. 249 (5453): 142–144. Bibcode:1974Natur.249..142F. doi:10.1038/249142a0. PMID 4151611. S2CID 275004.
^ ^a ^b Weeg, M. S.; Bass, A. H. (2002). "Frequency Response Properties of Lateral Line Superficial Neuromasts in a Vocal Fish, With Evidence for Acoustic Sensitivity". Journal of Neurophysiology. 88 (3): 1252–1262. doi:10.1152/jn.2002.88.3.1252. PMID 12205146.
^ ^a ^b Montgomery, J. C.; Bodznick, D. (1994). "An adaptive filter that cancels self-induced noise in the electrosensory and lateral line mechanosensory systems of fish". Neuroscience Letters. 174 (2): 145–148. doi:10.1016/0304-3940(94)90007-8. PMID 7970170. S2CID 15709516.
^ Maruska, K. P.; Tricas, T. C. (2009). "Central projections of octavolateralis nerves in the brain of a soniferous damselfish (Abudefduf abdominalis)". The Journal of Comparative Neurology. 512 (5): 628–650. doi:10.1002/cne.21923. PMID 19048640. S2CID 13604689.
^ Valera; Markov; Bijari; Randlett; Asgharsharghi; Baudoin; Ascoli; Portugues; Lopez-Schier (2021). "A neuronal blueprint for directional mechanosensation in larval zebrafish". Current Biology. 31 (7): 1463–1475. Bibcode:2021CBio...31E1463V. doi:10.1016/j.cub.2021.01.045. PMC 8044000. PMID 33545047.
^ ^a ^b King, Benedict; Hu, Yuzhi; Long, John A. (11 February 2018). "Electroreception in early vertebrates: survey, evidence and new information". Palaeontology. 61 (3): 325–358. Bibcode:2018Palgy..61..325K. doi:10.1111/pala.12346.
^ ^a ^b Bullock, T. H.; Bodznick, D. A.; Northcutt, R. G. (1983). "The phylogenetic distribution of electroreception: Evidence for convergent evolution of a primitive vertebrate sense modality" (PDF). Brain Research Reviews. 6 (1): 25–46. doi:10.1016/0165-0173(83)90003-6. hdl:2027.42/25137. PMID 6616267. S2CID 15603518.
^ Larsson, M. (2012). "Why do fish school?". Current Zoology. 58 (1): 116–128. doi:10.1093/czoolo/58.1.116.
^ Ziemer, Tim (2020). "Biology of the Auditory System". Psychoacoustic Music Sound Field Synthesis. Current Research in Systematic Musicology. Vol. 7. Cham: Springer. pp. 45–64. doi:10.1007/978-3-030-23033-3_3. ISBN 978-3-030-23033-3. S2CID 201209634.
^ Webb, J. (2011). "Hearing and Lateral Line: Lateral Line Structure". Encyclopedia of Fish Physiology. Elsevier. pp. 336–346. doi:10.1016/b978-0-12-374553-8.00010-1. ISBN 9780080923239.
^ ^a ^b "The origin of tetrapods". University of California Museum of Paleontology Berkeley. 28 April 2021. Retrieved 28 April 2023.
^ Budelmann, Bernd U.; Bleckmann, Horst (1988). "A lateral line analogue in cephalopods: Water waves generate microphonic potentials in the epidermal head lines of Sepia and Lolliguncula". Journal of Comparative Physiology A. 164 (1): 1–5. doi:10.1007/BF00612711. PMID 3236259. S2CID 8834051.

@@ Line 1: / Line 1: @@
+{{good article}}
 {{short description|Sensory system in fish}}
 [[File:Lateral line 01.JPG|thumb|upright=1.5|Oblique view of a [[goldfish]] (''Carassius auratus''), showing pored scales of the lateral line system]]
 The '''lateral line''', also called the '''lateral line organ''' ('''LLO'''), is a system of [[Sensory perception|sensory]] organs found in [[fish]], used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified [[epithelial cells]], known as [[hair cells]], which respond to displacement caused by motion and [[signal transduction|transduce]] these signals into electrical impulses via [[excitatory synapses]]. Lateral lines play an important role in [[School (fish)|schooling]] behavior, predation, and orientation.
-Early in the [[evolution of fish]], some of the receptive organs of the lateral line were modified to function as the [[electroreception|electroreceptors]] called [[ampullae of Lorenzini]].
+Early in the [[evolution of fish]], some of the sensory organs of the lateral line were modified to function as the [[electroreception|electroreceptors]] called [[ampullae of Lorenzini]]. The lateral line system is ancient and basal to the vertebrate clade, as it is found in fishes that diverged over 400 million years ago.
 == Function ==
-The lateral line system allows the detection of movement, vibration, and pressure gradients in the water surrounding an animal. It plays an essential role in orientation, predation, and [[School (fish)|fish schooling]] by providing spatial awareness and the ability to navigate in the environment.<ref name="Bleckmann Zelick 2009">{{Cite journal |last1=Bleckmann |first1=Horst |last2=Zelick |first2=Randy |date=2009-03-01 |title=Lateral line system of fish |journal=[[Integrative Zoology]] |volume=4 |issue=1 |pages=13–25 |doi=10.1111/j.1749-4877.2008.00131.x |pmid=21392273}}</ref> Analysis has shown that the lateral line system should be an effective passive sensing system able to discriminate between submerged obstacles by their shape.<ref>{{cite journal |last1=Bouffanais |first1=Roland |last2=Weymouth |first2=Gabriel D. |last3=Yue |first3=Dick K. P. |title=Hydrodynamic object recognition using pressure sensing |journal=[[Proceedings of the Royal Society]] A: Mathematical, Physical and Engineering Sciences |date=2 June 2010 |volume=467 |issue=2125 |pages=19–38 |doi=10.1098/rspa.2010.0095 |doi-access=free }}</ref><ref>{{cite journal |last1=Lakkam |first1=Sreetej |last2=Balamurali |first2=B. T. |last3=Bouffanais |first3=Roland |title=Hydrodynamic object identification with artificial neural models |journal=[[Scientific Reports]] |date=2 August 2019 |volume=9 |issue=1 |page=11242 |doi=10.1038/s41598-019-47747-8|pmid=31375742 |pmc=6677828 |arxiv=1903.00828 |bibcode=2019NatSR...911242L |doi-access=free }}</ref>
+The lateral line system allows the detection of movement, vibration, and pressure gradients in the water surrounding an animal. It plays an essential role in orientation, predation, and [[School (fish)|fish schooling]] by providing spatial awareness and the ability to navigate in the environment.<ref name="Bleckmann Zelick 2009">{{Cite journal |last1=Bleckmann |first1=Horst |last2=Zelick |first2=Randy |date=2009-03-01 |title=Lateral line system of fish |journal=[[Integrative Zoology]] |volume=4 |issue=1 |pages=13–25 |doi=10.1111/j.1749-4877.2008.00131.x |pmid=21392273 |doi-access=free}}</ref> Analysis has shown that the lateral line system should be an effective passive sensing system able to discriminate between submerged obstacles by their shape.<ref>{{cite journal |last1=Bouffanais |first1=Roland |last2=Weymouth |first2=Gabriel D. |last3=Yue |first3=Dick K. P. |title=Hydrodynamic object recognition using pressure sensing |journal=Proceedings of the Royal Society A: Mathematical, Physical and Engineering Sciences |date=2 June 2010 |volume=467 |issue=2125 |pages=19–38 |doi=10.1098/rspa.2010.0095 |s2cid=233793 |url=https://eprints.soton.ac.uk/349798/1/Bouffanais%25202011.pdf }}</ref><ref>{{cite journal |last1=Lakkam |first1=Sreetej |last2=Balamurali |first2=B. T. |last3=Bouffanais |first3=Roland |title=Hydrodynamic object identification with artificial neural models |journal=[[Scientific Reports]] |date=2 August 2019 |volume=9 |issue=1 |page=11242 |doi=10.1038/s41598-019-47747-8 |pmid=31375742 |pmc=6677828 |arxiv=1903.00828 |bibcode=2019NatSR...911242L |doi-access=free }}</ref> The lateral line allows fish to navigate and hunt in water with poor visibility.<ref name="Collin Marshall 2003">{{cite book |last1=Coombs |first1=Sheryl |last2=Braun |first2=Christopher B. |chapter=Information Processing by the Lateral Line System |chapter-url=https://link.springer.com/chapter/10.1007/978-0-387-22628-6_7 |editor1-last=Collin |editor1-first=Shaun P. |editor2-last=Marshall |editor2-first=N. Justin |title=Sensory Processing in Aquatic Environments |publisher=Springer Science & Business Media |year=2003 |isbn=978-0-387-95527-8 |pages=122–138 |doi=10.1007/978-0-387-22628-6_7 }}</ref>
-The lateral line system enables predatory fishes to detect vibrations made by their prey, and to orient towards the source to begin predatory action.<ref name="Coombs">{{cite journal |last1=Coombs |first1=S. I. |last2=Braun |first2=C. B. |last3=Donovan |first3=B. |title=The orienting response of Lake Michigan mottled sculpin is mediated by canal neuromasts |journal=Journal of Experimental Biology |date=2001 |volume=204 |issue=2 |pages=337–348 |doi=10.1242/jeb.204.2.337 |pmid=11136619 }}</ref> Blinded predatory fishes remain able to hunt, but not when lateral line function is inhibited by [[Cobalt(II) chloride|cobalt ions]].<ref>{{cite journal |last1=Karlsen |first1=H. E. |last2=Sand |first2=O. |year=1987 |title=Selective and Reversible Blocking of the Lateral Line in Freshwater Fish |journal=[[The Journal of Experimental Biology]] |volume=133 |issue=1 |pages=249–262 |doi=10.1242/jeb.133.1.249 }}</ref>
+The lateral line system enables predatory fishes to detect vibrations made by their prey, and to orient towards the source to begin predatory action.<ref name="Coombs">{{cite journal |last1=Coombs |first1=S. I. |last2=Braun |first2=C. B. |last3=Donovan |first3=B. |title=The orienting response of Lake Michigan mottled sculpin is mediated by canal neuromasts |journal=Journal of Experimental Biology |date=2001 |volume=204 |issue=2 |pages=337–348 |doi=10.1242/jeb.204.2.337 |pmid=11136619 }}</ref> Blinded predatory fishes remain able to hunt, but not when lateral line function is inhibited by [[Cobalt(II) chloride|cobalt ions]].<ref>{{cite journal |last1=Karlsen |first1=H. E. |last2=Sand |first2=O. |year=1987 |title=Selective and Reversible Blocking of the Lateral Line in Freshwater Fish |journal=[[The Journal of Experimental Biology]] |volume=133 |issue=1 |pages=249–262 |doi=10.1242/jeb.133.1.249 |doi-access=free }}</ref>
 The lateral line plays a role in fish schooling. Blinded ''[[Pollachius virens]]'' were able to integrate into a school, whereas fish with severed lateral lines could not.<ref>{{cite journal |last1=Pitcher |first1=T. |last2=Partridge |first2=B. |last3=Wardle |first3=C. |year=1976 |title=A blind fish can school |journal=Science |volume=194 |issue=4268|pages=963–965 |doi=10.1126/science.982056 |pmid=982056 |bibcode=1976Sci...194..963P }}</ref> It may have evolved further to allow fish to forage in dark caves. In Mexican blind cave fish, ''[[Mexican tetra|Astyanax mexicanus]]'', neuromasts in and around the [[Orbit (anatomy)|orbit of the eye]] are bigger and around twice as sensitive as those of surface-living fish.<ref>{{cite journal |last1=Yoshizawa |first1=Masato |last2=Jeffery |first2=William |last3=Van Netten |first3=Sietse |last4=McHenry |first4=Matthew |title=The sensitivity of lateral line receptors and their role in the behavior of Mexican blind cavefish, ''Astyanax mexicanus'' |journal=[[The Journal of Experimental Biology]] |date=2014 |volume=217 |issue=6 |pages=886–895 |pmid=24265419 |doi=10.1242/jeb.094599 |pmc=3951362 }}</ref>
-One function of schooling may be to confuse the lateral line of [[Predation|predatory]] fishes. A single prey fish creates a rather simple [[particle velocity]] pattern, whereas the pressure gradients of many closely swimming (schooling) prey fish overlap, creating a complex pattern. This makes it difficult for predatory fishes to identify individual prey through lateral line perception.<ref name="Larsson 2009">{{cite journal |last=Larsson |first=M. |year=2009 |title=Possible functions of the octavolateralis system in fish schooling |journal=Fish and Fisheries |volume=10 |issue=3 |pages=344-355 |doi=10.1111/j.1467-2979.2009.00330.x}}</ref>
+One function of schooling may be to confuse the lateral line of [[Predation|predatory]] fishes. A single prey fish creates a simple [[particle velocity]] pattern, whereas the pressure gradients of many closely swimming (schooling) prey fish overlap, creating a complex pattern. This makes it difficult for predatory fishes to identify individual prey through lateral line perception.<ref name="Larsson 2009">{{cite journal |last=Larsson |first=M. |year=2009 |title=Possible functions of the octavolateralis system in fish schooling |journal=Fish and Fisheries |volume=10 |issue=3 |pages=344–355 |doi=10.1111/j.1467-2979.2009.00330.x|bibcode=2009AqFF...10..344L |url=https://zenodo.org/record/895913 }}</ref>
 == Anatomy ==
-[[File:LateralLine Organ.jpg|thumb|center|upright=3|The lateral line system consists of canals that run along a fish's body, connecting rows of openings through the skin and scales to the water outside the body. Small sense organs, neuromasts, are positioned both at intervals along the canals, and on the surface of the body.]]
+[[File:LateralLine Organ.jpg|thumb|center|upright=3|The lateral line system consists of canals that run along a fish's body, connecting rows of openings (pores) through the skin and scales to the water outside the body. Small sense organs, neuromasts (inset), are positioned both at intervals along the canals, and on the surface of the body. Each hair cell contains a bundle of sensory hairs.]]
 <!--[[File:RutilusRutilusScalesLateralLine.JPG|thumb|Some scales of the lateral line of a ''[[Rutilus rutilus]]'']]-->
+[[File:Gasterosteus aculeatus with stained neuromasts.png|thumb|A [[Gasterosteus aculeatus|three-spined stickleback]] with stained neuromasts<ref>{{cite journal |last1=Wark |first1=Abigail R. |last2=Mills |first2=Margaret G. |last3=Dang |first3=Lam-Ha |last4=Chan |first4=Yingguang Frank |last5=Jones |first5=Felicity C |last6=Brady |first6=Shannon D. |last7=Absher |first7=Devin M |last8=Grimwood |first8=Jane |last9=Schmutz |first9=Jeremy |last10=Myers |first10=Richard M. |last11=Kingsley |first11=David M. |last12=Peichel |first12=Catherine L. |display-authors=3 |year=2012 |title=Genetic Architecture of Variation in the Lateral Line Sensory System of Threespine Sticklebacks |journal=G3: Genes, Genomes, Genetics |volume=2 |issue=9 |pages=1047–1056 |doi=10.1534/g3.112.003079 |pmid=22973542 |pmc=3429919 }}</ref>]]
-[[File:Gasterosteus aculeatus with stained neuromasts.png|thumb|A ''[[Gasterosteus aculeatus|three-spined stickleback]]'' with stained neuromasts]]
-Lateral lines are usually visible as faint lines of pores running lengthwise down each side of a fish's body.<ref name="Bleckmann Zelick 2009"/> The functional units of the lateral line are the neuromasts, discrete [[mechanoreception|mechanoreceptive]] organs that sense movement in water. There are two main varieties: canal neuromasts and superficial neuromasts. Superficial neuromasts are on the surface of the body, while canal neuromasts are along the lateral lines in subdermal, fluid-filled canals. Each neuromast consists of receptive [[hair cells]] whose tips are covered by a flexible jellylike cupula. Hair cells typically possess both [[glutamatergic]] [[afferent nerve fiber|afferent connections]] and [[acetylcholine|cholinergic]] [[efferent nerve fiber|efferent connections]].<ref>{{cite journal |last1=Russell |first1=I. J. |year=1971 |title=The Pharmacology of Efferent Synapses in the Lateral-Line System of Xenopus Laevis |journal=Journal of Experimental Biology |volume=54 |issue=3 |pages=643–659 |doi=10.1242/jeb.54.3.643 |pmid=4326222 }}</ref> The receptive hair cells are modified [[epithelial cells]]; they typically possess bundles of 40-50 [[microvilli]] "hairs" which function as the mechanoreceptors.<ref name="Flock 1967">{{cite book |last=Flock |first=Å. |year=1967 |chapter=Ultrastructure and function in the lateral line organs |title=Lateral Line Detectors; proceedings of a conference held at Yeshiva University, New York, April 16-18, 1966 |editor=P. Cahn |publisher=[[Indiana University Press]] |pages=163–197 }}</ref> These bundles are organized in rough "staircases" of hairs of increasing length order.<ref name="Peach">{{cite journal |last1=Peach |first1=M. B. |last2=Rouse |first2=G. W. |year=2000 |title=The Morphology of the Pit Organs and Lateral Line Canal Neuromasts of ''Mustelus Antarcticus'' (Chondrichthyes: Triakidae) |journal=Journal of the Marine Biological Association of the United Kingdom |volume=80 |issue=1 |pages=155–162 |doi=10.1017/s0025315499001678 |s2cid=85963954 }}</ref>{{-}}
+Lateral lines are usually visible as faint lines of pores running along each side of a fish's body.<ref name="Bleckmann Zelick 2009"/> The functional units of the lateral line are the neuromasts, discrete [[mechanoreception|mechanoreceptive]] organs that sense movement in water. There are two main varieties: canal neuromasts and superficial neuromasts. Superficial neuromasts are on the surface of the body, while canal neuromasts are along the lateral lines in subdermal, fluid-filled canals. Each neuromast consists of receptive [[hair cells]] whose tips are covered by a flexible jellylike cupula. Hair cells typically possess both [[glutamatergic]] [[afferent nerve fiber|afferent connections]] and [[acetylcholine|cholinergic]] [[efferent nerve fiber|efferent connections]].<ref>{{cite journal |last1=Russell |first1=I. J. |year=1971 |title=The Pharmacology of Efferent Synapses in the Lateral-Line System of Xenopus Laevis |journal=Journal of Experimental Biology |volume=54 |issue=3 |pages=643–659 |doi=10.1242/jeb.54.3.643 |pmid=4326222 |doi-access= }}</ref><ref name="Flock 1967"/> The receptive hair cells are modified [[epithelial cells]]; they typically possess bundles of 40–50 [[microvilli]] "hairs" which function as the mechanoreceptors.<ref name="Flock 1967">{{cite book |last=Flock |first=Å. |year=1967 |chapter=Ultrastructure and function in the lateral line organs |title=Lateral Line Detectors; proceedings of a conference held at Yeshiva University, New York, April 16-18, 1966 |editor=P. Cahn |publisher=[[Indiana University Press]] |pages=163–197 }}</ref> Within each bundle, the hairs are organized in a rough "staircase" from shortest to longest.<ref name="Peach">{{cite journal |last1=Peach |first1=M. B. |last2=Rouse |first2=G. W. |year=2000 |title=The Morphology of the Pit Organs and Lateral Line Canal Neuromasts of ''Mustelus Antarcticus'' (Chondrichthyes: Triakidae) |journal=Journal of the Marine Biological Association of the United Kingdom |volume=80 |issue=1 |pages=155–162 |doi=10.1017/s0025315499001678 |bibcode=2000JMBUK..80..155P |s2cid=85963954 }}</ref>{{-}}
 == Signal transduction ==
-[[File:Lateral line rate coding.svg|thumb|center|upright=2.5|Lateral line rate coding indicates direction and sometimes strength of stimulus.]]
+[[File:Lateral line rate coding.svg|thumb|center|upright=2.5|Lateral line rate coding indicates direction and sometimes strength of stimulus. One neuromast shown.]]
-The hair cells are stimulated by the deflection of their hair bundles in the direction of the tallest "hairs" or [[stereocilia]]. The deflection allows [[Ion|cations]] to enter through a [[Mechanosensitive channels|mechanically gated channel]], causing depolarization of the hair cell. This depolarization opens [[Cav1.3|Ca<big><sub>v</sub></big>1.3]] calcium channels in the [[Epithelial polarity|basolateral membrane]].<ref>{{Cite journal |last=Baker |first=Clare |date=June 11, 2018 |title=Insights into Electroreceptor Development and Evolution from Molecular Comparisons with Hair Cells |url=https://academic.oup.com/icb/article/58/2/329/5020173?searchresult=1#186667130 |url-status=live |journal=Integrative and Comparative Biology |volume=58 |issue=2 |pages=329–340 |doi=10.1093/icb/icy037 |pmid=29846597 |pmc=6927855}}</ref>
+The hair cells are stimulated by the deflection of their hair bundles in the direction of the tallest "hairs" or [[stereocilia]]. The deflection allows [[Ion|cations]] to enter through a [[Mechanosensitive channels|mechanically gated channel]], causing depolarization or hyperpolarization of the hair cell. Depolarization opens [[Cav1.3|Ca<big><sub>v</sub></big>1.3]] calcium channels in the [[Epithelial polarity|basolateral membrane]].<ref>{{Cite journal |last=Baker |first=Clare |date=June 11, 2018 |title=Insights into Electroreceptor Development and Evolution from Molecular Comparisons with Hair Cells |url=https://academic.oup.com/icb/article/58/2/329/5020173?searchresult=1#186667130 |journal=Integrative and Comparative Biology |volume=58 |issue=2 |pages=329–340 |doi=10.1093/icb/icy037 |pmid=29846597 |pmc=6927855}}</ref>
 [[Hair cells]] use a system of [[signal transduction|transduction]] with [[rate coding]] to transmit the directionality of a stimulus. The hair cells produce a constant, tonic rate of firing. As mechanical motion is transmitted through water to the neuromast, the cupula bends and is displaced according to the strength of the stimulus. This results in a shift in the cell's ionic permeability. Deflection towards the longest hair results in [[depolarization]] of the hair cell, increased neurotransmitter release at the excitatory afferent synapse, and a higher rate of [[signal transduction]]. Deflection towards the shorter hair has the opposite effect, [[Hyperpolarization (biology)|hyperpolarizing]] the hair cell and producing a decreased rate of neurotransmitter release. These electrical impulses are then transmitted along afferent lateral neurons to the brain.<ref name="Flock 1967"/>
-While both varieties of neuromasts utilize this method of transduction, their specialized organization gives them different mechanoreceptive capacities. Superficial organs are exposed more directly to the external environment. The organization of the bundles within their organs is seemingly haphazard, incorporating various shapes and sizes of [[microvilli]] within bundles. This suggests coarse but wide-ranging detection.<ref name="Peach" /> In contrast, the structure of canal organs allow canal neuromasts more sophisticated mechanoreception, such as of pressure differentials. As current moves across the pores of a canal, a pressure differential is created, inducing a flow in the canal fluid. This moves the cupula of the hair cells in the canal, resulting in a deflection of the hairs in the direction of the flow.<ref>{{cite book |last=Kuiper |first=J. W. |year=1967 |chapter=Frequency Characteristics and Functional Significance of the Lateral Line Organ |title=Lateral Line Detectors; proceedings of a conference held at Yeshiva University, New York, April 16-18, 1966 |editor=P. Cahn |publisher=[[Indiana University Press]] |pages=105–121}}</ref>
+While both varieties of neuromasts utilize this method of transduction, their specialized organization gives them different mechanoreceptive capacities. Superficial organs are exposed more directly to the external environment. The organization of the bundles within their organs is seemingly haphazard, incorporating various shapes and sizes of [[microvilli]] within bundles. This suggests coarse but wide-ranging detection.<ref name="Peach" /> In contrast, the structure of canal organs allow canal neuromasts more sophisticated mechanoreception, such as of pressure differentials. As current moves across the pores, a pressure differential is created, inducing a flow in the canal fluid. This moves the cupulae of the neuromasts in the canal, resulting in a deflection of the hairs in the direction of the flow.<ref>{{cite book |last=Kuiper |first=J. W. |year=1967 |chapter=Frequency Characteristics and Functional Significance of the Lateral Line Organ |title=Lateral Line Detectors; proceedings of a conference held at Yeshiva University, New York, April 16-18, 1966 |editor=P. Cahn |publisher=[[Indiana University Press]] |pages=105–121}}</ref>
 == Electrophysiology ==
-The mechanoreceptive hair cells of the lateral line structure are integrated into more complex circuits through their afferent and efferent connections. The synapses that directly participate in the transduction of mechanical information are excitatory afferent connections that utilize [[glutamate]].<ref>{{cite journal |last1=Flock |first1=A. |last2=Lam |first2=D. M. K. |year=1974 |title=Neurotransmitter synthesis in inner ear and lateral line sense organs |journal=[[Nature (journal)|Nature]] |volume=249 |issue=5453 |pages=142–144 |doi=10.1038/249142a0 |pmid=4151611 |bibcode=1974Natur.249..142F |s2cid=275004 }}</ref> Species vary in their neuromast and afferent connections, providing differing mechanoreceptive properties. For instance, the superficial neuromasts of the [[midshipman fish]], ''Porichthys notatus'', are sensitive to specific stimulation frequencies.<ref name="Weeg">{{cite journal |last1=Weeg |first1=M. S. |last2=Bass |first2=A. H. |year=2002 |title=Frequency Response Properties of Lateral Line Superficial Neuromasts in a Vocal Fish, With Evidence for Acoustic Sensitivity |journal=[[Journal of Neurophysiology]] |volume=88 |issue=3 |pages=1252–1262 |doi=10.1152/jn.2002.88.3.1252 |pmid=12205146 }}</ref> One variety is attuned to collect information about acceleration, at stimulation frequencies between 30 and 200 Hz. The other type obtains information about velocity, and is most receptive to stimulation below 30 Hz.<ref name="Weeg" />
+The mechanoreceptive hair cells of the lateral line structure are integrated into more complex circuits through their afferent and efferent connections. The synapses that directly participate in the transduction of mechanical information are excitatory afferent connections that utilize [[glutamate]].<ref>{{cite journal |last1=Flock |first1=A. |last2=Lam |first2=D. M. K. |year=1974 |title=Neurotransmitter synthesis in inner ear and lateral line sense organs |journal=[[Nature (journal)|Nature]] |volume=249 |issue=5453 |pages=142–144 |doi=10.1038/249142a0 |pmid=4151611 |bibcode=1974Natur.249..142F |s2cid=275004 }}</ref> Species vary in their neuromast and afferent connections, providing differing mechanoreceptive properties. For instance, the superficial neuromasts of the [[midshipman fish]], ''Porichthys notatus'', are sensitive to specific stimulation frequencies.<ref name="Weeg">{{cite journal |last1=Weeg |first1=M. S. |last2=Bass |first2=A. H. |year=2002 |title=Frequency Response Properties of Lateral Line Superficial Neuromasts in a Vocal Fish, With Evidence for Acoustic Sensitivity |journal=[[Journal of Neurophysiology]] |volume=88 |issue=3 |pages=1252–1262 |doi=10.1152/jn.2002.88.3.1252 |pmid=12205146 |doi-access= }}</ref> One variety is attuned to collect information about acceleration, at stimulation frequencies between 30 and 200 Hz. The other type obtains information about velocity, and is most receptive to stimulation below 30 Hz.<ref name="Weeg" />
 [[File:Lateral line circuits.svg|thumb|center|upright=3|The motion detection system in fish works despite "noise" created by the fish itself. The brain copies the efferent commands it gives to the swimming muscles to the lateral line, effectively suppressing swimming noise and revealing small signals from the environment, such as from prey.<ref name="Montgomery Bodznick 1994"/>]]
-The efferent synapses to hair cells are inhibitory and use [[acetylcholine]] as a transmitter. They are crucial participants in a [[corollary discharge]] system designed to limit self-generated interference. When a fish moves, it creates disturbances in the water that could be detected by the lateral line system, potentially interfering with the detection of other biologically relevant signals. To prevent this, an efferent signal is sent to the hair cell upon motor action, resulting in inhibition which counteracts the excitation resulting from reception of the self-generated stimulation. This allows the fish to retain perception of motion stimuli without interference created by its own movements.<ref name="Montgomery Bodznick 1994">{{cite journal |last1=Montgomery |first1=J. C. |last2=Bodznick |first2=D. |year=1994 |title=An adaptive filter that cancels self-induced noise in the electrosensory and lateral line mechanosensory systems of fish |journal=[[Neuroscience Letters]] |volume=174 |issue=2 |pages=145–148 |doi=10.1016/0304-3940(94)90007-8 |pmid=7970170 |s2cid=15709516 }}</ref>
+The efferent synapses to hair cells are inhibitory and use [[acetylcholine]] as a transmitter. They are crucial participants in a [[corollary discharge]] system designed to limit self-generated interference. When a fish moves, it creates disturbances in the water that could be detected by the lateral line system, potentially interfering with the detection of other biologically relevant signals. To prevent this, an efferent signal is sent to the hair cell upon motor action, resulting in inhibition which counteracts the excitation resulting from reception of the self-generated stimulation. This allows the fish to detect external stimuli without interference from its own movements.<ref name="Montgomery Bodznick 1994">{{cite journal |last1=Montgomery |first1=J. C. |last2=Bodznick |first2=D. |year=1994 |title=An adaptive filter that cancels self-induced noise in the electrosensory and lateral line mechanosensory systems of fish |journal=[[Neuroscience Letters]] |volume=174 |issue=2 |pages=145–148 |doi=10.1016/0304-3940(94)90007-8 |pmid=7970170 |s2cid=15709516 }}</ref>
-Signals from the hair cells are transmitted along lateral neurons to the brain. The area where these signals most often terminate is the medial octavolateralis nucleus (MON), which probably processes and integrates mechanoreceptive information.<ref name="Maruska">{{cite journal |last1=Maruska |first1=K. P. |last2=Tricas |first2=T. C. |year=2009 |title=Central projections of octavolateralis nerves in the brain of a soniferous damselfish (Abudefduf abdominalis) |journal=The Journal of Comparative Neurology |volume=512 |issue=5|pages=628–650 |doi=10.1002/cne.21923 |pmid=19048640 |s2cid=13604689 }}</ref> The deep MON contains distinct layers of basilar and non-basilar crest cells, suggesting computational pathways analogous to the electrosensory lateral line lobe of [[electric fish]]. The MON is likely involved in the integration of excitatory and inhibitory parallel circuits to interpret mechanoreceptive information.<ref>{{cite journal |last1=Valera |last2=Markov |last3=Bijari |last4=Randlett |last5=Asgharsharghi  |last6=Baudoin |last7=Ascoli |last8=Portugues |last9=Lopez-Schier |year=2021 |title=A neuronal blueprint for directional mechanosensation in larval zebrafish |journal=[[Current Biology]] |volume=31 |issue=7 |pages=1463–1475 |doi=10.1016/j.cub.2021.01.045 |pmc=8044000 |pmid=33545047 }}</ref>
+Signals from the hair cells are transmitted along lateral neurons to the brain. The area where these signals most often terminate is the medial octavolateralis nucleus (MON), which probably processes and integrates mechanoreceptive information.<ref name="Maruska">{{cite journal |last1=Maruska |first1=K. P. |last2=Tricas |first2=T. C. |year=2009 |title=Central projections of octavolateralis nerves in the brain of a soniferous damselfish (Abudefduf abdominalis) |journal=The Journal of Comparative Neurology |volume=512 |issue=5|pages=628–650 |doi=10.1002/cne.21923 |pmid=19048640 |s2cid=13604689 }}</ref> The deep MON contains distinct layers of basilar and non-basilar crest cells, suggesting computational pathways analogous to the electrosensory lateral line lobe of [[electric fish]]. The MON is likely involved in the integration of excitatory and inhibitory parallel circuits to interpret mechanoreceptive information.<ref>{{cite journal |last1=Valera |last2=Markov |last3=Bijari |last4=Randlett |last5=Asgharsharghi  |last6=Baudoin |last7=Ascoli |last8=Portugues |last9=Lopez-Schier |year=2021 |title=A neuronal blueprint for directional mechanosensation in larval zebrafish |journal=[[Current Biology]] |volume=31 |issue=7 |pages=1463–1475 |doi=10.1016/j.cub.2021.01.045 |pmc=8044000 |pmid=33545047 |bibcode=2021CBio...31E1463V }}</ref>
 == Evolution ==
@@ Line 52: / Line 53: @@
 The use of [[mechanosensation|mechanosensitive]] hairs is homologous to the functioning of hair cells in the [[auditory system|auditory]] and [[vestibular system]]s, indicating a close link between these systems.<ref name="Flock 1967"/> Due to many overlapping functions and their great similarity in ultrastructure and development, the lateral line system and the [[inner ear]] of fish are often grouped together as the octavolateralis system (OLS).<ref>{{cite journal |last1=Larsson |first1=M. |year=2012 |title=Why do fish school? |journal=Current Zoology |volume=58 |issue=1 |pages=116–128 |doi=10.1093/czoolo/58.1.116 |doi-access=free }}</ref> Here, the lateral line system detects particle velocities and accelerations with frequencies below 100&nbsp;Hz. These low frequencies create large wavelengths, which induce strong particle accelerations in the [[Near and far field|near field]] of swimming fish that do not radiate into the far field as acoustic waves due to an [[acoustic short circuit]]. The auditory system detects pressure fluctuations with frequencies above 100&nbsp;Hz that propagate to the far field as waves.<ref name="biologyllas">{{cite book |last1=Ziemer |first1=Tim |title=Psychoacoustic Music Sound Field Synthesis |volume=7 |chapter=Biology of the Auditory System |date=2020 |publisher=Springer |location=Cham |isbn=978-3-030-23033-3 |pages=45–64 |doi=10.1007/978-3-030-23033-3_3 |series=Current Research in Systematic Musicology |s2cid=201209634 }}</ref>
+The lateral line system is ancient and basal to the [[vertebrate]] clade; it is found in groups of fishes that diverged over 400 million years ago, including the [[lamprey]]s, [[Chondrichthyes|cartilaginous fishes]], and [[Osteichthyes|bony fishes]].<ref name="Webb 2011 pp. 336–346">{{cite book |last=Webb |first=J. |title=Encyclopedia of Fish Physiology |chapter=Hearing and Lateral Line: Lateral Line Structure |publisher=Elsevier |year=2011 |doi=10.1016/b978-0-12-374553-8.00010-1 |pages=336–346|isbn=9780080923239 }}</ref><ref name="UCMP"/> Most [[amphibian]] [[larva]]e and some fully aquatic adult amphibians possess mechanosensitive systems comparable to the lateral line.<ref>{{cite journal |doi=10.1007/BF00612711 |title=A lateral line analogue in cephalopods: Water waves generate microphonic potentials in the epidermal head lines of ''Sepia'' and ''Lolliguncula'' |year=1988 |last1=Budelmann |first1=Bernd U. |last2=Bleckmann |first2=Horst |journal=Journal of Comparative Physiology A |volume=164 |pages=1–5 |pmid=3236259 |issue=1 |s2cid=8834051 }}</ref> The terrestrial [[Tetrapod|tetrapods]] have secondarily lost their lateral line organs, which are ineffective when not submerged.<ref name="UCMP">{{cite web |title=The origin of tetrapods |date=28 April 2021 |url=https://evolution.berkeley.edu/what-are-evograms/the-origin-of-tetrapods/ |publisher=University of California Museum of Paleontology Berkeley |access-date=28 April 2023}}</ref>
-The [[Electroreception and electrogenesis|electroreceptive]] organs, called [[ampullae of Lorenzini]], appearing as pits in the skin of sharks and some other fishes, evolved from the lateral line organ. It is basal to the [[Gnathostomata|jawed fishes]].<ref name="King Hu Long">{{cite journal |last1=King |first1=Benedict |last2=Hu |first2=Yuzhi |last3=Long |first3=John A. |date=11 February 2018 |title=Electroreception in early vertebrates: survey, evidence and new information |journal=[[Palaeontology (journal)|Palaeontology]] |volume=61 |issue=3 |pages=325–358 |doi=10.1111/pala.12346 |doi-access=free }}</ref> Passive electroreception using ampullae is an [[Primitive (phylogenetics)|ancestral trait]] in the vertebrates, meaning that it was present in their [[Most recent common ancestor|last common ancestor]].<ref name="Bullock Bodznick Northcutt 1983">{{cite journal |last1=Bullock |first1=T. H. |last2=Bodznick |first2=D. A. |last3=Northcutt |first3=R. G. |date=1983 |title=The phylogenetic distribution of electroreception: Evidence for convergent evolution of a primitive vertebrate sense modality |journal=[[Brain Research Reviews]] |volume=6 |issue=1 |pages=25–46 |doi=10.1016/0165-0173(83)90003-6 |pmid=6616267 |hdl=2027.42/25137 |s2cid=15603518 |url=https://deepblue.lib.umich.edu/bitstream/2027.42/25137/1/0000573.pdf |hdl-access=free }}</ref>
+The [[Electroreception and electrogenesis|electroreceptive]] organs, called [[ampullae of Lorenzini]], appearing as pits in the skin of sharks and some other fishes, evolved from the lateral line organ.<ref name="King Hu Long">{{cite journal |last1=King |first1=Benedict |last2=Hu |first2=Yuzhi |last3=Long |first3=John A. |date=11 February 2018 |title=Electroreception in early vertebrates: survey, evidence and new information |journal=[[Palaeontology (journal)|Palaeontology]] |volume=61 |issue=3 |pages=325–358 |doi=10.1111/pala.12346 |bibcode=2018Palgy..61..325K |doi-access=free }}</ref> Passive electroreception using ampullae is an [[Primitive (phylogenetics)|ancestral trait]] in the vertebrates, meaning that it was present in their [[Most recent common ancestor|last common ancestor]].<ref name="Bullock Bodznick Northcutt 1983">{{cite journal |last1=Bullock |first1=T. H. |last2=Bodznick |first2=D. A. |last3=Northcutt |first3=R. G. |date=1983 |title=The phylogenetic distribution of electroreception: Evidence for convergent evolution of a primitive vertebrate sense modality |journal=[[Brain Research Reviews]] |volume=6 |issue=1 |pages=25–46 |doi=10.1016/0165-0173(83)90003-6 |pmid=6616267 |hdl=2027.42/25137 |s2cid=15603518 |url=https://deepblue.lib.umich.edu/bitstream/2027.42/25137/1/0000573.pdf |hdl-access=free }}</ref>
-The lateral line system is ancient and basal to the [[vertebrate]] clade; it is found in groups of fishes that diverged over 400 million years ago, including the [[lamprey]]s, [[Chondrichthyes|cartilaginous fishes]], and [[Osteichthyes|bony fishes]].<ref name="Webb 2011 pp. 336–346">{{cite book |last=Webb |first=J. |title=Encyclopedia of Fish Physiology |chapter=Hearing and Lateral Line: Lateral Line Structure |publisher=Elsevier |year=2011 |doi=10.1016/b978-0-12-374553-8.00010-1 |pages=336–346}}</ref><ref name="UCMP"/> Most [[amphibian]] [[larva]]e and some fully aquatic adult amphibians possess mechanosensitive systems comparable to the lateral line.<ref>{{cite journal |doi=10.1007/BF00612711 |title=A lateral line analogue in cephalopods: Water waves generate microphonic potentials in the epidermal head lines of ''Sepia'' and ''Lolliguncula'' |year=1988 |last1=Budelmann |first1=Bernd U. |last2=Bleckmann |first2=Horst |journal=Journal of Comparative Physiology A |volume=164 |pages=1–5 |pmid=3236259 |issue=1 |s2cid=8834051 }}</ref> The terrestrial [[Tetrapod|tetrapods]] have secondarily lost their lateral line organs, which are ineffective when not submerged.<ref name="UCMP">{{cite web |title=The origin of tetrapods |url=https://evolution.berkeley.edu/what-are-evograms/the-origin-of-tetrapods/ |publisher=University of California Museum of Paleontology Berkeley |access-date=28 April 2023}}</ref>
 {{clade
@@ Line 62: / Line 63: @@
    |label1=[[Lamprey]]s
    |sublabel1=>440 [[myr|mya]]
-   |1=[[File:Petromyzon marinus.jpg|100px]]
+   |1=<span style="{{MirrorH}}">[[File:Petromyzon marinus.jpg|100px]]</span>
    |label2=[[Gnathostomata|Jawed fishes]]
    |sublabel2=''[[Ampullae of Lorenzini]]''
    |2={{clade
       |1={{clade
-         |1=[[Chondrichthyes|Cartilaginous fishes]] <span style="{{MirrorH}}">[[File:White shark (Duane Raver).png|100px]]
+         |1=[[Chondrichthyes|Cartilaginous fishes]] [[File:White shark (Duane Raver).png|100px]]
          |sublabel1=430 [[myr|mya]]
          |label2=[[Osteichthyes|Bony fishes]]
@@ Line 78: / Line 79: @@
                |2={{clade
                   |1={{clade
-                     |1=[[Lungfish]] [[File:Barramunda coloured.jpg|70px]]
+                     |1=[[Lungfish]] [[File:Chinle fish Arganodus cropped cropped.png|70px]]
                      |2={{clade
                         |label1=[[Amphibian]]s
                         |sublabel1=(as larvae; some as adults)
-                        |1= [[File:Aquatic life (1916-1917) (19559021800) (cropped).jpg|80px]]
+                        |1= [[File:Salamandra salamandra (white background).jpg|80px]]
                         |2=Other [[tetrapod]]s
                         |sublabel2='''''(no lateral line)'''''

v t e Electric fish
Physiology	Electroreception and electrogenesis Jamming avoidance response
Anatomy	Electric organ Electroreceptors Ampullae of Lorenzini Knollenorgan
Groups	Gymnarchidae (African knifefish) Gymnotiformes (S. American knifefishes) Electrophorus (electric eels) Mormyridae (elephantfishes) Rajiformes (skates) Malapteruridae (electric catfishes) Torpediniformes (electric rays) Uranoscopidae (stargazers)
Related	History of bioelectricity Lateral line Magnetoreception

v t e Neuroethology
Concepts	Feedforward Coincidence detector Umwelt Instinct Feature detection Central pattern generator (CPG) NMDA receptor Lateral inhibition Fixed action pattern Krogh's Principle Hebbian theory Anti-Hebbian learning Sound localization Ultrasound avoidance in insects
People	Theodore Holmes Bullock Walter Heiligenberg Niko Tinbergen Konrad Lorenz Donald Griffin Donald Kennedy Karl von Frisch Erich von Holst Jörg-Peter Ewert Franz Huber Bernhard Hassenstein Werner E. Reichardt Eric Knudsen Eric Kandel Nobuo Suga Masakazu Konishi Fernando Nottebohm
Methods	Patch clamp Slice preparation
Systems	Animal echolocation Waggle dance Jamming avoidance response Vision in toads Frog hearing and communication Infrared sensing in snakes Caridoid escape reaction Vocal learning Surface wave detection Electroreception Mechanoreception Lateral line
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