Behaviour and tolerance to hypoxia in juvenile Norway lobster (Nephrops norvegicus ) of different ages

The annual occurrence of hypoxia (<25% oxygen saturation) in the bottom waters along the Swedish west coast coincides with the postlarval settlement of Norway lobster, Nephrops norvegicus (L.). This study investigates behaviour and the experimental effects of low oxygen concentrations in juvenile N. norvegicus of different ages. All experimental individuals were reared to the juvenile (postlarval) stage in the laboratory and then given sediment as a substratum. Behavioural responses to low oxygen concentrations were tested in early and late Postlarvae 1 exposed to normoxia (>80% oxygen saturation, pO₂ > 16.7 kPa), moderate hypoxia (30% oxygen saturation, pO₂ = 6.3 kPa) and hypoxia (25% oxygen saturation, pO₂ = 5.2 kPa). The experiments were run for a maximum period of 24 h or until individuals died. Behaviour was studied using sequential video recordings of four behavioural activities: digging, walking, inactivity or flight (escape swimming up into the water column). Behaviour and mortality changed with lowered oxygen concentrations; energetically costly activities (such as walking) were reduced, and activity in general declined. In normoxia, juveniles initially walked and then burrowed, but when exposed to hypoxia they were mainly inactive with occasional outbursts of escape swimming. To increase oxygen availability the juveniles were observed to raise their bodies on stilted legs (similar to adults in hypoxic conditions), but oxygen saturations of 25% were lethal within 24 h. The results suggest that the main gas exchanges of early postlarval stages occur over the general body surface. Burrowing behaviour was tested in Postlarvae 1 and 2 of different ages held in >80% oxygen saturation for 1 wk. The difference in time taken to complete a V-shaped depression or a U-shaped burrow was measured. The results showed a strong negative relationship between postlarval age and burrowing time, but all individuals made a burrow. Juveniles were more sensitive to hypoxia than adults. Thus, the possible consequences of episodic hypoxia for the recruitment of Nephrops norvegicus and for the recolonization of severely affected areas are discussed. Received: 4 August 1996 / Accepted: 11 October 1996     

Swimming performance and endurance of the Norway lobster Nephrops norvegicus

The swimming behaviour of the Norway lobster, Nephrops norvegicus (L.), was studied in the laboratory. The lobsters were stimulated to swim repeatedly until they failed to respond to a single tactile stimulus. The position of the initiating stimulus on the body determined the height of the escape path above the bottom of the tank. A stimulus to the rostrum resulted in a low swimming trajectory, rarely exceeding a height of 0.1 m. A similar tactile stimulus to the abdomen produced higher mean swimming trajectories up to 0.5 m. Several parameters of swimming performance and endurance were measured for each swimming sequence, including distance, duration, mean and maximum velocities and the frequency and number of tail beats. Results showed no significant differences in the average swimming performance and endurance of males and females, but there was a sex difference in the relationship between swimming performance and N. norvegicus size.     

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.     

Survival and growth of the Norway lobster Nephrops norvegicus in relation to light-induced eye damage

 The long-term survival and growth of Norway lobsters, Nephrops norvegicus (L.), were examined in relation to light-induced retina breakdown. The incidence of eye damage was first assessed in tagged N. norvegicus released in Loch Torridon, on the west coast of Scotland between 1978 and 1983. Of 34 recaptures examined, all but four showed evidence of eye damage, varying in extent from 1 to 63% of the retina. Additional tagged N. norvegicus were released in Loch Torridon in 1984 and 1985 in order to compare recapture and growth rates in lobsters with and without eye damage. Three groups of N. norvegicus were released; normal sighted, partially sighted (median retina damage = 68%) and fully blinded (retina damage = 100%). From 1217 lobsters released, 235 were recovered by September 1992, when the experiment was concluded. The final proportions of N. norvegicus recaptured were found to be independent of eye-damage condition, and there was no evidence that growth rate was affected by eye damage. Combining all categories of releases, the overall proportion of females recaptured (0.244) was significantly greater than the proportion of males (0.145), suggesting better long-term survival in females than in males. At the time of release, ∼80% of the females were carrying recently spawned eggs. Although the proportion of berried females in the recaptures was slightly reduced in the two eye-damaged groups compared with the normal sighted group, the difference was not statistically significant. It is concluded that light-induced eye damage in N. norvegicus is irreversible, but such damage does not seem to influence their long-term survival, growth or reproduction. Received: 21 October 1998 / Accepted: 26 October 1999     

Some direct observations on the ecology and behaviour of the Norway lobster Nephrops norvegicus

A detailed investigation of a small area of sea bed occupied by the Norway lobster Nephrops norvegicus (L.) was carried out by diving and television observations at depths of 30 m in Loch Torridon, Scotland. The density of burrows was 1/2 m², but only a proportion of these were occupied by N. norvegicus. Although about 70% of the larger burrows were occupied by N. norvegicus, giving a density of 1 lobster/8 m², very few juveniles (carapace length less than 30 mm) were found in the area. Many of the small burrows were occupied by the gobiid fish Lesueurigobius friesii (Collett). There was evidence that N. norvegicus frequently change their burrows, and fighting for burrows was observed. N. norvegicus remain within their burrows during the day, emerge around sunset to forage for food during the night, and then return to their burrows at dawn. This, and other aspects of their burrowing behaviour, have a marked effect on the commercial trawl catches of N. norvegicus which show large seasonal and diurnal variations in size and sex composition.     

Natural variation in the concentrations of haemocyanin from three decapod crustaceans, Nephrops norvegicus, Liocarcinus depurator and Hyas aranaeus

Natural variation in the concentrations of haemocyanin ([Hc]) is examined for three decapod species collected from two different locations (Gullmarsfjord and Kattegat) off the west coast of Sweden (August to September 1998). Only one of the frequency distributions for [Hc] is normally distributed, and median values differ both between and within species. Differences in [Hc] cannot be attributed to sex, reproductive condition or moulting condition (over the limited range of moult stages examined) for any of the species. While body size did not scale with [Hc] for the Norway lobster Nephrops norvegicus or the swimming crab Liocarcinus depurator, it did for the spider crab Hyas araneus. To our knowledge this is the first time a negative relationship between body size and [Hc] has been reported for any crustacean. Thus the right-skewed frequency distribution of spider crab [Hc] could be accounted for by the right-skewed body size distribution recorded. The shape of the frequency distributions for [Hc] from the other two species could not be accounted for through differences in the factors examined here, although it is suggested that the amount of food consumed (or not) may be important. Received: 19 July 1999 / Accepted: 4 October 1999     

Age and growth in Nephrops norvegicus (Decapoda: Nephropidae)

The growth of the lobster Nephrops norvegicus (L.) in laboratory tanks was studied. The moult increments of males and females were not significantly different. The growth per moult was of the progressive geometric type throughout the life of the lobster, including the larval instars. Moulting in both the laboratory and the natural population occurred throughout the year in males, but there was a peak of moulting activity in females from June to August. There was no indication of a terminal anecdysis. Size-frequency histograms from monthly samples taken 10 to 20 miles south-west of the Isle of Man over a period of 21/2 years provided data on the growth of the year classes. It is possible to recognise the year classes in females at least, until the lobsters are 3 to 4 years old. Instar determination based on the numbers of aesthetasc-bearing segments of the antennule and on the numbers of lamellae in the endocuticle was unsuccessful. Estimates of the numbers of moults occurring in each year, and the expected mean sizes of the year classes are given.     

Between-individual variation in haemocyanin concentrations in the Norway lobster<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> following exposure to hypoxia and manganese

Environmental hypoxia and the release of reduced manganese (Mn) are often combined stress factors in marine sediments. Previous investigations have shown that hypoxia induces variable responses in the concentration of the respiratory pigment haemocyanin (Hc) of crustaceans. A recent study demonstrated that the magnitude and direction of the changes was dependent on the initial level of Hc in individuals of the Norway lobster Nephrops norvegicus. The changes also took place within 24 h rather than days as suggested by previous studies. In this present study we investigated changes of Hc in individual N. norvegicus in uncontaminated sea water after exposure to and release from realistic hypoxic stress (PO ₂=6.0 kPa). Furthermore we investigated how concomitant exposure to realistic concentrations of manganese (20 mg l ^-1) modified the responses we observed. We confirmed a between-individual variation in response and a change in Hc towards an optimum level after hypoxia, but also showed that changes that took place post-hypoxic exposure occurred over a timescale similar to that of the hypoxic response itself. Manganese exposure resulted in no significant changes in Hc whereas Mn exposure combined with hypoxia resulted in a significant decrease (15%) in all individuals independent of initial Hc. N. norvegicus was thus unable to compensate for the effects of hypoxia by synthesis of Hc after exposure to naturally occurring Mn concentrations.     

Inter-individual and within-brood variability in the fatty acid profiles of Norway lobster, <Emphasis Type="Italic">Nephrops norvegicus</Emphasis> (L.) embryos

The present study investigated the existence of inter-individual and within-brood variability in the fatty acid (FA) profile of developing embryos of Nephrops norvegicus. In all surveyed females (n = 5), the quantitatively most important FAs were as follows: 22:6n-3 (20.8 ± 3.9% average of total FA ± standard error), 18:1n-9 (19.5 ± 2.0%), 16:0 (15.2 ± 3.4%), 20:5n-3 (10.2 ± 1.4%), 16:1n-7 (8.9 ± 1.6%), and 18:1n-7 (5.7 ± 1.3%). Differences in FA profiles of embryos in the same clutch were assessed using brooding chamber side (left and right) and pleopod (1st and 2nd, 3rd and 4th, and 5th) as predictive factors. There were no significant differences in the FA composition of embryos sampled from both sides of the brooding chamber in 4 of the 5 surveyed females. However, all females exhibited significant differences in the FA profiles of embryos sampled from different pleopods. Both saturated FA (SFA) and highly unsaturated FA (HUFA) present in developing embryos exhibited marked differences along the breeding chamber. Overall, FA reserves appeared to vary significantly within broods, which can ultimately be reflected on early larval survival. A potential cause for the within-brood variation recorded in the FA profile of developing embryos include (1) differential female investment during ovarian maturation, mainly due to variation in food quality/availability; (2) differential lipid catabolism during the incubation period of developing embryos, as a consequence of embryos position within the female’s brooding chamber; or (3) differential female investment during ovarian maturation amplified by differential lipid catabolism during the incubation period.     

Locomotory behaviour and swimming performance of the Norway lobster,Nephrops norvegicus,in the presence of an acoustic tag

Observations have been made on the locomotory behaviour and swimming performances of the Norway lobster, Nephrops norvegicus (L.), fitted with an acoustic transmitter ventrally under the cephalothorax. The walking behaviour of adult males (44 mm carapace length) appeared to be unaffected, but the tag caused significant reductions in certain measures of tail-flip swimming performance such as swimming speed and endurance. Flume-tank experiments in low water currents suggested that the transmitter would increase hydrodynamic drag during swimming by 9 to 32%, depending on lobster size. Given the weight and dimensions of the acoustic transmitters currently available, it is considered advisable to confine acoustic tracking studies to relatively large N. norvegicus.Correspondence to: C. J. Chapman     

The eye and some effects of light on locomotor activity in Nephrops norvegicus

The compound eye of Nephrops norvegicus (L.) is of the superposition type, well-adapted to the low levels of light prevailing at the sea bed during the activity periods of the species. Only the proximal retinal shielding pigment responds to light, the distal retinal shielding pigment being in the dark-adapted position at all times. The response of the proximal pigment appears to vary seasonally. Field observations compared light intensity at the sea bed with the numbers of N. norvegicus caught by trawl at various times of day in the Irish Sea in summer and winter. Laboratory experiments were combined with these field data to indicate that light is an important modulator of locomotor activity in this species.     

Effects of dusk and dawn on locomotor activity rhythms in the norway lobster Nephrops norvegicus

The endogenous locomotor activity rhythm of Nephrops norvegicus (L.) shows peaks during darkness, around 3 h after dusk. At the peaks, activity occurs mainly within the burrow, although the prawns repeatedly vacate the burrows during the same periods. The phase of the rhythm shifts with a change of light-dark regime and in response to changed timing of dusk when dawn is fixed. Activity is suppressed immediately at dawn, but recommences before complete darkness during gradually reduced light at simulated dusk. This difference in response to light at dusk and dawn indicates phase responsiveness which is characteristic of endogenous rhythms entrained by light. The light-entrained endogenous rhythm is complementary to the rhythm of emergence which, reflected in commercial catches, appears to be controlled by exogenous factors.     

Investigations of carotenoids in some fauna of the Adriatic Sea. III. Leander (Palaemon) serratus and Nephrops norvegicus (Crustacea: Decapoda)

Investigations have been carried out on the carotenoids in Leander (Palaemon) serratus (Pennant) and Nephrops norvegicus (L.) (Crustacea: Decapoda) from the Adriatic Sea. The presence of the carotenoids was determined by means of columnar and thin-layer chromatography. The following carotenoids were found: Leander serratus: -carotene, echinenone, isocryptoxanthin, cryptoxanthin, canthaxanthin, lutein, isozeaxanthin, zeaxanthin, astaxanthin, and astacene; Nephrops norvegicus: -carotene, lutein, zeaxanthin, astaxanthin, and astacene.     

Heavy Metals in Tissues of the Norway Lobster Nephrops Norvegicus: Effects of Sex, Size and Season

Concentrations of mercury, cadmium, copper, zinc and iron were determined in the carapace, hepatopancreas, gills, tail muscle, ovary and eggs of 288 Norway lobsters, Nephrops norvegicus, caught in the Clyde Sea area, south of the Isle of Cumbrae, Scotland. Metal levels in males and females were examined separately in relation to size and the season in which the animals were caught.

Cadmium, copper and zinc concentrations were highest in the hepatopancreas whereas mercury and iron concentrations were highest in the gill. Levels of metals showed variations between months, with highest levels tending to occur during molt. Metal levels were influenced by lobster size with pronounced size-related increases in mercury levels in the tail muscle and cadmium levels in the hepatopancreas. Levels of several metals in the various tissues differed between the sexes.     

Modelling spatial and temporal variability in a zero-inflated variable: The case of stone pine (Pinus pinea L.) cone production

Modelling masting habit, i.e. the spatial synchronized annual variability in fruit production, is a huge task due to two main circumstances: (1) the identification of main ecological factors controlling fruiting processes, and (2) the common departure of fruit data series from the main basic statistical assumptions of normality and independence. Stone pine (Pinus pinea L.) is one of the main species in the Mediterranean basin that is able to grow under hard limiting conditions (sandy soils and extreme continental climate), and typically defined as a masting species. Considering the high economical value associated with edible nut production, the masting habit of stone pine has been a main concern for the forest management of the species. In the present work we have used annual fruit data series from 740 stone pine trees measured during a 13 years period (1996-2008) in order: (a) to verify our main hypothesis pointing out to the existence of a weather control of the fruiting process in limiting environments, rather than resource depletion or endogenous inherent cycles; (b) to identify those site factors, stand attributes and climate events affecting specific traits involved in fruiting process; and (c) to construct a model for predicting spatial and temporal patterns of variability in stone pine cone production at different spatial extents as region, stand and tree. Given the nature of the data, the model has been formulated as zero-inflated log-normal, incorporating random components to carry out with the observed lack of independence. This model attains efficiencies close to 70-80% in predicting temporal and spatial variability at regional scale. Though efficiencies are reduced according to the spatial extent of the model, it leads to unbiased estimates and efficiencies over 35-50% when predicting annual yields at tree or stand scale, respectively. In this sense, the proposed model is a main tool for facilitating decision making in some management aspects such as the quantification of total amount of cones annually supplied to nut industry, design of cone harvest programs or the optimal application of seedling felling.     

Cadmium accumulation by green algae Cladophora glomerata (L.) Kutz. (Chlorophyta) in presence of Nile tilapia Oreochromis niloticus (L.)

Accumulation of metals by aquatic organisms is mostly affected by other biological components in environments. In this study, cadmium (Cd) accumulation in green algae, Cladophora glomerata (L.) Kutz., exposed to 0.1 and 1.0 mg L^?1 of Cd for 15 and 30 days was examined in laboratory conditions in the presence of Nile tilapia Oreochromis niloticus (L.). The green algae C. glomerata accumulated Cd concentrations as 690 ± 70 and 3430 ± 470 mg kg^?1 on day 15, and 1130 ± 180 and 6830 ± 1540 mg kg^?1 on day 30. There were significant increases (p < 0.05) in metal accumulation by green algae as the exposure time and metal concentration increased. The results also indicated that the presence of Nile tilapia in the medium led to a significant Cd accumulation in the green algae compared to control (p < 0.05).     

Behaviour of endosulfan in cabbage (Brassica oleracea L.)

Residue levels of endosulfan were determined in cabbage heads after being sprayed with a formulation containing the insecticide and upon storage at room temperature (15–27°C) and in a refrigerator (2–5°C). The half-life was calculated to be 4.8 and 5.5 days. Peeling removed the residues by 92%. Similarly, washing and washing followed by cooking removed 72% and 97% residue in cabbage stored at room temperature. These treatments were slightly less effective for samples stored in a refrigerator. Household processes such as peeling, washing, and washing followed by cooking should be regular practice before the vegetables are used.     

Risk-sensitive foraging in common shrews (Sorex araneus L.)

Summary Risk-sensitive foraging theory predicts that predators which face starvation if there is a temporary shortfall in their food supply should choose feeding sites on the basis of variation in as well as mean expected reward rate. For a given mean reward rate they should choose high variance feeding sites (be risk-prone) if they are running below energy requirement, but low variance sites (be risk-averse) if they are running above.Common shrews presented with a choice between constant and variable feeding stations were more likely to visit the variable station when they were running below energy requirement and more likely to visit the constant station when they were running above. However, the tendency towards risk-aversion above requirement was greater than that towards risk-proneness below.When all shrews were considered together, the probability of visiting the variable station correlated negatively and continuously with intake relative to requirement.     

Foraging behavior of juvenile Carcinus maenas (L.) and Cancer pagurus L.

Information concerning the way juvenile crabs choose their diet from a variety of prey types can be useful for a better understanding of community dynamics, as well as for the adequate management of natural resources. Prey size and species selection by juvenile Carcinus maenas (15-35 mm carapace width, CW) and Cancer pagurus (20-40 mm CW) feeding on four bivalves of contrasting shell morphology were investigated. When offered a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas, and Cerastoderma edule presented individually, crabs generally showed evidence of size-selective predation. Cancer pagurus selected larger mussels relative to the size of their chelae (relative prey size, RPS) than did Carcinus maenas of similar and even larger carapace width. However, the RPS of selected O. edulis and Cerastoderma edule were similar for all crabs, suggesting that certain prey features constitute effective barriers even to the powerful chelae of Cancer pagurus. When offered a wide size range of mussels and oysters simultaneously, all crabs consistently selected mussels. When offered O. edulis and Crassostrea gigas, crabs consumed both these oyster species in similar numbers. Carcinus maenas consumed similar numbers of mussels and cockles; Cancer pagurus, however, showed no preference for either prey in the smaller size classes but selected more mussels than cockles as prey increased in size. Although previous studies report that adult Carcinus maenas select prey species according to their profitability (amount of food ingested per unit of handling time, milligrams per second), consumption rates of the size classes of prey selected by juvenile shore crabs did not always parallel prey value. Although variations in crab strength can account for many of the differences between the foraging strategy of juvenile and adult C. maenas, our results suggest that juvenile crabs are less species selective than adults as a result of the restrictions imposed on small individuals that have limited access to larger prey.