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1.
Studies conducted on the distribution, fate and metabolism of DDT in a model ecosystem simulating a tropical marine environment of fish, Gobious nudiceps, Lethrinus harak, Gobious keinesis, Gobious nebulosis and white shrimp (Panaeus setiferus), show that DDT concentration in the water decreases rapidly within the first 24?h. Rapid accumulation of the pesticide in the biota also reaches a maximum level in 24?h before gradually declining. The bioaccumulation factors calculated for the fish species (G. keinesis) and white shrimp (P. Setiferus) were 270 and 351, respectively, after 24?h. There was a steady build up of DDT residues in the sediment during the first 24?h which continued to a maximum concentration of 6.66?ng/g in the seawater/fish/sediment ecosystem after 3 weeks and 5.27?ng/g in the seawater/shrimps/sediment ecosystem after 2.7 days. The depuration of the accumulated pesticide was slow with only 54% lost in G. nudiceps within 3 days of exposure in fresh sea water. By contrast, depuration was fast in the white shrimp, which lost 97% of the accumulated pesticide under the same conditions. DDT was found to be toxic to two of the fish species (G. nebulosis and L. harak) and to white shrimp, and the degree of toxicity was dependent on the particular species. The 24?h LC50 at room temperature for the fish species G. nebulosis and white shrimp was found to be 0.011 and 0.116?mg/kg, respectively. These levels are comparable to the ones recorded for the temperate organisms. Degradation of DDT to its primary metabolites, DDE and DDD, was found in all the compartments of the ecosystem with DDE being the major metabolite in the fish, shrimps and sediment, while in seawater, DDD dominated as the major metabolite.  相似文献   

2.
The sperm of the brown shrimp Penaeus aztecus are nonflagellated and non-motile. In vitro fertilization can be obtained when mature eggs removed from female shrimp are added to dense sperm suspensions in seawater and agitated in a manner simulating natural spawning.  相似文献   

3.
The results of toxicity experiments have shown that a level of 17 ppb ionic mercury is acutely toxic (96-h LC50) to postlarval white shrimp (Penaeus setiferus). In addition, the size of the shrimp, within the range tested (7 to 35 mm) did not significantly alter the toxicity values of mercury to this species. Fifty-seven days pre-exposure to low levels of mercury (0.5 and 1.0 ppb) had no effect on the acute toxicity (96-h LC50) of mercury to the shrimp. The effect of chronic exposure to low levels of mercury on the respiratory rate, growth, and molting rate of postlarval P. setiferus was studied, and the results showed that 60 days exposure to 0.5 and 1.0 ppb mercury did not significantly affect any of these parameters for the postlarvae.  相似文献   

4.
Field and laboratory experiments were conducted to identify the sources of food in the natural diet of postlarval brown shrimp (Penaeus aztecus Ives). A series of enclosures placed in East Lagoon (29°20N; 94°45W) on Galveston Island, Texas, USA, in May 1985, were used to evaluate the individual and combined contribution of Spartina alterniflora detritus, epiphytes of S. alterniflora, plankton, and demersal fauna in terms of differences in shrimp growth and carbon assimilation (stable carbon-isotope analysis). Demersal fauna (harpacticoid copepods, amphipods, tanaids and polychaete annelids), and plankton (>0.095 mm) accounted for approximately 53 and 47% of the growth of the postlarvae (11 to 22 mm rostrum-telson length), respectively, while the autochthonous plant substrates, S. alterniflora detritus and epiphytes, contributed little. Laboratory experiments confirm that a mixed diet consisting of both animal protein and phytoplankton promotes maximum growth. Our results indicate that plankton may be an important allochthonous source of carbon contributing to the growth and development of shrimp in the salt marsh.  相似文献   

5.
Enmin Zou  Ben Stueben 《Marine Biology》2006,149(6):1411-1415
The brown shrimp, Penaeus aztecus, in the northern Gulf of Mexico is faced with dual stresses of environmental hypoxia, which occurs as a result of oxygen depletion from microbial decomposition of organic materials from algal blooms, and pollution from polycyclic aromatic hydrocarbons (PAHs) from petroleum and gas production on the continental shelf of the northern Gulf of Mexico. This study investigated the effect of naphthalene, a PAH, on oxyregulating capacity of P. aztecus, when shrimps were subjected to progressive hypoxia. It was found that P. aztecus is an oxyregulator with a critical oxygen concentration of 2.53 mg/l at 19–21°C, below which the animal becomes an oxyconformer. Acute exposure to naphthalene at 2.0 mg/l significantly reduced the oxyregulating capacity by 112%. This is the first report on the alteration of an aquatic animal’s oxyregulating capacity by a PAH. Possible mechanism for the impaired oxyregulation in the presence of naphthalene was discussed.  相似文献   

6.
Symbiotic dinoflagellates, Symbiodinium microadriaticum (=zooxanthellae), may gain access to aposymbiotic hosts (i.e., those lacking zooxanthellae) by chemosensory attraction of the motile algae by the potential host or via an intermediate host. Laboratory experiments showed that motile zooxanthellae were attracted to intact aposymbiotic host animals, but not to starved symbiotic hosts. Fed symbiotic hosts and brine shrimp (Artemia sp.) nauplii also attracted motile zooxanthellae. The attraction of these zooxanthellae was directly correlated with nitrogen levels in the seawater surrounding the hosts; thus ammonia and possibly nitrate could be atractants. Brine shrimp nauplii, acting as intermediate hosts actively ingested both motile and non-motile zooxanthellae. the ingested zooxanthellae tended to remain morphologically unaltered during and after passage through the gut of the brine shrimp. Capture and ingestion of brine shrimp containing zooxanthellae by aposymbiotic scyphistomae of the jellyfish Cassiopeia xamachana led to infection of the scyphistomae with zooxanthellae. Zooxanthellae isolated from 17 different species of coelenterates and molluscs could be transferred via brine shrimp to the endodermal cells of the scyphistomae. However only 10 of these isolates persisted to establish a permanent association with C. xamachana. Scyphistomae in suspensions of motile zooxanthellae responded by a classical coelenterate feeding response, which may facilitate ingestion of the potential symbionts and establishment of a symbiosis.  相似文献   

7.
Summary Peromyscus melanotis is the only one of three mouse species that eats monarch butterflies at their overwintering sites in Mexico. I tested two hypotheses: 1)P. aztecus avoids monarchs because of a bitter taste aversion to cardiac glycosides (CGs) and an inability to reject CG-rich body parts; 2)Reithrodontomys sumichrasti avoids monarchs principally because of a bitter taste aversion to the CGs. None of the species are sensitive to the toxic effects of ingested CGs. Feeding responses of laboratory-reared mice of each species to monarchs with low, medium and high CG concentrations were compared. BothP. aztecus andR. sumichrasti ate significantly fewer of all three types of monarchs thanP. melanotis. ForP. aztecus andR. sumichrasti, the number of monarchs eaten decreased with increasing CG concentration, whereas forP. melanotis, the number remained constant.Peromyscus melanotis andR. sumichrasti developed a feeding technique for rejecting the CG-laden cuticular material, which reduced the bitterness of ingested monarch material. However,R. sumichrasti displayed the technique significantly less often thanP. melanotis; andP. aztecus never developed it. I conclude that high taste sensitivity to CGs and less versatile food handling preventP. aztecus andR. sumichrasti from overcoming the monarch's chemical defenses.  相似文献   

8.
Summary Of three common mouse species at the Mexican overwintering sites of the monarch butterfly, onlyPeromyscus melanotis eats monarchs. We hypothesized thatP. aztecus andReithrodontomys sumichrasti reject monarchs because they are more sensitive to the bitter taste and/or toxic effects of the cardiac glycosides (CGs) and pyrrolizidine alkaloids (PAs) in the butterflies. Two-choice preference tests revealed no difference in taste avoidance thresholds to free base and N-oxide forms of the PA, monocrotaline, but very different avoidance thresholds to the CG, digitoxin. Avoidance thresholds forR. sumichrasti andP. aztecus were, in respective order, 1020 and 34 times less than that forP. melanotis. We also tested the toxic sensitivity of juvenile mice by chronically feeding diets containing digitoxin or monocrotaline at concentrations similar to those used in the preference tests. No species developed CG toxicity, but bothP. melanotis andP. aztecus developed moderate PA toxicity (R. sumichrasti was not tested for PA toxicity).P. aztecus grew more slowly and manyP. melanotis had hepatic metabolic lesions. Thus, the three mouse species responded very differently to the taste and toxic properties of CGs and PAs at ecologically relevant concentrations: 1) CGs were taste rejected by all species exceptP. melanotis, while PAs were not; and 2) PAs were toxic, while CGs were not.  相似文献   

9.
The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.  相似文献   

10.
Sand shrimp, Crangon septemspinosa Say, are important to the trophic dynamics of coastal systems in the northwestern Atlantic. To evaluate predatory impacts of sand shrimp, daily energy requirements (J ind.–1 day–1) were calculated for this species from laboratory estimates of energy losses due to routine (RR), active (RA), and feeding (RSDA) oxygen consumption rates (J ind.–1 h–1), coupled with measurements of diel motile activity. Shrimp used in this study were collected biweekly from the Niantic River, Connecticut (41°33N; 72°19W) during late spring and summer of 2000 and 2001. The rates of shrimp energy loss due to RR and RA increased exponentially with increasing temperature, with the magnitude of increase greater between 6°C and 10°C (Q10=3.01) than between 10°C and 14°C (Q10=2.85). Rates of RR doubled with a twofold increase in shrimp mass, and RSDA was 0.130 J h–1+RR, irrespective of shrimp body size. Shrimp motile activity was significantly greater during dark periods relative to light periods, indicating nocturnal behavior. Nocturnal activity also increased significantly at higher temperatures, and at 20°C shifted from a unimodal to a bimodal pattern. Laboratory estimates of daily metabolic expenditures (1.7–307.4 J ind.–1 day–1 for 0.05 and 1.5 g wet weight shrimp, respectively, between 0°C and 20°C) were combined with results from previous investigations to construct a bioenergetic model and make inferences regarding the trophic positioning of C. septemspinosa. Bioenergetic model estimates indicated that juvenile and adult shrimp could meet daily energy demands via opportunistic omnivory, selectively preying upon items of high energy content (e.g. invertebrate and fish tissue) and compensating for limited prey availability by ingesting readily accessible lower energy food (e.g. detritus and plant material).Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by J.P. Grassle, New Brunswick  相似文献   

11.
Mixed zooplankton were collected in June and July of 1985 and 1986 from La Jolla Bay, California, USA, and experiments were conducted to determine how selected dinoflagellates affect development and survival of nauplius larvae of Calanus pacificus. We raised nauplii from eggs on nine species of dinoflagellates at concentrations generally >300 g C l-1, and compared their development and survival to controls reared using the diatom Thalassiosira weissflogii or filtered seawater. Experiments were conducted for 6 d at 17°C. Development and survival rates of the nauplii fell clearly into one of two groups, depending upon the phytoplankton used as food. The first group was characterized by high development rate (0.46 to 0.84 stage d-1), and by >27% of the original cohort surviving to at least Nauplius IV or V. The five species producing this result were Gymnodinium simplex, G. splendens, Exuviaella marie-lebourae, Gyrodinium dorsum, and T. weissflogii. The second group was characterized by a development rate similar to that in filtered seawater (0.21 to 0.34 stage d-1), and by nauplii generally failing to molt past the first feeding stage (Nauplius III), often accompanied by high mortality. The five species producing this result were Gyrodinium resplendens, Ptychodiscus brevis, Glenodinium sp., Amphidinium carterae, and Gonyaulax grindleyi. Development rate and survival were not related to cell size or cell carbon, nor to shape or texture (thecate vs athecate dinoflagellates). Poor growth could be related to the absence of some important, but unidentified, nutritional factors. Alternatively, it could be caused by the presence of plant secondary metabolites which are deleterious to growth, a factor we suspect in P. brevis in particular. Prefeeding nauplii exposed to P. brevis lost neuromuscular control prior to becoming lethargic and dying; nutritional deficiencies may not explain these effects. Methods employed in this study provide useful bioassays for detecting chemical interactions between marine plants and animals. Lethal or sublethal effects of dinoflagellates on their most likely potential predators — copepods — may partially explain why they form significant blooms.  相似文献   

12.
Some of the effets of dieldrin on the development of two species of crabs, Leptodius floridanus (Rathbun) and Panopeus herbstii (Milne-Edwards), were studied. It was found that the larvae of neither species were able to complete their development at 10 ppb dieldrin or higher in seawater. Groups of L. floridanus larvae reared in 1 ppb dieldrin in seawater had a 15 to 27% higher mortality during development to the postlarval stage than controls. The highest mortality occurred during the first zoeal stage. The time of development to the megalopal stage was as much as 11.4% longer among larvae reared in 1 ppb than among controls. The survival of L. floridanus larvae was not affected by 0.5 ppb dieldrin in seawater. The survival of P. herbstii larvae to the first crab stage was not affected by 1 ppb dieldrin in seawater at 28.5 °C, 30%.S. It was concluded that a comprehensive study of the toxicity of a given pesticide to all stages in the life cycle of a species is necessary to give even an incomplete idea of how the pesticide might affect the animal in its natural environment.  相似文献   

13.
Zostera marina L. was studied at the Izembek Lagoon, Alaska Peninsula. Two morphologically different forms, tidepool and subtidal, can be distinguished. Both show a high tolerance to different salinities and temperatures. The plasmatic resistance was found in a range of distilled H2O up to 3.0 seawater (24 h) and between-6° and 34°C (12 h). Within these resistance limits, the photosynthesis, which has its maximum in normal (1.0) seawater, decreases nearly to zero not only in distilled H2O but even in 2.0 seawater, and increases with the temperature in the tidepool form up to 35°C, but in the subtidal form up to 30°C only. At higher temperatures photosynthesis declines sharply in both forms. Respiration has its minimum in distilled H2O and at 0°C and increases with increasing salinity and temperature.Dedicated to Prof. Dr. Dr. h. c. mult. K. Mothes, Halle, Germany (DDR) on the occasion of his 70th birthday.  相似文献   

14.
G. Purschke 《Marine Biology》1981,63(3):257-267
Tolerance experiments on freezing and supercooling (without ice formation) were designed to determine correspondence between tolerance to low temperatures and spatio-temporal distribution of one turbellarian and five polychaete species from sandy beaches of the North Sea island of Sylt. Freezing tolerances are always less than supercooling tolerances. Dinophilus gyrociliatus is significantly more sensitive to freezing (LD50 value after 30 min of freezing:-3°C) than the others, whereas Stygocapitella subterranea is significantly less sensitive (50% mortality at-15.7°C after 30 min). The supercooling tolerances differ considerably among the species. The sequence of tolerances (LD50 values) is as follows: Microphthalmus sczelkowii (-2.9°C after 4 h); D. gyrociliatus (<1 h at-8°C); M. listensis (5.6 h at-8°C); Protodriloides symbioticus (8.2 h at-8°C); Notocaryoplanella glandulosa (66 h at-8°C); S. subterranea (72 h at-8°C). Species of sand flats (d. gyrociliatus, M. listensis, P. symbioticus) have lower tolerances than those of the beach slope (N. glandulosa, S. subterranea). Among the latter, tolerances increase with distance of the distributional area from low tide level. S. subterranea, a species occurring at the uppermost position in the intertidal, proves to be best adapted to both freezing and supercooling. Species preferring deeper regions of the beach (M. sczelkowii) show lower supercooling tolerances than surface dwelling forms. Northern species usually have higher tolerances to cold than southern ones, reaching their distribution limits near the island of Sylt.  相似文献   

15.
Acclimation of reef corals to environmental conditions has been related to metabolic response at large geographic scales, but regional relationships have rarely been described. Physiological responses to temperature increases of Montastraea annularis (Ellis and Solander 1786) from an inner lagoon and an outer barrier reef in the Gulf of Honduras, southern Belize, were compared in May 2003. The hypothesis that inferred differences in thermal history would result in contrasting responses to elevated temperature was tested. Ambient seawater temperatures adjacent to corals at 4–5 m depth were measured every 15 min at inner lagoon and outer barrier reef collection sites for 1 year (June 2002–May 2003). Monthly averages and 3-day running averages (warmest period, July–October 2002) of daily maximum seawater temperatures were significantly higher (by ∼0.5°C) at inner lagoon reef compared to outer barrier reef sites. M. annularis photosynthesis (P) and respiration (R) rates were measured in respirometers at six temperatures between 29°C and 35°C approximately every hour, with repeated measurements over 3 h. P and R were significantly lower across most temperature treatments for samples collected from the inner lagoon compared to outer barrier reef. Both inner and outer reef M. annularis displayed an increase in P and R with increasing temperature between 29°C and 32°C, but above 32°C P and R sharply declined. P/R ratio versus temperature showed a significant difference between the elevations of the regression lines suggesting that M. annularis from the outer barrier reefs may have been more physiologically stressed than those from the inner lagoon reefs when exposed to acute temperature changes. These results emphasize that thermal stress must be considered within the context of acclimation temperature, and that short-term exposures may have physiologically important effects on this species.  相似文献   

16.
The nutritional value of Artemia sp. as food for marine fish and crustacean larvae has been linked to the level of its polyunsaturated fatty acid (PUFA) content. Experiments in August 1984 were conducted to determine the effects of various artificial diets and algae on fatty acid composition of PUFA-deficient Artemia sp. (Utah GSL strain) and their resulting value as food for postlarvae of the prawn Penaeus monodon (Fabricius). Nauplii of the brine shrimp were grown on extracts of corn, copra, soybean and rice bran containing precursors (C18) to long-chain PUFA and also on algal species containing different levels of long-chain PUFA (C20). The nauplii were then used as food for P. monodon postlarvae. The results revealed that absence of C20 polyunsaturates from the feeds and their presence in the algae were reflected in the polyunsaturated fatty acid content of the tissues of Artemia sp. When fed with brine shrimp fed on algae, P. monodon displayed better postlarval survival and significantly higher growth; related to the content of polyunsaturated fatty acids in Artemia sp. A practical feeding approach in prawn hatcheries would be to grow Artemia sp. on a cheap diet such as rice bran, and then to enhance its nutritional value with a diet high in PUFA prior to harvesting, in order to improve hatchery production.  相似文献   

17.
The effects of body-size and temperature on moulting frequency of the shrimp Lysmata seticaudata Risso were examined under controlled laboratory conditions. Shrimp ranging in weight from approximately 0.4 to 1.14 g and maintained at 17°C exhibited intermoult periods of similar duration (20 d). On the other hand, small individuals (0.23 g) held at the same temperature moulted on the average every 14.5 d. Over a temperature range from 8° to 20°C mean intermoult periods were inversely and linearly related to temperature; an increase of 1 C0 resulted in a decrease in the intermoult period of approximately 1.5 d. Although temperature acted to regulate the intermoult period of each shrimp in increments of whole days, it had little effect on the time of moulting within any 24 h period, since moulting occurred more than 93% of the time between 18.00 and 08.00 hrs. The night-time moulting pattern also tended to be most pronounced at lower temperatures (8° and 13°C) and in larger individuals. Possible factors controlling this phenomenon and its occurrence in other crustacean species are discussed.  相似文献   

18.
Chondrus crispus (Stackhouse) is a perennial red seaweed, common in intertidal and shallow sublittoral communities throughout the North Atlantic Ocean. In the intertidal zone, C. crispus may experience rapid temperature changes of 10 to 20C° during a single immerison-emerision cycle, and may be exposed to temperatures that exceed the thermal limits for long-term survival. C. crispus collected year-round at Long Cove Point, Chamberlain, Maine, USA, during 1989 and 1990, underwent phenotypic acclimation to growth temperature in the laboratory. This phenotypic acclimation enhanced its ability to withstand brief exposure to extreme temperature. Plants grown at summer seawater temperature (20°C) were able to maintain constant rates of lightsaturated photosynthesis at 30°C for 9 h. In contrast, light-saturated photosynthetic rates of plants grown at winter seawater temperature (5°C) declined rapidly following exposure to 30°C, reached 20 to 25% of initial values within 10 min, and then remained constant at this level for 9 h. The degree of inhibition of photosynthesis at 30°C was also dependent upon light intensity. Inhibition was greatest in plants exposed to 30°C in darkness or high light (600 mol photons m-2s-1) than in plants maintained under moderate light levels (70 to 100 mol photons m-2s-1). Photosynthesis of 20°C-acclimated plants was inhibited by exposure to 30°C in darkness or high light, but the degree of inhibition was less than that exhibited by 5°C-grown plants. Not only was light-saturated photosynthesis of 20°C plants less severely inhibited by exposure to 30°C than that of 5°C plants, but the former also recovered faster when they were returned to growth conditions. The mechanistic basis of this acclimation to growth temperature is not clear. Our results indicate that there were no differences between 5 and 20°C-grown plants in the thermal stability of respiration, electron transport associated with Photosystems I or II, Rubisco or energy transfer between the phycobilisomes and Photosystem II. Overall, our results suggest that phenotypic acclimation to seawater temperature allows plants to tolerate higher temperatures, and may play an important role in the success of C. crispus in the intertidal environment.  相似文献   

19.
Intertidal organisms exposed to thermal stress normally experience other stresses simultaneously, but how these combined stresses modify tolerance to heat, especially for embryos, is poorly understood. Tolerance of fucoid algal embryos to heat, with and without acclimation to a sublethal temperature and with simultaneous exposure to hypersaline media, was examined. Embryos of Fucus vesiculosus L. (mid-intertidal zone) were less tolerant than embryos of Fucus spiralis L. (upper intertidal zone); without acclimation and with a growth temperature of 14°C, about half of the embryos survived 3 h exposure to 33°C in F. vesiculosus and of 35°C in F. spiralis. Conditions experienced by parental thalli (4°C versus 14°C storage) significantly affected the heat tolerance of embryos grown for 24 h post-fertilization at 14°C in F. vesiculosus, a result that is important for biologists using fucoid algae as model systems. Acclimation to a sublethal temperature (29°C) or exposure to the LT50 (33°C, F. vesiculosus; 35°C, F. spiralis) in 100 psu seawater (2850 mmol kg–1 osmolality) resulted in 30–50% higher levels of embryonic survival. Higher levels of HSP60s were found in embryos exposed to 29–33°C than to 14°C; lower levels of HSP60s were present in embryos exposed to the LT50 under hypersaline conditions than in normal seawater. Contemporaneous studies in 1995–1996 of substratum temperature and desiccation levels were made at Schoodic Point, Maine (USA) underneath F. spiralis and F. vesiculosus canopies and in Semibalanus balanoides patches. This study extends the bioindicator utility of heat-shock proteins in studies of intertidal organisms and demonstrates the importance of integrated stress responses in survival of a single stress factor (e.g. temperature).Electronic Supplementary Material Supplementary material is available in the online version of this article at  相似文献   

20.
S. Choe 《Marine Biology》1971,9(1):31-37
Individuals of the oriental brown shrimp Penaeus japonicus Bate, were raised separately (1 shrimp per rearing cage) with surplus food, in almost dark, non-sediment conditions. Results were obtained regarding growth of each part of the body in conjunction with exuviation and molting cycle. Increases in carapace, body length and body weight conformed to the general pattern discribed by Hiatt (1948), with no difference in growth resulting from sex, or inflexion point in juvenile stage. The molting cycle of shrimp weighing 1.5 to 15 g was 6 to 17 days at a water temperature of 20° to 28°C. The molting cycle was prolonged in proportion to the size of the shrimp; shortened as the water temperature increased.  相似文献   

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