Effect of Water
Temperature on Ovaprim-Induced Breeding Performance of Morakhi (Cirrhinus
mrigala) Under Hatchery Conditions
Mahdi Hassan 1, Ghulam
Rasool 2, Tofique Hassan Soomro3, Mukesh Kumar Bheel 4,
Khan Muhammad 5, Manzer Abbas Laghariv 6, Hamza Laghari 7,
Aqib Jarwar 8
¹’⁴Centre of Environmental Science, University of
Sindh, Jamshoro, Sindh, Pakistan
²’
³’ ⁵’ ⁶’ 7’ 8 Department of Fisheries and Aquatic Science,
University of Sindh, Jamshoro, Sindh, Pakistan
Corresponding
Email: abbasmanzer163@gmail.com
Abstract
Artificial propagation by hormone-induced breeding has
now become an essential means to produce seeds of economically important
freshwater fishes sustainably. Water temperature is one of the most important
environmental parameters affecting the reproductive physiology, maturation of
gametes, spawning activity and development of embryos during induced spawning.
The present study examined the effects of different
water temperatures on the breeding performance of Morakhi (Cirrhinus mrigala)
using Ovaprim in hatchery conditions. This
experiment was carried out in June 2026 at SFARI, three consecutive breeding
trials were used. Five pairs of mature
broodstock were used for each trial. Female
broodfish were given 0.5 mL kg-1 body weight of Ovaprim (Syndel) and males were
given 0.3 mL kg-1 body weight via intramuscular injection at the base of the
dorsal fin. Spawning began about 4-6 hours
post hormone treatment. As is common in
hatcheries, water temperature and total dissolved solids (TDS) were recorded
during each trial and the number of hatches was measured in volume (mL).
The water temperatures of the three trials were 26,
29 and 34°C while the TDS values ranged from 260 to 373 mg/l. The production of hatch varied across the three breeding
tests, totaling 1550, 1320 and 1750 mL respectively. The production of the hatches was highest in the third
trial when the temperature was 34°C and lowest in 29°C. The above observations indicate that it is possible for
breeding performance to be different under different environmental settings and
suggest that water temperature can affect reproductive success of the Morakhi
that are brooded in hatchery settings if they are stimulated to spawn using
Ovaprim. However, the results of the study
should be viewed as preliminary as it was only three observational breeding
events. Larger sample sizes, replicated
trials and detailed monitoring of water quality is suggested to determine the
best thermal conditions for induced spawning of C. mrigala. The present study serves as a useful baseline data for
hatchery managers for this commercially important freshwater fish and helps in
optimizing the artificial propagation practices for this species
Keywords: Artificial breeding, Ovaprim,
Cirrhinus mrigala, Morakhi, hatch production, water temperature, induced
spawning, hatchery management, freshwater aquaculture.
Introduction
Effect of Water
Temperature on Ovaprim-Induced Breeding Performance of Morakhi (Cirrhinus
mrigala) Under Hatchery Conditions
Mahdi Hassan 1, Ghulam
Rasool 2, Tofique Hassan Soomro3, Mukesh Kumar Bheel 4,
Khan Muhammad 5, Manzer Abbas Laghariv 6, Hamza Laghari 7,
Aqib Jarwar 8
¹’⁴Centre of Environmental Science, University of
Sindh, Jamshoro, Sindh, Pakistan
²’
³’ ⁵’ ⁶’ 7’ 8 Department of Fisheries and Aquatic Science,
University of Sindh, Jamshoro, Sindh, Pakistan
Corresponding
Email: abbasmanzer163@gmail.com
Abstract
Artificial propagation by hormone-induced breeding has
now become an essential means to produce seeds of economically important
freshwater fishes sustainably. Water temperature is one of the most important
environmental parameters affecting the reproductive physiology, maturation of
gametes, spawning activity and development of embryos during induced spawning.
The present study examined the effects of different
water temperatures on the breeding performance of Morakhi (Cirrhinus mrigala)
using Ovaprim in hatchery conditions. This
experiment was carried out in June 2026 at SFARI, three consecutive breeding
trials were used. Five pairs of mature
broodstock were used for each trial. Female
broodfish were given 0.5 mL kg-1 body weight of Ovaprim (Syndel) and males were
given 0.3 mL kg-1 body weight via intramuscular injection at the base of the
dorsal fin. Spawning began about 4-6 hours
post hormone treatment. As is common in
hatcheries, water temperature and total dissolved solids (TDS) were recorded
during each trial and the number of hatches was measured in volume (mL).
The water temperatures of the three trials were 26,
29 and 34°C while the TDS values ranged from 260 to 373 mg/l. The production of hatch varied across the three breeding
tests, totaling 1550, 1320 and 1750 mL respectively. The production of the hatches was highest in the third
trial when the temperature was 34°C and lowest in 29°C. The above observations indicate that it is possible for
breeding performance to be different under different environmental settings and
suggest that water temperature can affect reproductive success of the Morakhi
that are brooded in hatchery settings if they are stimulated to spawn using
Ovaprim. However, the results of the study
should be viewed as preliminary as it was only three observational breeding
events. Larger sample sizes, replicated
trials and detailed monitoring of water quality is suggested to determine the
best thermal conditions for induced spawning of C. mrigala. The present study serves as a useful baseline data for
hatchery managers for this commercially important freshwater fish and helps in
optimizing the artificial propagation practices for this species
Keywords: Artificial breeding, Ovaprim,
Cirrhinus mrigala, Morakhi, hatch production, water temperature, induced
spawning, hatchery management, freshwater aquaculture.
Introduction
Indian major carps are the
commercial fish species of Pakistan that live in running waters and spawn under
the influence of environmental stimuli. Mostly people prefer to culture and
consume Indian major carps including labeo rohita, Cirrhinus mrigala and catla
catla. Cirrhinus mrigala is commercially
significant due to its high consumer preference (Priyanka Mayank et al., 2016).
Before the induce breeding the seed was collected from natural riverain waters.
Ovaprim is a hormone commercially used for induced breeding overall the world.
Degree-hours is a standard unit that is used to measure the heat requirement
for spawning and hatching while data is scarce for many species (M.S Azhar et
al.,2022) Synthetic hormone triggers the brain to breed in confined area. Hormone contains 20μg of Salmon Gonadotropin
Releasing Hormone and dopamine (A M Khan et al., 2013). Fish brooders are identified by the
morphologically, male and female identification is done by the pectoral fins,
rough fin indicates the male and smooth fin for female. Eggs are light silver
colored and are non-adhesive, fertilized and hatched in circular tanks with low
water circulation. The fertilized eggs of C mrigala hatched within the 25 to 40
hours after fertilization (Rakesh Kumar 2023). The spawned eggs ranged in
diameter from 2.1 mm to 2.13 mm and possessed circular yolk sacs (Khan, S.A et
al., 2024). Cirrhinus mrigala with 3.5 kg body weight produced 227.8 ± 14.4 g
eggs (Mohammad Shoaib et al., 2014).
Environmental and anthropogenic factors decline the spawning of Indian
major carps (Kazi Rabeya Akther et al., 2024). Carps live and breed in
freshwater any biological and chemical change can effects on the gonadal
development and spawning, like increasing of temperature, TDS, pH and other primary
parameters. The spawning time was 62, 53.3 and 40 minutes for rohu, catla and
mrigal respectively at the temperature range between 27-34 ºC (B C Mohapatra et
al., 2018). Carps spawn during the monsoon season when cloudy weather appears.
June, July and August are conducive for the breeding of the carps (N.
Basavaraja et al., 2007). Temperature is
the most crucial environmental factor for embryonic development, 70% - 80%
hatching rates achieved between the 30 ºC and 32 ºC. Temperature exceeding 32ºC
caused denaturation of all eggs and larvae that resulted in reduced hatching
rate (S.A. Khan et al. 2022). At 22-30
ºC the L. rohita breeds successfully at Maland India (B.R KIRAN et al., 2006).
During the experiment 100 % ovulation rate has been found for Gonadoprim and
Gonopro-FH treated fishes followed by PGE treated fishes 90% (Kumar and Chauhan
2018). The carps with a pH value of 7-8 are crucial for success breeding
(Thirupathi et al., 2023). Hatching occurred after 18 to 24 at water
temperature 20 to 24.5°C (Naeem et al., 2013). Fecundity (Egg production) rate
was better under the Ovaprim treatment as compared to the Ovatide (A.M Khan et
al., 2006)
Materials and
Methods
Study
Area
The induced
breeding experiment was carried out at Sindh Fisheries Aquaculture Research
Institute, Sindh, Pakistan in June 2026. The study was conducted under normal
hatchery conditions, where the breeding performance of Morakhi (Cirrhinus
mrigala) was assessed after inducing them with the hormone Ovaprim at different
water temperatures
Experimental
Broodstock
The healthy, sexually
mature broodstock of Cirrhinus mrigala (Morakhi) were selected from the
broodstock ponds of the institute using external morphological features that
suggest reproductive maturity. Female broodfish were identified by the comparatively soft
and swollen abdomen and a reddish genital opening, while the male broodfish
were recognized by a comparatively narrow body, rough pectoral fins, and by the
release of milt from the genital opening after applying gentle pressure to the
abdomen. The experiment was conducted in
three consecutive trials, using five pairs of broodstock (five males and five
females) for each trial. New stocks of
broodstock were used for each brooding period. Weights of each male and female were measured with a
calibrated digital weighing balance before starting with the hormones.
Hormonal
Induction
Artificial breeding was
carried out by the use of the commercially available spawning hormone, Ovaprim®
(Syndel Laboratories Ltd.). A 0.5 mL kg−1 body weight intramuscular injection was
given to the female broodfish and 0.3 mL kg−1 body weight intramuscular
injection was given to the male broodfish. Sterile
disposable syringes were used to inject hormones at the base of the dorsal fin.
The broodfish were immediately moved to breeding
tanks after the hormone injection and were bred under normal hatchery
conditions. The spawning activity was
constantly observed and the spawning began about 4-6 hours after the hormone
injection.
Study
Area
The induced
breeding experiment was carried out at Sindh Fisheries Aquaculture Research
Institute, Sindh, Pakistan in June 2026. The study was conducted under normal
hatchery conditions, where the breeding performance of Morakhi (Cirrhinus
mrigala) was assessed after inducing them with the hormone Ovaprim at different
water temperatures
Experimental
Broodstock
The healthy, sexually
mature broodstock of Cirrhinus mrigala (Morakhi) were selected from the
broodstock ponds of the institute using external morphological features that
suggest reproductive maturity. Female broodfish were identified by the comparatively soft
and swollen abdomen and a reddish genital opening, while the male broodfish
were recognized by a comparatively narrow body, rough pectoral fins, and by the
release of milt from the genital opening after applying gentle pressure to the
abdomen. The experiment was conducted in
three consecutive trials, using five pairs of broodstock (five males and five
females) for each trial. New stocks of
broodstock were used for each brooding period. Weights of each male and female were measured with a
calibrated digital weighing balance before starting with the hormones.
Hormonal
Induction
Artificial breeding was
carried out by the use of the commercially available spawning hormone, Ovaprim®
(Syndel Laboratories Ltd.). A 0.5 mL kg−1 body weight intramuscular injection was
given to the female broodfish and 0.3 mL kg−1 body weight intramuscular
injection was given to the male broodfish. Sterile
disposable syringes were used to inject hormones at the base of the dorsal fin.
The broodfish were immediately moved to breeding
tanks after the hormone injection and were bred under normal hatchery
conditions. The spawning activity was
constantly observed and the spawning began about 4-6 hours after the hormone
injection.
Water
Quality Monitoring
Trail No
Temperature oC
TDS
1
26
373
2
29
260
3
34
313
|
Trail No |
Temperature oC |
TDS |
|
1 |
26 |
373 |
|
2 |
29 |
260 |
|
3 |
34 |
313 |
During all breeding trials,
basic water quality parameters were measured. Prior to the onset of
spawning, water temperature (°C) and total dissolved solids (TDS; mg/l⁻¹) were
recorded with portable digital water quality meters.
During all breeding trials,
basic water quality parameters were measured. Prior to the onset of
spawning, water temperature (°C) and total dissolved solids (TDS; mg/l⁻¹) were
recorded with portable digital water quality meters.
Hatch
Production Assessment
The total hatch production
recorded during each breeding trial was:
Breeding Trial
Hatch Production ml
Trial 1
1550
Trial 2
1320
Trial 3
1750
After successful spawning
and egg incubation, hatch production was determined according to the standard
hatchery practice employed at the Sindh Fisheries Aquaculture Research
Institute. Instead of counting individual hatchlings, the total quantity of hatch
produced was measured volumetrically (mL). This method is routinely used in
commercial carp hatcheries because it provides a rapid and practical estimate
of hatch output when large numbers of larvae are produced.
|
Breeding Trial |
Hatch Production ml |
|
Trial 1 |
1550 |
|
Trial 2 |
1320 |
|
Trial 3 |
1750 |
After successful spawning
and egg incubation, hatch production was determined according to the standard
hatchery practice employed at the Sindh Fisheries Aquaculture Research
Institute. Instead of counting individual hatchlings, the total quantity of hatch
produced was measured volumetrically (mL). This method is routinely used in
commercial carp hatcheries because it provides a rapid and practical estimate
of hatch output when large numbers of larvae are produced.
Experimental
Data
The body weights of broodfish used during each breeding
trial are presented below.
Trail
1
Male Fish
Weight kg
Female Fish
Weight kg
Male 1
1.8
Female 1
1.6
Male 2
2.3
Female 2
2.2
Male 3
1.4
Female 3
1.5
Male 4
2
Female 4
2.3
Male 5
1.5
Female 5
1.6
Trail 2
Male Fish
Weight kg
Female Fish
Weight kg
Male 1
1.5
Female 1
1.2
Male 2
2
Female 2
1.6
Male 3
2
Female 3
2
Male 4
1.2
Female 4
1.4
Male 5
2
Female 5
2
Trail 3
Male Fish
Weight kg
Female Fish
Weight kg
Male 1
2
Female 1
1.8
Male 2
2
Female 2
1.6
Male 3
1.5
Female 3
2
Male 4
1.8
Female 4
2
Male 5
1.5
Female 5
1.5
Statistical
Analysis
Descriptive statistics were
used for summarizing the collected data. The
mean weight of broodstock, water temperature, TDS and hatch production were
recorded for all three of the breeding trials so that comparisons could be
made. The results of the study were
interpreted descriptively as there were no replicated treatment groups and only
three observational breeding events. A
graphical approach and a comparative analysis were used to evaluate the
relationship between the environmental conditions and the hatch production;
there was no formal inferential statistical testing.
Results
Broodstock Characteristics
For the three
years, in June of 2026, a total of 15 pairs (30 broodfish) of Morakhi
(Cirrhinus mrigala) were utilized for three breeding trials. Five mature males and five
mature female broodfish were used for each trial. The body weight of the males ranged from 1.2 to 2.3 kg
while for females it ranged from 1.2 to 2.3 kg. Broodstock size was fairly similar in the three breeding
trials (Table 1) as were the average body weights of the broodfish.
Mean body weight of Morakhi
(Cirrhinus mrigala) broodstock used during the three breeding trials
shows in this table:
Breeding Trial
Mean Male Weight (kg)
Mean Female Weight (kg)
Trial 1
1.8
1.84
Trial 2
1.74
1.64
Trial 3
1.76
1.78
Water Quality during Breeding
The temperature of
the water used for the breeding trials varied from 26 to 34°C, with total
dissolved solids (TDS) ranging from 260 to 373 mg/l (Table 2). Trial 1 had the lowest
water temperature (26°C) and trial 3 had the highest water temperature (34°C).
On the other hand, the highest TDS value (373 mg/l)
was observed in Trial 1 while the lowest TDS value (260 mg/l) was recorded in
Trial 2.
Water quality parameters and
hatch production during Ovaprim-induced breeding of Morakhi (C. mrigala)
shows in this table:
Trial
Water Temperature (°C)
TDS (mg/l)
Hatch Production (mL)
1
26
373
1550
2
29
260
1320
3
34
313
1750
Hatch
Production
The consequences
of Ovaprim injection to all the broodfish were satisfactory and spawning
started about 4-6 hours after the injection of the hormone in all the breeding
trials. There were differences in hatch production between the
three trials (Table 2). The maximum mean
hatch output of 1750 mL was obtained in trial 3, the least mean hatch output of
1320 mL was obtained in trial 2 and trial 1 gave a mean hatch output of 1550
mL. Hatch production rose by around 32.6% and
17.4% for Trial 3 and Trial 1, respectively, as compared to Trial 2. The following observations suggest some differences in
breeding success between the three breeding events.
Relationship between Water Temperature and Hatch
Production
The water temperature
ranging from 26°C to 34°C was noticed to be increasing from the first breeding
trial to the third one, giving rise to an increasing trend in the hatch
production. The highest hatch production was obtained from the trial
carried out at 34°C while the lowest hatch production was obtained at 29°C.
This pattern indicates that under hatchery
conditions, water temperature might be a factor affecting the production of
hatches, but only three observational breeding trials were conducted in the
experiment. In conclusion, the relationship
observed should be interpreted with caution as a preliminary trend and not as a
statistically significant effect.
Figure 1. Relationship between water temperature
and hatch production during Ovaprim-induced breeding of Morakhi.
The body weights of broodfish used during each breeding
trial are presented below.
Trail
1
|
Male Fish |
Weight kg |
Female Fish |
Weight kg |
|
Male 1 |
1.8 |
Female 1 |
1.6 |
|
Male 2 |
2.3 |
Female 2 |
2.2 |
|
Male 3 |
1.4 |
Female 3 |
1.5 |
|
Male 4 |
2 |
Female 4 |
2.3 |
|
Male 5 |
1.5 |
Female 5 |
1.6 |
Trail 2
|
Male Fish |
Weight kg |
Female Fish |
Weight kg |
|
Male 1 |
1.5 |
Female 1 |
1.2 |
|
Male 2 |
2 |
Female 2 |
1.6 |
|
Male 3 |
2 |
Female 3 |
2 |
|
Male 4 |
1.2 |
Female 4 |
1.4 |
|
Male 5 |
2 |
Female 5 |
2 |
Trail 3
|
Male Fish |
Weight kg |
Female Fish |
Weight kg |
|
Male 1 |
2 |
Female 1 |
1.8 |
|
Male 2 |
2 |
Female 2 |
1.6 |
|
Male 3 |
1.5 |
Female 3 |
2 |
|
Male 4 |
1.8 |
Female 4 |
2 |
|
Male 5 |
1.5 |
Female 5 |
1.5 |
Statistical
Analysis
Descriptive statistics were
used for summarizing the collected data. The
mean weight of broodstock, water temperature, TDS and hatch production were
recorded for all three of the breeding trials so that comparisons could be
made. The results of the study were
interpreted descriptively as there were no replicated treatment groups and only
three observational breeding events. A
graphical approach and a comparative analysis were used to evaluate the
relationship between the environmental conditions and the hatch production;
there was no formal inferential statistical testing.
Results
Broodstock Characteristics
For the three
years, in June of 2026, a total of 15 pairs (30 broodfish) of Morakhi
(Cirrhinus mrigala) were utilized for three breeding trials. Five mature males and five
mature female broodfish were used for each trial. The body weight of the males ranged from 1.2 to 2.3 kg
while for females it ranged from 1.2 to 2.3 kg. Broodstock size was fairly similar in the three breeding
trials (Table 1) as were the average body weights of the broodfish.
Mean body weight of Morakhi
(Cirrhinus mrigala) broodstock used during the three breeding trials
shows in this table:
|
Breeding Trial |
Mean Male Weight (kg) |
Mean Female Weight (kg) |
|
Trial 1 |
1.8 |
1.84 |
|
Trial 2 |
1.74 |
1.64 |
|
Trial 3 |
1.76 |
1.78 |
Water Quality during Breeding
The temperature of
the water used for the breeding trials varied from 26 to 34°C, with total
dissolved solids (TDS) ranging from 260 to 373 mg/l (Table 2). Trial 1 had the lowest
water temperature (26°C) and trial 3 had the highest water temperature (34°C).
On the other hand, the highest TDS value (373 mg/l)
was observed in Trial 1 while the lowest TDS value (260 mg/l) was recorded in
Trial 2.
Water quality parameters and
hatch production during Ovaprim-induced breeding of Morakhi (C. mrigala)
shows in this table:
|
Trial |
Water Temperature (°C) |
TDS (mg/l) |
Hatch Production (mL) |
|
1 |
26 |
373 |
1550 |
|
2 |
29 |
260 |
1320 |
|
3 |
34 |
313 |
1750 |
Hatch
Production
The consequences
of Ovaprim injection to all the broodfish were satisfactory and spawning
started about 4-6 hours after the injection of the hormone in all the breeding
trials. There were differences in hatch production between the
three trials (Table 2). The maximum mean
hatch output of 1750 mL was obtained in trial 3, the least mean hatch output of
1320 mL was obtained in trial 2 and trial 1 gave a mean hatch output of 1550
mL. Hatch production rose by around 32.6% and
17.4% for Trial 3 and Trial 1, respectively, as compared to Trial 2. The following observations suggest some differences in
breeding success between the three breeding events.
Relationship between Water Temperature and Hatch
Production
The water temperature
ranging from 26°C to 34°C was noticed to be increasing from the first breeding
trial to the third one, giving rise to an increasing trend in the hatch
production. The highest hatch production was obtained from the trial
carried out at 34°C while the lowest hatch production was obtained at 29°C.
This pattern indicates that under hatchery
conditions, water temperature might be a factor affecting the production of
hatches, but only three observational breeding trials were conducted in the
experiment. In conclusion, the relationship
observed should be interpreted with caution as a preliminary trend and not as a
statistically significant effect.
Figure 1. Relationship between water temperature
and hatch production during Ovaprim-induced breeding of Morakhi.
Discussion
In order to enhance the seed
production of commercially important freshwater fishes, especially cyprinids
that are not capable of natural spawning due to environmental and physiological
factors, artificial breeding with synthetic hormonal agents is an indispensable
technique. In this study, it was found that
all the broodstock of Morakhi (Cirrhinus mrigala) responded positively after
administration of the hormone Ovaprim and started spawning within 4–6 hours
after injection of the hormone. The result of
this good spawning response suggests that the inducing agent, Ovaprim, works
well under hatchery conditions. In major
carps of India, use of a synthetic hormone in the form of GnRH analogue-based
drug, known as Ovaprim, has been shown to induce maturation, ovulation, and
synchronous spawning (Rath et al., 2007; Qureshi & Khan, 2015). The
latency period present in the present study falls within the range of observed
latency period in Indian carps under Ovaprim induced breeding, which occur
several hours after the hormonal stimulation depending upon species, broodstock
condition, temperature and management practice (Rath et al., 2007; Naik &
Mirza, 1994). Despite the use of equal doses of
hormones for females (0.5 mL kg-1) and males (0.3 mL kg-1), and the similar
body weight of brood fish, variation in hatch production was observed across
the three breeding trials. Hatch production
(1750 mL) was highest at the 34°C hatch incubation, whereas hatch production
(1320 mL) was lowest at the 29°C hatch incubation indicating that environmental
factors, particularly temperature, can affect reproductive performance and
hatch production. The temperature of the
water is a very important parameter affecting fish reproductive physiology as
it influences metabolic rate, endocrine control, gametogenesis, fertilization
rate and embryonic development. It has been
established in previous research that temperature changes affect the spawning
behaviour and reproductive efficiency of cyprinid fishes but the optimum
temperature ranges are different for different species and geographical areas.
The increase in hatch production in the third
breeding trial could be linked to increased physiological activity at higher
temperature; however, this is a tentative conclusion since only three
observations were conducted and no replicated temperature treatments were used.
Other studies on induced breeding have also
highlighted the importance of the state of maturity of the broodstock, its
nutritional status, the quality of the gametes and environmental handling
conditions during incubation, in addition to the dose of hormones used, in the
success of spawning and seed recovery (Rath et al., 2007). The average body weight of the broodfish in the three
trials was comparable, suggesting that large differences in broodstock size
were not a major factor in producing differences in the production of hatches.
However, differences in spawning performance could
have been caused by the variability of the gonadal maturity and reproductive
quality among the different fish because the physiological condition of the
broodstock is one of the most significant factors affecting the quality of the
eggs and the production of larvae. There have
been some previous studies related to carp breeding, which have indicated that
synthetic hormones can enhance the reliability of carp breeding, but the amount
and quality of the seed depends on the breeds of the broodstock and the skills
of the hatchery. The TDS values found during
the present study were between 260 and 373 mg/l⁻¹, which is freshwater and
favorable for carp production, but it was not possible to determine the effect
of individual TDS on hatch production due to the lack of other essential water
quality parameters, such as dissolved oxygen, pH, ammonia and hardness. The effective induction of spawning in all three trials
has established that the dosage of Ovaprim applied was effective in hatchery
conditions to spawn Morakhi. The results from
this study give preliminary data on the relationship between the environmental
temperature and the breeding performance of C. mrigala induced with Ovaprim.
Additional controlled experiments with larger
numbers of broodstock, repeated thermal treatment and other reproductive
parameters like the fertilization rate, hatching percentage, larval survival
and egg quality are needed to determine the best thermal conditions for
commercial seed production of this species.
Conclusion
The present study showed
that successful hatchery production of Morakhi (Cirrhinus mrigala) was achieved
by using Ovaprim induced breeding. All pairs
of broodstock responded well to the hormonal induction stimulus with spawning
achieved within 4-6 hours of injection of Ovaprim (0.5 mL kg-1 BW to females
and 0.3 mL kg-1 BW to males). The three
breeding trials with different water temperatures showed some variation in
hatch production. The highest hatch
production was obtained at 34°C (1750mL) and the lowest hatch production was
obtained at 29°C (1320mL), suggesting a relationship between temperature and
breeding performance and hatch production under hatchery condition. The observed relationship, however, is subject to the
limitations of the number of breeding trials and the lack of controlled
temperatures so that the relationship should not be used as a definitive
determination of optimum breeding temperature. The results are useful baseline data for hatchery
management and indicate that the environmental conditions under the induced
breeding operations need to be monitored carefully. Further studies with larger numbers of samples, replicated
studies, detailed water quality analysis, and additional reproductive factors
like fertilization rate, hatching rate, larval survival, and growth performance
should be recommended to determine exact requirement of temperature and for
better seed production efficiency of C. mrigala.
Declarations
Ethics
Approval and Consent to Participate
The present study was
conducted by observing the breeding performance of the fish under routine
aquaculture management conditions in the hatchery. All experimental procedures were carried out in accordance
with the standard hatchery procedures for induced breeding and handling of
broodstock. No unnecessary harm was done to
fish and all procedures were carried out appropriately and in good management.
In order to enhance the seed
production of commercially important freshwater fishes, especially cyprinids
that are not capable of natural spawning due to environmental and physiological
factors, artificial breeding with synthetic hormonal agents is an indispensable
technique. In this study, it was found that
all the broodstock of Morakhi (Cirrhinus mrigala) responded positively after
administration of the hormone Ovaprim and started spawning within 4–6 hours
after injection of the hormone. The result of
this good spawning response suggests that the inducing agent, Ovaprim, works
well under hatchery conditions. In major
carps of India, use of a synthetic hormone in the form of GnRH analogue-based
drug, known as Ovaprim, has been shown to induce maturation, ovulation, and
synchronous spawning (Rath et al., 2007; Qureshi & Khan, 2015). The
latency period present in the present study falls within the range of observed
latency period in Indian carps under Ovaprim induced breeding, which occur
several hours after the hormonal stimulation depending upon species, broodstock
condition, temperature and management practice (Rath et al., 2007; Naik &
Mirza, 1994). Despite the use of equal doses of
hormones for females (0.5 mL kg-1) and males (0.3 mL kg-1), and the similar
body weight of brood fish, variation in hatch production was observed across
the three breeding trials. Hatch production
(1750 mL) was highest at the 34°C hatch incubation, whereas hatch production
(1320 mL) was lowest at the 29°C hatch incubation indicating that environmental
factors, particularly temperature, can affect reproductive performance and
hatch production. The temperature of the
water is a very important parameter affecting fish reproductive physiology as
it influences metabolic rate, endocrine control, gametogenesis, fertilization
rate and embryonic development. It has been
established in previous research that temperature changes affect the spawning
behaviour and reproductive efficiency of cyprinid fishes but the optimum
temperature ranges are different for different species and geographical areas.
The increase in hatch production in the third
breeding trial could be linked to increased physiological activity at higher
temperature; however, this is a tentative conclusion since only three
observations were conducted and no replicated temperature treatments were used.
Other studies on induced breeding have also
highlighted the importance of the state of maturity of the broodstock, its
nutritional status, the quality of the gametes and environmental handling
conditions during incubation, in addition to the dose of hormones used, in the
success of spawning and seed recovery (Rath et al., 2007). The average body weight of the broodfish in the three
trials was comparable, suggesting that large differences in broodstock size
were not a major factor in producing differences in the production of hatches.
However, differences in spawning performance could
have been caused by the variability of the gonadal maturity and reproductive
quality among the different fish because the physiological condition of the
broodstock is one of the most significant factors affecting the quality of the
eggs and the production of larvae. There have
been some previous studies related to carp breeding, which have indicated that
synthetic hormones can enhance the reliability of carp breeding, but the amount
and quality of the seed depends on the breeds of the broodstock and the skills
of the hatchery. The TDS values found during
the present study were between 260 and 373 mg/l⁻¹, which is freshwater and
favorable for carp production, but it was not possible to determine the effect
of individual TDS on hatch production due to the lack of other essential water
quality parameters, such as dissolved oxygen, pH, ammonia and hardness. The effective induction of spawning in all three trials
has established that the dosage of Ovaprim applied was effective in hatchery
conditions to spawn Morakhi. The results from
this study give preliminary data on the relationship between the environmental
temperature and the breeding performance of C. mrigala induced with Ovaprim.
Additional controlled experiments with larger
numbers of broodstock, repeated thermal treatment and other reproductive
parameters like the fertilization rate, hatching percentage, larval survival
and egg quality are needed to determine the best thermal conditions for
commercial seed production of this species.
Conclusion
The present study showed
that successful hatchery production of Morakhi (Cirrhinus mrigala) was achieved
by using Ovaprim induced breeding. All pairs
of broodstock responded well to the hormonal induction stimulus with spawning
achieved within 4-6 hours of injection of Ovaprim (0.5 mL kg-1 BW to females
and 0.3 mL kg-1 BW to males). The three
breeding trials with different water temperatures showed some variation in
hatch production. The highest hatch
production was obtained at 34°C (1750mL) and the lowest hatch production was
obtained at 29°C (1320mL), suggesting a relationship between temperature and
breeding performance and hatch production under hatchery condition. The observed relationship, however, is subject to the
limitations of the number of breeding trials and the lack of controlled
temperatures so that the relationship should not be used as a definitive
determination of optimum breeding temperature. The results are useful baseline data for hatchery
management and indicate that the environmental conditions under the induced
breeding operations need to be monitored carefully. Further studies with larger numbers of samples, replicated
studies, detailed water quality analysis, and additional reproductive factors
like fertilization rate, hatching rate, larval survival, and growth performance
should be recommended to determine exact requirement of temperature and for
better seed production efficiency of C. mrigala.
Declarations
Ethics
Approval and Consent to Participate
The present study was
conducted by observing the breeding performance of the fish under routine
aquaculture management conditions in the hatchery. All experimental procedures were carried out in accordance
with the standard hatchery procedures for induced breeding and handling of
broodstock. No unnecessary harm was done to
fish and all procedures were carried out appropriately and in good management.
Availability
of Data and Materials
All data obtained and
analyzed in the present study are available from the corresponding author on
reasonable demand.
All data obtained and
analyzed in the present study are available from the corresponding author on
reasonable demand.
Competing
Interests
The authors declare that they have no competing
interests.
Funding
There was no specific
external funding for this research. This
study was carried out with available resources and facilities at SFARI.
Acknowledgements
The authors gratefully acknowledge the management and
technical assistance of the Sindh Fisheries Aquaculture Research Institute in
providing the facilities and technical assistance in the breeding experiment.
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Reproductive profile of Cirrhinus mrigala (Hamilton, 1822) and
suggestions for restoration from the Yamuna River, India. Bioved, 27(1),
115–120. https://www.researchgate.net/profile/Priyanka-Mayank/publication/303825845_Reproductive_profile_of_Cirrhinus_mrigala_and_suggestion_for_restoration_Hamilton_1822_from_the_Yamuna_river_India/links/5756b44008aec74acf5ae486/Reproductive-profile-of-Cirrhinus-mrigala-and-suggestion-for-restoration-Hamilton-1822-from-the-Yamuna-river-India.pdf
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Azhar, M. S., Anjum, M. Z., Akhter, S., Khan, M. Q.,
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4(1), 9–18. https://doi.org/10.33687/zoobiol.005.01.4487
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(2013). Evaluation of spawning efficacy of Ovatide for carp breeding in
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Sarangi, N. (2007). Comparative account of induced breeding of Indian major
carps with Ovaprim, Ovatide, WOVA-FH and carp pituitary extract. Indian
Journal of Animal Sciences, 77(10), 1057–1060. https://www.cabidigitallibrary.org/doi/full/10.5555/20073267688
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variations on breeding behavior of Cirrhinus mrigala during induced
spawning. Pakistan Journal of Zoology. https://www.researchgate.net/profile/Sohail-Ahmad-20/publication/355166342_Impact_of_Temperature_Variations_on_Breeding_Behavior_of_Cirrhinus_mrigala_during_Induced_Spawning/links/626569568e6d637bd1fa972b/Impact-of-Temperature-Variations-on-Breeding-Behavior-of-Cirrhinus-mrigala-during-Induced-Spawning.pdf
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M., Ghayasuddin, M., Shah, S. M. A., Waseem, R., & Abid, K. (2024).
Comparison of skin staples versus skin sutures after inguinal hernioplasty in
terms of surgical site infection. Pakistan Journal of Medicine and
Dentistry, 13(1), 44-49. https://ojs.zu.edu.pk/pjmd/article/view/2361
12. Purdom, C. E. (1992). Genetics
and fish breeding (Vol. 8). Springer Science & Business Media.
https://books.google.com/books?hl=en&lr=&id=9Al-fcFBRqwC&oi=fnd&pg=PR11&dq=fish+breeding&ots=1YBcSuY2qT&sig=WesAvTA8qHQNWnKVsb5j4IZBG_M
13. Harvey, B., & Carolsfeld, J.
(1993). Induced breeding in tropical fish culture. IDRC, Ottawa,
ON, CA. https://idl-bnc-idrc.dspacedirect.org/bitstreams/096731da-842e-4d58-bd48-63eeb92761ae/download
14. Kinghorn, B. P. (1983). A review of
quantitative genetics in fish breeding. Aquaculture, 31(2-4),
283-304. https://www.sciencedirect.com/science/article/pii/0044848683903204
15. Johnston, I. A., Kent, M. P., Boudinot,
P., Looseley, M., Bargelloni, L., Faggion, S., ... & Lien, S. (2024).
Advancing fish breeding in aquaculture through genome functional
annotation. Aquaculture, 583, 740589. https://www.sciencedirect.com/science/article/pii/S0044848624000504
16. Dupont-Nivet, M., Vandeputte, M.,
Haffray, P., & Chevassus, B. (2006). Effect of different mating designs on
inbreeding, genetic variance and response to selection when applying individual
selection in fish breeding programs. Aquaculture, 252(2-4),
161-170. https://www.sciencedirect.com/science/article/pii/S0044848605004655
17. Hu, J., Yang, J., & Liao, H. (2024).
Progress on stress resistance breeding in fish. Reproduction and
Breeding, 4(4), 267-278. https://www.sciencedirect.com/science/article/pii/S2667071224000449
18. Gall, G. A., & Bakar, Y. (2002).
Application of mixed-model techniques to fish breed improvement: analysis of
breeding-value selection to increase 98-day body weight in tilapia. Aquaculture, 212(1-4),
93-113. https://www.sciencedirect.com/science/article/pii/S0044848602000248
19. Villanueva, B., Woolliams, J. A., &
Gjerde, B. (1996). Optimum designs for breeding programmes under mass selection
with an application in fish breeding. Animal Science, 63(3),
563-576. https://www.cambridge.org/core/journals/animal-science/article/optimum-designs-for-breeding-programmes-under-mass-selection-with-an-application-in-fish-breeding/D7DE79522C43F3F3DA69EC18A03BD437
20. Villanueva, B., Woolliams, J. A., &
Gjerde, B. (1996). Optimum designs for breeding programmes under mass selection
with an application in fish breeding. Animal Science, 63(3),
563-576. https://www.cambridge.org/core/journals/animal-science/article/optimum-designs-for-breeding-programmes-under-mass-selection-with-an-application-in-fish-breeding/D7DE79522C43F3F3DA69EC18A03BD437
21. Lind, C. E., Ponzoni, R. W., Nguyen, N.
H., & Khaw, H. L. (2012). Selective breeding in fish and conservation of
genetic resources for aquaculture. Reproduction in domestic animals, 47,
255-263. https://www.tandfonline.com/doi/abs/10.4081/ijas.2007.1s.795
22. Lind, C. E., Ponzoni, R. W., Nguyen, N.
H., & Khaw, H. L. (2012). Selective breeding in fish and conservation of
genetic resources for aquaculture. Reproduction in domestic animals, 47,
255-263. https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1439-0531.2012.02084.x
23. Yan, S. Y., & Ozgunen, T. (1993).
Fish breeding and biotechnology. Journal of the Islamic Academy and
Sciences, 6(3), 121-129. https://jag.journalagent.com/z4/download_fulltext.asp?pdir=ias&plng=tur&un=IAS-44365
24. Gjedrem, T. (1997). Flesh quality
improvement in fish through breeding. Aquaculture international, 5(3),
197-206. https://link.springer.com/article/10.1023/A:1014546816984
25. Midtlyng, P. J., Storset, A., Michel,
C., Slierendrecht, W. J., & Okamoto, N. (2002). Breeding for disease
resistance in fish. BULLETIN-EUROPEAN ASSOCIATION OF FISH PATHOLOGISTS, 22(2),
166-172. https://www.researchgate.net/profile/Paul-J-Midtlyng-2/publication/266499208_Breeding_for_disease_resistance_in_fish/links/545a16e40cf2bccc49130371/Breeding-for-disease-resistance-in-fish.pdf
26. Nkalubo, W., Balirwa, J., Bassa, S.,
Muhumuza, E., Nsega, M., & Mangeni, R. (2018). Fish breeding areas as a
management tool for fisheries resources in Lake Victoria, East Africa. African
journal of Tropical hydrobiology and Fisheries, 16(1), 1-9.
https://www.ajol.info/index.php/ajthf/article/view/170909
27. De Verdal, H., Komen, H., Quillet, E.,
Chatain, B., Allal, F., Benzie, J. A., & Vandeputte, M. (2018). Improving
feed efficiency in fish using selective breeding: a review. Reviews in
Aquaculture, 10(4), 833-851. https://onlinelibrary.wiley.com/doi/abs/10.1111/raq.12202
28. Moorhead, J. A., & Zeng, C. (2010).
Development of captive breeding techniques for marine ornamental fish: a
review. Reviews in Fisheries Science, 18(4), 315-343.
https://www.tandfonline.com/doi/abs/10.1080/10641262.2010.516035
29. Bhat, I. A., Rather, M. A., Ahmad, I.,
Ahmad, I., Mir, I. N., & Hussna. (2025). Impact of shifting abiotic factors
in aquaculture on fish breeding and reproduction: a review. Blue
Biotechnology, 2(1), 3. https://link.springer.com/article/10.1186/s44315-025-00027-9
30. Tave, D. (1986). Genetics for
fish hatchery managers (pp. xv+-299pp). https://www.cabidigitallibrary.org/doi/full/10.5555/19870102800
31. Lal, J., Vaishnav, A., Singh, S. K.,
Meena, D. K., Biswas, P., Mehta, N. K., & Priyadarshini, M. B. (2024).
Biotechnological innovation in fish breeding: from marker assisted selection to
genetic modification. Discover Biotechnology, 1(1), 7.
https://link.springer.com/article/10.1007/s44340-024-00007-6
The authors declare that they have no competing
interests.
Funding
There was no specific
external funding for this research. This
study was carried out with available resources and facilities at SFARI.
Acknowledgements
The authors gratefully acknowledge the management and
technical assistance of the Sindh Fisheries Aquaculture Research Institute in
providing the facilities and technical assistance in the breeding experiment.
References
1.
Mayank, P., Dwivedi, A. C., & Tiwari, A. (2016).
Reproductive profile of Cirrhinus mrigala (Hamilton, 1822) and
suggestions for restoration from the Yamuna River, India. Bioved, 27(1),
115–120. https://www.researchgate.net/profile/Priyanka-Mayank/publication/303825845_Reproductive_profile_of_Cirrhinus_mrigala_and_suggestion_for_restoration_Hamilton_1822_from_the_Yamuna_river_India/links/5756b44008aec74acf5ae486/Reproductive-profile-of-Cirrhinus-mrigala-and-suggestion-for-restoration-Hamilton-1822-from-the-Yamuna-river-India.pdf
2.
Azhar, M. S., Anjum, M. Z., Akhter, S., Khan, M. Q.,
Ali, A., Farooq, M., & Bibi, S. (2022). Degree-hours to spawning response,
fertilization rate and hatching of Labeo rohita and Cirrhinus mrigala
through induced breeding under hatchery environment. Journal of Zoo Biology,
4(1), 9–18. https://doi.org/10.33687/zoobiol.005.01.4487
3.
Khan, A. M., Iqbal, A., Shakir, H. A., & Ayub, M.
(2013). Evaluation of spawning efficacy of Ovatide for carp breeding in
Pakistan. Punjab University Journal of Zoology, 28(2), 77–81. https://www.researchgate.net/profile/Abdul-Khan-193/publication/272794627_Evaluation_of_spawning_efficacy_of_ovatide_for_carp_breeding_in_Pakistan/links/5d399c4fa6fdcc370a5de463/Evaluation-of-spawning-efficacy-of-ovatide-for-carp-breeding-in-Pakistan.pdf
4.
Rath, S. C., Sarkar, S. K., Gupta, S. D., &
Sarangi, N. (2007). Comparative account of induced breeding of Indian major
carps with Ovaprim, Ovatide, WOVA-FH and carp pituitary extract. Indian
Journal of Animal Sciences, 77(10), 1057–1060. https://www.cabidigitallibrary.org/doi/full/10.5555/20073267688
5.
Naik, I. U., & Mirza, Z. S. (1994). Use of
Ovaprim-C induced spawning of Indian major carps in Punjab, Pakistan. Proceedings
of the Pakistan Congress of Zoology. https://www.cabidigitallibrary.org/doi/full/10.5555/19960100991
6.
Qureshi, M. S., & Khan, H. (2015). Efficacy of
synthetic hormones Ovatide and Ovaprim in induced breeding of major Indian and
Chinese carps. International Journal of Agricultural Technology. https://li04.tci-thaijo.org/index.php/IJAT/article/view/6520
7.
Sharma, A. P., & Singh, V. K. (2002). Induced
breeding response of Indian major carps using Ovaprim and carp pituitary
extract. Indian Journal of Animal Sciences. https://www.cabidigitallibrary.org/doi/full/10.5555/20023086690
8.
Dhawan, A., & Kaur, K. (2004). Comparative efficacy
of Ovaprim and Ovatide in carp breeding. Indian Journal of Fisheries. https://www.researchgate.net/profile/Kamaldeep-Kaur-10/publication/277103470_Comparative_efficacy_of_ovaprim_and_ovatide_in_carp_breeding/links/5f96ea70a6fdccfd7b7fad7c/Comparative-efficacy-of-ovaprim-and-ovatide-in-carp-breeding.pdf
9.
Naik, I. U., & Mirza, Z. S. (1992). Use of
Ovaprim-C in induced spawning of Indian major carps in Punjab, Pakistan. Proceedings
of the Pakistan Congress of Zoology. https://www.cabidigitallibrary.org/doi/full/10.5555/19960100991
10. Khan,
S. A., Sherzada, S., Ashraf, M., et al. (2022). Impact of temperature
variations on breeding behavior of Cirrhinus mrigala during induced
spawning. Pakistan Journal of Zoology. https://www.researchgate.net/profile/Sohail-Ahmad-20/publication/355166342_Impact_of_Temperature_Variations_on_Breeding_Behavior_of_Cirrhinus_mrigala_during_Induced_Spawning/links/626569568e6d637bd1fa972b/Impact-of-Temperature-Variations-on-Breeding-Behavior-of-Cirrhinus-mrigala-during-Induced-Spawning.pdf
11.
Ali,
M., Ghayasuddin, M., Shah, S. M. A., Waseem, R., & Abid, K. (2024).
Comparison of skin staples versus skin sutures after inguinal hernioplasty in
terms of surgical site infection. Pakistan Journal of Medicine and
Dentistry, 13(1), 44-49. https://ojs.zu.edu.pk/pjmd/article/view/2361
12. Purdom, C. E. (1992). Genetics
and fish breeding (Vol. 8). Springer Science & Business Media.
https://books.google.com/books?hl=en&lr=&id=9Al-fcFBRqwC&oi=fnd&pg=PR11&dq=fish+breeding&ots=1YBcSuY2qT&sig=WesAvTA8qHQNWnKVsb5j4IZBG_M
13. Harvey, B., & Carolsfeld, J.
(1993). Induced breeding in tropical fish culture. IDRC, Ottawa,
ON, CA. https://idl-bnc-idrc.dspacedirect.org/bitstreams/096731da-842e-4d58-bd48-63eeb92761ae/download
14. Kinghorn, B. P. (1983). A review of
quantitative genetics in fish breeding. Aquaculture, 31(2-4),
283-304. https://www.sciencedirect.com/science/article/pii/0044848683903204
15. Johnston, I. A., Kent, M. P., Boudinot,
P., Looseley, M., Bargelloni, L., Faggion, S., ... & Lien, S. (2024).
Advancing fish breeding in aquaculture through genome functional
annotation. Aquaculture, 583, 740589. https://www.sciencedirect.com/science/article/pii/S0044848624000504
16. Dupont-Nivet, M., Vandeputte, M.,
Haffray, P., & Chevassus, B. (2006). Effect of different mating designs on
inbreeding, genetic variance and response to selection when applying individual
selection in fish breeding programs. Aquaculture, 252(2-4),
161-170. https://www.sciencedirect.com/science/article/pii/S0044848605004655
17. Hu, J., Yang, J., & Liao, H. (2024).
Progress on stress resistance breeding in fish. Reproduction and
Breeding, 4(4), 267-278. https://www.sciencedirect.com/science/article/pii/S2667071224000449
18. Gall, G. A., & Bakar, Y. (2002).
Application of mixed-model techniques to fish breed improvement: analysis of
breeding-value selection to increase 98-day body weight in tilapia. Aquaculture, 212(1-4),
93-113. https://www.sciencedirect.com/science/article/pii/S0044848602000248
19. Villanueva, B., Woolliams, J. A., &
Gjerde, B. (1996). Optimum designs for breeding programmes under mass selection
with an application in fish breeding. Animal Science, 63(3),
563-576. https://www.cambridge.org/core/journals/animal-science/article/optimum-designs-for-breeding-programmes-under-mass-selection-with-an-application-in-fish-breeding/D7DE79522C43F3F3DA69EC18A03BD437
20. Villanueva, B., Woolliams, J. A., &
Gjerde, B. (1996). Optimum designs for breeding programmes under mass selection
with an application in fish breeding. Animal Science, 63(3),
563-576. https://www.cambridge.org/core/journals/animal-science/article/optimum-designs-for-breeding-programmes-under-mass-selection-with-an-application-in-fish-breeding/D7DE79522C43F3F3DA69EC18A03BD437
21. Lind, C. E., Ponzoni, R. W., Nguyen, N.
H., & Khaw, H. L. (2012). Selective breeding in fish and conservation of
genetic resources for aquaculture. Reproduction in domestic animals, 47,
255-263. https://www.tandfonline.com/doi/abs/10.4081/ijas.2007.1s.795
22. Lind, C. E., Ponzoni, R. W., Nguyen, N.
H., & Khaw, H. L. (2012). Selective breeding in fish and conservation of
genetic resources for aquaculture. Reproduction in domestic animals, 47,
255-263. https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1439-0531.2012.02084.x
23. Yan, S. Y., & Ozgunen, T. (1993).
Fish breeding and biotechnology. Journal of the Islamic Academy and
Sciences, 6(3), 121-129. https://jag.journalagent.com/z4/download_fulltext.asp?pdir=ias&plng=tur&un=IAS-44365
24. Gjedrem, T. (1997). Flesh quality
improvement in fish through breeding. Aquaculture international, 5(3),
197-206. https://link.springer.com/article/10.1023/A:1014546816984
25. Midtlyng, P. J., Storset, A., Michel,
C., Slierendrecht, W. J., & Okamoto, N. (2002). Breeding for disease
resistance in fish. BULLETIN-EUROPEAN ASSOCIATION OF FISH PATHOLOGISTS, 22(2),
166-172. https://www.researchgate.net/profile/Paul-J-Midtlyng-2/publication/266499208_Breeding_for_disease_resistance_in_fish/links/545a16e40cf2bccc49130371/Breeding-for-disease-resistance-in-fish.pdf
26. Nkalubo, W., Balirwa, J., Bassa, S.,
Muhumuza, E., Nsega, M., & Mangeni, R. (2018). Fish breeding areas as a
management tool for fisheries resources in Lake Victoria, East Africa. African
journal of Tropical hydrobiology and Fisheries, 16(1), 1-9.
https://www.ajol.info/index.php/ajthf/article/view/170909
27. De Verdal, H., Komen, H., Quillet, E.,
Chatain, B., Allal, F., Benzie, J. A., & Vandeputte, M. (2018). Improving
feed efficiency in fish using selective breeding: a review. Reviews in
Aquaculture, 10(4), 833-851. https://onlinelibrary.wiley.com/doi/abs/10.1111/raq.12202
28. Moorhead, J. A., & Zeng, C. (2010).
Development of captive breeding techniques for marine ornamental fish: a
review. Reviews in Fisheries Science, 18(4), 315-343.
https://www.tandfonline.com/doi/abs/10.1080/10641262.2010.516035
29. Bhat, I. A., Rather, M. A., Ahmad, I.,
Ahmad, I., Mir, I. N., & Hussna. (2025). Impact of shifting abiotic factors
in aquaculture on fish breeding and reproduction: a review. Blue
Biotechnology, 2(1), 3. https://link.springer.com/article/10.1186/s44315-025-00027-9
30. Tave, D. (1986). Genetics for
fish hatchery managers (pp. xv+-299pp). https://www.cabidigitallibrary.org/doi/full/10.5555/19870102800
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