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DOI: 10.4172/2155-9546.1000234
Research & Development
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ISSN: 2155-9546

Research Article Open

OpenAccess
Access

Fish Nutrition and Current Issues in Aquaculture: The Balance in

Providing Safe and Nutritious Seafood, in an Environmentally Sustainable
Manner
Stefanie M Hixson*
Department of Ocean Sciences, Memorial University of Newfoundland, St. John’s, Newfoundland and Labrador, Canada

Abstract
Global aquaculture production has increased in recent years and it is predicted that aquaculture will provide the
most reliable supply of seafood in the future. However, there are many controversial issues in aquaculture regarding
food safety, nutrition, and sustainability; many of which are directly related to the nutrition and feeds for farmed fish.
These nutrition-related issues must be considered in order to achieve balance in safe and nutritious food production
and sustainability in aquaculture. This review highlights recent studies and discusses new and innovative aspects
in fish nutrition. Some issues in the area of fish nutrition require consideration and improvement, such as: feed
and nutrient efficiency, overfeeding and waste, fish meal and fish oil replacements, fish health, biotechnology, and
human health concerns. The findings reviewed in this manuscript demonstrate promise toward improvement of the
aquaculture industry through nutrition. This review is an update in fish nutrition research, and provides insight on
the progression and evolution of this field in order to meet the needs of the industry with the purpose to achieve a
balance in seafood production and environmental sustainability. The outcome of this review encourages the use of
biotechnology as a tool to meet seafood production and environmental sustainability, in order to ensure global food
security in the future and to improve our resource use.

the minimum nutrient requirement to promote growth and prevent

Keywords: Aquaculture; Biotechnology; Feeds; Health; Nutrition; signs of nutrient deficiency [1]. Protein is required in the diet to
Sustainability obtain amino acids, which are utilized to synthesize new proteins or
maintain existing proteins in tissues while excess protein is converted
Introduction to energy. Lipids supply essential fatty acids and energy in the diet. The
Aquaculture requires optimisation of nutrition to efficiently raise requirement of essential fatty acids can only be met by supplying Long
fish for the purpose of food production. Fish nutrition is the study Chain (LC) Polyunsaturated Fatty Acids (PUFA) in the diet, specifically
of nutrients and energy sources essential for fish health, growth and α-Linolenic Acid (LNA, 18:3ω3) and Linoleic Acid (LA, 18:2ω6), with
reproduction. Global consumption of seafood is increasing, while varying requirements for eicosapentaenoic acid (EPA, 20:5ω3) and
the amount of captured fish is declining, therefore it is predicted that docosahexaenoic acid (DHA, 22:6ω3) depending on species. Dietary
aquaculture will provide the most reliable supply of seafood in the lipids are also important structural components of membranes, and act
coming years. With the world’s rapidly expanding population, it is as precursors of steroid hormones and prostaglandins in fish. Dietary
important to provide safe and nutritious fish; however there are many carbohydrates can be a source of energy for fish; however their ability to
issues related to fish nutrition that need to be considered in order to utilize dietary carbohydrate for energy varies depending on the species
achieve balance in food production and sustainability. Sustainability and their natural diet. Therefore, depending on species, protein and
of the aquaculture industry is an environmental, economic and social lipid are the main source of energy for fish. Feeds in aquaculture are
concern; however, this review focused specifically on environmental formulated with a balance of nutrients in order to meet specific nutrient
sustainability in regards to fish nutrition. Certain issues related to requirements for different species, life stages and other purposes.
fish nutrition have become controversial because they impact the
environment and/or affect the final product for consumption. Some The digestibility of nutrients in the feed can affect aquaculture
of these issues include: feed and nutrient efficiency, overfeeding and production efficiency and impact the environment. The bioavailability
waste, unsustainable feed ingredients, fish health issues, biotechnology or digestibility of the diet is the proportion of nutrients in the feed
and human health concerns. Ultimately, each of these issues can that is digested and absorbed by the fish. Data on the digestibility and
affect the final product for human consumption, either nutritionally, available digestible energy of feed ingredients in fish diets are essential
environmentally or economically. Achieving a balance between efficient
and safe food production with environmental sustainability will be a
challenge for the industry. The following is a review of several issues in *Corresponding author: Stefanie M Hixson, Department of Ocean Sciences,
farmed fish nutrition, relating to providing quality food products while Memorial University of Newfoundland, Marine Lab Road, St. John’s, Newfoundland,
maintaining environmental sustainability, emphasizing limitations of Canada A1C 5S7, Tel: +1 709 864 2521; E-mail: [email protected]
this balance and strategies for improvement. Received March 24, 2014; Accepted May 05, 2014; Published May 15, 2014

Fish nutrition and nutrient efficiency in aquaculture Citation: Hixson SM (2014) Fish Nutrition and Current Issues in Aquaculture:
The Balance in Providing Safe and Nutritious Seafood, in an Environmentally
The essential nutrients for fish are amino acids, fatty acids, Sustainable Manner. J Aquac Res Development 5: 234 doi:10.4172/2155-
vitamins, minerals and energy-yielding macronutrients (protein, lipid 9546.1000234
and carbohydrate). Diets for fish must supply all essential nutrients and Copyright: © 2014 Hixson SM. This is an open-access article distributed under
energy required to meet the physiological needs of growing animals. the terms of the Creative Commons Attribution License, which permits unrestricted
use, distribution, and reproduction in any medium, provided the original author and
Guidelines for nutrient adequacy for some farmed fish species suggest source are credited.

J Aquac Res Development

ISSN: 2155-9546 JARD, an open access journal Volume 5 • Issue 3 • 1000234
Citation: Hixson SM (2014) Fish Nutrition and Current Issues in Aquaculture: The Balance in Providing Safe and Nutritious Seafood, in an
Environmentally Sustainable Manner. J Aquac Res Development 5: 234 doi:10.4172/2155-9546.1000234

Page 2 of 10

for optimization of feed formulations [1]. Feeds that are poorly digested consequent feed wastage, a technology that has been used in recent
result in limited growth and feces with high nutrient content, which years by several commercial farms [2]. Nutritional strategies to reduce
pollutes the environment. Therefore, the digestibility of nutrients and waste include improvements in feed formulations without affecting
the potential for nutrient retention and waste must be considered for growth and production efficiency, inclusion of feed ingredients with
efficient and sustainable animal production when considering feed high phosphorus bioavailability, use of feed additives to improve
formulations. the apparent digestibility of phosphorus, and processing-refining of
ingredients [9]. These efforts have resulted in a significant reduction
Growing fish accrete new tissues and some of the energy supplied in
of waste outputs (per unit of fish produced) by fish culture operations
the diet is stored as protein, lipid and some glycogen. Protein deposition
in Canada over the past four decades [6]. However, feeds that are fully
depends on the balance of available amino acids in protein and the
digested by the fish cannot totally resolve the impact of fecal waste
digestible protein-to-digestible energy ratio. Excess energy intake
because the scope of digestion in fish is limited and there will always
and low protein levels result in the deposition of lipid as recovered
be a fraction of undigested feed [10]. Removing the solid waste before
energy, which does not equate to faster growth and is an inefficient
it is discharged can be a solution for reducing the environmental
use of nutrients. Fish have the ability to utilize lipids for energy, saving
impact of wastewater [9]. Eco-certification may be a tool to set
protein for deposition and growth [2]; therefore inclusion of lipids in
standards with criteria aimed at reducing eutrophication through
diets for fish is important for both growth and energy purposes. Most
the level of inclusion; e.g., specific allowed amounts of nitrogen and
commercial feeds today are formulated to increase growth performance
phosphorus release from the system and a set limit for solids in the
by exploiting the protein-sparing effect of high energy lipid, allowing
effluent water [11]. Integrated multi-trophic aquaculture (IMTA)
as much of the dietary protein as possible to be converted into muscle
of fish in combination with invertebrates and plants can help reduce
protein. As a result, the production efficiency of farmed salmon has
environmental impacts and maximize the usage of food input. IMTA
significantly improved over time [3]. Today, the use of more highly
shows the most promise in terms of new and innovative systems for
digestible nutrient-density extruded feeds (46 to 50% protein, 20 to
waste mitigation and production efficiency. The IMTA system reduces
24% fat) allows commercial farmers to achieve a Feed Conversion
organic waste by mimicking trophic relationships found in nature; the
Ratio (FCR) of about 0.9-1.2 for rainbow trout (Oncorhynchus mykiss)
waste from one organism is food for another, resulting in decreased
grown to market size [2]. The improved FCR over time has been due
organic particle concentrations with increased distance from farm sites
to increased digestible nutrient and energy content of the feeds, as well
as they are consumed by other farmed organisms [12]. The carbon,
as feed extrusion which has resulted in the production of feeds with
nitrogen, and phosphorus compositions of feed, fish and faeces were
higher lipid levels, improved starch gelatinization (increases digestible
studied at an Atlantic salmon (Salmo salar) farm to estimate the release
energy content and utilization), and improved pellet characteristics
rates of wastes from salmon cages and the qualities of particulate wastes
(durability, buoyancy, etc.) [2,3]. Further research to improve
as food resources for integrated multi-trophic aquaculture. The study
feed ingredients and feed efficiency in the future will help improve
found that both salmon feed and faeces were adequate food for blue
sustainability in aquaculture. For example, genetically modified (GM)
mussels (Mytilus edulis) and sea cucumbers (species unknown), and the
salmon have better FCR than non-GM salmon (see Biotechnology
nutrient content may meet their nutritional requirements, including
section).
DHA and EPA contents of feces, which were comparable to those of
Feeding and the environment some phytoplankton species [13]. Other studies have also reported
successful incorporation of nutrients from salmon waste feed into
Feed is the main source of waste and is responsible for most of the the tissue of blue mussels [14,15]. Choice of the extractive species and
environmental impact of aquaculture [4] feed composition and the distance from the feeding location is an important consideration. For
FCR affect the amount of waste produced, as well as its physical and example, in a study on a commercial salmon farm in British Columbia,
chemical composition. Excess feed results in immediate eutrophication Canada, mussels had significantly higher amounts of DHA compared
of the surrounding environment; while consumed feed is yields with other molluscs (chitons, clam, limpets, periwinkles and whelks),
products of metabolic processes, such as ammonia, phosphorus and which indicates their potential in IMTA. The levels of DHA in mussels
carbon dioxide [5]. The quantity and quality of the waste excreted by showed a significant breakpoint at 339 m from the farm, which suggests
fish depend on intake, digestion and metabolism of dietary compounds that distance should be considered to optimize certain nutrients [16].
[6]. Excess feed waste has a much greater capacity than fecal material However, organic fish waste captured by mussels is limited by the time
to impact the environment, in terms of energy content and degradation available to intercept solid wastes contained in the horizontal particle
rate. The particulate organic matter sinks and disperses, which results flux, the velocity of the current, available IMTA farm space, and any
in environmental toxicity and anoxia [7]. The degree of impact from negative feedback effects on fish culture from flow reduction caused by
effluent wastes depends on feed quality, digestion and metabolism of mussel culture [17]. New research is focused on expanding novel fish
the diet, species, culture method and the nature of the surrounding and extractive species. A study recently demonstrated that green sea
environment in terms of physics, chemistry and biology [8]. This has a urchins (Strongylocentrotus droebachiensis) actively ingest and absorb
direct impact on marine benthic habitats, with effects such as reducing organic material from the waste produced by sablefish (Anoplopoma
sediments, hypoxia in the water overlying the sediment, increased fimbria) culture. Further research was recommended to determine
sulphate reduction and changes in benthic fauna assemblages in terms the effect of the sablefish waste diet on green sea urchin survivorship,
of species number, diversity, abundance and biomass [7]. growth, and gonad quality for urchin production [18]. Multi-species
Feeding methods and technologies have advanced in recent years production in an IMTA system must be optimized and better utilized
to minimize and eliminate waste, but the issue remains. Improved in the future to increase productivity and improve sustainability.
digestibility, feed utilization and feeding practices should implemented
Sustainable feeds and ingredients
by the feed and production industry to reduce losses from waste. One
example is the use of devices to monitor feeding activity from below According to the Food and Agriculture Organization of the United
the surface in order to feed to satiation without overfeeding and Nations (FAO), at least half of the world’s recognized fish stocks are

J Aquac Res Development

ISSN: 2155-9546 JARD, an open access journal Volume 5 • Issue 3 • 1000234
Citation: Hixson SM (2014) Fish Nutrition and Current Issues in Aquaculture: The Balance in Providing Safe and Nutritious Seafood, in an
Environmentally Sustainable Manner. J Aquac Res Development 5: 234 doi:10.4172/2155-9546.1000234

Page 3 of 10

fully exploited and 32% are overexploited or depleted [19]. With the the fatty acid profile of terrestrial plant oils is not equivalent to that of
production of farmed fish surpassing that of wild fish, aquaculture’s FO, so research in this area has been extensive in recent years.
share of global Fish Meal (FM) and Fish Oil (FO) consumption has
more than doubled over the past decade [20]. For the past 25 years, Terrestrial plant oils in aquaculture feeds
annual FM and FO production has not increased beyond 1.5 million Terrestrial plant oils will likely be the main choice when replacing
tons per year; therefore the aquaculture industry cannot continue to FO in aquaculture diets; however they are fairly limited in their ability
rely on finite stocks of marine pelagic fish as a supply of FM and FO to fully replace FO in diets for fish. Most plant oils are relatively poor
[21]. The paradox of FM and FO use in aquaculture has sparked major sources of ω3 PUFA in comparison to marine FO, and completely
research into alternative feed ingredients to replace FM and FO over lack LC ω3 PUFA. Rather, they are rich sources of ω6 and ω9 fatty
the past two decades. The focus of this research has mainly been on acids, mainly LA and 18:1ω9, with the exception of some oilseeds.
terrestrial plant meals, concentrates and oils; and many of these studies Although considered an excellent energy source, feeding terrestrial
have led to an impressive reduction in the average inclusion of FM and plant oils inevitably results in lower levels of DHA and EPA in tissues
FO in commercial feeds [22]. The FAO predicts reductions in FM over of fish fed plant oils [21,28-32], which is detrimental to fish health and
the next decade for different fish diets: from 26% to 12 % for marine compromises the health benefits for humans that consume these fish.
fish, from 22% to 12% for salmon and from 3% to 1% for tilapia [3]. The Extensive replacement of FO with terrestrial plant oils, particularly
FAO also predicts that the sustainability of the aquaculture industry those high in LA, cause a high incidence of cardiovascular disorders in
will likely depend on using terrestrial plant oils for aquaculture feeds fish [33] and also has been suggested to be detrimental to human health
rather than solely depending on FO as a lipid source [19]. Other FM after consumption of fish fed soybean oil [34], which is discussed in
and FO alternatives include meat and bone meal, livestock tallow or fish nutrition and human health section in this review. As a result, the
fat, blood meal and poultry by-product meals; as well as fishery by- immediate thought is that plant oils best suited as a substitute for FO
products and zooplankton meals and oils [23]. Considering the use of should contain high levels of ω3 PUFA (LNA) and lower amounts of
alternative protein and lipid sources in commercial feeds [24], recent LA, in order to increase the ω3/ω6 ratio. However, a study by Francis
calculations using the Marine Nutrient Dependency Ratio (MNDR) et al. [35] contradicts this idea because it was found that ω6 PUFA
estimate marine nutrient input to marine nutrient output ratios that (sunflower oil diet) appeared to ‘spare’ the catabolism of ω3 LC PUFA
are less than one; indicating that farmed salmon can be net producers and, as such, resulted in the highest retention of these fatty acids by
of FM and FO [24]; and actually increases the supply of fish for human rainbow trout. These results suggest new nutritional approaches to
consumption by 7-8 million tonnes per year [25]. In fact, globally fed maximise the maintenance of the qualitative benefits of fish oils when
aquaculture is producing more than three times as much fish as it they are used in feeds for aquaculture species [35]. Other recent studies
uses in the feed; therefore reducing the amount of FM and FO in the have described that diets that contain high levels of LNA are relatively
feeds actually increases the effective global supply of fish for human wasteful, because the fish did not extensively utilize the LNA toward
consumption. However, it is a challenge to find sustainable feed ω3 PUFA biosynthesis, but rather catabolized the ω3 PUFA for energy.
ingredients that meet the nutritional requirements of many farmed fish A similar observation is true for some fatty acid classes, particularly
species. It is likely that some production of FM and FO will remain saturated fatty acids and monounsaturated fatty acids, which also
necessary due to nutritional and industry growth constraints. appear to enhance the retention of ω3 LC PUFA in the fillets of some
Terrestrial plant meals and oils will probably continue to be the fish species. These observations have been found in Atlantic cod (Gadus
main choice when replacing FM and FO in aquaculture diets; however, morhua) [31], sunshine bass (Morone chrysops x Morone saxatilis) [36],
they are fairly limited in their ability to fully replace FM and FO in Murray cod (Maccullochella peelii peelii) [37] and rainbow trout [35].
diets for fish in terms of meeting nutritional requirements. As a protein This approach has been given the more specific definition of the ‘ω3 LC
source, plant meals are often deficient in certain essential amino acids, PUFA sparing effect’ as found in Atlantic salmon [36]. Consequently, it
and contain high amounts of fibre, carbohydrate, and certain anti- could be argued that plant oils high in LNA do not have any nutritional
nutritional factors, which cause adverse effects on feed intake, digestion, advantage over other commercially available oils with lower levels of
absorption, leading to reduced growth [26]. A meta-analysis by Collins ω3 PUFA, provided that DHA and EPA are spared through catabolism
et al. [27] found differences in the effect of plant ingredients on the of other abundant fatty acids. These results also indicate that feeding
growth performance of salmonid fish and that increasing the inclusion a diet based on FO only is an inefficient practice anyway, because
level of pea meal, pea protein concentrate, soy protein concentrate, substantial amounts of the nutritionally valuable ω3 LC PUFA are
canola/rapeseed meal or canola/rapeseed concentrate decreased the oxidized for energy, particularly EPA [38]. Therefore, the use of plant
specific growth rate. The reduced growth performance can likely be oils in fish diets conserves LC ω3 PUFA in FO for critical physiological
explained by reduced protein, amino acid, lipid and energy digestibility, functions only. This has been the focus of studies in this area recently,
if not compensated by increased feed intake [27]. Some improvements specifically on the dynamics of dietary DHA/EPA/ARA and its effect
in the nutritional quality can be achieved by chemical and mechanical on fish performance, tissue concentration and immunity (Table 1).
processing, which can eliminate or reduce the concentration of certain The ability of fish to synthesize LC PUFA may allow for plant oils
anti-nutrients, as well as concentrate protein content [22]. In fact, in 25 to fully replace FO in aquaculture feeds without lowering levels of key
years of research, fish growth performance in response to plant protein fatty acids in the flesh such as DHA and EPA that are significant for
ingredients has improved due to changes in the chemical and physical fish and human health. The saturated fatty acids 16:0 and 18:0 can be
properties of the ingredients due to advances in plant breeding, such biosynthesized by all known organisms, including fish. Desaturases and
as reducing the glucosinolate content in crops like canola [27]. Several elongases are the critical enzymes in the pathways for the biosynthesis
studies have shown that replacing FO with various terrestrial plant oils of the LC PUFA from the shorter-chain fatty acids to longer, more
in diets for different fish species does not compromise health, growth unsaturated chains. Fish can desaturate 16:0 and 18:0 to yield 16:1ω7
or feed consumption and are that they readily catabolised by fish as an and 18:1ω9 by Δ9 desaturase. However, all vertebrates lack Δ12 and
energy source for growth, as reviewed by Turchini et al. [21]. However Δ15 desaturases, which are necessary to form LNA and LA, so these

J Aquac Res Development

ISSN: 2155-9546 JARD, an open access journal Volume 5 • Issue 3 • 1000234
Citation: Hixson SM (2014) Fish Nutrition and Current Issues in Aquaculture: The Balance in Providing Safe and Nutritious Seafood, in an
Environmentally Sustainable Manner. J Aquac Res Development 5: 234 doi:10.4172/2155-9546.1000234

Page 4 of 10

Lipid source Study duration (d) Diet total lipid (%) ω3/ω6 EPA/ARA DHA/EPA SGR1 Reference
Tuna oil/rapeseed oil 75 23.5 1.3 5 2.8 1 Codabaccus et al. [38]
Fish oil/chicken fat 75 23.5 2 16 0.5 1 Codabaccus et al. [38]
Linseed oil 112 20 3.6 4.9 1.8 2 Francis et al. [35]
Olive oil 112 20 0.8 5.2 2.1 2 Francis et al. [35]
Palm oil 112 20 0.8 5 2 2 Francis et al. [35]
Sunflower oil 112 20 0.2 5 2.2 2 Francis et al. [35]
Fish oil/vegetable oil+EPA 112 18 2.5 18 0.7 - Martinez-Rubio et al. [49]
Fish oil+EPA 112 18 3.6 18 0.9 - Martinez-Rubio et al. [49]
Fish oil/rapeseed oil: high fat 84 34 1.4 22 1.2 1.0 Martinez-Rubio et al. [50]
Fish oil/rapeseed oil: low fat 84 20 1.5 26 1.3 0.9 Martinez-Rubio et al. [50]
Camelina oil 112 19 1.2 15 1 0.9 Hixson et al. [32]
Olive oil+DHA 62 20 1.3 4 19 1.2 Glencross et al. [70]
Olive+DHA+ARA 62 18 0.5 0.1 6.8 1.2 Glencross et al. [70]
Rapeseed oil 109 27 0.8 16 1.7 0.9 Thomassen et al. [57]
Rapeseed oil+EPA 109 28 1.9 158 0.2 0.9 Thomassen et al. [57]
Rapeseed oil+EPA+DHA 109 28 1.9 114 0.6 0.9 Thomassen et al. 57]
Rapeseed oil 95 26 0.65 18 1.3 1 Hatlen et al. [81]
Rapeseed oil+GM yeast (high EPA source) 95 23 0.8 33 0.3 0.9 Hatlen et al. [81]
Soybean oil+GM yeast (high EPA source) 112 31 0.4 91 z0.1 0.66 Berge et al. [45]
1
Specific growth rate, % day -1

Table 1: Replacement of fish oil with terrestrial plant oils in diets for Atlantic salmon (Salmo salar), with particular focus on dietary DHA/EPA/ARA dynamics and the effect
on tissue fatty acid composition and/or immune function.

fatty acids are considered essential. Subsequently, LNA and LA can can be quantified using the fatty acid mass balance equation. The fatty
be desaturated and elongated to form the physiologically essential acid mass balance method was developed for fish by Turchini et al.
EPA, DHA and ARA. However, the degree to which an animal can [40] to quantify ω3 fatty acid synthesis in fish to determine the level of
synthesize these fatty acids from LNA and LA depends on the activities elongation and desaturation that occurred over the course of a feeding
of the elongase and desaturase enzymes (Δ6 and Δ5) in their tissues. experiment. The method involves computation of the fatty acid intake,
Fish species differ in the extent to which they can tolerate diets without accumulation, and appearance or disappearance of the selected fatty
FO, and this trait appears to be evolutionarily related to the fatty acid acids in the ω3 pathway and computes the percentage of synthesized
profile of the natural diet. Consequently, carnivorous marine fish have ω3 LC PUFA from dietary intake of LNA. An application of this
lost much of the capacity to synthesize these fatty acids during evolution method revealed that Atlantic cod synthesized 6% of their own LC ω3
since they remained in an environment where such a conversion is not PUFA [31], and 12% for rainbow trout [41] after fed a plant oil-based
necessary. Freshwater fish have a greater ability to biosynthesize EPA diet. Another interesting method to quantify LC PUFA biosynthesis is
and DHA from LNA, since the natural prey of many freshwater fish the use of compound specific stable isotope analysis. Fatty acid isotopic
is not rich in EPA and DHA, but rather LNA and LA. The fatty acid signatures are frequently used in food web studies to determine the
desaturation and elongation pathway has been extensively studied in transfer of fatty acids from prey to predator based on their 13C/12C
fish at both the molecular and enzymatic level, with fatty acyl elongase ratio [42]. Fatty acids from terrestrial plant oils have distinctly different
(ELOVL) and fatty acyl desaturase (FAD) identified and functionally isotopic signatures than the same fatty acids in marine sources like fish
characterized in several marine and freshwater species. The dietary oil due to differences in the source of carbon (terrestrial carbon in the
fatty acid profile is influential to the expression of ELOVL and FAD form of CO2 gas vs. marine carbon as carbonate) [43]. Using CSIA and a
genes. The tissue fatty acid profile has been found to be significantly mixing model calculation [42], the proportion of synthesized LC PUFA
correlated with FAD and ELOVL gene expression in Atlantic cod fed (i.e., DHA) from LNA can be determined. Using this method it has
diets without FO; while FAD and ELOVL expression were significantly been found that rainbow trout can synthesize up to 27% of the DHA
correlated with each other [38]. This is evidence that the regulation of in the muscle tissue from dietary camelina oil [31]. Quantifying fatty
these genes is signalled by a change in the fatty acid profile of the tissue. acid biosynthesis in fish using quick and efficient methods will become
However, up-regulation of these genes is not necessarily reflected even more important in subsequent years; particularly if breeding
phenotypically because levels of DHA and EPA in fish fed plant oil programs are designed to select fish that have superior ELOVL and
diets are significantly lower than DHA and EPA levels in fish fed a FO FAD expression and express these traits phenotypically.
diet. However, these PUFA could be present in even lower amounts if
Future research in this area will be dedicated to exploring new
ELOVL and FAD were not facilitating fatty acid biosynthesis. Often
plant resources that have high levels of ω3 fatty acids, particularly plant
this effort cannot fully compensate for low levels of ω3 LC PUFA intake
sources that contain substantial levels of EPA and DHA. Single-celled
from plant oil diets; although results show that it is this low dietary level
microalgae and yeast can produce their own EPA and DHA and are
that may trigger the up-regulation of genes involved in their synthesis.
renewable resources [44,45] ; however, high production costs make
Using molecular tools to identify ELOVL and FAD gene expression commercialization very limiting. Research in this area is needed to
has often been the centre of research studies on FO replacements. optimize time and cost of production. However, the most promising
However, there are other tools that can be used to quantify LC PUFA and upcoming FO replacements are genetically modified LC ω3
biosynthesis and can verify phenotypically the results found at the PUFA enriched crop production [46,47] and is further discussed in
gene expression level. Mathematically, the level of LC PUFA synthesis the Biotechnology section of this review. The use of plant ingredients

J Aquac Res Development

ISSN: 2155-9546 JARD, an open access journal Volume 5 • Issue 3 • 1000234
Citation: Hixson SM (2014) Fish Nutrition and Current Issues in Aquaculture: The Balance in Providing Safe and Nutritious Seafood, in an
Environmentally Sustainable Manner. J Aquac Res Development 5: 234 doi:10.4172/2155-9546.1000234

Page 5 of 10

in aquaculture inevitable; however it also raises questions regarding clinical symptoms and improving performance, notably the ω3/ω6
sustainability of crop production for aquaculture feeds. Measurements PUFA and EPA/ARA levels. Clinical nutrition through functional
to quantify the amounts of land, water, nutrients and energy required feeding has been shown to be an effective complementary therapy for
for crops per unit of fish production should be calculated to assess emerging salmon viral diseases associated with long-term inflammation
environmental impact and sustainable development, and also could [49]. Therefore, future dietary formulations containing plant oil should
be compared to other types of animal production. Nevertheless, the consider the ω3/ω6 ratio as it strongly affects eicosanoid production
conservation benefit of substituting plant meal and oil for FM and FO and can help deal with stress and disease; which has the focus of several
is obvious. recent studies related with fish oil replacements (Table 1).
Fish nutrition and fish health Achieving good health through nutrition is relatively simple
by providing essential nutrients in the right amount to cater to the
It could be argued that the activity of intensive fish farming is, by specific requirements of different species and life stages. However,
definition, a stressful procedure, but such a consideration has to be disease caused by infections and microorganisms is known to be one
balanced against the need to produce food for human consumption. of the major constraints in the aquaculture industry, and fish may
Therefore, fish health is of upmost importance, in terms of an ethical be susceptible to such diseases, regardless of nutritional status. Feeds
and production point of view. Proper nutrition plays a critical role in containing antibiotics are commonly used to treat diseases. However,
maintaining normal growth and health of aquatic organisms. Good sick fish generally do not feed vigorously and therefore may not
nutrition can help mitigate the effects of stress, decrease the susceptibility consume enough medicated feed to ensure proper dosage. The use of
to disease, and boost the immune system; therefore it is important to antibiotics in aquaculture is also problematic because of the potential to
consider their diet in terms of stress and disease prevention, while the develop antibiotic resistant bacteria, the presence of antibiotic residues
diet may also serve as a primary method of administering medications, in seafood, destruction of microbial populations in the environment
immuno-stimulants and functional ingredients for fish. and suppression of the fish immune system [51,52]. Therefore, research
It is well established that all essential nutrients are required in has focused on functional constituents in feeds to improve growth, feed
sufficient quantity to sustain normal health. A deficiency in any required efficiency, health status, stress tolerance and resistance to disease to
nutrient can adversely affect health by impairing metabolic functions reduce antibiotic utilization in farms.
and increasing susceptibility to disease. Lipids, for example, are directly The concept of functional feeds is an emerging paradigm in the
involved in aspects of the immune response. LC PUFAs are precursors aquaculture industry, which are diets supplemented with feed additives
and modulators for eicosanoid production, including prostaglandins to improve health and disease resistance with minimal negative
and thromboxane via a synthesis pathway involving cyclooxygenase effects on the fish, consumers and the environment [53]. Dietary
and lipoxygenase enzyme activity. Arachadonic acid (ARA; 20:4ω6) supplementation of different non-digestible feed additives, such as
and EPA compete for the same lipoxygenase or cyclooxygenase, immunostimulants, probiotics and prebiotics have been found to be
therefore the dietary ω3/ω6 ratio determines the ratio of high activity beneficial for improving immune status, feed efficiency and growth
(inflammation) to low activity eicosanoids (anti-inflammatory). performance in various different fish species. Feeding animals with
Given the recent focus on FO replacements, studies have compared immunostimulants prior to an infection or in situations known to
the immune and inflammatory response of fish fed different plant oil result in stress will elevate defences and thus provide protection against
diets after infection by a virus or pathogen. For example, barramundi otherwise potentially severe or lethal conditions [52]. Probiotics
(Lates calcarifer) fed a diet with either FO or echium oil had a longer are live microbial organisms, non-pathogenic and non-toxic, which
lasting and enduring response in eicosanoid concentrations following beneficially affect intestinal microbial products, reduce gut pH
a week after bacterial infection, compared with those fed on rapeseed and release chemicals with bactericidal effects on other microbial
oil. Echium oil has a comparatively higher ω3/ω6 ratio than rapeseed populations, thus improving the immune response. The improvement
oil, and proved more effective in compensating for stress [48]. Another in innate immunity in the fish helps against pathogenic bacteria, as well
study tested functional feeds for Atlantic salmon infected with heart as against environmental stressors [54]. One of the major bottlenecks in
and skeletal muscle inflammation, an emerging viral disease caused the industry is high mortality during larval first feeding. Probiotics may
by a novel salmon reovirus [49]. The functional feeds were formulated be administered to live feed which act as live carriers of the bacteria and
with reduced digestible energy levels achieved through lower lipid subsequently fed to fish larvae. This method has proven to be successful
content, and specifically increased the EPA level and increased ω3/ in European sea bass larvae (Dicentrarchus labrax) fed artemia cultured
ω6 PUFA and EPA/ARA levels. The LC PUFA biosynthesis activity with bacterial enrichment, which affected the larval immune response
in salmon fed the standard diet was low compared to those fed the and reduced mortality rates [55] Evidence of the beneficial effects of
functional feed, limiting the availability of anti-inflammatory LC ω3 probiotics gave rise to the concept of prebiotics, which are indigestible
PUFA, which highlighted the crucial role of the diet in maintaining oligosaccharides and dietary fibre that selectively stimulate growth
the availability of ω3 PUFA in membrane phospholipids. Both EPA and/or activate the metabolism of health-promoting bacteria in the
and ARA reduced considerably after 16 weeks with infection; these gut, and depress the proliferation of harmful microbes, thus improving
losses were mitigated by the functional feed in comparison to those fed intestinal balance in the fish [51]. Beta-glucan for example, can act
the standard feed, which likely contributed to the better outcome of as both an immunostimulant and a prebiotic. It is a polysaccharide
fish fed the functional feed. In addition, reduced dietary lipid content with glucose as its only structural component and forms the major
altered the expression of key genes (Δ5 and Δ6 FAD) in fish fed the constituents of the cell wall of some plants, fungi, bacteria, mushrooms,
functional feeds, potentially increasing their capacity for endogenous yeast, and seaweeds. Beta glucan appears to be the most promising
production and availability of anti-inflammatory EPA [49,50]. These feed additive to enhance growth, survival and immunity for a several
results highlight the benefits of using diets with reduced dietary lipid different of farmed fish species, supplemented at levels between 0.1
and increased EPA levels on disease outcome. Factors that modulate to 1.0% (Table 2). Functional feed additives, such as beta glucan, can
and dampen the inflammatory process might be the key to mitigating improve immunity, feed efficiency and growth performance of farmed

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Citation: Hixson SM (2014) Fish Nutrition and Current Issues in Aquaculture: The Balance in Providing Safe and Nutritious Seafood, in an
Environmentally Sustainable Manner. J Aquac Res Development 5: 234 doi:10.4172/2155-9546.1000234

Page 6 of 10

Compound Species Inclusion Result Reference

Immunogen® Rainbow trout 0.2% Up-regulated immune related genes (lysozyme, TNFα), lower heat shock protein Ahmadi et al. [80]
(Oncorhynchus (Hsp70) gene expression, stimulated humeral immune response, elevated
mykiss) disease resistance
Brown algae beta glucan Rainbow trout 0.2% Increase in phagocytic activity in head kidney macrophages and a significant Morales-Lange et al. [89]
Laminaria digitata increase in the production of TNFα and IL-8 in gill tissue
Yeast beta glucan 0.2-0.4% Increased compliment and immunoglobulin Rozita et al [91]
Saccharomyces cervisiae
Yeast beta glucan 0.2% Successfully induced inflammatory gene expression (interleukin-1β) in the head Poley et al [90]
(Aquate®) kidneys of fish infected with Lepeophtheirus salmonis; lice burdens were lower
on fish fed Aquate® (19%)
Yeast-derived beta glucan 0.1% Up-regulation of IL-1β after challenged with Vibrio anguillarum Lokesh et al. [88]
Beta glucan 0.1% Significantly increased growth, antioxidant activity, and digestive enzyme activity Guzman-Villanueva
after exposure to lipopolysaccharides et al. [89]
Oyster mushroom beta 0.5-2.0% Significant shifts in white blood cell, lymphocyte, monocyte, neutrophil Dobsikova et al. [84]
glucan Pleurotus ostreatus granulocytes-segments, as well as plasma concentrations of glucose, albumins,
cholesterol, antrium and chlorides
Macro Gard® 1% Differentiated the regulation of mRNA expression of claudin genes and prevented Syakuri et al. [93]
an intestinal inflammatory response post Aeromonas hydrophil intubation
Macro Gard® 1-2% Fish fed diets containing 1% and 2% MacroGard® showed significant Kuhlwein et al. [87]
improvements in weight gain, specific growth rate and feed conversion ratio
compared to fish fed both the control and the 0.1% MacroGard® containing diet
Mushroom beta glucan 0.2% Enhanced the lysozyme activity, alternative complement activity, phagocytic Chang et al. [82]
Ganoderma lucidum and activity and respiration burst; significantly enhanced the protection of grouper
Coriolus versicolor against Vibrio alginolyticus up to 16% compared to control group
Macrogard® 0.1% Significantly reduced mortalities after challenging with Aeromonas hydrophila Talpur et al. [94]
with significant improvement in red blood corpuscles, white blood corpuscles,
pack cell volume, haemoglobin concentration, erythrocyte sedimentation rate and
serum protein content in treated groups over the control
Table 2: Various forms of beta-glucan in functional feeds found to improve fish health and immunity for different farmed fish species.

fish [2] with minimal impact on the environment and the consumer; such as improving feed efficiency, performance, product quality and
therefore commercial feeds may standardize its use in feeds. health; and can indirectly benefit production, such as pest and disease
resistance, and increased yield [58].
Biotechnology to improve fish nutrition
A controversial and interesting topic of discussion is GM salmon.
Bridging the gap between problems and solutions in aquaculture Aqua Bounty Technologies was the first company to submit an
will require insight and creative applications. Biotechnology application to the Food and Drug Administration (FDA) for a GM
research and applications can help the aquaculture industry develop animal for food consumption in 1995 and has been in the regulation
sustainably, and efficiently produce good quality food for a growing system to date. In order to acquire FDA approval, Aqua Bounty has
human population. There are many new applications of biotechnology had to demonstrate the food’s safety, and gauge the environmental risk
in aquaculture; however this review will focus on how biotechnology of the sterile fish escaping its tanks and successfully breeding with wild
can improve fish nutrition for the purpose of safe, nutritious and salmon. By contrast, the FDA approved the first GM crop for human
sustainable food production. In particular, genetic modification will consumption (the Flavr Savr tomato) after just three years of regulatory
have the most significant impact on fish nutrition in aquaculture. consideration [59]. Research has shown that growth enhanced transgenic
Producing environmentally sustainable and nutritionally sound Atlantic and coho salmon (Oncorhynchus kisutch), have the capacity for
feeds for farmed fish has proved to be a challenge. The application increased daily feed intake, increased growth rates and improved feed
of biotechnology to produce feed ingredients that are specifically utilization due to an enhanced capacity for protein biosynthesis over
enhanced for aquaculture feeds can help alleviate pressure on wild non-transgenic fish [60,61,62]. Transgenic coho salmon demonstrate
fisheries that produce FM and FO. Research on GM crops that are an enhanced ability to utilize dietary carbohydrates (in addition to
enhanced specifically for fish feeds is in the forefront of both plant lipid) as a major source of non-protein energy, which is not typical for
and aquaculture research. The limiting factor in using plant oils is non-transgenic salmon [63]. The utilization of carbohydrate allows for
the lack of LC ω3 PUFA, namely DHA and EPA. GM plants that can greater amount of dietary protein to be spared for rapid tissue growth
produce their own DHA and EPA, and that can be grown sustainably, [62]. The altered gene expression and different metabolic enzyme
is certainly well sought after by the aquaculture industry. Development capacity in transgenic salmon allows for a shift towards the use of fatty
of such a crop is in the research and development phase, but is not yet acids and glucose for energy-production purposes, conserving amino
available commercially. A study by Ruiz-Lopez et al. [56] demonstrated acids for protein biosynthesis [64]. In a study by Tibbetts et al. [62],
fish oil-like levels of DHA and EPA produced by the transgenic transgenic Atlantic salmon consumed a significantly higher amount
oilseed camelina, while avoiding accumulation of undesirable fatty of feed on a daily basis, but also had better FCRs, higher nitrogen
acids. Atlantic salmon fed GM rapeseed oil and yeast containing high retention efficiency and achieved target weight gain in a shorter period
levels of DHA and EPA showed high retention of LC ω3 PUFA in the (40%) than non-transgenic fish. The transgenic salmon in this study
flesh [45,57]. The benefits of producing LC ω3 PUFA in plants are demonstrated a higher cellular capacity to direct dietary non-protein
clear: a sustainable and non-contaminated source of important fatty energy towards satisfying their daily metabolic energy requirements,
acids essential to human nutrition. GM crops may also have other allowing for a higher proportion of dietary amino acids to be directed
enhanced features that can directly benefit aquaculture production, towards protein biosynthesis rather than catabolised as a dietary

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Citation: Hixson SM (2014) Fish Nutrition and Current Issues in Aquaculture: The Balance in Providing Safe and Nutritious Seafood, in an
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Page 7 of 10

energy source. Dietary protein represents the highest proportion to the high level of ω6 fatty acids, terrestrial plant oils do not contain
of salmon feeds and also the major source of nitrogenous pollution EPA and DHA, which causes a lower concentration of these PUFA in
from salmon farming; therefore transgenic salmon could improve farmed seafood products compared to farmed fish that were fed a FO-
production efficiency through better feed utilization, which could based diet. Most consumers are not fully aware of the lower DHA and
result in more sustainable salmon production. Furthermore, transgenic EPA content now occurring in many farmed fish species as a result of
salmon were found to have a higher expression of Δ5 and Δ6 FAD, using alternative feeds in recent years. A study by Nichols et al. [70]
which might help compensate for a potentially higher demand for investigated the LC ω3 PUFA content in farmed Atlantic salmon and
EPA and DHA resulting from rapid growth [64]. This also has positive barramundi samples from 2002 compared to samples from 2010-2013.
implications for the use of alternative FO feeds for transgenic fish in In comparison to the 2002 samples, both species contained decreased
terms of their ability to synthesize EPA and DHA at an increased rate. levels and contents of LC ω3 PUFA. Atlantic salmon in 2002 contained
However, there are environmental concerns related to GM salmon, 2014 mg/100 g, decreasing to 975 mg/100g in 2013. Barramundi in 2002
particularly the potential impacts of escaped GM salmon on wild contained 1970 mg/100 g, decreasing to 790 mg/100 g in 2013 [70].
salmon; although GM salmon produced by Aqua Bounty are sterile, These changes have resulted from the use of new, sustainable and lower
thus eliminating the potential of interbreeding between escaped GM cost ingredients in the feed. Despite significantly lower LC ω3 PUFA
salmon and wild salmon. There are also health and safety concerns amounts in farmed salmon compared to a decade ago, a human study
that pertain to the immediate and long-term effects on human health found that addition of farmed Atlantic salmon to the diet twice per
after consumption of GM salmon products. Initial tests on the safety week for 4 weeks at portions of 180 g and 270 g modifies phospholipid
of consuming transgenic carp and tilapia show no apparent negative fatty acid proportions of ω3 and ω6 in a level associated with decreased
health impacts on mice and humans [65]. However, allergens or toxins risk for cardiovascular disorders [71]. Regardless, there are strategies
may be produced as a result of gene transfer if the transgene codes for to increase the ω3 PUFA content in the lipid profile of farmed fish. In
a protein that induces expression of a previously inactive toxin (e.g., the present time, even using plant oils that are high in LNA but do not
shellfish protein used to create a transgenic fish), and the expression of supply DHA and EPA is still beneficial for humans simply to achieve a
bioactive proteins which may continue to possess bioactive properties higher ω3/ω6 ratio in the diet. Finishing oil diets with higher inclusion
following consumption (e.g., growth hormone) [65]. Research on the of marine oils at the end of the production cycle can elevate DHA and
effect of GM fish on human health should be conducted, as well as the EPA levels immediately before harvest. This may result in grades of
effect on product quality. farmed seafood products that contain a higher content of LC ω3 PUFA
than standard (and lower cost) products grown using a mainly non-
Fish nutrition and human health marine oil based diet [70]. In the future it is likely that GM plants with
The world will be very different in 2050: the global population will endogenous DHA and EPA will be used in fish feeds. The effect of GM
likely increase to nine billion, and about 50% more food will be needed foods on human health is not well known, however, many different
to sustain the quality of human life [66]. Rapid population growth and types of produce and processed foods available today are non-labelled
increases in fish consumption has led to rapid increases in global fish GM, so many North American consumers have purchased GM foods
demand. The expansion of aquaculture will fill this gap and relieve on a regular basis for at least a decade. However, there have been no
pressure on capture fisheries, which have been steadily declining studies to date on the long term effect of GM ingredients fed to farmed
[67]. The benefits of consuming a diet rich in seafood are based on fish on human health. This will be a critical area of research in the
high levels of LC ω3 PUFA and high quality animal protein, essential future.
amino acids, vitamins, and minerals [67]. There are numerous benefits Dietary changes may also affect the product quality of the
of aquaculture being the main supply of seafood for the world, from fish, which may change public perception of farmed fish. Sensory
a nutritional, economic and social standpoint; however the industry evaluations, including taste, colour and texture are frequently part of
has received much public scrutiny regarding toxins, contaminants, studies that evaluate changes after feed ingredients have been altered
hormones, and antibiotics in farmed fish, which raises questions about in the diet, particularly replacements of FM and FO. Although many
public health and safety [68].The composition of the feed affects the studies have reported that alternative protein and lipid sources can
final composition of the product; therefore fish nutrition directly affect the sensory quality of farmed fish, results in the literature are
impacts consumers and must be considered in terms of safety and divided and contradictory [21], therefore it is difficult to pinpoint if
human nutrition. certain ingredients clearly affect sensory quality because it can depend
It is obvious that the use of FM and FO must be significantly on the study. Interestingly, some studies showed that salmon fed plant
reduced in aquaculture feeds in order to be environmentally oils had less rancid and marine characteristics and were preferred
sustainable. However, using sustainable alternative feed ingredients to over salmon fed FO [72]. Consumer perception and satisfaction
feed fish could potentially affect human health. For example, terrestrial are an important aspect of fish nutrition; therefore the use of GM
plant oils are typically high in ω6 fatty acids, specifically LNA, which ingredients or the production of GM salmon needs to be evaluated for
get stored in significant amounts in flesh of the fish that are consuming sensory quality due to possible biochemical changes that affect sensory
these feeds. From a human health standpoint, increases in ω6 fatty acids characteristics.
in salmon fillets are a concern. When mice were fed farmed salmon Contamination of fish tissues with organic and inorganic
raised on diets with 100% soybean oil, the excessive dietary LA elevated contaminants has been a pervasive environmental and public health
endocannabinoids in the liver, increased weight gain and counteracted issue. From a human health perspective, concerns regarding relatively
the anti-inflammatory properties of EPA and DHA [69] and also high levels of Persistent Organic Pollutants (POPs) (e.g. Polychlorinated
exaggerated insulin resistance and increased accumulation of fat in the Biphenyls, PCBs) and inorganic contaminants (e.g. heavy metals) in
liver in rats [34]. The typical “Western” diet is already dangerously high farmed salmon have raised questions regarding the relative health
in ω6 fatty acids, which is why it is so important to consume fish that risks and benefits of farmed salmon consumption [73]. However, the
are high in ω3 fatty acids to balance this ratio. However, in addition accumulation of some contaminants is the result of indirect exposure,

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Environmentally Sustainable Manner. J Aquac Res Development 5: 234 doi:10.4172/2155-9546.1000234

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Citation: Hixson SM (2014) Fish Nutrition and Current Issues in Aquaculture: The Balance in Providing Safe and Nutritious Seafood, in an
Environmentally Sustainable Manner. J Aquac Res Development 5: 234 doi:10.4172/2155-9546.1000234

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