ENVIRONMENTAL FACTORS INFLUENCING SPAWNING

Spawning is the culmination of the reproductive cycle and thus the factors which
influence reproductive maturity would indirectly or directly influence spawning for
which spawners sexual maturity is the basic pre-requisite. Adequate and proper diet,
suitable temperature/photoperiod, adequate living space and oxygen are well known
factors which effect sexual maturity. Yet given all these, certain species, though sexually
mature, do not spawn till proper environmental stimuli particularly temperature or
photoperiod or rainfall are fulfilled.

Fish such as medaka, Cryzias latipes, the honmoroko, Gnathopegon caerulescous, etc.
spawn during the long day season from spring to summer and it has been reported that
these species could be matured and spawned by means of increasing the illuminated
period (Yoshioka, 1963, 1966, 1970; Hibiya et al., 1972, 1973; Harrington, 1950, 1957).
Whereas, salmonids which spawn in the autumn when the diurnal illumination is short,
behave in the opposite way and were induced to spawn earlier by shortening the
illuminated period (Corson, 1955; Nomura, 1962; Henderson, 1963). It is interesting to
note that in case of Oryzias, light greatly influences spawning. They spawn before dawn
and when day and night effects were artificially altered spawning occurred at the
beginning of the artificially lit period after a few days. Yoshioka (1963) inhibited
spawning in Oryzias by keeping them in darkness and then provided them with various
amounts of artificial light. He found that in illumination greater than 5 lux all the fish
which were ovulating spawned but in illumination less than one lux some spawned and
some did not. Spawning was also induced by suddenly subjecting individuals which were
already well lit to a very strong light (more than 5000 lux) and all ovulating individual
spawned. This confirms that the occaional spawning of Cryzias during the day or in the
evening is the result of changes in illumination. Yet, the fish will not mature at a
temperature below 10°C, however, favourable the illumination conditions may be and so
is the case with minnow which will not spawn below 7°C (Bullough, 1939). The gold fish
will not spawn at any season unless the temperature is above 14°C. In contrast brook
trout (Salvelinus fontinalis) raised under natural illumination showed no influence of
temperature on maturation even with a temperature difference of 8.5°C to 16°C,
maturation was effected only when there was artificial change between long-day and
short-day period of illumination. Thus, it appears that both light and temperature are
limiting factors in maturation and spawning in certain species but there are species in
which temperature alone is the main factor in inducing maturation and illumination has
no role of importance. Shehadeh (1976) remarks that since most physiological functions
in poikilotherms are temperature dependent, photoperiodism seems to be temperature
sensitive in a majority of the species examined. However, certain factors may become
limiting in one situation while other factors may become limiting even for the same
species in different agroclimatic conditions.

In case of major carps such as Catla catla, Labeo rohita etc. monsoon rain has a
conducive effect on their breeding whether naturally breeding in wild or by
hypophysation. Therefore, factors associated with rain such as temperature, turbidity, pH,
O2 have been correlated to influence spawning of the carps but no critical factor was
assigned which was responsible for triggering spawning in nature or in induced breeding.
It has been observed that following injection of the pituitary extract fraction containing
gonadotropin (Sinha, 1969), the gonad of the recipient spawner always became
translucent and showed maximum percent of loss of weight after drying. Those fish
which had commenced ovulating after getting the injection of fish pituitary gonadotropin
lost 73.5% as compared to 63% in control. Males injected with this fraction showed the
maximum percentage loss of weight of the semen after drying, much higher than those of
control. The volume of semen increased to about twice the volume collected before the
treatment. Major electrolytic ions such as sodium, potassium, calcium and magnesium
and chloride in the blood and semen, showed that following injection of the pituitary
extracts the cl level increases significantly in the blood of the female. As with the female,
male serum cl significantly increased after the treatment but the level went down
significantly in the seminal fluid differing from serum cl (Sinha, 1971) indicating the
electrolytic concentration in the blood and dilution in the semen is because of the gonadal
hydration which appears to be a pre-requisite for spawning of those physiologically ripe
carps. In 1974, a hypothesis regarding spawning of carps suggested that during induced
Dr.C.V.Narasimha Murthy, JBDC, KAVALI, B.Sc., Zoology Notes 2010.
breeding gonadal hydration was achieved by the injection of the additional amount of
exogenous pituitary material, whereas in natural spawning in riverine condition a sudden
drop in the electrolytic level in the water by flood or heavy monsoon rain induced
hydration of the fish as a whole and consequently the gonad, and those fish which were
ripe in condition started spawning then in nature. It is common knowledge that when fish
go from water of higher electrolytic level to lower level, for example, from sea water to
freshwater, they gain water and thus the weight and reverse is true when they come from
lower electrolytic level to higher level. In rivers and reservoirs, there is a marked increase
of electrolytic level during mummer months and their level decreases suddenly after
monsoon rain.

Further, the total water content of the ovary, the appearance of fluid in the space between
the follicular cells, the loosening of the adhesion between the follicular cells and the egg
and increase in diameter of egg suggest that these morphological changes accompanying
ovulation are proceded by a remarkable absorption of water into the ovary and ovarian
egg. Whether the condition prevailing in ponds during spawning season do favour such
absorption of water or not needs investigation, especially because these carps do mature
in ponds but do not spawn there. The water in these ponds is perhaps not that much
diluted with rain or flood water as that of the rivers where they normally breed. In this
context it is important to note that the males do spermiate in ponds but the female do not
ovulate there. This may be because the sperm have no mechanical barrier to water
movement unlike the chorion which may mechanically oppose or support the entry of
ions and water to the egg for its final maturation (Sinha, 1980).

ENDOCRINCLOGICAL CONTROL OF SPAWNING

Wilhemi et al (1955) have shown that marked flexure and vibration of the body, which
they called a ‘spawning reflex’, could be induced in Fundulus hetroclitus by the injection
of highly purified vasopressin and oxytocin and by synthetic oxytocin. However, it is
difficult to decide whether such reflex is the true spawning reflex, since the experiments
were not planned to show the emittance of the genital products following the reflex.
Egami (1959) and Ishii (1961) claimed that neurohypophysial hormones were effective in
inducing spawning. However, Sinha (1971) found negative correlation between the
presence of isotocin/arginine vasotocin and spawning.

It is sometimes said that a two-component control system with two gonadotropins may be
needed: one for cogenesis and vitellogenesis and the other for ovulation. Thus, in case of
these carps which do not normally ovulate in ponds, though achieving gonadal
development up to fourth stage, it may be argued that they may not contain the other
gonadotropin which is responsible for ovulation. Yet, it is interesting to note that pituitary
of donor carps inhabiting ponds is effective on the recipient, which also inhabit the
ponds. Thus, if donor pituitary is lacking the ovulating gonadotropin it would not have
been effective in inducing spawning of others. In this regard, it is essential to point it out
that the same fraction containing pituitary gonadotropin (Sinha, 1969) induced courtship
behaviour as well as ovulation of viable eggs in the carps (Sinha, 1971) and also
maintained and initiated all the reproductive process in the hypophysectomised female
catfish indicating that some gonadotropin does both the functions in the carps
(Sundararaj, et al., 1972).

Further investigations indicated that qualitatively there appears to be no difference in the

gonadotropin of a free spawner like common carp and refractory species like Indian
major carp or Chinese carps. However, when quantitatively assessed it has been found
out that the gonadotropin content of the pituitary gland of common carp and Puntius
gonionotus is considerably more than that in the pituitary glands of the other carps which
do not breed in confined water presumably due to reduced accumulation of gonadotropin
in the pituitary gland and therefore the additional amount of exogenous pituitary material
is needed to induce those refractory fishes to spawn in confined water. The reason for
reduced accumulation of the gonadotropin needs thorough endocrinological investigation
in terms of ecological parameters of the environment and the reproductive biology of the
fish. However, researches are actively going on to induce spawning with the intervention
at the hypothalamic-pituitary interface through the administration of LH-RH. The
introduction of the synthetic LH-RH (luteinizing hormone-releasing hormone) for
Dr.C.V.Narasimha Murthy, JBDC, KAVALI, B.Sc., Zoology Notes 2010.
ovulation which causes an elevated plasma level of gonadotropin hormone in the
common carp, Cyprinus caroio (Breton and Weill, 1973) appears to be promising. Its use
along with fish pituitary extract is giving excellent results in carp breeding in China.

Many have started experimenting ovulation with sex steroids with conflicting results.
However, it can be fairly presumed that with the present knowledge of the spawning of
the fish much better results of induced spawning of cultivable species could be achieved
if ripe parent stock is available and they are subjected to hypophysation with simulation
of natural condition for the spawning.

Dr.C.V.Narasimha Murthy, JBDC, KAVALI, B.Sc., Zoology Notes 2010.