Effects of hypoxia on the respiratory and circulatory regulation of Nephrops norvegicus

The burrowing decapod Nephrops norvegicus (L.) was kept under various degrees of hypoxia in order to measure respiration, heart rate, scaphognathite rate, haemolymph oxygen content and pH. An emergence reaction to hypoxia occurred only in dim light (<10^-2 m-c) or darkness, but after 10 d of moderate hypoxia the decapods showed no emergence response at all. The weight specific respiration of quiescent individuals was relatively low and increased only slightly in hypoxia (P_wO₂=40 torr). Heart rate, about 50 beats min^-1, changed little during hypoxia, down to P_wO₂=40 torr, whereas scaphognathite rates rose from about 60 beats min^-1 at normoxia to peak at 120 beats min^-1 at P_wO₂=40 torr. The oxygen extraction efficiency (E) remained at 20 to 30% during the first hour of hypoxia then rose gradually to maximum values of 30 to 40%. A small respiratory alkalosis of the blood became evident only after 4h of hypoxia (P_wO₂=50 torr). Normoxic postbranchial O₂ tensions (P_aO₂) were low (25–30 torr) and showed only a small decline during hypoxia. Over 10 to 13 d in moderate hypoxia an effective biosynthesis of 0.024 mM haemocyanin individual^-1 d^-1 occurred in fed decapods, whereas controls (normoxic) showed no significant change in pigment levels. A linear relationship between oxygen carrying capacity and haemocyanin concentration was found. It is contended that N. norvegicus is better able to cope with periodic exposure to hypoxia when food of sufficient quantity and quality is available.     

Effects of ambient ammonia levels on blood ammonia, ammonia excretion and heart and scaphognathite rates of Nephrops norvegicus

During commercial handling of Nephropsnorvegicus (L.) there are a number of situations when the prawns may be exposed to very high ambient ammonia levels. These experiments evaluated the effects of increased levels of ambient total ammonia (TA = NH₃ + NH₄ ⁺) on␣blood ammonia, ammonia efflux rates and on the cardio-ventilatory performance of N. norvegicus. When prawns were taken from <1 to 2000 μmol TA l⁻¹ medium, blood TA concentrations increased rapidly for the first 2 h but tended to drop thereafter. Original blood TA levels were restored 6 h after the prawns were transferred back from seawater containing 2000 to <1 μmol TA l⁻¹. Sudden exposure to 500, 1000, 2000 or 4000 μmol TA l⁻¹ medium induced blood TA concentrations to increase respectively to 50, 30, 33 and 36% of external concentrations (normally, internal TA values are much higher than external levels). Immediately after transfer back to seawater with low ammonia concentration ( <1 μmol TA l⁻¹), excretion rates were higher than those of control prawns, and the absolute amounts of TA excreted were considerably higher than those calculated to have accumulated in the haemolymph. Heart rate (HR) and scaphognathite rate (SR) were not altered when prawns were subjected to sudden alterations in ambient ammonia ( <1 to 2000 to <1 μmol TA l⁻¹). When water ammonia concentrations were altered more gradually, both rates increased, but only at 4000 μmol TA l⁻¹. These results show that N. norvegicus is able to remove ammonia from the haemolymph and/or transform ammonia into some other substance when subjected to increased levels of ambient ammonia. Possible mechanisms involved (e.g. active transport across the gills; storage in some other tissue; glutamate synthe sis) are discussed. Received: 20 May 1996 / Accepted: 1 July 1996     

Nutrient transfer in a marine mutualism: patterns of ammonia excretion by anemonefish and uptake by giant sea anemones

Many symbioses involve multiple partners in complex, multi-level associations, yet little is known concerning patterns of nutrient transfer in multi-level marine mutualisms. We used the anemonefish symbiosis as a model system to create a balance sheet for nitrogen production and transfer within a three-way symbiotic system. We quantified diel patterns in excretion of ammonia by anemonefish and subsequent absorption by host sea anemones and zooxanthellae under laboratory conditions. Rates of ammonia excretion by the anemonefish Amphiprion bicinctus varied from a high of 1.84 μmole g⁻¹ h⁻¹ at 2 h after feeding, to a basal rate of 0.50 μmole g⁻¹ h⁻¹ at 24–36 h since the last meal. Conversely, host sea anemones Entacmaea quadricolor absorbed ammonia at a rate of 0.10 μmole g⁻¹ h⁻¹ during the daytime in ammonia-enriched seawater, but during the night reduced their absorption rate to near zero, indicating that ammonia uptake was driven by zooxanthella photosynthesis. When incubated together, net ammonia excretion was virturally zero, indicating that host anemones absorbed most of the ammonia produced by resident fish. Adult anemonefish weighed about 11 g under laboratory conditions, but on the coral reef may reach up to 64 g, resulting in a maximal potential ammonia load of >200 μmole h⁻¹ produced by two adult fish during daylight hours. In contrast, host sea anemones weighed about 47 g in the laboratory, but under field conditions, large individuals may reach 680 g, so their maximal ammonia clearance rates may reach about 70 μmole h⁻¹ during the daytime. As such, the ammonia load produced by adult anemonefish far exceeds the clearance rate of host anemones and zooxanthellae. Ammonia transfer likely occurs mainly during the daytime, when anemonefish consume zooplankton and excrete rapidly, and in turn the zooxanthellae are photosynthetically active and drive rapid ammonia uptake. We conclude that zooplanktivorous fishes that form mutualisms with coral reef cnidarians may serve as an important link between open water and benthic ecosystems, through the transfer of large quantities of nutrients to zooxanthellate hosts, thus enhancing coral reef productivity.     

Haemolymph constituent levels and ammonia efflux rates of Nephrops norvegicus during emersion

Nephropsnorvegicus (L.) were subjected to 8 h of emersion, either between layers of seawater-soaked hessian with periodical (20 min) flushes of seawater (high humidity, HH) or to unprotected emersion (low humidity, LH). Blood ammonia levels rose during emersion in both groups but reached higher levels under LH conditions. Ammonia efflux rates after re-immersion were higher than those of control prawns, and amounts of ammonia excreted at such times were considerably higher than those calculated to have accumulated in the blood during emersion. Possible explanations for such differences are discussed. C _aO2 and C _vO2 decreased rapidly to ca. 10% normoxia values within 2 h of HH and LH emersion and remained low throughout the remaining emersion time. Emersion-induced tissue hypoxia increased blood concentrations of glucose and lactate. Lactate accumulation was higher during LH emersion, compared with HH emersion. Blood pH dropped ca. 0.40 units but increased again after 2 h of re-immersion. Acidosis was probably related more to respiratory difficulties (CO₂ accumulation) than to lactate accumulation, as blood lactate values remained high after 2 h of re-immersion. The ability of N. norvegicus to cope with emersion appears to be little influenced by high humidity conditions. Received: 26 June 1996 / Accepted: 5 August 1996     

Lethal effects of ammonia and nitrite onPenaeus chinensis juveniles

Juveniles of the prawnPenaeus chinensis (3.96 ±0.18 cm, 0.36±0.06 g) reared in Taiwan in 1989 were exposed to different concentrations of ammonia and nitrite, by a static renewal method in 33 seawater at pH 7.94 and at 26 °C. The 24, 48, 96 and 120 h LC₅₀ (median lethal concentration) of ammonia were 3.29, 2.10, 1.53 and 1.44 mg l^–1 for NH₃-N (un-ionized ammonia as nitrogen) and 79.97, 51.14, 37.00 and 35.09 mg l^–1 for ammonia-N (un-ionized plus ionized ammonia as nitrogen). The 24, 96, 120, 144 and 192 h LC₅₀ of nitrite-N were 339, 37.71, 29.18, 26.98 and 22.95 mg l^–1. The LC₅₀ decreased with increasing exposure time. During the first 96 h,P. chinesis juveniles were more susceptible to ammonia than nitrite. However, prawns were less tolerant to nitrite than ammonia when exposed for more than 96 h. The threshold was found at 120 and 192 h for ammonia and nitrite, respectively, on the toxicity curves. Incipient LC₅₀ was 1.44 mg l^–1 for NH₃-N, 35.09 mg l^–1 for ammonia-N and 22.95 mg l^–1 for nitrite-N. The safe value forP. chinensis juveniles was 0.14, 3.51 and 2.30 mg l^–1, respectively.     

A rapid,highly sensitive technique for the determination of ammonia in seawater

A reliable and sensitive technique for the rapid determination of ammonia in seawater is described. The procedure is based on the conversion of NH ₄ ⁺ in seawater to NH₃ and the subsequent diffusion of NH₃ across a hydrophobic membrane using flow-injection analysis. Sixty determinations can be made per hour on a flowing stream of seawater or discrete samples. The lower limit of detection is 0.05 M. Results of two applications that demonstrate the potential of this technique are presented, a laboratory excretion experiment employing the crabPachygrapsus crassipes, and small-scale vertical mapping of ammonia in the ocean.     

Effects of ammonia on survival and osmoregulation of various development stages of the shrimp Penaeus japonicus

In Penaeus japonicus, the tolerance to ammonia increased with the development from nauplius to late juvenile. The 48-h LC₅₀ of ammonia in nauplii (III–V), 96-h LC₅₀ in zoeae (I–III), mysis (I–III), post-larvae (PL1) and late juveniles (10.4±1.1 g) were respectively 5.0, 6.1 to 8.1, 9.4 to 10.9, 15.5 and 52.7 mg Nl^-1 (0.5, 0.6 to 0.7, 0.9, 1.3 and 3.1 mg NH₃–Nl^-1). In a chronic experiment (20 d), the LC₅₀ in post-larvae (PL1) was 19.1 (1.4) at 96 h and 16.2 mg Nl^-1 (1.3 mg NH₃–Nl^-1) at 480 h. Osmoregulatory capacity (OC) was calculated as the osmotic gradient between the hemolymph and the external medium at given salinities. The effects of ammonia on OC, Na⁺ and Cl^- regulation and gill Na⁺–K⁺ ATPase activity in late juveniles were examined in fullstrength seawater, SW (1050 mosm kg^-1, 36 S) and in dilute SW (450 mosm kg^-1, 15%.), after 48 or 96 h exposure to various concentrations of ammonia. Ambient ammonia disrupted both hypo- and hyper-osmoregulation; decreased OC resulted from impaired Na⁺ and Cl^- regulation. Gill Na⁺–K⁺ ATPase activity increased in SW and was not affected in dilute SW. The decrease of OC was ammonia-dose-dependent. The threshold ammonia concentrations affecting hypo-OC and hyper-OC were, respectively, 16 (1.3) and 32 mg Nl^-1 (2.3 NH₃–Nl^-1) for a 48 h exposure; these concentrations were lower than the 48-h LC₅₀ value, 65.3 mg Nl^-1 (3.5 NH₃–Nl^-1). The time course of exposure to sublethal ammonia (48 mg Nl^-1) demonstrated that the effect on osmoregulation was time-dependent. This effect was also temporary, and the exposed shrimps recovered control OC values after removal of excessive ambient ammonia. The possibility of using OC as an indicator of physiological condition in osmoregulating crustaceans and the acting mode of ammonia on osmotic and ionic regulation are discussed.     

Effect of ammonia on survival and osmoregulation in different life stages of the lobsterHomarus americanus

Acute toxicity of ammonia was determined for cultured larval, postlarval, and wild adult lobsters (Homarus americanus) in 1988. Ammonia tolerance was found to increase with ontogenetic development. Based on 96-h LC₅₀ values of 58 mg l^–1 NH₄ ⁺ + NH₃ l^–1 seawater (0.72 mg NH₃ l^–1) for Stage I larvae, 87 mg NH₄ ⁺ + NH₃ l^–1 (1.7 mg NH₃ l^–1) for Stage II larvae, 125 mg NH₄ ⁺ + NH₃ l^–1 (2.13 mg NH₃) for Stage III larvae, 144 mg NH₄ ⁺ + NH₃ l^–1 (2.36 mg NH₃ l^–1) for Stage IV postlarvae, 377 mg NH₄ ⁺ + NH₃ l^–1 (5.12 mg NH₃ l^–1) for adult lobsters at 5°C and 219 mg NH₄ ⁺ + NH₃ l^–1 (3.25 mg NH₃ l^–1) for adult lobsters at 20°C, recommendations for safe levels of total ammonia and un-ionized ammonia were calculated using an application factor of 0.1. Effects of ammonia on osmoregulatory capacity were studied on postlarvae and adults. Ability of postlarvae and adults to hyper-regulate in low-salinity media decreased after exposure to ammonia. In postlarval lobsters, osmoregulatory capacity was significantly affected in ammonia concentrations exceeding 32 mg l^–1. Osmoregulatory capacity in adult lobsters (5 and 20°C) was affected at 150 mg l^–1. In postlarval lobsters, a minimum exposure time of 12 h was required to impair osmoregulatory capacity. The decrease in hemolymph osmotic pressure was caused by lower hemolymph sodium concentrations. The presence of ammonia in the external medium could markedly affect the Na⁺/NH₄ ⁺ transport mechanism by permanently, temporarily, or partially impairing the transport sites for sodium.     

Effect of salinity on hypoxia tolerance of resting green crabs, Carcinus maenas, after feeding

Mechanisms that can influence the tolerance of hypoxia in brackish waters were studied in resting and fed crabs, Carcinus maenas, at 15?°C. Mortality, blood oxygenation, acid-base status and lactate concentration were analysed in fed crabs held in full-strength normoxic seawater (32.5‰?S) and then transferred for 24?h to a partial pressure of oxygen (P_o2) of 3?kPa (1.4?mg?l^?1) and various salinities (17, 12.5, 10, 8‰?S). At salinity levels >10‰, fed crabs tolerated P_o2 values as low as 3?kPa in the ambient water and 0.5?kPa in their arterial blood for 24?h without switching to anaerobic metabolism. Only below 10‰?S did their blood-lactate content rise, leading to their death despite the fact that their blood O₂-content was twice the control value measured in full-strength normoxic seawater and their blood P_o2 did not decrease below values recorded at higher salinity levels. Addition of CO₂ to 8‰?S water (CO₂ partial pressure increasing from 0.1 to 0.3?kPa) decreased blood-lactate production and mortality, suggesting that at 10‰?S impairment of the O₂ supply is limited by an excessive blood O₂-affinity. The results are discussed in terms of the distribution (?10‰?S) of C. maenas along salinity gradients in estuaries and bays.     

Microsensor studies of photosynthesis and respiration in the symbiotic foraminifer Orbulina universa

Oxygen and pH microelectrodes were used to investigate the microenvironment of the planktonic foraminifer Orbulina universa and its dinoflagellate endosymbionts. A diffusive boundary layer surrounds the foraminiferal shell and limits the O₂ and proton transport from the shell to the ambient seawater and vice versa. Due to symbiont photosynthesis, high O₂ concentrations of up to 206% air saturation and a pH of up to 8.8, i.e. 0.5 pH units above ambient seawater, were measured at the shell surface of the foraminifer at saturating irradiances. The respiration of the host–symbiont system in darkness decreased the O₂ concentration at the shell surface to <70% of the oxygen content in the surrounding air-saturated water. The pH at the shell surface dropped to 7.9 in darkness. We measured a mean gross photosynthetic rate of 8.5 ± 4.0 nmol O₂ h⁻¹ foraminifer⁻¹. The net photosynthesis averaged 5.3 ± 2.7 nmol O₂ h⁻¹. In the light, the calculated respiration rates reached 3.9 ± 1.9 nmol O₂h⁻¹, whereas the dark respiration rates were significantly lower (1.7 ± 0.7 nmol O₂ h⁻¹). Experimental light–dark cycles demonstrated a very dynamic response of the symbionts to changing light conditions. Gross photosynthesis versus scalar irradiance curves (P vs E _o curves) showed light saturation irradiances (E _k) of 75 and 137 μmol photons m⁻² s⁻¹ in two O. universa specimens, respectively. No inhibition of photosynthesis was observed at irradiance levels up to 700 μmol photons m⁻² s⁻¹. The light compensation point of the symbiotic association was 50 μmol photons m⁻² s⁻¹. Radial profile measurements of scalar irradiance (E _o) inside the foraminifera showed a slight increase at the shell surface up to 105% of the incident irradiance (E _d). Received: 26 January 1998 / Accepted: 11 April 1998     

Effects of body size and temperature on the metabolic rate of Penaeus monodon

Laboratory measurements of oxygen consumption were made on Penaeus monodon (Fabricius) from protozoea to adult stage at temperatures between 15° and 35°C. The logarithmic relationship between weight-specific respiration rate (WRt) and temperature (T) for two size groups, Protozoea 1 (PZ1) to Postlarva 1 (PL1) and PL to adult, are given as; WRt=10^{0.431+0.0146 (T)} (ml O₂ g^-1 h^-1) and WRt=10^{-0.948+0.0338 (T)} (ml O₂ g^-1 h^-1), respectively. Additionally, equations relating metabolic rate, temperature and size for the two size groups are; PZ1-PL1: log M=0.431+0.0146T+(1.25 (log TL)+0.579), and PL1-adult: log M=-0.948+0.0338T+(2.60(log CL)-0.683), where M=oxygen consumption in ml O₂ individual ^-1h^-1, T=temperature in °C, TL=total length in cm, and CL=carapace length in cm. Activation energies of 6 186.75 J for PZ1-PL1 and 14 066.62 J for PL-adults point to different metabolic pathways or to differences in the ratio between the metabolic pathways used.     

Growth and development of nauplii and copepodites of the estuarine copepod Acartia tonsa from southern Europe (Ria de Aveiro, Portugal) under saturating food conditions

A temperature-dependent growth model is presented for nauplii and copepodites of the estuarine calanoid copepod Acartia tonsa from southern Europe (Portugal). Development was followed from egg to adult in the laboratory at four temperatures (10, 15, 18 and 22°C) and under saturating food conditions (>1,000 μg C l⁻¹). Development times versus incubation temperature were fitted to a Belehradek’s function, showing that development times decreased with increasing incubation temperature: at 10°C, A. tonsa need 40.3 days to reach adult stage, decreasing to 8.9 days when reared at 22°C. ANCOVA (homogeneity of slopes) showed that temperature (P<0.001) and growth phase (P<0.01) had a significant effect on the growth rate. Over the range of temperatures tested in this study, highest weight-specific growth rates were found during naupliar development (NI–NVI) and varied from 0.185 day⁻¹ (10°C) to 0.880 day⁻¹ (22°C) with a Q ₁₀ equal to 3.66. During copepodite growth (CI–CV), the weight-specific growth rates ranged from 0.125 day⁻¹ (10°C) to 0.488 day⁻¹ (22°C) with a Q ₁₀ equal to 3.12. The weight-specific growth rates (g) followed temperature (T) by a linear relationship and described as ln g=−2.962+0.130 T (r ²=0.99, P<0.001) for naupliar stages and ln g=−3.134+0.114T (r ²=0.97, P<0.001) for copepodite stages. By comparing in situ growth rates (juvenile growth and fecundity) for A. tonsa taken from the literature with the temperature-dependent growth model defined here we suggest that the adult females of A. tonsa are more frequently food limited than juveniles.     

Respiratory quotient and ammonia excretion in Tilapia mossambica

Tilapia mossambica (Peters), acclimated to and tested in fresh water at 30°C, maintained a routine respiratory quotient (R Q) of about unity and an ammonia quotient (A Q) (Vol. NH₂/Vol. O₂) of about 0.2 at high ambient oxygen concentrations. At low oxygen concentrations (below 2 ppm) R Q and A Q increased sharply to values of 8 and 1, respectively (at 0.6 ppm), indicating a close relationship of increase in anaerobic energy utilization and increase in protein metabolism at inadequate oxygen concentrations. T. mossambica (8 cm), exercised continuously with intervening sampling and flushing stops for 6 h, at a swimming speed of about 2 body lengths/sec, derived some anaerobic energy throughout the exercise (R Q: 1.2), utilizing more protein the longer the exercise. The coupling of the increased protein metabolism and anaerobic energy utilization may be of advantage in preventing acidosis and also in conserving sodium (Na⁺) in fish.Part of this work was included in a paper presented at the Centennial Meeting of The American Fisheries Society, September 13–16, 1970, New York.     

Acute toxicity of ammonia to a freshwater teleost,Labeo bata larvae

Ammonia toxicity tests were performed with Labeo bata (bata) larvae of three different size groups. One hundred percent survival of larvae (500.0?±?4.0?mg) was recorded when exposed to ammonia concentrations of 1.0–13.56?mg?L^?1 at 96?h of exposure. Bata larvae exposed to ammonia concentrations of 15.8–25?mg?L^?1 showed 10–74% mortalities. The 96?h LC₅₀ value for 200 (±5), 250 (±2) and 500 (±4) mg bata larvae were 11.5, 16.8 and 22.5?mg?L^?1 un-ionised ammonia concentrations, respectively. When fish were exposed to different doses of ammonia, behavioural changes immediately occurred even at the lowest dose. At first, the fish became hyperexcitable, the skin darkened and they showed an increased ventilation frequency, fish behaviour became normal, 24?h after exposure. A 96?h LC₅₀ value of un-ionised ammonia showed direct relationship with the increasing size of bata larvae.     

Diel cycles of activity,metabolism, and ammonium concentration in tropical holothurians

Movement rate, oxygen consumption, and respiratory tree ammonium concentration were measured in situ in the holothurians Pearsonothuria graeffei and Holothuria edulis in the Agan-an Marine Reserve, Sibulan, Philippines (9°20′30″N, 123°18′31″E). Measurements were made both day and night for both species during April–July 2005. P. graeffei had significantly higher movement rate during the day than at night (1.14 and 0.27 m h⁻¹, respectively; three-way ANOVA, P < 0.05) while H. edulis had higher movement rate at night compared to the day (0.83 and 0.07 m h⁻¹, respectively), spending the daylight hours sheltering under coral. More than 80% of H. edulis had movement rate of zero during the day. Oxygen consumption of P. graeffei was significantly higher during the day than at night (1.61 and 0.83 μmol O₂ g⁻¹ h⁻¹, respectively; two-way ANCOVA, P < 0.05), but the reduction at night was not as pronounced as the reduction in movement. H. edulis had a 75% reduction in oxygen consumption during the day compared to night (0.51 and 1.96 μmol O₂ g⁻¹ h⁻¹, respectively), matching this species’ reduced movement rates during the day. Ammonium concentration in water withdrawn from the respiratory trees of P. graeffei during the day (12.0 μM) was three times higher than in respiratory tree water sampled at night (4.3 μM) and 15 times higher than ambient seawater (0.8 μM; three-way ANOVA, P < 0.05). Ammonium concentration in the respiratory tree water of H. edulis was six times higher at night (14.6 μM) than during the day (2.2 μM) and 16 times higher than that of ambient seawater (0.9 μM). Even though H. edulis and P. graeffei are found within the same coral reef environment, they may affect different substrates and reef organisms due to their different habitats and distinct but opposite diel cycles.     

The burrows and physiological adaptations to a burrowing lifestyle of Natatolana borealis (Isopoda: Cirolanidae)

The isopod Natatolana borealis Lilljeborg constructs U-shaped burrows in soft mud, the bore of which closely approximates the width of the occupant. Within artificial burrows, the isopods are largely quiescent and often adopt a position close to one of the burrow openings. Conditions within burrows constructed in the laboratory are moderately hypoxic [11.7 to 14.9 kPa (88 to 112 torr)], with isopods showing discontinuous irrigation behaviour (pleopod beating). Rates of oxygen consumption (measured at 10°C) are maintained approximately constant over a wide range of oxygen partial pressure (P_{O 2}) due, in part, to a pronounced increase in pleopod beat rate. Values for the critical partial pressure of oxygen (P_c), the P_{O 2} at which can no longer be maintained independent of P_{O 2}, were 2.0 to 3.3 kPa (15 to 25 torr). N. borealis can survive lengthy periods (65 h at 5°C) of anoxia, during which there is a significant reduction in the carbohydrate concentration and an increase in the l-lactate concentration of the tissues. The oxygencarrying capacity of the haemolymph of N. borealis was low. The haemocyanin showed a relatively high oxygen affinity [P₅₀=0.39 kPa (2.99 torr) at 10°C at the in vivo pH of 7.80] and a pronounced Bohr effect (-1.22). These characteristics may be advantageous to a burrowing mode of life and also for the conditions likely to be encountered in fish carcasses into which they burrow en masse to feed.     

Metabolic characteristics of midwater zooplankton: Ammonia excretion,O:N rations,and the effect of starvation

The nature of protein catabolism in a wide range of species of midwater zooplankton was investigated. The weight-specific ammonia excretion rates (g NH₃–N g^–1 dry wt h^–1, y) decline exponentially with minimum depth of occurreece (MDO, x), y=163.4 x^{–0.479±0.212} (95%ci) (CI=confidence interval), when temperature is held constant. The change in ammonia excretion can be partially explained by the decrease in percent protein (%P) with MDO, %P=80.17 MDO^{–0.148±0.122 (95%ci)} The atomic O:N ratio of freshly caught zooplankters ranged from 9.1 to 91, with most measurements between 9 and 25. Detailed studies were carried out on the response of one of the species studied (Gnathophausia ingens) to starvation (28 d). After 14 d of starvation the average ammonia excretion rate declined by more than 75% to less than 1 g NH₃–N g^–1 wet wt h^–1, although the average oxygen consumption declined by only 13% within the first 7 d of starvation and then remained stable. This differential response of oxygen consumption and ammonia excretion to starvation resulted in an increase in the average O:N ratio of starved animals from an initial 33 to 165 after 21 d. The average O:N ratios of fed mysids remained below 38 during the experiment. G. ingens maintains a relatively uniform metabolic rate during starvation by relying more heavily on its large lipid stores than when being fed.     

Microsensor studies of photosynthesis and respiration in larger symbiotic foraminifera. I The physico-chemical microenvironment of Marginopora vertebralis, Amphistegina lobifera and Amphisorus hemprichii

 The physico-chemical microenvironment of larger benthic foraminifera was studied with microsensors for O₂, CO₂, pH, Ca²⁺ and scalar irradiance. Under saturating light conditions, the photosynthetic activity of the endosymbiotic algae increased the O₂ up to 183% air saturation and a pH of up to 8.6 was measured at the foraminiferal shell surface. The photosynthetic CO₂ fixation decreased the CO₂ at the shell down to 4.7 μM. In the dark, the respiration of host and symbionts decreased the O₂ level to 91% air saturation and the CO₂ concentration reached up to 12 μM. pH was lowered relative to the ambient seawater pH of 8.2. The endosymbionts responded immediately to changing light conditions, resulting in dynamic changes of O₂, CO₂ and pH at the foraminiferal shell surface during experimentally imposed light–dark cycles. The dynamic concentration changes demonstrated for the first time a fast exchange of metabolic gases through the perforate, hyaline shell of Amphistegina lobifera. A diffusive boundary layer (DBL) limited the solute exchange between the foraminifera and the surrounding water. The DBL reached a thickness of 400–700 μm in stagnant water and was reduced to 100–300 μm under flow conditions. Gross photosynthesis rates were significantly higher under flow conditions (4.7 nmol O₂ cm⁻³ s⁻¹) than in stagnant water (1.6 nmol O₂ cm ⁻³ s⁻¹), whereas net photosynthesis rates were unaffected by flow conditions. The Ca²⁺ microprofiles demonstrated a spatial variation in sites of calcium uptake over the foraminiferal shells. Ca²⁺ gradients at the shell surface showed total Ca²⁺ uptake rates of 0.6 to 4.2 nmol cm⁻² h⁻¹ in A. lobifera and 1.7 to 3.6 nmol cm⁻² h⁻¹ in Marginopora vertebralis. The scattering and reflection of the foraminiferal calcite shell increased the scalar irradiance at the surface up to 205% of the incident irradiance. Transmittance measurements across the calcite shell suggest that the symbionts are shielded from higher light levels, receiving approximately 30% of the incident light for photosynthesis. Received: 6 July 1999 / Accepted: 28 April 2000     

Metabolism,enzymic activities and cold adaptation in Antarctic mesopelagic fishes

Several species of Antarctic mesopelagic fishes that have different minimal depths of occurrence but the same environmental temperature were collected in November–December 1983 and in March 1986 between 0 and 1 000 m in the open water near the marginal ice zone in the vicinity of 60°S 40°W (1983) and 65°S 46°W (1986), and oxygen consumption rate (V _{O 2}) and the activity of two metabolic enzymes, lactate dehydrogenase (LDH, an indicator of the anaerobic potential of locomotory muscle) and citrate synthase (CS, an indicator of citric acid cycle activity or aerobic potential), were determined. In four dominant species, whole-individual oxygen-consumption rate (y, ml O₂ individual^–1 h^–1) varied with weight (X, g) according to the equation y=aX ^b, with b values falling between 0.889 and 1.029. The relation of weight-specific LDH activity (y, U g^–1 wet wt) with weight (x, g) was also described by the equation y=aX ^b, with b values varying between 0.229 and 1.025. Weight-specific CS activity declined with weight, with b values from-0.031 to-0.369. V _{2 O}, LDH activity and CS activity all declined markedly with increased species' minimum depth of occurrence (the depth below which 90% of a species' population lives). Comparisons with previous studies on ecologically equivalent species of the California Borderland indicate that depth-related decreases in metabolism are the result of adapted traits of deeper-living species, not declining temperature within the water column. The metabolic rate of Antarctic mesopelagic fishes is approximately twice that of California species at equivalent temperatures; similar rates were found at the normal habitat temperatures of the two groups. Thus, a well-developed compensation for temperature is present in the Antarctic fishes: cold adaptation. Differences in enzymic activity among species, and among different sized individuals of a species are related to differences in metabolic rate and locomotory capacity. Enzymic indices can be used to estimate metabolic rates and evaluate ecological parameters such as predatory strategies and niche separation.     

Oxygen consumption and ammonia excretion of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> (Cephalopoda) in relation to body mass and temperature

The common octopus, Octopus vulgaris Cuvier, is of great scientific and commercial importance and its culture is becoming an area of increasing interest. In this study, the combined effects of temperature (T) and body mass (M) on the routine oxygen consumption rate (R) and ammonia excretion rate (U) in O. vulgaris were quantified. The experiments were conducted in a closed seawater system, and great care was taken to reduce handling stress of the animals. Temperature, salinity, pH and ammonia, nitrite, nitrate and phosphate concentrations were monitored and controlled during the experiment. The following predictive equations were evaluated: at temperatures between 13°C and 28°C and at temperatures between 15.5°C and 26°C (T_a is degrees Kelvin and M in gram). O/N ratios showed that O. vulgaris has a protein-dominated metabolism. No significant relationship between the O/N ratio and body mass or temperature was found. Sex had no significant effect on the oxygen consumption rate or on the ammonia excretion rate. For other octopod species, the dependence of metabolic rate on temperature does not differ with that for O. vulgaris.Communicated by O. Kinne, Oldendorf/Luhe