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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Page 1: 2.1. Breeding - Shodhgangashodhganga.inflibnet.ac.in/bitstream/10603/95793/9/09_chapter 2.pdf · feeding, population dynamics, breeding- maturation, spawning, fecundity etc. from

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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Review of Literature

11

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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Review of Literature

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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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23

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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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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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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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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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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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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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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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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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.