2.1. breeding - shodhgangashodhganga.inflibnet.ac.in/bitstream/10603/95793/9/09_chapter 2.pdf ·...
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Review of Literature
10
tudy on feeding ecology, maturation and spawning of fish are
aimed at understanding and predicting the changes in
composition of the population, which are expected to undergo
fluctuations from time to time. Further such studies are also useful
for the judicial management of the fishery. A number of researchers
have worked on various ecological and biological aspects including
feeding, population dynamics, breeding- maturation, spawning,
fecundity etc. from time to time.
2.1. Breeding
Mathews (1938) studied seasonal cycle of the gonads of Fundulus and
observed that in case of Fundulus, the follicular epithelium, after the
discharge of the ovum, shows hypertrophy.
James (1946) studied the development of sex cells and hatching of
young fishes in the large mouth bass (Huro salmoides) and the blue gill
(Lepomis macrochinus rafinesque) and found that structural and histological
change in fishes correlated with seasonal changes in the gonads.
S
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Jones (1946) has worked on the fecundity of different fishes in Indian
waters. In Notopterus chitala, the number of eggs varied from 300 to 500,
while in Carracius auratus there were 2,000 to 70,000 eggs depending on the
size of the fish.
Mathews (1950) found as many as six million ova in the ovary while
studying the fecundity of Cetorhinus maximus with an average size of 61
inches long of 54 pounds weight, contained as many as 28,361,000 eggs.
Alikhuni (1952) also worked on the aspect of fecundity and calculated
the number of ova in Ophicephalus striatus at 2997 in an ovary weighing 2.7
grams; and 20,070 eggs in another ovary weighing 30.0 grams.
Ghosh and Kar (1952) studied the reproduction of Heteropneustus
fossilis and found that the ovary attains maximum weight during the summer
months, after which it recedes to a quiescent phase. They divided the ovarian
activities into three phases: - 1. Preparatory phase, here a gradual increase in
the weight of the gonad takes place followed by initiation of cytosomal
differentiation. 2. Active phase, maturing maximum ovarian weight with ova
becoming ripe, after which the ovulation occurs. 3. Quiesent phase, here the
ovaries gradually recede to their infantile condition.
Marshal and Lofts (1956) made investigations on the testes of teleosts
and reported that the testes can be differentiated into two basic structural
types. In the first type (Gasterosteus, Tilapia) the lobules are formed by
fibrous connective tissue, and the interstitial cells are located between the
lobules. In the second type (Esox, Labeo) the lobules are bounded by cells. As
there are no interstitial cells, the lobule boundary cells perform the function of
the leydig cells of other vertebrates. This conclusion was drawn on the
discovery made by them i.e., the lobule boundary cells of Esox under go a
seasonal production of cholesterol- positive lipids in a pattern much similar to
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that of avian leydig cells. Accumulation of lipid inclusions within the lobule
boundary cells was found during the three months proceeding spawning. At
the end of spawning the inclusions were gradually discharged into the lumina
of the tubules. It was found that the lobule boundary cells were derived from
modified fibroblasts, and show seasonal accumulation of cholesterol –
positive lipids similar to the condition in the interstitial cells.
Bagenal (1957) investigated the fecundity of Long Rough Dab
(Hippoglossoides platessoides) and found it to be related to weight linearly, to
the length at less than unity.
Gokhale (1957) studied the seasonal histological changes in the gonads
of the whiting (Gadus merlangus) and the Norway Pout (Gadus esmarkii). He
reported that the increase in the size of the testes is due to the increase in the
number of spermatogonia and sperms. It was also revealed that after spawning
no spermatogonia was left over. Formation of new spermatogonia takes place
in the month between January and May, and then the fish spawns in May and
June, thereafter gonads remain small until the following January.
Rounsefell (1957) investigated the fecundity, as the number of ripening
eggs found in the ovaries prior to spawning of North American Salmonidae.
Nair (1958) made investigations relating to the seasonal changes in the
oocytes of Hilsa ilisha. He worked on the histological changes in the ovary of
the fish and found that very low and very high temperature have an inhibitory
effect on the oogenetic activities.
Hodder (1963) studied the fecundity of Grand band haddock, and
stated that fecundity increased at a rate proportional to about the fifth power
of the body length and to the square of the age.
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Qasim and Qayum (1963) studied the fecundities of Ophicephalus
punctatus, Callichrous pabda and Mystus vittatus and reported 23,000 to
29,600 eggs in Ophicephalus punctatus, 25,000 to 40,000 eggs in Callichrous
pabda and much higher number of eggs in Mystus vittatus.
Das (1964) also worked on the fecundity of four fresh water fish
species found in the Indian waters and the average number of eggs found
were Wallagonia attu (42,000), Mystus bleeker (16,000), Gonialosa
manninna (13,000) and Rita rita (12,000).
Das and Malhotra (1964) investigated the occurrence of corpus luteum
in Schizothorax niger and revealed that the discharged follicle undergoes
hypertrophy to form a solid corpus luteum. The luteal cells are derived from
the granulosa, with theca retaining original form, undergoing only an increase
in size. The hypertrophy of the cells is due to the response of the cells to LH
factor in gonadotrophin.
Higham and Nicholson (1962) made studies on the spawning of
Brevoortia tyrannus in Atlantic regions and reported that the fish spawns
during summer in the north Atlantic area; in spring and again in autumn in the
middle Atlantic; and during winter in the south Atlantic area.
Pitt (1964) found that fecundity in American Plaice is directly related
to the length, weight, and to a little extent the age of the fish.
Sivaprakasam (1965) made investigations on the maturation stages
spawning habits and occurrence of juveniles of the Brown pomfret,
Parastromateus niger (Bloch). The spawning habits were studied by the ova-
diameter frequency method, and the distribution of various stages of maturity
in time, and the occurrence of juveniles. Spawning occurred during July to
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October. The study on sex ratio in various size groups during different months
was also worked out.
Ahsan (1966) studied the cyclical changes in the activity of the testes
of Couesius plembeus, from north temperature latitude, by histological and
histochemical methods, with a quantitative evaluation of the different
spermatogenetic stages. He divided the testicular cycle into five stages in
relation to the changing environmental factors, comparable to the five stages
in the ovaries. He further reported that the annual cycle consists of a
prolonged mitotic phase during the fall, and a shorter meiotic phase during the
spring.
Lofts and Pickford (1966) demonstrated that androgens elicit a
spermatokinetic response in Fundulus heteroclitus, as marked stimulation of
spermatogenetic activity results after administration of methyl testosterone to
hypophysectomised and sexually regressed specimens. It has also been
suggested (based on the spermatogenetic condition in the gonad and the
nuptial colouration as index of androgen production) that the endogenous
androgen has a role in accelerating post-spermatogonial development of germ
cells in the testicular cycle of Fundulus.
Chan and Philips (1967) investigated the seasonal changes in the
distribution of gonadial lipids and spermatogenetic tissue in the male phase of
Monopterus albus, which undergoes cyclical changes after the reversal of sex
from female to male. The seasonally variable events include a prenuptial
accumulation of cholesterol - positive lipid droplets in the cytoplasm of the
interstitial cells, when spermatogenetic activity is resumed in late February
and early march. There occurs a sudden depletion of interstitial lipids during
the breeding season in June at a time when male animals exhibit active nuptial
behaviour.
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Malhotra (1967) gave the relation between the ovarian cycle and
feeding of Schizothorax and Botia, and showed that during the maturation of
gonads, feeding slackened in these fishes.
May (1967) studied the fecundity of Atlantic Cod from the New
Foundland and Labrador area in relation to other parameters like length,
weight, and age of the fish.
Moser (1967) while working on the seasonal histological changes in
the gonads of Sebastoides paucispinis, found that the ovaries undergo greater
seasonal changes as compared to those of the testes. He found that the ovaries
are small during the post –extrusion period from March to June. Vitllogenesis
begins in July and continues through out the summer, this activity causes the
ovaries to enlarge and become opaque yellow. Ovulation probably precedes
fertilization since spermatozoa were not found within Pre-ovulatory or post
ovulatory follicles. After ovulation the regression of the follicular epithelium
takes place whereas capillary beds remain intact, resulting in the mechanism
for foetal – material exchange of gas and nitrogenous waste.
Hussain and Das (1968) studied the effects of mammalian hormones
on the gonads of Schizothorax esocinus (Heckel). The hormone (bi-anayI.
CC.hormone and 0.2% saline H2O) brought premature spawning in
Schizothorax esocinus. Duogynon for the female and Testoviron for the male
was found most effective in spawning of fish, prematurely.
Hussain and Das (1968) studied the effects of mammalian hormones
on the gonads of Cyprinus carpio specularis (Mirror carp) and it was
confirmed that the mature ova and sperms were obtained pre-maturely by
induced spawning after employment of mammalian hormones. Macroscopic
examination showed a rapid maturation from stage III to IV and V.
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Malhotra (1968) investigated seasonal variations in the morphology of
the ovaries of Botia birdi and ascertained that its ovary recovered after
spawning. It remained in stage I (Immature) upto end of April (Diapause) and
enters stage II during May. The increase in the ratio of the weight of ovary to
that of the fish was sudden restricted from May to mid of August.
Bagenal (1969) experimentally determined the relationship between
the food supply and the fecundity of brown trout, Salmo trutta, and reported
that a reduced diet led to a lower percentage of maturity and lower fecundity.
However, something other than diet responsible for the predetermined egg –
size was not ruled out.
Das (1969) and Malhotra (1968, 1970) reported on the seasonal
variation in the morphology of the ovaries of Bottia birdii and Schizothorax
niger.
Das and Singh (1969) studied the comparative fecundity of Kashmir
fishes and found that Barbus conchonius had the highest comparative
fecundity (4,569) but in total fecundity it stood 4th
among the five fishes.
Similarly, Labeo dero was getting rarer due to its low comparative fecundity
(422) and Nemachilus was going up.
Singh and Das (1969) worked on the morphology of the ovaries of
Crossochilus lateus punjabensis (Heckel) and reported that the ovaries
underwent vitellogensis from August to December, immediately after
spawning (May-end or June). The gonado-somatic index showed a gradual
change with change in the weight and maturity of the ovary due to the
external environmental conditions such as food, water, temperature, day light
and shoaling.
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Stephon Dadzic (1969) studied the cyclical changes in the testes of
Tilapia mosambica. His investigation revealed that the immature virgin testes
contain primary germ cells and primary spermatogonia enclosed within a
connective tissue. A pre spawning testes is packed with sperms, with other
cells at all stages of spermatogenesis. After spawning the unexpelled
spermatozoa undergo phagocytosis and resorption like that of atretic ova.
Hyder (1970) while working on the gonadial and reproductive pattern
in Tilapia leucosticta, found that the ovarian development was lowest
between the months July –August. In the month of October slighting but
fluctuating breeding activity and gonadial development is resumed following
increased sunlight and temperature from September. The period December -
February is marked by sustained maximum environmental factors and is also
the period of rising gonadial development in most females. Sustained heavy
rainfall on the other hand seem to check breeding.
Bhatt (1971) studied the Biology of fresh water fish Mystus vittatus
(Bloch) and reported that the length frequency distribution of Mystus vittatus
gave evidence of 2 to 3 modes. At maturity the males are 8.0 cm. and the
females 8.5 cm long. Seasonal changes in the gonad maturity revealed that
this fish spawns from August to September. The variation in K-values seems
to be correlated with feeding intensity. Its main food items include insect
larvae, copepods, daphnids, and rotifers etc. The food items showed little
variation from season to season.
Barlow and Victor (1972) made investigations on the ovarian condition
of the long jaw goby Gillichthys mirabistis, changing with the season in the
Salton sea. The protracted spawning period extends from Dec. to Jan. upto
June. Each female spawns more than once, and the population does not spawn
synchronously. Regress of the ovaries start by late June, and recrudescence
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begins in September or October. The timing of ovarian development at
different localities in the lake varies slightly. Ovarian cycling appears to be
correlated with temperature changes.
Malhotra et al. (1978) studied the ovarian cycle and spawning season
of Ophiocephalus punctatus and reported that the oocyte showed six well
differentiated stages of maturation, the fish had prolonged spawning season
extending from May to August and was bimodal in character. The stages in
the formation of the yolk nucleus of Balbaini was described and the origin of
the cytoplasmic inclusion was also discussed.
Vass et al. (1980) made observations on the breeding behaviour of
Schizothorax niger and reported that the fish breeds in shallow parts of Dal
Lake. Breeding extended over a period of two months from the first week of
March till end of April.
Toshio Okazaki (1984) studied age composition, growth, sex ratio and
gonad development of the steelhead trout and the Kamchatkan trout in the
north Pacific. The differentiation of mature and immature forms of fish was
estimated from the seasonal changes in the gonad weight distribution,
indicating high concentrations of immature fish. The migration patterns of
both the fish was also discussed.
Lund et al. (1990) studied interpopulation variation in reproductive
traits of anadromus female brown trout, Salmo trutta and found a positive
significant correlation between fish length and fecundity in all populations
and between fish length and egg diameter in five populations. The ratio of
gonadial weight to somatic weight varied significantly among populations.
Seishi and Kiriyama (1993) worked on the embryonic, larval and
juvenile development of the Labrid fish, Halichoeres poecilopterus. The eggs
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measuring 0.60-0.70mm in diameter, were pelagic and spherical. Hatching
occurred 18h 48 min after spawning. The newly hatched larvae measured
1.46-1.70mmTL. The yolk was completely absorbed after three days at 2.52-
2.72mmTL.
Mihelakakis and Kitajima (1995) studied the spawning of the Silver
sea beam, Sparus sarba, in captivity at ambient temperature and photoperiod.
They divided brood stock into group A and B of 6 and 7 individuals
respectively. Spawning lasted 65 days from April 8 to June 11 in group A,
and 43 days from April 10 to May 22 in group B, with in the temperature
range between 13.5 and 21.3oC. The total number of eggs released during the
spawning period was estimated at 18.1 million in-group A and 10.8 million in
group B.
Grier and Taylor (1998) investigated testicular maturation and
regression in the common snook and reported that during the annual
reproductive cycle, the lobular testes of Centropomus undecimalis undergoes
height, width, and morphological changes which reflect five reproductive
classes: regressed, early, mid, and late maturation; and regression. Differences
between continuous and discontinuous germinal epithelia, are used to
distinguish these five classes. Throughout the year, periodic acid –schiff-
positive macro – melanophore centres and PAS- positive granulocytes were
observed in the testes. They were most abundant after the breeding season,
and might be involved in tissue degradation.
Dadzie et al. (2000) made seasonal observations on the distribution of
maturity stages and variations in fluctuations in the gonadosomatic index
(GSI) of the silver pomfret, Pampus argenteus (Euphrasen). There were two
spawning peaks, the first in May and the second in August. Variations in GSI
relative to fish length indicated that females and males were most fecund at
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about 24.5-26.4cm and 20.5-2204cm length classes, respectively. The males
matured earlier than females.
Yildirin et al. (2000) studied some reproductive characteristics of
Copoeta tinca living in the oltic stream of Coruh basin. The spawning was
found to occur in the month of May and July and fecundity was related to fork
length, total length, total weight, age and gonad weight.
Parsley et al. (2000) observed the spawning areas of white sturgeon
Acipenser transmontanus in the lower snake river from the mouth to the lower
Granite Dam. Spawning location were determined by the collection of
fertilized eggs in artificial substrates or in plankton nest.
Morita and Morita (2000) studied individual growth and maturation
histories, age and size at maturity of resident white spotted charr, Salvinus
leucomaenis and found that slow growing fish reached sexual maturity not
only at an older age but also at a smaller size then fast growing fish. Such
patterns results from adaptive phenotypic plasticity that depends on individual
growth conditions.
Holcombe et al. (2000) conducted reproductive studies with brook
trout Salvinus fontinalis at times other than the normal fall of spawning
period. These studies demonstrated that photoperiod alterations could be used
to manipulate brook trout reproductive cycles for the purpose of conducting
studies at times other than the fall.
Liley et al. (2002) examined aspects of the fertilization dynamics of
rainbow trout (Oncorhynchus mykiss) that may play a role in determining
reproductive success of males of different age and status competing for
spawning. There was no differences in the gonadosomatic indices and relative
yields of milt of adult and precocious male rainbow trout collected from a
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wild population. It was also examined that short gamete longevity and the
speed with which fertilization occurs indicate that the timing and position of
sperm release may play a critical role in determining the reproductive success
of males in competition for spawning with a single female.
Brandao et al. (2003) while investigating the ovary maturation stages
and oocyte features in three species of the Neotropical fish Hemiodus and
reported that there was group synchronous oocyte development in H.
microlepis, H. ternetzi and H. unimaculatus. It was also reported that the
remarkable thickness of zona radiata layer and the large size of vitellogenic
oocytes of H. ternetzi distinguished this species from the other.
Kumar et al. (2003) investigated annual reproductive cycle of male
rohu, Labeo rohita (Ham.) and reported that its testicular development under
ambient environmental conditions was associated with increasing day length
and temperature, whereas testicular maturity and spermatogenesis during
spawning phase seem to be correlated with the lowering of water temperature.
A few lobules with spermatozoa and spermatids were observed in preparatory
phase. The peak spermatogenesis having active interstitial leydig cells was
recorded during pre-spawning phase. Decreased interlobular space with some
inactive interstitial cells was noticed during post – spawning phase.
Narejo et al. (2003) worked on the reproductive biology of an air –
breathing fresh water mud eel. Monepterus cuchia (Hamilton) with respect to
sex- dimorphism, gonadosomatic index, ova diameter and fecundity. Peaks of
gonadosomatic index (GSI) recorded during April to June for female
(7.521.51) and male (5.501.15) indicated that the fish has only one
breeding season during summer. The number of ova per gram body weight
was 4.61 and the number of ova present per gram of ovary weight was 74.27.
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Chellapa et al. (2004) while studying reproductive cycle of male
Amazonian red discus, examined the morphological and histological features
of the testicular development in males. Fish in different stages of gonadal
development have been used in the study and their gonads weighed and
examined to observe the maturation stage. Macroscopic observations showed
the immature, maturing, mature and partially spent stages of the testicular
development. Microscopic examination of the testes revealed five different
stages and phases of testicular development.
Zacharia and Kakati (2004) while studying the effect of serotonin (5-
hydroxytryptamine, 5-HT) on the histological structure of ovaries of the
banana prawn, Fenneropenaeus merguiensis concluded that the
gonadosomatic index (GSI) increased significantly in the treated females,
with a majority of oocytes reaching early vitellogenic stage, indicating that
ovarian maturation could be induced and accelerated by 5-HT.
Bisht et al. (2005) carried out the studies on maturation biology and
spawning ecology of Schizothorax plagiostomus and concluded that the fish
spawns in shallow areas of side water on the pebbly substratum that was rich
in algal carpets and preferred a temperature range between 12.5 to 16.6oC, pH
range of 7.8-8.3 and higher dissolved O2 contents (10-13 ppm). The male fish
attained maturity little earlier (33.8cm) than the female fish (35.2cm) and
spawning was twice a year during April-May and October-November.
Ingram et al. (2005) studied the induced spawning, larval development
and rearing of two indigenous Malaysian mahseer, Tor tambriod douronensis
and found that the exploitation of natural stocks because of high demand and
the deteriorating environmental conditions marked decline of mahseers in the
wild. A captive breeding programme was investigated to attempt to propagate
these two species. Pre-treatement of fish with ovaplant (28-68 weeks before
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spawning induction) greatly improved the success rate of spawning induction.
Use of formalin baths improved hatching fungal infections. Juveniles were
reared in static gap ponds and reached 142- 179g (max 270g) in six weeks.
Madan (2005) investigated spawning biology of snow trout, and
noticed that species spawns twice in a year (July-October and January-
February) as was evident from the occurrence of ripe fishes, high GSI values
during the mentioned period. Besides it was also found that fecundity ranged
from 2248 to 2726 in fishes of 160 -245 mm TL and 40-110gm weight.
Kumar (2005) studied the maturation and spawning period of Indian
scad Decapterus russelli and reported that its spawning period was prolonged
extending from March to December with peak in April – May and September.
Mature fishes with ripe and spent gonads were noticed from March to
December. The relative condition faction factor (Kn) and gonad somatic
index (GSI) values ranged between 0.93-0.98 and 0.75-3.85 respectively, with
peaks in March – May. The fecundity varied from 33,159 to 1,96,392 and the
number of ova per gram ovary varied from 8,270 to 28,300 increasing with
increase in length of the fish. The females dominated over males in the
population.
Musallam et al. (2006) made observations on the fecundity and
gonadosomatic index (GSI) of about 70 mature Omani Indian oil sardines.
The fecundity ranged between 5500 and 60000 with an average of 19000.
Linear regression was found between F and fish total length (TL); fish weight
(FW) and gonad weight (GW). The results of the correlation regression were
0.966437; 0.796241 and 0.877952, respectively.
Tarkan (2006) investigated the reproductive activities of two dominant
fish species, roach Rutilus rutilus and rudd Scardinius erythrophthalmus from
August to July in an oligotrophic lake. A total of 907 roach and 593 rudd
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individuals were examined. The ratio of males to females was 1.3:1 and 1.2:1
for roach and rudd, respectively. The average condition factor K was 1.21 for
rudd and 1.14 for roach. Absolute fecundity varied from 7829 to 95,387 eggs
between ages 3 and 9 for roach and from 1807 to 35,629 eggs between ages 1
and 7 for rudd.
Yamaguchi et al. (2006) studied the reproductive biology of white
Croaker by measurement of gonadosomatic indices (GSI) and histological
examination of ovaries. Ovaries were classified into five maturity stages:
immature\resting stage; developing stage; maturing stage; ripe\spawning
stage; spent stage. The spawning season lasted from April to September but
most activity was observed from May to August. Age at sexual maturity was
assumed to be 1 year. Croaker was found to spawn more than once in a single
spawning season.
2.2. Food and Feeding
Lang ford and Martin (1940) gave importance to the study of seasonal
variations in the food of fishes and its changes with respect to growth and
condition of gonads.
Hiatt (1947) studied the food chains and food cycles in HAWAIIAN
pond fishes, Mugil cephalus and milk fish (Chanos chanos). The biotic
interaction between the fishes and their ponds was also discussed.
Smith (1947) gave the differences between the food present in the
alimentary canal of white – perch and to its availability in nature.
Pillay (1953) worked on the food, feeding habits and alimentary tract
of the gray mullet, Mugil tade Forsk and illucidated that the length of the
alimentary canal varies with the nature of the food consumed in different
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environments. Where algae formed the predominant food, the alimentary
canal is found to be longer with a well developed pyloric stomach.
Vasisht (1959) made investigations regarding the food and feeding
habits of Labeo calbasu, Labeo rohita, Catla catla and Cirrhina mrigala in
Punjab, and reported that the organs of feeding vary in relation to food and
the food is the controlling factor in local distribution of fishes, while the fish
food in nature is dependent upon the depth, nature of bottom, currents and the
physical condition of the waters.
Chandy and George (1960) studied alimentary canal of the milkfish
Chanos chanos in relation to its food and feeding habits.
Moitra (1963) classified fishes according to their feeding habits into
herbivores, omnivores and carnivores and also into surface feeders, mid
feeders, column feeders and bottom feeders; after studying seasonal variation
in the food components of 24 fresh water fishes of U.P.
Qayum and Qasim (1964) studied the food habits of Ophicephalus
punctatus Bl. and found that the food habits of adults, immature and larval
forms varied accordingly. The larval forms feed entirely on planktonic
organisms, and the adult is completely carnivorous and predatory in nature.
Saigal (1964) worked on the biology of the catfish Mystus seenghala
(osteobagarus) of the Ganga river system. The adult was found to be
carnivorous, feeding on teleosts and the juvenile fish to be mostly
insectivorous.
Saxena and Bakshi (1964) studied the anatomy of the alimentary canal
of Botia birdi (Choudhuri) and differentiated it into a, bucco-pharyngeal
cavity, oesophagus, stomach, intestine and rectum. The intestine was long
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tube having three loops and liver was large having three well-defined lobes.
Pancreas was diffused.
Kamal (1966) investigated the gut contents of the juveniles of Labeo
rohita and reported that its principal food item was insect larvae, showing its
food preference as of a typical larvicidal fish.
Das (1967) made investigations on the ecology of fishes and reported
that the plankton and bottom fauna comprising of Pedonic, Limnetic and
paralimnic fauna, are important constituents of fish food He further added that
the percentage of herivorous fishes increase while proceeding from South to
the North in India.
Das and Nath (1967) studied the comparative morphology of the
alimentary tract of fresh water fishes and found significant variations in the
morphology of the alimentary tracts of different fish species that can be
correlated with the differences in the nature of food taken by these species and
with their feeding habits. The presence or absence of stomach, length of the
intestine etc. was also discussed in herbivorous, omnivorous and carnivorous
fishes.
Kamal (1967) also studied the food and alimentary canal in Labeo
rohita (Ham.) whose juveniles feed on Zooplankton and in Cirrhina mrigala
whose fry feed on phytoplankton and zooplankton, the main part of the food
was found to that of cladocerans .
Moitra and Bhowmick (1967) while studying the functional histology
of the alimentary canal of the young Catla catla reported that the feeding
changed from the young to the adult stage, with respect to development of
feeding apparatus.
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Subla (1967) studied the structure of the feeding apparatus of four
fishes: Cyprinous carpio specularis, Oreinus plagiostomus, Schizothorax
esocinus and Nemachilus stoticzkae. The first three being herbivorous with
edentulous buccal cavity, having well developed tongue, filter feeding
mechanism formed by interdigitating gill –rakers.
Das and Subla (1969) found that feeding mechanism in fishes varied
highly depending on the type of food whether herbivores, carnivores and
omnivores and mode of feeding. Besides they gave an illustrated account of
the mechanisms of jaws and the associated skull bones, along with the
muscles.
Jan and Das (1971) studied in detail the food of some Kashmir fishes
and found a definite periodicity in the intake of food in different seasons of
these fishes, which depends on the availability of the food material in nature.
The food spectra of commercially important fishes was extremely useful for
determination of niches in fish biome.
De Silva et al. (1977-80) reported that the diet of the fish varies in
response to increase in size in Puntius imaculatus and P. tithya.
Chatterji et al. (1978) studied the food and feeding habits of Labeo
gonius (Ham.) and found that the fish feeds mainly on phytoplanktonic
organisms e.g. diatoms, green algae and decayed organic matter. Feeding was
found to be maximum during post – monsoon and winter months (October –
February) and low during post –winter and monsoon months (March –
August). Maturation of gonads also adversely affected the feeding intensity in
both the sexes.
Sharma and Tandon (1982) worked on the food and feeding habits of
Mastacembelus armatus (lacepede), Oreineus richardsonii (gray) and
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Glyptothorax pectinopterus in order to understand various biological
problems relating to growth and behaviour. It was concluded that
Mastacembelus armatus was omnivorous, becoming predominantly
piscivorous in large size groups; Oreineus richardsonii is herbivorous and
Glyptothorax pectinopterus is carnivorous.
Prakash and Lal (1984) studied the food and feeding habits of the
fingerlings and juveniles of Tor putitora inhabiting river Nayar, and
concluded that the fingerlings and juveniles were “monophagic” and “column
feeder”. Inspite of the fact that the RGL values supported its omnivorous
habit, the percental value of insect food item and their occurrence in 5% of the
fishes clearly indicated a “carnivorous” habit.
Sunder et al. (1984) examined the gut contents of Cyprinus carpio
specularis (Linn.) in Dal Lake and found the fish to be a bottom feeder,
chiefly feeding on diatoms, algae, desmids and crustaceans. Decayed organic
matter was also reported in insignificant quantities in the gut of the fish.
Baloni and Tilak (1985) studied food and feeding, sexual dimorphism
and breeding behaviour of Schizothorax richardsonii and reported that the
fish is predominantly herbivorous. Feeding intensity decreased during
spawning period (July to September). The mature males and females can be
well differentiated. Males were smaller in size than females. It breeds in
monsoon (July to early October). The diameter of ova ranged from 2.99-
3.992mm.
Sunder and Subla (1985) investigated food of juveniles of Schizothorax
curvifrons from Bringhi stream in Kashmir and descried its planktophagus
nature of feeding.
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Sunder (1985) while studying Puntius conchonius from Dal lake
catagorised it a herbivorous and illophagic.
Amma and Suryanarayan (1988) carried out their research on
Cyanaglossus punticeps and found it to be a bottom-feeding carnivore, having
a preference for crustaceans and polychaetes. They also reported a clear
relationship existing between feeding breeding and condition of stomach.
Khan (1988) investigated food and feeding habits of afresh water carp,
Cyprinion macrostomus (Heckel) and reported the intensity of feeding and its
variation with season, size and sex etc. The fish was found to be a bottom
feeder. Seasonal variation in the intensity of feeding was higher in maturing
individuals than the ripe ones.
Das and Mishra (1990) worked on the biology of Psettode erumei
(Bloch and Schn) an Indian Hallibut and found it to be highly predacious and
predominantly piscivorous in habit. Molluscus and arthropods only
supplement the diet to some extent.
Das Gupta (1990) studied the food and feeding habits of the Mahseer
Tor tor (Ham.) and found it to be an omnivore.
Dedual and Collier (1995) studied the aspects of juvenile rainbow trout
diet in relation to food supply during summer and found the importance of
diptera in the diet of the juvenile trout during summer. It was further reported
that the stomach fullness index was similar during three periods between
dawn and dusk, indicating the feeding to be continuous.
Casas et al. (1996) Investigated changes in the feeding of Cod (Gadus
morhua) off the Flemish cap, Newfoundland (1989-93) and found that
juvenile cod diet was dominated by crustaceans, mainly hyperiids and
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30
polychaetes, while in adult cod diet the most important prey was fish, mainly
red fish.
Azuma et al. (1998) made research on the feeding habits of large
mouth bass (Micropterus salmoides) in a non-native environment. Regardless
of the size the principal food of the bass consisted of two numerically
dominant gobiids, the trident gobi, Tridentiger obscures and paradise goby,
Rhinogobius giurinus.
Sutela et al. (1998) studied the prey selection and the density of suitale
food for Vendace (coregonus albula) larvae in lake Lentua and found that
vandace larvae preffered certain Rotatoria species, Copepoda, nauplii and
small Bosmina longispina individuals and avoided hard spiny Rotatoria
species and neglected Cyclops, Eudiaptomus and large Bosmina individuals.
Basudha and Vishwanath (1999) recorded percentage composition of
food items in the gut contents of Osteobrama belangeri and found that major
food items in various size groups of fish was different. In juveniles (6-10
cmsTL) zooplankton contents were the highest (51.9%). Aquatic plants
formed the main food items (40-60%) in grown up fishes (10-26 cm TL).
Kurian and Inasu (2001) made a comprehensive study on the food and
feeding habits of Horabagrus brachysoma (Gunther) and revealed that the
fish was predominantly a carnivore, mainly feeding on crustaceans. Further it
was found that the feeding intensity was maximum in the post spawning
period of the fish and stomach was found empty during the spawning season
(June). A significant role of pharyngeal teeth was found in the feeding of live
prey.
Somashekhar et al. (2003) worked on the aquatic weeds incorporated
feed for carp fry and found that the growth of carp fry grown on pistia leaf
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31
based feed (PF) was marginally superior to that obtained with LF (Lemna
based feed) and CF (Conventional feed).
Appa and Sreeramulu (2006) made investigations on the food and
feeding habits of Pseudapocryptes lanceolatus (Bloch and Schneider, 1801).
The diet of males and females did not differ. Qualitative analysis of the diet
composition revealed that the fish was phytoplanktonic bottom feeder.
2.3. Length and Weight Relation
Dixit (1959) investigated the seasonal variations in the length and
weight of the ovaries in Mystus scenghala and Wallogonia attu, and has
described the changes undergone by the ovaries in their length and weight.
Das and Subla (1964) gave description of oral valves of nine fishes of
Kashmir for the first time and reported the absence of the mandibular valve in
about six species of fishes. Besides the crescentric and U-shaped valves
reported by Saxena (1958), a new V-shaped valve and the occurrence of
dorso-ventral muscles joining maxillary and mandibular valves was also
reported.
Johal and Tandon (1983) studied the age, growth and length-weight
relationship of Catla catla and Cirrhina mrigala from Sukhna lake and found
that the total length-scale radius relationship in both species was linear.
Udupa (2003) compared the length-weight relationship in fishes using
dummy variables. Regression analysis was used to fit multiple linear
regression equation of log weight n, log length by including dummy variables.
Tiwari et al. (2004) studied the morphology of the exotic, fresh water,
larvicidal, aquarium fish, Gambusia affinis (Baird and Girard, 1853). The
observations revealed that body form, shape and size acted as the important
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32
secondary sex-determining characteristic. Males were considerably smaller 3-
4cm (1.2 to 1.6 inches), while females were 5-7 cm (2-2.8 inches).
Gerritsen and McGrath (2007) used length-weight relationship of fish
to estimate the biomass of length distributions and indices of condition. They
examined length-weight relationship of 1334 haddok (Melanogrammus
aeglefinus) and 1186 whiting (Merlangius merlangius) collected on a ground
fish survey in the waters around Ireland in 2004. The length-weight regression
showed a significant area effect but no significant difference existed between
the sexes. The condition indices showed a moderate spatial structure for oth
species.
Shah et al. (2008) carried out an experiment on length weight
relationship of Schizothorax esocinus in cold water bodies of Kashmir valley
India and concluded that maximum body weight occurs just before spawning
period.
2.4. Age and Growth
Le Cren (1947) while studying opercular bones of perch Perca
fluviatilis from windermere lake, England observed that the age determination
by opercular bones had similarity with frequency distribution method.
Opercular bones showed allometric growth in relation to length of fish and it
was concluded that opercular bones can be used for age determination.
Chacko and Zobiri (1948) concluded that the number of rings
represents the body length of the fish in inches and also probably the age in
months.
Raj (1951) studied the annuli on the scales of Hilsa ilisha and
described the varying temperature, amount of food and other physiological
factors responsible for the formation of annual growth rings on scales. The
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number of these rings describe the age of fish and the length of the scale was
proportional to the length of the fish in all growth stages.
Lagler (1956) provided an elaborate account of the classification of
fishes on the basis of scale counts along the lateral line; between the lateral
line and pelvic fin; and between dorsal fin and lateral line. Further
classification on the basis of type and arrangement of scales was also given.
Pantulu et al. (1962) stated that a critical examination of the periodic
markings on otoliths helped to establish that the markings could be used for
the estimation of age and growth of the eel, Anguilla nebulosa.
Regier (1962) explained that the scale method for the study of age and
growth was one of the best methods in blue gill, Lepomis macrochirus. He
pointed out the use of combined correction factor and was of the view that
formation of an annulus was advanced with the increase in latitude.
Jhingran (1963) while studying the biology of Catla catla observed the
formation of annuli in the months of April-August with a high frequency in
June-July. It was concluded that annulus formation was influenced by
spawning stress and low feeding intensity.
Natarajan and Jingran (1963) gave an account of differentiation
between false and true annuli in the scales of Catla (Ham.). However, it was
stated that the density variation in the basic material often provides the optical
illusion of a band or groove, which can easily be differentiated from true
annulus as it does not confirm the properties of a true annulus.
Das (1964) worked on the scales of fresh water fishes of India and
found that the scales studied show annuli in one form or the other. The
location of the nucleus, the comparative extent of the scale fields, the
prominence of the shoulders, the mode of arrangement of the circuli, and the
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34
presence or absence of species in the posterior field, were all scale characters
specific for a fish species. Scale formula given for each fish was also a
constant specific feature. The investigations also revealed the importance of
age and growth studies in the fresh water fishery biology in India with the
help of scales after proper study and interpretation.
Das and Dulari (1968) worked on the age determination of Salmo
trutta fario Linn. and Cyprinus carpio specularis Linn. and showed the
annual growth rings in the form of discontinuation of circuli, cutting over of
circuli, opaque rings, or even as thin, clear streaks surrounding the nucleus
concentrically.
Singh and Rege (1968) determined the age of Tachysurus sona (Ham.)
using vertebrae and concluded that vertebrae can be employed for the
determination of age of fishes in the absence of scales and not as substitute for
scales.
Qasim (1973) described the difficulties encountered while studying the
age and growth of marine fishes of India.
Balon (1974) successfully used the growth data obtained from the scale
studies of common carp, Cyprinus carpio from central Europe for the
calculation of absolute and relative growth values.
Dudley (1974) explained the formation of annulus in three species of
Tilapia in October-January and confirmed that in young fishes the formation
of annuli was sooner than in the older fishes and considered three factors
responsible for it, viz. Low temperature, food conditions and spawning.
Monchizuki (1979) examined the age and growth of the two
representatives of the Scombropidae, Scombrops boops and Scombrops
gilberti, by means of otolith reading. Marginal growth of the otolith showed
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35
the availability as annual rings of the outer margins of hyaline ones formed in
the period from November to January. It was further revealed that Scombrops
gilberti is larger than Scombrops boops in standard length upto two years of
age. The estimated maximum standard length is 880mm for Scombrops boops
and 514mm for. Scombrops gilberti. Growth in terms of body weight was also
examined.
Brauhn and Kincaid (1982) studied survival, growth, and catchability
of rainbow trout of four strains and it was found that the survival rate was
higher in the natural strain as compared to the domestic strain, but growth was
slower. It was further revealed that the rainbow trout of different strains vary
in their suitability for different fishery management practices.
Rubec and Qadri (1982) studied age and growth of brown bull-head,
Ictalurus nebulosus in sections of the Ottawa river, Canada and concluded
that vertebrae were more reliable than other hard parts.
Sunder and Subla (1984) observed the occurrence of growth rings on
the vertebrae, opercular bones and otoliths of Schizothorax curvifrons and
reported that the periodic markings on these parts were laid annually.
Casey et al. (1985) studied age and growth of the Sandbar shark,
Carcharhinus plumbeus from the rings on the vertebrae and concluded that
one ring was formed each year between September and June.
L’ Abee-Lund (1985) worked on the age and growth of tench, Tinca
using opercular bones, otoliths and scales and concluded that opercular bones
were more suitable for age determination than scales.
Abbas and Siddiqui (1987) studied the age and growth of Channa
punctatus by using opercular bones.
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36
Kohli (1989) used pectoral spines as an age indicator in
Heteropneustes fossilis.
Dobriyal and Singh (1990) successfully employed otoliths and scales
for the determination of age in Barilus bendelisis.
David and Pancharatna (2003) studied age determination of the Indian
Whiting, Sillago indica using length–weight relationship and otolith
dimensions. Condition factor (k) and relative condition factor (Kn) was
calculated. Otolith dimensions such as diameter (OD), weight (OW) and
growth ring count (ORC) were recorded from sagittae. Otoliths of the
different sized fishes revealed the presence of 1-5 growth rings.
Phelps et al. (2007) studied precision of five structures for estimating
age of common carp, they analysed that otoliths have been validated as an
accurate structure for estimating age of common carp, Cyprinus carpio. They
estimated age using scales, vertebrae, operculas, pectoral fin rays and otoliths.
Although pectoral fin rays gave accurate age estimates however it still
requires evaluation. Accurate fish ages are important for growth analysis, age
structure analysis and mortality rate. They concluded that otoliths have gained
favour over other body structures because of their lack of resorption and
because their growth is a cellular rather than by calcification.
2.5. Declining Reasons
Fotedar and Qadri (1974) worked on the impact of cyprinus carpio L.
on endemic fish. It was concluded that the three varieties of Cyprinus carpio
L. viz., scale carp, mirror carp and leather carp introduced in the valley led to
the predominance of exotic species over the indigenous fish population.
Raizada (1985) studied the breeding, development and culture
prospects of the Schizothorax plagiostomus (Heckel). It was found that the
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37
fish has importance as a fishery of balancing link in the cold waters where the
exotic carnivore salmonids (trouts) was introduced. Population of
schizothoracine was an indicator of the population of trouts and mahaseer in
these waters. Over-exploitation on unrestricted bag limit besides killing
through illegal means including use of explosives and poisoning with plant
extracts such as (Artemisia) and Euphorbia (Thor) have reduced the
population affecting adversely the fishery.
Zutshi and Ticku (1990) studied the impact of mechanical deweeding
on Dal lake ecosystem and reported that due to deweeding practice some
undesirable changes have taken place disturbing the balance of the lake
ecosystems.
Sandhu et al. (1994) studied growth of an endangered fish, Tor
puttitora (Hamilton), from Govind Sagar (Himachal Pradesh). The studies
revealed that the Cyprinus carpio and Hypothalmichthyc molitrix had
alarmingly reduced the Mahseer (Tor Putitora) fishery in Govindsagar
reservoir.
Dehadrai and Poniah (1997) worked on the fish diversity and reported
that the principle factors for the decline of fishes include physical habitat loss,
degradation or alteration, chemical pollution or alteration.
Nurulamin et al. (2001) studied the population dynamics of Labeo
rohita and reported that over fishing mortality was found to be (E > 0.50) and
this was attributed to the lowering of population of the same fish.
Jha (2004) worked on the toxicological impact of the house hold
detergent, surf on digestive tissues (Stomach, Intestine, Liver) of the fish.
Investigation revealed higher degree of atrophy, disorganization of the
mucosal fold, severe dilation of blood vessel, necrosis of the columnar
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38
epithelial cells. Hepatic cords were disorganised and irregular. Cell membrane
was autolysed, in addition vacuolar degeneration of cytoplasm, splitting of
Hepatocytes, oedema, moderate dilation of sinusoidal space as also hyperemia
and liqufactive necrosis were apparent.
Khan (2004) worked on the various anthropogenic pressures
endangering the Kashmir lchthyofauna. Anthropogenic stresses, over the
years has remarkably affected the fish diversity.
Rao et al. (2005) studied the effect of the industrial effluents on the
length weight relationship and condition factor of the mullet, Lizia parsia.
The results of the study revealed that the length - weight relationship and
condition factor were significantly less in the polluted harbour and coastal
waters. The study further reveals the impact of the pollutants on the growth
and fishery of Lizia parsia.
Dhamendra and Kosygin (2006) studied the water quality, flora and
fauna of Merakhong river during September 2000-June 2002. Well marked
seasonal variations were recorded in respect of physico-chemical
characteristics. The river water was considered polluted as it had high
concentration of free CO2 (11.9 mg/l), nitrite nitrogen (0.035mg/l), inorganic
phosphorus (0.110 mg/l) and aquatic flora and fauna, which can thrive well in
polluted environments.