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Chapter 13. Diet and Fish Husbandry
Richard T. Lovell
Nutrient requirements for normal growth and function of the major cultured ﬁsh species are essentially known and the amounts required for optimum performance under standard conditions have been fairly well identiﬁed. However, optimum methods of formulating, processing, and feeding
diets for a diverse group of commercial aquaculture species, in a variety
of culture environments, and for different production functions present a
myriad of challenges to the practicing ﬁsh nutritionist.
This chapter deals with feeding strategies for several important commercial aquaculture species with diverse culture environments and technologies. Feed costs account for over 50% of the variable costs in most
aquaculture operations, therefore applying the best feeding strategy can
have a signiﬁcant impact on optimizing proﬁt, which is the primary goal
of commercial aquaculture. Since commercial aquaculture and livestock industries have similar goals, to provide a high-quality consumer product at
a proﬁt, it may be well to compare feeding ﬁsh with land animals. Subsequently, different levels of ﬁsh farming are examined and then applicable
feeding practices are discussed.
13.1.1. Levels of Aquaculture
Several authors have recommended different systems of classiﬁcation of
the various stages of aquaculture. Some classify according to the level of
intervention. i.e., how much the culture environment has been modiﬁed.
This ranges from the impoundment of natural waters and harvest of any
and all animals therein, without adding seed stock or nutrients, to the use
of a closed system in which water is recirculated. Some classify on the basis
of the quantity and quality of nutrients utilized by aquaculturists, such as
extensive, where no nutrients are added; fertilization, to enhance production of aquatic organisms; supplemental feeding, using incomplete feeds;
intensive feeding using nutritionally balanced feeds; and hyperintensive
feeding, where high inputs of concentrated, nutritionally complete feeds
are used. Some classify on the basis of the energy input (labor, fossil fuel,
feed) or technology input (harvesting, stocking, feeding, pumping, aeration, bioﬁltration). Generally, the higher the level of intervention in the
production of aquatic animals, the more important is the feed to the success of the operation.
220.127.116.11. Production of Fish Exclusively from Natural Aquatic Foods
Some ﬁsh obtain their food exclusively from plankton. These ﬁsh are usually continuous grazers and have mechanisms for ﬁltering or concentrating
13. Diet and Fish Husbandry
Tilapia grown in combination with ducks. The pond receives only manure from the
ducks. The ﬁsh yield is approximately 1200 kg/ha annually.
the suspended animal and plant organisms from the water. An example is
the silver carp. Others, such as some of the tilapias, have the ability to feed
on plankton but also feed on bottom materials. The common carp is an
efﬁcient bottom feeder. Some ﬁsh, such as grass carp, have herbivorous appetites and consume large quantities of higher aquatic plants. Such ﬁsh can
be cultured without artiﬁcial feeds, as shown in Fig. 13.1, but usually with
pond fertilization. This level of production is most applicable in environments where supplemental feeds are expensive or unavailable.
18.104.22.168. Supplementation of Natural Foods with Feed
This level of ﬁsh farming essentially involves taking full advantage of natural aquatic productivity and using various feedstuffs or prepared feeds
as a supplement to increase the yield further. An example is semiintensive
shrimp culture in Central and South America, where large ponds are fertilized to enhance natural productivity and also receive pelleted feeds. Usually
with species that will accept supplemental feeds, the additional yield of ﬁsh
Richard T. Lovell
resulting from the additional feeding is proﬁtable. For example, the yield
of common carp in fertilized ponds was 390 kg ha−1 , the addition of grain
or grain by-products increased the yield to 1530 kg ha−1 , and formulated
feed further improved the yield to 3000 kg ha−1 (Lovell et al., 1978).
Where natural aquatic food may make a relatively small contribution to
the total protein and energy requirements of the cultured ﬁsh, it can provide
essential micronutrients that will allow nutritionally incomplete supplemental feeds to be used. As the biomass of ﬁsh in the pond increases, however,
the ﬁsh will become more dependent on the supplemental feed for all nutrients. Channel catﬁsh grown in earthen ponds to maximum standing crops of
2000 kg ha−1 grew normally and showed no deﬁciency signs when vitamin C
was deleted from their feed. However, when the ﬁsh density was increased
to 4000 kg ha−1 and above, growth was normal but resistance to bacterial
infection was reduced and subclinical deﬁciency signs occurred when no
vitamin C was fed (Lovell and Lim, 1978).
22.214.171.124. Intensive Culture of Fish in Highly Modiﬁed Environments
With these systems, natural foods are an insigniﬁcant source of nutrients.
Maximum yield per unit of space and effort and minimum accumulation
of unretained nutrients in the culture system are primary concerns. Thus,
highly concentrated, nutritionally complete feeds are justiﬁed. Examples of
this type of production are rainbow trout cultured in spring-fed raceways
and Atlantic salmon grown in net pens in the coastal areas of the sea. Also,
channel catﬁsh or marine shrimp in intensively stocked ponds may obtain
negligible amounts of nutrients from natural foods.
13.1.2. Feeding Fish versus Feeding Land Animals
Feeding ﬁsh in their aqueous environment takes on dimensions beyond
those considered in feeding warmblooded food animals. These include
the nutrient contribution of natural aquatic organisms in pond cultures,
the effects of feeding on water quality, and the interaction between feed
allowance and optimum dietary nutrient concentration. Because ﬁsh cannot be fed ad libitum, the feeder, not the ﬁsh, decides how much feed is fed,
and thus a higher level of management is required to feed ﬁsh. However,
the concept of feeding is the same as that applied in feeding other food
animals; to nourish the animal to the desired level or form of productivity
as proﬁtably as possible. Thus, application of knowledge of the nutritional
requirements of ﬁsh and the husbandry of feeding various cultured species
is essential to successful aquaculture.
Unlike intensively reared livestock and poultry, which are fed ad libitum,
ﬁsh are given a restricted feed allowance that will minimize waste. Because
13. Diet and Fish Husbandry
feedlot farm animals eat as much and as often as they want, their nutrient
allowances are based upon satiation feeding. Fish are often (perhaps usually) not fed to satiation and the daily feed allowance has been shown to
affect the ﬁsh’s response to various dietary nutrient concentrations. Li and
Lovell (1992) found that the optimum dietary protein allowance for channel
catﬁsh fed to satiety was 26%, while ﬁsh fed to less than satiation responded
to higher protein concentrations. For example, a study at the Tunison Fish
Nutrition Laboratory in Cortland, New York (Rumsey, 1993), showed that
the arginine requirement for maximum growth for young rainbow trout
was signiﬁcantly lower when the ﬁsh were fed to satiation than when the
ﬁsh were given a restricted feed allowance. This interaction between feed
allowance and optimum dietary nutrient concentration makes the formulation of commercial feeds more difﬁcult for ﬁsh than for farm animals.
Because ﬁsh are fed in water, feed that is not consumed within a reasonable time not only represents an economic loss, but can reduce the water
quality. Therefore, feed allowance, feeding method, and water stability of
the feed are factors that the ﬁsh culturist must consider but the livestock
feeder does not. The culture environment may make valuable nutrient contributions to the ﬁsh. For example, most waters contain enough dissolved
calcium to provide most of the ﬁsh’s requirement. For ﬁsh that feed low on
the food chain, such as shrimp and some tilapias, the pond environment
can be a valuable source of protein, energy, and other nutrients.
Fish convert practical feeds into body tissue more efﬁciently than do
farm animals. Cultured catﬁsh can gain approximately 0.84 g of weight
per g of practical diet, whereas chickens, the most efﬁcient warmblooded
food animal, gain about 0.48 g of weight per g of diet (Table 13.1). The
Efﬁciency Utilization of Feed and Dietary Protein and Metabolizable Energy (ME)
by Fish, Chicken, and Cattlea
Weight gain Protein gain ME required
ME–protein per g of food per g of protein
per g of
(%) (kcal/g) (kcal/g)
From Lovell (1989).
Richard T. Lovell
reason for the superior food conversion efﬁciency of ﬁsh is that they can
economically assimilate diets with higher percentages of protein, apparently
because of their lower dietary energy requirement. Fish have a lower energy
requirement than terrestrial animals because of their lower maintenance
requirement and lower heat increment. Fish, however, do not hold an advantage over monogastric farm animals in protein conversion; as shown in
Table 13.1, poultry convert dietary protein to body protein at nearly the
same rate as ﬁsh. The primary advantage of ﬁsh over land animals is the
lower energy cost of protein gain rather than the superior food conversion
efﬁciency. The metabolizable energy requirement per g of protein gain is
21 for channel catﬁsh versus 23 for the broiler chicken.
13.2.1. The Industry
Culture of channel catﬁsh (Fig. 13.2) accounts for about two-thirds of the
commercial aquacultural production in the United States. The production
of farm-raised catﬁsh reached 225,000 tons in 1998 (USDA, 1999). Once
considered to have primarily a regional appeal as a food, farm-raised catﬁsh
are now in national and international markets. Catﬁsh reach the processing
plant alive and are kept alive until they are slaughtered, which takes less
than 30 min. Farm-raised catﬁsh are fed grain-based feeds, which give the
ﬁsh a mild ﬂavor, with the absence of a “ﬁshy” odor. The ﬂesh is mostly white
muscle, which is free of intramuscular bones. Nutritionally, an 8.4-g serving
of farm-raised catﬁsh contains approximately 140 kcal, 17 g of protein, 9 g
of fat, 50 mg of cholesterol, 40 mg of sodium, and a number of essential
vitamins and minerals. The delicate ﬂavor, light ﬂesh, high nutritional value,
and year-round availability make farm-raised catﬁsh an appealing choice to
the food service industry and to consumers.
Channel catﬁsh possess several qualities that make it amenable to culture.
The ﬁsh normally does not reproduce in culture ponds, it is easy to spawn
under hatchery conditions, and it produces a large number of fry, readily accepts a variety of prepared feeds, and tolerates water temperatures from near
freezing to 34◦ C and wide ﬂuctuations in water quality in production ponds.
Channel catﬁsh grow rapidly; a 10-g ﬁngerling reaches a harvestable size of
0.5 kg in about 6 months as long as the water temperature remains above
23◦ C. Also, it converts feed efﬁciently; feed conversion ratios (feed/grain)
of 1.4–1.5 can be achieved.
In the early 1970s, when catﬁsh farming was in its infancy, farmers stocked
earthen ponds at rather low densities, ranging from 2500 to 5000 ﬁngerlings
13. Diet and Fish Husbandry
Channel catﬁsh weighing approximately 0.5 kg harvested from a 5-ha pond that
constrained approximately 7500 kg of ﬁsh per hectare.
per hectare, in the spring and harvested the ﬁsh in the fall. The ﬁsh were fed
a pelleted, concentrated feed, and yields of 1000 to 2000 kg per hectare were
typical. Today, yields range from 4000 to 7000 kg per hectare. The increased
yields can be attributed to higher stocking densities and to improvements in
feeds, feeding practices, water quality management, and disease control. In
addition, a multiple-batch cropping system is used in which ﬁsh of different
sizes and ages are present in the pond simultaneously. Harvest-size ﬁsh are
removed several times during the year, and ponds are restocked with ﬁngerlings without draining. Figure 13.3 shows catﬁsh that have been collected
by seine and are waiting to be loaded onto a truck for transport to the
Catﬁsh farming has become a major industry located primarily on the
Mississippi River ﬂood plain. A typical farm is several hundred hectares in
size, although some may be 1000 ha or more, with individual ponds of 5 to
10 ha. Large farms coupled with the development of specialized feed mills
Richard T. Lovell
Channel catﬁsh harvested by seine from a 5-ha pond are held in a “live sock” until
loaded onto a truck and hauled alive to the processor.
13. Diet and Fish Husbandry
and catﬁsh processing plants have made catﬁsh farming a proﬁtable, stable
industry. Ninety-ﬁve percent of the catﬁsh produced in the United States is
produced in Mississippi, Arkansas, Alabama, and Louisiana in approximately
160,000 ha of catﬁsh ponds (USDA, 1999).
Although nitrogenous and phosphorus compounds excreted from the
ﬁsh or from uneaten feed are of concern, management of dissolved oxygen
is the most critical pond environment problem. During periods of heavy
feeding dissolved oxygen levels drop precipitously during the night. Permanent electrical aerators located in the ponds as well as temporary aeration
devices powered by tractors are used to provide oxygen in emergencies.
13.2.2. Feeding Practices
Even though catﬁsh have been cultured for many years and considerable
research has been conducted on nutrition and feeding of catﬁsh, feeding is
far from an exact science. There is considerable variation in feeding practices among commercial catﬁsh farms. Some farmers feed a ﬁxed amount
of feed in the pond daily, usually the maximum amount that the pond can
safely “metabolize,” and have the pond overstocked with ﬁsh so no feed will
be wasted. Other farmers feed what the ﬁsh in each pond will efﬁciently
consume each day, using a ﬂoating feed so feeding activity can be observed.
However, water quality or ﬁsh health problems may cause farmers to restrict
the daily feed allowance or to feed less frequently.
Newly hatched catﬁsh fry, which are only about 2.5 mm in total length, are
usually held in indoor troughs or tanks for about 10 days before releasing
into outdoor nursery ponds. Initially, catﬁsh fry use their yolk sac as an
energy and nutrient source. Once the yolk sac is absorbed, at approximately
3 to 5 days after hatching, fry begin to seek food. In the hatchery, fry are
fed ﬁnely ground, meal-type feeds containing 45 to 50% protein, supplied
primarily from ﬁsh meal, at a daily rate equal to about 25% of the body weight
divided into 8 to 10 feedings. Table 13.2 presents a formula for fry feed.
The best way to ensure good growth and survival of recently stocked fry is
to ensure that plenty of natural food is available in the nursery pond when
the ﬁsh are stocked. Natural foods for channel catﬁsh fry include microcrustaceans, insect larvae, and zooplankton. Even though fry presumably meet
their nutrient needs from natural food organisms for the ﬁrst 3 or 4 weeks,
they are fed once or twice daily using ﬁnely ground feed at a rate equal to 25
to 50% of the fry biomass. Since the feed is a supplement to natural pond
foods, it is not necessary to feed a high-protein feed as used in the hatchery.
Fines from 32% grow-out feeds are suitable for catﬁsh fry during this phase.
Richard T. Lovell
Catﬁsh Fry, Fingerling, and Production Feed
Menhaden meal (61%)
Soybean meal (48%)
Cottonseed meal (41%)
Catﬁsh vitamin mixa
Trace mineral mixa
Vegetable or ﬁsh oilb
Commercial mix that meets the requirements for catﬁsh.
Sprayed on after extrusion to reduce feed dust.
After a few weeks, when the fry reach 2.5 to 5 cm in length and are generally referred to as ﬁngerlings, they will come to the pond surface to
accept food. Initially, the small ﬁngerlings are fed once or twice daily to
satiation using a crumbled feed or small pellets (3-mm diameter) containing 35% protein (Table 13.2). The feed should contain some ﬁsh meal or
other animal protein source. Fingerlings are generally fed according to this
regimen until they reach about 12 to 15 cm in length, at which time they are
stocked in grow-out ponds to be grown as production ﬁsh to harvestable size.
126.96.36.199. Production Fish
Catﬁsh grown to harvestable size are typically fed a 28 to 32% ﬂoating
feed (Table 13.2), with a pellet diameter of approximately 4 to 5 mm. Lowprotein feeds (28%) can be used if the ﬁsh are fed to satiation. Li and
Lovell (1992) found that channel catﬁsh grew maximally when fed 24 or
26% balanced protein feeds, if fed as much as they would consume, but if
they were fed to less than satiation, they required higher levels of protein for
optimum growth. Because most farmers feed conservatively (underfeed) to
avoid waste, a 32% protein feed is considered practical for most operations.
Catﬁsh are fed expanded feeds that are manufactured by extrusion processing, which allows them to ﬂoat. This type of feed is advantageous because
the farmer can see how much the ﬁsh are consuming. On most commercial
catﬁsh farms the feed is typically blown onto the surface of the water using
13. Diet and Fish Husbandry
pneumatic dispensers mounted on or pulled by vehicles. Feed should be
scattered over a large area to provide feeding opportunities for as many ﬁsh
as possible. It is desirable to feed on all sides of the pond, but this is generally
not practical because prevailing winds dictate that feed must be distributed
along the upwind side to prevent it from washing ashore.
Typically, catﬁsh producers feed once a day, 7 days a week. Feeding twice
a day when the water temperature is above 25◦ C has been shown to allow for
a higher rate of feed consumption and a correspondingly faster growth rate
(Lovell, 1979); however, the additional time and management required for
multiple daily feedings make the practice unattractive to most farmers. During disease episodes or during extremely hot weather when feeding activity
is poor, it may be beneﬁcial to feed every other day or every third day.
Feed allowance is affected by several factors including ﬁsh standing crop,
ﬁsh size, water temperature, and water quality. Water temperature and ﬁsh
size have a profound affect on feed consumption by channel catﬁsh. Feed
consumption increases as the water temperature increases until a temperature of about 32◦ C is reached and subsequently begins to decrease. As the
ﬁsh size increases, the feed consumption as a percentage of the body weight
decreases and the feed conversion efﬁciency is reduced (Table 13.3).
When catﬁsh are cultured using a multiple-harvest production system in
which several sizes of ﬁsh are present in the pond simultaneously, they should
be fed to satiation. Offering as much feed as possible, without wasting feed,
provides a better opportunity for the smaller, less aggressive ﬁsh to get feed.
Satiation feeding appears to be particularly important when catﬁsh are fed
less frequently than on a daily basis. Although it is recommended that catﬁsh
Feed Consumption and Feed Conversion for
Different Sizes of Channel Catﬁsha
(% body weight)
a At optimum temperature (27–29◦ C). Adapted
from Lovell (1989).
Richard T. Lovell
typically be fed as much feed as they will consume, at high standing crops of
ﬁsh it may be impossible to satiate the ﬁsh and maintain the water quality.
Feeding rates should not exceed what can be assimilated by organisms in the
pond and not require excessive use of aeration or cause toxic concentrations
of waste metabolites, such as ammonia. Generally, the long-term daily feed
allowance should not exceed 100 to 120 kg per hectare. Overfeeding should
deﬁnitely be avoided because wasted feed reduces feed efﬁciency and also
contributes to deterioration of the water quality.
The best time of day to feed is still debated, but the point is more or less
academic. On large catﬁsh farms, the time at which ﬁsh are fed is largely
dictated by the logistics required to feed many hectares of ponds in a limited
time period. As a result, many catﬁsh farmers start feeding in early morning, as soon as the dissolved oxygen levels begin to increase. Figure 13.4
shows the diurnal variation in dissolved oxygen in typical catﬁsh ponds.
Research has shown that there are no signiﬁcant differences in weight gain,
feed consumption, feed conversion, and survival among catﬁsh fed to satiation at 0800, 1600, or 2000 hr (Robinson et al., 1995). There were also
no differences in emergency aeration time among treatments. However,
Typical diurnal variation in dissolved oxygen content in intensively stocked