A laboratory study of behavioral interactions of the Antarctic keystone sea star Odontaster validus with three sympatric predatory sea stars

The circumpolar sea star Odontaster validus is ubiquitous in the nearshore marine benthos of Antarctica. Despite its ecological importance, little is known of its behavioral interactions with other common sympatric sea stars. To examine these interactions we employed time-lapse video analyses conducted in a large laboratory tank (1.8 m diameter circular tank, 1,629 L). In each experimental trial, 34 adult O. validus were placed in a tight circular grouping on one side of the tank, and one adult individual of one of three common sympatric species of predatory sea star (Labidiaster annulatus, Diplasterias brandti, or Perknaster aurorae) was placed on the opposite side of the tank. Digital images of sea star movements were then captured at one min intervals over a 24 h period and aspects of sea star movements subsequently analyzed. Each 24 h treatment was replicated three times, as was a control treatment consisting only of O. validus. O. validus had significantly elevated levels of activity in the presence of P. aurorae when compared with the other two sea stars (potential chemically mediated response), and displayed a distinct “flight response” (change in direction and twofold to sixfold increase of speed) upon tactile contact with this species. Moreover, an “alarm response” was detected when individuals of O. validus that encountered a fleeing conspecific also fled the vicinity. In contrast, our results indicated that O. validus displays virtually no chemical or tactile behavioral responses to the large multi-armed L. annulatus and only weak tactile responses to D. brandti.     

Encounters between Antarctic limpets, <Emphasis Type="Italic">Nacella concinna</Emphasis>, and predatory sea stars, <Emphasis Type="Italic">Lysasterias</Emphasis> sp., in laboratory and field experiments

Antarctic limpets, Nacella concinna, from the Admiralty Bay (King George Island, South Shetlands) for at least part of the year (austral winter) co-exist with predatory sea stars Lysasterias sp. Our laboratory and field experiments established that the presence of Lysasterias sp. or its odour had considerable influence upon their behaviour. Limpets’ responses, consisting of shell mushrooming, shell rotation and flight, were distinctly different from their reaction to other stimuli, such as food and conspecific odours, or mechanical stimulation. Moreover, a significant impact of sea star presence on limpets’ activity was observed, with limpets fleeing to a distance of 60 cm from the predator. Such reactions allow limpets to lower the incidence of sea star predation, but at the cost of presumptive disrupting of foraging and an additional energy expended for locomotion. A visible difference was noted between two limpet populations, with the rockpool limpets responding only after physical contact with being touched by a sea star, and the subtidal ones responding at a distance of up to 20 cm.     

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.     

Gametogenic and reproductive cycles of the sea anemone, <Emphasis Type="Italic">Entacmaea quadricolor</Emphasis>

Host sea anemones are ecologically important as they provide habitat for obligate symbiotic anemonefish in many areas of the Indo-Pacific. Despite their importance, no information is available on their gametogenic cycles. This study aimed to address this lack of knowledge by determining the gametogenic cycles of Entacmaea quadricolor. Gonad samples were taken from January 2003 to February 2005 at North Solitary Island, Solitary Islands Marine Park, Australia using a specially developed non-lethal field biopsy sampling technique. Sampling was done 17 times during the study period, with 15–20 individuals being sampled on each occasion. Samples were examined prior to fixation, and then histologically sectioned to determine the reproductive activity of each individual. Female anemones were significantly more abundant than males, and had asynchronous oocyte development both within and among individuals. Male anemones showed a single annual cycle of spermary growth, development and spawning. Data from the 26-month study indicated that spawning occurred in the austral summer and autumn between January and April, which coincided with the observed spawning periods that have previously been documented for this species in outdoor flow-through seawater tanks at the study location. The biopsy sampling technique used during this study provides an opportunity to gain a more thorough understanding of the gametogenic cycles and sexual pattern of host sea anemones throughout their distribution.     

Symbiont photosynthesis increases both respiration and photosynthesis in the symbiotic sea anemone Anemonia viridis

Dark respiration of the symbiotic sea anemone Anemonia viridis (Forskäl) was observed to increase by 34% when anemones were exposed to hyperoxic sea water (150% oxygen saturation) overnight, and by 39% after exposure to 6 h in the light at a saturating irradiance of 300 E m^-2 s^-1 at normoxia (100% oxygen saturation). No increase due to light stimulation was observed in aposymbiotic control anemones. In darkness, the oxygen concentration of the coelenteric fluid was hypoxic. However, within 10 min of anemones being illuminated, coelenteric fluid was hyperoxic, and it remained elevated throughout a 12 h light period. When measured over a 24 h period (12 h light: 12 h dark), the dark respiration rate increased gradually over the first 6 h of the light period until it was 35% above the dark night-time resting rate. It remained elevated throughout the remaining light period and for 2 h into the following dark period, after which it fell back to the resting rate. Gross photosynthesis (P ^gross) increased significantly when anemones were exposed to either hyperoxia (150% oxygen saturation) or 300 E m^-2 s^-1 at normoxia. This increase was not observed when symbiotic anemones were illuminated at a low-light intensity of 100 E m^-2 s^-1. The results of this study suggest that respiration in the dark is limited by oxygen diffusion and that normal respiration is restored in the daytime by utilisation of the oxygen released by photosynthesis. Furthermore, it appears that the increased respiration following exposure to high-light intensities provides a CO₂-rich intracellular environment which further enhances the photosynthetic rate of the zooxanthellae.     

Chemical mediation of invertebrate defensive behaviors and the ability to distinguish between foraging and inactive predators

When the predatory sea star Pycnopodia helianthoides was placed upstream, the sea urchin Strongylocentrotus purpuratus responded defensively by extending and opening its globiferous pedicellariae. No pedicellaria response was given in control seawater or when the sea star was downstream. The snail Tegula funebralis responded by moving up vertical surfaces when Pycnopodia helianthoides or when Pisaster ochraceus were placed upstream. When these sea stars were introduced downstream, the snail's response was not significantly different from that in control seawater. Water collected from an aquarium containing a single sea star was sufficient to trigger the response of S. purpuratus and T. funebralis; the physical presence of the sea star was not essential. This indicated that a chemical stimulus was involved, and the lack of responses when sea stars were downstream argued strongly against the possible additional involvement of visual or vibrational stimuli. S. purpuratus gave stronger pedicellaria responses to water flowing over an active Pycnopodia helianthoides than to water flowing over the same sea star when it was inactive. The significance of the ability to distinguish between actively foraging and inactive predators is discussed, and a mechanism is proposed to explain differences in the amount of stimulatory chemicals released by active and inactive sea stars.     

Effects of starvation,and light and dark on the energy metabolism of symbiotic and aposymbiotic sea anemones,Anthopleura elegantissima

Rates of oxygen and carbon-dioxide exhange were measured in symbiotic and aposymbiotic specimens of the sea anemone Anthopleura elegantissima while fed and starved under light or dark conditions. Respiratory quotients indicated that fed anemones switched from a carbohydrate to a fat catabolism when starved, with the exception that symbiotic individuals starved in the light showed a pronounced carbohydrate catabolism for over 1 month. The source of the carbohydrate was probably photosynthate translocated by the dinoflagellate Symbiodinium (=Gymnodinium) microadriaticum (Freudenthal) living in the anemones' tissues. The starved symbiotic anemones maintained in the light had lipid levels not significantly different from fed controls and 44 to 61% higher than starved aposymbiotic anemones after 1 month. Thus, the quality and quantity of the metabolic flux from the symbionts to the sea anemone were sufficient to conserve the host's lipid reserves.     

Feeding aggregations of sea stars (<Emphasis Type="Italic">Asterias</Emphasis> spp. and <Emphasis Type="Italic">Henricia sanguinolenta</Emphasis>) associated with sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) grazing fronts in Nova Scotia

Migrating feeding aggregations (or fronts) of sea urchins can dramatically alter subtidal seascapes by destructively grazing macrophytes. While direct effects of urchin fronts on macrophytes (particularly kelps) are well documented, indirect effects on associated fauna are largely unknown. Secondary aggregations of predators and scavengers form around fronts of Strongylocentrotus droebachiensis in Nova Scotia. We recorded mean densities of the sea stars Asterias spp. (mainly A. rubens) and Henricia sanguinolenta of up to 11.6 and 1.7 individuals 0.25 m⁻² along an urchin front over 1 year. For Asterias, mean density at the front was 7 and 15 times greater than in the kelp bed and adjacent barrens, respectively. There was strong concordance between locations of peak density of urchins and sea stars (Asterias r = 0.98; H. sanguinolenta r = 0.97) along transects across the kelp–barrens interface, indicating that sea star aggregations migrated along with the urchin front at rates of up to 2.5 m per month. Size–frequency distributions suggest that Asterias at the front were drawn from both the barrens (smaller individuals) and the kelp bed (larger individuals). These sea stars fed intensively on mussels on kelp holdfasts and in adjacent patches. Urchin grazing may precipitate aggregations of sea stars and other predators or scavengers by incidentally consuming or damaging mussels and other small invertebrates, and thereby releasing a strong odor cue. Consumption of protective holdfasts and turf algae by urchins could facilitate feeding by these consumers, which may obtain a substantial energy subsidy during destructive grazing events.     

Prey defense,predator preference,and nonrandom diet: The interactions between Pycnopodia helianthoides and two species of sea urchins

Interactions between the predatory sea star Pycnopodia helianthoides (Brandt, 1835) and two of its natural prey, the sea urchins Strongylocentrotus purpuratus (Stimpson, 1857) and S. franciscanus (Agassiz, 1863), are examined with regard to predator preference, predator diet, and prey defenses. The sea star is able to detect both species of sea urchin upstream in a Y-trough, but does not consistently choose one over the other (i.e., no preference). However, when the sea star is presented with equal numbers of similar-sized specimens of the two species of sea urchin, its diet is markedly nonrandom, since S. purpuratus is eaten almost 98% of the time. The defensive responses of the two species of sea urchin differ in form and effectiveness. S. franciscanus employs its long spines as defensive weapons, pinching the rays of an attacking sea star. This defensive response is more effective than the pedicellarial response used by S. purpuratus. The nonrandom diet of the predator seems to result primarily from prey defensive responses that differ in effectiveness, rather than from an intrinsic, behavioral preference of the predator at an earlier stage in the predator/prey interaction.     

Effect of feeding regime and irradiance on the photophysiology of the symbiotic sea anemone Aiptasia pulchella

To determine how the animal and algal components of the symbiotic sea anemone Aiptasia pulchella respond to changes in food availability and culture irradiance, sea anemones from a single clone were maintained at four irradiance levels (320, 185, 115, and 45 E m^-2 s^-1) and either starved or fed for 5 wk. Changes in protein biomass of sea anemones maintained under these conditions were not related to the productivity of zooxanthellae, since the protein biomass of fed A. pulchella decreased with increase in irradiance and there was no difference in protein biomass among starved sea anemones at the four irradiance levels. Except for the starved high-light sea anemones, the density of symbiotic zooxanthellae was independent of culture irradiance within both starved and fed. A. pulchella. Starved sea anemones contained over twice the density of zooxanthellae as fed sea anemones. Within both starved and fed individuals, chlorophyll per zooxanthella increased with decreasing culture irradiance while algal size remained constant (in fed sea anemones) at about 8.80 m diameter. Chlorophyll a: c ₂ ratios of zooxanthellae increased with decreasing culture irradiance in zooxanthellae from starved sea anemones but remained constant in zooxanthellae from fed sea anemones. As estimated from mitotic index data, the in situ growth rates of zooxanthellae averaged 0.007 d^-1 and did not vary with irradiance or feeding regime. Photosynthesis-irradiance (P-I) responses of fed A. pulchella indicated an increase in photosynthetic efficiency with decreasing culture irradiance. But there was no consistent pattern in photosynthetic capacity with culture irradiance. Respiration rates of fed sea anemones also did not vary in relation to culture irradiance. The parameter I _k , defined as the irradiance at which light-saturated rates of photosynthesis are first attained, was the only parameter from the P-I curves which increased linearly with increasing culture irradiance. The daily ratio of net photosynthesis to respiration for A. pulchella ranged from 1.6 to 2.8 for sea anemones maintained at the three higher irradiances, but was negative for those maintained at 45 E m^-2 s^-1. Since the final protein biomass was greatest for sea anemones maintained at the lowest irradiance, these results indicate that sea anemone growth cannot be directly related to productivity of zooxanthellae in this symbiotic association.     

A comparative analysis of the photobiology of zooxanthellae and zoochlorellae symbiotic with the temperate clonal anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis> (Brandt). III. Seasonal effects of natural light and temperature on photosynthesis and respiration

The sea anemone Anthopleura elegantissima hosts two phylogenetically different symbiotic microalgae, a dinoflagellate Symbiodinium (zooxanthellae, ZX) and a chlorophyte (zoochlorellae, ZC). The photosynthetic productivity (P), respiration (R), and contribution of algal carbon translocated to the host (CZAR) in response to a year’s seasonal ambient changes of natural light and temperature are documented for both ZX- and ZC-bearing anemones. Light and temperature both affect photosynthesis, respiration, and CZAR, as well as various algal parameters; while there are evident seasonal differences, for the most part the relative effects on P, R, and CZAR by the two environmental variables cannot be determined. Net photosynthesis (P_n) of both ZX and ZC was significantly higher during spring and summer. During these seasons, the P_n of ZX was always greater than that of ZC. Regardless of algal symbiont, anemone respiration (R) was significantly higher during the spring and summer. The annual net carbon fixation rate of anemones with ZX and ZC was 325 and 276 mg C anemone⁻¹ year⁻¹, respectively, which translates to annual net community productivity rates of 92 and 60 g C m⁻¹ year⁻¹ for anemones with ZX or ZC, respectively. CZAR did not show a clear relationship with season; however the CZAR for ZX was always significantly greater than for ZC. Lower ZX growth rates, coupled with higher photosynthetic rates and higher CZAR estimates, compared to ZC, suggest that if A. elegantissima is simply carbon limited, ZX-bearing anemones should be the dominant symbiont in the field. However ZC-bearing anemones persist in low light and reduced temperature microhabitats, therefore more than the translocation of carbon from ZC must be involved. Given that global climate change will increase water temperatures, the potential for latitudinal range shifts of both ZC and ZX (S. californium and muscatinei) might be used as biological indicators of thermal shifts in the littoral zone of the Pacific Northwest.     

Crushing predation of the spiny star Marthasterias glacialis upon the sea urchin Paracentrotus lividus

Literature data report that only fish predators are able to crush sea urchin tests in Mediterranean rocky reefs. This experimental study showed that the spiny star Marthasterias glacialis is able to break Paracentrotus lividus tests and that the breaking event is more likely to occur for small-sized sea urchins than for big ones. Our results show that the role of M. glacialis in regulating P. lividus population density can be important in specific locations. They may have important implications, moreover, for the use of tethering techniques aimed at identifying predator types of sea urchins.     

Rates of fission, somatic growth and gonadal development of a fissiparous sea star, Allostichaster insignis, in New Zealand

Seasonal changes in population structure and incidence of fission were measured in intertidal and subtidal populations of Allostichaster insignis, a fissiparous sea star. Population size structure was stable over the course of the 1-year study. Sea stars in the subtidal zone attained greater maximum size (mean arm length, R = 35 mm) than those in the intertidal population (20 mm). Fission rates were greatest among small individuals (R < 20 mm). The frequency of fission ranged from 5 to 32% with peaks in early austral summer in the intertidal zone, and in autumn and winter in the subtidal zone. Sexual reproduction occurred in early spring in sea stars larger than 12 mm. The populations were heavily biased toward males. In the laboratory, A. insignis of three size classes (small, R = 9–13 mm; medium, 19–21 mm; and large, 29–31 mm) were fed mussels ad libitum or starved (not fed macroscopic food) for ∼1 year in a 3 × 2 factorial experiment. Small and medium-sized sea stars divided throughout the experiment and the ramets of most individuals regenerated sufficiently to divide again after 6–9 months. Unfed sea stars did not undergo fission (with one exception), had a higher mortality rate, and did not grow. Small, fed sea stars grew significantly faster than medium-sized or large individuals. At the end of the experiment, the pyloric caeca index (a measure of nutritional condition) was greater in fed than in unfed animals. Gonads (only testes were observed) developed in medium-sized and large, fed sea stars. Our field and laboratory results indicate that asexual reproduction in A. insignis predominantly occurs in small, well-nourished individuals. Ramets grow gradually through repeated fission and regeneration to a size (mean length of regenerating arms, R _r ∼ 20 mm) at which they begin to switch to sexual reproduction as the dominant reproductive mode.     

The effects of symbiotic state on heterotrophic feeding in the temperate sea anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis>

The temperate sea anemone Anthopleura elegantissima is facultatively symbiotic with unicellular algae. Symbiotic A. elegantissima can supplement heterotrophic feeding with excess photosynthate from their algal partners, while asymbiotic individuals must rely solely on heterotrophy. A. elegantissima individuals were collected from Swirl Rocks, Washington (48°25′6″ N, 122°50′58″ W) in July 2010, and prey capture and feeding characteristics were measured to determine whether asymbiotic individuals are more efficient predators. Feeding abilities were then measured again after a 3-week exposure to full sunlight or shaded conditions. Freshly collected asymbiotic anemones had larger nematocysts, but symbiotic individuals showed greater nematocyte sensitivity. Sunlight enhanced digestion and reduced cnida density in all anemones regardless of symbiotic state. Results suggest that the phototropic potential of A. elegantissima, as influenced by symbiotic condition, has little effect on heterotrophic capacity. The anemones appear to maximize heterotrophic energy input independent of the presence or identity of their algal symbionts.     

Calanoid copepod escape behavior in response to a visual predator

Calanoid copepods typically exhibit escape reactions to hydrodynamic stimuli such as those generated by the approach of a predator. During the summers of 2000, 2001 and 2004, two small calanoid species, Temora turbinata Dana, 1849 and Paracalanus parvus Claus, 1863 were exposed to a visual predatory fish, the blenny Acanthemblemaria spinosa Metzelaar, 1919, and their predator–prey interactions were recorded using both high-speed and standard videographic techniques. Copepod escape reaction components, including swimming pattern, reactive distance, turning rate, and jump kinetics, were quantified from individual predation events using motion analysis techniques. Among the observed escape reaction components, differences were noted between the species’ swimming patterns prior to attack and their response latencies. Temora turbinata was a continuous cruiser and P. parvus exhibited a hop-and-sink swimming pattern. During periods of sinking, P. parvus stopped beating its appendages, which presumably reduced any self-generated hydrodynamic signals and increased perceptual abilities to detect an approaching predator. Response latency was determined for each copepod species using a hydrodynamic stimulus produced by a 1 ms acoustic signal. Response latencies of T. turbinata were significantly longer than those of P. parvus. Despite some apparent perceptual advantages of P. parvus, the blenny successfully captured both species by modifying its attack behavior for the targeted prey.     

Physiological energetics of the intertidal sea anemone Anthopleura elegantissima

Energy budgets were calculated for individuals of the sea anemone Anthopleura elegantissima (Brandt), collected in 1981 and 1982 from Bodega Harbor, California, USA. Rates of ammonium excretion were measured in high-and low-intertidal, symbiotic and aposymbiotic sea anemones within 24 h of collection. Among symbiotic and aposymbiotic individuals, no differences in excretion rate were found on the basis of intertidal height. However, rates of ammonium excretion in aposymbiotic anemones (2.14 mol NH ₊ ⁴ g^-1 h^-1) were significantly higher than in symbiotic ones (0.288 mol NH ₊ ⁴ g^-1 h^-1). Rates of excretion were used with estimated rates of oxygen uptake to calculate nitrogen quotients (NQ). NQ and RQ values from the literature were used to calculate an oxyenthalpic equivalent [501 kJ (mol O₂)^-1 for R+U], and mass proportions of protein (54%), carbohydrate (44%) and lipid (2%) catabolized during routine metabolism in this species 24 h after feeding. Integrated energy budgets of these experimental anemones were calculated from data on ingestion, absorption and growth, and estimates of translocated energy from the symbiotic algae. Contribution of zooxanthellae to animal respiration based on translocation=90% and RQ=0.97 are 41 and 79% in high-and low-intertidal anemones, respectively. Calculated scope for growth is greater than directly measured growth in both high-and low-intertidal individuals. The deficit, estimated as 30% of assimilated energy in high-intertidal anemones, is attributed to unmeasured costs (specific dynamic effect) or production (mucus). Low-intertidal anemones lost mass during the experiment, implying that the magnitude of the deficit was greater in these anemones than in upper intertidal individuals. Anemones from both shore levels lost zooxanthellae during the experiment, which contributed to energy loss since the contribution of the zooxanthellae is greater in low-intertidal anemones. Scope for growth is preserved in high-intertidal anemones (29% of assimilated energy) because metabolic demands are lower due to aerial exposure, and prey capture rate is higher compared to lowshore anemones. Although possibly underestimated, lower scope for growth in low-shore anemones may result from continuous feeding and digestion processes that are less efficient than those of periodically feeding high-intertidal anemones.     

Seasonal variations in biochemical constituents during the reproductive cycle of the female dog cockle Glycymeris glycymeris

In an attempt to describe the biochemical events associated with the main stages of the annual and reproductive cycles of the female dog cockle Glycymeris glycymeris L., we studied seasonal variations in the various stages of oocyte development of the ovaries, and the glycogen, total protein and total lipid content of five body tissues – adductor muscle, foot, tunic coat, visceral mass and mantle. From November 1991 to November 1994, microscopic examination of the ovaries and measurement of the tissue concentrations of glycogen, total proteins and total lipids in these five body tissues were made monthly on ten female dog cockles originating from the sea area around Douarnenez (south Brittany, France). Morphological studies revealed that in the population investigated the annual cycle is characterised by three major periods: a first period of vitellogenesis extending from February/March to April/May and preceding a spawning in spring; a second period of vitellogenesis extending from May/June to September/October and leading to either no spawning, a single autumnal spawning event, or to two spawning events in summer and autumn; and a third period extending from October/November to February/March and characterised by a high level of oocyte lysis. In the muscular body tissues of the dog cockle, i.e. the adductor muscles, the foot and the tunic coat (the muscular envelope containing the visceral mass), the concentrations of glycogen, total proteins and total lipids underwent very similar variations during the annual cycle. During each stage of vitellogenesis, a typical glycogen–protein–lipid sequence was observed in the muscular tissues that was characterised firstly by a peak of glycogen concentration 2 to 3 mo before spawning, followed by a peak in total proteins 1 mo before spawning, and finally by a peak in lipid content just before spawning. A similar glycogen–protein–lipid sequence was also recorded in the first half of the winter period. However, these events were followed by general atresia affecting all oocytes in the gonad. Maximum energetic value of biochemical constituents in females coincided with peaks in lipid content in the visceral mass and mantle. These biochemical events occurred principally immediately before and at the end of oocyte lysis (December/January). A drop in the total energetic value, affecting mainly the visceral mass and the mantle, was recorded each year during the period January to March, coinciding with the period of shell growth in this species. Our data clearly indicate that in female G. glycymeris all muscular tissues contribute to the storage of glycogen and proteins, and suggest that glycogen may be the source of energy triggering vitellogenesis. Biochemical and microscopic observations revealed that oocyte development takes place during the first half of winter, but that these oocytes undergo atresia in December/January. The metabolites produced from oocyte lysis could contribute to somatic growth, which occurs in late winter. Received: 3 March 1997 / Accepted: 23 July 1997     

Behavioural responses of the Dungeness crab, <Emphasis Type="Italic">Cancer magister</Emphasis>, during feeding and digestion in hypoxic conditions

The Dungeness crab, Cancer magister, inhabits areas that are frequently subject to periods of hypoxia. This species can employ physiological mechanisms that allow it to cope with acute hypoxic episodes. When crabs feed there is a general increase in physiological variables; these may pose an additional physiological burden on crabs already attempting to maintain adequate oxygen uptake in hypoxia. In Barkley Sound, British Columbia, the inshore habitats of C. magister ranged in dissolved oxygen from 28 kPa at the water surface to less than 1.0 kPa just above the sediment–water interface. During short-term hypoxic events, crabs reduced both the amount of food eaten and the amount of time spent feeding. Crabs tended to cease feeding below 3.2 kPa oxygen, but resumed feeding when the dissolved oxygen tensions were rapidly raised to 6 kPa. In a high (10.5–21 kPa) oxygen gradient, both unfed and fed crabs showed no preference for any area of the gradient. In a low (2.5–10.5 kPa) dissolved oxygen gradient, both unfed and fed crabs preferred the highest oxygen regime. In the laboratory, crabs were less likely to enter hypoxic waters (below 3.2 kPa oxygen) to obtain and consume food; those that did moved the food to a higher oxygen regime prior to feeding and settled in higher oxygen regimes for digestion. Crab behaviour was also monitored in the field. Fed and unfed crabs were fitted with ultrasonic telemetry tags and tracked during a tidal cycle. Unfed crabs remained mobile, travelling up to 1,370 m within 6 h, while postprandial crabs settled in areas of high oxygen and moved very little during the first 48 h after release. The present study suggests that C. magister exhibits behavioural responses in order to minimise the use of physiological mechanisms, and maximise foraging and digestive processes. Thus the nutritional state of the individual may be important in regulating both its behaviour and distribution in its natural environment.     

Vertical distribution and mortality of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during overwintering in oceanic waters southwest of Iceland

The seasonal vertical distribution and the predatory regime encountered by Calanus finmarchicus were studied along a transect across the Reykjanes Ridge in the oceanic area southwest of Iceland from data collected during four cruises between November 1996 and June 1997. The mortality for the overwintering period was estimated using linear regressions of density estimates from November 1996 to April 1997. In addition, we also estimated the mortality of the oldest population stages (C4 and older) in April and June by applying the vertical table method. During winter (November/December–January/February), the animals mainly resided at a depth of ∼300–1,500 m in the water of Atlantic origin. Ascent to upper layers took place mainly during March and April, and continued until May. During all cruises, continuous deep-scattering layers were observed, mainly within the range of 400–500 m to 700–800 m depth. Based on sampling with a Harstad pelagic trawl in April, the scattering was mainly ascribed to jellyfish (mainly Periphylla periphylla), small mesopelagic oceanic fishes (several species but Benthosema glaciale and Maurolicus muelleri were most abundant), euphausiids (mainly Meganyctiphanes norvegica) and shrimps (mainly Sergestes arcticus). These species may represent a predatory threat to overwintering C. finmarchicus. From November to April, daily per capita mortality rates were estimated to be (mean ± 95% CL) 0.004 ± 0.0028 for the total data set, and 0.004 ± 0.0033 and 0.004 ± 0.0023 (day^-1) for the Iceland Basin and Irminger Basin, respectively. Mortality rates were higher later in life (mean ± 95% CL) for C5/females (0.13 ± 0.044) and C5/males (0.19 ± 0.051) than for C4/C5 (0.00 ± 0.035) when averaged over all samples taken in April and June 1997. We discuss how the observed distribution and mortality rates of overwintering C. finmarchicus might be related to predatory regime.     

Ultrastructure and anaerobic metabolism of mitochondria in the marine oligochaete Tubificoides benedii : effects of hypoxia and sulfide

It has often been suggested that ultrastructural properties of mitochondria are correlated with oxygen and sulfide levels from the environment, although careful analyses of this question are rare. In this study the ultrastructure and distribution of mitochondria in Tubificoides benedii, a marine oligochaete from sulfide-rich sediments, were investigated after a series of oxic, hypoxic and hypoxic–sulfidic (200 μM H₂S) incubations up to 24 h. Succinate, one of the key endproducts of an anaerobic metabolism, was used as an indicator of mitochondrial anaerobiosis. Consistent differences in mitochondrial ultrastructure were not observed in any of the incubations, even after 24 h. Stereological parameters of mitochondria (volume density, surface density of the outer mitochondrial membrane, and specific surface) in epidermal and intestinal tissues of T. benedii were not affected by hypoxia or sulfide either. On the other hand, succinate concentrations increased significantly within 24 h under hypoxic and hypoxic–sulfidic conditions. Thus, experimental hypoxia and sulfide clearly caused mitochondrial anaerobiosis without affecting ultrastructure or distribution of mitochondria in T. benedii. Distinct differences in ultrastructural and stereological parameters were common between different tissues and between individuals, showing that different forms of mitochondria can occur within one species. Our results imply that a mitochondrial ultrastructure specific to thiobiotic animals does not appear to exist. Received: 4 August 1996 / Accepted: 20 September 1996