March | April 2012 Feature title: The Quest To Keep Salmon In The Pink Naturally

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FEATURE modulate various selected muscle quality parameters, including the degree of pigmentation in salmon was noted by Nordgarden et al. (2003). These investigators concluded that rapid period of growth achieved under continuous lighting compared to natural photoperiod resulted in elevated growth of salmon and increased oxidative stress with marked reductions in fillet vitamin E levels (a-tocopherol) and astaxanthin. achieve acceptable results. A minimum size threshold exists to initiate pigmentation in salmon and trout and maintain deposition during the initial freshwater stages of growth in fish of above 80-100 grams mean body weight. This extends with subsequent transfer as smolts to sea water with salmon typically fed 65mg/kg total carotenoids either as astaxanthin or in combination with canthaxanthin with the latter not exceeding 25mg/kg in the mixture (EU directives). Maximum permitted levels for astaxanthin in the USA is 80mg/kg of feed (FDA, 2010). However, revision of such levels is the basis of much scientific activity to optimise their use and minimise wastage and cost. Over the last decade, feed manufacturers have been able to significantly lower the inclusion of pigment in feed. Instead, it is recommended by some, that a three-phase pigmentation strategy with post-smolts being fed pigment at a relatively high level of 60-75mg/ kg with a transition to an intermediate level of 40-50mg/kg from a body weight of 2-3kg before a final regime maintained on a finishing diet prior to harvest of 25-35mg/kg of dietary carotenoids concentration (Sinnot, 2006). There are some scientific rationales for supporting higher diet pigment levels in the later stages of growth for the feeding of large adult fish. This is based on the view that larger salmon can pigment more efficiently than smaller fish. Since pigmentation develops quite rapidly from post- seawater transfer, pigmentation rate generally slows down when fish get bigger although the absorptive efficiency may actually increase. It is still therefore wise to promote a strategy based on using higher levels in early phase feeding with a reduction to a maintenance level in fish approaching harvest.

FEATURE Some evidence suggests reduced pigmentation efficiency may result in fish fed diets with elevated plant oils over extended periods during growth. However, consumer tests seem to indicate that there are no significant differences in public perception regarding the overall appearance of salmon colour when fed diet regimes that include appreciable levels of vegetable oil sources to replace fish oils (Rosenlund et al. 2003). This is of importance given the trend of using higher amounts of plant oil blends for a major part of the production and the strategy of using fish oil enhanced wash out diets in the final stage to harvest to achieve high omega 3 fatty acid concentrations in the flesh of salmon prior to market. Quinton et al (2005) conducted trials with salmon to ascertain the influence that genetics can play in affecting the absorption of dietary carotenoids, metabolism and the efficiency of flesh deposition. These workers in Canada evaluated the genetic parameters in different year classes of Atlantic salmon reaching harvest. They determined sexual maturation characteristics and associations to colour score, astaxanthin, canthaxanthin, oil, and moisture contents of flesh. Positive genetic correlations were found between body weight and with pigment retention levels. Obviously, there is the scope to exploit more efficient stocks for their pigmentation capacity using selective breeding programs. Such findings could result in more even and consistent flesh colour for salmon and trout and tailored to different production systems.

The quest to keep Salmon in the pink

naturally

Environmental factors
Other environmental factors that may affect the efficiency of pigmentation would likely include water quality and flow rates that would interact to modulate growth and nutrient retention. Stocking density and exercise are known to affect fish performance and optimum fish stocking densities will produce superior growth and feed conversion leading to improved fish quality. Similarly, adequate flow rates, water exchange rates will promote firmer fleshed fish and may enhance pigmentation in fish under intensive production systems as described previously in relation to muscle quality. Colouration and its relationship to flesh quality in farmed trout and salmon was extensively reviewed by Davies (2008) which addressed carotenoid function in fish as well as the physiological, biochemical aspects and aesthetic attributes leading towards optimal pigmentation of salmonid fish. Our knowledge of fish nutrition has expanded considerably in recent times and there have been numerous investigations that have addressed the effects of dietary levels of both astaxanthin, canthaxanthin separately or in combination on the flesh colouration of trout and salmon. The type of feed, level of feeding and matrix effects on carotenoid uptake will affect the degree of pigmentation in and can significantly alter the colour characteristics resulting in possible tainting with background colour. An optimum dietary level of about 65-mg/ kg astaxanthin is preferable in general to

by Simon J Davies, School of Biosciences, Plymouth University England, UK

he colour of farmed salmon has attracted so much attention within the aquaculture industry as it can be appreciated that the typical pink-reddish colour of salmon is symbolic of quality and value, with retailers demanding stringent criteria for farmed fish such as Atlantic salmon and trout to meet consumer expectations. Indeed this has often proved controversial in the media when the farming of fish has been criticised on many grounds with negative statements alluding to the use of synthetic agents and even dyes suggested as the source of artificial colours being prominently expressed by the misinformed. Carotenoids comprise a complex group of xanthophylls and carotenes, which are diverse in their chemical structure to provide the host of yellow to orange red, and pink colours found extensively in nature as also seen in the plumage of birds such as flamingos. The main carotenoids of importance to salmon and trout are astaxanthin and canthaxanthin, which are specific in their mode of activity and in their manner of metabolism. Astaxanthin is recognised to be the predominant red carotenoid found in salmonids in nature with canthaxanthin also used commercially to a more limited extent.

derivatives in the skin such as red sea bream and red tilapia (Gouveia et al, 2002). This is also a characteristic of high value and acceptability to consumers who perceive this to be superior compared to un-pigmented fish. The efficacy of flesh colouration by different carotenoids is a function of complex physiological processes followed by a series of biochemical events that involve metabolism primarily in the liver as well as the intestinal tract (Page et al 2005, Page and Davies, 2006). Evidence from the research investigations of White et al (2003a) suggests that a major fraction of absorbed astaxanthin is transformed into vitamin A in the intestinal tissue of rainbow trout and most likely salmon as well. In addition there is increasing evidence that carotenoids are involved in gene regulation and signal transduction thereby affecting the entire metabolism of fish (Azzi, 2007;

viewed as conditionally essential nutrients for salmonid fish species. Davies (2005) previously reviewed several aspects of biochemical and physiological parameters affecting salmonid pigmentation with particular emphasis on post-prandial absorption kinetics and retention efficiency and explained that considerable amounts of astaxanthin and canthaxanthin is excreted by fish and therefore efficiency of utilisation is quite low (~20 percent) compared to other nutrients such as proteins, amino acids, vitamins and minerals. There are many production related factors that can influence the degree of pigmentation of farmed salmonids. These include species, race or stock type, intra-population variation, age of fish, type and quantity of carotenoids ingested over specific period, seasonal effects, maturation, health and state of physiological stress, dietary formulation, regulated feed dep-

Less intensive conditions for farmed fish

Traditional use of commercially synthetic sources of astaxanthin in compounded feeds adds greatly to their costs and the value of the resulting products. The annual sales of synthetic astaxanthin for aquaculture alone were estimated at more than US$200 million at 2010.

The impact of plant oils on pigmentation

Choubert et al (2006) have reported that the pigmenting efficacy of astaxanthin fed to rainbow trout was affected by the composition of dietary oil present.

A dietary requirement
Salmon and trout do not possess the inherent metabolic ability to synthesise these pigments but instead require them within the diet as preformed molecules that are absorbed and subsequently deposited in tissues (Bjerkeng, 2000). It should also be noted that pigmentation leading to the reddish-pink and orange colour of integument is also favoured for certain fish that can deposit carotenoids or metabolic

Lordan et al 2008). They are also potent anti-oxidants that can interact with vitamin E (-tocopherol) and can prevent fatty acid oxidation due to free radical generation within tissues. There are also positive benefits on the post mortem quality of fish under frozen storage conditions when fed high levels of carotenoids thus prolonging shelf life of products (Jacobsen et al (2011). In this way astaxanthin and canthaxanthin have pro-vitamin and metabolic functions well beyond their standard role for pigmentation and could therefore be
16 | InternatIonal AquAFeed | March-april 2012

rivation periods maintenance feeding, slaughter conditions, visualisation of colour, processing conditions: storage, smoking, cooking etc. The high growth rates achieved in modern salmon production can result in a reduction of pigmentation and uneven distribution within the flesh. It is well known that considerable variations can occur in the ability of fish to retain carotenoids within the muscle and this can sometimes manifest as very low pigmentation or indeed excessive levels in different regions of muscle. Indeed the fact that seasonal changes can

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March-april 2012 | InternatIonal AquAFeed | 17

FEATURE Recently, however, there has been a growing demand in the production of farmed fish under less intensive conditions, and with more emphasis on natural additives and supplements in the diet. As such, a number of investigations have reported the feasibility of various single cell products such as the red yeast Phaffia rhodozyma and Haematococcus pluvialis algae with respect to their pigmentation ability compared to the synthetic form currently available to the industry (Lagocki, 2001). Investigations have confirmed that Phaffia can effectively pigment salmonid fish but only few products are currently available for this purpose. Choubert et al (2006) found differences with respect to the colour of rainbow trout fed Haematococcus pluvialis compared to synthetic astaxanthin with higher reported colour and flesh retention. The prevailing consensus has been that the synthetic commercial astaxanthin is readily available for assimilation by fish, and is a more stable and consistent product. It should also be noted that the astaxanthin present in H pluvialis is found as complex esters which confer different assimilation properties compared to other sources adding to variable results under practical conditions (Bowen et al, 2002; White et al, 2003b), there is current interest in sea weeds and extracts from macro-algae which can contain appreciable levels of carotenoids as well as waste products from krill and shrimp processing. However, these materials are inconsistent in availability and carotenoid levels may vary with seasonality. However, speciality seaweeds may have a promising future as feed additives combining functionality as prebiotics and contributing to natural carotenoid intake. mentation to fish that deposit these in muscle and skin. The red carotenoidrich bacterium Paracoccus carotinifaciens is permitted as a sensitive additive for use in salmon and trout with a maximum content of 100mg, expressed as the sum of astaxanthin, adonirubin and canthaxanthin per kg complete feed. The current carotenoid composition of the product is specified as 35g canthaxanthin, 1015g adonirubin and 2023g astaxanthin/kg. The applicant proposes to modify the ranges of canthaxanthin to 15g, and that of adonirubin to 715g/kg product, while maintaining the astaxanthin range. The European Food Safety Authority (EFSA) of the European Commission has expertly reviewed the terms of authorisation of the product for fish and verified its efficacy and safety. Indeed a number of trials in Scotland by leading feed manufacturers and salmon producers have proved most encouraging with favourable results obtained in production cages. Flesh pigment deposition and stability comparable to that of synthetic pigmentation regimes proved acceptable and quite comparable with orthodox products. The feedback received so far on fish pigmented with Panaferd -AX has been consistently good from the retailer and consumer standpoint. "To have all the colour in our salmon derived from naturally occurring organisms is the culmination of years of hard work," said managing director Nick Joy of Loch Duart salmon in Scotland who have been pioneering the product in their bespoken feeds. "As a farmer, I am proud that we rear well-nourished salmon that look and taste great," he has stated.

fleshed salmon in the same manner as a wild salmon accumulates pigment is a widely accepted principle advocated by a number of market surveys in which the pink flesh colour of salmon scores consistently higher as a desirable factor second only to fish freshness (Baker, and Gnther, 2004) In terms of feed costs it is well known that the added costs of including astaxanthin in feeds amounts to an extra 10-15 percent, which manifests as an additional 4-6 pence per kilogram produced, that is a cost of UK40,000-UK60,000 per 1000 tonnes of production. In Europe, it is common practice to pigment rainbow trout to a defined level at portion size although this is not so usual in the United States where un-pigmented (white) fleshed rainbow trout is more favoured by the consumer. Several feed companies have evaluated the costs of pigmenting rainbow trout towards harvest with strategies for more economic approaches advocated by several feed companies and fish farming operations.

Colouration is an emotive issue

The question of fish colouration in farmed aquatic species will always be an emotive issue since it is easily visualised and can be measured by suitable techniques ranging form direct chemical analysis in feeds and flesh or by a selection of optical methods based on colour scale assessments and advanced image analysis. The aquafeed industry and fish nutritionist must be responsive and embrace the opportunities for innovative products including natural pigmenting agents. With the need to consider organic certification for farmed fish and the quest for sustainable ingredients, research is needed to evaluate the efficacy of carotenoids from a wide spectrum of potential sources. This will be a continuous challenge as feed formulations become refined and as new candidate species become available for aquaculture.
References available on request

Algal and yeast sources

Despite the potential of both algal and yeast sources of carotenoids being able to effectively pigment salmon and trout, these have been prohibitive in terms of their considerable costs and variable qualities compared to synthetic products with consistent characteristics. Consequently, an exciting development is the product Panaferd-AX produced by a leading Japanese company Nippon Oil Corporation. Panaferd-AX contains the dessicated cells of Paracoccus carotinifaciens, a soil-inhabiting bacterium which naturally contains carotenoids. The bacteria are cultured by fermentation and have been selected to yield high carotenoid concentrations without the need for genetic modification. Although astaxanthin is the major pigment within Panaferd-AX, it can also express appreciable levels of natural adonirubin and canthaxanthin. Both of these carotenoids can also support additional pig-

Consumers and retailers are driving the agenda

It seems that Panaferd will benefit salmon producers globally by offering opportunities for diversification in the marketplace. Since the product has full FDA and EU approval for use and is available for application in the industry its likely to be used increasingly in main- stream production. It is the requirements of the consumer and retailers that are driving the agenda and generating the need for more information in this area with media attention to colouration of farmed fish and crustacean species. The question of producing a pink/ red
18 | InternatIonal AquAFeed | March-april 2012

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