Fish intelligence
Fish intelligence is "the resultant of the process of acquiring, storing in memory, retrieving, combining, comparing, and using in new contexts information and conceptual skills"[1] as it applies to fish. Due to a common perception amongst researchers that Teleost fish are "primitive" compared to mammals and birds, there has been much less research into fish cognition than into those types of animals, and much remains unknown about fish cognition, though evidence of complex navigational skills such as cognitive maps is increasing.[2]
Compared to similarly sized fish, mammals and birds typically have brain sizes fifteen times larger, though some species of fish such as elephantnose fish have very large brain-to-body ratios. However, fish still display intelligence that cannot be explained through Pavlovian and operant conditioning, such as reversal learning, novel obstacle avoidance, and passing simultaneous two-choice tasks.[3] Some fish also match mammals and birds in the executive functioning capability of inhibitory motor control.[4] Furthermore, some fish also have an oxytocin-enabled ability to sense another's fish's fear. Australian biologist Culum Brown has argued that fish may give the appearance of being less intelligent than they are due to differences between aquatic and terrestrial environments.[5]
Fish hold records for the
Brain
Fish typically have quite small brains relative to body size compared with other vertebrates, typically one-fifteenth the brain mass of a similarly sized bird or mammal.[9] However, some fish have relatively large brains, most notably mormyrids and sharks, which have brains about as massive relative to body weight as birds and marsupials.[10]
The cerebellum of
In mormyrid fish (a family of weakly electrosensitive freshwater fish), the cerebellum is considerably larger than the rest of the brain put together. The largest part of it is a special structure called the valvula, which has an unusually regular architecture and receives much of its input from the electrosensory system.[13]
Memory
Individual
A number of studies have shown that fish can retain information for months or years. Anecdotally,
Several fish species are capable of learning complex spatial relationships and forming
Tool-like behavior
Several species of
Archerfish (family Toxotidae) squirt jets of water at insects on plants above the surface to knock them into the water. They can adjust the size of the squirts to the size of the insect prey.[34] They can even learn to shoot at moving targets.[35][36]
Whitetail damselfish clean the rock face where they intend to lay eggs by sucking up and blowing sand grains onto the surface.[37] Triggerfish blow water at sea urchins to turn them over, thereby exposing their more vulnerable underside.[38] River stingrays create water currents with their fins to suck food out of a PVC pipe.[39] Banded acaras (Bujurquina vittata) lay their eggs on a loose leaf and carry the leaf away when a predator approaches.[40]
In one laboratory study, Atlantic cod (Gadus morhua) given access to an operant feeding machine learned to pull a string to get food. The researchers had also tagged the fish by threading a bead in front of their dorsal fin. Some fish snagged the string with their bead, resulting in food delivery. These fish eventually learned to swim in a particular way to repeatedly make the bead snag the string and get food. Because the fish used an object external to their body in a goal-oriented way, this satisfies some definitions of tool use - though in a more rudimentary fashion, a human blowing air wouldn't qualify as tool usage just like a fish using water in the same fashion, or even in the case of certian wrasse, the bivalve is not a tool but the prey itself. If the wrasse had of used a rock to access the bivalve then it would fall under that classification. There appear to be very few to no observed behaviors of fish using tools in a fashion similar to humans, but instead altering water flows in ways that benefits their survival.[41]
Construction
As for tool use, construction behaviour may be mostly innate. Yet it can be sophisticated, and the fact that fish can make judicious repairs to their creation suggests intelligence. Construction methods in fishes can be divided into three categories: excavations, pile-ups, and gluing.[42]
Excavations may be simple depressions dug up in the substrate, such as the nests of
Mounds are easy to build, but can be quite extensive. In North American streams, the male
Male pufferfish, Torquigener sp., also build sand mounds to attract females. The mounds, up to 2 m in diameter, are intricate with radiating ridges and valleys.[49]
Several species build up mounds of coral pieces either to protect the entrance to their burrows, as in
Male sticklebacks are well known for their habit of building an enclosed nest made of pieces of vegetation glued together with secretions from their kidneys. Some of them adorn the entrance of the nest with unusually colored algae or even shiny tinfoil experimentally introduced in the environment.[56]
Foam nests, made up of air bubbles glued together with mucus from the mouth, are also well known in
Social intelligence
Fish can remember the attributes of other individuals, such as their competitive ability or past behavior, and modify their own behavior accordingly. For example, they can remember the identity of individuals to whom they have lost in a fight, and avoid these individuals in the future; or they can recognize territorial neighbors and show less aggression towards them as compared to strangers.[57] They can recognize individuals in whose company they obtained less food in the past and preferentially associate with new partners in the future.[57]
Fish can seem mindful of which individuals have watched them in the past. In an experiment with Siamese fighting fish, two males were made to fight each other while being watched by a female, whom the males could also see. The winner and the loser of the fight were then, separately, given a choice between spending time next to the watching female or to a new female. The winner courted both females equally, but the loser spent more time next to the new female, avoiding the watcher female.[58] In this species, females prefer males they have seen win a fight over males they have seen losing,[59] and it therefore makes sense for a male to prefer a female that has never seen him lose as opposed to a female that has seen him lose.
Social interactions also provides the context for a test of transitive inference, that is figuring out that if A > B and B > C, then A > C. In a study with the cichlid Astatotilapia burtoni, observer fish could watch aggressive interactions between pairs of other individuals. They witnessed individual A beat individual B, then individual B beat individual C, then C beat D, and D beat E. The observer fish were then given a choice of associating with either B or D (both of which they had seen win once and lose once). All eight observer fish tested spent more time next to D. Fish in this species prefer to associate with more subordinate individuals, so the preference for D showed that the observers had worked out that given that B was superior to C, and C to D, then D had to be subordinate to B.[60]
A 2023 study found that zebrafish utilize oxytocin to react to the fear of other fish. Oxytocin is also used by humans in empathy. It found that zebrafish that had oxytocin production removed by gene editing cannot respond to the fear of other fish. When oxytocin is injected back into the fish, they react in a way that suggests they may have empathy in regards to this emotion. Furthermore, because the same regions of the brain were involved as in mammals, the study suggests oxytocin-based empathy may have evolved from a common ancestor many millions of years ago.[61]
Deception
There are several examples of fish being deceptive, suggesting to some researchers that they may possess a theory of mind. However, most of the observations of deception can be understood as instinctive patterns of behavior that are triggered by specific environmental events, and they do not require a fish to understand the point of view of other individuals.
Distraction display
In the
False courtship behaviour
In the Malili Lakes of Sulawesi, Indonesia, one species of
Death feigning
Death feigning as a way to attract prey is another form of deception.[67] In Lake Malawi, the predatory cichlid Nimbochromis livingstonii have been seen first remaining stationary with their abdomen on or near sand and that then dropping onto their sides. In a variant behaviour, some N. livingstonii fell through the water column and landed onto their side. The fish then remained immobile for several minutes. Their colour pattern was blotchy and suggested a rotting carcass. Small inquisitive cichlids of other species often came near and they were suddenly attacked by the predator. About a third of the death-feigning performances led to an attack, and about one-sixth of the attacks were successful.[68] Another African cichlid, Lamprologus lemairii, from Lake Tanganyika, has been reported to do the same thing.[69] A South American cichlid, the yellowjacket cichlid Parachromis friedrichsthalii, also uses death feigning. They turn over onto their sides at the bottom of the sinkholes they inhabit and remain immobile for as long as 15 minutes, during which they attack the small mollies that come too close to them.[70] The comb grouper Mycteroperca acutirostris may also be an actor, though in this case the behaviour should be called dying or illness feigning, rather than death feigning, because while lying on its side the fish occasionally undulates its body. In 1999, off the coast of southeastern Brazil, one juvenile comb grouper was observed using this tactic to catch five small prey in 15 minutes.[71]
Cooperation
Cooperative foraging reflects some mental flexibility and planning, and could therefore be interpreted as intelligence. There are a few examples in fishes.[72]
Yellowtail amberjack can form packs of 7-15 individuals that maneuver in U-shaped formations to cut away the tail end of prey shoals (jack mackerels or Cortez grunts) and herd the downsized shoal next to seawalls where they proceed to capture the prey.[73]
In the coral reefs of the Red Sea,
Similarly,
Numeracy
Mosquitofish (Gambusia holbrooki) can distinguish between doors marked with either two or three geometric symbols, only one of which allows the fish to rejoin its shoalmates. This can be achieved when the two symbols have the same total surface area, density and brightness as the three symbols.[77] Further studies have shown that this discrimination extends to 4 vs 8, 15 vs 30, 100 vs 200, 7 vs 14, and 8 vs 12 symbols, again controlling for non-numerical factors.[78] Other species similarly tested with some measure of success include goldfish (discriminating between 2 vs. 3, and 10 vs. 15),[79] guppy (3 vs. 4, and 4 vs. 5),[80][81] and zebrafish (2 vs. 3, 3 vs. 4, and 4 vs. 5, but not 5 vs. 6, nor 6 vs. 7).[82]
Many studies have shown that when given a choice, shoaling fish prefer to join the larger of two shoals. It has been argued that several aspects of such choice reflect an ability by fish to distinguish between numerical quantities.[83][84][85][86]
A laboratory study with zebra mbuna and ocellate river stingray has demonstrated that these fish can add and subtract 'one' from 2, 3, or 4. The fish had to learn that if presented with, say, 3 yellow symbols, then in a subsequent choice test between 2 and 4 symbols they had to choose 2 to get a food reward (thus, yellow meant "choose one less"); but if presented with 3 blue symbols, then they had to choose 4 rather than 2 (thus, blue meant "choose one more"). The fish easily learned this task (the success criterion was 70% correct choice). Crucially, if after 3 yellow symbols they were given a choice between 1 and 2, they chose 2; conversely, if after 3 blue symbols the fish were given a choice between 4 or 5 symbols, they chose 4. This latter test showed that the fish had learned the concept of "one less" or "one more", and not simply "less" or "more".[87]
Social learning
Fish can learn how to perform a behavior simply by watching other individuals in action.
Through cultural transmission, fishes could also learn where good food spots are. Ninespine stickleback, when given a choice between two food patches they have watched for a while, prefer the patch over which more fish have been seen foraging, or over which fish were seen feeding more intensively.[90] Similarly, in a field experiment where Trinidadian guppies were given a choice between two distinctly marked feeders in their home rivers, the subjects chose the feeder where other guppies were already present, and in subsequent tests when both feeders were deserted, the subjects remembered the previously popular feeder and chose it.[91]
Through social learning, fishes might learn not only where to get food, but also what to get and how to get it. Hatchery-raised salmon can be taught to quickly accept novel, live prey items similar to those they will encounter once they will be released in the wild, simply by watching an experienced salmon take such prey.[92][93] The same is true of young perch.[94] In the laboratory, juvenile European seabass can learn to push a lever in order to obtain food just by watching experienced individuals use the lever.[95]
Fishes can also learn from others the identity of predatory species. Fathead minnows, for example, can learn the smell of a predatory pike just by being simultaneously exposed to that smell and the sight of experienced minnows reacting with fear, and brook stickleback can learn the visual identity of a predator by watching the fright reaction of experienced fathead minnows.[96] Fish can also learn to recognize the odor of dangerous sites when they are simultaneously exposed to it and to other fish that suddenly show a fright reaction.[97] Hatchery-raised salmon can learn the smell of a predator by being simultaneously exposed to it and to the alarm substance released by injured salmon.[98]
Latent learning
Latent learning is a form of learning that is not immediately expressed in an overt response; it occurs without any obvious reinforcement of the behaviour or associations that are learned. One example in fish comes from research with male three spot gouramis (Trichopodus trichopterus).[99] This species quickly form dominance hierarchies. To appease dominants, subordinates adopt a typical body posture angled at 15-60º to the horizontal, all fins folded and pale body colors. Individuals trained to associate a light-stimulus with the imminent arrival of food exhibit this associative learning by approaching the surface where the food is normally dropped immediately the light-stimulus is presented. However, if a subordinate is placed in a tank with a dominant individual and the light-stimulus is presented, the subordinate immediately assumes the submissive posture rather than approaching the surface. The subordinate has predicted that going to the surface to get food would place it in competition with the dominant, and to avoid potential aggression, it immediately attempts to appease the dominant.
Cleaner fish
The
Mirror test
A 2019 study on bluestreak cleaner wrasses found they can successfully pass the mirror test. According to the authors, their findings mean that either prior conceptions of animal self-awareness or the validity of the mark test need to be reexamined.[104] Gordon Gallup, the originator of the test, believes that the results are best explained by the fish having instincts to scrape off parasites.[105] In 2022, a larger study was performed with positive results, but the fish only passed when the mark resembled a parasite. The authors then reiterated their conclusion that either self-awareness in animals or the validity of the mark test as a method need to be reconsidered.[106]
Play
Play behaviour is often considered a correlate of intelligence. One possible example in fish is provided by the electrolocating Peters' elephantnose fish (mentioned above as having one of the largest brain-to-body weight ratios of all known vertebrates). One captive individual was observed carrying a small ball of aluminum foil (a good conductor of electricity) to the outflow tube of the aquarium filter, letting the current push the ball away before chasing after it and repeating the behaviour.[107] The same species has been reported trying to balance tubes vertically at the water surface until they fell, and then try again.[108] Captive white-spotted cichlids have also been seen hitting a floating thermometer hundreds of times to make it wobble and bob.[109]
Food stocking
Food stocking can be viewed potentially as an animal planning for the future. One example of short-term stocking involves climbing perch (Anabas testudineus). Individuals were kept singly in aquaria and fed with pellets dropped at the surface. When the pellets were dropped one after the other at 1-s intervals, the fish took them as they reached the surface and stocked them inside the mouth. On average, the fish placed 7 pellets in their mouth before moving away to consume them. When starved for 24-h before the feeding test, they doubled the number of pellets stocked (14 on average); the underside of their heads bulged under the load. The behaviour may be an indication that competition for food is normally severe in this species and that any adaptation to secure food would be beneficial.[110]
See also
- Animal cognition
- Bird intelligence
- Cephalopod intelligence
- Cetacean intelligence
- Pain in fish
- Shoaling and schooling
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- ^ Helfman, G., and Collette, B. 2011. Fishes: The Animal Answer Guide. Johns Hopkins University Press, Baltimore.
- ^ Page 289 in: Yong, E. (2023) An immense world: How animal senses reveal the hidden realms around us. Vintage, London.
- ^ Burghardt, G.M., Dinets, V., and Murphy, J.B. (2014) Highly repetitive object play in a cichlid fish (Tropheus duboisi). Ethology 121: 38-44.
- ^ Binoy, V.V., and Thomas, K.J. 2008. The influence of hunger on food-stocking behaviour of climbing perch Anabas testudineus. Journal of Fish Biology 73: 1053-1057.
Works cited
- Helfman, G.; Collette, B.; Facey, D. (1997). The Diversity of Fishes (1st ed.). Wiley-Blackwell. ISBN 978-0-86542-256-8.
Further references
- Braithwaite, Victoria A (2005) "Cognitive ability in fish" Fish physiology, 24: 1–37.
- Brown C, Laland K and Krause J (Eds) (2011) Fish Cognition and Behavior John Wiley & Sons. ISBN 9781444342512.
- Brown C, Laland K and Krause J (2003) Learning in fishes: why are they smarter than you think? Fish and Fisheries, 4:197–288.
- Brown C and Laland K (2003) "Social learning in fishes: A review" Fish and Fisheries, 4(3), 280-288.
- Bshary R, Wickler W and Fricke H (2002) "Fish cognition: a primate's eye view". Animal Cognition, 5 (1): 1–13.
- Laland K, Brown C and Krause J (2003) "Learning in Fishes: An introduction" Fish and Fisheries, 4(3): 199-202.
- Reebs, Stephan (2001) Fish Behavior in the Aquarium and in the Wild. Cornell University Press. ISBN 0-8014-8772-2
- Schultz, Nora (2007) "When fish get emotional" New Scientist.
External links
- Researchers find fish that can count up to four The Guardian, 26 February 2008.
- Fish aren't thick: scientist The Sydney Morning Herald, September 12, 2006.
- The Times Smart school of fish expose stupidity of a popular myth TimesOnline, 22 November 2006.
- Fishy Machiavellis swim rings around anglers TimesOnline, 1 September 2003.