Types of Feeding Mechanism in Animals
Copepoda
Janet W. Reid , Craig E. Williamson , in Ecology and Classification of North American Freshwater Invertebrates (Third Edition), 2010
2. Feeding Mechanisms
The feeding mechanisms of suspension-feeding calanoids have been of great interest through the years as they are some of the most abundant metazoans on the planet, dominating the plankton of the world's oceans. Feeding mechanisms of calanoids appear to be similar for marine and freshwater species. Food is brought in toward the copepod on microcurrents created by rapid vibrations of the second antennae, mandibular palps, first maxillae, and maxillipeds [210,405]. Smaller food particles are captured passively, and are funneled into the mouth through the setae of the second maxillae[287,405]. During passive captures, the vibrations of the feeding appendages continue uninterrupted. Larger food particles are captured actively with an outward fling of the second maxillae followed by an inward squeeze to remove the excess water surrounding the food particle.
When microzooplankton such as rotifers and nauplii are brought in on the feeding currents of suspension-feeding diaptomids, the copepods will often attack these small animal prey from a distance with an active thrust response in which the antennae and swimming legs are used to orient and pounce toward the prey[428, 430 ,437]. Particles less than 5 μm are generally captured passively by diaptomids, while those greater than 50 μm are generally captured actively or by attack; in intermediate size ranges, the frequency of active captures increases with increasing particle size[405,437]. While chemical cues appear to be important in prey detection[260], high-speed video studies with inert particles have demonstrated that mechanoreception alone can be used to capture larger particles[33]. Many of the larger calanoid species such as Heterocope, Epischura, Limnocalanus, and some of the large species of diaptomids have predatory tendencies and feed on other zooplankton as well as algae. These predators cruise through the water, attack their prey with a pounce, and grasp them with their first and second maxillae. If a capture attempt fails, the copepod may swim in a vertical loop to try again to capture the prey[204].
Cyclopoid copepods do not create currents to aid their feeding, but instead grasp their food directly with their first maxillae or, to a lesser extent, with their second maxillae and maxillipeds. These appendages push food between the mandibles. By oscillating rapidly during feeding bouts, the mandibles tear prey into pieces and stuff them into the esophagus[ 130 ]. Cyclopoids detect their prey with the help of mechanoreceptors on their first antennae. Prey which generate mechanical disturbance in the water as they swim can be detected at distances of several millimeters. Cyclopoids actively orient and attack prey with considerable precision[204]. Larger prey such as cladocerans, copepods, and chironomids may be handled for 30 min or more before ingestion[ 130 ,139,422]. Small, generally less-active prey such as algae, protozoans, rotifers, and nauplii may be detected only after contact, and are generally eaten whole. Cyclopoid nauplii use their mandibles and second antennae to capture and ingest food[256].
Harpacticoid copepods are primarily surface feeders that use their mouthparts for scraping food from a diversity of substrates. Suspension feeding has been reported in two primitive marine families, the Longipediidae and the Canuellidae[180]. Some harpacticoids can use both surface-feeding and suspension-feeding mechanisms[180]. In surface feeding, the second maxillae and second antennae grasp the prey while the maxillipeds aid in anchoring the copepod to the substrate. Once the food is grasped, the first maxillae shred and stuff the food into the vibrating mandibles. The second maxillae and maxillipeds are generally not used in food handling. An exception is the primitive soil-dwelling Phyllognathopus viguieri, which uses its modified leaflike maxillipeds to grasp nematodes and push them into its mouth[222]. Harpacticoid nauplii have modified second antennae that are employed to grasp, masticate, and push food into the mouth[114]. The mandibles of nauplii serve only for locomotion and not for feeding. Suspension-feeding harpacticoids feed by lying on their dorsal or lateral sides and vibrating their first and second maxillae and mandibles. These vibrations create feeding currents that bring water and food in toward the mouth and out posteriorly along the ventral surface of the copepod.
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COPEPODA
Craig E. Williamson , Janet W. Reid , in Ecology and Classification of North American Freshwater Invertebrates (Second Edition), 2001
2. Feeding Mechanisms
The feeding mechanisms of suspension-feeding calanoids appear to be similar for marine and freshwater species. Food is brought in toward the copepod on microcurrents created by rapid vibrations of the second antennae, mandibular palps, first maxillae, and maxillipeds ( Koehl and Strickler, 1981; Vanderploeg and Paffenhöfer, 1985). Smaller food particles are captured passively and are funneled into the mouth through the setae of the second maxillae (Vanderploeg and Paffenhöfer, 1985; Price and Paffenhöfer, 1986). During passive captures, the vibrations of the feeding appendages continue uninterrupted. Larger food particles are captured actively with an outward fling of the second maxillae followed by an inward squeeze to remove the excess water surrounding the food particle.
When microzooplankton such as rotifers and nauplii are brought in on the feeding currents of suspension-feeding diaptomids, the copepods will often attack these small animal prey from a distance with an active thrust response in which the antennae and swimming legs are used to orient and pounce toward the prey (Williamson and Butler, 1986; Williamson, 1987; Williamson and Vanderploeg, 1988). Particles less than 5 μm are generally captured passively by diaptomids, while those greater than 50 μm are generally captured actively or by attack; in intermediate size ranges, the frequency of active captures increases with increasing particle size (Vanderploeg and Paffenhöfer, 1985; Williamson and Vanderploeg, 1988).
Many of the larger calanoid species such as Heterocope, Epischura, Limnocalanus, and some of the large species of diaptomids have predatory tendencies and feed on other zooplankton as well as algae. These predators cruise through the water, attack their prey with a pounce, and grasp them with their first and second maxillae. If a capture attempt fails, the copepod may swim in a vertical loop to try again to capture the prey (Kerfoot, 1978).
Cyclopoid copepods do not create currents to aid their feeding but instead grasp their food directly with their first maxillae or, to a lesser extent, with their second maxillae and maxillipeds. These appendages push food between the mandibles. By oscillating rapidly during feeding bouts, the mandibles tear prey into pieces and stuff them into the esophagus (Fryer, 1957a).
Cyclopoids detect their prey with the help of mechanoreceptors on their first antennae. Prey which generate much disturbance as they swim can be detected at distances of several millimeters. Cyclopoids actively orient and attack prey with considerable precision (Kerfoot, 1978). Larger prey such as cladocerans, copepods, and chironomids may be handled for 30 min or more before ingestion (Fryer, 1957a; Gilbert and Williamson, 1978; Williamson, 1980). Small, generally less active prey such as algae, protozoans, rotifers, and nauplii may be detected only after contact, and are generally eaten whole. Cyclopoid nauplii use their mandibles and second antennae to capture and ingest food (Monakov, 1976).
Harpacticoid copepods are primarily surface-feeders that use their mouthparts for scraping food from a diversity of substrates. Suspension-feeding has been reported in two primitive marine families, the Longipediidae and the Canuellidae (Hicks and Coull, 1983). In surface-feeding, the second maxillae and second antennae grasp the prey while the maxillipeds aid in anchoring the copepod to the substrate. Once the food is grasped, the first maxillae shred and stuff the food into the vibrating mandibles. The second maxillae and maxillipeds are not used in food handling. Harpacticoid nauplii have modified second antennae that are employed to grasp, masticate, and push food into the mouth (Fahrenbach, 1962). The mandibles of nauplii serve only for locomotion and not for feeding.
Suspension-feeding harpacticoids feed by lying on their dorsal or lateral sides and vibrating their first and second maxillae and mandibles. These vibrations create feeding currents that bring water and food in toward the mouth and out posteriorly along the ventral surface of the copepod. Some harpacticoids can use both surface-feeding and suspension-feeding mechanisms (Hicks and Coull, 1983).
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ORIGIN OF THE AMNIOTE FEEDING MECHANISM: EXPERIMENTAL ANALYSES OF OUTGROUP CLADES
George V. Lauder , Gary B. Gillis , in Amniote Origins, 1997
SYNTHESIS: CONCLUSIONS AND UNRESOLVED ISSUES
Bramble and Wake (1985) presented a general model of kinematic and electromyographic patterns for tetrapod feeding mechanisms. This model has been of important heuristic value because it has provided a hypothesis against which empirical data from extant tetrapods can be tested. At present, Bramble and Wake's model has received some support (see, for example, Schwenk and Throckmorton, 1989). However, this model has also come under review where experimental results do not match predictions. Based on the results from analyses of individual taxa, various authors have examined specific predictions of this model (Deheusy and Bels, 1992; Reilly and Lauder, 1990a, 1991b; So et al., 1992).
Althugh there is no doubt that, in amniotes, several general characteristics of the Bramble and Wake model do describe features of jaw function common to many amniote clades, the experimental results summarized above for amniotes and anamniote tetrapods also point out a number of complications that render their description of a "generalized tetrapod" functional pattern problematic.
First, the pattern of jaw movement and muscle function during prey transport in ambystomatid salamanders (the only anamniote taxon for which quantitative data are available on prey transport) is quite different than expected under the general tetrapod model. For example, there is no slow opening phase, hyoid protraction thus does not immediately precede fast opening, motion of the head and neck does not occur in the predicted manner (there is often very little horizontal skull movement), and there is no electrical activity in the depressor mandibulae and hyoid muscles just prior to the fast opening phase. In fact, many features of ambystomatid transport systems instead appear to be primitive traits inherited from aquatic ancestors. However, recall that ambystomatids show relatively distinct kinematic patterns during prey capture relative to most other salamanders, and it is indeed possible that this is true of their transport behavior as well. Further examination of transport behaviors in other urodeles is required before the accuracy of Bramble and Wake's model can be assessed relative to such primitive tetrapods.
Second, results from a variety of amniote taxa also suggest that kinematic and electromyographic data do not tightly fit predicted patterns. For example, in many transport and manipulation cycles there is no clear SO phase, and electromyographic patterns in the depressor mandibulae, sternohyoideus, adductor mandibulae, and pterygoideus show unpredicted patterns (Herrel et al., 1995). In addition, comparing prey capture in iguanians to the proposed model has been done by several investigators (e.g. Kraklau, 1991; Schwenk and Throckmorton, 1989), and Delheusy and Bels (1992: p. 184)s summarize their results by noting that "Our data do not support the model of Bramble and Wake or their speculation about the relationship between SO II duration and the size of the prey."
Given the limited experimental data available in 1985, it is perhaps not surprising that more recent results have called many of our previous concepts of amniote jaw function into question. However, even these additional data are insufficient to do more than suggest the outlines of a new view of amniote feeding function. Given the diversity of both anamniote and amniote taxa, the number of taxa for which we have both kinematic and electromyographic data is surprisingly few. We probably do not have a complete set of kinematic and electromyographic data for more than ten taxa of anamniote tetrapods and squamates. Furthermore, such data are rarely available for the full range of behavioral diversity exhibited by the feeding mechanism. In order to evaluate biomechanical models of jaw function and to produce evolutionary hypotheses of functional transformation, a much larger data set is needed. Such experimental data will permit a more quantitative assessment of the diversity of feeding system function in primitive amniotes and provide a better understanding of how plesiomorphic functional traits combined with novel features to form the basal amniote feeding mechanism.
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Controlling Soil-Borne Plant Pathogens
T.S. BELLOWS , in Handbook of Biological Control, 1999
Predacious Nematodes
Predatory nematodes are found in four main taxonomic groups (Monochilidae, Dorylaimidae, Aphelenchidae, and Diplogasteridae), each with a distinct feeding mechanism and food preferences ( Stirling, 1991). The monochilids have a large buccal cavity that bears a large dorsal tooth; all species are predaceous, feeding on protozoa, nematodes, rotifers, and other prey, which may be swallowed whole or pierced and the body contents removed. The dorylaimids are typically larger than their prey and possess a hollow spear that is used either to pierce the body of the prey or to inject enzymes into the food source and to then remove the predigested contents. The group is considered omnivorous, but the feeding habits are known for only a few species (Ferris & Ferris, 1989). Almost all the predatory aphelenchids are in the genus Seinura. Although small, they can feed on nematodes larger than themselves by injecting the prey with a rapidly paralyzing toxin via their stylet. The diplogasterids, typically a bacteria-feeding group, have a stoma armed with teeth, and the species with large teeth prey on other nematodes. Species in all these groups are generally omnivorous, feeding on free-living as well as plant-parasitic nematodes. The role of individual species in the population dynamics of plant-parasitic nematodes in the soil has been difficult to quantify, but it is possible that a number of species may act together to produce a significant impact (Stirling, 1991).
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Gelatinous Zooplankton
L.P. Madin , G.R. Harbison , in Encyclopedia of Ocean Sciences, 2001
Doliolida
This order of the Thaliacea comprises six genera and 23 species of small (2–10 mm), barrel-shaped animals with circumferential muscle bands. The filter feeding mechanism is similar to that of pyrosomes, with currents generated by ciliary beating passing through a mucous net supported on the branchial basket. The life cycle involves five asexual stages and one sexual stage, several of which occur together as parts of large colonies of thousands of zooids. These colonies may attain lengths over 1 m, but are fragile and are rarely collected intact. In most genera of doliolids, the life cycle begins with a sexually produced larva, which becomes the oozooid stage. This stage feeds initially, but then begins budding off the trophozooid and phorozooid stages, thus forming the colony. During this process the oozooid loses its branchial basket and gut, and transforms into the 'old nurse' stage, whose function is to swim by jet propulsion and pull the attached colony along behind it. Contractions of the body muscles produce short exhalent pulses that move the colony rapidly. The trophozooids in the colony filter-feed to support themselves and the nurse. The phorozooids grow attached to the colony, but then break free to lead independent lives and produce asexually a small group of gonozooids. These eventually break free from the phorozooid, and become the sexually reproducing stages (hermaphrodites?) that produce the larvae and begin the whole cycle again (Figure 4B).
Figure 4. Pelagic Tunicates. (A) Megalocercus huxleyi, a larvacean of about 5 mm body length, house length about 4 cm. (B) Dolioletta gegenbauri, portion of a colony showing gastrozooids and phorozooids, individuals 2–5 mm long, colonies up to 1 m. (C) Salpa maxima, solitary generation salp, up to 25 cm long. (D) Salpa maxima, chain of aggregate generation salps; orange dots are guts of salps; individuals are to 15 cm, chains up to 10 m long. (E) Pegea socia, aggregate generation salp with attached embryo of solitary generation; aggregate 7 cm, embryo about 1 cm. (F) Traustedtia multitentaculata, solitary generation salp with appendages of uncertain function, about 3 cm long. (All photographs by L. P. Madin.)
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Free-Living Protozoa
Genoveva F. Esteban , ... Alan Warren , in Thorp and Covich's Freshwater Invertebrates (Fourth Edition), 2015
Ciliated Protozoa
This diverse and distinctive group of protozoa uses cilia for locomotion and feeding (Figures 7.1, 7.4, and 7.5 ). They demonstrate a considerable adaptive radiation of feeding mechanisms and cell morphologies, e.g., the ribbon-shaped forms Tracheloraphis and Geleia adapted for life in the marine interstitial (Fenchel, 1987). The smallest species tend to feed on bacteria-sized particles, and the larger species on unicellular algae, filamentous cyanobacteria (Figure 7.1), other protozoa, and even rotifers and other microzooplankton. The raptorial feeders (e.g., Prorodon) use a simple mouth to catch diatoms, dinoflagellates, and other large food items individually; some (e.g., Lacrymaria) kill motile prey, whereas others (e.g., Chilodonella) ingest diatoms and other elongate food particles from surfaces. The filter feeders (e.g., Cyclidium) use fine-mesh filters to sieve suspended bacteria, and some of these ciliates (e.g., Paramecium and Tetrahymena) thrive in habitats with very high bacterial concentrations. Other ciliates (e.g., Pleuronema and Tintinnopsis) use coarser filters to collect small algae (Figure 7.5). In many ciliates (e.g., Oxytricha, Aspidisca, and Strombidium), a row of membranelles generates a water current and acts as a relatively coarse feeding filter, and some of these ciliates (e.g., Euplotes) feed most efficiently if they are raised on cirri (fused cilia). Many ciliates are typically sessile and aligned perpendicular to the substrate (e.g., Stentor and Vorticella). Diffusion feeders (e.g., Podophrya and other Suctoria) catch swimming prey (usually other protozoa) that collide with their sticky tentacles.
FIGURE 7.5. A selection from the variety of form and function in ciliated protozoa.
(a) A bacteria-feeding tetrahymenid (Tetrahymena; length ∼0.05 mm). (b) A small scuticociliate (Cyclidium; length ∼0.02 mm) filter-feeding on bacteria. (c) A bacteria-feeding colpodid ciliate (Colpoda; length ∼0.02 mm) from soil. (d) An anaerobic, typically benthic plagiopylid feeding on bacteria (Plagiopyla; length ∼0.1 mm). (e) A large haptorid (Dileptus; length ∼0.4 mm) with toxicysts used to kill motile prey (flagellates and other ciliates) prior to ingestion. (f) A planktonic, loricate, diatom-feeding tintinnid (Tintinnopsis; length ∼0.1 mm). (g) A gymnostome ciliate (Coleps; length ∼0.07 mm) about to ingest a dinoflagellate. (h) A diatom-feeding karyorelictid (Remanella; length ∼0.12 mm) typical of the marine interstitial. (i) A "hoover-feeding" cyrtophorid (Chilodonella; length ∼0.04 mm) ingesting a diatom. (j) A planktonic oligotrich (Halteria; diameter ∼0.04 mm) filter-feeding on small algae. (k) A "diffusion-feeding" suctorian (Podophrya; diameter ∼0.04 mm) with a flagellate trapped on a feeding tentacle.
More is probably known about the biodiversity and ecology of free-living ciliates than any other protozoan group. There are several reasons for this, including the distinctive and immediately recognizable ciliate morphology and swimming behavior and the relative ease with which many species can be cultured. They are a group of predominantly free-living protozoa. A few are parasites (e.g., Ichthyophthirius infects fish and Balantidium coli is a human endoparasite), but most species are either free-living or harmless commensals of aquatic invertebrates. For example, the ciliate epifauna of crustaceans is particularly diverse. Free-living species are known from all natural aquatic habitats in which temperatures are <45 °C and where there is sufficient food, including freshwater and marine sediments, oceanic sinking detritus, anaerobic municipal landfill sites, and sewage treatment plants. They are abundant (up to 105/ml) in activated sludge plants, in which they consume bacteria and also flocculate bacteria and other suspended particulate matter. These activities aid the clarification of the effluent and the formation of sludge. Ciliates also have a role as indicators of the level of organic pollution in river water (Corliss, 1979, 2000; Lynn, 2010).
Most ciliates are in the size range 0.02–2 mm, so they are generally larger than the heterotrophic flagellates and other nanoplankton (0.002–0.02 mm) on which many of them feed. Planktonic ciliates are relatively abundant (1–100/ml) and important grazers of nanoplankton in marine waters and fresh waters. They are key grazers within the "microbial loop" that is responsible for the rapid remineralization of organic matter in the water column. The diet of planktonic metazoans includes ciliates, although the quantitative significance of this link is unclear. Benthic ciliates are often abundant (>1000/ml) and usually the most important grazers in freshwater sediments (especially in lakes) and marine sandy sediments of inshore waters.
Many ciliate species harbor prokaryotic and/or eukaryotic symbionts. At the oxic–anoxic boundary in the water column of lakes, most ciliates may harbor sufficient endosymbiotic algae (Chlorella) to render the consortia capable of net photosynthesis in dim light. The marine interstitial ciliate Kentrophoros carries ectosymbiotic chemolithotrophic sulfide-oxidizing bacteria, and most anaerobic ciliate species harbor methanogenic bacteria that act as a sink for (potentially inhibitory) H2 produced by the ciliate.
Many ciliate species are microaerobic, and they seek out the (microbe-rich) oxic–anoxic boundary in sediment or the water column of ponds and lakes. Some of these species are facultative anaerobes (in Cyclidium, Euplotes, Strombidium, and Paranophrys). There are less than 100 known species of anaerobic free-living ciliate species, mainly in the genera Metopus, Caenomorpha, Saprodinium, Epalxella, Trimyema, and Plagiopyla. It is likely that all these contain hydrogenosomes (Fenchel and Finlay, 1995).
Anaerobic ciliates also live as endocommensals in the enlarged forestomach (rumen) of ruminants and in the cecum of other mammals with postgastric fermentation. It is unlikely that any of these ciliates (e.g., in Dasytricha, Entodinium, and Polyplaston) are capable of a free-living existence. Rumen ciliates are typically extremely abundant (105–106/ml of rumen liquor). They consume bacteria and microscopic fragments of grass and other plant material. Some species can degrade cellulose and other structural carbohydrates, and the endosymbiotic methanogenic bacteria in some species contribute to the methane emitted from ruminants. The economic significance of such large numbers of ciliates living in the rumen is unclear.
Ciliates also live in soil, in which all species can probably produce desiccation-resistant cysts (e.g., Colpoda). The abundance of active ciliates in soil is extremely variable and reflects repeated cycles of cyst formation and excystment in response to fluctuating physical factors such as the level of soil moisture. Many species found in soil are frequently found in other (predominantly freshwater) habitats.
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Marine Biogeochemistry
Laurence P. Madin , G. Richard Harbison , in Encyclopedia of Ocean Sciences (Third Edition), 2019
Doliolida
This order of the Thaliacea comprises 6 genera and more than 20 species of small (2–10 mm), barrel shaped animals with circumferential muscle bands. The filter feeding mechanism is similar to that of pyrosomes, with currents generated by ciliary beating passing through a mucous net supported on the branchial basket. The life cycle involves up to five asexual and one sexual stage, several of which occur together as parts of large colonies comprising thousands of zooids. These colonies may attain lengths over 1 m, but are fragile and rarely collected intact. In most genera of doliolids, the life cycle begins with a sexually produced larva, which becomes the oozooid stage. This stage feeds initially, but then begins budding off the trophozooid and phorozooid stages, thus forming the colony. During this process the oozooid loses its branchial basket and gut, and transforms into the "old nurse" stage, whose function is to swim by jet propulsion and pull the attached colony along behind it. Contractions of the body muscles produce short exhalent pulses that move the colony rapidly. The trophozooids in the colony filter-feed to support themselves and the nurse. The phorozooids grow attached to the colony, but then break free to lead independent lives and produce asexually a small group of gonozooids. These eventually break free from the phorozooid, and become the sexually reproducing stages that produce the larva and begin the whole cycle again. Recently several unusual doliolid species with enigmatic life histories have been described from the mesopelagic zone (Fig. 4B ).
Fig. 4. Pelagic Tunicates. (A) Megalocercus huxleyi, a larvacean about 5 mm body length, house length about 4 cm; (B) Dolioletta gegenbauri, portion of a colony showing gastrozooids and phorozooids, individuals 2–5 mm long, colonies up to 1 m; (C) Salpa maxima, solitary generation salp, up to 25 cm long; (D) S. maxima, chain of aggregate generation salps, orange dots are guts of salps, individuals to 15 cm, chains up to 10 m long; (E) Pegea socia, aggregate generation salp with attached embryo of solitary generation, aggregate 7 cm, embryo about 1 cm; F. Traustedtia multitentaculata, solitary generation salp with appendages of uncertain function, about 3 cm long.
Source: All photos by L.P. Madin.Salpida. This order (with 12 genera and about 40 species) is of larger filter feeding animals, also with circumferential muscle bands. The salps alternate between two forms, an asexually budding solitary (oozooid) stage and a sexually reproducing aggregate (blastozooid) stage. The aggregate salps usually remain connected together in chains or whorls of various types. Swimming is by jet propulsion, produced by a pulsed water current generated by rhythmic contraction of body muscles. Food particles are strained from the water passing through the body cavity by a mucous filter, which is continuously secreted and ingested. The individual animals range in size from 5 to over 100 mm, and chains can be several m long. Some species of salps develop dense populations and have a significant impact on ocean food chains by their grazing. Some are also strong vertical migrators, moving from the surface zone to midwater depths on a daily basis (Fig. 4C–F).
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Ecomorphology of Feeding in Coral Reef Fishes
Peter C. Wainwright , David R. Bellwood , in Coral Reef Fishes, 2002
2. FUNCTIONAL MORPHOLOGY OF PREY CAPTURE IN TELEOST FISHES
Whether capturing prey by ram, suction, or manipulation all teleost fishes use a common feeding apparatus constructed of a homologous network of muscles, bones, and soft connective tissue. A dominant feature of the fish feeding mechanism that sets it apart from other vertebrate groups is the large number of moving elements. More than 20 major skeletal components are put into motion by about 40 muscles. Fortunately, it is possible to focus on a relatively small number of elements to embody the major features of feeding mechanics.
Ram and suction feeding involve the rapid expansion of the buccal cavity by nearly simultaneous elevation of the neurocranium, lateral expansion of the cheek bones (suspensorium), ventral depression of the floor of the buccal cavity by depression of hyoid elements, and frequently some anterior expansion by depression of the lower jaw and protrusion of the upper jaw (Figs. 3 and 4). The major muscles that participate in these actions include the epaxialis, which dorsally rotates the neurocranium on the vertebral column; the sternohyoideus, which retracts the hyoid bar; the hypaxialis, which retracts and stabilizes the pectoral girdle; and the levator arcus palatini, which laterally rotates the suspensorium (Fig. 6) (Liem, 1970; Sanderson, 1988; Wainwright and Turingan, 1993). Muscles that produce the reverse actions (adduction of the jaws and suspensorium) include the adductor mandibulae and adductor arcus palatini muscles (Fig. 6) (Ballintijn et al., 1972; Friel and Wainwright, 1999). During suction feeding buccal expansion may be coupled with strong adduction of the gill bars to prevent communication between the buccal and opercular cavities (Lauder 1980, 1983a). Adduction of the jaws during manipulation behaviors involves the actions of the adductor mandibulae complex in a variety of rasping, nipping, scraping, and forceful biting actions (Wainwright and Turingan, 1993; Ralston and Wainwright, 1997; Alfaro and Westneat, 1999).
FIGURE 6. Plots of scores of 16 Great Barrier Reef labrid species in the mechanical space of the four-bar linkage systems of the oral jaws and the hyoid apparatus. Separate Principal Component Analyses were run on each linkage system from a data set of 228 specimens from 81 wrasse species of the GBR region. In each analysis the first principal component (PC) was a size factor and the second PC was the major shape axis. Mean PC2 scores were calculated for each species and examples of several major trophic types are shown. In most (but not all) cases, the species shown for each trophic group are thought to represent independent origins of that trophic habit. Note that most trophic groups tend to occupy specific regions of four-bar space. The one exception is the zooplanktivores, which are represented by considerable mechanical diversity in both four-bar systems. In order of increasing anterior jaws PC2 score, the species shown are Cheilio inermis, Oxycheilinus digrammus, Anampses neuguinaicus, Labropsis australis, Labrichthys unilineatus, Pseudojuloides cerasinus, Macropharyngodon meleagris, Hemigymnus melapterus, Halichoeres ornatissimus, Leptojulis cyanopleura, Thalassoma jansenii, Choerodon jordani, Cheilinus fasciatus, Bodianus loxozonus, and Cirrhilabrus punctatus.
Data are from Wainwright, et al. (2002b). With permission from Wainwright and Friel, 2000, Wiley-Liss, Inc., a subsidiary of John Wiley & Sons, Inc. Copyright © 2002Teeth function differently during ram/suction feeding than they do during manipulation feeding. For ram/suction feeding the teeth mainly act as a friction device, preventing captured prey from escaping back out of the mouth. Large teeth in ram/suction feeders are typically raptorial and associated with capture of particularly large and elusive prey, such as other fishes. Teeth are more diverse in manipulators, often reflecting the method used by the fish to extract prey from the substratum. In some taxa, such as many wrasses, the teeth are relatively large and recurved, though not sharp, as seen in piscivores, and are used in gripping relatively large invertebrate prey from within the reef. Manipulating predators (e.g., the Chaetodontidae) of smaller invertebrate prey possess smaller teeth, sometimes arranged in pads. Herbivores show variation from the beaklike structures of the parrotfish to single rows of complex crowned teeth as seen in the surgeonfish. Teeth of manipulators frequently have significant iron deposits (Motta, 1984b, 1987; Suga et al., 1989) that appear to enhance tooth strength.
Aspects of the functional morphology of feeding have been studied in representatives of many of the major coral reef groups, including the Blenniidae (Goldschmid and Kotrschal, 1985; Kotrschal, 1988, 1989a,b), the Labridae (Rognes, 1973; van Hasselt, 1978, 1979a,b, 1980; Tedman, 1980a,b; Sanderson, 1988, 1990, 1991; Wainwright, 1988; Westneat and Wainwright, 1989; Westneat, 1990, 1991, 1994, 1995; Clifton and Motta, 1998), the Scaridae (Monod, 1951; Board, 1956; Tedman, 1980a,b; Clements and Bellwood, 1988; Gobalet, 1989; Bellwood and Choat, 1990; Bellwood, 1994; Bullock and Monod, 1997; Alfaro and Westneat, 1999), the Chaetodontidae (Motta, 1982, 1984b, 1985, 1987, 1988, 1989; Sano, 1989), the Serranidae (Mullaney and Gale, 1996; Viladiu et al., 1999), the Mullidae (Gosline, 1984; McCormick, 1993, 1995; McCormick and Shand, 1992), the Acanthuridae (Jones, 1968; Purcell and Bellwood, 1993), Tetraodontiformes (Sarkar, 1960; Turingan and Wainwright, 1993; Wainwright and Turingan, 1993, 1997; Turingan, 1994; Turingan et al., 1995; Friel and Wainwright, 1997, 1998, 1999; Ralston and Wainwright, 1997; Wainwright and Friel, 2000), the Pleuronectiforms (Gibb, 1995, 1996), and several elasmobranchs (Motta and Wilga, 1999; Wilga and Motta, 2000).
An interesting aspect of the ram/suction vs. manipulation categorization of prey capture mechanisms is that the performance attributes that are expected to enhance each feeding mode are different. In general, ram/suction feeding is expected to emphasize speed and power of jaw and head motion. Manipulation highlights forcefulness of movements, and fine motor control of the jaws in the case of taxa that pick at individual items. As we shall see in the next section, force and speed of motion trade off in the mechanical systems that underlie motion in the fish prey capture apparatus. This design trade-off reveals a major dimension of fish skull diversity, and has considerable ecological consequences.
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Scallops
Shawn M.C. Robinson , ... Norman J. Blake , in Developments in Aquaculture and Fisheries Science, 2016
Spat Husbandry
Recent work has focused on the dietary requirements and feeding regimes of raising post-metamorphic spat in the nursery phase. Metamorphosis results in structural changes including the loss of the velum, which is used in larval feeding, and the development of a filter-feeding mechanism involving the new gills and cilia. The ingestion rate of the alga Isochrysis galbana by juvenile scallops was described by the relationship IR=1.5×104 W×1.22 where IR=ingestion rate (cells min−1) and W=body weight (g) (Strickland and Dabinett, 1993); however, Lesser et al. (1991) demonstrated that juvenile scallops feed selectively. Juveniles fed on an unialgal diet selected particles based on size, whereas for those fed on multialgal diets, selection did not appear to be based on size, but rather on other characteristics of the algae or pre-ingestive sorting. A mixed algal diet of three species resulted in better growth than a unialgal or paired algal diet for 2 mm spat (Gillis and Dabinett, 1989). Similar findings were reported by Parrish et al. (1999) and Milke et al. (2004) who found that mixed species diets worked better for growing juveniles and that omega-3 and omega-6 fatty acids were essential for providing an optimal diet. Mallet (1989) found analogous results for larvae (see below). Hollett and Dabinett (1989) have found that a ratio of 45 cells·µL−1 of a multialgal diet provided the best growth and highest spat growth efficiencies. Their experimental range of algal densities was 12 to 87 cells·µL−1.
Several studies have examined the physical conditions under which juveniles should be grown. Frenette and Parsons (2001) and Frenette (2004) showed that lethal temperatures were above 18 °C and lethal salinities below 25 PSU. The highest filtering rates for juveniles were found at 13 °C and a salinity of 32 PSU (Frenette et al., 2002; Frenette 2004). Un-ionised ammonia levels in the water were found to inhibit feeding of juvenile scallops at levels of 0.54 mg·L−1 and above (Dabinett et al., 1998; Grecian et al., 2001b). There was some evidence of size-related tolerances of ammonia as well as temperature-dependent sensitivity (Abraham et al., 1996).
Kean-Howie et al. (1989, 1991) studied the nutritional physiology of juvenile sea scallops and successfully developed a microparticulate food capsule upon which juveniles will feed. These studies have provided the basis for a more complete understanding of the specific dietary requirements of scallops, which have enabled researchers to manipulate the different dietary components of the microparticles. Coutteau et al. (1996) have used lipid emulsions as feed additives to increase the levels of n-3 highly unsaturated fatty acids in Placopecten and resulting in 20% more lipid being incorporated into juvenile sea scallops compared to the algal-fed controls.
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Source: https://www.sciencedirect.com/topics/earth-and-planetary-sciences/feeding-mechanism
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