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User:Amhwarren/Cleaner fish

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Two bluestreak cleaner wrasses removing dead skin and external parasites from the potato grouper.
Video of bluestreak cleaner wrasse cleaning the gills of an elongate surgeonfish.

Cleaner fish are fish that show a specialist feeding strategy[1] by providing a service to other species, referred to as clients[2], by removing dead skin, ectoparasites and infected tissue from the surface or gill chambers[2]. This example of cleaning symbiosis represents mutualism and cooperation behaviour[3], an ecological interaction that benefits both parties involved. However, the cleaner fish may consume mucus or tissue, thus creating a form of parasitism[4] called cheating. The client animal is typically a heterospecific fish [3], But can also involve aquatic reptiles (sea turtles and marine iguana), mammals (manatees and whales) or octopuses.[5][6][7] A wide variety of fish including wrasse, cichlids, catfish, pipefish, lumpsuckers and gobies display cleaning behaviors across the globe in fresh, brackish, and marine waters but specifically concentrated in the tropics due to high parasite density[2]. Similar behavior is found in other groups of animals, such as cleaner shrimps.

There are two types of cleaner fish, obligate full time cleaners and facultative part time cleaners [1] where different strategies occur based on resources and local abundance of fish.[1] Cleaning behaviour takes place in pelagic waters as well as designated locations called cleaner stations[8]. Cleaner fish interaction durations and memories of reoccurring clients are influenced by the neuroendocrine system of the fish, involving hormones Arginine Vasotocin, Isotocin and serotonin[3].

Conspicuous coloration is a method used by some cleaner fish, where they often displaying a brilliant blue stripe that spans the length of the body.[9] Other species of fish, called mimics, imitate the behavior and phenotype of cleaner fish to gain access to client fish tissue.

The specialized feeding behaviour of cleaner fish has become a valuable resource in salmon aquaculture in Atlantic Canada, Scotland, Iceland and Norway[10] for prevention of sea lice out breaks[2] which is beneficial to the economy and environment by minimizing use of chemical delousers. Specifically cultured for this job are lumpfish (Cyclopterus lumpus) and ballan wrasse (Labrus bergeylta)[11]. The most common parasites that cleaner fish feed on are gnathiidae and copepod species[1].

Diversity and Examples edit

Marine Fishes edit

The following is a selection of few of the many marine cleaner species.

Commonly studied cleaner fish are the cleaner wrasses of the genus Labroides found on coral reefs in the Indian Ocean and Pacific Ocean [12].

Neon gobies of the genera Gobiosoma and Elacatinus provide a cleaning service similar to the cleaner wrasses, though this time on reefs in the Western Atlantic, providing a good example of convergent evolution[13] of the cleaning behaviour.

Lumpfish who are utilized as salmonid cleaner fish in aquaculture, but it is unknown if they display cleaning behaviour on salmon in the wild.[14]

Brackish Freshwater Fish edit

Brackish water refers to aquatic environments that have a salinity in between salt and fresh water systems. Cleaning symbiosis has also been observed in these areas between two brackish water cichlids of the genus Etroplus from South Asia. The small species Etroplus maculatus is the cleaner fish, and the much larger Etroplus suratensis is the host that receives the cleaning service[16].

Freshwater Fish edit

Cleaning has infrequently been observed in fresh waters compared to marine waters. This is possibly related to fewer observers (such as divers) in the former habitat compared to the latter[17]. One of the few known examples of cleaning is juvenile striped Raphael catfish cleaning the piscivorous Hoplias cf. malabaricus. In public aquariums, Synaptolaemus headstanders have been seen cleaning larger fish[18].

Mechanisms edit

Facultative Cleaner Fish edit

A facultative cleaner fish does not rely solely on specialized cleaning behaviour for nutrient consumption[2]. Facultative cleaners can be further divided by stationary vs. wandering facultative cleaners[1] .Facultative cleaners may display cleaning behaviour through their whole life history or solely during juvenile stages for additional nutrients during rapid growth[1][2]. Examples of facultative cleaners are commonly wrasse species such as the blue headed wrasse, noronha wrasse (Thalassoma noronhanum) and goldsinny wrasse (Ctenolabrus rupestris), sharp nose sea perch in Californian waters,[2] and the lumpfish (Cyclopterus lumpus).

Using the example of the blue wrasse from Caribbean waters, their alternative feeding strategy is described as being a generalist forager, meaning they eat a wide variety of smaller aquatic organisms based on availability[1]. When displaying cleaning behaviour, it has been noted that the blue wrasse inspects potential clients and only feeds on some, implying that the wrasse is seeking out a particular type of parasite as diet supplementation. It has also been quantified that the blue wrasse foraging behaviour does not change proportionate to cleaning opportunities, again promoting the idea that the cleaning behaviour in this facultative fish is for diet supplementation and not out of necessity[1].

Obligate Cleaner Fish edit

An obligate cleaner fish relies solely on specialized cleaning behaviour for nutrient consumption[2]. Therefore, obligate cleaners have a higher output of cleaning on a wider range of parasites in comparison to facultative fish. To maximize nutrient consumption, obligate cleaners utilize a higher proportion of cleaning stations[1]. Obligate cleaner fish may also be divided by stationary and wandering and these life history choice are made based on the amount of interspecific competition from other obligate cleaners in the area [19] An example of an obligate cleaner is the shark nose goby (Elacatinus evelynae) in the Caribbean Reef, where it has been observed to perform up to 110 cleanings per day.[1]

 
Client Mulloidichthys flavolineatus at a cleaning station.

Cleaner Stations edit

Cleaning stations are a strategy used by some cleaner fish where clients congregate and perform specific movements to attract the attention of the cleaner fish. Cleaning stations are usually associated with unique topological features, such as those seen in the coral reef[1] and allow a space where cleaners have no risk of predation from larger predatory fishes, due to the mutual benefit from the cleaners service [8].

Interactions are initiated by the client and terminated by the cleaner, implying that the client is seeking out the service where the cleaner has control[2].

Cheating edit

Cheating parasitism occurs when the cleaner eats mucus or healthy tissue from the client. This can be harmful to the client as mucus is essential for preventing UV damage, and open wounds can increase risk of infection [2]. Cleaner fish maintain a balance between consuming ectoparasites and mucus or tissue because of the respective nutritional benefits, sometimes despite the risk to the client [20]. For example, theCaribbean cleaning goby (Elacatinus evelynae) will eat scales and mucus from the host during times of ectoparasite scarcity to supplement its diet. The symbiosis relationship between client and host does not break down because the abundance of these parasites varies significantly seasonally and spatially, and the overall benefit to the larger fish outweighs any cheating on the part of the smaller cleaner[21].

Memory edit

Cleaner fish (especially facultative cleaners) asses value of possible clients when deciding whether to invest in a client or cheat and consume mucus or tissue [1][3]. Observations of cleaner and client interactions have noted that cleaners may provide the client with tactile stimulation as a way to establish a relationship and gain the clients 'trust'. This interaction is at cost to the cleaner as it is time that it is not feeding[3]. This physical interaction demonstrates a cleaner fish tradeoff. The cleaner minimizes feeding time to establish a memorable relationship with the client that also contributes to conflict management with a possible predatory client[3].

Neurobiology edit

 
Protein structure of non-mammalian specific hormone, vasotocin, from the posterior pituitary.

The cleaner fish neuroendocrine system has been studied specifically in reference to Arginine Vasotocin (AVT) and Isotocin. These are fish specific hormones that are analogous to human hormones involved in sociality[3]. In laboratory experiments, during conditions of low AVT, cleaners are more engaged in interspecific interactions. High AVT conditions tend to show high client interactions but more instances of cheating. This implies that AVT expression acts as a switch for cleaner fish feeding behaviour, showing less client interactions (but more honest cleaning) or increased client interactions (with less honest cleaning)[3]. It has also been observed that obligate cleaners have higher overall brain activity, and specifically in the cerebellum, likely related to the movements involved in cleaning [3].

Serotonin has also been noted to influence cleaning behaviour. High serotonin increases motivation to interact with clients, and a lack of serotonin decreases client interaction and slows learning [3].


Mimicry edit

 
The bluestriped fangblenny is an aggressive mimic of the cleaner wrasse.

Mimic species have evolved body forms, patterns, and colors which imitate other species to gain a competitive advantage [22]. One of the most studied examples of mimicry on coral reefs is the relationship between the aggressive mimic Plagiotremus rhinorhynchos (the bluestriped fangblenny) and the cleaner wrasse model Labroides dimidiatus. By appearing like L. dimidiatus, P. rhinorhynchos is able to approach and subsequently feed on the tissue and scales of client fish while posing as a cleaner.[23][24]

The presence of the cleaner mimic, P. rhinorhynchos, has a negative impact on the foraging success of the cleaner model L. dimidiatus.[24] P. rhinorhynchos feeds by eating the tissue and scales of client fish, making client fish much more cautious while at cleaning stations. More aggressive mimics have a greater negative impact on the foraging rate and success of the cleaner fish.[24] When mimics appear in higher densities relative to cleaners, there is a significant decline in the success rate of the cleaner fish. The effects of the mimic/model ratio are susceptible to dilution, whereby an increase in client fish allows both the mimics and the models to have more access to clients, thus limiting the negative effects that mimics have on model foraging success.[25][26]

Similar species also include Plagiotremus tapeinosoma (the Mimic blenny),Aspidontus.

Implications edit

Salmonid Aquaculture edit

 
An example of an outdoor aquaculture facility in Chile.

Aquaculture is the farming of aquatic organisms, where salmon farming is growing in the North Atlantic [10]. Cleaner fish are used to eat unwanted sea lice from salmon to reduce outbreaks which cause disease in populations. The two most commonly used cleaner fish are the lumpfish, Cyclopterus lumpus, and the ballan wrasse Labrus bergeylta.[11]. Lumpfish are distributed across the Atlantic ocean, ranging from Greenland to France, Hudson's Bay to New Jersey, and in high concentrations in the Bay of Fundy and St. Pierre Coast, near Newfoundland. [27] Ballan wrasse are distributed widely across the Northeast Atlantic Ocean [28]. The switch towards lumpfish has been preferred as wrasse are less active feeders during winter months [29].

Methods edit

Cleaner fish are commercially cultured and introduced into salmonid sea cages. Salmon and lumpfish are able to coexist, where the lumpfish spend a certain amount of time foraging for supplemented food and only a portion of their time delousing salmon. With significant ratios of cleaner to client, the efforts are sufficient to minimize louse outbreaks.[29][11] Sea cages are designed with additional substrate for lumpfish to attach to during periods of inactivity to minimize stress levels in the cleaner fish and maximize delousing abilities.[29]

Challenges of using cleaner fish edit

 
Sea lice causing morphological damage on cultured salmon in New Brunswick, Canada.

The cleaner fish used in North Atlantic Aquaculture facilities are facultative cleaners (Cyclopterus lumpus, and Labrus bergeylta) in order to control the nutrients they receive during culturing, before their use in aquaculture. One of the challenges that comes along with utilizing facultative cleaners is that parasite feeding from salmon must be maximized while also balancing additional nutrients from supplemented feed to ensure the health of the cleaner fish and the safety of the salmonid clients. [1] Another challenge that arises in management of cleaner fish behaviour is balancing the number of cleaners to the number of clients. With a low cleaner to client ratio, the risk of lice infestation increases. With a high cleaner to client ratio, competition among cleaners increases and there is a higher risk of cheating and consumption of salmonid mucus and flesh thereby increasing their risk of infection. [1][11]

Minimizing disease in commercial lumpfish stocks is critical for the continuation of their usage in aquaculture. Vaccine development for the lumpfish is a current area of research as lumpfish demand is increasing in the aquaculture industry.[29] In an effort to minimize disease in the cleaner fish, commercial lumpfish stocks are supplemented with wild individuals during the breeding season to minimize inbreeding depression. The lumpfish genome has not yet been fully sequenced so subtle details between populations is not yet appreciated.[29]

Another consideration in using cleaner fish in aquaculture is minimizing escapees from sea cages. If escaped cleaner fish spawn with natural populations in the environment it may decrease the wild fishes natural survival abilities. [29]

Environment edit

Cleaner Fish have taken over lice reduction strategies, which were based upon chemical delousers in the past. This decreases the degree of effluent waste affecting the surrounding habitats in outdoor aquaculture. [11] Introducing cleaner fish into salmonid aquaculture cages has also been studied to be less stressful on salmonids than medical intervention for sea louse outbreaks. [29]

Cleaner fish in the wild contribute to the overall health of aquatic communities by reducing morphological and physiological injuries by parasites to other heterospecifics. Maintenance of these populations of fish help the complex web of interactions remain stable.[2]

Economic edit

Sea lice outbreaks are detrimental to the survival of cultured salmonids and cause the majority of revenue loss in the aquaculture business. By employing the cleaner fish behaviour, aquaculture farmers save money in comparison to medical intervention for sea louse management. [29]

See also edit

References edit

  1. ^ a b c d e f g h i j k l m n Dunkley, Katie; Cable, Jo; Perkins, Sarah E. (2018-02-01). "The selective cleaning behaviour of juvenile blue-headed wrasse (Thalassoma bifasciatum) in the Caribbean". Behavioural Processes. 147: 5–12. doi:10.1016/j.beproc.2017.12.005. ISSN 0376-6357.
  2. ^ a b c d e f g h i j k Morado, Nadia; Mota, Paulo G.; Soares, Marta C. (2019). "The Rock Cook Wrasse Centrolabrus exoletus Aims to Clean". Frontiers in Ecology and Evolution. 7. doi:10.3389/fevo.2019.00182. ISSN 2296-701X.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  3. ^ a b c d e f g h i j "The Neurobiology of Mutualistic Behavior: The Cleanerfish Swims into the Spotlight". www.frontiersin.org. Retrieved 2019-10-27.{{cite web}}: CS1 maint: url-status (link)
  4. ^ Gingins, Simon; Werminghausen, Johanna; Johnstone, Rufus A.; Grutter, Alexandra S.; Bshary, Redouan (2013-06-22). "Power and temptation cause shifts between exploitation and cooperation in a cleaner wrasse mutualism". Proceedings of the Royal Society B: Biological Sciences. 280 (1761): 20130553. doi:10.1098/rspb.2013.0553. ISSN 0962-8452. PMC 3652443. PMID 23615288.{{cite journal}}: CS1 maint: PMC format (link)
  5. ^ Grutter, A. S. (2002). "Cleaning symbioses from the parasites' perspective". Parasitology. 124 (7): 65–81. doi:10.1017/S0031182002001488. ISSN 0031-1820. {{cite journal}}: no-break space character in |first= at position 3 (help)
  6. ^ Sazima, Cristina; Grossman, Alice; Sazima, Ivan (2010-02-05). "Turtle cleaners: reef fishes foraging on epibionts of sea turtles in the tropical Southwestern Atlantic, with a summary of this association type". Neotropical Ichthyology. 8 (1): 187–192. doi:10.1590/S1679-62252010005000003. ISSN 1982-0224.
  7. ^ "Manatee gets 'haircut' from gill fish". 2010-02-26. ISSN 0307-1235. Retrieved 2019-10-28.
  8. ^ a b Helfman, Gene S. (1997). The diversity of fishes. Collette, Bruce B., Facey, Douglas E. Malden, Mass.: Blackwell Science. ISBN 0865422567. OCLC 36051279.
  9. ^ Cheney, Karen L.; Grutter, Alexandra S.; Blomberg, Simon P.; Marshall, N. Justin (2009). "Blue and Yellow Signal Cleaning Behavior in Coral Reef Fishes". Current Biology. 19 (15): 1283–1287. doi:10.1016/j.cub.2009.06.028.
  10. ^ a b "Cleaner fish – what do they do?". Lochduart. 2017-06-08. Retrieved 2019-10-27.
  11. ^ a b c d e Brooker, Adam J; Papadopoulou, Athina; Gutierrez, Carolina; Rey, Sonia; Davie, Andrew; Migaud, Herve (2018-09-29). "Sustainable production and use of cleaner fish for the biological control of sea lice: recent advances and current challenges". Veterinary Record. 183 (12): 383–383. doi:10.1136/vr.104966. ISSN 0042-4900.
  12. ^ Helfman, Gene S. (1997). The diversity of fishes. Collette, Bruce B., Facey, Douglas E. Malden, Mass.: Blackwell Science. ISBN 0865422567. OCLC 36051279.
  13. ^ Fenner, Robert M. (1998). The conscientious marine aquarist : a commonsense handbook for successful saltwater hobbyists. Shelburne, Vt.: Microcosm. ISBN 1890087033. OCLC 38168280.
  14. ^ Powell, Adam; Treasurer, Jim W.; Pooley, Craig L.; Keay, Alex J.; Lloyd, Richard; Imsland, Albert K.; Leaniz, Carlos Garcia de (2018). "Use of lumpfish for sea-lice control in salmon farming: challenges and opportunities". Reviews in Aquaculture. 10 (3): 683–702. doi:10.1111/raq.12194. ISSN 1753-5131.
  15. ^ "Cleaner fish – what do they do?". Lochduart. 2017-06-08. Retrieved 2019-10-28.
  16. ^ Wyman, Richard L.; Ward, Jack A. (1972-12-29). "A Cleaning Symbiosis between the Cichlid Fishes Etroplus maculatus and Etroplus suratensis. I. Description and Possible Evolution". Copeia. 1972 (4): 834. doi:10.2307/1442742. ISSN 0045-8511.
  17. ^ Carvalho, L.N (2007). "Natural history of Amazon fishes". In Encyclopedia of Life Support Systems (ed.). Tropical Biology and Natural Resources Theme. 1. Oxford: Eolss Publishers. pp. 1–24.
  18. ^ Planet, Den Blå; Fortlingsvej 1, Address: Jacob; Kastrup, Address: 2770; info@denblaaplanet.dk, Send email:; Phone: +45 44 22 22 44 (2016-10-03). "Broadband red headstander". Den Blå Planet. Retrieved 2019-10-27.{{cite web}}: CS1 maint: extra punctuation (link) CS1 maint: numeric names: authors list (link)
  19. ^ "Memorial University Libraries - Proxy Login". login.qe2a-proxy.mun.ca. doi:10.1007/s00338-012-0933-9. Retrieved 2019-11-13.
  20. ^ Gingins, Simon; Werminghausen, Johanna; Johnstone, Rufus A.; Grutter, Alexandra S.; Bshary, Redouan (2013-06-22). "Power and temptation cause shifts between exploitation and cooperation in a cleaner wrasse mutualism". Proceedings of the Royal Society B: Biological Sciences. 280 (1761): 20130553. doi:10.1098/rspb.2013.0553. ISSN 0962-8452. PMC 3652443. PMID 23615288.{{cite journal}}: CS1 maint: PMC format (link)
  21. ^ Cheney, Karen L; Côté, Isabelle M (2005-05-16). "Mutualism or parasitism? The variable outcome of cleaning symbioses". Biology Letters. 1 (2): 162–165. doi:10.1098/rsbl.2004.0288. ISSN 1744-9561.
  22. ^ Cheney, Karen L.; Grutter, Alexandra S.; Marshall, N. Justin (2008-01-22). "Facultative mimicry: cues for colour change and colour accuracy in a coral reef fish". Proceedings. Biological Sciences. 275 (1631): 117–122. doi:10.1098/rspb.2007.0966. ISSN 0962-8452. PMC 2596177. PMID 17986437.
  23. ^ Cheney, Karen L.; Grutter, Alexandra S.; Marshall, N. Justin (2008-01-22). "Facultative mimicry: cues for colour change and colour accuracy in a coral reef fish". Proceedings. Biological Sciences. 275 (1631): 117–122. doi:10.1098/rspb.2007.0966. ISSN 0962-8452. PMC 2596177. PMID 17986437.
  24. ^ a b c Cheney, Karen L. (2012-02-23). "Cleaner wrasse mimics inflict higher costs on their models when they are more aggressive towards signal receivers". Biology Letters. 8 (1): 10–12. doi:10.1098/rsbl.2011.0687. PMC 3259977. PMID 21865244.{{cite journal}}: CS1 maint: PMC format (link)
  25. ^ Cheney, Karen L; Côté, Isabelle M (2007-09-07). "Aggressive mimics profit from a model–signal receiver mutualism". Proceedings of the Royal Society B: Biological Sciences. 274 (1622): 2087–2091. doi:10.1098/rspb.2007.0543. PMC 2706197. PMID 17591589.{{cite journal}}: CS1 maint: PMC format (link)
  26. ^ Cheney, Karen L; Côté, Isabelle M (2005-12-22). "Frequency-dependent success of aggressive mimics in a cleaning symbiosis". Proceedings of the Royal Society B: Biological Sciences. 272 (1581): 2635–2639. doi:10.1098/rspb.2005.3256. ISSN 0962-8452. PMC 1559983. PMID 16321786.
  27. ^ "Lumpfish: Emerging species profile list. DFO" (PDF).{{cite web}}: CS1 maint: url-status (link)
  28. ^ "Labrus bergylta : Ballan Wrasse | NBN Atlas". species.nbnatlas.org. Retrieved 2019-10-28.
  29. ^ a b c d e f g h Powell, Adam; Treasurer, Jim W.; Pooley, Craig L.; Keay, Alex J.; Lloyd, Richard; Imsland, Albert K.; Leaniz, Carlos Garcia de (2018). "Use of lumpfish for sea-lice control in salmon farming: challenges and opportunities". Reviews in Aquaculture. 10 (3): 683–702. doi:10.1111/raq.12194. ISSN 1753-5131.
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