




































 Agricultural Science; Vol. 2, No. 1; 2020 
ISSN 2690-5396   E-ISSN 2690-4799 

https://doi.org/10.30560/as.v2n1p1 

1                             Published by IDEAS SPREAD 
 

A Review on Embryonic Development of Inland Fishes of Bangladesh 
Md. Borhan Uddin Ahmed1, Jobayda Sifat1, Md. Fazla Rabbi1, Md. Ashraful Islam1, H.M. Al Kabid Rafin1 & 

Md. Kamal Uddin1 
1 Department of Fisheries Biology and Genetics, Bangladesh Agricultural University, Memensingh-2202, 
Bangladesh 
Correspondence: Md. Borhan Uddin Ahmed, Department of Fisheries Biology & Genetics, Bangladesh 
Agricultural University, Mymensingh-2202, Bangladesh. E-mail: mbuasiam@gmail.com, 
borhan.fbg15@bau.edu.bd 
 
Received: January 5, 2020   Accepted: February 14, 2020   Online Published: February 18, 2020 
 
Abstract 
The early developmental pattern of inland fishes of Bangladesh are not well studied though it has a great 
importance in fisheries and aquaculture sector. The embryonic study provides interesting information on further 
growth and health of the fish and considered as an essential component for optimization of fish seed production 
by natural and induced breeding. Therefore, the current review work has been undertaken to provide a detail 
information on embryonic development of important inland fishes of Bangladesh. Information was collected from 
published scientific papers, un-published Masters and PhD dissertations from universities, popular articles and 
other published and grey literature. Diameters of unfertilized egg of the reviewed fish species were found to be 0.5 
to 1.3 mm and fertilized egg were 0.49 to 1.6 mm. Shapes of the egg were also variable from species to species. 
There is little information available on egg activation and egg micropyle of fish species of Bangladesh. The 
fertilization rate of different fishes ranged from 40.1% to 93.9%. There are different stages of early development 
in different species and time needs to complete the stages also vary. The timing of post hatching development by 
metamorphosis was found to vary based on the fish species from several days to weeks. Different factors like 
temperature, photoperiod, DO, seasonality and presence of chemicals in water were found to affect the early 
development of fish. The review included eighteen inland fishes and unearthed useful insights of their embryonic 
development and influence of different factors. As we expect, the outcome of the study would provide a baseline 
and would be very useful in conducting further research on the embryology of indigenous fishes of Bangladesh.   
Keyword: Fish embryo, ontogenic development, early life stage, hatching, larvae 
1. Introduction 
Bangladesh is a land with massive potential water bodies with a wide diversity of fishes. During several decades’ 
fishes of Bangladesh has declining due to various natural causes. Different man-made activities are also 
influencing the process. IUCN (2015) reported that 64 fish species are threatened which comprises 9 Critically 
Endangered, 30 Endangered and 25 Vulnerable fish species. Therefore, the conservation of this fishes is 
considerably important. The knowledge on early developmental pattern is essential for establishment of proper 
conservation measure. However, there is a very little knowledge available about the early embryonic and larval 
development of fishes of Bangladesh. Embryonic development is a complex process in which cellular 
differentiation and proliferation occur simultaneously at different rate (Hall, 2003). Changes in the pattern of the 
entire structure of an organ or of specific organ in relation to the environment are decisive for evaluating the 
developmental pattern of a species. Information on early life history is an essential requirement for optimization 
of mass seed production, culture and management of fishes. Embryonic development and larval development 
providing remarkable information in itself are imperative and consequential to the successful rearing of larvae for 
large scale seed production. Therefore, it is indispensable to conduct study to characterize different embryonic and 
larval stages of fish. In addition, embryonic developmental stages of fish life are also used in various 
investigational studies; especially in aquaculture as well as toxicological studies (Rahman et al., 2009).  
1.1 Fish Embryo 
Embryo is the earliest developing stages of fish from the time when the fertilized egg starts to divide, while it is 
continued within the egg until hatching. The embryo goes through several complex stages before hatching. Embryo 
is the result of fusion of male and female gamete and the first stage of life in fish as well as other animals 
(Langeland & Kimmel, 1997). 



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1.2 Gametogenesis 
The process of gamete formation in the sexually reproducing animals is gametogenesis. The male gamete is known 
as spermatozoon or sperm, and the female gamete is known as ovum or egg. Fish produce gametes directly through 
meiosis in organs called gonads (testis in males and ovaries in females). Gametogenesis are of two types; 
spermatogenesis and oogenesis (Andrade et al., 2001). 
1.2.1 Spermatogenesis 
Spermatogenesis involves two distinct process known as formation of the spermatids and Spermiogenesis (Sharma 
et al., 2018). The primordial germinal cells called spermatogonia undergoes repeated mitotic divisions to maintain 
a supply of cells for the production of sperm. After meiosis primary spermatocytes form secondary spermatocytes; 
after the second meiotic division, they are spermatids (Avidor-Reiss et al., 2015). The metamorphosis or 
differentiation of the spermatids into the sperm is called spermiogenesis. Spermiogenesis is characterized by 
changes in the nucleus, acrosome formation and Centrioles (Avidor-Reiss et al., 2015). 
1.2.2 Oogenesis 
The primordial germinal cells divide repeatedly to form the oogonia which multiply by the mitotic divisions and 
form the primary oocytes and pass through the growth phase. In the primary oocyte, large amount of fats and 
proteins become accumulated in the form of yolk and due to its heavy weight, it is concentrated towards the lower 
portion of the egg (forming vegetal pole). After this process cytoplasm of egg divides unequally forming three 
polar body and one egg (Lubzens et al., 2010). 
1.3 Different Embryonic Developmental Stages of Fishes 
1.3.1 Fertilized Eggs  
Prior to fertilization, the egg is in a quiescent state, arrested in metaphase of the second meiotic division. Upon 
binding of a sperm, the egg rapidly undergoes a number of metabolic and physical changes. The yolk is usually 
translucent and yellowish in color; the oil droplets are unchanged (Kinsey et al., 2007). Aerobic respiration 
increases in the egg (Nakano, 1953). Enzyme systems become activated. In most animals, a burst of protein 
synthesis begins and the nucleus undergoes the second division of meiosis after fertilization. 
1.3.2 Cleavage 
The zygote experiences a quick cell cycles with no significant growth, producing a cluster of cells within few 
minutes of fertilization is called cleavage (Sperber, 1995). Cleavage is basically occurring in the blastodisc region 
of the animal pole of the eggs which further converts into embryo (Fukazawa et al., 2010). After several successive 
cell division, the egg forms a thick layer of cell known as germ ring and is made up of superficial layer, ectoderm, 
endoderm mesoderm (Lee et al., 2004). A dorsal-ventral axis forms at that time that may be referred as pre-
notochord from which a neural plate is formed. Cells of the neural plate fold to form the neural groove and the 
surrounding neural folds which fuse, forming a hollow neural tube (Forgacs & Newman, 2005). 
1.3.3 Morula, Blastula and Gastrula 
A series of cleavage of zygote forms a solid ball of cell is called morula which occurs within few hours of 
fertilization. Soon after development of the 8-cell or 16-cell embryo (depending on the species), the blastomeres 
begin to form mulberry-shaped mass of cells called a morula. This change in shape of the embryo is called 
compaction (Forgacs & Newman, 2005). This compaction leads to form a hollow sphere called blastula, 
surrounding by a hollow blastocoel (Forgacs & Newman, 2005). The blastula enhances gastrula formation in which 
embryo form germ layers (Gilbert, 2010). Gastrula is a dramatic rearrangement of the cells of the blastula. Initially 
blastoderm cells move outwardly to intercalate with the more superficial cells which leads to the formation of 
gastrula (Warga & Kimmel, 1990). At this stage the yolk syncytial layer starts expansion around the yolk cell 
(Trinkaus, 1984). By the end of gastrulation, embryonic cells have rearranged into three layers’ ectoderm, 
mesoderm and endoderm. 
1.4 Importance of Embryonic Developmental Study of Fishes of Bangladesh 
Different organs of fish develop in different embryonic stage. Knowledge about the timing of different organ 
development of fish can be extended by embryonic study. Knowledge on cleavage pattern of Bangladeshi fish can 
be achieved by embryonic study. Embryonic study can be helpful to optimize survival and growth rate of fish 
larvae in our country. Embryonic development besides, providing interesting information are imperative and 
substantial to the successful rearing of larvae for large scale seed production of any species and thus important for 
hatchery operators. 



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1.5 Factors Affecting Embryonic Development 
A wide-ranging literature is present on the problem of the effect employed by some environmental factors upon 
the embryonic stage in fishes. These factors are like temperature, salinity, light, and some mechanical factors. 
Among this factors temperature is the most prominent factor as it is well known, the rate of the embryonic 
development of a given species is directly related to the temperature. Salinity is another important factor that affect 
the embryonic development of fishes. Every fish species has a tolerance limit of salinity beyond which in cannot 
survive. Light is not always controls the development of fish embryo. However, it has been shown in case of some 
species e.g. some salmonids, light usually has a negative impact on the early development of embryo (Eisler, 1957). 
Rather than these factors, some mechanical factors like pressure, shock etc. can sometime influence the embryonic 
development of fishes (Ciechomski, 1964). 
1.6 Justification and Objectives of the Study 
The embryonic and larval stages are considered very sensitive indicators of environmental disturbances 
(Marimuthu and Haniffa 2007). They are also indispensable in the study on ontogeny and phylogeny of their 
families (Legendre and Teugels 1991; Verreth et al., 1992). In addition, such studies on the embryonic 
development of any cultivable species can be useful in directing the husbandry efforts of fish farmer. In Bangladesh 
the embryonic study of fish has been not yet gained so much attention though it has a huge potential in the sector 
of fish and fisheries. The current review work is therefore has been undertaken to provide a details knowledge 
about the embryonic development of fishes of Bangladesh, its current status and future research importance on 
embryonic development of fishes of Bangladesh. 
2. Materials and Methods 
2.1 Literature Collection and Review 
Literature were collected from different journals, published paper, Magazines related to fish and fisheries. 
Unpublished research from masters and PhD research were also considered. Personal communication with the 
experts on the field of fish embryology were done for collection of some information. The fish species under this 
review are given in Table 1. 
 
Table 1. Fish species under review. 

Local name Scientific name Family Author 
Rui Labeo rohita Cyprinidae Das et al., 2006 
Catla Gibelion catla Cyprinidae Tumbahangfe et al., 2014 
Mrigel Cirrhinus cirrhosus Cyprinidae Chakraborty and Murty, 1972 
Desi Sarputi Puntius sarana Cyprinidae Chakraborty et al., 2007 
Bata Labeo bata Cyprinidae Hossain et al, 2007; Miah et al., 2009 
Silver berb Barbodes gonionotus Cyprinidae Basak et al., 2014 
Tara Baim Macrognathus 

aculeatus 
Mastacembelidae Farid et al., 2008 

Guchi baim Mastacembelus 
pancalus 

Mastacembelidae Hasan, M.R. et al., 2016; Rahman et al., 
2009 

Shol Channa striatus Channidae Roy et al, 2016; Marimuthu & Haniffa, 
2007 

Taki Channa punctatus Channidae Banerji, 1975 
Tengra Mystus cavasius Bagridae Rahman et al., 2004 
Rita Rita rita Bagridae Molla et al., 2008; Mollah et al., 2011 
Pabda Ompok pabo Siluridae Purkayastha et al., 2012; Sarma et al., 

2012 
Kani Pabda Ompok bimaculatus Siluridae Raizada et al., 2013 
Pangas Pangasius pangasius Pangasiidae Khan & Mollah, 2004; Ferosekhan et al., 

2015 
Local Koi Anabas testudineus Anabantidae Karim et al., (2012) 
Gutum Lepidocephalicthys 

guntea 
Cobitidae Sayeed et al, 2009 

Meni, Bheda Nandus nandus Nandidae Pal, et al., 2003; Das et al., 2002 
Shing Heteropneustes fossilis Heteropneustidae Puvaneswari et al., 2009; Nesa et al., 2017



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2.2 Data Analyses, Tabular and Graphical Representation 
All collected data were subjected in computer software MS Excel v2016 for analysis and graphical representation. 
3. Results and Discussion 
3.1 Fertilization 
The egg and spermatozoa are the main component for fertilization. The eggs and spermatozoa of many fish have 
an extremely short functional life after spawning. Marimuthu & Haniffa, (2007) showed in case of Channa striatus 
fertilized egg were free floating, spherical, non-adhesive translucent and yellow in color. In case of Rita rita, 
Mollah et al., (2011) reported the same result but it was demersal and brownish in color. They reported a redish 
spot in the fertilized egg which indicates the blastodisc. Same blastodisc has been reported in case of Macrognathus 
aculeatus, Labeo bata, Anabas testudineus, Nandus nandus, Heteropneustes fossilis, Mystus cavasius, by Farid et 
al., (2008); Miah et al., (2009); Karim et al., (2012); Das et al., (2002); Nesa et al., (2017); Rahman et al., (2004). 
The diameter of fertilized and unfertilized eggs is varying in species to species. Fertilization rate, hatching rate, 
diameter of fertilized and unfertilized egg of different fish species are given in the Table 2. 
The fertilization rate of different fish species is different. Fertilization rate are also season dependent in case of 
many species. Fertilization rate also differs depending on the natural or striping. 
 
Table 2. Fertilization rate, hatching rate and egg diameter of different inland fishes of Bangladesh 

Species name Highest 
fertilization 
rate (%) 

Highest 
hatching 
rate (%) 

Egg diameter Author 
Unfertiliz
ed (mm) 

Fertilized 
(mm) 

Catla Catla 93.9 90.98 - 4.5 Tumbahangfe et al., 2014 
Labeo bata 92.33 88.33 0.7±0.01 0.8±0.01 Hossain et al, 2007; Miah et 

al., 2009 
Mystus cavasius - - - 0. 49-

0.51 
Rahman et al., 2004 

Rita rita 71.66±7.64 48.33±7.6
4 

1.0 to 1.3 1.3 - 1.6 Molla et al., 2008; Mollah 
et al., 2011 

Mastacembelus 
pancalus 

75.23± 1.13 55.12± 
1.07 

0.50±0.00 0.70±0.0
2 

Hasan, et al., 2016; 
Rahman et al., 2009 

Macrognathus 
aculeatus 

- - 0.7±0.11 0.8±0.11 Farid et al., 2008 

Ompok pabo 75.5 60.5 0.99-1.1 1.0-1.3 Purkayastha et al., 2012; 
Sarma et al., 2012 

Ompok bimaculatus 75–90 80–90 - - Raizada et al., 2013 
Heteropneustes fossilis 62.33±4.51 41.33±5.6

9 
1-1.1 1.3 -1.4 Nesa et al., 2017 

Anabas testudineus - - 0.6±0.01 0.7±0.0 Karim et al., (2012) 
Lepidocephalicthys 
guntea 

78.60±3.21 65.49±5.2
3 

- - Sayeed et al, 2009 

Nandus nandus 92±5% 90±2 0.6 0.8 Pal, et al., 2003; Das et al., 
2002 

Pangasius pangasius 40.1±3.1 65.0±2.3 1.09-1.28 1.2-1.45 Khan & Mollah, 2004; 
Ferosekhan et al., 2015 

Channa striatus 59.5±2.50 67.5±2.50 - 1.20 -
1.40 

Roy et al, 2016; Marimuthu 
& Haniffa, 2007 

 
3.1.1 Egg Activation 
The event of egg activation is thought to be the result from the introduction of protein or other component from 
the sperm plasma to the egg. The existence of some factors which are capable of triggering egg for activation has 
been reported by Coward et al., (2003). The majority of the evidence to date indicates that the critical sperm 
component is a phospholipase isoform (phospholipase zeta) (Cox et al., 2002; Saunders et al., 2002). One of the 
exceptions of this was found in the study by Lee et al. (1999), which described the apparently normal activation 



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of zebrafish egg in the absence of sperm. This may be defined as parthenogenic activation egg. In general, some 
morphological changes occur during fertilization while the egg is being activated. According to Kinsey et al. 
(2007), some morphological changes occur in the fish egg at fertilization like progressive disintegration of cortical 
alveoli, reduction in volume, transformation of the chorion, expulsion of second polar body, bipolar differentiation. 
Kusa (1953) says that the first observable change after fertilization involves the outline of the alveolus becoming 
indistinct. The alveolus then disappears. Till date no researches were recorded on egg activation of fishes of 
Bangladesh. Details study is needed for the proper knowledge about the activation of fish egg of Bangladesh. 
3.1.2 Micropyle 
There is no significant record of micropyle study of Bangladeshi fish. At the internal aperture of the micropylar 
canal a site of sperm attachment was found on the egg membrane (Kudo, 1982), recognized as a gentle cytoplasmic 
swelling, bearing 19-27 cytoplasmic finger-like projections (Kudo, 1982). 
3.2 Embryonic Developmental Stages and Timing of Development in Different Fish Species 
The timing of development is species specific and also stage specific, that means different developmental stage 
needs different time for each species. In Bangladesh, the developmental timing for all species is not well known. 
However, in case of some species, research shows some good findings. A complete chart on the timing of 
development is shown in the Table 3. 
 
Table 3. Timing of early developmental stages of inland fishes of Bangladesh 

Stages/Species Labeo 
bata 
(Miah 
et al., 
2009) 
 

Ompok 
pabo 
(Sarma et 
al., 2012) 

Heteropneustes 
Fossilis 
(Nesa et al., 2017) 
 

Channa 
striatus 
(Marimuthu 
and Haniffa 
2007) 

Macrognathus 
aculeatus 
(Farid et al. 
2008). 

Anabas 
testudineus 
(Karim et al., 
2012) 

Barbodes 
gonionotus 
(Basak et al., 
2014) 

Fertilized egg 00 min 00 min 00 min 00 min 00 hrs 00 hrs 00 hrs 

Two cells 45 min 36 min 20.60 min 15-20 min - 1.20 hrs 0.35hrs 

Four cells 55 min 46 min 39.8 min - - 1.50 hrs 0.50 hrs 

Eight cells 80 min 60 min 71.67 min - - 2.20 hrs 1.00 hrs 

Sixteen cells - 1.08 hrs - 30-50 min - - 1.20-1.40 hrs 

Thirty-two 
cells 

- - - 1.00-1.20 min  - 2.00 hrs 

Morula 45-55 
min 

2.06 hrs 2.20 hrs 1.3-2.0 hrs 5.00hrs 5.10hrs 3.00hrs 

Blastula 4.30 hrs 3.30 hrs 4.15 hrs 5.00-6.00 hrs 9.00 hrs - 5.00 hrs 

Gastrulation 8.30-
12hrs 

- 6.35 hrs 8.00-9.00 hrs 15.10hrs - 6.15-6.25 hrs 

Germinal ring 
formed 

- - 5.1 hrs - - 8.20 hrs - 

14 somites - -  - - 32.30 hrs - 

16-18 somites - -  - -- 37.30 hrs - 

Yolk plug 
stage 

16 hrs 5.0 0hrs 8.25 hrs - - - -- 

Organogenesis 16-
18hrs 

14.00hrs  - - - - 

Just before 
hatching 

18-20 
hrs 

- 22.5 hrs - 36-40 hrs 75.20 hrs - 



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Fully active 
embryo 

- 17.0-18.0 
hrs 

-  - - - 

Hatchling 
competed 

20-21 
hrs 

20 hrs 39.5 hrs 23.30-24.0 hrs 40 hrs 80.30 hrs 13.40-14.00 
hrs 

 
3.3 Organogenesis and Hatching Temperature 
In organogenesis stage different organs of the fish are formed. In case of Bangladeshi fish, organogenesis is not 
well studied except only a limited number of species. In an experiment conducted by Farid et al. (2008) reported 
that organogenesis in case of Tara baim starts almost 11 hours after fertilization. Both tail and head ends were 
clearly differentiated and the beating heart was visible in this stage. Gills and pectoral fins start to appear. Auditory 
and optic vesicle develops. These aspects of organogenesis were similar to L. rohita ((Khan, 1943) and C. mrigala 
(Chakraborty and Murty, 1972). In another study in case of Labeo bata conducted by Miah et al. (2009) concluded 
that, at the organogenesis stage, appearance of heart rudiment pectoral fin buds and gill rudiment occurs. 
Notochord becomes visible, auditory and optic vessels developed in 16-18 hours after fertilization. Das et al. (2002) 
on the other hand, reported that organogenesis occurs 13 hours after fertilization. At this stage both tail and head-
end were clearly visible and heart beat starts in case of Nandus nandus. Out of this species discussed, there are 
many commercially important fish species which organogenesis are not well defined in the previous research. 
However, the pattern of organogenesis is found more or less similar in the species of the country. It may be due to 
similarity in environmental condition, food supply and geographical location. 
Optimum incubation temperature for different fish species is different found. A list of optimum incubation 
temperature reported for different fish species of Bangladesh is presented in Table 4. 
 
Table 4. Optimum incubation temperature of different fish species of Bangladesh 

Name of the species Scientific name Temperature Author 
Bata Labeo bata 27-31°C Miah et al., 2009 
Sharputi Barbonymus gonionotus 26.6-27.5 °C Chakraborty et al., 2007 
Mrigal Cirrhinus cirrhosus 25-30 °C Chakrabatry & Murty., 1972
Taki Channa punctatus 26-30 ºC Ramanathan et al., 1985 
Local Koi Anabas testudineus 27-29 ºC Karim et al., 2012 
Shing Heteropneustes fossilis 29 ºC Puvaneswari et al., 2009 
Pabda Ompok pabo 29.3 ºC Sarma et al., 2012 

Rita Rita rira 27-29 °C Mollah et al., 2011 
 
3.4 Post Hatching Development of the Hatched Larvae 
Post hatching development means the development of the larvae from the hatching till metamorphosis. The pattern 
of post hatching development of different species is different. Timing of development is also dependent on some 
environmental factors, like temperature of the water, available DO of the water, salinity etc. A typical chart on the 
development timing of different fish species reported for Bangladesh is given in the Table 5. 
 
Table 5. Timing of post hatching development of different fishes of Bangladesh 

Species name Time after hatching Author 

Twelve hours Twenty-four hour Thirty-six hour Forty-eight hour Seventy-two hour 

Labeo bata Larvae size 

3.0±0.05 mm. 

Chromatophores 

seen in the eye. 

Ventral embryonic 

fin fold more 

prominent. 

Larvae length 4.4±0.01 

mm. Operculum & 

Myomeres visible. 

Prominent pectoral and 

pelvic fins fold. 

Eye size increased 

with pigmentation. 

Pectoral fin more 

prominent. Brain 

lobe visible mouth 

cleft formed. 

Larvae reached to 

Larvae length 

5.9±0.02 mm. Yolk 

sac convex interiorly, 

air bladder distinct 

elliptical. Large black 

chromatophores on 

head, prominent gills. 

Larvae silver-blackish 

and transparent, 

6.5±0.02 mm in size. 

Myomere visible. 

Larvae swim and feed 

actively. 

Harun et al., 

2009 



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Pectoral fin bud 

appeared.  

5.5±0.05 mm in 

size. 

Puntius sarana Larvae increased 

3.2±0.05 mm in 

size. Pectoral fin 

bud appeared. 

Melanophore 

bands prominent 

at the posterior 

end of the body, 

also appeared 

above the eye and 

around the yolk 

sac.  

Larva length 5.0 ± 0.01 

mm. Colour changed to 

silvery-yellow. 

Myomeres visible and 

mouth cleft formed.  

Larve color 

whitish-black. 

Pectoral and 

pelvic fin buds 

found. Length of 

the larva 5.2 ± 

0.04 mm. 

Air bladder distinct. A 

few black 

chromatophores found 

in the area posterior to 

the auditory and large 

black chromatophores 

observed on head. 

Larvae size 5.7 ± 0.02 

mm. 

Larval length 5.8 ± 

0.01 mm. Eyes fully 

pigmented and 

pectoral fin bud more 

pronounced. 

Chakraborty 

et al., 2007 

Channa 

striatus 

Length 4.2 mm, 

Caudal fin begins 

to separate and 

pectoral fin buds. 

Swim bladder 

formed and heart 

positioned in front 

of the yolk. 

 Average length 

5.1 mm. Pectoral 

fin round shaped, 

mouth formed, the 

lower jaw less 

developed, vent 

formed.  

Rudimentary gill 

opening and pits 

differentiate. 

Thick band of 

melanophore 

observed. 

Average length 5.4 

mm. Pectoral fins 

paddle shaped, mouth 

formed with well-

developed jaw. Vent 

and gill rudiments 

clearly visible. Larvae 

move at water surface 

and feeding 

exogenously. 

 Marimuthu 

and Haniffa, 

2007 

Rita rita. Larvae length 2.2 

mm. Yolk sac 

partially reduced. 

Alimentary canal 

tube like. Eye spot 

with a dark 

pigmented area, 

barbels found.  

Larvae length 2.5 mm. 

Pectoral fin buds seen. 

Pigmentation gradually 

extended all over the 

body, blood circulation 

system fully developed 

Length 2.8 mm. 

Distinct heart 

visible, functioned 

actively and 

reddish blood seen 

around the heart. 

Dark pigmented 

eyes and upper 

and lower jaws 

visible. Gills 

covered by the 

operculum. Mouth 

and anal pore 

found with small 

opening.  

Larvae length 3.0 mm. 

Pectoral fin folds 

became distinct and 

the rudimentary rays 

developed in the 

caudal fin. Mouth 

well-formed and 

barbels became 

elongated. Alimentary 

tract straight and 

distinct and a small 

pouch like stomach 

formed. Larvae 

swimming smoothly 

and feeding 

exogenously at end of 

this stage. 

 Mollah et 

al., 2011 

Macrognathus 

aculeatus 

Larvae length 

2.8±0.05 mm. 

Pectoral fin bud 

appeared. Large 

number of 

melanophores 

appeared above 

the eye and around 

the yolk sac.  

Pectoral and pelvic fin 

bud appeared. Air 

bladder visible. Anus 

became distinct. Larvae 

increased to 4.2±0.01 

mm. 

Length of the larva 

5.2±0.04 mm. 

Colour whitish-

black. Eyes 

became whitish 

black. 

Distinct air bladder 

seen. Few black 

chromatophores found 

on the caudal fin. 

Large black 

chromatophores 

observed on head. 

Larvae increased to 

5.7±0.02 mm in size 

Larvae length 

6.0±0.09 mm and the 

silver-blackish and 

transparent in colour. 

Eyes fully pigmented. 

Dorsal and ventral fin 

folds persistent. Larva 

swims actively. 

Farid et al., 

2008 

Anabas 

testudineus 

 Pectoral fins paddle 

shaped and the 

movements of fins were 

 Larva 2.7 mm in 

length. Yolk sac 

reduced to half. Brain 

Larvae length 3.5mm 

and mouth gap quite 

large. Head broadened 

Karim et 

al., 2012 



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marked. Mouth 

elliptical in shape and 

inferior in position. Gill 

appeared in the form of 

comb and the supply of 

yolk diminished 

gradually. 

formed completely and 

continuous heart beat 

visible. Mouth turned 

in terminal position. 

than body and became 

round in shape. Larva 

started feeding at the 

end of this period. 

Heteropneustes 

fossilis 

 Larvae length 4.0±0.2 

mm, reduced yolk sac 

seen. Eyes dark 

pigmented and 

prominent. Pectoral fin 

buds seen, jaws formed 

and the alimentary tract 

distinct. Heart visible 

and the blood 

circulatory system fully 

functional. Barbells 

appeared in the form of 

tiny knobs. 

Length of larvae 

4.3±0.3 mm. 

Pectoral fin oval 

shaped with a 

membranous flap. 

Mouth formed. 

Rudimentary gill 

openings and 

olfactory pits 

differentiated. The 

yolk reserve 

further 

diminished. 

Larvae length 4.6±0.2 

mm. Barbells became 

elongated and 

prominent around the 

mouth. Anal aperture 

and opercula well-

formed and distinct. 

Blood circulation 

observed in the heart, 

tail and opercula 

region. 

 

Larvae length 5.0±0.2 

mm, brownish in 

color. Mouth and anus 

became fully 

functional. Head 

prominent and four 

pairs of barbells 

noticed. Pectoral fins 

vascularized and the 

caudal fin had 5 

rudimentary rays. The 

yolk material 

completely absorbed 

and the larvae 

exhibited vigorous 

movements 

Nesa et al., 

2017 

 
3.5 Factors That Influence Embryonic Development 
There are many factors that influences embryonic development of fish has been identified by different scientist. 
These factors may be categories into some category. These are internal factors like endocrine regulation of the fish 
or may be other external physico-chemical factors. 
3.5.1. Endocrine Regulation 
Study on endocrine regulation of the developing embryo is not well studied in Bangladeshi fishes except Shing 
and Koi. In case of Shing, Nesa et al., (2007) reported that a dorso-ventral unpaired fin, and some melanophores 
appeared on the head region, ventral side of the notochord and dorsal side of the body, probably by influence of 
some endocrine glands. Karim (2012) also reported some endocrine regulatory development of the Koi embryo, 
but didn’t mentioned any special type of name of the gland. This two research clearly indicates that the 
development of early stages of the fishes are somehow regulated by the endocrine gland. But there is a lack of 
clear information that which gland is responsible for development of which organ. More research on this aspect is 
necessary in case of fish species of Bangladesh. 
3.5.2 Physico-Chemical Factors 
Embryonic stage is very crucial stage of a fish life. In this stage certain factors plays important role in the 
development of the embryo. There are many external factors that are responsible for regulation of the embryonic 
development of fish. These factors may be physical or may be chemical also. Temperature is the most vital factor 
that determines the proper development of the embryo. It has been proved as one of the major factors that regulate 
the development of the early embryonic stages of fish. Legendre & Teugels, (1991) indicated that temperature 
have an influence on the development of the embryo. In a study in case of Labeo bata embryonic and larval 
development were reported optimum from 27 °C-31°C (Miah et al., 2009). Similar influence of temperature was 
observed in case of Shol, koi, indigenous Magur, Shing, Taki, pabda, Pangas in discrete study conducted by 
Ramanathan et al., 1985; Karim et al., 2012; Singh & Vidyarthi, 1990; Benerji, 1975; Sarma et al., 2012; 
Ferosekhan et al., 2015, respectively. In a study conducted by Chattopadhyay and Chattoraj (2017), reported that 
gonadal development and spawning stops with the fall of temperature that happen with the approach of winter. 
This provides a straight indication that temperature along with photoperiod is the key controlling factor for 
maturation and spawning in fish. In case of spawning of fish such as carp and many other cyprinids, gonadal 
maturation begins in late winter or early spring. Therefore, it can easily be said that increasing temperature 
influence the maturation of gonad as well as influence the early developmental stages of fish which is also agreed 
by the result showed by Chakraborty et al., (2007) in case of Puntius sarana where the author found significant 



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variation in the larval development in different temperature treatments. In case of Australian strain of Lates 
calcarifer the rate of embryonic development was positively correlated to the increase in incubation temperature 
and the thermal tolerance range for the Australian strain of L. calcarifer eggs was found to be 28–34 °C in a study 
conducted by Thepot and Jerry (2015). In case of Bangladeshi strain of L. calcarifer it may be more or less similar 
however it needs further research on Bangladeshi strain. In a study conducted by Das et al., (2006) reported highest 
hatching rate and least time for attaining each ontogenic stage at 31 > 33 > 26 > 36 °C and were significantly 
different (p < 0.05) in case of Labeo rohita. The lowest hatching percentage and maximum time duration for 
attaining a given ontogenic stage for L. rohita were observed at 36 °C and also resulted in malformed embryos. 
Another important factor that affect the embryonic development of fishes is salinity. In case of L. rohita, Pillai et 
al., (2003) reported that survivability limit of L. rohita embryo in waters up to 8 ppt salinity but best embryonic 
development was obtained at 0 to 2 ppt.  
Deprived of temperature and salinity, there are some other factors that may have influence on embryonic 
development of the fishes in Bangladesh. This are sound, light, chemical compounds of water such as DO, pH, 
Alkalinity etc. (Rahman et al., 2011). But those factors are not well studied yet. Study on this factors that may 
influence the embryonic development is very important for Bangladeshi fish. 
4. Conclusion 
One of the major goals of fisheries biology is to inspect a fish stock and in terms of fisheries biology it is vital to 
know the embryonic and larval development. As these type of studies are essential to define the spawning periods 
and areas, to determine the chronological variations of the spawning period, to predict the mature stock of fish, to 
predict the rate of death of fish at the end of spawning period and to inspect the relation of the growth with its 
environment. In Bangladesh we need more research on the developmental biology of fishes, especially native or 
indigenous fish needs more concentration on this aspect. 
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    /HRV (Za stvaranje Adobe PDF dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke.  Stvoreni PDF dokumenti mogu se otvoriti Acrobat i Adobe Reader 5.0 i kasnijim verzijama.)
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    /NLD (Gebruik deze instellingen om Adobe PDF-documenten te maken die zijn geoptimaliseerd voor prepress-afdrukken van hoge kwaliteit. De gemaakte PDF-documenten kunnen worden geopend met Acrobat en Adobe Reader 5.0 en hoger.)
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    /ENU (Use these settings to create Adobe PDF documents best suited for high-quality prepress printing.  Created PDF documents can be opened with Acrobat and Adobe Reader 5.0 and later.)
  >>
  /Namespace [
    (Adobe)
    (Common)
    (1.0)
  ]
  /OtherNamespaces [
    <<
      /AsReaderSpreads false
      /CropImagesToFrames true
      /ErrorControl /WarnAndContinue
      /FlattenerIgnoreSpreadOverrides false
      /IncludeGuidesGrids false
      /IncludeNonPrinting false
      /IncludeSlug false
      /Namespace [
        (Adobe)
        (InDesign)
        (4.0)
      ]
      /OmitPlacedBitmaps false
      /OmitPlacedEPS false
      /OmitPlacedPDF false
      /SimulateOverprint /Legacy
    >>
    <<
      /AddBleedMarks false
      /AddColorBars false
      /AddCropMarks false
      /AddPageInfo false
      /AddRegMarks false
      /ConvertColors /ConvertToCMYK
      /DestinationProfileName ()
      /DestinationProfileSelector /DocumentCMYK
      /Downsample16BitImages true
      /FlattenerPreset <<
        /PresetSelector /MediumResolution
      >>
      /FormElements false
      /GenerateStructure false
      /IncludeBookmarks false
      /IncludeHyperlinks false
      /IncludeInteractive false
      /IncludeLayers false
      /IncludeProfiles false
      /MultimediaHandling /UseObjectSettings
      /Namespace [
        (Adobe)
        (CreativeSuite)
        (2.0)
      ]
      /PDFXOutputIntentProfileSelector /DocumentCMYK
      /PreserveEditing true
      /UntaggedCMYKHandling /LeaveUntagged
      /UntaggedRGBHandling /UseDocumentProfile
      /UseDocumentBleed false
    >>
  ]
>> setdistillerparams
<<
  /HWResolution [2400 2400]
  /PageSize [612.000 792.000]
>> setpagedevice

