This unedited article first appeared in the July/August 1990 issue of Aquarium Fish Magazine.

Nutritional Requirements of Reef Systems

by J. Charles Delbeek M.Sc. The nutritional requirements of the various organisms that can occur in reef systems is extremely varied and/or difficult to ascertain. We are no longer dealing only with the different feeding habits of the fish that we are keeping but also various orders of invertebrates, each with their own peculiarities. Add to this mass of confusion the fact that very little is known about the nutritional requirements of these organisms and one can quickly see that the topic of nutrition in reef systems is a most intimidating one. That is why I find the type of advertising used by some manufacturers of "reef foods" rather distressing. Very few of these products are based on any form of original research and some appear to contain additional chemical additives (e.g. glutamic acid, proline or alanine) that will elicit a feeding response in some invertebrates but have questionable nutritional value. It will not be the purpose of this article to tell you what to feed each organism but more to outline the various feeding modes that one will encounter and to help you decide how best to deal with them. To go into detail on the diet of each organism would require a complete book. There are a few references that I will give at the end of this article that will help you get started. Read as much as you can, ask questions of your dealer and friends. Don't be afraid to experiment when it comes to feeding but remember to ALWAYS FEED IN MODERATION. The general approach taken in feeding organisms in captivity is to closely study the diets of these organisms in the wild and to duplicate this as much as possible. This approach works quite well with most fish and some invertebrates. However, many natural diets are almost impossible to duplicate e.g. coral polyps and sponges which are fed on by a multitude of fish and invertebrates. Another problem comes with corals where there are few studies published on the diets of corals and indeed, much debate on how much and even if corals need to feed. There are various feeding mechanisms used by the inhabitants of our reef aquariums. In some instances the same organism may use more than one feeding strategy which is probably an adaptation to ensure that as much nutrition as possible can be extracted from the nutrient poor area that a reef is. This is the case with many coral species and for this reason, coral feeding strategies will be dealt with separately. Many organisms actually trap and/or catch and digest other organisms such as shrimp, copepods and marine worms. Some organisms, commonly known as filter feeders, feed by trapping much smaller particles ranging in size from copepods and other zooplankton to detritus and feces. Those that feed on detritus either digest the bacteria living on them or the particulate organics that coat such particles (Wotton 1988). The method of feeding that has probably elicited the most debate is the one involving the relationship between organisms such as corals, bivalves, sponges and the symbiotic algae that live in their tissues. Contrary to popular belief these algae (usually referred to as zooxanthellae) are not green but are in fact gold-coloured dinoflagellates of the genus Symbiodinium. At one point it was thought that there was only a single species, Symbiodinium microadriaticum but it has recently been shown that there are in fact several strains of zooxanthellae; some fast growing, some slow (Blank and Trench, 1985; Trench 1979). The final method of feeding that we will examine in this article is the direct uptake of nutrients such as ammonia, phosphate, proteins and dissolved organics from the water, through the body wall. The majority of the animals that we keep in our aquariums, such as fish, shrimp, starfish and urchins fall into the first category of feeding. These organisms may be omnivorous, herbivorous or carnivorous in nature and depending on their size and appetite may be able to fend for themselves in a reef tank. More common though is the situation where additional feedings are required. For those organisms that are carnivorous or omnivorous good quality flake foods with low amounts of carbohydrates, large amounts of protein and the proper lipids (fish oils as opposed to plant oils) should suffice. The same goes for frozen foods. The problem with any prepared food is the issue of vitamins, pigments and other micronutrients. Many of these are destroyed during the manufacturing process and need to be added to the feed later. The problem is that these substances have a limited shelf life. By the time the food has gone through the hands of the manufacturer, wholesaler, retailer and finally to you, the majority of these substances may have been lost through oxidization. As a result, your organisms will not be getting the full nutritional value that you may think. Fortunately, recent developments in aquaculture are making their way into the pet industry and many foods are now available that contain vitamins stabilized against deterioration. The main vitamin available in this form is vitamin C and there are a number of foods and vitamin supplements now advertising its use. There are a number of precautions you should take when purchasing and using prepared foods. First of all only buy food from a dealer who regularly rotates their stock and who buys direct from the manufacturer or from a wholesaler who rotates their stock. Most flake foods have a shelf live of about 6 months if not refrigerated. Secondly, as soon as you open your food make sure you do not leave it close to the tank or in other warm, humid areas. Keep the container tightly sealed and in the frige. Some hobbyists soak their foods in vitamin supplements before feeding. This is a good idea only if the vitamins are fresh of course. Baby vitamins have been used with some success and have the added benefit of having expiry dates on the jars. Of course the best foods are those that are as fresh as possible such as raw shrimp, clams, mussels and marine fish. For large predators such as groupers and lionfish, it is extremely important to avoid feeding them freshwater fish as a staple food. Try and wean them from these foods and onto prepared marine foods or fresh shrimp etc. The reason for this is that freshwater fish lack certain essential fatty acids that are mandatory for marine fish. A lionfish may live for years on a diet of goldfish then one day it will inexplicably die. When these fish are examined they are usually found to have fatty livers caused by the abundance of improper fatty acids in their diet. Most omnivorous invertebrates can be fed on prepared foods supplemented with pieces of shrimp, mussels or clams. Organisms that fall into this category include Brittle Stars, Fromia sp. starfish, sea urchins, shrimps and crabs. Some of these will supplement their diet with the algae found in the system. Herbivores, in some systems, can get enough to eat from the various micro- and macroalgae in the aquarium. In some cases though, such as with tangs, surgeons and sea urchins, they will quickly strip the tank of algae and will require additional feedings. These animals can be fed a variety of fresh greens such as green peas, spinach, romaine lettuce, bok choy and various marine algae that can be found in oriental food markets. I have found that blanching the greens for 30 seconds or so helps to soften them and fish that would not touch them before will devour the same greens when blanched. Additional feedings of vitamin enriched prepared foods are recommended too. Organisms that feed on smaller particles such as detritus and its associated components, as well as zooplankton, can either be fed or, in some cases, ignored entirely. It has been my experience that many forms of fanworm will exist quite well in aquariums, where the fish are fed regularly, without any additional feedings. If you find that the crowns of these fanworms decrease in size, then this may be a sign that they are not getting enough to eat and supplemental feeding would then be required. Liquid foods work well for the smaller species while live baby brine shrimp will be accepted by the larger species i.e. Sabellastarte magnifica. Other such feeders include Sea Cucumbers, clams (other than Tridacna spp.), tunicates and crinoids. Given that large amounts of detritus are produced in reef systems and that this detritus is often found as a fine suspension in the water due to the actions of burrowing worms in the rock, fish, bacterial action and water movement, most filter feeders receive more than enough to eat and do not need any additional feeding. Zooxanthellae bearing corals (hermatypic), both hard and soft, including mushroom anemones, zoanthids, anemones and gorgonians, can utilize a wide variety of feeding techniques. Not only can they utilize the photosynthetic products of their algal symbionts but they can also feed directly on plankton, bacteria, detritus and fish feces. Some corals have even been shown to be able to directly absorb glucose from the water (Stephens, 1962). Other zooxanthellae bearing organisms include some sponges and Tridacna spp. clams. As mentioned above, most corals contain symbiotic algae in their tissues which can supply some of their nutritive needs. I say some because the degree to which zooxanthellae contribute to a coral's nutrition has been the subject of much research over the past 40 years. It seems that the amount varies between species. In some species of zoanthids over 90% of their nutrition can be met by the zooxanthellae while in others this figure is much lower (60%) (Steen and Muscatine, 1984). However, the general consensus is that zooplankton do not contribute a major portion of the caloric or carbon requirements of hermatypic corals (Muscatine and Porter, 1977). The general rule seems to be the smaller the polyps, the more important zooxanthellae are in the diet (Porter, 1976). Corals feed in a variety of ways. The larger polyped forms (e.g. Euphyllia spp.) can actually feed on shrimp-sized prey which they capture with their tentacles. Other forms collect the slime that forms on the polyps and swallow the microorganisms and detritus trapped in it (Kuhlmann, 1985). Still others can directly absorb nutrients (ammonium, nitrate and phosphate used by the zooxanthellae as well as various amino acids) from the water (Franzisket, 1974; Muscatine and Porter, 1977; D'Elia, 1978; Muscatine and D'Elia, 1978). The problem comes when deciding when and how much to feed. In my own personal opinion many coral species do not need direct feeding. Many get more than enough from natural sources in the tank. Every time you feed your fish, particles of food and nutrients are added to the water. The presence of live rock and their associated algae and bacteria produce copious amounts of nutrients, vitamins and other products. Even in the presence of a highly efficient skimmer, Wilkens (1987) found that the levels of amino acids in the aquarium were many times higher than on the reef. It would be safe to assume that many other "nutrients" are just as abundant, despite our best efforts. Those polyps that are large enough to be fed small pieces of shrimp can be fed once a week or so by directly placing pieces of food on some of the polyps. Zooxanthellae require phosphate and although they may be able to absorb this from the water it is generally felt that the main source is from the prey captured by the polyps. However, some soft corals (e.g. Xenia) have never been observed feeding. Lacking stinging cells in their tentacles, Xenia may absorb phosphate and other nutrients directly from the water. Judging from the large number of successful aquariums that I have seen in which the corals are never directly fed, indicates to me that most zooxanthellae bearing corals do not require direct feeding to survive, grow and multiply. If you do decide to try and feed your corals be very careful about overfeeding i.e. feed SPARINGLY. An occasional feeding of live baby brine shrimp or one of the better liquid foods, may be appropriate for some specimens but not others. Pay careful attention when feeding, if it looks like the coral is not ingesting any food then perhaps it does not require additional feeding. For example, some species of mushroom anemones will feed if food is placed on their discs but others are never observed to feed. A general rule with mushroom anemones is that if they have large bumps or tentacle-shaped protrusions, you should try and feed them. Smooth surfaced forms, however, generally obtain enough nutrition from their zooxanthellae. However, Elliott and Cook (1989) showed that the Caribbean corallimorpharian Discosoma sanctithomae relied both on their zooxanthellae and on nocturnal morphological changes to capture prey. Even in those species that have short tentacles, the tentacles are non-retractile, non-motile, practically devoid of musculature and lack significant numbers of nematocysts (Den Hartog 1980). Those mushroom anemones (e.g. Elephant Ear Mushrooms, Rhodactis howesii sp.) that do feed, do so primarily by envelopment where the prey is trapped in the disc cavity and digested by mysenterial filaments extruded through the mouth. When one is dealing with corals that do not contain zooxanthellae (ahermatypic) then feeding takes on extreme importance. Examples of such organisms include certain gorgonian species, Dendronepthya sp. soft corals and Orange Cup Coral (Tubastrea aurea). In these cases live or prepared foods should be used. Live foods such as baby brine shrimp and rotifers are excellent for most gorgonians and Dendronepthya sp. while Tubastrea should be fed larger items such as live adult brine shrimp or small pieces of shrimp, scallop or fish. Prepared foods can be used as well. Dried or freeze-dried foods can be finely ground and soaked in a vitamin preparation. This sludge is then fed directly to the coral through a pipette or baster. You should not feed such food by simply placing it into the water. This only results in added pollution as most of it ends up in the filter, in the gravel or under rocks. The final method of feeding in reef tanks is through the direct uptake of organic compounds through the body walls of various marine worms, ascidians (i.e. Tunicates), bryozoans, etc. (Sepers, 1977). The mechanisms, importance and role of such feeding in marine ecosystems is not well understood and certainly bears more extensive research. Whether or not such feeding is an important component of our aquariums is certainly an unknown area but I feel that the majority of the nutrient needs of these organisms can be met by other means. Certainly, the addition of extra amounts of nutrients would be both unfounded and unnecessary given the lack of knowledge in this area. It would first have to ascertained exactly what substances were required by each organism and if these were depleted in an aquarium before any consideration could be given to additions. As I mentioned at the beginning of this article the topic of nutrition in aquariums is poorly understood at best. This is an area where the experiences of hobbyists can be of utmost importance to scientific researchers. There are a lot more hobbyists out there than there are people actively researching this area. It would be a shame if this tremendous pool of information and experience went unused. Share your information with others, write articles for club or national magazines, keep detailed notes on each of your specimens, spread your knowledge and experience, we will be all better off in the long run. The more we can demonstrate the value of our hobby, the less likely that it will be shut down.

References

Blank, R. and R.K. Trench 1985. Speciation in symbiotic dinoflagellates. Science 229:656-658. D'Elia, C.F. 1977. The uptake and release of dissolved phosphorus by reef corals. Limnol. Oceanogr. 22:301-315. Den Hartog, J.C. 1980. Caribbean shallow water corallimorpharia. Zool. Verhand. 176:1-83. Elliott, J. and C.B. Cook 1989. Diel variation in prey capture behaviour by the Corallimorpharian Discosoma sanctithomae: Mechanical and chemical activation of feeding. Biol. Bull. 176:218- 228. Franzisket, L. 1974. Nitrate uptake by reef corals. Int. Rev. Gesamten Hydrobiol. 59:1-7. Kuhlmann, D. 1985. Living Coral Reefs of the World. Arco Publ. Muscatine, L. and C.F. D'Elia 1978. The uptake, retention, and release of ammonium by reef corals. Limnol. Oceanogr. 23:725-734. Muscatine, L. and J.W. Porter 1977. Reef Corals: Mutualistic symbioses adapted to nutrient-poor environments. BioScience 27:454-460. Sepers, A.B.J. 1977. The utilization of dissolved organic compounds in aquatic environments. Hydrobiologia 52:39-54. Steen, R.G. and L. Muscatine 1984. Daily budgets of photosynthetically fixed carbon in symbiotic zoanthids. Biol. Bull. 167:477-487. Stephens, G.C. 1962. Uptake of organic material by aquatic invertebrates. I. Uptake of glucose by the solitary coral Fungia scutaria. Biol. Bull. 123:648-657. Trench, R.K. 1979. The cell biology of plant-animal symbiosis. Annu. Rev. Plant Physiol. 30:485-532. Wilkens, P. 1987. Niedere Tiere: Steinkorallen, Scheiben- und Krustenanemonen. Engelbert Pfriem Verlag, Wuppertal, Germany. Wotton, R.S. 1988. Dissolved organic material and trophic dynamics: What is the food of filtering collectors in aquatic ecosystems? BioScience 38: 172-178. The following references contain information on feeding marine invertebrates: Colin, P.L. 1978. Caribbean Reef Invertebrates and Plants. TFH Publ., Inc., Neptune City, NJ. de Graaf, F. 1973. The Marine Aquarium Reference. The Pet Library, LTD., Harrison, NJ. Freshwater and Marine Aquarium Magazine. R.C. Modeller Corp., Sierra Madre, CA. Haywood, M. and S. Wells 1989. The Manual of Marine Invertebates. Tetra Press. Kaplan, E.G. 1982. A Field Guide to Coral Reefs, Caribbean and Florida. Houghton Mifflin Co., Boston, MA. Marine Fish Monthly. Publ. Concepts Corp., Luttrell, TN. Marine Reef Newsletter, Aardvark Press, Bridgeport, CT. Moe, M.A. Jr. 1989. The Marine Aquarium Reference: Systems and Invertebrates. Green Turtle Publ., Plantation, FL. Teh, Y.F. 1974. Keeping live corals. Marine Aquarist 5(1):19-24. Wilkens, P. 1973. The Saltwater Aquarium for Tropical Marine Invertebates. 2nd. Extended Ed., Engelbert Pfriem, Wuppertal- Elberfeld, Germany. Wilkens, P. 1976a. Mini-Reef. Marine Aquarist 7(5):37-42. Wilkens, P. 1976b. Flower animals. Marine Aquarist 7(9):32-43. Wilkens, P. 1976c. More flower animals. Marine Aquarist 7(10):31-44. Wilkens, P. and J. Birkholz 1986. Niedere Tiere: Rohren-, Leder- und Hornkorallen. Engelbert Pfriem Verlag, Wuppertal, Germany. Wilkens, P. 1987. Niedere Tiere: Steinkorallen, Scheiben- und Krustenanemonen. Engelbert Pfriem Verlag, Wuppertal, Germany.

This is a rather old article and should be taken as such. Although much of what I wrote back then still applies to today's systems, you should not take the recommendations as gospel .