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sure in the buccal cavity causing the water in front of the mouth to rush into the oral cavity 3 entrapping the prey in this flow this mode of feeding has two main phases expansion and compression 2 the expansion phase involves the initial opening of the jaws to capture prey these movements during the expansion phase are similar across all suction feeders with the kinesis of the skull leading to slight variations during the compression phase the jaws close and water is compressed out of the gills though suction feeding can be seen across fish species those with more cranial kinesis show an increase in suction potential as a result of more complex skull linkages that allow greater expansion of the buccal cavity and thereby create a greater negative pressure most commonly this is achieved by increasing the lateral expansion of the skull in addition the derived trait of anterior protrusion via the premaxillary bone in the upper jaw is acknowledged to increase the force exerted on the prey to be engulfed 4 protrusible jaws via a mobile premaxilla can only be seen in fishes within the teleostei clade 2 however a common misconception of these fishes is that suction feeding is the only or primary method employed 5 in micropterus salmoides ram feeding is the primary method for prey capture however they can modulate between the two methods or use both as with many teleosts 5 6 also it is commonly thought that fishes with more primitive characteristics also exhibit suction feeding although suction may be created upon the mouth opening in such fishes the criteria for pure suction feeding includes little or no bodily movement towards their prey 2 tradeoffs edit the morphologies and behaviors during suction feeding have led to three main proposed tradeoffs that determine the success of prey capture 7 the rate of jaw opening and closing the mobility of the bony elements in the skull and the ratio of ram to suction feeding behavior the first two qualifications center around the situation that results from a highly kinetic skull 8 having a highly mobile skull introduces a tradeoff between the ability to have high speed jaw opening high kinesis or higher bite transmission lower kinesis while there is a more complex relationship between mechanical advantage and the speed of lower jaw depression 9 10 11 there is consensus that species using high speed attacks have more cranial kinesis compared to species that exhibit low speed attacks 12 13 10 species that have a durophagous diet have also evolved skull morphologies to crush the hard shelled prey that is a part of their diet 14 15 durophagous species skulls consistently have more fused skulls and shorter jaw lengths this morphology leads to the skulls being less kinetic than their piscivorous counterparts 13 8 16 having shorter jaw lengths with a more akinetic skull allows for an individual to have a higher bite force compromising the ability to have a faster jaw opening when the jaw lengths are longer the third main tradeoff within suction feeding occurs with the incorporation of ram feeding with suction feeding behaviors ram feeding involves movement of the predator with its mouth open to engulf the prey 6 most species use ram feeding combined with suction feeding to increase the chances of capturing elusive prey 6 by swimming towards their prey while using suction to draw prey into the mouth this diversity in relative use is quantified using the ram suction index rsi that calculates the ratio of use for ram and suction during prey capture 6 the rsi ratio can be influenced by the morphology of the predator and by the elusiveness of the prey ram feeding and suction feeding are on opposite sides of the feeding spectrum where extreme ram feeding is when a predator swims over an immobile prey item with open jaws to engulf the prey extreme suction feeding is demonstrated by sit and wait predators that rely on rapid depression of the jaws to capture prey e g frogfish antennariidae there is wide diversity on how much of each feeding strategy an individual uses especially when body ram movements are considered 17 the relative use of ram and suction feeding is species dependent but it can help determine the accuracy of prey capture 18 the mouth aperture represents another tradeoff between the ability to capture large elusive prey with more chances of failure large gape or to capture smaller elusive prey with greater success smaller gape a predator with a small mouth aperture can generate strong suction force compared to an individual with a wider gape 19 18 this was demonstrated by wainwright et al 2007 by comparing the feeding success of the bluegill sunfish lepomis macrochirus and the largemouth bass micropterus salmoides l macrochirus has a smaller gape and was found to have higher accuracy with higher flow velocity and acceleration while m salmoides has a larger gape with lower accuracy and lower flow velocity and acceleration 18 however with the larger gape the largemouth bass were able to capture larger elusive prey using ram feeding in combination with suction feeding can also influence the direction of water into the mouth of the predator with use of ram predators are able to change the flow of water around the mouth and focus the flow of water into the mouth 20 but with too much ram a bow wave is created in front of the predator which can push the prey away from the predator s body 20 the mouth aperture and rsi represent the overall tradeoff between having a large gape with lower accuracy but being able to capture larger prey vs having a smaller gape with increased accuracy but the size of prey is limited the three main tradeoffs within the fish skull have occurred because of the high kinesis in the skull and the elusiveness of some prey types however having kinesis in the skull can enable a predator to evolve new techniques on increasing the performance of prey capture ram feeding edit ram feeding is a method of feeding underwater in which the predator moves forward with its mouth open engulfing the prey along with the water surrounding it during ram feeding the prey remains fixed in space and the predator moves its jaws past the prey to capture it the motion of the head may induce a bow wave in the fluid which pushes the prey away from the jaws but this can be avoided by allowing water to flow through the jaw this can be accomplished by means of a swept back mouth as in balaenid whales 21 or by allowing water to flow out through the gills as in sharks and herring a number of species have evolved narrow snouts as in gar fish and water snakes 22 herrings often hunt copepods if they encounter copepods schooling in high concentrations the herrings switch to ram feeding they swim with their mouth wide open and their opercula fully expanded every several feet they close and clean their gill rakers for a few milliseconds filter feeding the fish all open their mouths and opercula wide at the same time the red gills are visible in the photo below click to enlarge the fish swim in a grid where the distance between them is the same as the jump length of the copepods citation needed foraging mobula alfredi ram feeding swimming against the tidal current with its mouth open and sieving zooplankton from the water 23 herring ram feeding on a school of copepods school of adult indian mackerel ram feeding on macroplankton lunge feeding edit a humpback whale straining water through its baleen after lunging rorquals feed on plankton by a technique called lunge feeding 24 lunge feeding could be regarded as a kind of inverted suction feeding during which a whale takes a huge gulp of water which is then filtered through the baleen 24 biomechanically this is a unique and extreme feeding method for which the animal at first must accelerate to gain enough momentum to fold its elastic throat buccal cavity around the volume of water to be swallowed 25 subsequently the water flows back through the baleen keeping back the food particles the highly elastic and muscular buccal rills are a specialized adaptation to this feeding mode jaw protrusion edit external videos video of a slingjaw wrasse catching prey by protruding its jaw main article jaw protrusion jaw protrusion is the outward movement of the premaxilla or mouthparts towards the prey which is achieved via more mobile mechanical linkages in the articulations of the mouth vertebrate jaw protrusion is known only among modern bony fishes which possess many forms of coupled linkages in their head 26 remarkable examples are the slingjaw wrasse and the sand eel which can protrude their mouth by several centimeters this is usually done to extend the striking range of suction feeding and the retraction of the jaw after protrusion can also help retrieval once the prey has been engulfed another example of mouthpart protrusion is seen in dragonfly larvae nymphs which have hydraulic lower mandibles that can extend rapidly protruding forward to seize prey and bring it to the top jaw 27 pivot feeding edit pivot feeding is a method to transport the mouth towards the prey by an upward turning of the head which is pivoting on the neck joint pipefish such as sea horses and sea dragons are specialized on this feeding mechanism 28 with prey capture times of down to 5 ms shrimpfish centriscus scutatus this method is used by the fastest feeders in the animal kingdom the secret of the speed of pivot feeding is in a locking mechanism in which the hyoid arch is folded under the head and is aligned with the urohyal which connects to the shoulder girdle a four bar linkage at first locks the head in a ventrally bent position by the alignment of two bars the release of the locking mechanism jets the head up and moves the mouth toward the prey within 5 10 ms the trigger mechanism of unlocking is debated but is probably in lateral adduction seahorses rely on stealth to ambush small prey such as copepods they use pivot feeding to catch the copepod which involves rotating their snout at high speed and then sucking in the copepod 29 30 filter versus suspension feeding edit krill feeding under high phytoplankton concentration slowed down by a factor of 12 these are contrasting methods for the removal of food particles from a water flow for example by the gill rakers of fish the baleen of whales or the ostia of sponges filter feeding edit in filter feeding the water flow is primarily generated by the organism itself for example by creating a pressure gradient by active swimming or by ciliary movements suspension feeding edit in suspension feeding the water flow is primarily external and the particles themselves move with respect to the ambient water flow such as in sea lilies see also edit cleaner fish feeding behaviour of fish lepidophagy list of feeding behaviours paedophagy references edit alexander r mcneill 1967 functional design in fishes london hutchinson isbn 0 09 084770 9 oclc 456355 1 2 3 4 lauder george v may 1982 patterns of evolution in the feeding mechanism of actinopterygian fishes american zoologist 22 2 275 285 doi 10 1093 icb 22 2 275 issn 0003 1569 lauder george v march 1980 evolution of the feeding mechanism in primitive actionopterygian fishes a functional anatomical analysis of polypterus lepisosteus and amia journal of morphology 163 3 283 317 doi 10 1002 jmor 1051630305 issn 0362 2525 pmid 30170473 s2cid 26805223 holzman roi day steven w mehta rita s wainwright peter c 10 june 2008 jaw protrusion enhances forces exerted on prey by suction feeding fishes journal of the royal society interface 5 29 1445 1457 doi 10 1098 rsif 2008 0159 pmc 2607355 pmid 18544504 1 2 gardiner jayne m motta philip j 28 january 2012 largemouth bass micropterus salmoides switch feeding modalities in response to sensory deprivation zoology jena germany 115 2 78 83 doi 10 1016 j zool 2011 09 004 pmid 22285791 1 2 3 4 norton s f brainerd e l 1993 03 01 convergence in the feeding mechanics of ecomorphologically similar species in the centrarchidae and cichlidae journal of experimental biology 176 1 11 29 doi 10 1242 jeb 176 1 11 issn 0022 0949 gidmark nicholas j pos kelsie matheson bonne ponce esai westneat mark w 2019 functional morphology and biomechanics of feeding in fishes feeding in vertebrates springer international publishing pp 297 332 doi 10 1007 978 3 030 13739 7_9 isbn 978 3 030 13738 0 s2cid 150135750 1 2 westneat m w 2004 11 01 evolution of levers and linkages in the feeding mechanisms of fishes integrative and comparative biology 44 5 378 389 doi 10 1093 icb 44 5 378 issn 1540 7063 pmid 21676723 bellwood d r wainwright p c fulton c j hoey a s 2005 10 12 functional versatility supports coral reef biodiversity proceedings of the royal society b biological sciences 273 1582 101 107 doi 10 1098 rspb 2005 3276 issn 0962 8452 pmc 1560014 pmid 16519241 1 2 oufiero c e holzman r a young f a wainwright p c 2012 08 01 new insights from serranid fishes on the role of trade offs in suction feeding diversification journal of experimental biology 215 21 3845 3855 doi 10 1242 jeb 074849 issn 0022 0949 pmid 22855615 wainwright peter c bellwood david r westneat mark w grubich justin r hoey andrew s 2004 04 22 a functional morphospace for the skull of labrid fishes patterns of diversity in a complex biomechanical system biological journal of the linnean society 82 1 1 25 doi 10 1111 j 1095 8312 2004 00313 x issn 0024 4066 lauder george v liem karel f november 1981 prey capture by luciocephalus pulcher implications for models of jaw protrusion in teleost fishes environmental biology of fishes 6 3 4 257 268 bibcode 1981envbf 6 257l doi 10 1007 bf00005755 issn 0378 1909 s2cid 24315046 1 2 martinez christopher m mcgee matthew d borstein samuel r wainwright peter c 2018 07 10 feeding ecology underlies the evolution of cichlid jaw mobility evolution 72 8 1645 1655 doi 10 1111 evo 13518 issn 0014 3820 pmid 29920668 s2cid 49311313 archived from the original on 2020 07 21 retrieved 2020 09 06 collar david c reece joshua s alfaro michael e wainwright peter c mehta rita s june 2014 imperfect morphological convergence variable changes in cranial structures underlie transitions to durophagy in moray eels the american naturalist 183 6 e168 e184 doi 10 1086 675810 issn 0003 0147 pmid 24823828 s2cid 17433961 durie c j turingan r 2001 relationship between durophagy and feeding biomechanics in gray triggerfish balistes capriscus intraspecific variation in ecological morphology florida scientist 64 20 28 westneat mark w 2005 skull biomechanics and suction feeding in fishes fish biomechanics fish physiology vol 23 elsevier pp 29 75 doi 10 1016 s1546 5098 05 23002 9 isbn 978 0 12 350447 0 longo sarah j mcgee matthew d oufiero christopher e waltzek thomas b wainwright peter c 2015 11 23 body ram not suction is the primary axis of suction feeding diversity in spiny rayed fishes the journal of experimental biology 219 1 119 128 doi 10 1242 jeb 129015 issn 0022 0949 pmid 26596534 1 2 3 wainwright p c carroll a m coll...