January | February 2012 Feature title: Redefining mineral requirements: Why its necessary?

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FEATURE for many higher organisms, but for fish only the commercially valuable species have received significant attention. Numerous studies have been carried out on rainbow trout, Atlantic salmon and Channel catfish; these species have well defined and frequently cited requirement levels. These requirement levels tend to be calculated using purified (non-realistic) diets and inorganic forms of the minerals. In reality, aquaculture diets contain anti-nutritional factors (ANFs); these are components of the feed that inhibit the uptake or utilization of another part of the feed. When concerned with mineral digestibility and availability two of the main ANFs are tricalcium phosphate and phytate (phytic acid). Tricalcium phosphate is found in the bone tissue of animals and phytate in many plant proteins. These ANFs bind to minerals such as zinc and effectively render them unavailable to the fish. An example of the effect of these ANFs can be seen in rainbow trout. Rainbow trout have a requirement of 15-30mg Zn/kg diet (Ogino and Yang, 1987). This was calculated using a purified egg albumin diet, with no ANFs, and using an inorganic zinc sulphate. When a practical diet containing fishmeal was used an additional 40mg Zn/kg diet (probably bringing the total dietary zinc level closer to 80-100mg Zn/kg diet) was required to maintain normal growth. Similarly, Atlantic salmon fed a fishmeal diet containing 65mg Zn/kg could not maintain their normal zinc status (Lorentzen and Maage, 1999). The higher the bone content of the fishmeal the more zinc needs to be added, especially when using an inorganic zinc salt. The replacement of fishmeal with plant protein may exacerbate this effect. Rainbow trout fed a soyabean meal based diet required 150mg Zn/ kg to achieve optimal growth. The increased use of sustainable fishmeals, often from trimmings high in bone content, and plant protein sources high in phytate may mean that a set of minimum requirement levels for fish fed more realistic diets will be of more practical use to the industry. The development of more advanced feed supplements such as proteinate sources of minerals may reduce the effect of ANFs on mineral availability. Mineral proteinates bind the mineral within their structure, protecting the mineral from the ANFs. This relationship between the protein and the mineral is complex. The mineral has to be bound tight enough not to be released in the gut where it would be a free mineral ion, susceptible to the ANFs, just like an inorganic salt, but the mineral still needs to be available to the animal once it has been taken into the cells. If the correct protein/mineral complex is used the level of the mineral used in the diet can be reduced by as much as 70 percent (Paripatananont and Lovell, 1995; channel catfish with zinc methionine). If research into the type of protein/mineral

FEATURE complex is carried out for each species the efficiency of mineral supplementation can be hugely improved. It would also reduce the problems (lower availability and excessive mineral excretion) associated with higher levels of mineral inclusion, which is required when using more sustainable animal and plant based protein sources.

he increasing development of aquafeed technologies embraces a new generation of feed ingredients and additives leading to changes in the specification of diet formulations. This necessitates a new understanding of mineral nutrition and the need to redefine trace element requirements in keeping with intensive production whilst promotion of fish health. This short review gives a basic outline of the biological mechanisms involved in one of these trace elements, zinc and describes why its important to re-evaluate the mineral requirements for salmonids.

Redefining mineral requirements: necessary? t Why is i

by Dan Leeming PhD Research Student, Aquaculture and Fish Nutrition Research Group, University of Plymouth, Uk

References:
Bury, NR, Walker, PA, Glover, CN, 2003. Nutritive metal uptake in teleost fish. J Exp Biol. 206, 11-23. Davies, SJ, Rider, S, Lundebye, A-K, 2010. Selenium and zinc nutrition of farmed fish: new perspective in feed formulation to optimise health and production. In: Bury, NR, and Handy, RD (Eds) surface chemistry, bioavalability and metal homeostasis aquatic organisms: An integrated approach. SEB, London, pp. 159-181. Lorentzen, M, Maage, A, 1999. Trace element status of juvenile Atlantic salmon Salmo salar L fed a fishmeal based diet with or without supplementation of zinc, iron, manganese and copper from first feeding. Aquac. Nutr. 5, 163-171. Ogino, C, Yang, G.Y, 1978. Mineral requirements in fish.4. Requirement of rainbow-trout for dietary zinc. Bulletin of the Japanese Society of Scientific Fisheries. 44, 1015-1018. Paripatananont, T, Lovell, RT, 1995. Responses of Channel Catfish Fed Organic and Inorganic Sources of Zinc to Edwardsiella ictaluri Challenge. Journal of Aquatic Animal Health. 7, 147-154.-

How does the fish obtain zinc?

How and where is zinc used?

Why is zinc important?

Micro-nutrients is a generic term for dietary components required in small quantities. Minerals such as copper, zinc, iron, manganese and selenium are all micronutrients although they are usually categorised as trace minerals, and are essential for the health of all animals, including fish. In aquaculture these dietary essentials are often supplemented as part of a vitamin/ mineral premix due to the inadequate supply obtained from many commercially used feed ingredients. Zinc is the most abundant trace mineral found in fish. It is essential for growth, the development and maintenance of healthy bones, and over 300 proteins require zinc as either a structural of functional co-factor. These include approximately 20 metalloenzymes such as alkaline phosphatase (required for bone mineralisation/formation), alcohol dehydrogenase (required for fructose metabolism) and carbonic anhydrase (required to aid the removal of CO2 from cell respiration). Fish deficient of zinc shows growth retardation, cataracts, fin and skin erosion, increased mortality rates and taste dysfunction resulting in reduced appetite and feed conversion.

Fish have two routes of zinc uptake, first from the diet and second from the surrounding water. There is the potential for waterborne zinc to be absorbed in both the gut, from the swallowed external water, and also directly from the external aqueous environment via the gills. Salmonids drink very little, especially when in freshwater; freshwater zinc levels are usually less than 10g/l and saltwater levels even lower. This is considered too low to make any significant contribution to the whole body zinc levels even though the gills affinity for zinc is extremely high. However, even with this high affinity the uptake rate of zinc from the gill is three to four times lower than from the gut (Bury et al, 2003). The uptake mechanism in fish is described as high affinity low capacity in the gills and low affinity but high capacity in the gut. This supports the theory that despite the high affinity for zinc in the gill, dietary uptake is the major contributor to the body zinc status. Free zinc ions (i.e. not bound to other compounds) are potentially very toxic to many biological processes, yet the incorporation of these free zinc ions in numerous proteins is vital for these very same biological processes to function. Thankfully, from a toxicological stance, only a very small fraction of the total zinc in the environment is in this free state. Unfortunately, from a nutritional stance, the majority of zinc in the environment is therefore unavailable. For the zinc to become available these compounds need processing in some way. This processing occurs when the compound is digested, freeing the potentially toxic zinc ion, which can now cross the intestinal barrier; or breaking the large compounds down into their smaller components, which can cross the intestine and take the zinc with it. Once inside the organism any free zinc is usually bound to another compound generally termed a chaperone, ready to be used or transferred around the body.

Zinc is very highly regulated in all aspects of the fishs body: its uptake from the water or the diet; its excretion by the gills, the intestine, the urine or the integument; and also by its distribution within the body. This regulation means that even a dietary level of 1700mg/kg ZnSO4 is still non-toxic to the fish. The ability to regulate this appears to come from the intestine, it is thought that excess zinc is simply not absorbed and passes through the fish in its faeces, however, it hasnt been proved that the high levels of zinc remaining in the faeces hasnt been processed by the liver and excreted back into the faeces in the bile. Either way, excess zinc in the diet does not seem to present a problem. Low dietary zinc levels are however more serious and the regulatory mechanism more complex. Every tissue of the fish can be broadly grouped into one of two categories; either functional or exchangeable. A functional zinc pool, such as the liver, fins, eyes, gills and skin are generally considered metabolically important. It is these tissues, which maintain their zinc concentration regardless of the dietary levels. Exchangeable pools seem to be less important metabolically but it is in these tissues (bone, muscle, intestine) we see fluctuations in zinc levels corresponding to the dietary levels. When the dietary supply exceeds requirement these tissues increase in zinc concentration and act as a storage facility and when the diet is deficient it is these pools that decrease quickly and allow the metabolically important tissues to maintain their zinc levels. Regardless of the ability of the fish to regulate zinc within its body the turnover of zinc is relatively fast (~1% per day). This means that in order to avoid deficiency a continual supply of dietary zinc is essential (Davies et al., 2010)

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28 | InternatIonal AquAFeed | January-February 2012

January-February 2012 | InternatIonal AquAFeed | 29

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