Introduction to Fish Bioenergetics

The flow of energy through a biological system and the system's inherent energy
requirements are covered by the term 'bioenergetic'. In biology energy is required to
maintain life, grow and reproduce.From a fish farming point of view, growth is the
main area of interest. The main energy source for consumers (as opposed to producers
such as plankton and plants) is their diet or food. This provides the energy to drive
chemical processes giving rise to new tissues, to help in osmoregulation, to aid
digestion (the means by which consumers unlock energy stored in food) and so forth.
[Note: energy cannot be made or destroyed but can only be converted from one form
to another].

Energy taken in by fish through digestion of food is used ultimately in one of three
ways - that is for growth, metabolic processes and that lost to the fish through waste.
During digestion the main components of the diet (protein, fat, carbohydrate) are
broken down into carbon dioxide and water with heat as a by-product, the latter being
rapidly dissipated to the surrounding water. The energy liberated is temporarily stored
in special 'energy compounds', the main one being adenosine triphosphate (ATP),
which beome cellular level energy sources for functions such as protein manufacture
(i.e muscle growth), swimming and so on.

Energy Flow in Fish

All food, whether natural or man-made, has a total or gross energy value. However
not all of this is accessible or usable by fish - in other words it is indigestible and will
be lost as faecal energy material. Of the digestible components of the food, not all can
be metabolised completely and further energy is lost through excretion in urine and
through respiration across the gills. The remaining metabolisable energy can then be
used for maintenance, movement and growth, although a small amount is lost through
heat production. The figure below shows this flow of energy along with some
approximate percentage values.
Thus approximately one third of a diet is used by fish for growth. As muscle is
essentially protein, more than 33% of the diet needs to be in the form of protein to
ensure maximum conversion and growth

Metabolism of Food
Following digestion and absorption in the gut, the base components of the nutrients
which constitute the fishes' diet are used in different ways. Chemical reactions result
in either liberation of their energy through final breakdown - this is known as
catabolism - or production of new molecules and tissues - known as anabolism. The
sum of these catabolic and anabolic reactions is referred to as metabolism.

It is the metabolism of the main dietary ingredients of protein, fat and carbohydrate
that is of prime importance in terms of energy for maintenance and growth. Through
digestion proteins are broken down into amino acids, carbohydrates to simple sugars,
and fats to glycerol and free fatty acids (FFA). On absorption these smaller
components end up in the liver, although fats take a circuitous route through tissue
fluids and part of the lymphatic system known as the lacteals.

A number of metabolic processes occur in the liver. Amino acids are removed from
the blood and stored temporarily prior to transport through the body for the synthesis
of new fish proteins. There is a continual breakdown and synthesis of proteins which
are used in cell and tissue structures, and enzymes. Any excess will be used for
maintenance/movement energy as it is broken down to carbon dioxide, water and
ammonia. Absorbed sugars (from dietary carbohydrates) are converted to glucose in
the liver which can then either be burned to supply energy or recombined to form
glycogen which is stored in the liver as an energy reserve. The processes involved in
the synthesis of proteins, fats and carbohydrates are shown in each of the diagrams
below.

Nutrient metabolism showing inter-relationships and links into Glycolysis,

Krebs, etc

Each of these links is shown in more detail below.

SOURCE: De Silva S.S & Anderson, T.A. (1995)
Fish Nutrition in Aquaculture
Chapman & Hall

Glycolysis
Krebs/Citric acid cycle
Fats are readily catabolised as a source of daily maintenance and movement energy
needs. If there is too much fat in the diet, or fish are overfed for a long period of time,
fat deposition will occur. Fats can be synthesized from amino acids and glucose as
well as from glycerols and free fatty acids - it is thus easy for fish to accumulate fat
from their diet. Fat is also the most economical way to store energy as it has an
inherent higher energy latency. The above metabolic pathways are shown in the
diagram.
Feed Metabolism

<TD&NBSP;<
TD>
Amino
Glucose Triglycerides
Acids
Protein Glycogen Fatty Acids
Free Fatty
Carbohydrate
Acids

Feeding and Oxygen Demand

All chemical processes, including metabolism, are energy-consuming.The burning or
combustion of nutrients requires oxygen as a catalyst (in the same way as a fire
requires oxygen to stay alight). A high energy consumption necessarily means a high
oxygen demand. Digestion and metabolisation require energy so there is an increased
oxygen demand by fish after feeding. It is therefore important to consider a potential
site with respect to dissolved oxygen levels in terms of this increase in demand for
oxygen at feeding times. Due to the phenomenon of scaling, this problem does not
directly increase as the weight/size of fish does - larger fish require less energy per
unit body weight than smaller fish do. However as oxygen and temperature are
inextricably linked (the higher the temperature, the lower the solubility of oxygen),
temperature regimes need to be carefully considered to ensure optimal growth of
stock.

Feeding and Oxygen Demand

Diet Composition
Most people will be familiar with the main components of food from reading the
cornflake box at breakfast time. Diets comprise three major constituents in the form of
proteins , fats and carbohydrates plus three or four minor ingredients in the shape of
vitamins, minerals, water and, for older salmonids in particular, pigments.

Just as nutritional requirements change as animals age, so the composition of fish

diets alters as the fish goes through its various developmental stages. Naturally those
involved in fish feed formulation need to know the nutritional requirements of a
particular stock species and the means by which these can best be met to give
maximum growth in a cost-effective manner. An obvious starting point is to look at
what the stock would eat in the wild at various life cycle stages, to ascertain
nutritional profiles and attempt to replicate them. This is indeed what happens - most
proprietary fish diets are based on piscine raw materials.

Within modern aquaculture, a stocks' nutritional needs can be met using different
types of diet depending on the stock being cultured and what life cycle stage is
involved. For herbivorous and omnivorous species (such as cyprinids) under
extensive or semi-intensive culture, a natural food supply is utilised with, in the latter
type, some supplemental fertilisation of holding units to encourage plant and plankton
growth. A variety of diet types can be utilised for carnivorous species ranging from
wet (minced/diced raw fish) through moist (basic fish meal and additives mixed on
site with freshwater) to the well-known modern pelleted diets. For some species a live
diet is required during larval development, as in the feeding of Artemia to larval
halibut and cod.

Whatever diet is used, it must be nutritionally balanced to meet the energy budget of
the stock so as to optimise growth and profit. Total energy value of diets will vary
with the proportion of its various ingredients. As fish regulate their energy intake to
match their energy needs, they in effect regulate the absolute amount of proteins that
they ingest. Thus the balance of ingredients in a diet may be more important than
definitive levels of one particular nutritional component. Consequently diets deemed
to be 'low energy' ones may result in as much growth as a 'high energy' one, provided
it is nutritionally balanced.
Dietary Components
The main components of fish diets, man-made or natural are:

Protein
Carnivorous fish, such as salmonids, require diets rich in protein (about 40 - 50%
content) compared to omnivorous or herbivorous fish such as carps (typically 25 -
35% content) for optimal growth.

Proteins are composed of a number of building blocks known as amino acids.

Different proteins are formed from around 20 known amino acids, the calorific value
(energy worth) depending on the amino acids present, their arrangement(s) and
availability following absorption. Fish appear to be incapable of synthesizing 10
amino acids totally or in sufficient quantities to meet their needs - these are referred to
as essential amino acids and must be present in their original, pure form in any diet.
The non-essential amino acids can be either synthesized from their basic raw
materials (carbon, oxygen, hydrogen and nitrogen) or interconverted from other
amino acids. These essential and non-essential amino acids are listed in below.
 Essential Amino Acids Non-essential Amino Acids

Arginine Adenosine

Histidine Alanine

Isoleucine Aspartic Acid

Leucine Cystine

Lysine Glutamic Acid

Methionine Glycine

Phenylaline Proline

Threonine Serine

Trytophan Tyrosine

Valine ----

Provided the correct ratio and amount of essential amino acids are provided, protein
synthesis can occur and new tissue can be laid down. As amino acids cannot be
stored, they must be present in the correct amounts. If one amino acid is limiting or
the protein component of the diet is in excess, proteins may be used as an energy
source for maintenance and movement. This process, known as deamination, is
energy- expensive and also leads to increased nitrogen waste production which can
have environmental consequences.

Protein sources, mainly as fish meals, are the most expensive components of diets and
are also poor as an energy source. It therefore pays to utilise or spare all dietary
protein for growth and supply maintenance and movement energy from a cheaper and
more readily accessible source. This is the accepted approach with lipids, and to a
lesser extent carbohydrates, meeting daily energy requirements and demands. Levels
of protein in formulated pelleted diets have changed little over the last 20 or so years
since they were introduced, remaining at around levels of 40 - 47%.
Lipids/fats

Of the major nutrients, lipids are the group most easily digested and metabolised. On
a unit -for-unit basis they also provide more energy than carbohydrates or proteins.
Fish also require certain essential fats for the correct structural arrangement of
membranes such as cell walls - long-chain polyunsaturated fats are necessary for
carnivorous fish, particularly salmonids. Over the years the fat/oil content of diets has
steadily increased, from around 6% some 20 years ago, to 10 - 12% 15 years ago, 18 -
21% five or six years ago to current levels of 25 - 33%. These so-called 'high energy'
diets have to be given with care as problems can and do occur.

Dietary lipids also serve as carriers for absorption of other nutrients including fat-
soluble vitamins and pigments for flesh coloration in salmonids. As with protein,
dietary lipids are provided from marine fish oils which are rich in essential fatty acids.
Care has to be taken with regard to storage as PUFAs readily oxidise and become
rancid.

Carbohydrates
These act as a third source of dietary energy. In addition to being cheap they also act
as binding agents for feed and increase the palatability of proprietary diets.

Carbohydrates are divided into two main groups - the sugars and the non-sugars. The
former includes such things as glucose, sucrose and lactose whilst the latter contains
the more complex materials such as starch and various polysaccharides. From a fish
feed aspect, glucose and starch are of importance. As carnivorous fish, salmonids lack
sufficient quantities of the enzymes necessary for efficient digestion and metabolism
of most carbohydrates. Levels in proprietary diets are consequently low,
carbohydrates giving way to increases in other components such as oil. For
herbivorous fish, the carbohydrate content of man-made diets is correspondingly
higher.

Vitamins

Fish require a certain number of vitamins for good growth and health, the amount
varying according to the age (and health) of the fish. They are all present in the
dietary raw materials but processing and subsequent storage can result in variable
levels being left when the time comes to feed the fish. As a result the raw material is
assumed to contain no vitamins and sufficient amounts of synthetic vitamins are
added during the manufacturing process.

Vitamins fall into two categories - water-soluble and fat-soluble. The former includes
Vitamin C and various B vitamins (B1, B2, B6, B12) and the latter Vitamins A, D, E,
and K. All these vitamins and several others are added to feeds in small amounts (e.g.
10-500 ug per kg dry weight of food).
Note: the fat-soluble vitamins can accumulate in fish and cause vitamin poisioning of
hypervitaminosis. A lack of vitamins in the diet can lead to growth and health
problems.

Minerals
Fish require a supply of essential minerals for healthy growth. In the wild these are
obtained from the surrounding water and the tissues of prey items. The main ones are
calcium and phosphorus which are added to diets in the form of carbonates and
phosphates. As synthetic minerals are much cheaper to produce than vitamins and are
much more stable, mineral additives do not contribute greatly to higher feed costs.
Other minerals required include iron, magnesium, manganese, sodium, potassium and
cobalt.

Pigmentation

Wild salmon and trout are characterised by a pink-red flesh colour derived from the
algae and crustacea that form a large part of their diet. The colour results from the
accumulation of carotenoid pigments in the muscle. Salmonids cannot synthesize
these pigments themselves. From a marketing point of view, it is necessary to provide
pigment in the diet - customers are not going to buy grey-looking salmon!

The predominant pigment in wild salmon is astaxanthin which is also available

commercially. A cheaper alternative is canthaxanthin which produces an orangey red
colour rather than a "salmon pink" one. It is also added in smaller amounts to
feedstuffs. Another source of dietary pigment is the red yeast Phaffia rhodozyma
which contains astaxanthin in concentrations ten times higher than that found in
shrimp and prawn waste. Very little use has been made of this yeast because of
production problems and consequent cost. Whatever the source, pigmentation may
account for as much as 25% of the total cost feed.

Of the dietary pigment available to the fish, 40 - 60% (depending on age, time of year,
diet) is absorbed through the gut whilst the rest is lost via the faeces. The absorbed
pigment is transported by the blood throughout the body. 30-40% of the absorbed
pigment is absorbed by the skin, some is processed by the liver whilst some is
deposited in the muscle. Overall, only 7-10% of the total dietary pigment available
appears in the flesh.

Pigmentation is influenced by a variety of factors. Different species absorb more than

others - e.g. Pacific salmon and trout species absorb more than Atlantic salmon. As
fish get bigger, more pigment is absorbed -it is unclear whether this is due to age or
increased food intake.

Fish Weight and Pigmentation

Young salmonids do not absorb pigment very well. Pigmentation is only laid down
and improves colour from weights of 200gm for salmon and 100 gm for trout .
Consequently pigmented diets are only fed to trout from six months of age and to
salmon following smolt transfer and usually at the point where fish are switched to a
grower feed from a smolt feed.

Pigmentation and age

During maturation, pigment in the flesh is re-absorbed and re-deposited in the skin,
giving salmon their characteristic reddish breeding colour. In females, pigment is also
transported to the ovary and deposited in the eggs - hence their red colour. Overall
60% of muscle pigment can be lost during maturation.

Sexual maturation and pigmentation

Although canthaxanthin is cheaper to produce and therefore keeps the cost of feed
down, the industry switched to the exclusive use of astaxanthin in late 1989 when
high levels of canthaxanthin were found in salmon samples taken from supermarket
shelves. Although banned within the UK as a direct food additive (colouring) for
human consumption, it is still legal as an animal feed additive. However, to avoid
potential problems the salmon farming industry switched to astaxanthin in 1990.
Obviously this meant that the farmer was having to pay more for his food. The ban on
canthaxanthin followed from its use in "oral-tanning" pills, primarily in America,
which, when taken in large, prolonged doses caused severe eye problems in a few
cases. More recently there has been a switch to mixed (asta/cantha) and pure
canthaxanthin diets following a relaxation of the ban, although the end buyer
(supermarkets) still demand fish fed only on astaxanthin.

Incidences of fish with a high tissue fat level exhibiting poor colour have been
reported. As both pigments are fat-soluble this is perhaps not surprising. Recent
reports appear to indicate that both very low and very high fat tissue levels can result
in poor colour - the latter situation could occur in farmed fish fed to satiation
throughout the production cycle. Attempting to solve this potential problem by
increasing feed pigment levels beyond those legally permitted would be of no benefit
as any differences would not be visually discernible. Starting pigmented diets earlier,
e.g. in August rather than in September or October, as is often the case, may be
beneficial in establishing a deeper background colour.

Alternative dietary sources of nutrients

Of the world's total production of fish meal and fish oil, some 5% and 27%
respectively goes into the salmon feed production market. Increased concerns about
the sustainability of industrial fishing and the morality of using fish to grow more fish
has intensified the search for alternative sources. Scarcity of fish in some parts of the
world has led to increases in fish meal prices which are naturally passed down the line
to the farmer.

Fish oil is characterised by a large proportion of long chain, polyunsaturated fatty

acids (PUFAs) which are essential for growth. High levels of PUFAs in the diet of
humans are encouraged from a health aspect. It is known that oils derived from plants
(soya, sunflower, rapeseed) also contain some PUFAs but do not have the range found
in fish oils nor do they have a high proportion of linolenic (n-3) and linoleic (n-6)
fatty acids. Thus it would not be possible to use plant oils alone. However, trials in
which dietary lipids are made from combined fish and plant oil sources (3:1 ratio
respectively) indicate no problems with fish growth, health or body composition and
carcass quality. The only area of contention is that of taste - argument continues as to
whether there is an alteration of taste to a less 'fishy' one.

Replacement of the protein component of farmed fish diets is not so straightforward

due to the requirement of the 10 essential amino acids. Alternative sources are usually
lacking in one or more of these, meaning that they have to be either specifically
isolated from fish meal or manufactured. This is an expensive process. Work
continues on finding appropriate substitutes.
Dietary Problems
A diet deficient in either the major or minor nutrients will ultimately result in health
problems in the stock. Feed formulation is such that pelleted or other manufactured
diets provide stock with a good nutritionally balanced and sufficient food supply.
However things can go wrong if environmental factors exceed normal tolerance limits
for the stock (e.g., higher than expected water temperatures, poor quality raw
ingredients are used or feedstuffs are incorrectly stored).

Dietary problems concerned with carcass quality rather than fish health through the
production cycle have on occasion come to the fore in recent years. Among the
problems manifesting themselves have been excess, running/flowing oil in flesh,
gaping of fillets and a loss of fillet colour/pigmentation. Fish farmers blamed all these
on the feed on the grounds that nothing else within the industry had changed other
than the relatively rapid increase in the percentage oil content of diets. This was rather
unfair in that, over the same period of time, there had been additional vast
improvements in husbandry techniques and overall fish health through increased
availability and use of vaccines.

One of the major outlets for farmed salmon is the fish smoking industry. Fish smokers
complained in the early 1990s that fillets were difficult to slice, colour was poor and
oil was leaching from fillets into packs compromising seals, shelf life and freshness.
Product traceability within quality control systems meant the product could be traced
back to the farms where they were reared. Results of inquiries did not present a very
clear picture but the finger was pointed at diet as the main culprit. This was on the
basis of a 10% or more increase in the oil content of pelleted diets (22/24% to
30/33%). The link of high oil in diet and high oil in fillet was easily made.

Whilst dietary fats will be utilised by fish as an energy source, fish will accumulate fat
if they are present in excess of its needs. This is a natural biological reaction by fish -
these energy (fat) stores can be utilised in times of scarcity. However it did not and
does not fit in with what the fish farmer was trying to produce, namely a low fat
(14%), lean fish for smoking. Investigations revealed that it was not the high oil in the
diet per se that was the problem but rather the way in which these feeds were being
administered to stock. Satiation feeding was the norm to promote appetite, better
growth and feed conversion efficiency. As energy expenditure in a cage is fairly
limited, stock were being presented with an excess of oil at each meal leading, over
time, to accumulation in the flesh. Additional associated problems included gaping in
fillets and apparent less strong pigmentation.

Proposed Dietary Solutions

Proposed solutions took two main routes. First was to switch to a lower energy/lower
oil content diet so that, to maintain existing energy expenditure, fish had to call on the
oil reserves within their muscle tissue. An alternative was to continue with the same
diet but to use a reduced ration, again to encourage use of in-built stores. In short,
changes to feeding management were deemed to be the answer rather than wholesale
changes to the type of diet.
Reduction of energy input aided the gaping problem. This was thought to be a
consequence of too rapid growth in muscle tissue resulting in incorrect tissue
formation and insertion of muscle myofibrils into the myosepta. Thus, on filleting the
muscle tissue tore very easily. The colour problem was associated with the fact that
pigments used in fish diets are fat-soluble and with excess around, some pigment was
being re-absorbed. These changes to feed management resulted in a resolution to the
problem in time - it took several months as opposed to being an overnight cure.

The range of diets now available to farmers mean that they have the capability to
produce the quality of fish required by the various market outlets through diet
manipulation.

Specialist Diets
Specialist diets include those containing pigments, medications , hormones and new,
so-called finishing diets.

Medicated Diets.

Incorporation of medications into feeds is not so common these days with the advent
of vaccinations, although they are still used. Their main use was to combat bacterial
infections and a few internal parasite problems.

One of the main drawbacks of medicated feeds is that they obviously need to be
consumed to be effective - most health problems usually result in a loss of appetite
plus the presence of the drug/chemical may make the food unpalatable, thus
exacerbating the problem. Fish size, water temperature, extent of disease all influence
feeding rates and by association the efficacy of any treatment. It should also be borne
in mind that fish eat at different rates and there is no guarantee that each individual
fish will obtain the required dose to combat the disease.

The commonest medicated diets are those containing antibiotics or other anti-bacterial
comounds such as sulphonamides and nitrofurans. Sulphonamides have been around
for a long time and were used 30 years ago to control furunculosis in salmon
populations. One (sulphamerazine) is still in use today. They have their drawbacks in
that they are unpalatable to fish and can cause kidney damage in high doses. They
have been superseded by "potentiated sulphonamides" - i.e. a general sulphonamide
plus another compound which increases the remediative effects but reduces any side
effects. A common example is tribrissen which is a combination of sulphadiazine and
trimethroprim.

Nitrofurans are similar to sulphonamides but are obviously nitrogen-based rather than
sulphur-based. They are cheap to manufacture and readily available - a common
example is furazolidine.

Most effective are the true antibiotics such as oxytetracycline and oxolinic acid. These
are readily available as they are the same as those used in human medicine - this
means that they are only available in prescription from a vet. As with human bacterial
infections, repeated use of the same antibiotic can rapidly induce resistance in
baterical strains so that the subsequent treatments are useless. A second antibiotic has
therefore to be used in such cases. In some fish farming areas triple-resistant bacterial
strais have appeared, that is strains resistant to the three main antibiotic treatments.
Sulphonamides and/or nitrofurans have then had to be used.

Dosage rates vary according to the type of antibiotic being used but is generally
expressed as so many grammes per 100kg of fish per day. In order to calculate dosage
one needs to know the number and average weight of stock

For example:

No. of fish =12 000

Average weight = 1.5kg
Treatment - oxolinic acid = 1.0gm/100kg fish/day
Amount oxolinic needed =12 000 x 1.5 x 1/100 = 180gm/day

This quantity of antibiotic needs to be added to the daily feed for this stock, which is
approximately 125kg per day - i.e. 36 gm of oxolinic per bag per day over the
treatment period which lasts 10-14 days. For relatively small quantities like this the
farmer can do the mixing himself using a cement mixer, pellets, antibiotic and fish oil
to get the powder to stick. There is no guarantee with this method that each pellet gets
the correct coating - some will get none, others will get an overdose - and
consequently some fish will not get treated. Large quantities will need to be obtained
from the feed manufacturer.

Dosing rates for the common antibiotics in use are shown in A and B. When selecting
a feeding rate for treatment, select the one which is 0.5% lower than the one in use to
allow for the reduction in appetite which usually accompanies a health problem

A. Medicated feed calculations

Tribrissen Aqualinic
di-
40% powder 20%
Feeding Oxytetracyclin Furazolidone Sulphamerazin nbutyl
(Sulphadiazin powder
rate % e pure e pure tin
e& (Oxolinic
oxide
Trimethoprim) acid)
0.5 15.4 22.0 44.0 50.0 15.0 10.0
1.0 7.7 11.0 22.0 25.0 7.5 5.0
1.5 5.1 7.3 14.6 15.0 5.0 3.3
2.0 3.9 5.5 11.0 12.5 3.8 2.5
2.5 3.1 4.4 8.8 10.0 3.0 2.0
3.0 2.6 3.7 7.4 7.5 2.5 1.7
3.5 2.2 3.2 6.4 7.1 2.1 1.4
4.0 1.9 2.8 5.6 6.3 1.9 1.3
4.5 1.7 2.5 5.0 5.6 1.7 1.1
 5.0 1.5 2.2 4.4 5.0 1.5 1.0
5.5 1.4 2.0 4.0 4.5 1.4 0.9
6.0 1.3 1.8 3.6 3.8 1.3 0.8

B. Antibiotic dosing rates

Drug Dose (gm/100 kg fish/day

Oxytetracycline 7.5 for 5 - 10 days
Furazolidine 11 for 5 - 10 days
Severe: 22 for 1 day, 11 for 10 days Mild: 20 for 3 days, 0 for 2 days,
Sulphamerazine
20 for 2 days
Tribrissen 7.5 for 5 - 7 days
Oxolinic acid 1 for 10 days

Whichever method is used to incorporate the antibiotics, it is esential that all relevant
Health and Safety practices are observed. In addition it is essential that a withdrawal
period is observed after treatment prior to any fish being harvested. As all drugs are
controlled under the Medicines Act and they have the potential to accumulate in flesh
tissue, they all have a legal withdrawal period which must be adhered to. many areas
add an additional safety period - for example, in Shetland an extra 40 degree days for
every 100 legal degree days in added.

Hormone-treated Diets
Used in hatcheries to manipulate the phenotype sex of fish in the very early stages of
development. The sex of fish is genetically determined at some point during its
embryonic development and it is possible to influence the sex up to this point -
thereafter only radical measures work e.g surgery! With salmonids phenotype is
determined after hatching - by feeding hormone- laced food for the first 40 to 60 days
of first feeding the phenotype can be pre-selected, i.e. you can decide whether you
want all male or all female. The genotype remains fixed within the genes, thus a
genetic female (XX) can be forced to become a phenotypic male (grow testes which
produce functional milt with XX genotype) which when crossed with normal eggs
(XX) produce all female progeny.

The levels of hormone or steroid added to the diet are low (2-20 mg/kg of food) and
once the phenotype is set they can be discontinued so that they are not present in the
harvest stock. Use of low levels of steroids also improve growth and feed efficiency
in salmonids but there are marketing problems with their use.

Finishing Diets
These are a relatively recent development in stock feeding. The rationale behind them
is that they are used during the last few months prior to harvest to improve one or
more quality factors, mainly colour or tissue fat levels. This is achieved by altering
the dietary components of the feed - reduction (small) of oil levels, increased pigment
levels and increased levels of vitamins E and C. Examples of these diets are Trouw's
Harvest and Focus diets.

These diets should not be used to obtain further growth from the stock but should be
used once the selected harvest weight has been attained to round off the product so
that it is in the best possible shape.

Introduction to Feeding Management

It makes economic, environmental and common sense to feed any stock of fish in a
well-managed fashion. Any method employed has to satisfy the appetite demands of
the stock, promote good growth, be economically efficient and be capable of being
easily delivered to the fish. The 'what, when and how' of feeding has become very
important over the past 5 years or so as the price differentials of production and sale
prices have reduced.

Feeding can be divided into a number of areas - how much should be fed (ration size),
how does the ration fit in with growth potential, how often should rations be given
and at what time of day (feeding regimes), how are rations best delivered and how to
monitor the efficiency of the diet and its performance. These factors all interact and
need consideration in order to get the best out of both the diet and the stock. If one
factor is outside the optimal range, stock quality and income will ultimately suffer.
There may also be a need to utilise specialist diets at certain points of the production
cycle.

Ration Size in Feeding Management

The economy and productivity of a farm operation is influenced by both ration size
and the way the ration is fed. This is particularly true for aquaculture. Use of the best
diets, equipment and stock can still result in poor growth rates and food conversion
ratios if the stock is not properly fed. Feeding revolves around two main factors:

(1) How much food should be given with reference to the numerous biotic and
abiotic criteria which determine the appetite and nutritional requirements of
fish.
(2) How should feed be delivered with respect to times of day, the number of
feeds per day, the duration of each meal and the rate of application of feed.
Growth-ration relationship
Growth rate and ration size interact to determine FCR and are used to determine the
daily ration for a particular fish stock. The diagram below shows a typical growth-
ration-FCR relationship for salmonids.

Like all animals fish will lose weight when their nutrient intake rate falls below that
required for daily maintenance (Rmaint). As food availability increases, the quantity
consumed by the fish will also increase, giving a linear increase in specific growth
rate (SGR as % body weight per day) up to the point of maximum voluntary food
intake (Rmax). The mathematical relationship between food intake and growth rate
determines that gross FCR decreases from infinity in fish fed at a level that satisfies
their maintenance requirement, to some maximum value at the maximum
consumption rate. If fish are fed above their appetite, then the extra food will be
wasted and an artificially high FCR will result. So high FCRs can result from both
over- and underfeeding - it is essential therefore that the farmer interprets FCR data
correctly.

Growth-ration interaction

High growth rates and high FCRs tend to indicate overfed fish, i.e. the ration fed is to
the right of the Rmax line in the diagram above. Thus overfeeding means delivering a
ration in excess of the fishes' appetite. It doesn't mean giving more than is good for
the fish in terms that some physiological problems may ensue. Fish have the ability to
regulate their appetite in relation to their requirements at any particular time.

Low growth rates and high FCRs could indicate underfed fish, i.e. the ration fed is to
the left of the R<SUBMAX< SUB> line. Underfeeding a stock has the potential for
the problem to be compounded. If a farmer considers that a low growth rate combined
with a high FCR is due to some environmental or biological problem causing the
former and resultant feed wastage causing the latter, he may be tmpted to reduce the
ration in order to correct the situation. Obviously this will compound the problem
causing growth rates to decrease and FCRs to increase further. This means that ration
is driven towards that required for maintenance alone.

Feeding Rate and Growth Efficiency

Studies indicate that, with fish, maximum growth efficiency does not necessarily
coincide with maximum food intake. Instead the growth-ration relationship shows
growth rates which initially increase with increasing ration but then plateau out at the
highest feeding rates (Figure A). In some quarters it is felt that the optimum ration
(Ropt), that is the point just prior to growth starting to plateau, is achieved if fish are
fed at restricted ration levels rather than to appetite.

Figures A + B showing Ration level and growth rate

This plateau may result from the differences in the ration delivered to the fish
and the actual ration consumed and indicate a decreased growth efficiency. The
situation may be closer to that shown in Figure B where the optimum ration for
growth and FCR is the same as the maximum ration - anything above this results
in wasted feed.

The appetite of fish is not constant, varying from feed to feed under the influence
of different factors such as the presence of predators, stormy weather, etc. Thus
the appetite at any one meal can be more or less than the preceding or
succeeding one. This meal -to- meal variation is compounded by longer term
seasonal variations in temperature, photoperiod and body size. In order to match
appetite to the food ration to be fed, there is therefore a requirement to have
some means of monitoring feeding by the fish in any particular cage. Such
systems include detecting feed falling through the bottom of the cage (sonar,
video) or mechanical retrieval systems which return uneated feed to the surface.
None of these is cheap and are only cost-effective in large-scale production units.
Smaller units will have to rely on visual surface operation which is not very
effective with regard to knowing when feeding has stopped. This implies that
feeding tables are not sufficiently accurate when feeding stock to appetite on a
meal-to-meal basis.

Feeding Regimes
Feeding efficiency is affected by the rate at which food is dispersed, the number of
feeds per day and the duration of each feed. These factors also play a part in
determining satiation. Analysis of the feeding process in terms of the number of
pellets delivered is one way of developing an optimal feeding regime. Based on fish
size and pellet diameter the number of pellets a fish requires for its daily ration can be
estimated and used to examine the influence of feeding techniques.

Number of feeds per day

Using the example of a 1kg fish fed at 1% of body weight per day (31
pellets/fish/day), some consequences of splitting the daily ration into a different
number of feeds can be seen above.

Feeding once a day would result in each fish obtaining its full ration (31 pellets) and
all the fish would be satiated. If fed twice a day, there would be 15.5 pellets per fish at
each meal. In this case, either 50% of the fish would be satiated at each meal or each
fish obtains only 50% of its daily requirement in the first feed and the other half at the
second meal. At the other extreme, feeding 31 times in a day means that there is one
pellet per fish at each meal - either 3% of the fish could be satiated at each meal or
each fish obtains around 3% of its daily need at each feed.

Increasing the number of meals thus reduces feeding opportunities through reduction
in feed availability. In addition many small meals will result in the formation of
dominance hierachies and increased competition leading large size/weight variations,
which the farmer wishes to avoid. The dominant fish will ensure that all the food
offered with many small meals will mean that the use of waste feed detection systems
is not feasible as there will be no waste feed at any of the meals. Similarly,
comparision of growth rates and FCRs will have no meaning.

Effect of increasing number of feeds per day.

Feeding varies within the industry from one to three meals per day for those using
hand feeding to automatic systems delivering hundreds of small quantities of feed
throughout the day. Either system can produce high growth rates and low FCRs as the
fish adapt to the preferred system, physiologically and behaviourally.

Depending on fish size, the maximum number of meals per day would be the one
which ensures at least one pellet per fish per meal. From numerous studies the
optimum number of meals appears to be in the range 1-6 per day to satiation, the
actual number depending on the species and water temperature. Except for first-
feeding fry, the number of meals does not relate to size as gut evacuation rate appears
to be independent of body weight.

With a few large meals stomach capacity could limit the amount of food that could be
eaten. However research indicates that salmonids maintain their stomach capacity
well below the maximum (0.1-0.25g per 100g body weight as opposed to 5g) in the
presence of unlimited food supply. Thus salmonids could consume more than the
highest daily ration they are likely to be given in a single meal. Appetite will increase
as the stomach empties so that too long a period between meals could lead to reduced
growth rates. Feed intake and gut evacuation rates are temperature-linked; as the latter
increases so do the former. Consequently more meals should be offered at higher
water temperatures - this is common industry practice as may be deduced from the
table below.

Number of feeds and water temperature

Water Temperature (oC) Feeds per day

0-5 1
5-10 2
10-15 3
> 15 4

Rate of feeding and meal duration

The rate at which feed is offered to fish and the length of each meal is related to the
number of pellets a fish will take per minute. This is a function of how hungry the fish
are and the number of available pellets at any one time. Studies show that feeding
rates are initially high but slow down as the fish attain satiation - as demonstrated by
the tailing off of the normal feeding response.

Cage and laboratory studies indicate that hungry fish consume around two pellets a
minute except at high (>18oC) and low(<5oC) water temperatures when rates of 0.5
pellets a minute are the norm. Knowing the number of pellets an individual fish will
eat for a given daily ration size and the number of feeds per day for a given
temperature , the number of pellets eaten per feed can be calculated. Dividing the
number of pellets per feed by either 2 or 0.5 (above) gives the duration of the meal
time. Consider the following examples:

1.

Fish weight:1kg
Temp: 4oC
Feed rate:0.2% per day
Meals/day: 1
Pellets/fish/day: 6-7
Pellets/fish/feed: 6-7
Feeding rate of fish: 0.5 pellets/min
Application rate/meal: 2.0kg in 12-14 minutes per tonne of fish

2.
Fish weight:1kg
Temp: 16oC
Feed rate:1.2% per day
Meals/day: 4
Pellets/fish/day: 46-47
Pellets/fish/feed: 11-12
Feeding rate of fish: 2 pellets/min
Application rate/meal: 3.75 kg in 5-6 minutes per tonne of fish

For different biomasses of fish the feed quantity will differ but the duration of the
meal will remain the same. Thus a cage holding 50 tonnes of 1 kg fish at 16oC will
require 187.5 kg of food to be delivered in 5-6 minutes. Reducing the number of
meals from 4 to 3 for the same cage would result in the need to deliver around 250 kg
of feed in 7-8 minutes. These rates may appear high to the husbandry person but from
the fishes' point of view, eating one pellet every 30 seconds, there is plenty of food
available and competition is much reduced. For delivery of these quantities at short
time intervals some high capacity dispensing system is required - it is not possible to
achieve these levels using a scoop. Cannons or automatic systems are necessary.

Timing of Meals
Salmonids readily adapt to operational feeding regimes. Obviously the best time to
feed is when the fish are hungry! If the time interval between meals varies
considerably the fish will adapt in terms of how much they will eat at each meal. The
time interval between satiation and return of appetite appears to be relatively constant
for any given temperature within a range of +-2oC. Naturally fish should not be fed
when environmental factors are limiting, e.g. when it is too dark.

Smolt feeding
It is frequently observed that smolts do not feed very well, following sea water
transfer. Studies suggest that several weeks elapse before 100% of survivors are
feeding post transfer, see diagram below. Thus if a farmer is feeding to tables and
only a certain percentage are feeding, he is going to waste a lot of food. If the daily
ration is split into a number of small meals the problem may not be too bad as only a
small percentage are feeding so these fish will obtain their daily feed requirement.
However as more come onto feeding there will be an increase in competition for
available food and hierachies will evolve. It is possible that 'failed smolts' or
'pinheads' may be socially disadvantaged fish rather than ones of poor quality or
perhaps diseased.

The differentials in time to feed post transfer may result in differing growth rates
within the stock and it can be advantageous to feed two feed sizes to one cage several
weeks after transfer. An example is shown in table at the foot of the screen.

Smolt feeding and post transfer

 Post-transfer feeding regime

Weeks after transfer % feeders Pellet size

1 10 100% as before transfer (A)
2-3 50 100% as before transfer (A)
3-6 75 50% A / 50% next size up (B)
7-8 100 25% A / 75% B
9 onwards -- 100% B

Feed Delivery

The best known and simplest form of feed delivery is through the use of the scoop.
This is perfectly adequate for small production units (up to 12 metres square or 50
metres diameter) and producing small tonnages (up to 100 tonnes maximum). Minor
increases in cage size and production output will force a change in technique - this
may simply be use of a larger scoop, that is, use a wheelbarrow adn shovel rather than
1 kg scoop and a feed bag!

Large-scale increases in cage size and/or production necessitate radical changes in

feed delivery by virtue of the amount of food potentially required and the optimum
delivery times as outlined in previously. One of the simplest ways to step up feed
delivery rates is to use a feed cannon. These consist of a hopper, which can hold up to
5 bags of feed at a time, connected to an archimedes screw delivery system through
which water is pumped, either from a stand alone pump or from the work boat. Pellets
drop from the hopper into the screw where water is introduced and the pellet/water
mix is forced out under pressure through a nozzle/cannon. The whole assembly can be
rotated so that spray can be angled up or down and from side to side allowing the
pellets to be spread over the cage surface. With sufficient water pressure feed can be
fired across a 100 metre diameter circular cage with ease. Obviously, as with scoops,
feed has to be delivered down wind for it to be effective.
Another jump up the production scale (>150 tonnes) will require use of almost totally
automated feeding with supplementary hand feeding. Feed wil be stored in huge silos,
either located onshore close to the site or on a floating barge adjacent to the site, from
which pipes lead to each cage within the system. Food can be delivered either in air or
in water. In the case of air delivery, the cage end of the pipe is out of the water so that
the pellets are sprayed over the surface; with a water delivery system the end of the
pipe can be in the water or sticking out of it. These systems are usually computer
controlled with manual override with alarms for low feed store levels, etc. with direct
information relay to feed programmes. Farmers will still have to sample weigh and so
forth to utilise these feed programmes effectively.

Monitoring of Feed Performance

Whatever feed a farmer utilises, there is a need to monitor how effective it is

achieving pre-set production targets/figures. Ways to do this include looking at the
growth rate of the fish on a particular diet or seeing how efficient the stock is at
converting food into growth. These are respectively referred to as the specific growth
rate (SGR) and the food conversion efficiency rate(FCR), which are discussed under
the next 2 subheadings.

Growth Rate

The rate at which fish grow is dependent on a number of factors including species,
age, genetic potential, water temperature, health, and quantity and quality of food.

The simplest modes for fish growth can be obtained by saying that all newly laid-
down tissue is itself capable of equal growth thereby producing an exponential growth
curve. However this only holds true if the percentage of body weight gained per unit
time remains constant throughout the life of the fish. This is not the case - young fish
are capable of doubling their weight in a much shorter time than when they are older
due to a decrease in potential growth rates.

It is therefore useful to be able to ascertain the rate at which fish are growing. The
best method of doing this is to calculate the specific growth rate (SGR or SPGR),
which is a measure of the percentage body weight increase per day. The SGR can be
calculated in the formula:

At first feeding, SGRs can be greater than 3 while fish over 1.0kg have average values
of 1. This is because smaller fish are capable of eating a much greater percentage of
their body weight per day. Temperature also has an effect on the SGR - the higher the
temperature, the higher the growth rate.

19. Growth rate calculation

Consider the following example:

Weight of fish on Day = grams
Weight of fish on Day = grams

Time between weighings = days

SGR = ((ln - ln )/ ) x 100 =

i.e this fish is increasing its body weight by % per day

Food Conversion Efficiency

The efficiency with which food is converted into fish weight is usually expressed as a
feed conversion ration(FCR):

Commercially available pelleted diets usually give FCRs between 1 and 2 depending
on fish size, temperature, feeding rate and feed composition. An FCR of 1 means that
every kg of food fed is converted to 1 kg of fish flesh; an FCR of 1.5 means that every
1.5 kg of feed will produce 1 kg of fish flesh.

It should be remembered that this is comparing a feed with a 10% moisture content
producing a wet weight gain in the fish which is about 70% water. On a dry weight
basis (dry feed to dry flesh), the FCR would be about 5:1, i.e 5 kg of food for 1 kg of
flesh.

Because of the factors which influence FCRs, feed manufacturers tend not to quote
them in their literature other than perhaps to give a general indication. In order to
calculate FCRs, several things need to be measured, calculated and understood. It also
requires the maintenance of good, accurate records whether they be paper or computer
generated. Without these, the calculations are impossible.

Accurate records of daily water temperatures, mortalities - so that number of fish in a

cage are known - and weight of fish need to be kept. One year's records can then ne
used to predict/ calculate the following year's food requirements based on that year's
smolt intake.

The next thing to understand is feeding charts and how to use them. They all give the
same basic information. Each manufacturere provides a chart detailing fish weight,
feed type and size and the feeding rate as a percentage of body weight/ day for
different water temperatures. There will obviously be differences in feeding rates
between feed companies for the same size fish and water temperature - but these are
relatively minor.
To calculate daily, weekly, or monthly feed requirements, FCRs, etc. the following
steps should be followed.

1. Sample weigh the fish in the cage or holding unit. The sample should be as
representative as possible and should not be based on a single fish - use at
least 10 (the more the merrier!).
Weigh the fish using an appropriate method e.g. place a water-filled bin/large
tank on a scale and note its weight. Place fish in receptacle - increase in weight
= weight of fish. If fish are placed directly on scale, the reading will be hard to
take and fish will suffer damage/stress.
2. Having kept accurate records of both the number of fish put into the cage at
the outset and the number of mortalities, the number of fish in the cage at the
time of weighing will be known. From (1) and (2), the total biomass (or
weight) of the cage can be calculated.
3. Take the water temperature (to the nearest 0.5oC).
4. From the feed tables, ascertain the feeding rate for the temperature and size of
fish.
5. Using the information from (1)-(4) the daily feed requirements can be
calculated. Multiplying by 7 gives the weekly feed requirement.
6. If the fish are weighed at a later date, e.g. 7 or 14 days on, the weight gain of
the fish can be measured. Knowing this and the amount of food fed in the time
period allows the FCR to be calculated.
7. Knowing the FCR allows future daily/weekly feed requirements to be
calculated. This allows a farmer to order his feed well in advance of its being
needed and so to avoid potential distribution problems.

The examples show how this is done in practice.

Being able to understand feed tables, calculate feeding rates and FCRs allows the
farmer to check how his stock is doing and to budget his money with regard to how
much feed is required over a certain length of time.

Introduction to Fish Feeding and the Environment

Aquaculture operations are influenced by surroundings, intrinsic environmental
factors whilst simultaneously impacting on the environment. Factors such as water
temperature and oxygen levels influence fish feeding and digestion (and ultimately
growth) whilst fish feed, faecal material and chemicals impact on the surrounding
environment. It is the interaction between these two sides of the coin which ultimately
determine how well stock will do in any particular aquatic environment.

Temperature and Oxygen

Fish are poikliothermic or cold-blooded, that is they cannot generate their own
internal body heat but have to derive it from the surrounding environment. As the
temperature of the surrounding aquatic environment is not constant due to the latent
heat capacity of water, so most fish do not have stable internal temperatures. Fish tend
to have optimal temperature ranges within which they can function normally, and
outside this range things can and do go wrong.

Within any range, increases and decreases in external temperature have an effect on
internal metabolic and physiological rates. The warmer, the faster: these go up to the
point where they enter overdrive and collapse. Conversely, when temperatures are too
low, processes slow right down adn may be stopped.

With respect to feeding, the process of digesting food results in the release of heat
irrespective of whether the animal if poikliothermic or homeothermic. In
homeotherms, the levels of insulation in body tissues only allow slow release of this
heat to the outside environment. In aquatic poikliotherms this heat can be readily lost
to the water. As a consequence it is possible for fish to gorge themselves to the point
of the stomach and throat being completely full of food. Mammals, etc. cannot do this
as the buildup of heat becomes uncomfortable and the animal will stop feeding long
before the stomach alone is full.
Environmental oxygen levels and the impact on feeding are discussed in the
metabolism of food section.

Environmental Impacts of Feeding

Finfish (and shellfish) farming operations, can have noticable impacts on both water
quality and the surrounding substrates, with resulting consequences for the water
column. Waste feed, phosphorus and nitrogen waste also impact on the environment.

Solid wastes, from waste feed etc., as well as soluble wastes such as ammonia and
dissolved oxygen have severe consequences for the surrounding substrates.

(a) Changes to the water column by fish farming:

1. the addition of waste products, principally nutrients and suspended solids, into
the water column;
2. the reduction of ambient dissolved oxygen levels
3. disruption caused by cage structures;
4. impediment to water flow

(b) Changes to the water column by shellfish farming:

1. the accumulation of the solid waste material from the farming operation;
2. the influence of fish farming structures on "natural water currents";
3. the use of toxic chemical treatments.

Impacts on the water column - solid and soluble

wastes
Impacts on the water column are influenced by the nature and amount of wastes
produced, and the characteristics of the recipient environment.

Wastes from fish culture can be subdivided into two major fractions, solid and soluble
waste.The solid fraction is composed of uneaten food pellets and faecal particles,
together with smaller quantities of fish scales, mucus and other detritus, most of
which have a density greater than water and will tend to sink out of the water column.
The soluble fraction consists of materials excreted in urine or across the gills and
leached from the solid fraction as it sinks. The most significant characteristics of these
wastes as they affect water quality and sediments are the suspended solids, the various
nitrogen and phosphorus compounds and the oxygen-consuming properties.

Waste feed
The actual quantity of waste feed produced varies considerably, depending on feed
type, feeding method, feed quantity and other environmental and physiological
factors, the effects of which are largely undetermined. Feed type is important, for
example, estimates from freshwater trout culture suggest a wastage of 1-5% for dry
feed, 5-10% for moist feed and 10-30% for wet feed (trash fish). Common estimates
of feed wastage, including dust and uneaten food, from salmonid farms using dry
feeds vary from 5% to 20%. Fish faeces are the second source of solid waste and
estimates from laboratory studies suggest that 25-30% of the dry weight of consumed
feed is voided as faeces.

Phosphorus and nitrogen wastes

There are major differences between the fate of phosphorus and nitrogen wastes in the
environment. Phosphorus wastes are concentrated within the solid fraction, whereas
the vast majority of nitrogenous wastes are found as dissolved nitrogen, particularly
ammonia, which is excreted through the gills as an end product of protein
metabolism. The waste loadings of dissolved nitrogen are particularly important
because nitrogen is generally regarded as the limiting nutrient within marine waters,
i.e., the nutrient which controls the growth of primary producers. Phosphorus is
important as a limiting nutrient in freshwater, and is not regarded as critical in some
northern coastal waters. The limited information on the waste loadings from marine
salmonid culture is given in the table below.

S. salar1 80 kg dissolved nitrogen

43 kg particulate nitrogen
2
S. salar 47.9 kg total ammonia-nitrogen
0.99 kg nitrite-nitrogen
13.1 kg nitrate-nitrogen
61.9 kg inorganic-nitrogen
1
theoretical calculations
2
data from several studies on a commercial marine tank farm in Scotland growing
Atlantic Salmon (hand feeding, commercial dry pellet, food conversion ratio 1.5:1)
(units are kg/tonne fish production per year)

Impacts of ammonia on the environment

Ammonia is an important waste product because it can be directly toxic to aquatic

organisms. Ammonia exists in water in two forms, unionised ammonia (NH3) and the
ammonium ion (NH4+); the relative proportion of the total ammonia pool in these
different fractions depends on pH, temperature and salinity. Unionised ammonia is
generally regarded as being most toxic to fish and other aquatic animals and in
Scottish seawater a maximum of 2.5% of the total ammonia will be found in this toxic
form. It is useful, therefore, to assess whether the concentrations of unionised
ammonia found in and around cage farms are likely to be toxic to marine life. The
maximum "safe" level of unionised ammonia is 0.025 mg/l, which is equivalent to
around 1.0 mg/l of total ammonia under the worst case Scottish conditions (high
temperature). The concentration is well above any of the recorded concentrations
within and outside marine salmonid cages. Direct ammonia toxicity to both farmed
and wild fish populations, therefore, is considered extremely unlikely at the current or
future predicted cage farming levels.

The oxygen content of water will be reduced by the fish themselves and by bacteria
which decompose farm wastes (food, faeces). This may cause a reduction in oxygen
content on a very localized basis. The amount of oxygen used in any particular
situation will depend on fish size, activity, age, physiological state, feeding regimes
and external environmental parameters.

Impacts of dissolved oxygen

Measurements of dissolved oxygen within salmon cages commonly show
concentrations of between 0 and 2.0 mg/l less than the background level, depending
on tidal flow, stocking density and the degree of fouling. Adjacent to cages, changes
in dissolved oxygen content have been either undetectable or very small. The surface
waters within most Scottish sea lochs are normally close to saturation thus minimum
concentrations of around 7.7 mg.l are predictable in worst case conditions of high
temeratures (18oC) and full strength seawater (34ppt). Dissolved oxygen
concentrations of between 5 and 6 mg/l are considered to be the minimum acceptable
concentrations for marine animals and most salmon cages are stocked at a level which
will keep dissolved oxygen within the cages above 6-7 mg/l. Thus traditional stocking
policies should not cause significant oxygen depletion of saturated surface waters
outside of cage farms. If there is a deterioration in ambient concentrations as a result
of phytoplankton blooms or accumulation of oxygen consuming sediments in shallow
waters, then there may be more problems, although such effects will probably be most
significant for the caged stock.

In the absence of oxygen, hydrogen sulphide, ammoni and methane are generated
within the sediments and may be released to the water column. Hydrogen sulphide is
highly toxic to fish, slightly soluble in water and has been implicated in gill damage
and mortalities. Methane is highly toxic to fish. As yet these two types of self- or
auto- pollution have not seriously affected fish farms - those located in shallow water
with a poor tidal exchange will be most at risk.

Impacts of solid wastes on the environment

The fate of solid wastes from fish farms depends on their settling velocity and the
current velocity, turbulence and depth at individual sites. The settling velocity of
marine salmonid feed pellets is greater than faecal particles because of their higher
density. Consequently there is a tendency for faecal waste to spread over a wider area
than feed pellets, although most waste solids tend to accumulate in the immediate
vicinity of the cage site. The amount of solid accumulating below cages is very
variable, from up to 40cm of soft, light brown flocculent material in low velocity sites
to no visible accumulation on high velocity sites.

The solids falling to the seabed below fish cages are enriched in carbon, nitrogen and
phosphorus relative to the natural sediments, hence fish farming may alter the
physico-chemical nature of sediments below and adjacent to the operation but is
usually limited to the vicinity (50 m) of the cages. This increase in carbon
sedimentation results in an increase in oxygen consumption by bottom-living animals.
The sediments will become anoxic (i.e.contain no oxygen) if this additional oxygen
demand exceeds oxygen supply, at which point there may be severe consequences for
both benthic organisms and the fish farming operation itself.

Changes in Water Circulation

The presence of fish cages, like any other structure placed in a moving fluid, affects
the velocity and direction of water movement. The magnitude of the effect depends on
the size and mechanical properties of the structure, which for aquaculture equipment
is related to the design characteristics and the degree of fouling. The velocity drop
within fish cages varies from 35-81% of the external current velocity, depending on
stocking density, net mesh size and structure and the growth of fouling organisms.

Local changes in water circulation are, therefore, expected, although predicting the
magnitude and extent of effects is impossible because of the extremely variable
hydrographic regimes around the coast. Possible water quality impacts could include
preventing the dispersal of metabolic wastes, reducing oxygen supply and affecting
the distribution of planktonic organisms. These potential effects have their most
severe impact on the farmed stock. Given maintenance of good conditions for the
farmed stock and selection of sites with good water exchange, the impacts of water
quality changes due to poor water circulation per se are considered minimal. The
introduction of cages, moorings and other structures also offers new substrates for the
attachment of organisms which may affect water quality through respiration or
excretion or alter the structure of local food chains.

Nutrient loading

As yet there is little evidence to suggest that the increased nutrient loading of the
water column from aquaculture operations directly causes algal blooms - however
problems may ultimately arise in enclosed, slow-flushing waters. Heavily polluted
farms may cause alterations to the bottom-living plants and animals in the immediate
vicinity of cages. As the oxygen levels decrease, mobile animals will move out - those
that can't will die (i.e. plants, shellfish). These are replaced by bacteria (particularly
one called Beggiatoa which covers the sea-bed in a white film) - if these disappear,
the oxygen is completely used up and hydrogen sulphide and methane may start to
bubble out of the sediments.

Pollution
Aquaculture is also a source of chemical pollution through materials incorporated in
fish feed, chemicals and pharmaceuticals in the treatment of diseases and treatments
applied to the equipment which can leach into the water. The first-mentioned does not
yet appear to have any effects on the environment although vitamins in food may have
minor effects on a very local basis. Recently, problems over the use of canthaxanthin
as a pigment have been raised with regard to its carcinogenic effects on humans. This
has now been replaced by astaxanthin, a naturally occurring pigment.

Fish farming operations have the potential to be major polluters. However to do so

would be self-defeating as the first thing to suffer would be the farmers' own stock.
Thus, to ensure a healthy and profitable stock, a farmer must exercise good
husbandry. This in turn will minimise any environmental pollution.

Introduction to Feed Selection

As most farmers know, feed costs constitute a very large proportion of the overall
production costs, typically in the range of 30-60%. Increasing the level of production
technology has the added benefit of reducing feed costs and manpower costs.
Anything which reduces feed costs is going to benefit the farmer.

A number of inter-related issues need consideration with respect to feed management

and cost reductions. On the consumer side there is a need to consider such things as
food safety, health aspects and the product's nutritional value. Producers need to
evaluate such things as food and stock performance and availability, stock health,
environmental impacts and legislation, and the implicit costs of these. Despite these
potential hurdles, there is opportunity to reduce feed costs through feed selection,
reduction of wastage and use of appropriate ration sizes - all these influence the cost-
effectiveness of a diet.

Feed selection
Farmers usually select a diet on the basis of cost and perceived quality with additional
inputs from extras such as feed company backup, computer feed programmes and so
forth. Very little detail exists which compares one diet with another, either within one
company or between companies. The only comparison a farmer can make between
diets is historical - that is he can compare a new one with one he used previously by
analysis of his farm records (growth rates, FCRs, etc.)

When a new diet becomes available, the benefits from it must be obvious in terms of
improved growth, FCRs or product quality. If a new diet is more expensive than
existing ones, there must be an accompanying decrease in the FCR if feed cost per
unit of fish produced is not to go up. This is illustrated in the diagram on the screen
which also shows that small changes in FCR can have a large impact on feed costs.

Effect of changes in price and FCR on fish production costs

Comparison can also be made between diets where potential FCRs and price are
known.
For example, a new diet (ND) promises an FCR of 1.0 but at a price of 1100 pounds
per tonne. How does this compare to an old diet (OD) at 950 pounds per tonnes giving
an FCR of 1.2?

Diet ND has a feed cost per tonne of fish produced of 1100 x 1.0 = 1100 pounds,
whilst cost of diet OD is 950 x 1.2 = 1140 pounds. Thus the newer, more expensive
diet is actually cheaper in terms of production costs.

Feed wastage

Quite naturally, any reduction in the wastage of feed will have a significant effect on
costs. It is essential that overfeeding is avoided and that feeding practices do not
become lax. Continual monitoring and review of feed amounts and feeding regimes is
necessary. Use of performance indicators (growth rates, FCRs) both currently and
historically is recommended so that any problems can be picked up at the earliest
opportunity.

For large operators there may be financial benefit (long term) to be gained from the
installation of some form of feed retrieval system. Any food retrieved can be re-used
in addition to the visual clues indicating what is happening to the stock. Fully
automated systems are prone to overfeeding - there is a need either for a feed retrieval
system or for visual feedback, perhaps by supplementary hand feeding, to prevent
this. Delivery amounts in automatic systems need constant re-appraisal.

Cost Effective Rations

It is possible to calculate the cost of producing fish from feed on its own (using feed
costs and FCR values. However when feed cocts are analysed against the backdrop of
farm/site production, other factors come into play. As a consequence of fixed holding
unit volumes and/or water exchange rates, the production capacity from a farm is
reasonably well fixed also. Through hard work it may be possible for an operator to
increase his production therby reducing the operating cost-revenue ratio.

Rations of any particular diet must be such as to allow production targets to be met in
a cost-effective manner. Appetite or satiation feeding may optimise growth and
holding unti capacity but at the expense of feed wastage. Conversely restricted
feeding may reduce wastage but at the expense of site production potential. The
farmer thus has to select an appropriate feeding strategy for each operation being run.

Summary

As with all industries it is possible to waste money through a combination of a lack of

thought, knowledge and lax practices.

Given thought, knowledge and good attention to husbandry practices it is possible to

utilise fish diets extremely cost-effectively. As they form one of the largest portions of
the production costs-pie, this becomes a necessity. To maximise this cost-
effectiveness, a farmer has to ensure that he uses the best diet in such a way that
growth and feed conversion are maximised and wastage is minimised within the
physiological limits of the stock. This necessitates constant monitoring and continual
re-evaluation of what is happening. There is no room for complacency.

There is still room for improvements in fish feed and its formulations. Some of the
waste in feed (dust, etc.) can be removed, unit feed production costs can perhaps be
reduced through alternative ingredients and so on. Through these methods it may be
possible to reduce FCR values below 1.0, perhaps down to 0.9-0.95. At these levels
feed becomes very cost-effective, particularly in the light of recent low prices for
some aquaculture products. It is felt there is a biological FCR limit which has a value
of 0.85 but it is unlikely to be achievable in practice.

References
Alsted, N (1991). Better rations for better fish. Fish Farmer, 14(5): 48-49

Christiansen, J.S, Jobling, M. and Amin, A.B. (1994). Growth associated

degenerations in the swimming muscle of a salmonid fish. Norsk Fiskeoppdreft.

Cowey, C.B. (1991). Some effects of nutrition on flesh qualitity of cultured fish. Fish
Nutrition in Practise. Biarritz (France) June 24-27, 1991: 227-236.

DeSilva, S.S. and Anderson, T.A. (1995). Fish Nutrition in Aquaculture. Chapman
and Hall

Ecoline (1994). New diets put emphasis on opyimum feeding levels. Fish Farmer,
17(1): 48-49

Hillestad, M and Johnsen, F. (1994). High-energy/low-protein diets for Atlantic

salmon:effects on growth, nutrient retention and slaughter quality. Aquaculture,
124(1/4): 109-116

Holmes, M. (1995). Comparision of low and high oil content diets on the growth rate
and fat levels in the flesh of the atlantic salmon (Salmo salar). NAFC Fisheries
Development Note No. 3

Howgate, P., Mackie, P.R., Whittle, K.J., Farmer, J., McIntyre, A.D. and
Eleftheriou, A. (1977). Petroleum tainting in fish. Rapp. P:-v. Reun. Cons. Cons. int.
Explor. Mer., 171:143-146.

Jobling, M. (1994). Fish Bioenergetics. Chapman and Hall

Lavety, J. (1993). Soft salmon. Torry Research Station Newsletter 24:5.

Luzzana, U., Serrini, G., Moretti, V.M., Gianesini, C. and Valfre, F. (1994).
Effect of expanded feed with high fish oil content on growth and fatty acid
composition of rainbow trout. Aquaculture International 2, 239-248.

Mackie, P.R., McGill, A.S. and Hardy, R. (1972). Diesel oil contamination of
brown trout (salmo trutta L.). Environ. Pollut. (3):9-16
Sinnott, R. (1995). Pers. comm.

Springate, J. (1991). Extruded diets - worth the extra. Fish Farmer, 14(2):45

Springate, J. (1994). Watch oil and protein levels. Fish Farmer, 17(1):47

Springate, J. and McKinney, R. (1994). High energy diets. Scottish Fish Farmer,
Dec. 1994:16

Talbot, C. (1994). How growth relates to ration size. Fish Farmer, 17(1): 45-46

Talbot, C. (1995). Fish Feeding and flesh quality. Outlook No. 3: 4-5

Talbot, C. (1997). Feed management for salmon in sea cages. Outlook No. 8: 10-11

Thomassen, M.S. and Rosjo, C. (1989). Different fats in feed for salmon: influence
on sensory parameters, growth rate and fatty acids in muscle and heart. Aquaculture,
79: 129-135.

Waagbo, R., Sandnes, K., Torissen, O.J., Sandvin, A. and Lie, O. (1993).
Chemical and sensory evaluation of fillets from Atlantic salmon (Salmo salar) fed
three levels of n-3 polyunsaturated fatty acids at two levels of vitamin E. Food
Chemistry, 46:361-366.

Watanabe, T. (1982). Lipid nutrition in fish. Comp. Biochem. Physiol. 73B/1;3-15

Energetics Overview

by James F. Kitchell

The field of fish bioenergetics includes temporal scales that range from those
of evolutionary time to cellular metabolism (Tytler and Calow 1985). It also
includes spatial scales ranging from nutrition and growth in controlled
aquaculture systems (Jobling 1994) to predator-prey systems in the largest
ecological context (Adams and Breck 1990). Among the several reviews of
the field, those by J. R. Brett offer the most insightful combination of basic
laboratory studies and their application in the context most pertinent to
fisheries science (Brett and Groves 1979). We recommend Brett’s lead
chapter (Brett 1995) in the new volume edited by Groot et al. (1995) for a
thorough review of the extensive work conducted on energetics of Pacific
salmonids and for an insightful assessment of areas where knowledge of
energetics should be improved.

The underpinnings of energetics have a firm theoretical base in the laws of

thermodynamics (Kleiber 1975). Working from an energy budget requires that
you satisfy the terms of a simple equation; outputs must equal inputs and the
budget must balance. As detailed in Chapter 2, the terms of the energy
budget for fishes are well known and each can be measured independently.
The model allows the user to specify the important external regulators:
temperature and diet. For fishes, the most easily measured component of the
energy budgeting process is expressed as growth. Growth integrates the
array of environmental variables affecting an individual fish. Thus the
evidence provided in the observed growth rate is the rich and varied
foundation of scientific inquiry and the basis for better understanding.

The modeling approach presented in this manual derived from the extension
of energetics principles used in ecosystem-scale models of trophic
interactions developed during the International Biological Programme
(Kitchell et al. 1974). These models focused on biomass dynamics. They
often included formulations requiring an estimate of carrying capacity which
was used to characterize density-dependent constraints for growth rates of a
given trophic level. While those kinds of models have utility in an ecosystem
context, they had three important shortcomings when applied to fishes. First,
units of biomass per area or volume did not allow for resolution of cause and
effect at the species or individual scale. After all, it is individual fish that feed,
grow, reproduce and die. Further, as a fish grows from first-feeding larvae to
reproductive adult, it may ascend through three or four trophic levels.
Second, biomass models did not allow an effective interface with either the
long history of population-based models in fisheries science or the models of
predator-prey interactions developed in the ecological sciences. Third,
biomass models required an estimate of environmental carrying capacity.
The latter is difficult to do and, more importantly, likely to change as a
consequence of the ecological effects due to fishery exploitation and/or
anthropogenic effects on fish habitats.

An alternative to biomass models is an energetics-based approach focused

on the processes that regulate growth by individual fish (Kitchell et al. 1977).
This model assembled individuals in age- or size-based populations,
 separated the agents of mortality (natural vs. fishing) and specified the
trophic ontogeny of predator-prey interactions. It focused on using the kinds
of data most frequently collected by biologists  the habitat that is occupied
(thermal history), size at age (growth curves), stomach contents, size or age
at sexual maturity, and size- or age-related mortality rates. The development
of size- or age-based cohorts is elaborated in Chapter 3.

1.1 The Modeling Strategy

Starting the process with observed growth rate is different from that of many
kinds of modeling practices. In this case, the strategy of model building is
based on specifying rules that define the limit conditions, i.e., the maximum
and minimum possible rates of growth for members of a population. The
physiological parameters used to represent the rules derive from readily and
oft-measured processes such as temperature dependence, thermal
tolerance, thermal preference, size dependence, assimilation efficiency, etc.,
that can be accurately measured in the laboratory. Those physiological
parameters are assembled as empirical rules that define the effect of
temperature, body size and food quality on maximum feeding rates. The
minimum is similarly defined by rules describing the effect of temperature and
body size on metabolic rates when food consumption is set to zero. These
limits define the boundaries of the scope for growth. Observed growth is
somewhere between those limits and allows the user to estimate how that
growth rate is being regulated.

The hierarchy of energy allocation is an important component of this

modeling approach. Consumed energy is first allocated to catabolic
processes (maintenance and activity metabolism), then to waste losses
(feces, urine and specific dynamic action) and that left over is allocated to
somatic storage (body growth and gonad development). This hierarchy is
analogous to practical economics. The first costs paid are those for rent or
mortgage (metabolism) that sustain the organism. The second set of costs
(waste losses) are like taxes  they are proportional to income (food
consumption) and must be paid. The energy resource remaining may then be
allocated to savings (growth) or invested in the next generation (gonad
development). In an ecological or evolutionary context, it is easy to imagine
selection for behaviors that maximize benefits (growth rate or gonad
development) and minimize costs. Like an account balance, a record of
growth reveals how well the organism has resolved the complexities of its
environment.

In a thorough review of previous energetics work, Brett and Groves (1979)

presented a generalization about energy budgets for two classes of fishes. If
the energy budget is stated in the following terms:

Energy Consumed = Respiration + Waste + Growth,

and normalized to percentages when energy consumption = 100, then fishes

growing at "typical" rates would have energy budgets approximated as below.

Consumption = Respiration + Waste + Growth

For carnivores: 100 = 44 + 27 + 29

For herbivores: 100 = 37 + 43 + 20

These budgets reveal two important features. First, as expected, herbivores

exhibit lower growth rates and higher waste-loss rates per unit of energy
consumed. That is the logical consequence of eating foods of lower energy
density and higher indigestible content. Second, both types of fishes
demonstrate high rates of growth efficiency compared to those known for
mammals and birds. Although these budgets can serve as a first
approximation, the 95% confidence intervals for each component are
substantial (e.g., plus or minus 20% of the mean). Of course, the energy
budget for an average fish in a typical habitat may be very different from that
of fishes in some unique ecological context. Fishes are known to exhibit
among the highest growth efficiencies recorded (approaching 50%) and are
known to exhibit strikingly negative energy budgets, as in the case of
migrating salmon (Brett 1995). Note, too, that the hierarchy of energy
allocation operates in all cases. Growth efficiency is not a constant, and
growth rates in fishes are highly variable. Observed growth is the integrated
answer to a complex question about prey resources and environmental
conditions. Deducing the quantitative components of cause and effect is the
significant challenge.

In most of its applications, model users will seek an answer to questions

about factors that constrain growth (e.g., diet quality or environmental
stressors) or use the measured growth to estimate how much effect a
predator has had on its prey populations. Assembled as a population, the
model allows answers to those questions at the larger scales of ecological
and management interest. This approach does not provide for feedback to
future generations. Predator or prey population dynamics are not
represented. Those must be characterized as simulations using specified
assumptions about prey availability, mortality rates and environmental
conditions.

We view the modeling process as having two general components. First is

the "nuts-and-bolts" process of assembling the parameter tables and the
input data. Much of the former is available in the manual or formatted in ways
that welcome site-specific input. Second is the "arts-and-crafts" process of
structuring analyses in ways that pose key questions and provide instructive
answers. In these cases, it is often valuable to use the model as a way to
create boundary conditions such as those for maximum possible growth or
for maintenance requirements. Using the model in this way allows it to serve
as a "deductive engine" in the more creative and challenging process of
science (Walters 1986).