Ommochrome pigments in the eyes of Euphausia superba (Crustacea,Euphausiacea)

Summary Ommin and a xanthommatin-like ommochrome pigment have been extracted from the eyes of Euphausia superba, and identified by their absorbance spectra and redox reactions invarious solvents. The absorbance spectrum of ommin is relatively stable, but that of the xanthommatin pigment is sensitive to light, pH, and hydroxylamine. Both pigments are readily soluble in 2% digitonin. They are therefore likely contaminants of visual pigment extracts, and may consequently introduce artifacts into visual pigment characterizations. Microspectrophotometric measurements show that both pigments are components of the distal and proximal screening pigment granules in E. superba ommatidia.This paper is dedicated to the memory of Professor Mary Alice McWhinnie, to whom I owe a great debt of gratitude for her patient tutelage and the opportunity to participate in her Antarctic research, for the encouragement to pursue my own interests, and for her generous friendship. It was a cherished association which ended sadly with her untimely death in March, 1980. She will always be fondly remembered     

Partial characterization of chitin degrading enzymes from two euphausiids,Euphausia superba and Meganyctiphanes norvegica

Summary Chitinase and N-acetyl--D-glucosaminidase have been demonstrated in Meganyctiphanes norvegica and in Euphausia superba and partly characterized. The enzymes from both species have broad pH-optima (maxima around pH 5.0) and temperature optima between 40 and 50°C. The enzymes are relatively stable; even at about 45°C half of the enzyme activity is retained after 30 min incubation. The presence of fluoride does not affeet enzymatic activity. Chitinase activity appears in three different molecular masses, N-acetyl--D-glucosaminidases in two different forms. pH and temperature optima, thermal stability and kinetic properties of the two enzymes are strikingly similar in the polar E. superba versus the boreal euphausiid M. norvegica. Enzyme activity in the lower temperature range is still high, whereas activation energies are low in both euphausiids. This suggests a functional adaptation to a low temperature range in seawater.     

Relationships between stomach contents and vertical migration in Meganyctiphanes norvegica, Thysanoessa raschii and T. inermis (Crustacea Euphausiacea)

Stomach contents of Thysanoëssa raschii, T. inermis andMeganyctiphanes norvegica collected in the Gulf of St. Lawrenceshowed the Thysanoëssa species fed primarily during thenight in the upper 75 m of water. M. norvegica fed on copepodswhile at their daytime depth of about 125 m and on phytoplanktonand copepods during the night above 75 m depth. M. norvegicafed on a wider range of organisms than the other two speciesduring the three sample periods of spring, fall and early winter.There was no difference in the minimum size of the cells ingestedbetween any of the species. No evidence was found of any ofthe species feeding on bottom deposits during the sample periods.     

The photophores of Meganyctiphanes norvegica (M. Sars) (Euphausiacea): mode of operation

The photophores of Meganyctiphanes were investigated with regard to the control of light production and with respect to their role in a hitherto unknown communication system using light flashes which became evident from observation of specialised signalling behaviour. To that purpose the light production was recorded during presentation of a range of stimuli delivered to the intact, tethered shrimp. Stimuli used were changes in ambient light, water turbulence, simulated predator approach and light flashes, as well as electric shocks and serotonin injections. Strong negative light gradients, exaggerating the natural sunset signal, reliably elicited light production, the peak of which lasted on average 2 min. In the late phase of this light production, low frequency water oscillations and turbulent flow (assumed intraspecific communication signals at close range) elicited transient increases in light production. Artificial light flashes presented to a group of shrimp evoked a signalling behaviour in which the animal points the light of its photophore beamers (positioned at the ventral side and normally directed downwards) for a fraction of a second at observers within the same depth level. The responses produced by the signalling behaviour indicate a fixed delay with respect to the triggering flash. Electric stimulation of the ventral nerve cord via implanted electrodes resulted in a strong light production with a latency of 160 ms. Injection of serotonin, resulting in haemolymph concentrations of 10^–5 M and higher, initiated increasingly strong and increasingly long-lasting continuous light production. Implications for the control of the photophores are discussed. Electronic Publication     

Microscopic anatomy,functional morphology,and ultrastructure of the stomach of Euphausia superba Dana (Crustacea,Euphausiacea)

Summary Microscopic anatomy, functional morphology, and ultrastructure of the stomach of the antarctic krill Euphausia superba Dana were investigated by means of serial sections, scanning and transmission electron microscopy, and video technique. A separation of the stomach into an anterior part, called cardia, and a posterior part, called pylorus, became evident. Protrusions of the stomach into the midgut form the third region, called the funnel. The interior of the cardia is dominated by the two lateralia, originating from the side walls of the stomach. At their undersurface, they bear the primary filter. It separates the dorsal food channel from the ventral filtration channel, which is divided into two channels by a ventromedian ridge, the anteromedianum. Within the pylorus, the inferolateralia act in sealing the food channel from the filtration channel. In contrast to many other Malacostraca, the inferomedianum bears no secondary filter. During live observations, the stomach of Euphausia superba shows distinct pumping phases. A comparison of the structure of the stomach with data obtained from other Crustacea will lead to a better understanding about the relationships between the Malacostraca.     

The effect of lunar eclipse on the vertical migration behaviour of Meganyctiphanes norvegica (Crustacea: Euphausiacea) in the Ligurian Sea

The vertical migration of a zooplankton community dominated by the euphausiid Meganyctiphanes norvegica was monitored between 16 and 23 September 1997 with a 153 kHz Acoustic Doppler Current Profiler (ADCP) and a MOCNESS net. The sampling period covered a phase in the lunar cycle when the rise of the moon (full moon) coincided initially with sunset and then became progressively later. On 16 September 1997, a lunar eclipse occurred 45 min after sunset, lasting for 2 h. At dusk, the ADCP observed the upward vertical migration of two principal backscattering bands 10 min apart with vertical velocities of up to 7 cm s^-1. After a period at the surface, a more diffuse band subsequently sank at a slower rate (1-2 cm s^-1) to a depth of 75-100 m. Net samples showed that the earlier band consisted mainly of the pteropod Cavolinia inflexa, whilst the later band was mostly euphausiids, predominantly M.norvegica. This species was also the major constituent of the band that sank. The timing of upward migration was relatively constant over the sampling period, but there was an increasing delay of the secondary sinking until 21 September. This showed as a strong correlation between the onset of sinking and the time of moonrise. The lunar eclipse on 16 September perturbed this pattern, such that animals did not sink soon after their arrival at the surface, as occurred on 17 September but remained at the surface until the end of the umbra. This suggests that M.norvegica can perceive moonlight and that this influences vertical migration. Evidence that the behaviour is not solely mediated by this erogenous factor, however, is seen in the pattern that emerged after 21 September, when midnight sinking occurred at a relatively constant time after sunset and before moonrise. These observations support the hypothesis that moonlight is a Zeitgeber for an endogenous rhythm that synchronizes secondary sinking behaviour with the lunar cycle.      

Fluoride in tissues of Krill Euphausia superba Dana and Meganyctiphanes norvegica M. Sars in relation to the moult cycle

Summary The fluoride content of whole animals and different tissues of the euphausiid species Euphausia superba and Meganyctiphanes norvegica was analyzed by two different and improved methods of isolation and determination. In contrast to other authors our findings show that the internal organs (muscle, hepatopancreas and hemolymph) contain less than 6 ppm d.w. fluoride this being the same order of magnitude as for vertebrates. The high concentrations reported by other authors must be mainly due to contamination of the soft tissue during storage (post-mortem migration of fluoride from shell) and/or contamination caused by minute fractions of cuticle during dissection. Over 99% of the total fluoride content is located in the cuticle (i.e. integument) of the euphausiids (2600 ppm/d.w. in E. superba and 3300 ppm/d.w. in M. norvegica in pleon cuticle). Analysis of F^- levels in relation to the moulting cycle showed that the uptake in both euphausiids occurs at a comparable and fast rate during the same physiological phase shortly after moult, parallel to the general construction of the cuticle. The internal organs show homeostasis in respect to fluoride. Accordingly, no internal deposition takes place, and F^- is reaccumulated from the external medium at each moult.This work was supported by grants from the DFG No. Ad 24/9 and Bu 548/1     

Experiments on the physiology of southern and northern krill, Euphausia superba and Meganyctiphanes norvegica, with emphasis on moult and growth - a review

Physiological data are needed for life history studies on krill, and as parameters for input into energy budgets and models. In conjunction with moult and growth data, these may also prove useful for assessing the fishable biomass of krill. Here, the development of physiological concepts in experimental krill research is briefly evaluated, with emphasis on the gaps to be filled. Krill growth is very flexible, as well as strongly temperature and nutrition dependent. The polar Antarctic krill Euphausia superba grows as fast as the boreal species Meganyctiphanes norvegica, at least during the first 2.5 years, and the species are comparable in terms of physiological plasticity. Accordingly, as krill appear to adjust quickly to specific laboratory conditions, short-term experiments are essential if field conditions are to be reflected as closely as possible. Furthermore, direct comparisons between laboratory experiments and swarming studies in the field are advantageous. For these, M. norvegica is particularly well-suited, as swarms can be followed over longer times and more easily than in E. superba. For example, processes of moult and reproduction were found to be highly coordinated in swarms and populations of Northern krill. For this species a conceptual model of reproduction was developed based on a combination of short-term laboratory observations coupled with field data on moult and ovary stages. In further physiological experiments krill should be studied as groups when swarming. Using proxies, that is applying physiological and/or biochemical methods side by side, is a promising way to enhance the reliability of life history data.     

Spatial distribution of Euphausia pacifica (Euphausiacea: Crustacea) in the Yellow Sea

Euphausia pacifica were collected in the Yellow Sea in summer(August, 1997) and winter (February, 1998), and their distributionwas investigated in terms of the developmental stages in relationto environmental factors (temperature, salinity and chlorophylla). In summer, the water column was highly stratified with athermocline between 10–30 m depth, whereas the water columnwas well mixed in winter. Seasonal variation in temperaturewas large, ranging between 6.3 and 28.8°C in summer and3.9 and 12.2°C in winter. Chlorophyll a concentration changedlittle seasonally but a high concentration was observed in coastalareas. Salinity in the two seasons varied little spatio-temporally.In the studied area and sampling period, E.pacifica was themost dominant euphausiid species (99.7 and 99.8% in summer andwinter, respectively), and comprised adults and juveniles (74.6%in summer and 41.9% in winter), furcilia (17.5 and 44.9%) andcalyptopis (7.9 and 13.1%). The spatial distribution of theE.pacifica population varied seasonally depending on developmentalstage and appeared to be related primarily to the seawater temperatureand secondly to the chlorophyll a concentration, but not tosalinity. In summer, adults were abundant in areas below 10°C,and furcilia and calyptopis above 9°C. In winter, adultswere confined to areas between 7 and 10°C, and furciliaand calyptopis, to areas between 6 and 8°C. Furcilia andcalyptopis were concentrated in the vicinity, or at the centerof chlorophyll a-rich water masses, whereas adults seemed toavoid those water masses.     

Field observations on the developmental ascent of larval Euphausia superba (Crustacea)

Summary Marr's (1962) hypothesis on the descent of eggs and ascent of early larvae of Euphausia superba is examined in the light of investigations prior to and including field investigations in January 1985. Two deep stations (4100 m) west of the South Orkney Islands yielded more than 6600 eggs and early larvae per 100 m³ in the 1000 to 2000 m layer; also below 2000 m eggs and naupliar stages occurred. Sinking speed and sinking depth of krill eggs are discussed in the light of laboratory experiments of Marschall (1983), Quetin and Ross (1984), and George and Strömberg (1985). Hatching depths calculated from the experimental data on sinking rate and incubation time tend to be shallower than the deep occurrence of krill eggs at sea. Possibly krill may spawn not only near the surface but also at depths of several hundred meters. The high abundance of naupliar stages in smaples from our deepest layer corrobrate the presumption of Marschall and Hirche (1984) that the nearly immobile nauplii still sink and that the larvae begin to rise only when they have reached the metanaupliar stage.     

Larval development of Euphausia superba Dana, 1852 and a phylogenetic analysis of the Euphausiacea

Literature data and new investigations by SEM of selected ontogenetic stages of the Antarctic Krill, Euphausia superba Dana, 1852 revealed morphological characters that are either missing in, or significantly changed towards, the adult. Besides adult features, such ontogenetic characters enabled us to propose a hypothesis of the phylogenetic relationships of and within the Euphausiacea on the basis of a computer aided cladistic analysis. These are of the form (Bentheuphausia amblyops + Euphausiidae = ('Thysanopoda' + Nematobrachion + Euphausiinae = (Meganyctiphanes norvegica + Euphausiini, new name + Nematoscelini, new name = (Nyctiphanes + Nematoscelina, new name)))). From this analysis the taxon names 'Euphausiina', 'Nematoscelini', and 'Nematoscelina' are introduced for in-groups of the taxon Euphausiacea as representing monophyletic units. The position of a set of ontogenetic characters remains relatively uncertain due to the still unknown larval development of Bentheuphausia amblyops (G. O. Sars, 1883).     

Embryo and early larval stages of the Humboldt Current krill Euphausia mucronata (Crustacea: Euphausiacea)

The Humboldt Current krill, Euphausia mucronata (Crustacea: Euphausiacea), is an endemic and keystone species in the food web of this highly productive eastern border current ecosystem. The morphology and ontogeny of E. mucronata is known from calyptopis I to the adult phase, but the embryonic and early‐life stages (nauplius and metanauplius) of this broadcast spawning species are unknown. We describe the morphology and development time of these life stages to complete the knowledge of its life cycle. Embryos were obtained from purple‐gonad gravid females collected off Dichato, central Chile, during November 2012. Eight gravid females (mean=16 mm total length) were incubated in seawater at 12°C under laboratory conditions. The average development time from single‐cell embryos to the metanauplius stage was 2.2 d and hatching occurred between 20 and 25 h. The average growth rate was 0.35 mm d^?1 from the late limb bud to the metanauplius stage (range=0.21–0.48 mm d^?1). Embryos of E. mucronata had a mean chorion diameter of 0.460 mm, embryo diameter of 0.343 mm, and perivitelline space of 0.056 mm. Our biological information reported here constitutes a baseline for future ecological studies on distribution and temporal variability of spawning activity and reproductive strategies of E. mucronata in the highly variable and productive Humboldt Current ecosystem.     

Studies on metabolic properties in the Northern Krill, Meganyctiphanes norvegica (Crustacea, euphausiacea): influence of nutrition and season on pyruvate kinase

The specific activity and the kinetic properties of partly purified pyruvate kinase (PK) (EC 2.7.1.40) from the Northern Krill, Meganyctiphanes norvegica, were investigated in relation to varying food resources. In order to evaluate the effect of starvation on the total energy metabolism, the respiration rates of fed and unfed krill were determined. The FPLC-elution profile of PK displayed two distinct peaks - PK I and II. The first isoform represented 80% of the total PK activity in the organism, and 20% was contributed by the second isoform. PK I was inhibited by ATP but was not influenced by fructose-1,6-bisphosphate (FBP). In contrast, PK II showed ATP inhibition and up to 2.5-fold increased activity by addition of 17 micromol.l(-1) FBP. The Michaelis-Menten constants of both isoforms were 2-10-fold higher for ADP than for phosphoenolpyruvate (PEP). Alanine showed no regulatory effect on PK I and II. In specimens starved for 7 days oxygen consumption decreased by 20%. Neither the feeding experiments nor the animals captured in the field during low and high productive seasons indicate that PK properties of M. norvegica are modified in relation to food supply. Accordingly, alternative mechanisms are involved in the depression of the metabolic rate in terms of oxygen consumption.     

Feeding appendages of the Euphausiacea (Crustacea)

The Euphausiacea comprises about 85 species and the structure of the feeding appendages of 68 of these species is discussed here. A considerable uniformity is apparent in the appendages throughout the order but generic, and even in some cases, specific differences are evident. It is concluded from the study of the morphology of the appendages and the analyses of stomach contents that the majority of species in the genera Bentheuphwsia, Thysanopoda, Meganyctiphanes, Nyctiphanes, Pseudeuphausia, Euphausia, Tessara-brachion and Thysanoessa are omnivorous; that is, they can feed by filtering material from the water and act as predators of small zooplankton, especially copepods. Species in the genera Nematoscelis, Nematobrachion and Stylocheiron can feed on bottom deposits and also by predation of zooplankton but the amount of filter-feeding which they do may be limited because their mouthparts are not so well adapted for filtering as those of the previous group of genera.
It is suggested, from the study of the appendages and several other features of the animals, that the genera Thysanopoda, Meganyctiphanes, and Euphausia are closely related to one another and that a similar relationship exists between the genera Nematoscelis, Nematobrachion and Stylocherion. The genera Nyctiphanes and Pseudeuphausia are probably more closely related to the Thysanopoda group of genera and the genera Tessarabrachion and Thysanoessa to the Nematoscelis group of genera.     

The organization of the olfactory lobes in Euphausiacea and Mysidacea (Crustacea,Malacostraca)

Summary The structural organization of the olfactory lobes in representatives of euphausiid and mysid crustaceans was investigated and compared, also with these structures described in other crustaceans and in insects. In the investigated euphasiid and mysid species, the olfactory-globular tract and the position of cell clusters associated with the olfactory lobes show a similar arrangement. This arrangement is in agreement with that described in decapod crustaceans. The olfactory lobe neuropil in representatives of both taxa shows glomerular arrangement. These glomeruli are partly enclosed in a glial wrapping, and they represent the only site where synaptic contacts are established within the olfactory lobes. This glomerular arrangement appears similar to that described in the antennal lobe of insects, but differs from the columnar arrangement described in decapod crustaceans. Furthermore, about 15–20 FMRFamide-like immunoreactive globuli cells were labeled and they occupy a similar position in the investigated species. Neurites from these cells terminate only in the olfactory lobe glomeruli, and they are therefore regarded as intrinsic interneurons. The absence of serotonin-like immunoreactivity in the olfactory lobes is a feature only ascribed to the euphausiids and mysids. A specific neuropil area is present in male mysids, and it occupy a position forward of the olfactory lobe. The male-specific neuropil in mysids and the macro-glomerular in insects complex are interpreted as analogous structures.     

On the physical oceanography of the Kattegat and Clyde Sea area, 1996–98, as background to ecophysiological studies on the planktonic crustacean, Meganyctiphanes norvegica (Euphausiacea)

 The euphausiid, Meganyctiphanes norvegica (M. Sars), has an extremely wide distribution around the North Atlantic, from Canada to the Arctic to the western Mediterranean. It appears to be adaptable to life under quite different oceanographic conditions, yet no morphological features have been identified that distinguish any of the populations occurring in different areas between which exchange must be supposed to be restricted. The genetic, physiological, behavioural and ecological investigations that are being carried out on M. norvegica in the Kattegat, the Clyde Sea area and the Ligurian Sea (the EU MAST III PEP Project) in order to study its adaptability must be closely related to the physical environment within which the respective populations occur. This paper presents a summary of the hydrographic conditions found in the Kattegat and the Clyde Sea area in summer and in winter, during each research cruise, and places them in the context of current understanding of the hydrography in each area. An account of the hydrography of the Ligurian Sea is in preparation (Dallot et al., in preparation). The data will be used in the further analysis of genetic, physiological and behavioural adaptability of M. norvegica, and will be available for use as appropriate to supplement other hydrographic studies in the regions concerned. Received: 2 February 1999 / Accepted: 6 April 1999     

The physiological response of Northern krill (Meganyctiphanes norvegica) to temperature gradients in the Kattegat

The Alkor-Deep (140 m), which forms part of a depression system in the northern Kattegat channel east of the island of Læsø (Denmark), is the location of a self sustaining population of Northern krill, Meganyctiphanes norvegica (Euphausiacea). This population is exposed to one of the most pronounced thermal gradients within the distributional range of this pelagic crustacean. During summer, the temperature of the water column ranges between 4 and 6 in the deep to 16 °C near the surface which results in the krill being exposed to temperature differences of 8–10 °C during diel vertical migration. Oxygen consumption rates were used to investigate the physiological adaptation of the animal to such gradients in temperature. The rates were found to increase exponentially from 31 mol O₂ h^-1 g_dw ^-1 at 4 °C to 72 mol O₂ h^-1 g_dw ^-1 at 16 °C, giving a Q ₁₀-value of 2.0, and indicating that physiological adaptation to varying thermal conditions does not take place. Behavioural adaptations are discussed which may help the krill to cope with large temperature gradients in their environment.