82-95

Transcription

82-95

Advances in Environmental Biology, 10(5) May 2016, Pages: 82-95
AENSI Journals
Advances in Environmental Biology
ISSN-1995-0756
EISSN-1998-1066
Journal home page: http://www.aensiweb.com/AEB/
Reproduction cycle of Donax trunculus
(Mollusca, Bivalvia) in the gulf of Annaba
(Northeast Algeria)
1Imed
Hafsaoui, 1Rym Bouaziz, 2Brahim Draredja, 3Hayet Beldi
1
Department of Marine Science. University Badji Mokhtar – Annaba. 23000 Annaba, Algeria
Laboratory of Coastal and Marine Ecobiology. University Badji Mokhtar – Annaba. 23000 Annaba, Algeria
3
Laboratory of Applied Animal Biology. University Badji Mokhtar – Annaba. 23000 Annaba, Algeria
2
Address For Correspondence:
Hafsaoui I., Department of marine Sciences. University Badji Mokhtar – Annaba. 23000 Annaba, Algeria.
E-mail: [email protected]
This work is licensed under the Creative Commons Attribution International License (CC BY).
http://creativecommons.org/licenses/by/4.0/
Received 22 March 2016; Accepted 28 May 2016; Available online 12 June 2016
ABSTRACT
This study is interested in knowing the cycle of oogenesis of the bivalve Donax trunculus colonizing Shallow waters of Sidi Salem and
Echatt beaches in the Gulf of Annaba, during 2012. The histological analysis of gonads was complemented by changes in the condition
index and associated with the temperature and salinity measured monthly. Thermal extremes varied from 15.02 (February) to 29.04 °C
(July). The haline variations are lower and range between 37.07 (July) and 38.04 psu (September). The condition index fluctuates between
0.20 (July and August) and 0.32 (April). The spatial differences of this index are in favor of Sidi Salem site where individuals have higher
biomass compared to those of Echatt. This would be in relation to trophic factor of this bivalve. Indeed, this site is close to the discharge of a
chemical fertilizers complex which fertilizes the waters of the beach. During the period of sexual activity; fluctuations in the sex ratio at
both sites generally show a predominance of males. The study of oogenesis through histological analyzes in D. trunculus reveals that the
reproduction begins in December (stage I) with a first emit of gametes in March (stage III). This sexual activity continues until September,
when we witness the highest rate of emission of gametes into the surrounding environment. Moreover, monitoring of oogenesis through
microscopic observations shows a lag in the oocyte development between the two sites. Thus it detects a delay of the evolution of stage III
during the month of March in the ovaries of individuals taken from the Sidi Salem site. This phenomenon could be explained by the
exposure of the site to different sources of pollution from the port and other effluents carrying sewage and industrial waste in the region.
KEYWORDS: Bivalvia, Donax trunculus, Reproduction, Oogenesis, Gulf of Annaba.\
INTRODUCTION
Donax trunculus is a strictly intertidal bivalve that colonizes sandy beaches [55] of the AtlanticMediterranean coasts [2]. Its geographical distribution extends from the French Atlantic coast [43] to the coasts
of Senegal [37]; it is widespread in the Mediterranean [8] and Black Sea [37]. This species inhabits the turbulent
environments on beaches exposed to the rhythms of the tides or under intense wave action [20]. In these
environments, D. trunculus is able to reach very high densities [40,84] Moreover, this Donacidae has a tolerance
to changes in physical and chemical environmental factors; at the condition that these changes are not too brutal
[53].
The studies of ecology, biology and population dynamics of D. trunculus are several either at the
Mediterranean level [59,17,2,3,27,64,69,25,26] or Atlantic [3,43,6,8,42,9]. But those who treat reproduction are
Copyright © 2016 by authors and Copyright, American-Eurasian Network for Scientific Information (AENSI Publication).
83
Imed Hafsaoui et al, 2016
few; in the Mediterranean [60,5,58,31,83,15] and in the Atlantic [7,40]. D. trunculus is very abundant in the
Gulf of Annaba [11] and is the subject of artisanal exploitation in the region. Despite its abundance and its use,
no research on the reproductive cycle was performed [46] which dealt with the biochemistry of the gonads of
this bivalve). Other works in the Gulf of Annaba were interested to use this mollusk species as bioindicator of
pollution [12,77,75,33,1,44,79,10,76]. The ecological and economic importance of D. trunculus and lack of
information about the trigger period of reproduction and the arrival of young recruits motivated us to undertake
this investigation. Moreover, the study of the sexual cycle in organisms is of great importance for population
dynamics to better knowledge and understanding of specificity and biogeographical distribution. In the case of
this bivalve used for commercial purposes, the study of the sexual cycle could enable improved performance by
determining the period during which the Edge shell may be harvested.
This work treats the oogenesis of D. trunculus and its reproductive cycle in the Gulf of Annaba; to better
understand the biology and dynamics of this bivalve in its natural environment and to preserve the natural
harvested stock.
MATERIALS AND METHODS
Study zone:
The Gulf of Annaba is located at the north-east of Algeria between two capes; Western Cape de Garde
(7°16’E - 36°68’N) and Cape Rosa to the east (8°15’E - 36°38’N), distant of about 40 km. The maximum depth
does not exceed 65 m. The Gulf receives fresh water through two rivers; Mafragh (South) and Seybouse
(Southwest) (Fig. 1).
N
Mediterranean
Sea
Algeria
Cape Garde
Cape Rosa
Gulf of Annaba
Annaba
S.S
Seybouse
river
Ech.
Mafragh
river
Mellah
lagoon
10 Km
Fig. 1: Geographical position of the Gulf of Annaba and location of the sampling sites (S.S: Sidi Salem, Ech.:
Echatt).
Measurements of temperature and salinity:
The temperature and salinity were measured by a thermo-salinometer type "WTW Cond.197i" thermal
accuracy and respective haline 0.01°C and 0.01 psu. Measurements were performed monthly and parallel to the
sampling of the bivalve Donax trunculus in the two study sites.
Sampling of Edge shell:
Sampling was done manually with a large rake commonly called "Cope". It is dragged for a distance of 5 m
by the angler which travels backwards at depths varying between 1 and 1.5 m. Individuals are thus recovered in
a bag made of a net of 5 mm mesh size; in order to collect individuals of different sizes.
Condition index:
In an environmental monitoring study, condition index (C.I) assesses the general condition of the animal,
its bioenergetic status and reproductive status [68]. It was measured for 50 individuals per site during the study
period after passage in the oven at 60°C for 48 h. The condition index (C.I) was calculated according to the
equation of Beninger [14]:
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C.I = (dry flesh weight/weight of the shell)*100
Its monitoring allows to know the stages of gametogenesis and periods of gamete emissions.
Sex ratio:
The distinction between sexes is only possible during the period of gametogenesis. Indeed, the
differentiation between sexes is based on the color of the gonads. Those of male have a viscous appearance,
whitish to orange, while the ovaries in females are dark bluish. Thus, the two valves are opened and carried to
the macroscopic observations for determining the sex of individuals.
The relative frequency of the sexes (males, females and immatures) is calculated and expressed as a
percentage. Gender percentages were determined during the months of sexual activity (i.e, when the sex of
individuals is recognizable).
Oogenesis study:
Collection and preservation of gonads:
After collecting the samples, mature female individuals are selected (> 10 mm) freshly collected. To take
the female gonad, both valves are opened, then using a micro-scissor, ovarian are dissected. Collected gonads
are preserved in formalin solution 10%, for further use in the histological study.
Histological techniques:
The steps of histological preparation of oogenesis D. trunculus are those described by Martoja and Martoja
[54], using an automated apparatus "Fabric Tech. 11". The thinning is done by immersion in 3 baths of xylene
[47,21]. This is followed by soak and inclusion in both paraffin baths to 60°C. Semi thin sections (2-3 microns)
are achieved through a microtome from the blocks obtained. These sections were mounted on slides held in a
small drop of gelatin, and placed on a hotplate at 60°C to remove the fold. They are then dewaxed to hydrate the
tissue; they are subject to hematoxylin-eosin baths.
Scale adopted for the study of oogenesis:
As in most bivalves, in D. trunculus, we identified four stages of the cycle of reproduction. These stages
have been identified and characterized based on morphological and cytological criteria described by several
authors: Mouneyrac et al. [62] in the semelidae Scrobicularia plana, Delgado and Pérez-Camacho [30] in the
veneridae Ruditapes decussatus, Gaspar and Monteiro [39] in the pharidae Ensis siliqua and the veneridae
Venus striatula and Gaspar et al. [40], Tlili [82] in the Donacidae Donax trunculus. These stages are as follows:
stage 0 (sexual rest), stage I (Resumption of genital activity), stage II (Gametogenesis) and stage III (genital
maturity).
Results:
Temperature and salinity:
Thermal water variations in both sample sites are important (Fig. 2), where the thermal extremes passes
from 15.02 (February, Echatt) to 29.04°C (July, Echatt), an amplitude of 14.02°C. The inter-station thermal
differences are generally low and hardly exceed 2.56°C (June). Generally temperatures of Sidi Salem waters are
slightly higher compared to those observed in Echatt (Fig. 2).
Fig. 2: Spatio-temporal variations of temperature in the surveyed stations during 2012.
The salinity fluctuations (Fig. 3) are less pronounced in comparison with those of the temperature. Indeed,
haline variations range between 35.71 (April, Echatt) and 38.05 psu (September Sidi Salem), an annual
amplitude of 2.34 psu.
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Fig. 3: Spatio-temporal variations in salinity at the surveyed stations during 2012.
Condition index:
Monthly monitoring of the condition index (C.I) in Donax trunculus taken from the two selected sites,
reveals significant spatio-temporal fluctuations (Fig. 4). The most important values of this index are in spring
and autumn with a peak of 8.40 (April, Echatt). Furthermore, the lowest values are recorded during the summer
season with a minimum of 4.98 (Jul, Echatt). In the beach of Sidi Salem, condition index goes from 5.66 (June)
to 8.28 (February). Overall, the values of this index show a slight increase for the individuals collected from the
beach of Sidi Salem; while reflecting slightly higher biomass compared to those of individuals collected in
Echatt.
Fig. 4: Monthly changes of condition index (C.I) in Donax trunculus fished in Echatt and Sidi Salem during
2012.
Sex ratio:
At Donax trunculus, sexual differentiation is possible only during the period of sexual activity. Therefore,
the sex ratio was calculated during this period which lasts from February to October, where sex is
macroscopically identifiable.
Overall, the relative frequencies show no difference between males and females in both study sites (Fig. 5).
However, in the range of Echatt, there was a slight dominance of males in February (56.64%) in July (56.50%)
and August (54.17%). While in the Sidi Salem beach, males dominate especially in August (61.82%) in
September (54.31%) and October (69.58%). Immature individuals are identified during the period which runs
from March to October. Their presence remains low (0.33 to 3.03%), except in July in Echatt (9%) and June at
the beach of Sidi Salem (20%).
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Fig. 5: Relative Frequency of sexes in both surveyed stations during 2012. (A: Echatt, B: Sidi Salem, M: male,
F: female, I: undifferentiated).
oogenesis study:
Histological examination of the female gonad Donax trunculus at both sites, allowed the identification of
four stages of gametogenic development: (1) sexual rest, (2) recovery of genital activity, (3) gametogenesis (4)
genital maturity. Morphological and cytological features of differentiation between these stages are:
Stage 0: sexual rest, oocytes are very small. There is a gonadal tissue mainly composed of a connective
tissue (Fig. 6). This phase is characterized by the accumulation of reserves and adipogranuleuses vesicular cells.
The cloak is so homogeneous and transparent. Therefore, it is impossible to identify the sex of D. trunculus at
this stage.
Stage I: Resumption of genital activity, Gonies multiply hence the increased volume of oocytes at the
expense of connective tissue. Sex of Edge shell can then be identified. oogonia can be observed on the wall and
some vitellogenic oocytes (Fig. 7). We can also see an appearance of follicles dispersed in connective tissue and
having oocytes maturation and glued to the wall. At this point, the mean oocyte diameter is about 79 microns.
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Fig. 6: Stage 0 "sexual rest" appearance of a reduced size acini within which we distinguish oogonia. A: Ovary
in October; B: Ovary in November; UC: undifferentiated cell; OG: oogonia. Scale bar: 87 µm.
Fig. 7: Stage I: "Resuming sexual activity." A: Ovary in December; B: ovary in January; C & D: Biometrics;
OA: adhered oocyte; OG: oogonia; VO: vitéllogénétique oocyte; CT: conjunctival tissue. Scale bar: 87
µm.
Stage II: Gametogenesis, we can observe on the wall of the pedunculate oocytes during detachment and
mature oocytes that acquire a polygonal shape; because of their overcrowding in the light of gonadal tubules.
During this phase, the mean oocyte diameter is about 120 microns. Connective tissue has shrunk in favor of
developing follicles (Fig. 8). At this point, individuals are easily excitable and release their gametes under the
action of external stimuli.
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Fig. 8: Stage II: Maturation of gonads. A: ovary in February; B: ovary in March; C & D: Biometrics; MO:
mature oocyte. Scale bar: 87 µm.
Stage III: Genital maturity, at this stage oocytes are partially or completely emptied of their contents.
Sometimes there is the presence of residual oocytes with an average diameter of about 122 microns, not issued
at the end of the cycle (Fig. 9). Acini have a torn appearance and will be resorbed by the body. Gonadal tissue
will be invaded again by the connective tissue. It is the restoration of the gonad (the gametic renewal takes
place). During this stage the germ activity stops completely. Important phenomena occurs in the cloak. The
follicles degenerate and crumble. Amoebocytes attack gametes not laid. The animal is again at the stage of
sexual rest.
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Fig. 9: Stage III: Maturation of oocytes and gamete emission. A: ovary in March (GX100) B: ovary in April
(GX400) C & D: Biometrics (GX400). RO: residual oocyte. Scale bar: 87 µm.
Discussion:
In this study the temperature readings performed at the beach of Sidi Salem and Echatt in a cycle [35],
reveal a certain similarity between the two sites where the temperature ranges between 15.02°C (February ) and
29.04°C (July) with an amplitude of 14.02°C. The comparison between the average temperatures of the two
sites shows that the waters of Sidi Salem are slightly warmer than 21.76°C to 21.28°C Echatt, probably due to
water dicharge used to cool the complex engines of (Fertial). These results are similar to the work done in the
bay of Annaba, where the amplitudes are close to 15°C [12,49,75,45]. Temperature is one of the major factors
that control reproduction of bivalve molluscs, it is considered the main factor affecting gonadal development in
marine bivalves. The food availability and the temperature are principal factors that affect the gametogenesis of
marine bivalves in general [29,26] and of Donacidae in particular [32,48]. For D.trunculus Ruiz-Azcon et al.
[71] show that at a temperature of 20°C the larvae have a large percentage of survival. Among other molluscs
numerous studies show that the temperature can affect the hatching of eggs of molluscs and their rate of sexual
maturity [63,15,23]. According to Serdar et al. [73], Gamete maturation and spawning in bivalves can continue
throughout the year, so the rest period will be reduced and may even be absent whenever trigger factors (i.e,
food availability and temperature) are appropriate for gametogenesis. In oysters Crassostrea gigas Lubert [52]
and Chavez-Villalba et al. [23] indicate a trigger of gametogenesis from a temperature (8-9°C) in the Channel
and 10°C in the Bay of Veys; with an induced emission of gametes by a threshold value (> 19-20°C). Indeed,
the Edge shell in the Gulf of Annaba starts the emission of gametes into the surrounding environment from the
month of March-April; parallel to the beginning of the warming of waters at the end of the winter and early
spring.
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The salinity of shallow waters is influenced by seasonal temperature variation; it depends on the
evaporation, and arrivals of fresh water whose flow rate can vary significantly depending on rainfall. In this
study the monthly salinity fluctuations do not show remarkable variation between the two study sites with
average salinity of 37.08 psu for the site of Sidi Salem and 37.05 psu for the site of Echatt. The salinity varies in
the range of 35.71 (April, Echatt) and 38.05 psu (September, Sidi Salem). These results are consistent with
other work in the Gulf of Annaba, the haline variations oscillate between 35 and 38 psu [12,38,75,45], and close
the work of Ounissi et al. [65], who report salinities between 32 and 37 psu. In the Gulf of Annaba, the haline
variations do not seem to influence either the trigger or the conduct of reproduction in Donax trunculus since
the differences of this ecological factor are relatively low during a cycle, taking into account the wet season and
the dry season. For D. trunculus, sexual cycle may be punctuated by seasonal climatic factors. Reproduction
would be triggered by sudden variations in temperature and / or salinity [53].
The condition index is one of the indices that affect the cycle of reproduction in marine organisms in
general and bivalve molluscs in particular. The study of this index can provide additional information on this
process [16,17]. Monthly monitoring of this index shows significant seasonal variations with an increase in
spring and autumn and a decrease in summer. These variations are due to several parameters including the
status of breeding and nutrition [18,78,66]. Indeed, the condition index is an indicator of the physiological state
of bivalves. The decrease in index values condition which begins in May coincides with the emission of
gametes, hence the importance of the gametes mass in the whole mass of individuals. Indeed, the issue of
gametes causes weight loss inducing lower values of the CI. On the other hand, the increase of this index which
begins in the fall may can be caused by a maturation or redevelopment of the ovary. Histological examination
of the gonads allowed us to confirm this. Several authors have observed similar results [5,56,83]. In the
Mediterranean, the condition index of Donax has strong values from February to June [2], reflecting the
expansion of the period of reproduction. According to Roméo et al. [70], the condition index is associated with
the water temperature and correspond to filling ratio of the shell by soft tissue and by that, informs on the
nutritional and physiological status of the animal. The change in the condition index may also be caused by
trophic conditions that are influenced by environmental conditions including pollution [28,57,35].
The study of the sex ratio in the population of Donax trunculus in the two study sites during the period of
sexual activity (February to October), allowed us to reveal differences over time and depending on location
where the strongest female domination is in April for the site of Echatt but shifts from one to two months in the
Sidi Salem site. The rest of the cycle is characterized by a slight predominance of males. Indeed, after laying the
percentage of females decreased, and there has been an increase in male individuals that extends through the
winter. These results are in agreement with those of Gaspar [40] in Portugal, Deval [31] in the Turkish coast of
the Marmara Sea in the eastern Mediterranean, from Tlili et al. [83] in Tunisia. Several works on D. trunculus
showed a slight predominance of males compared to females during the rest period and the early gametogenesis
[40,50,19]. The decrease in females after laying could be explained as differential mortality due to the effort of
reproduction [40]. However, Badino and Marchionni [5] in Italy report some balance and fairness between the
sexes.
The study of the sexual cycle of Donax trunculus in the Gulf of Annaba by histological analysis of female
gonads during the period which runs from January to December, allowed us to see that the gonadal development
starts from December in both sites. This period coincides with the cooling water (17.22°C in Sidi Salem and
Echatt 16.63°C). This demonstrates the importance of abiotic factors in the reproduction of marine invertebrates
in general [67], and in bivalve molluscs in particular [34,36,72]. According to Mouëza [61], low temperatures
triggers gonad maturation and the emission of gametes is undertaken when it increases until a proximate degree
of (20°C). Furthermore, the author reports that the temperature is not the only factor triggering the reproduction
or the only limiting factor. Moreover, this factor acts differently between adult individuals and young ones,
because in juveniles (from 8 to 16 mm), maturity intervene later than in adults [61].
In both study sites, the emission of gametes begins in March and continues through September with two
major spawning periods: an emission peak of gamete in spring and the other in summer. This period coincides
with the sharp decline in index condition values. These results agree with those reported by Tlili et al. [83] in
the Gulf of Tunis. They are also close to those of Mouëza and frenkiel-Renault [58] and Ansell et al. [3] also
note that two main shift laying periods; one in autumn and one in winter. However, we note that this
physiological activity is closely related to the location where this species lives. Moreover, in the same species in
Spain [81] and in Turkey [31] notes a single spawning period during the summer season. These differences can
be explained by changes in environmental conditions that affect the frequency and duration time of
gametogenesis. Furthermore, it should be noted that we have identified a delay (about a month) of stage III at
the Sidi Salem site. This time lag in this site near Seybouse river is induced by the negative effects of the
surrounding water pollutants and contaminants impacting consequently on the reproductive physiology of this
bivalve. This same finding was reported by Tlili [82], which noted a delay in stage III at a polluted site
compared to that which is less exposed to pollution. The negative influence of pollutants on the physiology of
reproduction in marine bivalves, has already been reported in several studies [80,13,41,74].
91
Conclusion:
The monitoring of the temperature and salinity in the two selected sites in the Gulf of Annaba during a
cycle (2012), shows that the thermal fluctuations of the water of the study area are important and under the
direct influence of changes in the atmospheric temperature. Overall, the beach of Sidi Salem more sheltered and
under the influence of cool waters of the engines of the complex Fertial, is characterized by relatively warmer
waters compared to waters of the beach of Echatt more exposed (roughest) to dominant winds of Northwest
sector.
However, warming waters in the late winter and early spring helps the first trigger (March) of gametes
emission in the surrounding environment. The haline variations are less important, with annual amplitudes not
exceeding 1.26 psu in the range of Echatt and 1.97 psu in Sidi Salem. These small variations do not allow us the
highlight of this parameter in the course of reproduction for the bivalve Donax trunculus in the Gulf of Annaba.
The study of the condition index of D. trunculus showed that Sidi Salem site individuals possess a higher
body weight compared to those of Echatt with decreasing values of this index during the period of release of the
gametes in both sites of the study. Indeed, the weakest condition index values coincide with the release of the
gametes in the spring and summer, hence the importance of the gametes mass in the body mass of individuals
for this bivalve Donacidae.
For Donax trunculus, sexual differentiation is possible only during the period of sexual activity. In general,
both sexes are equitably distributed and the differences between sexes across a cycle are low: <10%,
exceptionally 22.81% (Echatt, July) and 38.16% (Sidi Salem, October). Overall, the immature individuals are
poorly represented in the two sites. Histological study of oogenesis of D. trunculus, allowed us to locate the
breeding season which runs from February to October. This study also allowed us the timing and conditions of
gonadal maturation and their emission into the environment. Furthermore, this analysis allowed us to detect a
slight delay in the development of oogenesis in both sites. Indeed, there is a delay of stage III, for individuals
taken from the beach of Sidi Salem. This could be caused by negative effects of various pollutants from
discharges from Wadi Seybouse (industrial pollution) and port (wastewater and hydrocarbons) on the
reproductive physiology of this bivalve.
ACKNOWLEDGEMENTS
This study was supported by the Algerian Fund for Scientific Research and by the Ministry of High
Education and Scientific Research of Algeria (CNEPRU project F.1120120112).
REFERENCES
[1] Amira, A., K. Sifi and N. Soltani, 2011. Measure of environmental stress biomarkers in Donax trunculus
(Mollusca, Bivalvia) from the gulf of Annaba (Algeria). European Journal of Experimental Biology, 1(2):
7-16.
[2] Ansell, A.D. and A. Bodoy, 1979. Comparison of events in the seasonal cycle for, Donax vittatus and D.
trunculus. In: Cyclic phenomena in marine plants and animals, Proc. Europ. mar. Bisl. Symp., Naylor, E. et
Hartno, R.G. (eds). Pergamon Press, Oxford & New York: 191-198.
[3] Ansell, A.D., L. Frankiel and M. Mouëza, 1980. Seasonal changes in tissue weight and biochemical
composition for the bivalve Donax trunculus L. on the Algerian coasts. J. exp. mar. Biol. Ecol., 45: 105116.
[4] Ansell, A.D. and F. Lagardère, 1980. Observations on the biology of Donax trunculus and Donax vittatus
at Ile d'Oléron (French Atlantic Coast). Mar. Biol., 57(4): 287-300.
[5] Badino, G. and B. Marchionni, 1972. Neurosecretion and gonad maturation in a population of Donax
trunculus L. from Leghorn (Italy). Bull. Zool., 39: 321-326.
[6] Bayed, A. and J. Guillou, 1985. Contribution à l’étude des populations du genre Donax : La population de
D. trunculus L. (Mollusca, Bivalvia) de Mehdia (Maroc). Ann. Inst. Oceanogr., Paris, 61(2): 139-147.
[7] Bayed, A., 1990. Reproduction de Donax trunculus sur la côte atlantique marocaine. Cah. Biol. Mar., 31:
159-169.
[8] Bayed, A., 1991. Étude écologique des écosystèmes de plages de sable fin de la côte atlantique marocaine :
Modèles de zonation, Biotypologie, Dynamique des populations. Thèse Doct. d'État. Univ. Mohammed V,
Rabat, pp: 229.
[9] Bayed, A., 1998. Variabilité de la croissance de Donax trunculus sur le littoral marocain. CIHEAMOptions méditerranéennes, 35: 11-23.
[10] Belabed, S. and N. Soltani, 2013. Acute toxicity of cadmium on Donax trunculus acetylcholin-esterase,
glutathione S-Transferase activities and pattern of recovery. Eur. J. Exp. Biol., 3(2): 54-61.
[11] Beldi, H., F. Gimbert, S. Maas, R. Scheiffler and N. Soltani, 2006. Seasonal variations of Cd, Cu, Pb and
92
Zn in the edible mollusc Donax trunculus (Mollusca, Bivalvia) from the gulf of Annaba, Algeria. African
Journal of Agricultural Research, 1(4): 085-090.
[12] Beldi, H., 2007. Etude de Gambusia affinis (Poisson, Téléostéen) et Donax trunculus (Mollusque,
Pélécypode): écologie, physiologie et impacts de quelques altéragènes. Thèse de Doctorat d’état en
Sciences de la Mer, option: Biologie et écologie marines, Université d’Annaba., pp: 105.
[13] Benelli, A., R. Bacchetta, G. Vailati, S. Galassi and A. Provini, 2001. DDT contamination in Lake
Maggiore (N. Italy) and effects on zebra mussel spawning. Chemosphere, 45: 409-415.
[14] Beninger, P.G., 1984. Seasonal variations of the major lipid classes in relation to the reproductive activity
of two species of clams raised in a common habitat: Tapes decussatus L. (Jeffreys, 1863) and T.
philippinarum (Adams & Reeve, 1850). Exp.Mar. Biol. Ecol., 79: 79-90.
[15] Benmassaoud, F., 1987. Étude de deux mollusques lamellibranches: Cerastoderma edule (L., 1767) et
Venerupis decussata (L., 1758) dans l’estuaire du Bou-Regreg. Thèse de 3 ème cycle, Univ. Mohammed V,
Rabat., pp: 119 (+ annexes).
[16] Beukema, J.J. and W. De Bruin, 1977. Seasonal changes in dry weight and chemical composition of soft
parts of the telline Macoma balthica in the Dutch Wadden Sea. Neth. J. Sea. Res., 11(1): 42-55.
[17] Bodoy, A. and Massé, 1978. Quelques paramètres permettant de suivre la production organique d'un
Mollusque bivalve au cours d'un cycle saisonnier. Publ. sci. tech. CNEXO, Actes colloq., 7: 753-766.
[18] Boscolo, R., M. Cornello and O. Giovanardi, 2003. Condition index and air survival time to compare three
kinds of Manila clam Tapes philippinarum (Adams & Reeve) farming systems. Aquacult. Int., 11: 369-374.
[19] Boussoufa, D., N. Ghazali, I. Rabeh, N. Soudani, J. Carlos Navarro and M. El Cafsi, 2015. Reproductive
cycle and gonad development of the commercial clam Donax trunculus from the Bay of Tunis (Northern
Tunisia, South Mediterranean coast): effects of environmental variability. Cah. Biol. Mar., 56(4): 369-380.
[20] Brown, A.C. and A. McLachlan, 1990. Ecology of sandy shores. Elsevier Publishers. Amsterdam. pp: 328.
[21] Bucke, D., 1972. Some histological technique applicable to fish tissues. In: Diseases of fish, procceding of
symposium. (L.E. Mawdesley-Thomas, ed). Acad . Press and Zool. Soc., New York and London, 30: 135189.
[22] Chávez-Villalba, J., J. Barret, C. Mingant, J.C. Cochard and M. Le Pennec, 2003a. Influence of timing
broodstock collection on conditioning, oocyte production, and larval reaing of the oyster, Crassostrea gigas
(Thunberg), at a six production sites in France. J. Shellfish. Res., 22(2): 465-474.
[23] Chávez-Villalba, J., J.C. Cochard, M. Le Pennec, J. Barret, M. Enriquez-Diaz and C. Caceres-Martinez,
2003b. Effects of temperature and feeding regimes on gametogenesis and larval production in the oyster
Crassostrea gigas. J. Shellfish. Res., 22(3): 721-731.
[24] Çolakoğlu, S. and M. Palaz, 2014. Some population parameters of Ruditapes philippinarum (Bivalvia,
Veneridae) on the southern coast of the Marmara Sea, Turkey. Helgoland Marine Research, 68: 539-548.
[25] Çolakoğlu, S. and A. Tokaç, 2011.Some population parameters of the wedge clam (Donax trunculus L.,
1758) in the west Marmara Sea. Ege Journal of Fisheries Aquatic Sciences, 28(2): 65-70.
[26] Çolakoğlu, S., 2014. Population Structure, Growth and Production of the Wedge Clam Donax trunculus
(Bivalvia, Donacidae) in the West Marmara Sea, Turkey. Turkish Journal of Fisheries and Aquatic
Sciences, 14(1): 221-230.
[27] Costa, C., M. Bianchini, P. Ceccarelli, P. Orecchia, E. Rambaldi and L. Volterra, 1987. Indagine sui
molluschi bivalvi di interesse commerciale (telline, cannolicchi e vongole) delle coste della Toscana, del
Lazio e della Campania, 1985–1987. Quaderni dell’Istituto di Idrobiologia e Acquacoltura G. Brunelli, 7:
58.
[28] Damiens, G., M. Gnassia-Barelli, F. Loqués, M. Roméo and V. Salbert, 2007. Intgrated biomarker response
index as a useful tool for environmental assessment evaluated using transplanted mussels. Chemosphere.,
66: 574-583.
[29] Delgado, M. and A. Pérez-Camacho, 2007. Comparative study of gonadal development of Ruditapes
philippinarum (Adams and Reeve) and Ruditapes decussatus (L.) (Mollusca: Bivalvia): Influence of
temperature. Scientia Marina, 71: 471-484.
[30] Delgado, M. and A. Pérez-Camacho, 2005. Histological study of the gonadal development of Ruditapes
decussatus (L.) (Mollusca, Bivalvia) and its relationship with available food. Sci. Mar., 69: 87-97.
[31] Deval, M.C., 2009. Growth and reproduction of the wedge clam (Donax trunculus) in the Sea of Marmara,
Turkey. Journal of Applied Ichthyology, 25: 551-558.
[32] Dhaoui Ben Kheder, R., N. Aloui Bejaoui and M. Le Pennec, 2003. Cycle sexuel de Donax trunculus
(Mollusque, bivalve) du Golfe de Tunis. Bulletin de la Société Zoologique de France, 128: 9-20.
[33] Drif, F. and C. Abdennour, 2010. Trace Metals in the Mussel Donax trunculus Linnaeus 1758 from urban
and industrial contaminated locations. J. Appl. Sci. Res., 6(12): 2063-2067.
[34] Enriquez-Daiz, M.R., 2004. Variabilité et bioénergétique de la reproduction chez l’huître creuse
Crassostrea gigas. Thèse Doct. Univ. Bretagne, pp: 216.
[35] Erica, G., A.D. Claudia and A.A. Oscar, 2010. Biomarkers and heavy meatl bioaccumulation in mussels
93
transplanted to coastal waters of the Beagle Channel. Ecotoxicology and Environmental Safety, 73: 270279.
[36] Fabioux, C., S. Pouvreau, F. Le Roux and A. Huvet, 2004. The oyster vasa-like gene: a specific marker of
the germline in Crassostrea gigas. Biochem. Bioph. Res. Com., 315: 897-904.
[37] Fischer, W., G. Bianchi and W.B. Scott, (eds) 1981. Fiches FAO d’identification des espèces pour les
besoins de la pêche. Atlantique centre - est : zone de pêche 34, 47 (en partie) J. Canada Fonds de Dépôt.
Ottawa, Ministère des Pêcheries et Océans Canada, en accord avec la FAO, vol. 6 (Bivalves).
[38] Fréhi, H., 2007. Hydrobiologie de la baie d’Annaba: Etude descriptive des Dinoflagellés et écologie des
espèces nuisibles. Thèse de Doctorat d’état, Université d’Annaba, Algérie, pp: 166.
[39] Gaspar, M.B. and C.C. Monteiro, 1998. Reproductive cycles of the razor clam Ensis siliqua and the clam
Venus striatula of Vilamoura, southern Portugal. J. Mar. Biol. Ass. UK. 78: 1247-1258.
[40] Gaspar, M.B., R. Ferreira and C.C. Monteiro, 1999. Growth and reproductive cycle of Donax trunculus L.,
(Mollusca: Bivalvia) off Faro, southern Portugal. Fish. Res., 41: 309-316.
[41] Gauthier-Clerc, S., G. Pellerin, C. Blaise and F. Gagné, 2002. Delayed gametogenesis of Maya arenaria in
the Saguenay fjord (Canada): a consequence of endocrine disruptors? Comp. Biochem. Physiol., C131:
457-467.
[42] Guillou, J. and A. Bayed, 1991. Contraintes du milieu sur les populations de Donax trunculus L. et Donax
venestus Poli, du littoral atlantique marocain. Oceanologica Acta., 14(3): 291-298.
[43] Guillou, J. and Y. Le Moal, 1980. Aspects de la dynamique des populations de Donax vittatus et Donax
trunculus en baie de douarnenez. Ann. Inst. Océanogr., 56(1): 55-64.
[44] Hamdani, A. and N. Soltani-Mazouni, 2011. Changes in Biochemical Composition of the Gonads of Donax
trunculus L. (Mollusca, Bivalvia) from the Gulf of Annaba (Algeria) in Relation to Reproductive Events
and Pollution. Jordan Journal of Biological Sciences, 4(3): 149-156.
[45] Hamdani, A., 2012. La reproduction chez un mollusque bivalve Donax trunculus L. dans le golfe
d’Annaba. Rapport entre les aspects cytologiques et biochimiques et la pollution. Thèse Doct. Univ.
Annaba, pp: 165.
[46] Hamdani, A., N. Soltani Mazouni and N. Soltani, 2014. Quantitative and Qualitative Analysis of Proteins
in Gonads of Donax trunculus from the Annaba Bay: Effects of Site, Season and Sex. Adv. Environ. Biol.,
8(13): 740-749.
[47] Headden, G.F. and M.C. Williams, 1968. A comparison of embedding waxes. J.Med. Lab. Technol., 25:
250-262.
[48] Herrmann, M., C. De Almeida Rocha Barreira, E. Arntz, W. Rgen, J. Laudien and P.E. Penchaszadeh,
2010. Testing the habitat harshness hypothesis: reproductive biology of the wedge clam Donax Hanleyanus
(Bivalvia: Donacidae) on three argentinean sandy beaches with contrasting morphodynamics. Journal of
Molluscan Studies, 76: 33-47.
[49] Khati Hadj moussa, W., 2009. Etude de la qualité des eaux du golfe d'Annaba par l'utilisation d'un
mollusque sentinelle, la moule Perna perna : essai in vivo et in situ. Application a la biosurveillance de
l'environnement marin. Thèse de doctorat. Univ. Annaba. Laboratoire écosystème marins et littoraux, pp:
87.
[50] La Valle, P., 2005. Donax trunculus (Bivalvia: Donacidae) as a biological indicator of coastal dynamic and
sedimentary balance. École doctorale de Sciences de l’environnement Université de Rome "La Sapienza":
Rome, pp: 148.
[51] Laziak, T.A. and T.C.E. Barnard, 1995. Recruitment of the bown mussel Perna perna on to natural
substrata: a refutation of the primery/secondry settlement hypotheseis. Mar.ecol.Prog Ser., 120: 147-153.
[52] Lubet, P., 1976. Ecophysiologie de la reproduction chez les mollusques lamellibranches. Haliotis, 7: 49-55.
[53] Marobin, D., M. Prat and J.Y. Jouvenel, 2007. Étude globale sur la Telline Donax trunculus (Linnée, 1767)
en Camargue. Parc Naturel Régional de Camargue. Rapport Intermédiaire de la campagne d’automne.
Contrat BIOTOPE/P2A Développement, pp: 113 (+ annexes).
[54] Martoja, M. and R. Martoja, 1967. Initiation aux techniques de l’histologie animale. Masson & Cie (Eds)
pp: 347.
[55] Mazé, R.A. and A.J. Laborda, 1988. Aspectos de la dináimica de poblacion de Donax trunculus (Linnaeus,
1758) (Bivalvia: Donacidae) en la Ria del Barquero (Lugo, NO España). lnvestigaciones Pesqueras., 52:
299-312.
[56] McQuaid, C.D. and S.M. Lawrie, 2005. Supply-side ecology of the brown mussel Perna perna: an
investigation of spatial and temporal variation in, and coupling between, gamete release and larval supply.
Mar. Biol., 147(4): 936-955.
[57] Merzouki, M., N. Talib and J. Sif, 2009. Indice de condition et teneurs de quelques métaux (Cu, Cd, Zn et
Hg) dans les organes de la moule Mytilus galloprovincialis de la côte d’El Jadida (Maroc) en mai et juin
2004. Bulletin de l’Institut Scientifique, Rabat, section Sciences de la vie, 31(1): 21-26.
[58] Mouëza, M. and L. Frenkiel-Renault, 1973.Contribution á l'étude de la biologie de Donax trunculus L.
94
(Mollusques : Lamellibranches) dans l’Algérois : la reproduction. Cah. Biol. Mar., 14: 261-283.
[59] Mouëza, M. and D. Chessel, 1976. Contribution à l'étude de la biologie de Donax trunculus L. (Mollusque
Lamellibranche) dans l'Algérois : analyse statistique de la dispersion le long d'une plage en baie de Boulsmaïl. J. Exp. Mar. Biol. Ecol., 21: 211-221.
[60] Mouëza, M. and L. Renault, 1970. Contribution à l'étude de la sexualité chez Donax trunculus. Haliotis, 1:
21-22.
[61] Mouëza, M., 1971. Contribution à l’étude de l’écologie et de la biologie de Donax trunculus L. (Mollusque
Lamellibranche). Thèse Doct. Sciences Biologiques. Univ. Marseille, pp: 130.
[62] Mouneyrac, C., S. Linot, J.C. Amirad, C. Amirad-Triquet, I. Métais, C. Durou, C. Minier and J. Pellerin,
2008. Biological indices, energy reserves, steroid hormones and sexual maturity in the infaunal bivalve
Scrobiclario plana from three sites differing by their level of contamination. Gen. Comp. Endocrinol., 157:
133-141.
[63] Muranaka, M.S. and J.E. Lannan, 1984. Broodstock management of Crassostrea gigas: environmental
influences on broodstock conditioning. Aquaculture, 39: 217-228.
[64] Neuberger-Cywiak, L., Y. Achituv and L. Mizrahi, 1990. The ecology of Donax trunculus Linnaeus and
Donax semistriatus Poli from the Mediterranean coast of Israël. J. Exp. Mar. Biol. Ecol., 134: 203-220.
[65] Ounissi, M., H. Frehi and M. Khelifi-Touhami, 1998. Composition et abondance du zooplancton en
situation d’eutrophisation dans un secteur côtier du golfe d’Annaba (Algérie). Annal. Inst. Océano. Paris.,
73(1): 53-67.
[66] Pampanin, D.M., E. Volpato, I. Marangon and C. Nasci, 2005. Physiological measurements from native
and transplanted mussel (Mytilus galloprovincialis) in the canals of Venice. Survival in air and condition
index. Comp. Biochem. Physiol., 140: 41-52.
[67] Pearse, J.S., 1999. Seasonal reproduction, marine invertebrates. In: Encyclopedia of reproduction (Ed.
Knobil E., Neill J.D). Academic Press. San Diego, pp: 352-360.
[68] Pellerin-Massicotte, J.B., Y. Vincent and E. Pelletier, 1989. Evaluation de la qualité de l'estuaire du SaintLaurent. Etudes spatiales. Rapport remis au Ministère de l'Environnement du Québec, pp: 130.
[69] Ramón, M., P. Abelló and C.A. Richardson, 1995. Population structure and growth of Donax trunculus
(Bivalvia: Donacidae) in the western Mediterranean. Marine Biology, 121: 665-671.
[70] Roméo, M., N. Bennani, M. Gnassia-Barelli, M. Lafaurie and J.P. Girard, 2000. Cadmium and copper
display different responses towards oxidative stress in the kidney of the sea bass Dicentrarchus labrax.
Aquat Toxicol., 48: 185-194.
[71] Ruiz-Azcona, P., R. Rodriguez-Sierra and J.B. Martin, 1996. Culture of coquina clam, Donux trunculus,
larvae. Aquaculture, 139: 151-155.
[72] Samain M.S., G. Mathieu, L. Vanderschaeve, S. Wanke, C. Neinhuis and P. Goetghebeur, 2007.
Nomenclature and typification of subdivisional names in the genus Peperomia (Piperaceae) Taxon, 56:
229-236.
[73] Serdar, S., A. Lök, A. Kirtik, S. Acarli, A. Küçükdermenci, M. Güler and S. Yiğitkurt, 2010. Comparison
of gonadal development of crpet shell clam (Tapes decussatus Linnaeus, 1758) in inside and outside of
Çakalburnu Lagoon, Izmir Bay. Turkish Journal of Fisheries and Aquatic Sciences, 10: 395-401.
[74] Siah, A., G. Pellerin, J.C. Amiard, E. Pelletier and L. Viglino, 2003. Delayed gametogenesis and
progesterone levels in soft-shell clams (Maya arenaria) in relation to in situ contamination organotins and
heavy metals in the St. Lawrence River (Canada). Comparative Biochemistry and Physiology Part C:
Toxicology & Pharmacology, 135(2), 145-156.
[75] Sifi, K., 2009. Biosurveillance de la qualité des eaux du golfe d’Annaba : croissance, copmposition
biochimique et dosage des biomarqueurs du stress environnemental chez Donax trunculus (Mollusque,
Bivalve). Thèse de Doct. Biologie Animale, Univ. Annaba, pp: 229.
[76] Sifi, K., A. Amira and N. Soltani, 2013. Oxidative stress and biochemical composition in Donax trunculus
(Mollusca, Bivalvia) from the gulf of Annaba (Algeria). Advances in Environmental Biology, 7(4): 595604.
[77] Sifi, K., S. Chouahda and N. Soltani, 2007. Biosurveillance de l’environnement par la mesure de
biomarqueurs chez Donax trunculus dans le golfe d’Annaba (Algérie). Mésogée, 63: 11-18.
[78] Smolders, R., L. Bervoets, W.M. De Coen and R. Bulst, 2004. Cellular energy allocation in zebra mussels
exposed along a pollution gradient: linking cellular effects to higer levels of biological organization.
Environ. Pollut., 129: 99-112.
[79] Soltani, N., A. Amira, K. Sifi and H. Beldi, 2012. Environmental monitoring of the Annaba gulf (Algeria):
measurement of biomarkers in Donax trunculus and metallic pollution. Bull. Soc. zool. Fr., 137(1-4): 4756.
[80] Steele, S. and M.F. Mulcahy, 1999. Gametogenesis of the oyster Crassostrea gigas in southern Ireland. J.
Mar. Biol. Assoc. U.K., 70: 673-686.
[81] Tirado, C. and C. Salas, 1998. Reproduction and fecundity of Donax trunculus L., 1758 (Bivalvia:
95
Donacidae) in the littoral of Málaga (Southern Spain). Journal of Shellfish Research, 17(1): 169-176.
[82] Tlili, S., 2012. Approche multi-marqueurs pour l’évaluation de l’état de santé su golfe de Tunis : Etude des
réponses biochimique, physiologique, cytologique des mollusques bivalve exposé aux effluents des oueds.
Thèse Doct. Univ. Nantes/Univ. Monastir, pp: 249 (+ annexes).
[83] Tlili, S., I. Metais, N. Ayache, H. Boussetta and C. Mouneyrac, 2011. Is the reproduction of Donax
trunculus affected by their sites of origin contrasted by their level of contamination? Chemosphere., 10(84):
1362-1370.
[84] Wilson, J.G., 1999. Population dynamics and energy budget for a population of Donax variabilis (Say) on
an exposed South Carolina beach. J. Exp. Mar. Biol. Ecol., 239: 61-83.

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