Behavioural differences in relation to pycnoclines during vertical migration of the euphausiids Meganyctiphanes norvegica (M. Sars) and Thysanoessa raschii (M. Sars)

An observation on differences in the vertical migration patternbetween Meganyctiphanes norvegica (M. Sars) and Thysanoessaraschii (M. Sars) caused by the presence of a thermocline isreported. Generally, M.norvegica did not migrate through a thermoclineat 50–60 m depth while T.raschii did. This observationis discussed in the context of how the combination of differencesin vertical migration and advection over sill depth may affectthe abundance of fjord stocks of these two euphausiid species.     

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.      

Euphausiid visual pigments. The rhodopsins of Euphausia superba and Meganyctiphanes norvegica (Crustacea, Euphausiacea)

The rhabdoms of Euphausia superba contain one digitonin-extractable rhodopsin, lambda max 485 nm. The rhodopsin undergoes unusual pH- dependent spectral changes: above neutrality, the absorbance decreases progressively at 485 nm and rises near 370 nm. This change is reversible and appears to reflect an equilibrium between a protonated and an unprotonated form of the rhodopsin Schiff-base linkage. Near neutral pH and at 10 degrees C, the rhodopsin is partiaLly converted by 420-nm light to a stable 493-nm metarhodopsin. The metarhodopsin is partially photoconverted to rhodopsin by long-wavelength light in the absence of NH2OH; in the presence of NH2OH, it is slowly converted to retinal oxime and opsin. The rhodopsin of Meganyctiphanes norvegica measured in fresh rhabdoms by microspectrophotometry has properties very similar to those of the extracted rhodopsin of E. superba. Its lambda max is 488 nm and it is partially photoconverted by short wavelength irradiation to a stable photoconvertible metarhodopsin similar to that of E. superba. In the presence of light and NH2OH, the M. norvegica metarhodopsin is converted to retinal oxime and opsin. Our results indicate that previous determinations of euphausiid rhodopsin absorbance spectra were incorrect because of accessory pigment contamination.     

Ecological investigations on the zooplankton community of Balsfjorden, northern Norway: population dynamics of the euphausiids Thysanoessa inermis (Kroyer), Thysanoessa raschii (M.Sars) and Meganyctiphanes norvegica (M.Sars) in 1976 and 1977

The population dynamics of the euphausiids Thysanoessa inermis(Kröyer), T. raschii (M.Sars), and Meganyctiphanes norvegica(M.Sars) have been followed in Balsfjorden in 1976 and 1977.Seasonal variations in length-frequency distributions, growthin carapace length, sex-ratio, and spermatophore productionand attachment are presented and discussed in relation to changesin hydrography and phytoplanlcton standing-crop. An annual generationof T. inermis and T. raschii was spawned in April and May. Eggsand larvae of M. norvegica were not found in Balsfjorden, indicatingthat recruitment occurs from outside the fjord. T. inermis andT. raschii reached maximum carapace lengths of 7–8 mmand 6–7 mm respectively and had life-spans of c. 2 years3 months. M. norvegica had a life-span of c. 2 years 6 monthsand reached a maximum carapace length of c. 12.5 mm. In bothT. inermis and T. raschii 0-group underwent the greatest lengthincreases from May to October, I-group from March to Augustand II-group from April to June. The population structure, growthpatterns and growth periods of M. norvegica were difficult todiscern. The phytoplanlcton cycle appears to be the dominantfactor regulating both growth and spawning of the Thysanoessaspp in Balsfjorden, while temperature has no obvious influence.     

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     

Quantitative analysis of the invasion with gregarinids (Sporozoa: Gregarina) of the euphausiid Thysanoessa raschii (Crustacea: Euphausiacea) from the Barentsev sea

An occurrence of the gregarinids in the gastrointestinal tract of euphausiid Thysanoessa raschii was studied in summer 1982 and winter 1983 in the southern part of the Barents Sea. The infection rates of T. raschii with gregarinids was 0% in winter and 89.4% in summer. The highest level of infection was 1035 gregarinida per crustacean. The level of infection was found to be related mainly to be food composition (no gregarinids was found in euphausiids with copepods in stomach, the high level--in crustaceans with phytoplankton in the gastrointestinal tract).     

Effects of temperature on metabolism, growth and growth efficiency of Thysanoessa longipes (Crustacea: Euphausiacea) in the Japan Sea

Growth (assessed from intermolt period and molt increment) andmetabolism (oxygen consumption) of juvenile and adult Thysanoessalongipes from central Japan Sea were determined at eight differenttemperatures ranging from 0 to 14°C. The intermolt periodshortened progressively as temperature increased from 0 to 14°C.The molt increment was not assessed satisfactorily in the rearingexperiments, and therefore this was estimated from the naturalgrowth curve and habitat temperature, combined with laboratory-obtainedintermolt data. Oxygen consumption rates increased exponentiallyfrom 0 to 8°C and then leveled off. From these results,the growth was expressed as a function of temperature and bodysize and metabolism as a function of temperature. Because ofthe differential effects of temperature on growth and metabolism,the net growth efficiency [NGE: 100 x growth/(growth + metabolism)]changed with temperature. The temperature at which T. longipesattained maximum NGE varied from 0 to 8°C, depending onthe body length of specimens of 5, 10, 15 and 20 mm. The presentresults are compared with our previous data on Euphausia pacificain relation to the body composition and habitat usage of thesetwo trophically important species in the food web of the JapanSea.     

Vertical distribution, population structure and life history of Thysanoessa longipes (Crustacea: Euphausiacea) around Yamato Rise, central Japan Sea

The diel vertical migration, growth and spawning season of theeuphausiid, Thysanoessa longipes, were investigated using seasonalsamples collected from waters around the Yamato Rise, centralJapan Sea, during the period 1987 to 1999. Thysanoessa longipeswas present throughout a broad bathymetric layer extending downas deep as 1000 m. There was a clear trend for larger specimensto occur at deeper depths. The peak of abundance of the totalpopulation occurred at depths of 30–300 m at night, and150–500 m during the day, and the distance of the dielvertical migrations of the total population was estimated tobe between 100 and 150 m. Population structure analysis revealedthe occurrence of three cohorts aged 0+, 1+ and 2+ years, withfemales attaining a larger body size than males. Growth as determinedby body length was found to fit well to the von Bertalanffygrowth equation. The estimated life span for males and femalesis 3 years, and females reach maturity in 2 years. Based onthe occurrence of calyptopis larvae, spawning of T. longipeswas estimated to occur over only a limited period of the yearbetween April and May.     

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.     

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.     

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     

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     

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.     

Distribution and diel vertical migration of the euphausiid Thysanoessa macrura in Gerlache Strait,Antarctica

Thysanoessa macrura was found throughout Gerlache Strait, Antarctica, during four surveys carried out from 30 October to 23 November 1989, with the highest abundance being 332 individuals m^–2 (0–290 m). Reproduction had begun just before the surveys took place, as indicated by the presence of females with attached spermatophores and of larvae. Thirteen-month old females were reproductive. Larvae in 9 depth strata between 0–290 m were dominated by calyptopis stages, and developed from calyptopis 1 to furcilia 1 during November. Larval abundance was not correlated to chlorophyll a concentration, which showed a consistent east-west gradient in Gerlache Strait with highest concentrations (>30 mg chlorophyll a m^–3) in bays of the Antarctic Peninsula. Survival of larvae appeared to not be affected by phytoplankton abundance. Older T. macrura showed strong diel vertical migration between the surface at night and depths to 120 m during mid-day. Larvae were consistently found in the chlorophyll a-rich upper 50 m during night (90%) and day (81%), while adults and juveniles were found in the upper 50m at night (83%), but only 16% remained there during the day.     

Feeding behaviour in Meganyctiphanes norvegica (M. Sars) (Crustacea: Euphausiacea)

Meganyctiphanes norvegica (M. Sars) will feed upon the centric diatom Thalassiosira weissflogii (Grunow) G. Fryxell & Hasle but cannot fulfil its energy requirement for metabolism on this food. Its daily metabolic requirement can be exceeded when the euphausiid feeds upon the copepods Calanus finmarchicus Gunner or Centropages typicus Krøeyer, but not when feeding upon the smaller copepods Pseudocalanus spp. or Acartia spp. When feeding upon a natural copepod assemblage Meganyctiphanes norvegica requires high concentrations of copepods to achieve its metabolic requirements, suggesting that the euphausiid may exploit vertically patchy concentrations of prey. Short-term predation rates on Pseudocalanus spp. were also used to estimate feeding rates. Feeding in Meganyctiphanes norvegica appears to be adapted to a spatially variable food supply and rapid exploitation of food sources concentrated into patches or layers. The filter area of the feeding basket of M. norvegica is proportionally smaller than the filter area of Euphausia superba Dana, but has the same allometric length exponent. The filter area probably reflects the difference between the primarily carnivorous diet of Meganyctiphanes norvegica and herbivorous diet of Euphausia superba.     

Some observations on moulting, growth and maturation of krill (Thysanoessa inermis) from the Barents Sea

The moulting interval of the euphausiid Thysanoessa inermisfrom the Barents Sea was found to be temperature dependent withinthe range of 1.5–8.0°C. Each moult accounted for a2.1% loss of dry body wt, equivalent to a 1.2–1.3% lossof body carbon or nitrogen. During 5 months of observation bothbody length and secondary sexual characters (thelycum and petasma)of specimens varied greatly, including positive and negativegrowth periods and progression and regression of maturity stages.Characteristics of eggs spawned by one female are described.     

Relationships between geotaxis/phototaxis and diel vertical migration in autotrophic dinoflagellates

Marine dinoflagellate diel vertical migrations are often conceptuallyexplained by a species' geotactic and phototactic preferences,but actual simultaneous measurements are rare. Newly collectedsimultaneous measurements on Heterocapsa (Cachonina) illdefina(Herman and Sweeney) and Gymnodinium breve (Davis) are combinedwith similar literature information on Amphidinium carterae(Hulbert), Peridinium faeroense (Paulsen) and Prorocentrum micans(Ehrenberg) to explore several examples of the actual relationshipsbetween diel vertical migration and geotaxis/phototaxis. Amphidiniumcarterae does not migrate, but it exhibits a negative geotaxisthat may counter a small sinking velocity. The four other speciesall exhibit diel vertical migrations that yield surface aggregationsduring daylight, but the associated combinations of geotaxisand phototaxis precision (which is strongest when every cellin a population exhibits the same response to a stimulus andweakest when the response is random) and sign [which is positive(negative) when motion is toward (away from) the stimulus] aredifferent in each case. These different taxis combinations maybe related to species-specific sensor structure and/or placement.Furthermore, variations in the different biochemical pools overa species' cell cycle may contribute to structural/mechanicalchanges that influence how a given sensory array functions ata given time. If so, this coupling may be an important linkin the growth optimization mechanisms and occasional bloom successesof different autotrophic dinoflagellate species under varyingenvironmental conditions.     

Egg Size and Formation of the Geographical Range in <Emphasis Type="Italic">Thysanoessa inermis</Emphasis> (Kroyer, 1846) (Crustacea: Euphausiacea) in the Norwegian and Barents Seas

Using materials on the euphausiacean Thysanoessa inermis collected in the Norwegian and Barents seas (from 60°N to 73°N) we studied the variability of size characteristics in the eggs (diameter of embryos and egg capsules and the width of the perivitelline space) and their correlations with water temperature and salinity. The size of embryos showed almost no variability irrespective of the location of the sampling site and the water temperature and salinity; the perivitelline space performs a protective function, and its width showed a tendency to decrease with increasing water temperature and salinity. The similarities in the size of embryos in T. inermis populations thousands of kilometers away from each other might be explained, first, by the relatively small age of the populations (the age of the population from the Barents Sea did not exceed 6000–7000 years) and second, by movement of the crustaceans with currents from the Norwegian Sea to the Barents Sea.