REPRODUCTIVE BIOLOGY OF FRESHWATER PRAWN … · REPRODUCTIVE BIOLOGY OF FRESHWATER PRAWN...

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Banats Journal of Biotechnology

2012, III(6),

86

REPRODUCTIVE BIOLOGY OF FRESHWATER PRAWN Macrobrachium

vollenhovenii (Herklot, 1857) CAUGHT IN WARRI RIVER

DOI: 10.7904/2068 – 4738 – III(6) – 86

Nkeonyeasua Florence OLELE1, Prekeyi TAWARI-FUFEYIN2 and Josiah Chidiebere

OKONKWO3

1Fisheries Department, Delta State University, Asaba Campus, P.O. Box 1110 Asaba, Nigeria

GSM: 08062905921 E–mail: [email protected] or [email protected] 2Department of Animal and Environmental Biology, University of Benin, PMB 1154. Benin City.

Nigeria. GSM: 08076072363, E–mail: [email protected] 3Department of Animal Science and Technology, Nnamdi Azikiwe University, PMB 5025 Awka,

Nigeria. E–mail: [email protected]

Abstract: A total number of 338 specimens of Macrobrachium vollenhovenii were bought from fisher folks who fished from Ubeji, Jala and Soroghagbene; but marketed their landings at Ubeji. Specimens were collected at monthly intervals between March and November, 2011. The male to female sex ratio established was 1:2:3. The size at first maturity was 9.0 cm for males and 10.0 cm for females. The length–weight relationship revealed an increase in total length of each specimen as body weight increased. The regression coefficient ‘r’ was significant at p< 0.05 for male (0.901) and female (0.644) specimens. An average condition factor of 3.9 and 2.74 was recorded for males and females respectively. The Gonado–Somatic–Index (GSI) was generally higher, especially during the month of July for females (0.60) than for males (0.50). Distinct histological changes were observed in the gonads of both sexes. Gonad development revealed five distinct stages (dominant, developing, developed, ripe and spent) for both sexes. Fecundity ranged between 5,540–28,470 eggs, and was highly correlated with total length (0.979) but not with body weight (0.185). Fecundity was highest in the month of July (20,113) and lowest in the month of August (6,140). It could be concluded that matured specimens probably spawn, in March and in July, when GSI were at their peaks. This pattern of breeding suggests that they were capable of multiple spawning within a single reproductive season.

Key words: length–weight relationship, condition factor, GSI, maturity stages, fecundity and

Macrobrachium vollenhovenii

Introduction The biology of prawns attracts

attention of biologists, due to their economic importance although; their detailed scientific study is at its infancy SHARMA and SUBBA, 2004.

Macrobrachium vollennovenii, of family Palaemonidae is among the largest genus in the Nigeria waters.

A maximum size of 13.50 cm and 19.50 cm attainable by this specie is reported MARIOGHAE, 1995; HOLTHIUS, 1980 respectively. The specie is bigger than M. macrobrachion whose maximum size is 12.50 cm. According to MARIOGHAE, 1992

Macrobrachium vollennovenii accounted for 60 % of the prawn landing from the

Lagos Lagoon. Reproduction is important in the life of organisms, because through it, ripe females give birth to their young’s.

Knowledge on gonad development and reproductive performance was important for culture systems to be improved. Hence, this idea has been used for estimating mean length at first maturity. This parameter which was used for management procedures indicate ecological processes when monitored through time and space IRECE et al., 2009.

Spawning was linked to seasons of the year, which were influenced by environmental variables such as food availability, temperature and photoperiod ISMEAL and NEW, 2000.

Banat’s University of Agricultural Sciences and Veterinary Medicine from Timisoara,

Contact: web: http://www.bjbabe.ro, e-mail: bjb@usab–tm.ro

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According to SPRINGATE, 2002, prawns require food to grow and reproduce; hence food deprivation was reported to reduce their rate of maturation and total fecundity.

The successful culturing of any animal requires a basic understanding of its key biological processes.

The most important of these biological processes was the reproductive cycle and formation of gametes.

In addition to the limited information available on prawn fisheries in Warri environ, non-have described gametogenesis, reproductive cycle and sex ratio, because of restive youth activists, in an area where oil exploration has become a lucrative venture.

The study therefore investigated the length/weight relationship, condition factor, gonado–somatic index, fecundity and histological presentation of maturity stages, of the species of study.

Materials and methods Description of study area Warri River is one of the most

important coastal rivers of the Niger–Delta region of Nigeria. It lies within longitude

5.30 to 5.40E and latitude 5.27 to 5.32N (Figure 1).

Taking its source few kilometers away from the first station at Ubeji, the river flows’ south–west as a main channel as it flows into Forcados Estuary, which finally empties into the Atlantic Ocean?

The river covers a surface area of about 255 km2 with a length of about 150 km2, and is located in the rain forest region of Nigeria.

There are two recognizable seasons of variable durations: the sunny and the rainy seasons OLOMUKORO and EGBORGE, 2004.

The study area is not entirely fresh water most periods of the year. During the sunny months (November–April) the water becomes brackish due to the incursion of marine waters from Forcados River. The reverse condition is the case during the rainy months of May–October.

The region is surrounded by tropical rainforest and swampy vegetation zone, while the landing stations (Ubeji, Jala and Soroghagbene) are presently secretly subjected to fishing activities, due to frequent youth restiveness.

Figure 1. Map of Warri River and its environing showing the regions (Souarogaene, Jala and Ubeji) where prawn samples were purchased collected on monthly basis.

Specimens were purchased from the fisher folk’s at Ubeji, sorted out and packed into collecting ice–chest cooler provided, before there were transported to the laboratory (located at the Aquaculture and Fisheries Department of the Delta State University, Asaba Campus). Sampling was carried out once

a month between 0900–1200 hours on each sampling day, for nine months (March to November, 2011).

In the laboratory, specimens were dissected, while gonads were isolated from the visceral region.

They were preserved in Bouin’s fixative, in readiness for histological



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preparations. When eggs were observed in matured females, they were neatly collected into different vials and preserved in 5 % formalin for fecundity estimation. Length measurements were recorded to the nearest 0.01cm using a meter rule, while weight measurement was conducted with an electronic balance, Model LP 302A LARK C. R.) and recorded to the nearest 0.01g SHARMA and

SUBBA, 2004. Sex ratio The sex of each prawn was

determined by visual examination of the region where the second pereopods or chelipeds was attached at specific positions at the abdominal region. A pair of cheliped was either joined together at the abdominal segment (in males) or separated (in female) specimens. Sex ratio was computed using chi–square.

Length/weight relationship The length–weight relationship was

calculated by the formula aLb, but because body weight increase more rapidly than total length, the formula was linearised for the purpose of regression analysis using logarithm transformation where W = Log a + b log L BERNACSEK, 1984.

Condition factor The well-being of the specimens

otherwise known as the condition factor BERNACSEK, 1984 was determined by the formula: K=100W/L3, where K=Condition factor; W=Weight of prawn in gram (g) and L=Length of prawn in centimeters (cm).

Size at first maturity In order to determine the length at

which the prawns attained maturity, it was essential to ascertain at what length, 50% and above of the prawn had gonads at maturity stage 111 and above. This length was observed for both sexes of the prawns according to DUMONT et al. 2007.

Gonado somatic index The gonado–somatic–index was

calculated from the formula: GSI=100(GW)/BW; where:

GW=Gonad weight and BW=Body weight. FERNANDEZ et al., 1998.

Fecundity estimation Fecundity is the number of ripened

eggs in the Ovary. Ripe ova were used for fecundity estimation by the gravimetric method. Eggs preserved in 5 % formalin were dried on layers of filter paper while the ovarian tissues were physically removed by hand. Ovary and sub–sample weights (g) were taken with the aid of Bran Scientific weighing balance (LA 164 Model). By simple proportion, the total number of eggs in the ovary was calculated by the formula of FERNANDEZ et al.,

1998, written thus: F=nG/g, where F=Fecundity; n=number of eggs, G=ovary weight, g=weight of sub samples.

Relative fecundity was calculated by the formula. F=Total no of eggs in the brood pouch/ Total weight of prawn, Where: F = Fecundity, FERNANDEZ et al., 1998.

Gonad histology Transverse sections of gonads

preserved in Bouin’s fixative were dehydrated in several changes of alcohol (70, 80, 90, 95 and 100 %) for 45 minutes each. Gonad sections were cleared in two changes of Xylene for 45 minutes each.

The tissues were infiltrated in paraffin wax at a temperature of 56C in the oven and embedded in molten paraffin wax. They were immersed in ice to solidify, cut (at 5.0µm) using the rotary microtome, and floated in the water bath.

Sections were picked with cleaned slides, placed on hot plate to dry and stained with Haematoxylin and Eosin.

They were dried thoroughly again, in the air and viewed under the binocular microscope (x 10 magnification) connected to a digital photomicrograph (positioned at x 5000 magnification).



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Result Sex ratio Two hundred and thirty–five females

and one hundred and three males making up 338 specimens of Macrobrachium vollenhovenii were examined during a nine months monthly study.

Out of this number, the male specimens made up 31.0 % while the females made up the remaining 69.0 %.

A 1:2:3 male to female sex ratio was established.

Length–weight relationship The length weight relationship

revealed that the longest male specimen measured 16.80 cm in total length and 86.7 g in body weight.

The shortest specimen was in the size class of 0.10–0.60 cm. It measured

0.52 cm in mean total length and 5.61g in mean body weight.

A specimen in the size class of 1.30–1.90 cm for females measured 7.08 cm in mean total length and 14.95 g in mean body weight.

Another specimen caught within the size class of 0.70–1.20 cm for females was 0.84 cm in mean total length and 10.6 g in mean body weight.

The highest numbers of specimens were caught within the size class, 0.70–1.20 cm for both sexes while the least number was caught in the size class 8.5–9.0 cm for both sexes (Table 1).

The regression coefficient ‘r’ was significant for female (0.644) and male (0.901) specimens at P < 0.05, (Figures 2 and 3).

Table 1 Percentage occurrence of specimens at different stages of maturity in the various size

groups Male Female Size Class (TL cm) F 1 11 111 IV V F 1 11 111 IV V 04–5 4 100.0 00 00 00 00 8 100.0 00 00 00 00 06–7 5 80.3 19.7 00 00 00 120 83.9 16.1 00 00 00 08–9 33 56.5 32.5 8.2 2.8 00 39 9.7 90.3 00 00 00 10–11 30 42.3 20.5 14.4 22.8 00 35 40.8 41.1 14.8 3.3 00 12–13 22 60.2 20.5 9.0 1.1 9.2 24 50.2 30.7 10.0 9.1 00 14–15 3 00 68.3 00 31.7 00 4 00 29.3 70.7 00 00 16–17 4 00 00 50.0 50,0 00 3 00 00 00 60.8 39.2 18–19 1 00 00 100.0 00 00 2 00 00 100.0 00 00 >20 1 00 00 00 00 100.0 0 00 00 00 00 00 Total 103 235

Key: F= frequency of occurrence

Figure 2. Log of total length versus body weight for Female Macrobranchium

vollenhovenii



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Figure 3. Log of total length versus body weight for Male Macrobranchium vollenhovenii

Condition Factor Generally, mean condition factor

(K) was higher in juvenile prawns than in adults.

Within the juvenile males, a high condition factor 3.99 was observed when the mean total length (TL) was 0.52 cm, whereas, a mature male with mean total length 1.44 cm, had a lower mean condition factor 0.67.

The females also showed the same pattern of higher condition factor in juvenile than in adults.

Size at first maturity Table 1 show that the specimens

were grouped into 2 mm total length size class, to enable the percentage of occurrence computation of various stages of maturity.

Ovaries and testis in maturity stages 1 to 11 were classified as immature, while those in stages 111 to V were classified as matured.

The table revealed that the size class of 4.0 to 7.0 cm had immature specimens for males.

While those for females appeared in the size class of 4.0 to 9.0 cm.

In the size class of 14–15 cm, only three males were caught, while in the size class of 18–19 cm, only two females were caught.

The biggest/largest spent male specimen was caught in August.

In the size class of 10–15 cm (for male) and 10–13 cm (for female), the percentage of immature specimens were more than the matured ones.

Granted that some specimens mature at a size class of 8–9 cm for males and 10–11 cm for females, maturity of the specimens attained maturity at a size class of 10–11 cm.

The size at first maturity was therefore 9 cm for males and 10 cm for females.

The distinctions amongst different stages of maturity were based on the size of the sex cells, their staining affinity, presence/absence of yolk globules and the general changes in somatic development.

Five stages of gonad developments were observed.

The stages are shown in Table 2, plate’s M1–M5 and F1–F5.



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Table 2 Gonad developmental stages

Maturity stages

Male Female

Immature dominant phase (undeveloped). March

Spermatogonia cells appear compact under the digital photomicrograph. Testis is physically translucent in color. Plate M1

Ovary small and translucent (white) and difficult to distinguish through the carapace. Oogonia cells are aggregated, as seen through the microscope. Plate F1.

Immature non–dominant phase (developing). April to May

Cells are visible without aids, graduating into spermatocyte stage. Note the presence of the melanomacrophage centers and lacanae cells in Plate M2.

Oogonia cells have advanced into the oocyte stage. Developing ovaries larger, opaque, and yellowish, with scattered melanophores over the surface; Their nucleolus is not visible. Plate M2.

Pre–vitellogenic phase. June to July

Dominated by black dots representing spermatocytes and spermatids. Gonad is physically light orange in colour. Plate M3

Vitilogenesis shown by eosin stained oocytes. Pre–vitilogenic oocytes with basophilic cytoplasm. Nucleus appears. Vesicular bodies appear with chromatin clumps. Ovary is nearly ripe. Its physically dark orange in color. Plate F3

Mature spawning phase. August to September

Dominated by Spermatids and spermatozoa cells. Gonad is physically light brown in colour. Plate M4

Numerous cortical rods. Follicle cells appear. This is the ripe spawning stage. Characterized by fully matured ovum whose centre is opaque. It bears very thin cell wall. Ripe ovaries, filling virtually the whole space among other organs. This is the vitellogenic stage. Ovary is physically light yellow in color. Plate F4

Hydrated/spent phase. October to November.

This is the spent or hydrated phase, characterized by irregularly shaped/empty lumens that look flaccid after the release of spermatozoa. Gonad is physically dark brown in color.

Post vitellogenic stage. It is characterized by shrunken and irregularly shaped cells observed after ova release. Degeneration of oocytes by the appearance of vacuoles. Ovary is physically dark yellow in color. Plate F5

Gonado somatic index (gsi) The gonado somatic index revealed

that the somatic activity in the sexes were high in the month of March: 0.17 for males and 0.29 for females, indicating not only the first peak period, but also continuation of yolk deposition and build-up of other sex cells. This activity decreased in the month of April: 0.11 for males and 0.13 for females. Soon after these decreases, there was another gradual increase in gonad development in subsequent months. Somatic activities peaked again in the month of July as shown for both sexes:

0.60 for females and 0.50 for males. A higher mean gonado somatic index was recorded for the females (0.25) than the males (0.19) (Table 3).

There was a sharp decreased in gonado somatic index during the months of April and August for both sexes, which characterized the months immediately after the release of ova/spermatozoa. From the pattern of somatic activity, it could be concluded that the specimens spawned in March and July and recovered in April and August respectively (Table 3).

Table 3 Gonado Somatic Index of specimens used for the study

Months Male Female March 0.17 0.29 April 0.11 0.13 May 0.25 0.30 June 0.28 0.38 July 0.50 0.60 August

0.20

0.24 Sept

0.14

0.20 Oct

0.08

0.14 Nov

0.05

0.07 Mean 0.19 0.25



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Male maturity stages

Mag. X 400 Sp1=Spermatogonia; Sp2=Spermatocyte; Sp3=Spermat id; Sp4=Spermatozoa; Sp5=Empty irregular cells.

Female maturity stages

Mag. X 400 O= Oogonia; NR=Nuclear ring; BB=Balbiani body; N=Nucleus; CV=Cortical vessicle; DF=Disintegrating follicle; MO=Matured oocyte; CR=Cortical rod and OC=Ovarian cavity.

Fecundity Fecundity estimates varied between

6,140–20,113 eggs. The largest females had the highest body weight.

Table 4 Means of parameters used for the study

Month Mean Total Length (cm)

Mean Body Weight (g)

Mean Fecundity

Mean relative Fecundity

March 1.08 22.13 6,644 3,951 April 1.16 20.71 – – May 1.19 20.34 – – June 1.28 30.85 18,775 13,789 July 1.39 33.45 20,113 15,546 August 1.37 20.00 6,140 2,213 September 1.41 18.41 – – October 1.23 17.37 – – November 1.04 17.41 – –



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The highest mean fecundity was

recorded in the month of July. The reverse was the case in the

month of August, when the lowest mean (body weight/fecundity) were recorded (Table 4).

DISCUSSION Sex Ratio The sex ratio observed during the

present study, revealed that the females were significantly more than the males.

Similar sex ratio differentials were observed by GEORGE and RAO, 1967 in respect to Penaeus. indicus, Metapenaeus dobsoni, Machrobrancium affinis and Parapenaeopsis stylifera from the trawl catches off Cochin. These observations were not in line with the reports of MENON,

1957; INYANG, 1981 and MARIOGHAE, 1982 where the sex ratio was the same.

The reason for the present observation may be that the females were better prone or vulnerable to catch in nature than the males who migrate into deeper waters soon after spawning.

According to TAWARI–FUFEYIN et al., 2005 sex ratios may not always be static, as they vary from season to season or from year to year within the same population.

Length Weight Relationship/

Maturity stages Weight relationship revealed a

higher body weight increase in male than in female specimens.

This observation agrees with that expressed by EDOKPAYI, 1990.

Differences in weight relationships have been attributed to sex, season and time of breeding MARIOGHAE, 1982.

A low size at maturity observed in the present study was not comparable with that reported for female (15.7 cm total length) P. semisulcatus in the Bushehr area KING, 1995.

That of the same spp harvested from the Gulf of Carpentaria CROCOS, 1987 was even higher (17.0 cm) than that reported by KING, 1995.

Both reported estimates were comparable with each other, considering the geographic and temporal separation of their studies.

Fishing pressure was reported to reduce the size of not only target species but also their size at first maturity.

However, according to SPARRE and

VENEMA, 1992 fishing pressure apparently has influence on organisms, although to a lesser effect on short–lived species such as crustaceans or cephalopods.

Only the size at first maturity as reported by CROCOS, 1987 for female specimens of P. semisulcatus (13.4 cm TL) and the length at 50% maturity 17.2 cm TL could be compared to that of present study.

Condition Factor In the present study, a number of

berried prawns were found with empty stomachs which agree with those of the present study.

Food availability influenced condition factor in this study because the juveniles, who were more active and so scavenged more food, were in a better condition than was otherwise the case for the adults.

This is a confirmation of an earlier statement where the berried specimens had empty stomachs.

High condition factor was recorded during the rainy season months (May–August) than during the dry season months (March–April) as was also reported by earlier researchers INYANG, 1981;

MARIOGHAE 1982; HLA and OHTOMI, 2005; MURPHY and

AUSTIN 2005. It could be that biological

interactions experienced in intra–species competition for food and space were probably the reasons for the reduction in the value of condition factor during the dry season months.

These observations were in line with those of ZAR, 1984.

Variation of condition factor in respect to size class was not uniform,



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indicating that their well–being did not depend on size. This was also the observation of HLA et al., 2005 and TAWARI – FUFEYIN

et al., 2005. Gonado–Somatic Index Generally, increases were

observed in the cyclical changes in gonad developments; evident in the monthly changes of gonado somatic index, MURPHY and AUSTIN, 2005 in both sexes throughout the study period.

During the immature stage, the germ cells (oocytes and spermatogonia cells) gradually enlarged.

This was the reason for the increase in gonado somatic index which climaxed in March and July and decreasing soon after.

The variation in gonado somatic index followed the same pattern in both sexes. This observation was also evident in the report of HLA and OHTOMI, 2005.

During this study, except in September, (March to August) where maturing as well as mature individuals were observed, those in premature condition were observed in February, very few in April and May and altogether absence from June, July and September.

It can therefore be said that the period from October to February was the resting/immature period for this species, at which time they migrated into deeper waters.

Fecundity Fecundity estimation in this study

ranged between 5,540 and 22,740 an observation not comparable with those of MARIOGHAE, 1995 where fecundity was 1,000–10,000 for Macrobrachium macrobrachion and 45,000–400,000 for Macrobrachium vollenhovenii.

The reasons adduced for the differences in fecundity could be attributed to differences in season, size of the specimens and or habitat where they were caught INYANG, 1981; MARIOGHAE,

1982; HLA and OHTOMI 2005; MURPHY and AUSTIN 2005. Few gravid females were caught in

March (dry season) while an appreciable number was caught in June and July during the rains.

Abundant food available during the rainy season was probably the reason for appreciable number of spawner during that time.

Only the rainy season spawning activity in the present study was in line with those reported INYANG 1981; MARIOGHAE

1982; HLA and OHTOMI; 2005; MURPHY and AUSTIN 2005. In contrast to the present observation, NAGABHUSHANAM et al., 1987 reported that rainy season reduced breeding activity in M affini.

Again PILLAY and NAIR, 1971, reported that the rainy season was a constraint to breeding for estuarine and shore crabs.

Observations by GYANANATH, 1982;

VICTOR, 1984, SAROJINI and RAJANI, 1987 (that rainfall contributed very little to breeding activities of fresh water prawns) did not support the present investigation.

Spent ovaries were observed in the catches in April and August, and together with the presence of more than one size of oocytes in the ovary, suggests continuous breeding pattern and confirms that the specie was a partial spawners.

In the present study, spermatogenic tubules were observed to contain two or three developmental stages.

This was also the situation with M. choprai, and different species of crab FASTEN, 1926; RYAN, 1967; SENGUPTA and CHATTERJI,

1976 equally noted the presence of both spermatocytes and spermatids) SENGUPTA

and CHATTERJI, 1976 in the tubule and reported that of all these cells, the spermatocytes were always at the same stage of differentiations. SAROJINI and GYANANATH, 1984 made a similar observation in M. larnarrel. References 1. Based on the review of the ecological

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Received: September 2, 2012

Accepted: September 29, 2012

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