Osmotic and chloride regulation in the hemolymph of the tiger prawn Penaeus monodon during molting in various salinities

The effect of molting on osmotic and chloride concentrations in the blood of the prawn Penaeus monodon Fabricius (20±3 g) at various salinities was investigated. Prawns were obtained from ponds in Iloilo, Philippines, in 1984. They were stocked in salinities of 8, 20, 32 and 44, and their hemolymph was sampled during molt (Time 0), and then 0.125, 0.25, 0.5, 1, 2, 4, 6, 10 and 14 d after molting. Prawns during and immediately after molt tended to conform to the environmental osmolality. Subsequent postmolt (0.5 d) stages displayed more divergence from external salinity. The isosmotic point was higher (940±30 mOsm kg^-1) during molt than during intermolt (663±8 mOsm/kg^-1), suggesting different osmotic requirements in early molt. Hyperregulation of hemolymph chloride below 20 S, as well as isoionic point (301±6 mM), were independent of molting stage. At 20 S and above, newly molted (0 to 0.25 d post-molt) individuals tended to conform to the external chloride concentration while intermolt (0.5 d) post-molt individuals did not. Contribution of hemolymph chloride to hemolymph osmolality was greater during intermolt than during ecdysis, suggesting an important role for other negatively charged ions during molt. When molt occurred in 20 S (the test salinity most similar to the isoionic salinity), there was little or no change in hemolymph osmolality or chloride concentration from 0 to 14 d postmolt. At 8, 32 and 44 S, the change from molt to intermolt values in hemolymph osmotic and chloride concentrations was hyperbolic. Non-linear least-squares regression showed that prawns generally achieved intermolt values within 1 d after molting. Prawns at intermolt regulated hemolymph osmolality (620 to 820 mOsm kg^-1) and chloride concentration (300 to 450 mM) at a much narrower range than during molt (520 to 1 170 mOsm kg^-1 and 250 to 520 mM, respectively). Hemolymph osmolality was a more sensitive indicator of physiological response than hemolymph chloride concentration. Distribution and culture of P. monodon might be limited in low salinities by its ability to maintain a hemolymph osmolality 500 mOsm kg^-1 during molt and 600 mOsm kg^-1 in intermolt, and in high salinities by its capacity to reduce the hemolymph osmolality from values at molt to those in intermolt. Osmotic and chloride concentrations in the blood of P. monodon clearly varied with both molt stage and salinity of the medium. Dependence on external factors, however, gradually declined in older molt stages, suggesting a reduction in integument permeability and greater development of ion absorption/secretion mechanisms as the exoskeleton hardened.SEAFDEC Contribution No. 197     

Effect of salinity on hemolymph calcium concentration during the molt cycle of the prawn Penaeus monodon

Prawns (Penaeus monodon) were obtained from ponds in Iloilo, Philippines, in 1984 and 1985 and maintained in salinities from 8 to 44. Total hemolymph calcium was largely affected by molt stage and less so by salinity. A sharp, transient increase in hemolymph calcium occurred 3 to 6 h postmolt, followed by an equally rapid decrease from 6 h postmolt to intermolt. This biphasis response was limited to prawns in 8, 20 and 32S; in 44S, hemolymph calcium remained the same throughout the sampling period. Peak concentrations of total calcium were greater in low (8 and 20S) than in high salinities. Salinity had no effect on the duration of molt cycle nor on time of occurrence of molt. Almost half of molting incidents occurred between 18.01 and 0.00 hrs, and one-third between 0.01 and 06.00 hrs.     

Use of nucleic acids as indicators of growth in juvenile white shrimp,Penaeus vannamei

The use of nucleic acids to estimate crustacean growth is not well documented, and may be complicated by biochemical changes associated with their molt cycle. The objectives of this study were to assess the effects of molt stage on nucleic acid concentrations in abdominal muscle tissue of juvenile white shrimp,Penaeus vannamei, and to examine the relationship between nucleic acid concentrations and growth rates of shrimp exposed to different feeding regimes throughout a 12 d feeding experiment. RNA and DNA concentrations and RNA:DNA ratios were not significantly different among the five major molt stages early postmolt, late postmolt, intermolt, early premolt, and late premolt. In the feeding experiment, RNA concentrations and RNA:DNA ratios accounted for >70% of the variation in shrimp growth on three different sampling days. In addition, RNA concentrations and RNA:DNA ratios were used successfully to discriminate between unfed, moderately-fed, and well-fed shrimp. These variables exhibited significant treatment differences in <24 h after the initiation of the different feeding regimes, whereas significant changes in whole-body weight took longer to detect. Rapid detection of significant treatment effects can be useful in ecological studes, especially those concerned with food-web interactions.     

Respiratory and behavioral responses of the grass shrimp Palaemonetes pugio to cadmium and reduced dissolved oxygen

Adult grass shrimp (Palaemonetes pugio) were collected during July and August, 1973, and exposed for 48-h periods to a series of sublethal cadmium (Cd) concentrations: 0.1 and 0.56 mg l^-1 for metabolic studies; and 0.01, 0.05, 0.1, 0.3, 0.56 mg l^-1 for locomotor studies. Treatment effects were tested singly and in combination at selected dissolved oxygen (DO) concentrations ranging from near saturation to 0.8 mgl^-1. Shrimp not exposed to Cd exhibited metabolic regulation between 6.5 and, 4.6 mg DOl^-1, with progressively lower O₂ consumption rates at lower DO concentrations. Oxygen consumption of Cd-exposed shrimp was suppressed overall relative to the non-Cd treated shrimp, and with one exception, the magnitude of metabolic suppression directly corresponded to Cd concentration. Locomotor activity of non-Cd exposed shrimp was relatively independent of DO concentration down to 2.8 mg DO l^-1 and became significantly suppressed at 1.8 mg DO l^-1 and below. Significant suppression of locomotion was evident in shrimp exposed to 0.56 mg Cd l^-1, but was not seen in treatment groups of 0.3 mg Cd l^-1 or lower. No interaction on locomotor activity of shrimp was detected between Cd exposure and oxygen concentration. The reduction of locomotor activity and metabolism at 0.56 mg Cd l^-1 exposure is similar to that occurring under reduced oxygen condition. It is hypothesized that Cd acts in some manner to suppress O₂ availability to P. pugio, a suggestion consistent with literature reports on adverse effects of Cd on fish respiratory enzymes and gill tissue.Contribution No. 351 of the U.S. Environmental Protection Agency, Environmental Research Laboratory, Narragansett, Rhode Island. Although the research described in this article was funded by the U.S. Environmental Protection Agency, it has not been subjected to Agency review and therefore does not necessarily reflect the views of the Agency and no official endorsement should be inferred     

Mineral concentration in tissues during ovarian development of the white shrimp Penaeus vannamei (Decapoda: Penaeidae)

Levels of calcium, magnesium, zinc, copper, iron, and manganese were examined in the muscle, hepatopancreas, and gonad of white shrimp, Penaeus vannamei, females at different stages of ovary maturation. Eyestalk-ablated shrimp were sampled during the intermolt stage and classified using the following maturation stages evaluated histologically: cortical stage, vitellogenesis, previtellogenesis, and without gonadic development. In muscle, the copper concentration was significantly lower in the vitellogenic stage than in other stages of ovary development. In the hepatopancreas, manganese levels were higher in the cortical and vitellogenic stages than in animals in previtellogenesis and without gonadic development. No other influence of ovary maturation on muscle and hepatopancreas mineral content was found. In contrast, in the ovary, significant differences of some mineral concentrations were detected. Zinc, iron, and magnesium concentrations were significantly increased in the cortical stage compared to the vitellogenic and previtellogenic stages. Our results indicate that these elements are selectively accumulated during ovary development, especially when the ovaries reach the cortical stage. Received: 14 March 2000 / Accepted: 30 November 2000     

Kinetics of osmoregulation in the crab Carcinus maenas

When shore crabs Carcinus maenas are transferred from 11 to 38 S at 11°C, new constant levels of hemolymph freezing points and of concentrations of Na, K, Ca, and Mg in the hemolymph are accomplished within 24h. From a decrease in serum protein and in serum free amino acids and an increase in the relative amounts of individual essential free amino acids in the body fluids of whole crabs, a participation of proteolytic activities and a transport of amino acids from hemolymph to cell is deduced; the stationary concentration of total intracellular free amino acids increases up to a nearly two-fold value, compared with the concentration in crabs remaining in diluted sea water. Also, the low molecular neutral sugars increase in whole crabs after high salinity stress, reaching values of more than two-fold initial concentrations within a period of 10 days. This increase is fully accounted for by a 6.7-fold increase in trehalose levels. The processes of increasing the concentrations of low molecular organic material seem to be slower than those of establishing new osmotic pressures in the hemolymph. The oxygen consumption decreases by 30 to 45% to new constant values within 8 to 12 h after the salinity change from 11 to 38, reflecting similar kinetics as the establishment of new osmotic pressures in the hemolymph.     

Respiration,pumping activity and heart rate in Ciona intestinalis exposed to fluctuating salinities

Specimens of Ciona intestinalis L. were exposed to both gradual (sinusoidal) and abrupt (square-wave) salinity fluctuations and the changes in pumping activity and oxygen sonsumption monitored. Heart rate was monitored under steady state conditions using a new in vivo method, and pumping activity was monitored as spontaneous squirting by use of a pressure transducer. Oxygen consumption was also monitored under steady state conditions and under conditions of declining oxygen tension. It was found that during periods of regular beating the mean heart rate for 5 ascidians was 13.2 beats min^-1 for the branchial pacemaker and 32.6 beats min^-1 for the visceral pacemaker. Regular periods of beating lasted for approximately 1 min, followed by periods of irregular activity prior to heart beat reversal. The heart beat became irregular and reversal occurred sporadically when the ascidians were exposed to dilute seawater. The ascidians showed a pumping rate of 14 squirts per hour in full-strength seawater. This rate declined during decreasing salinity and ceased entirely when the external seawater concentration reached approximately 60% seawater (100%=32% S). During periods of decreased salinity, the siphons were tightly closed and oxygen consumption was zero. The rate of oxygen consumption by C. intestinalis decreased during decreasing salinities and there was no evidence of an oxygen debt. Oxygen consumption under steady state conditions varied with body weight according to the following: oxygen consuption =0.515 W ^0.831 (where W is body weight). The rate of oxygen consumption was found to be dependent on the external oxygen tension.     

Effect of dissolved oxygen level on respiratory metabolism, nutritional physiology, and immune condition of southern king crab Lithodes santolla (Molina, 1782) (Decapoda, Lithodidae)

Episodes of hypoxia are common in the marine environment, and their ecological effects depend, in part, on their severity and duration. Many species of decapod crustaceans reside in areas with fluctuating oxygen regimens. Physiological mechanisms enhance the ability of these crustaceans to cope with acute episodes of hypoxia. Southern king crab, Lithodes santolla, fishery is important in the south of South America, and some data describe fishing zones with low dissolved oxygen (DO) levels (3.5 mgO₂ l⁻¹, i.e., 8.3 kPa). Our main objective was to evaluate the effect of dissolved oxygen level on respiratory metabolism, nutritional physiology, and immunological condition of L. santolla juveniles. Individual animals were exposed for 10 days to different oxygen tensions (2.1, 4.2, 8.5, 12.7, and 21.1 kPa) to quantify the oxygen consumption rate; thereafter, blood oxyhemocyanin (Hc), protein concentration, as well as hemocytes, were sampled. Freeze-dried animals were dissected, and digestive gland metabolites (glycogen, protein, glucose, cholesterol, acylglycerol, and lactate) and digestive enzyme activity (general protease, trypsin, and chymotrypsin), as well as gill lactate dehydrogenase (LDH) activity, were quantified. In the present study, Lithodes santolla showed a critical oxygen tension between 4 and 9 kPa, indicating that this crab species is more sensitive to DO than other crustacean species. Protein and Hc concentrations followed a similar pattern to that of oxygen consumption. Digestive gland glycogen and protein concentration did not change after 10 days at different oxygen exposures, but glucose, cholesterol, and acylglycerol concentrations decreased linearly and proportionally to the available oxygen in the water. As in other decapods, chymotrypsin showed over 90% of the total quantified proteases activity. Chymotrypsin activity together with total proteases and trypsin was not affected by the environmental oxygen tension. Gill LDH and digestive gland lactate followed a similar increase at lower environmental oxygen tension but dropped sharply at the lowest tension (2.1 kPa). Dissolved oxygen affected also the immune system through reduction of hemocytes. This could provide a critical window for opportunistic pathogens to become established when crabs are exposed to hypoxic conditions. L. santolla juveniles show a moderate tolerance to low oxygen availability by modifying the concentration of hemolymph proteins, mainly OxyHc, some digestive gland metabolites, and by activating the anaerobic metabolism. This allows L. santolla juveniles to inhabit temporarily low oxygen zones in the deep ocean and suggests an advantage for culture conditions.     

Osmoregulation of the grass shrimp Palaemonetes pugio exposed to polychlorinated biphenyls (PCBs). I. Effecton chloride and osmotic concentrations and chloride-and water-exchange kinetics

Grass shrimp, Palaemonetes pugio, were capable of hypo- and hyper-osmotic regulation of body fluids. Hemolymph chloride and osmotic concentrations were maintained at relatively stable levels over a wide salinity range. Following an abrupt transfer from intermediate (14 and 17) to high (31 and 35) or low (1 and 2) salinities, hemolymph chloride levels exhibited initial overshoot and undershoot, respectively, of new steady-state levels. Osmotic concentrations exhibited an initial undershoot at low, but not overshoot at high salinity. Chloride space in salinity-acclimated shrimp was relatively stable at salinities from 1 to 35. Changes in chloride space following salinity transfer paralleled those of hemolymph chloride levels, and are discussed in the light of alterations in intracellular sodium concentrations reported earlier. Rate constants for chloride turnover indicated independent exchanges of sodium and chloride ions. Water-turnover measurements showed that permeability of P. pugio was greatest at the isosmotic salinity (17) and reduced at salinities which were associated with active osmoregulation. Exposure to sublethal and 96-h LC₅₀ levels of Aroclor^® 1254 did not seriously alter hemolymph chloride and osmotic concentrations, chloride space or chloride-exchange kinetics in adult shrimp. Disruption of hemolymph chloride regulation in juvenile shrimp was associated with large mortalities not observed in adults. Shrimp exposed to Aroclor 1254 at 17 S exhibited reduced water permeability similar to levels previously observed in controls at high and low salinities in response to osmotic or ionic gradients. Exposure to PCBs did not result in further reduction in permeability at the latter salinities.     

Comparative study of the effects of temperature,salinity and oxygen tension on the rates of oxygen consumption of nauplii of different strains of Artemia

The effects of temperature, salinity and oxygen tension on the rates of oxygen consumption of three different strains of Artemia nauplii have been studied. When acclimated to a salinity of 30, nauplii from each of the three strains were able to maintain approximately constant rates of oxygen consumption over a wide range of oxygen tension. The ability to maintain respiratory independence during hypoxia was reduced, however, with an increase in either temperature or salinity. Nauplii of two of the strains (parthenogenetic diploid and tetraploid) showed a progressive increase in the rate of oxygen consumption with increasing temperature up to 35°C. Nauplii of the bisexual strain appeared to be less tolerant of exposure to temperatures >30°C, since at higher temperatures their oxygen consumption declined slightly. The differences between the nauplii of the different strains in their physiological responses to changing environmental conditions appear to correlate well with their seasonal occurrence in the field.     

Effects of environmental factors and body size on rates of oxygen consumption in Archaeomysis grebnitzkii and Neomysis awatschensis (Crustacea: Mysidae)

The rates of oxygen consumption in relation to oxygen tension, temperature, salinity and body size were determined for Archaeomysis grebnitzkii Czerniavsky and Neomysis awatschensis (Brandt). Oxygen uptake was regulated by both species down to an oxygen tension of about 50 mm Hg (30% of saturation value); below this level, it was related to the oxygen tension in the medium. Oxygen tension below 20 mm Hg was lethal. A statistically significant interaction was noted for the effects of species, temperature, and salinity. In general, oxygen-uptake rate decreased with decreasing salinity in both species. The effect of size is expressed as a power function of body weight. The regression coefficients of oxygen consumption on body weight, 0.70 for A. grebnitzkii and 0.62 for N. awatschensis, were found to be significantly different from each other.Contribution No. 726 from the Department of Oceanography, University of Washington, Seattle, USA.     

Effects of sudden changes in salinity on endogenous rhythms of the spotted sea bass Lateolabrax sp.

The endogenous rhythm of oxygen consumption in juvenile spotted sea bass (Lateolabrax sp.) was measured to test the effects of sudden changes in salinity on the metabolic activity. Mean oxygen consumption rates of this euryhaline fish decreased by 13.5 to 16.0% and 25.3 to 36.4% when they were transferred from 31.5 to 15‰ seawater and to fresh water (0‰), respectively. The maximum rate of oxygen consumption was observed between 18:00 and 19:00 hrs local time, 1 to 2 h before sunset, even though they were kept in constant darkness. The peaks of oxygen consumption occurred in 23.2- and 23.3-h intervals, which correspond with a circadian rhythm, as revealed by maximum entropy spectral analysis. A markedly weakened rhythm in oxygen consumption occurred from 8 to 10 d after onset of the experiments. This study indicates that spotted sea bass can withstand sudden drops in salinity from 31.5‰ to fresh water, and yet maintain a regular though somewhat dampened endogenous rhythm of oxygen consumption. Received: 16 June 1997 / Accepted: 3 February 1998     

Effects of sulfide exposure history and hemolymph thiosulfate on oxygen-consumption rates and regulation in the hydrothermal vent crab Bythograea thermydron

The hydrothermal vent crab Bythograea thermydron is exposed to high environmental concentrations of sulfide and low levels of oxygen for extended periods of time. It has previously been shown that hydrogen sulfide is oxidized to the relatively non-toxic thiosulfate (S₂O ₃ ^2– ), which accumulates in the hemolymph. Hemolymph thiosulfate levels in freshly captured crabs vary significantly among crabs from different hydrothermal vent sites as well as between crabs from different microhabitats within the same site. Hemolymph thiosulfate concentrations were not significantly different between crabs captured at the same site 6 mo apart. Hemolymph thiosulfate concentrations ranged from 66 mol 1^–1 in a crab captured at a site with relatively low sulfide venting, to 3206 mol 1^–1 in an individual that was netted from an active smoker vent with much higher sulfide exposure. The differences in hemolymph thiosulfate between sites and the stability of hemolymph thiosulfate in crabs captured at the same site at different times suggest that sulfide exposure is significantly different between sites and that this exposure may not vary significantly over the course of a few months. B. thermydron experimentally exposed to sulfide had high levels of thiosulfate in their hemolymph and increased abilities to regulate oxygen consumption in conditions of low oxygen. This enhancement of regulatory abilities suggests that the previously demonstrated increased hemocyaninoxygen (Hc–O₂) affinity due to elevated thiosulfate may be adaptive in vivo. Average oxygen-consumption rates were much higher in crabs experimentally exposed to sulfide than in unexposed crabs. Crabs injected with isosmotic thiosulfate did not have increased oxygen-consumption rates as did the sulfide-exposed individuals, but did show a similar reduction in P _c (the critical partial pressure of oxygen at which crabs can no longer regulate oxygen consumption). This suggests that it is the sulfide exposure and/or detoxification rather than the elimination of thiosulfate that causes the increase in metabolic rate. Thiosulfate diffuses into dead crabs and into live crabs exposed to 15 mmol S₂O ₃ ^2- l^–1, indicating substantial permeability, and yet live crabs are able to eliminate thiosulfate when incubated in sea water containing 1.5 mmol S₂O ₃ ^2- l^–1, suggesting a process that has an active component.     

Quantification of arginine requirements of juvenile marine shrimp,Penaeus monodon,using microencapsulated arginine

Using microencapsulated L-arginine, the quantitative requirement for amino acid has been determined for the first time for a shrimp species, which can not effectively utilize crystalline amino acids. In an 8 wk feeding trial (1990), juvenile Penaeus monodon were fed casein-based purified diets containing one of six levels (13.1, 17.7, 22.3, 26.9, 31.5 and 36.1 g/kg diet) of arginine. In addition to the protein-bound arginine already present in the casein of the test diets, pure arginine was supplemented by L-arginine microencapsulated in cellulose acetate phthalate, which is easily assimilated. The arginine level required for optimal growth was determined by brokenline model analysis of weight gain data to be 25.0 g/kg diet (=54.7 g/kg protein). Examination of the hemolymph 3 h after feeding revealed that the free arginine level in the hemolymph had not increased as a result of increasing levels of the dietary arginine. There was an abrupt increase of urea in the hemolymph when the arginine requirement of the shrimp had been met.     

Amino and fatty acid dynamics of<Emphasis Type="Italic"> Lysmata seticaudata</Emphasis> (Decapoda: Hippolytidae) embryos during early and late reproductive season

The present study investigates amino and fatty acid dynamics of embryos of different-sized simultaneous hermaphrodite shrimp (SH) (Lysmata seticaudata) during early (ERS) and late reproductive seasons (LRS). A significant relative decrease in total amino acids and essential amino acids (EAA) was recorded (P<0.05) during the development of embryos produced by shrimp collected during ERS and LRS. The content of non-essential amino acids (NEAA) showed a smaller variation, without a marked decrease. During the last embryonic stage, the major EAAs of embryos were, in decreasing magnitude, lysine and arginine, while the major NEAAs were glutamic acid and valine. A substantial decrease in lipid content (P<0.05) was observed, and the quantitatively more important fatty acids were the saturates 16:0 and 18:0, the monounsaturates 18:1n-9 and 18:1n-7 and the polyunsaturates 20:4n-6 (arachidonic acid, ARA), 20:5n-3 (eicosapentaenoic acid, EPA) and 22:6n-3 (docosahexaenoic acid, DHA). Monounsaturates were used at a higher rate, and embryos produced by SH shrimp displayed similar consumption rates of saturated and polyunsaturated fatty acids. Considering individual fatty acids, no clear utilization pattern between different-sized SH shrimp in ERS and LRS was recorded. The inexistence of consistent differences between amino and fatty acid utilization during embryogenesis among different-sized SH shrimp in ERS and LRS emphasizes the variability affecting offspring in decapod crustaceans.Communicated by S.A. Poulet, Roscoff     

Effects of kelthane on the estuarine shrimp Crangon tranciscorum

The chlorinated hydrocarbon pesticide Kelthane was assayed for effects on food consumption, molting rate, cannibalism, respiration, mortality and behavior of the estuarine shrimp Crangon franciscorum Stimpson. The test system was a single-pass pulse-flow apparatus employing a modified Mount-Brungs style diluter. Treatment levels were 0, 14, 33 and 81 g l^-1 Kelthane at 16.6°C and 19 S over an exposure period of 317 h. Exposure to Kelthane reduced food consumption, molting rate, cannibalism, and respiration. Behavior also was abnormal in exposed shrimp. They spent more time swimming, were less coordinated and had feeble escape reactions. Kelthane was very toxic to C. franciscorum with LT₅₀ values of 163, 196 and 264 h for shrimp exposed to 81, 33 and 14 g l^-1 respectively. The incipient lethal level of Kelthane is probably below 14 g l^-1.     

Harpacticoid copepod respiration: Enhydrosoma propinquum and Longipedia helgolandica

Oxygen consumption rates of 2 harpacticoid copepods, one benthic, Enhydrosoma propinquum (Brady) and one benthopelagic, Longipedia helgolandica (Klie), were determined and compared with those of the pelagic harpacticoid Euterpina acutifrons. The respiratory rates of the 3 species differed significantly and could be correlated with activity patterns and habitat preferences. E. propinquum consumed the least oxygen, E. acutifrons an intermediate amount, and L. helgolandica the most per unit dry weight. Gravid L. helgolandica had a lower metabolic rate than the non-gravid females. The lower values for the gravid females most likely reflected the metabolic activity of the developing egg mass. Harpacticoids may be important contributors to total meiobenthic metabolism, since they respire more per unit weight than do nematodes.Supported by NSF Grant GB-8189 to Duke University Cooperative Program in Biological Oceanography (BCC) and NSF Grant 7435 (WBV).     

Daily energy requirements and trophic positioning of the sand shrimp<Emphasis Type="Italic"> Crangon septemspinosa</Emphasis>

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 (R_R), active (R_A), and feeding (R_SDA) 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 R_R and R_A 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 R_R doubled with a twofold increase in shrimp mass, and R_SDA was 0.130 J h^–1+R_R, 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     

Feeding ecology of the three juvenile phases of the spiny lobster<Emphasis Type="Italic"> Panulirus argus</Emphasis> in a tropical reef lagoon

The three juvenile phases of the spiny lobster Panulirus argus (algal phase: 5-15 mm carapace length, CL; postalgal phase: 15-45 mm CL, and subadults: 45-80 mm CL) occur in the reef lagoon at Puerto Morelos, Mexico. The algal phase abounds in this lagoon, which is covered by extensive seagrass-algal meadows, but the density of postalgal and subadult juveniles is low, owing to the scarcity of crevice-type shelters suitable for these phases. The feeding ecology of the three juvenile phases was investigated to examine whether spatial or temporal differences in food intake, diet composition, or nutritional condition occurred among phases and could partially account for the low abundance of the larger juveniles. Juveniles were collected by divers at night, from January to November 1995, throughout the mid-lagoon and back-reef zones. Percent stomach fullness, relative weight of the digestive gland (RWDG, an index of nutritional condition), percent frequency of occurrence and percent volume of food categories in the diet were compared between sexes, juvenile phases, molt stages (postmolt, intermolt, premolt), seasons, and sampling zones (mid-lagoon and back-reef zones). Significant differences in stomach fullness occurred only among molt stages, mainly because postmolt individuals had emptier stomachs. The main food categories in all juvenile phases were crustaceans (mostly hermit crabs and brachyurans) and gastropods, but the food spectrum was wide, including many other animal taxa as well as plant matter. In June 1995, the epibenthic macrofauna was sampled in five sites in the lagoon that differed in their amount of vegetation. The most abundant taxa in all sites were decapods and gastropods, but density and diversity measures showed that the distribution of these potential prey taxa for juvenile P. argus was rather patchy. Diet overlap in juvenile lobsters was high between sexes, juvenile phases, sampling zones, seasons, and molting stages, indicating that all juveniles fed on the same general food categories throughout time. The only factor that affected the RWDG was the juvenile phase. RWDG was significantly lower in subadults than in algal and postalgal phases, suggesting a poorer nutritional condition in the largest juveniles. This may be related to the scarcity of suitable shelters for large juveniles throughout the lagoon, which may preclude subadults from exploiting food resources in areas of the lagoon where shelter is limited.     

Effect of cadmium on survival and osmoregulation of various developmental stages of the shrimp Penaeus japonicus (Crustacea: Decapoda)

The object of this study was to evaluate the acute toxicity of cadmium in different post-embryonic stages of the penaeid shrimp Penaeus japonicus (Bate, 1888) and to determine the effect of sublethal cadmium on the osmoregulatory capacity used as an indicator of physiological condition. Tolerance to cadmium increases with the developmental stage. The least tolerant stages are the nauplii (48 h LC₅₀: 124 g Cd l^-1) and the zoeae (96 h LC₅₀: 10 to 30 g Cd l^-1). The most tolerant stages are the postlarvae (96 h LC₅₀: 200 to 3500 Cd l^-1) and juveniles (96 h LC₅₀: 5500 g Cd l^-1). In juvenile shrimp, 2000 g Cd l^-1 significantly reduce hypo- and hyper-OC. The effect of cadmium on hypo- and hyper-osmoregulatory capacity illustrates a dose- and time-dependent response. Surviving shrimp recover their hypo-osmoregulatory capacity after 6 d of readaptation in cadmium-free seawater.