Aquaculture

45
Elsevier Publishing Company, Amsterdam - Printed in the Netherlands

A REVIEW OF RECENT DEVELOPMENTS IN Tilapia CULTURE,

WITH SPECIAL REFERENCE TO FISH FARMING IN THE HEATED
EFFLUENTS OF POWER STATIONS

R.G. KIRK
Ministry ofAgriculture, Fisheries and Food,
Marine Hatchery, Port Erin, Isle of Man (U.K.)

Preliminary studies suggest that Tilapia species are suitable for culture
in the heated effluents of power stations. Growth and reproduction may
be normal or near normal in saline conditions, and Tilapia are able to
tolerate low oxygen levels and moderately low temperatures (6- 12 C).
Population control techniques and food requirements are described and
discussed. Although Tilapia may have a secondary role as sport fish or in
clearing excessive plant growth in power stations, their main value would
seem to be as food fishes.

INTRODUCTION

The culture of Tilapia in recent times was probably initiated in Keyna

in 1924. Experiments in the former Belgian Congo commenced in 1937,
and intensive culture of Tilapia species has been carried out in the Congo
since 1946 (Huet, 1955, quoted by Chimits, 1957).
The discovery of T. mossambica Peters in Java in 1939, and the sub-
sequent dispersion of the species, initially throughout the Far East, and
later to South and Central America, has been well documented by Chimits
(1955 and 1957). In many of the countries to which it has been intro-
duced, T. mossambica has proved itself to be a useful addition to the local
fauna, and yields of from 270 pounds per acre in rice fields to 6-8 tons
per acre with heavy fertilization and feeding have been quoted by Swingle
( 1960).
Tilapia species are eaten by Europeans in the former colonies of Africa
(Cole, 1968) and it is possible that there is a potential market for Tilapia
in Europe. In anticipation of this market, a commercial fish-pond interest
46

has commenced an investigation into the suitability of cooling-water

ponds in Germany for Tilupia culture (Denzer, 1968). In Israel, where
many of the present inhabitants are of European origin, the production of
pond-reared Tilapia rose from 380 tons (or 316% of the total pond-reared
fish) in 1965 to 1178 tons (or 11 .S% of all pond-reared fish) in 1969. The
total Israeli Tilupia production in 1969 was 1558 tons, 215 tons more
than in the previous year, and more than double the total production for
1964. Marketing of Tilupia in 1969 was 16% higher than in 1968, and this
increase was supplied entirely from the pond-fish sector; pond-grown
Tilapia represented 75% of the totaimarketed in 1969. In spite of the
steady annual increase in Tilapia production, market shortages occurred in
the summer of 1969, although these were less pronounced than in the
previous year (Sarig, 1970a). It is of interest that the increase in Tilupiu
production has been largely at the expense of carp, a traditional favourite
in central and eastern Europe.
In the early 1960s work on. the culture of Tilapia in impoundments
heated by the inflow of water from power stations and other installations
appears to have been undertaken in at least three countries (Iles, 1963;
Kiener and Lamarque, 1969; Krayev, 1966). These pioneer experiments
have since been followed by studies on the suitability of T. gulilaea Artedi
and T. nilotica L. for culture in factory cooling ponds in Germany
(Denzer, 1968). All the investigations undertaken clearly demonstrated
the suitability of Tilapia species for culture under factory conditions,
although work in this field appears to have been abandoned in both
Britain and France.
Since power stations use large quantities of water for cooling purposes,
they are normally situated near inland waters or the sea. In the United
Kingdom, nearly all nuclear power stations now operating or under con-
struction are sited on the coast, at least partly because of the need for vast
quantities of water. Investigations into the feasibility of large-scale fish
culture using heated water in Great Britain have therefore been directed
mainly towards the study of marine fish, and cooling waters from power
stations have been shown to be suitable for Qhe culture of plaice (Pleuro-
nectes pkztessa) and sole (Solea solea) (Shelbourne, 1970). Both these
species have a high rate of growth in heated waters, and it has been
suggested that theycould be cultured to a marketable size in half the time
taken in natural waters.
A list of the fish species commanding the highest price on markets in
the UK has recently been prepared (A. Jones, Ph.D. thesis), and three of
these - ,turbot (Scophthalmus maximus L.), Dover sole (Solea soleu L.)
and halibut (Hippoglossus hippoglossus L,) - are outstanding in holding a
high price throughout the year. Their estimated growth rates under
 47

natural conditions and minimum marketable sizes are given below:

Turbot (both sexes) 30 cm 614g 28 years

Dover sole (both sexes) 24 cm 197g 2.8 years
Halibut (both sexes) 32 cm 271 g 2.0 years

Assuming that one or more of the above species would respond well to
artificial culture, and that the growth rate would be of the order of twice
that of growth under natural conditions, it seems probable that a market-
able size could be obtained in from 1 to 1% years.
Growth rates of Tilapia have been found to vary considerably between
the sexes, from species to species, and with the environmental conditions
in which culture is carried out (Chimits, 1955 and 1957). In parts of the
Far East and in Israel, where culture techniques are highly sophisticated, a
number of Tikzpia species are marketed after one years growth only, and
it is likely that such growth rates could be obtained in power station
cooling water in temperate climates. In addition to reaching a marketable
size in about the same time, the cost of food for Tilapia (which grow well
on cheap vegetable waste and waste grains) would almost certainly be less
than that for any truly marine. species commanding a reasonable market
price.
Tilapia species respond well to an aquarium environment, and, thanks
to this characteristic and to their importance as food fishes, there is a
considerable body of data on the biology of the genus. The following
review of the literature deals with those aspects of the physiology and
breeding behaviour of Tilupiu which are relevant to the development of
culture techniques in both fresh and saline waters.

SURVIVAL, GROWTH AND REPRODUCTION IN SALINE CONDI-

TIONS

It has been assumed by Myers (1938) and Steinlitz ( 1954) that Tilapia
species evolved from a marine ancestor which penetrated fresh water.
Such an ancestor would certainly account for the marked euryhalinity of
certain Tilapia species (Chervinski, 196 la).
Tilapia nilotica L. is able to survive direct transfer from fresh water to
60-70~ sea water (20.2-25.0 ho salinity), and through gradual adap-
tion is able to withstand a medium of up to 150% sea water (53.5 Oloo
salinity) according to Lotan (1960). The rate of growth of T. nilotica at
high levels of salinity has been studied by Chervinski (196 la), and his
results show that growth in 50% sea water is not significantly different
from that of the same species in fresh water. Chervinski (196 lb) also
48

notes that T. nilotica are able to breed in their first year in 50% sea water,
although the number of young produced may be smaller than in fresh
water.
Observations by Vaas and Hofstede (195 2) established that T. mossam-
bica tolerates brackish water in Java, and Brock and Takata (1955) have
shown that this species is able to grow and reproduce in brackish waters of
up to 20 Oh0 in Hawaii. More recently, Canargaratnam (1966) had
demonstrated that T. mossambica has a higher rate of growth in 50% sea
water than in either fresh water or undiluted sea water (Table 1).

TABLE 1

Initial average weight of Tilapia mossambica and the percentage increase in weight at the end of the
4th and 8th week. The number of fish surviving at each stage is indicated in parentheses. (Canar-
garatnam, 1966)

Salinity Initial At end of 4th week At end of 8th week

condition average
weight Average % Average %
W weight increase weight increase
(9) (g)

Fresh water 0.20 (15) 0.40 (10) 100 0.70 (7) 250
50% sea water 0.24 (15) 0.60 (8) 150 2.20 (2) 817
100% sea water 0.25 (15) 0.50 (8) 100 1.53 (7) 512

Vaas and Hofstede (1952) reported that although T. mossambica grows

well and reproduces in water of up to 30 ho salinity, reproduction did
not occur at salinities of between 30 o/oo and 40 o/oo , although growth
was satisfactory. Hora (1955), on the other hand, found that reproduction
occurred in sea water of up to 35 ho salinity, and Canargaratnam ( 1966)
has confirmed that T. mossambica will breed in water of this salinity, and
that the young are capable of survival. The closely related T. hornorum
Trewavas appears to be equally tolerant of saline conditions, and has been
reared in marine fish ponds on Zanzibar Island (Talbot and Newell, 1957).
According to El Saby (195 1), T. nilotica, T. galilaea and T. zillii are all
known to breed in the Great Bitter Lakes of Egypt at salinities of from
13.5 to 29.0 o/oo . Of the above three species, T. zillii appears to be the
most tolerant of high salinities, since in Lake Qarun (a formerly fresh-
water lake which has now become highly saline) it survived and bred at
29 oh0 salinity after all other species, including T. galilaea and T. nilotica,
had disappeared (El Zarka, 1956). More recent observations have shown
that T. zillii has established itself in the Red Sea at a salinity of 42.7 o/oo
(Bayoumi, 1969).
 49

OXYGEN DEMAND AND TEMPERATURE TOLERANCE

Tilapia species are able to survive in extremely adverse environmental

conditions, and are frequently to be found in habitats which most other
fish genera are unable to tolerate. Beadle (1932) has pointed out that the
last three genera of fish to survive in shallow African lakes during periods
of drought are Clarias, Barbus and Tilapia.
Denzer ( 1968) has shown experimentally that T. galilaea and T. dotica
have a comparatively low oxygen demand per 100 g live weight; in T.
galilaea it was 26.9 cm3 for a fish of average weight 10.2 g, corresponding to
38.42 mg/h, and in T. nilotica (average weight 2.0 g) 14.8 cm3, corres-
ponding to 2 1.4 mg/h. These values ire considerably lower than those for
European fish.
Small fish need on average more oxygen than larger ones; for example.
trout of 10 g weight require 34 cm3 for 100 g weight/h, and trout of 40 g
only 5.3 cm3 per 100 g weight/h. Comparing these figures for those
obtained for T. rzilotica, it is clear that the oxygen demand of the latter
species is very low indeed, while the relatively high figure for T. galilueu
suggests that there may be considerable variation in oxygen demand bct-
ween Tilapia species.
T, mossambica is also undemanding in relation to the oxygen content
of the water, and a laboratory stock has withstood a reduction in dissolved
oxygen down to 1 mg/litre, according to Mironova (1969). In aquaria
with a high population density of up to 47 g/litre (the number and size of
the fish concerned are not given) the introduction of a considerable
amount of food caused substantial pollution of the water (pH ranged from
6.0-6.5, permanganate oxidizability reached 17.3, and dichromate oxi-
dizability, 67); nevertheless the fish tolerated this degree of pollution.
Worthington (1932) recorded shoals of Tihpia skittering, i.e., sucking
at and breaking the water surface, in Lake Edward, East Africa; these fish
were apparently feeding on the lakefly corpses thickly scattered on the
water surface at the time. Shoals of 4- 15 cm long T. nilotica have also
been observed to skitter on a number of occasions, but apparently for
respiratory reasons. Aquarium observations suggest that skittering occurs
both to collect food from the surface and when the water is deoxygenated
(Lowe (McConnell), 1958).
Despite their normal preference for high temperatures, many Tilaplu
species, particularly those at the southern and northern limits of the distri-
bution of the genus are able to withstand relatively cold conditions. T.
sparmanm A. Smith is capable of surviving the low temperatures of the
Cape winter in South Africa, and is known to have withstood a tempera-
ture of 7 C, although. spawning does not take place below 16 C (Hof-
50

stede, 1955).
According to Chimits (1957), T. mossambica is killed by temperatures
of between 8 and 10 C, although the same species in aquaria in Leather-
head in England has withstood a tempertature of 7 C (R. S. A. Beau-
champ, personal communication). Li Gang Long et al. ( 196 1) found the
lower lethal limit for T. mossambica to be 5.5 C in Hanoi, although
normal vital activity was affected below 20 C.
The effect of low temperatures on T. nilotica and T. galilaea is known
in some detail. T. galilaea has been shown to be more sensitive to low
temperatures than T. nilotica, although both species took food and grew
at a temperature of 14 C. T. galilaea in Lake Kinneret stores body fats in
winter at temperatures of 16-l 1 C before the spring spawning, and
grows in length under these conditions. The effect of temperature on T.
nilotica was shown schematically by Y ashuov ( 1960).
Denzer (1968) gives a temperature of 11 C for the lower lethal limit of
T. nilotica, a somewhat higher figure than that given by Yashuov. Cases of
mortality of T. zillii have been recorded from Lake Huleh in Israel, at a
time when temperatures were known to range from 6-8 C (Yashuov,
1960).
Recent work on the hybridization of Tilapia in Israel has demonstrated
that low temperature tolerance may be transmitted to Tilapia hybrids. T.
vulcani (= T. nilotica) introduced to Israel from Lake Rudolf, Kenya, were
found to die at temperatures of 1 l- 13 C, and local T. aurea at 8.5-8.0
OC, but the hybrid of these two species has a low temperature tolerance of
9.0-8.0 C (Sarig, 1970b).
It is probable that all Tilapia species are tolerant of temperatures up to
35 C. T. grahami Blgr. normally inhabits hot spring water of between 35
and 40 OC, although very small increases in temperature above 40 C
prove fatal (Coe, 1966), and Denzer (1968) gives an upper lethal limit of
42 C for T. nilotica. Allanson and Noble ( 1964) suggest on the ba& of
experimental data that the highest temperature tolerated by T. mossam-
bica is 38 C, but according to Li Kuang Long et al. (1961) the upper
lethal limit is 42.2 C, although normal vital activity is affected above 38
C.

POPULATION CONTROL

One of the main problems in Tilapia culture is over-population brought

about by the high rate of survival of the young. This leads to a marked fall
in the growth rate, frequently as a direct result of inadequate food sup-
plies. Experiments in Malaya (Chen, 1965) show that even when food
availability is not a limiting factor, growth of T. mossambica in ponds of
 51

half an acre or less may still be poor, and Chen has suggested that this may
be the result of either a build-up in the level of excretory metabolites, or
the outcome of a social hierarchy of dominance. Iles (in press) attributes
the size reduction to an adverse response to the external environment, and
regards this stunting as an adaptive mechanism involving both reproduc-
tive and growth characteristics.
Restricted growth rates are not confined to pond populations, and
stunted Tilapiu from shallow lakes and lagoons have been described in the
literature (Lowe (McConnell) 1958 and Coe, 1966). Several of the Tilapia
species most suitable for culture breed at a.very small size in a restrict-
ed environment. T. mossambica in ponds may breed at 8-9 cm (Chimits,
1955); T. nilotica, with normally breeds at well over 20 cm total length
in large lakes, bred at 17 cm in experimental ponds (Lowe (McConnell)
1958), and T. galilaea at 15 cm (Yashuov, 1956).
Growth after maturity is slow in T. variabilis Blgr. (Fryer, 196 1) and
this is almost certainly true of all Tilapia, although males of the T. mos-
sambica complex have a higher rate of growth than females (Whitehead,
1962). As a result of these factors, work on controlled stocking of ponds
has largely been directed towards the isolation of males from females,
either by physical separation .of the sexes (the females being retained
solely as a breeding stock) or the development of all-male hybrids. Preda-
tory fish have also been stocked with Tilapia in order to restrict the
numbers of the latter, and it has been shown that confinement of Tilapiu
in cages can effectively prevent the fertilization of any eggs spawned
(Pagan, 1969).

Monosex culture and hybridization

Work on the monosex culture of TiEapia nigra in Kenya has been

described by Brown and van Someren (1953). T. nigra fry of unspecified
size were cultured separately from adult fish at a density of 2500 per ha.
Three months after stocking, when the young Tilapia were of an average
size of 10 cm, they were removed from the fry pond. At about 10 cm the
differential in size between the sexes is believed to be about 2 cm, and the
larger (theoretically male) fish were then selected and placed in a separate
pond. Following separation the males were reared to an economically
acceptable size of 20 cm, giving a yield of 300 kg. per ha. This practice has
been followed with T. mossambica in Indonesia on a limited scale (Huet,
1955) and in culture experiments in the West Indies (T.W. Burdon, per-
sonal communication to Chimits, 1957).
Monosex culture following physical separation of the sexes now appears
to have been generally superseded by the culture of all-male hybrids. The
52

first successful experiment in hybridization was made by Hickling ( 1960)

between male T. hornorum Trewavas (the Zanzibar Tilapiu) and female
T. mossambica (the Malayan Tilapia). The offspring were almost certain-
ly 100% males; a few females were found to be present, but it is unlikely
that these arose from the experimental crossing. Later work showed that,
with necessary precautions, 100% male offspring could be obtained in
every case (Hickling, 1968). The hybrid males grow faster than fish of
either parent stock, and brood sizes are larger (Hickling, 1959). Pruginin
and Kanyike (1965) have also succeeded in producing a T. hornorum
hybrid, by crossing male T. hornorum with female T. nilotica.
Work on the hybridization of T. nilotica females with T. urea males
has resulted in a cross producing mostly males, the F, generation having a
higher rate of growth than either of the parent species. The hybrids reach-
ed a weight of 600-650 g in one year in aquaria, whereas fish of the
parent stock grew to only 400-450 g over the same period (Fishelson,
1962). The hybridization experiments were later transferred to the field at
Gan Shmuel in Israel, and observations over three years showed that re-
sults obtained in the laboratory could be replicated under field conditions
(Fishelson, 1967). Although the first experimental crossing of T. nilotica
and T. aurea gave 100% males, later field work showed that hybrid
females may also result from the cross, and that these females had a higher
rate of growth than pure males (Fig. 1).
A comparison of the growth rates of hybrids from a cross between T.
hornorum males and T. mossumbica females and the hybrid offspring of

n.
1 Hybrids $
232 specimens

:x
::::
::::
R
a
::::
::::
::::
:::: :A
:::: ::::
:::: ::::
:::: ::::
.g ::::
::::: ::::
::::: R
::::: ::::
::::: ::::
::::: a
::::: ::::
::::: ::::
# q:
150-180 181- 210
Group of total length in mm

Fig. 1. The percentage of various length groups in the total T. aureu and female hybrids, measured
from the fish ponds of Gan-Shmuel in 1965 (Fishelson, 1967).
 53

male T. hornorum and female T. nilotica is given by Bard (1969). Growth

rates of the two hybrids were similar but the T. hornorum x T. nilotica
cross was described as being the more difficult and time-consuming.
Bard (1964) has summarized the results of hybridization of a number
of other Tilapia species, and Yashuov and Chervinski ( 1960) have reported
a cross between T. galilaea and T. nilotica. None of these hybrids appears
to possess any more favourable features than the parent stocks.

Predator control

Although T. sparmanni has long been used as a forage fish for black
bass (Micropterus salmoides Lacepede) according to Chimits (1957), there
appears to have been little deliberate exploitation of predators as a means
of cropping excess numbers of Tilapia in fish ponds (Swingle, 1960). By
stocking black bass with T. mossambica and supplying a supplementary
feed to the Tikzpia, high production of marketable fish was obtained
(Swingle, 1960). Black bass have been stocked with T. shirana ssp. and T.
melanopleura Dum. in estate ponds in the southern region of Malawi, and
have generally proved to be at least moderately effective as a predator of
juvenile Tilapia (Kirk, unpublished data). Semakulu and Makora ( 1968)
found that black bass were not effective in controlling population growth
in fish ponds in Kenya; they appeared to show a preference for the tad-
poles which were present in great numbers in ponds used for experiments
in predator control.
The Nile perch (Lates niloticus Cuvier and Valenciennes) was also em-
ployed by Semakulu and Makora as a predator in mixed cultures with
Tilapia, and proved to be highly effective. Because of difficulties in
breeding Nile perch, this experiment was abandoned. Research carried out
by the Uganda Fisheries Department has shown that the catfish Bagrus
docmac is as effective a predator as the Nile perch in controlling Tilapia
populations. B. docmac breeds freely in ponds and its growth rate when
stocked with T. nilotica or T. zillii is reported to be quite satisfactory
(Kanyike, 1969).

Cage culture

According to the way the young are reared, Tilapia can be divided into
two main groups, guarders and brooders (Lowe (McConnell), 1955). In
the former group the eggs are deposited on specially cleared areas of the
substrate, to which they adhere; they are then guarded by the parents
until they hatch. Only four species are known to be guarders: all other
Tilapia are brooders. The eggs of these species are picked up by one of
54

the parents (in nearly all known cases, the female) almost immediately
after they are laid, and are brooded in the mouth until the larvae are fully
developed.
In order to suppress breeding in Tikzpia, experimental culture of T.
aurea in floating cages has been carried out in the United States. When
raised in cages T. aurea does not reproduce, whereas under pond condi-
tions at the same rates of stocking (from 7000 to 15 OOO/ha) its reproduc-
tion rate is high (Pagan, 1969). Pagan gives two main reasons for the lack
of reproduction in cages: (a) even if spawning took place (spent females
were collected) the reproductive behaviour would be altered in such a
manner that the eggs would not be fertilized; (b) the female T. aurea is a
mouthbreeder, but under cage conditions such parental care is elimi-
nated, because the eggs fall through the bottom of the cage. It seems
probable that, even if some of the eggs were fertilized, lack of parental
care would preclude their normal development.
The experimental data obtained from cage culture experiments (Pagan,
1970) clearly show that this technique is more efficient in terms of pro-
duction per unit volume than is traditional pond culture. Further, the
absence of small fish in the cage at the end of the experiment demon-
strated that breeding was totally inhibited.

FEEDING

It has long been established that Tilapia mossambica is omnivorous

(Vaas and Hofstede, 1952; Chimits, 1955), and recent work has demon-
strated that this fish will grow well on a variety of cheap, easily obtainable
foodstuffs. Verigin (1963) has shown that blue-green algae are readily
consumed by this fish in ponds heated by power station cooling water,
and Krayev (1966) has discovered filamentous algae, Charophyfa, mos-
quito larvae, mayflies,.beetles, worms and Gambusia in the stomachs of T.
mossambica from power station cooling ponds. Other feeding regimes
which have proved successful are raw minced meat, fish, mollusc tissue,
and oatmeal cakes (Atz, 1954).
Soviet research workers have tested the response of T. mossambica to a
wide range of foods and found that the leaf material from many common
vegetables and dried Chlorella were readily consumed by both adults and
larvae (Mironova, 1.965, 1967a and 1967b; Mironova and Skvortsova,
1967). In an experiment set up to study the effects of Chlorella-con-
taining granulated feed on growth, Antsyshkina et al. ( 1968) found that high
rates of growth in both weight and length were achieved with granules of
Chlorella alone. Growth was further improved by the addition of small
quantities of yeast, but. the presence of too much yeast,in the.diet was
 55

liable to induce functional disturbances attended by some mortality.

The daily food requirement of T. mossambica has been established in
some detail by Mironova (1969). At the age of three months, juvenile
food consumption had reached 20-40% of body weight per day where
the diet had an animal basis, and 30-60% where plant food was given By
the age of six months, consumption had declined to between 5-15% and
1O-250/, of the body weight for animal and plant tissues respectively. At
one year of age, food intake was about 3%, and in old fishes it was
I- 1.5% of body weight.
Lowe (1958) was of the opinion that T. niloticu feeds mainly on algae,
and concluded that the species had more generalized feeding habits than T.
esculentu Graham, which feeds almost entirely on phytoplankton. Daget
( 1954) found that T, nilotica feeds on phytoplankton and bottom animals
and also swallows mud particles. Yashuov and Chervinski (1961) suggest
that, in lakes, T. niloticu feeds on phytoplankton and epiphytic diatoms,
and when confined in ponds and lagoons, on bottom organic debris.
In an experiment set up to determine the range of naturally-occurring
food organisms eaten by T. nilotica, rotifers, copepods, insect larvae,
blue-green algae, green algae, diatoms and nannoplankton were found in
fish .of all sizes, but cladocerans were found only in the guts of fish up to
50 mm total length. Euglena, filamentous algae and higher plants were
found in the gut contents of all but the smallest fish of up to 20 mm
(Yashuov and Chervinski, 196 1).
In an evaluation of the various food items in the diet of T. niloticu, the
fish were fed on phytoplankton, water bugs (Corixa), chironomids and
cotton-seed cakes, either singly or in various combinations. It was not
possible to make valid comparisons of the nutritional value of various
food items, but it was clear from the results that the natural food spec-
trum of T. nilotica is very wide. Even juveniles of only 2.5 g weight were
found to feed on plankton, water bugs, other insects and cotton-seed
cakes.
Two species of Tilupia which are commonly reared in ponds in Africa,
T. melanopleura and T. zillii, are predominantly weed-eating in habit. T.
zillii is reported to have consumed a thick growth of Valisneria in an
enclosure with an area of about 6 m2 in the coolant reservoir of a power
station in the USSR (Verigin, 1963). T. melunopleura is reputed to feed
largely on Cerutophyllum in Lake Nyasa (Jackson, 1961) but is capable of
utilizing a very wide range of higher plant life (Chimits, 1955).

DISCUSSION

At least three species of Tilapia (T. mossambica, T. nilotica and T. zillii)

56

are known to grow well and to breed at high salinity levels, and could
therefore presumably be cultured in sea water. Although T. niloticu is
unable to tolerate direct transfer from fresh water to 70% sea water, it is
capable of withstanding salinities of up to 150% sea water if the salt
concentration is gradually increased over a period of 27 days (Lotan,
1960).
Since there is usually a high rate of flow through cooling ponds of
power stations, it seems unlikely that oxygen depletion would occur
during normal circumstances. Even if such conditions were to occur,
Tilapiu species are able to tolerate very low levels of dissolved oxygen.
Gribanov et al. (1968) note that the temperature of water discharged
from certain Soviet power stations is from lo- 15 C higher than the
temperature of neighbouring natural waters, and that the water tempera-
ture at some power stations in the central region of the USSR may be in
excess of 20 C even in winter. It is extremely unlikely that this tempera-
ture is below the lower lethal limit of any Tilapia species, and at temper-
atures lo- 15 C above ambient, good growth may be expected over the
greater part of the year. The hardier species of Tilapia (e.g., T. mossam-
bicu and T. zillii) would almost certainly withstand much lower tempera-
tures than those recorded by Gribanov et al. ; moreover, hybrids (between
T. mossambica and T. nilotica) not only grew faster with a better food
conversion than the parent species, but were also found to be intermediate
between the latter in their tolerance of low temperatures (Avault and
Shell, 1968).
Avault et at. (I 968) found it necessary to overwinter large numbers of
Tilapia in troughs at a temperature of about 21 C. It was found that,
even allowing for the cost of this operation, Tilapia can be cultured on an
economic basis, provided that a good summer growth rate is maintained.
By using fast-growing hybrids, it should prove possible to obtain annual
yields in excess of 1000 kg of Tilapia per ha in heated ponds. Since water
temperatures outside the ponds would be too low to permit the devel-
opment of wild populations, there would be no risk of cross-breeding.
Gaucher (1970) has pointed out that the concept of production per unit
area has little or no meaning where there is a strong flow of water, and
where food supply is not a limiting factor. He suggests that extremely high
production could be achieved by growing fish in raceways at heavy
stocking rates. This has been confirmed for carp by experiments in high
density culture (Meske, 1968); fish stocked in a 40-litre aquarium reached
a weight of 800 g in a year, and the growth rate of five fish confined to
one half of a 40-litre aquarium was only slightly less than that of the same
number of fish occupying a whole tank. The flow rate of the tanks was
very high (total volume replaced in 15 min) and Meske concludes that the
 57

ideal stocking rate is more dependent on the flow rate of water than on
the size of the tank, provided that a suitable diet is fed. Although no
studies have been made on the growth rate of Tilapia in raceways, recent
population density experiments in America (Shell, 1968) indicate that
male T. nilotica in monosex cultures grow equally well at stocking
densities of 2500 per ha. and 5000 per ha., when a suitable diet is fed at
optimum feeding rates.
The growth of algae in the cooling ponds of coastal power stations may
lead to the development of conditions detrimental to fish farming. Bowers
(1970) has suggested the use of the herbivorous ormer (Haliotis sp.) and
mullet (Mugil sp.) for clearing excess weed from cooling ponds, but
neither appears to have been successful in dealing with algal growths at an
experimental fish farm at Hunterston in Scotland (M. Cheetham, personal
communication). Work in the Soviet Union on the control of both algae
and higher plant species in the cooling ponds of inland power stations
using fresh water has shown that both T. mossambica and T. zillii are very
effective in clearing large areas of plant growth (Verygin, 1963; Krayev
1966). Krayev also notes the added bonus of the production of edible fish
(T. mossambica) at insignificant cost.
In addition to the role of weed control, Tilapia species might also be
grown for recreational purposes, either as sport fish in their own right, or
as fodder for larger carnivorous fish. Some years ago Iles (1963) pointed
out the need for developing bodies of water for angling, and since then,
while the number of anglers has grown, the area of unpolluted water in
the United Kingdom has, if anything, diminished. Coarse fishing is one of
the major sporting interests in the industrial Midlands, an area with some
of the most polluted waterways in England, and there seems to be little
doubt that the use of power station cooling ponds for coarse fishing
would yield a high revenue in licence fees, probably greatly in excess of
the market value of the fish cultured.
While Tilapia species might be expected to serve a subsidary role as
consumers of unwanted vegetation, their main value - other than for
angling purposes - would seem to be as food fishes. In the late 1950s
Tilapia were air-freighted from Kenya to London, where they were
supplied to luxury hotels and restaurants. Although the Tilapia were well
received, problems in maintaining a constant supply occurred, and their
export from Kenya was discontinued (P. Watson, personal communica-
tion). Israel is at present exporting Tilapia fillets to the USA, following
the first consignment of 25 tons in 1969 (Sarig, 1970a).
Although nearly all but the most recent work on Tilapia culture was
carried out in the Middle East or in Africa, most of the techniques
acquired could be applied to fish farming in power station cooling water,
58

either directly, or with relatively minor modifications, and evidence from

Europe and North America indicates that there might be a market for
Tirclpia in both continents.

REFERENCES

Allanson, B. and Noble, R.G.( 1964) The tolerance of Tilapia mossambicu Peters to high tempera-
ture. Trans. Amer. Fish. Sot., 94, 323-332.
Antsyshkina, L.M., Kirilenko, N.S., Melnikov, G.B. and Ryabov, F.P. (1968) Dynamics of weight,
nutritional state and body length of Tilaprb mossambica Peters, reared on granular, Chlorella-
containing feed. Robl. Ichthyol. 8(4) 576-580.
Atz, J.W. (1954) The peregrinating Tilapia. Anim. Kingdom, 57(S) 148-155.
Avault, J.W. and Shell, E.W. (1968) Preliminary studies with the hybrid Tilapia Tilnprir niloticu x
Tilapia mossambica. FAO (Food Agr. Organ. U.N.) Fish. Rep., 44(4) 237-242.
Avault, J.W., Shell, E.W. and Smitherman, R.O. (1968) Procedures for overwintering Tilapia. FAO
(Food Agr. Organ. L!N.) Fish. Rep., 44(4) 343-345.
Bard, J. (1964) Ou en est actuellement la pisciculture africaine? Bull. Fr. Piscicult. ,214, S-28.
Bard, J. (1969) Tikrpio hybridization. FAO Fish. Bull., l(3) 5-6.
Bayoumi, A.R. (1969) Notes on the occurence of Tilapia zillii (Pisces) in Suez Bay. Mar. Biol.
N. Y., 4, 255-256.
Beadle, L.C. (1932) Scientific results of the Cambridge Expedition to the East African Lakes,
1930-31. 4. The waters of some East African Lakes in relation to their fauna and flora. J.
Linn. Sot., ZooL, 38, 157-211.
Bowers, A.B. (1970) Whatever happened to fish farming? New Sci., 3, 380-381.
Brock, E. and Takarta, M. (1955) Contribution to the problems of boat capture and mortality
together with experiments in the use of T&pin as live bait. Industrial Research Advisory
Council Hawaii, Final Report, 49, 39-40.
Brown, J.M. and van Someren, V.D. (1953) New fish culture method for Tikzpia in East Africa.
Nature (Land.), 172, 330-332.
Canargaratnam, P. (1966) Growth of Tilapia mossambicu Peters in different salinities. Bull. Fish.
Res. Sta. Ceylon, 19(1-2) 47-50.
Chen, -F.Y. (1965) Report of the Tropical Fish Culture Research Institute, Malacca.
Chervinski, J. (1961a) Laboratory experiments on the growth of Tikzpia nilotica in various saline
concentrations. Bamidgeh, 13( 1) 8- 13.
Chervinski, J. (1961b) On the spawning of Tilapia nilotica in brackish water during experiments in
concrete tanks. Bamidgeh, 13( 1) 30.
Chimits, P. (195.5) Tilapti and its culture. A preliminiary bibliography FAO Fish. Bull., 8(l)
l-33.
Chimits, P. (1957) The tilapias and their culture. A second review and bibliography. FAO Fish.
Bull. 10(l) l-24.
Coe, M. (1966) The biology of Tilapia gruhami Boulenger in Lake Magadi, Kenya. Actu @op., 23,
146-177.
Cole, H.A. (1968) The scientific cultivation of sea fish and shellfish. J. Roy. Sot. Arts, 116,
590-603.
Daget, J. (1954) Les poissons du Niger superieur. Mem. Inst. jk Afr. noire, 36, 341-344.
Denzer, H.W. (1968) Studies on the physiology of young Tilapia. FAO (Food Agr. Organ. U.N.)
Fish. Rep., 44(4) 357-366.
El Saby, M.K. (195 1) The lake fisheries of Egypt. PToc. Un. Sci. Conf., 7, 126.
El Zarka (1956) Breeding behaviour of the Egyptian cichlid fish Tilapia zilZii. Cope& 2, 112- 113.
Fishelson, L. (1962) Hybrids of two species of fishes of the genus Tikzpia (Cichlidae, Teleostei).
Fishermans Bull Haifu., 4( 2) 14- 19 (in Hebrew with English abstract).
Fishelson, L. (1967) Cichlidae of the genus Tilapia in Israel. Bamidgeh 18(3-4) 67-80.
 59

Fryer, G. (1961) Observations on the biology of a cichlid fish Tilapia variabilis Boulenger in the
northern waters of Lake Victoria (East Africa). Rev. Zool. Bot. Afi., 64(1-2) l-33.
Gaucher, T.A. (1970) Thermal enrichment and marine aquiculture. Marine aquiculture. Oregon
State University Press, pp. 141-151.
Cribanov, L.V.A., Korneev, A.N. and Korneeva, L.A. (1968) Use of thermal waters for commercial
production of carps in floats in the USSR. FAO (Food Agr. Organ U.N.) Fish. Rep., 44.
Hickling, C.F. (1959) Some marking experiments with the Tilapio fish. Malaysian Agr. J., 42(l)
21- 30.
Hickling, C.F. (1960) The Malacca Tilapia hybrids. J. Genef., 57( 1) l-- 10.
Hickling, C.F. (1968) Fishhybridization. FAO (Food Agr. Organ. UN.) Fish. Rep., 44(4) I- 11.
Hofstede, A.E. (1955) Report to the Government of Israel on inland fisheries. FAO (Food Agr.
Organ. iJ.N.) Fish. Rep., 327, 56 pp.
Hora, S.L. (1955) Handbook of fish culture in the Indo-Pacific Region. (Provisional edition) FAO,
Rome, Italy. Mimeo II, 264 pp.
Huet, M. (1955) Culture of Tilapia. FAO Third Infernafional Inland Fisheries I955 Training Cenfre
at Bogor. Lecture 5.65. FAO, Rome, Italy.
Iles, R.B. (1963) Cultivating fish for food and sport in power station cooling water. New Scientist,
17, 227-229.
Iles, T.D. (in press) Dwarfing or stunting in the genus Tilopia (Cichlidae); a possibly unique
recruitment mechanism. Rapp. I+oces- Verbaux Rdunions Cons. perma. inf. Explor. Mer, 164.
Jackson, P.B.N. (1961) Checklist of the fishes of Nyasaland. Oct. Pap. natn. MUX Sfh. Rhod., 258.
25B.
Jones, A. (1970) Some aspects of the biology of fhe turbof (Scophthalmus maximus I,.) with
special reference fo feeding and growth at the juvenile stage. Thesis submitted to the University
ofXast Anglia for the Ph.D. degree.
Kanyike, F.S. (1969) Predator of Tikzpia. FAO Fish Cuff. Bull., l( 3) 9.
Kiener, A. et Lamarque, P. (1969) Note sommaire sur un klevage de Tilapia dans un milieu
particulaire du Sud-Ouest de la France. Verh. Internat. Verein. I.imnol., 17, 662-664.
Kraycv, N.P. (1966) Tikzpia in the coolant ponds of a State Regional Flectric Station. Ryb. Khoz.,
3, 24-25.
Li Kuang Long, Nguen Tin Zau and Nguen Kuang Win (1961) The first results of investigation of
the effect of tefnperature on the fish ~ibpia mossambica Peters acclimatized in Vietnam since
1961. Zh. Obshch. Biol., 22(6).
Lotan, R. (1960) Adaptability of Tilapia nilotica to various saline conditions. Bamidgch l?(4)
96- 100.
Lowe (McConnell), R.H. (1955) The fecundity of Tilapia species. E. Afr. Agric. Ebresf .I., ?I( 1)
45-52.
Lowe (McConnell), R.H. (1958) Observations on the biology of Tilapia nilotica L. in East African
waters (Pisces Cichhdac). Rev. 2001. Bot. Afi., 57(1-2) 129.- 170.
Meske, Ch. (1968) Breeding carp for reduced number of inter-muscular bones and growth of carp
in aquaria. Bamidgeh, 20(4) 105- 119.
Mironova, N.V. (1965) The growth of Tilapia mossambicu Peters under aquarium conditions.
Doklady AN SSR, 165(5).
Mironova, N.V. (1967a) On selection of the animal components of closed ecological systems.
Problemy kosmich. biol., I.
Mironova, N.V. (1967b) Comparative evaluation of the growth of Tikzpia mossambica Peters when
fed on Chlorella and on other food. Problemy kosmich. biol. 7.
Mironova, N.V. (1969) The biology of Tilapia mossambica Peters under natural and laboratory
conditions. PTobl. Ichthyol.
Mironova, N.V. and Skvortsova, T.A. (1967) Growth of Tilapzizmossambica Peters and its relation-
ship to feeding. Problemy kosmich. biol., 7.
Myers, G.S. (1938) Freshwater fishes and West lndian zoogeography. Smifhs. Inst. Publ., 3465.
339.
Pagan, F.A. (1969) Cage culture of Tilapia. FAO Fish Cult. Bull., 2( 1) 6.
 60

Pagan, F.A. (1970) Cage culture of Tilapia. FAO Aquaculture Bull., 3( 1) 6.

Pruginin, Y. and Kanyike, E.S. (1965) Monosex culture of Tilapia through hybridization. SRTC
Symposium on Fish Farming, Nairobi, 1- 3.
Sarig, S. (1970a) Fisheries and fish culture in Israel in 1969. Bamidgeh 22(3) 51-68.
Sarig, S. (1970b) Winter storage of Tilapia. FAO Fish BUN., 2(2) 8.
Semakulu, S.N. and Makoro, J.T. (1968) The culture of Tilapia species in Uganda. FAO (Food Agr.
Organ. UN.) Fish. Rep., 44( 2) 161- 164.
Shelbourne, J.E. (1970) Marine fish cultivation: priorities and progress in Britain. Marine Aqui-
culture. Oregon State University Press, 15536.
Shell, E.W. (1968) Monosex culture of male Tilapia nilotica Linnaeus in ponds stocked at three
rates. FAO (Food Agr. Organ. U.N.) Fish. Rep., 44(4) 353-356.
Steinlitz, H. (1954) The distribution and evolution of the freshwater fishes of Palestine. Istanbul
Univ. Fen. Fak. Hidrobiologi B., I(4) 225.
Swingle, H.S. (1960) Comparative evaluation of two tilapias as pondfishes in Alabama. Trans.
Amer. Fish. Sot., 89(2) 142-148.
Talbot, F.H. and Newell, B.S. (1957) A preliminary note on the breeding and growth of Tilapia in
marine fish ponds in Zanzibar island. E. Afr. Agr. Forest J., 22, 118-121.
Vaas, K.F. and Hofstede, A.E. (1952) Studies on Tikzpia mossambica Peters (ikan mudjair) in
Indonesia. Pember. Ba@i Penjel. Perik. Darat., 1, l-88.
Verygin, B.V. (1963) The problem of the biological improvement of the bodies of water in the
cooling water systems of thermal power stations and their fisheries utilization. In Problemy
rybokhozyaystvennogo ispolzovaniya rastitelnoyanykh ryb v vodoyemakh SSSR (Problems of
the fisheries utilization of phytophagous fishes in bodies of water in the USSR). Ashkabad.
Whitehead, P.J. (1962) The re_lationship between TiZapia n&a (Gunther) and T. mossambica Peters
in the eastern rivers of Kenya. Proc. Zool. Sot., Lond., 138, Pt. 4, 605-637.
Worthington, E.B. (1932) A report on the fisheries of Uganda. Crown Agents, Land., l-88.
Yashuov (Wirszubsky), A. (1956) Report on the growth of Tilapia galikzea Artedi at the Fish
Culture Research Station - Dor. Bamidgeh, 8(2) 31-34.
Yashuov, A. (1960) Effect of low temperatures on Tilapia nilotica and Tilapia galilaea. Bamtdgeh,
12(3) 62-66.
Yashuov, A. and Chervinski, J. (1960) Evaluation of the various food items in diet of T. nilotica.
Bamidgeh., 12(3) 71-78.
Yashuov, A. and Chervinski, J. (1961) The food of Tilapta nilotica in ponds of the Fish Culture
Research Station at Dor. Bamtdgeh 13(2) 33- 39.