45
Elsevier Publishing Company, Amsterdam - Printed in the Netherlands
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
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.
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)
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
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
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).
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
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
DISCUSSION
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
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