The Reef Aquarium Consequences of Suspension-Feeding as a Way of Life
By Ronald L. Shimek
Probably the most common way that organisms make a living in the marine environment is by feeding on particulate material suspended in the water. This way of gathering nutrition is fundamental to marine and aquatic ecosystems and probably due to the density of water and its relationship to the density of living tissue. Simply put, many organisms can float or easily swim in water, and this waterborne life constitutes a potential food source to many animals. The exploitation of floating food has been the driving force of much of the evolution occurring in marine ecosystems. Many of the organisms maintained by reef aquarists are suspension-feeders and an understanding of their morphology and natural history and the constraints this places on the organisms is important to successful husbandry of these organisms.
-The Methodology of Passive Suspension Feeding
There are passive and active suspension-feeding organisms, and these two ways of feeding are fundamentally different, particularly as regards the amount of energy expended in the capture of prey. In passive suspension-feeding, the organism may extend a body part to allow a food item to impinge on it and then expend some energy transferring the food to the mouth, but if so, those activities may be the extent of the energy expenditures of the critter. In active suspension-feeding the organism expends significant energy in procuring food by pumping, secreting filter materials or actively grabbing prey.
Passive suspension-feeding animals generally depend on ambient water motion to bring food to them. While these animals expend very little energy to bring water to themselves, they may expend a lot of energy in the actual capture of the food item once it has reached them. Numerous groups of organisms can be considered to contain passive suspension-feeding organisms. Among them are the cnidaria, such as the soft corals, sea anemones, and corals, many annelid worms and some echinoderms such as sea cucumbers and brittle stars (Lewis, 1977, 1981, 1992; Kozloff, 1990, Ruppert and Barnes, 1994, Nybakken, 1997).
Orientation to currents is important in gorgonians. These animals are dependent on food collection from water flowing over them, and their orientation to that water flow is important. Mobile animals such as sea anemones, annelid worms or sea cucumbers may orient themselves or their filters in the water to maximize their filtering efficiency, but immobile animals such as corals and gorgonians cannot change the orientation. In nature these animals are often found specifically oriented to prevailing currents. They grow in these particular orientations after they metamorphose from the larval state. As they fasten themselves securely to the substrate, they will not be able to change their orientation once they grow into it (Leversee, 1976; Warner, 1976; Lasker, 1981; Patterson, 1984, 1991; Sebens, and Johnson, 1991; Johnson, and Sebens; 1993; Lesser, et al., 1994; Sebens et al., 1996, 1997).
Unfortunately for these animals, after collection we hobbyists are often unable to discern what is the correct orientation for optimal feeding. The animals are placed haphazardly, but perhaps aesthetically, in our systems and we hope they will thrive. Particularly for animals such as gorgonians, the water flow direction and strength are important. Fortunately, finding an acceptable water flow regime is not too difficult, but it does take a little time and patience. After the animal has been placed and acclimated sufficiently to the tank that it extends its polyps, newly hatched brine shrimp should be placed in the tank, and the behavior of the gorgonian closely monitored. Generally, a magnifying glass helps to observe the behavior of the ones with smaller polyps. If the orientation to the tank's current flow is acceptable, it will be possible to watch the polyps catch and eat the small crustaceans. If the water flow is not acceptable, then either few crustaceans will move by the polyps or they will move by in eddies or other mini-currents in such a manner that the polyps simply cannot catch them. If the latter situation occurs, the gorgonian, or perhaps the powerhead that generates the current, will need to be moved until the proper water movement occurs. The cnidarians capture, subdue and adhere to prey by the use of nematocysts, and these can be quite effective in maintaining a grip on the prey, even in strong currents.
Many animals such as sea cucumbers often use mucus to catch and hold on to prey. Mucus is a material made by chemically combining sugars and proteins, and as we all know it is sticky. Suspension-feeding sea cucumbers are able to feed by secreting mucus on expanded tentacles located around the mouth. These tentacles are typically repeatedly branched until they can look like miniature tree branches. This tree-like branching pattern is referred to as dendritic branching, and the suspension-feeding cukes are known as "Dendrochirotes," a name which literally means "tree-like fingers." The dendrochirotes extend their sticky tentacles up into the water and leave them there for a while, then bend them back, one at a time, and stick them in the mouth where the mucus and adherent food are licked off (Bakus, 1973; Kozloff, 1990; Ruppert and Barnes, 1994).
Possibly the most unusual way for organisms to capture food is by the use of aerosol properties. This is the method by which numerous brittle stars capture small particulate materials. The brittle stars extend their arms up into the water currents. As the water flows past the arms, it gives the arms a static electrical charge, much as one can get a static charge by shuffling across certain carpets. Particles in the water are also charged and if they are close enough to the arm they will tend to be attracted to it. These small food particles literally are electrostatically pulled out of the water column to stick to the arm, similar to dust particles in an electrostatic air filter (LaBarbera, 1978). Once on the brittle star's arm, the food particles are passed from tube foot to tube foot down to the mouth.
-The Methodology of Active Suspension Feeding
Active suspension-feeding animals generate a current of their own to bring food into their traps, and unlike the passive feeders can be located in areas of weak currents. These animals include many larvae, some rotifers, annelid worms, crustaceans, many mollusks, but especially bivalves, and tunicates (Kozloff, 1990; Ruppert and Barnes, 1994). Most of them use some kind of ciliary-mucus type feeding.
Basically the animal makes some kind of mucus net and moves water through the net by the use of cilia. Probably the most familiar of the ciliary-mucus suspension feeders are clams. Almost all shallow water bivalves feed by using a ciliary-mucus feeding process. This process of feeding can be used for deposit feeding on sediments as well as feeding on material suspended in the water column, and many bivalves are sediment eaters (Olaffson, 1986).
Most clams, however, feed on particulate material suspended in the water. The bivalves have modified gills that, in addition to having a respiratory function, are used as filter-feeding structures. The structure of the clam gill varies a bit, but it can be visualized as having filaments with very thin openings between them. In many species there are cross connections between the filaments which creates a sieve-like mesh. Virtually all surfaces of the gills are covered in cilia and bathed in mucus. These cilia move the water into the clam, often through a siphon, and the water is forced through the mesh. Particles collect on the gills and get conveyed to the mouth in a conveyor belt of mucus moved by yet other cilia (Kozloff, 1990; Ruppert and Barnes, 1994).
The gills are generally visible in a clam if it gapes, and look like sheets of tissue located in the middle of the cavity between the shells. An aquarist can see the gills in a tridacnid clam simply by looking into the internal cavity through one of the large siphons. Incidentally, all clams need to feed, including the tridacnids. These species can get a lot of their caloric needs from their zooxanthellae, but particularly when the clams are small they don't have enough tissue to support the large populations of the algae necessary to produce sufficient food. Consequently in tridacnids, feeding is more important for smaller clams than larger ones (Klumpp and Lucas; 1994; Griffiths, and Klumpp, 1996; Hoegh-Guldberg, 1996).
A different type of ciliary-mucous feeding is seen in feather-duster worms. Here there is an elaborate crown of highly-branched tentacles on the head. These tentacles are lined with cilia which pull the water through the tentacle crown. Food is caught on the tentacles and moved to a food groove in the center of each tentacle. Moving mucus in the food groove transports the food to the mouth (Fauchald and Jumars, 1979; Kozloff, 1994; Ruppert and Barnes, 1994).
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By Ronald L. Shimek
Probably the most common way that organisms make a living in the marine environment is by feeding on particulate material suspended in the water. This way of gathering nutrition is fundamental to marine and aquatic ecosystems and probably due to the density of water and its relationship to the density of living tissue. Simply put, many organisms can float or easily swim in water, and this waterborne life constitutes a potential food source to many animals. The exploitation of floating food has been the driving force of much of the evolution occurring in marine ecosystems. Many of the organisms maintained by reef aquarists are suspension-feeders and an understanding of their morphology and natural history and the constraints this places on the organisms is important to successful husbandry of these organisms.
-The Methodology of Passive Suspension Feeding
There are passive and active suspension-feeding organisms, and these two ways of feeding are fundamentally different, particularly as regards the amount of energy expended in the capture of prey. In passive suspension-feeding, the organism may extend a body part to allow a food item to impinge on it and then expend some energy transferring the food to the mouth, but if so, those activities may be the extent of the energy expenditures of the critter. In active suspension-feeding the organism expends significant energy in procuring food by pumping, secreting filter materials or actively grabbing prey.
Passive suspension-feeding animals generally depend on ambient water motion to bring food to them. While these animals expend very little energy to bring water to themselves, they may expend a lot of energy in the actual capture of the food item once it has reached them. Numerous groups of organisms can be considered to contain passive suspension-feeding organisms. Among them are the cnidaria, such as the soft corals, sea anemones, and corals, many annelid worms and some echinoderms such as sea cucumbers and brittle stars (Lewis, 1977, 1981, 1992; Kozloff, 1990, Ruppert and Barnes, 1994, Nybakken, 1997).
Orientation to currents is important in gorgonians. These animals are dependent on food collection from water flowing over them, and their orientation to that water flow is important. Mobile animals such as sea anemones, annelid worms or sea cucumbers may orient themselves or their filters in the water to maximize their filtering efficiency, but immobile animals such as corals and gorgonians cannot change the orientation. In nature these animals are often found specifically oriented to prevailing currents. They grow in these particular orientations after they metamorphose from the larval state. As they fasten themselves securely to the substrate, they will not be able to change their orientation once they grow into it (Leversee, 1976; Warner, 1976; Lasker, 1981; Patterson, 1984, 1991; Sebens, and Johnson, 1991; Johnson, and Sebens; 1993; Lesser, et al., 1994; Sebens et al., 1996, 1997).
Unfortunately for these animals, after collection we hobbyists are often unable to discern what is the correct orientation for optimal feeding. The animals are placed haphazardly, but perhaps aesthetically, in our systems and we hope they will thrive. Particularly for animals such as gorgonians, the water flow direction and strength are important. Fortunately, finding an acceptable water flow regime is not too difficult, but it does take a little time and patience. After the animal has been placed and acclimated sufficiently to the tank that it extends its polyps, newly hatched brine shrimp should be placed in the tank, and the behavior of the gorgonian closely monitored. Generally, a magnifying glass helps to observe the behavior of the ones with smaller polyps. If the orientation to the tank's current flow is acceptable, it will be possible to watch the polyps catch and eat the small crustaceans. If the water flow is not acceptable, then either few crustaceans will move by the polyps or they will move by in eddies or other mini-currents in such a manner that the polyps simply cannot catch them. If the latter situation occurs, the gorgonian, or perhaps the powerhead that generates the current, will need to be moved until the proper water movement occurs. The cnidarians capture, subdue and adhere to prey by the use of nematocysts, and these can be quite effective in maintaining a grip on the prey, even in strong currents.
Many animals such as sea cucumbers often use mucus to catch and hold on to prey. Mucus is a material made by chemically combining sugars and proteins, and as we all know it is sticky. Suspension-feeding sea cucumbers are able to feed by secreting mucus on expanded tentacles located around the mouth. These tentacles are typically repeatedly branched until they can look like miniature tree branches. This tree-like branching pattern is referred to as dendritic branching, and the suspension-feeding cukes are known as "Dendrochirotes," a name which literally means "tree-like fingers." The dendrochirotes extend their sticky tentacles up into the water and leave them there for a while, then bend them back, one at a time, and stick them in the mouth where the mucus and adherent food are licked off (Bakus, 1973; Kozloff, 1990; Ruppert and Barnes, 1994).
Possibly the most unusual way for organisms to capture food is by the use of aerosol properties. This is the method by which numerous brittle stars capture small particulate materials. The brittle stars extend their arms up into the water currents. As the water flows past the arms, it gives the arms a static electrical charge, much as one can get a static charge by shuffling across certain carpets. Particles in the water are also charged and if they are close enough to the arm they will tend to be attracted to it. These small food particles literally are electrostatically pulled out of the water column to stick to the arm, similar to dust particles in an electrostatic air filter (LaBarbera, 1978). Once on the brittle star's arm, the food particles are passed from tube foot to tube foot down to the mouth.
-The Methodology of Active Suspension Feeding
Active suspension-feeding animals generate a current of their own to bring food into their traps, and unlike the passive feeders can be located in areas of weak currents. These animals include many larvae, some rotifers, annelid worms, crustaceans, many mollusks, but especially bivalves, and tunicates (Kozloff, 1990; Ruppert and Barnes, 1994). Most of them use some kind of ciliary-mucus type feeding.
Basically the animal makes some kind of mucus net and moves water through the net by the use of cilia. Probably the most familiar of the ciliary-mucus suspension feeders are clams. Almost all shallow water bivalves feed by using a ciliary-mucus feeding process. This process of feeding can be used for deposit feeding on sediments as well as feeding on material suspended in the water column, and many bivalves are sediment eaters (Olaffson, 1986).
Most clams, however, feed on particulate material suspended in the water. The bivalves have modified gills that, in addition to having a respiratory function, are used as filter-feeding structures. The structure of the clam gill varies a bit, but it can be visualized as having filaments with very thin openings between them. In many species there are cross connections between the filaments which creates a sieve-like mesh. Virtually all surfaces of the gills are covered in cilia and bathed in mucus. These cilia move the water into the clam, often through a siphon, and the water is forced through the mesh. Particles collect on the gills and get conveyed to the mouth in a conveyor belt of mucus moved by yet other cilia (Kozloff, 1990; Ruppert and Barnes, 1994).
The gills are generally visible in a clam if it gapes, and look like sheets of tissue located in the middle of the cavity between the shells. An aquarist can see the gills in a tridacnid clam simply by looking into the internal cavity through one of the large siphons. Incidentally, all clams need to feed, including the tridacnids. These species can get a lot of their caloric needs from their zooxanthellae, but particularly when the clams are small they don't have enough tissue to support the large populations of the algae necessary to produce sufficient food. Consequently in tridacnids, feeding is more important for smaller clams than larger ones (Klumpp and Lucas; 1994; Griffiths, and Klumpp, 1996; Hoegh-Guldberg, 1996).
A different type of ciliary-mucous feeding is seen in feather-duster worms. Here there is an elaborate crown of highly-branched tentacles on the head. These tentacles are lined with cilia which pull the water through the tentacle crown. Food is caught on the tentacles and moved to a food groove in the center of each tentacle. Moving mucus in the food groove transports the food to the mouth (Fauchald and Jumars, 1979; Kozloff, 1994; Ruppert and Barnes, 1994).
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