210Pb as a dietary indicator in the Antarctic pelagic community

The naturally occurring radionuclide ²¹⁰Po is useful as a tracer of diets of marine organisms. Data for ²¹⁰Pb, the grandparent of ²¹⁰Po, are less abundant: we therefore report here ²¹⁰Pb concentrations in a substantial collection of marine biota obtained from the Antarctic Ocean during the SIBEX II cruise in 1985. The general levels of ²¹⁰Pb are within the range of previously published data but, as in the case of ²¹⁰Po, there are differences which can be associated with variations in the diet of the organism. The levels of ²¹⁰Pb in the krill Euphausia superba show seasonal changes which can be interpreted in terms of changing diet. In a simple classification of the pelagic food chain, the levels of ²¹⁰Pb tend to increase from omnivores to filter-feeders to predators.     

Polonium-210 in euphausiids: A detailed study

A detailed study of ²¹⁰Po, the predominant alpha-emitting nuclide found in most marine organisms, has been undertaken in a particular zooplanktonic species, the euphausiid Meganyctiphanes norvegica. The purpose was to obtain information concerning the origin, the localization and the flux of the nuclide in and through this organism. Measurements of ²¹⁰Po were made in euphausiids of different sizes, in dissected organs and tissues, and in excretion products. The results show higher concentrations in the smaller specimens; this fact cannot be explained on the basis of surface adsorption, but is probably related to the ingestion of food. Dissection results show that the distribution of ²¹⁰Po in euphausiids is not homogeneous, but that the majority is concentrated in the internal organs, the alimentary tract and the hepatopancreas in particular. The natural radiation dose received by these organs is in consequence much higher than that received by the whole animal. Use of a dynamic model allowed the flux of ²¹⁰Po through M. norvegica to be calculated. The calculations confirm that food is the principal source of ²¹⁰Po for this species, and clearly show that fecal pellets constitute the major elimination route. Extrapolation of the data to zooplankton in general leads to the conclusion that zooplankton metabolic activity plays an important role in transporting ²¹⁰Po from the surface layers of the ocean to depth.     

Polonium-210 and lead-210 in marine food chains

²¹⁰Po and ²¹⁰Pb have been measured systematically in whole animals, muscle and hepatopancreas of crustaceans and of molluscan cephalopods representative of a pelagic and benthic food chain. The same nuclides were also measured in liver, pyloric caecum, stomach contents and muscle of tuna. The concentration factors from sea water to whole animals were approximately constant along both food chains, being of the order of 10⁴ for ²¹⁰Po and 10² for ²¹⁰Pb. The highest concentration factors were found in shrimp of the genus Sergestes. In muscle, the concentration factors were an order of magnitude less; in the hepatopancreas, they were an order of magnitude higher, reaching 10⁶ in shrimp of the genus Sergestes. Such concentrations imply alpha-radiation doses of the order of 10 rem per year and more in this organ, which contains about 50 to 90% of the ²¹⁰Po in the whole animal in the 11 species analyzed. A detailed study of the intracellular behaviour of ²¹⁰Po in the hepatopancreas is clearly indicated. ²¹⁰Po can be used as a sensitive natural tracer in biological systems. Thus, feeding Meganyctiphanes norvegica in the laboratory on food low in ²¹⁰Po led to an approximate value of about 61/2 days for the biological half-life of ²¹⁰Po in the hepatopancreas of this euphausiid. Furthermore, the data on ²¹⁰Po and ²¹⁰Pb in the cephalopod hepatopancreas allowed the time of conservation of frozen squid which had been bought at the market to be estimated.     

210Po and 210Pb in zooplankton fecal pellets

²¹⁰Po and ²¹⁰Pb concentrations in fecal pellets from the zooplanktonic euphausiid Meganyctiphanes norvegica are reported. The ²¹⁰Po:²¹⁰Pb activity ratio is 2.2±0.3, a value in good agreement with that found in suspended particulate matter in surface seawater. Estimates of ²¹⁰Po and ²¹⁰Pb removal times from the mixed layer by fecal pellets alone yield values which are of the same order of magnitude as the removal times for these nuclides by all routes. It is suggested that there is a high probability that zooplanktonic fecal pellets play a significant role in the removal of both these nuclides from the surface layers of the ocean.     

Polonium-210 content of marine shrimp: Variation with biological and environmental factors

The concentrations of ²¹⁰Po and ²¹⁰Pb were determined in more than 30 species of marine shrimp. Samples were collected in the years 1977 to 1980; most were from the Mediterrancean Sea near Monaco, the remainder from Kuwait, South Africa, USA and Great Britain. The median ²¹⁰Po concentration was 8.0 pCi g^-1 dry wt in the whole shrimp and 85 pCi g^-1 dry in the hepatopancreas; the corresponding ²¹⁰Po:²¹⁰Pb activity ratios were 77 and 138, respectively. The range of concentrations covered more than two orders of magnitude for both nuclides. The variations in the levels of ²¹⁰Po, in particular, could be related to biological and environmental factors; thus, penaeid shrimp had ²¹⁰Po at median concentrations some three to four times higher than carid shrimp, and there was a steady increase in the median ²¹⁰Po concentration as one moved from estuarine to coastal to pelagic to deep-pelagic species. ²¹⁰Po levels were highest in 3 species of deep-pelagic penaeids, where a median ²¹⁰Po concentration in the whole shrimp of 43 pCi g^-1 dry wt was found. The natural radiation dose received by the hepatopancreas of such individuals will be of the order of 100 rem per yr. Elevated ²¹⁰Po was also found in 4 species collected near Kuwait. The ²¹⁰Po data are interpreted as indicating changes in the food regime of different shrimp; specifically it is suggested that ²¹⁰Po-rich items such as hepatopancreas, faecal pellets and organic particulates are likely to be important in the diet of shrimp which feed at depth in the ocean. It is suggested further that the ultimate availability of ²¹⁰Po to the foodchain depends on its form in sea water, and the need for information concerning the degree of organic binding of ²¹⁰Po in natural waters in stressed.     

Lipid metabolism of the Antarctic krill Euphausia superba and its ecological implications

 Various developmental stages (early larvae to adults) of Euphausia superba have been collected in different seasons in the Weddell Sea, the Lazarev Sea and off the Antarctic Peninsula to investigate the role of lipids and fatty acids in the life cycle of the Antarctic krill. The total-lipid data for E. superba exhibited seasonal variations, with low lipid levels in late winter/early spring and the highest levels in autumn. Seasonal changes were most pronounced in the immature and adult specimens, increasing from about 10% lipid of dry mass to more than 40%. The fatty-acid compositions of the younger stages were dominated by 20:5(n-3), 22:6(n-3) and 16:0. These are typical phospholipid fatty acids, which are major biomembrane constituents. The phospholipid composition was similar in the older stages. With increasing storage of triacylglycerols in the lipid-rich immature and adult stages, the fatty acids 14:0, 16:0 and 18:1(n-9) prevailed, comprising about 70% of total triacylglycerol fatty acids. The trophic-marker fatty acids 16:1(n-7) and 18:4(n-3), indicating phytoplankton ingestion, were less abundant. They reflected, however, the dependence of the larvae on phytoplankton as well as the seasonal changes in algal composition. The generally close linear relationships between fatty acids and lipid suggest that the fatty-acid compositions of the collected specimens were largely independent of the respective developmental stage, season and region. The linear fit indicates that triacylglycerol accumulation started at a level of about 5% of total lipid. Considering the various overwintering scenarios under discussion, the life cycle and reproductive strategies of krill are discussed in the context of the lipid metabolism and fatty-acid composition of E. superba. Lipid production is effective enough to accumulate large energy reserves for the dark season, but E. superba does not exhibit the sophisticated biosynthetic pathways known from other Antarctic euphausiids and copepods. Although important, lipid utilisation appears to be just one of several strategies of E. superba to thrive under the extreme Antarctic conditions, and this pronounced versatility may explain the success of this species in the Southern Ocean. Received: 16 June 2000 / Accepted: 18 December 2000     

210Po content in sea water and its accumulation in southern Baltic plankton

This paper presents the results of an investigation from 1980 to 1985 on the content of ²¹⁰Po in sea water and its accumulation in southern Baltic plankton. The mean value of the ²¹⁰Po concentration in sea water has been determined to be about 0.5 mBq dm^-3, with the proportion of the dissolved forms being ca 80%. It has been proved that ²¹⁰Po is accumulated by plankton organisms. The mean ²¹⁰Po concentration factors calculated on the basis of its content in plankton fall within the range from 5×10³ to 4.2×10⁴ and increase as follows: phytoplankton210Po concentration in mesozooplankton collected at the Slupsk Trough and the Gdask Basin have been observed. The above differences have been related to intense blue-green alga blooming in the Gdask Basin.     

Natural tracers in dietary studies: data for 210Po and 210Pb in decapod shrimp and other pelagic organisms in the Northeast Atlantic Ocean

Measurements of the concentrations of the naturally-occurring radioactive nuclides ²¹⁰Po and ²¹⁰Pb in a large number of pelagic organisms from the Atlantic Ocean made in the spring/summer of 1984 and 1985 are reported. Marine shrimp are particularly well represented, and in many cases data were obtained for shrimp hepatopancreas, stomach-contents, and posterior intestine plus contents, as well as for whole individuals. The wide ranges found in ²¹⁰Po concentrations group into categories: the shrimp, for example, divide into four categories, two penaeid and two carid. These groupings are explained on the basis of variations in diet between the different categories. A clear difference was also observed between the ²¹⁰Po levels in shrimp of similar species and size from opposite sides of an oceanographic front; this difference too can be explained plausibly in terms of a change in certain penaeid shrimp, and attention is drawn to the need for investigation of cytological and/or genetic effects which could possibly be caused by the very high natural radiation doses to which such organisms are exposed.     

Comparative lipid dynamics of euphausiids from the Antarctic and Northeast Pacific Oceans

To better understand the feeding and reproductive ecology of euphausiids (krill) in different ocean environments, lipid classes and individual lipid components of four different species of euphausiids from Northeast Pacific (temperate species) and Southern Ocean (Antarctic species) were analyzed in animals from multiple life stages and seasons. The dominant krill species in the Northeast Pacific Euphausia pacifica and Thysanoessa spinifera, were compared to the two major Antarctic species, Euphausia superba and E. crystallorophias. Analysis comprised total lipid and lipid classes together with individual fatty acid and sterol composition in adults, juveniles, and larvae. Antarctic krill had much higher lipid content than their temperate relatives (10–50 and 5–20% of dry mass for Antarctic and temperate species, respectively) with significant seasonal variations observed. Phospholipids were the dominant lipid class in both temperate krill species, while neutral storage lipids (wax esters and triacylglycerols for E. crystallorophias and E. superba, respectively) were the major lipid class in Antarctic krill and accounted for up to 40% of the total lipid content. Important fatty acids, specifically 16:0, 18:1ω9, 20:5ω3, and 22:6ω3, were detected in all four krill species, with minor differences between species and seasons. Detailed lipid profiles suggest that krill alter their lipid composition with life stage and season. In particular, larval Antarctic krill appear to utilize alternate food resources (i.e., sea-ice associated organisms) during austral winter in contrast to juveniles and adults (i.e., seston and copepods). Lipid dynamics in krill among krill in both systems appear closely linked to their life cycle and environmental conditions including food availability, and can provide a more complete comparative ecology of euphausiids in these environmentally distinct systems.     

Omnivorous feeding behavior of the Antarctic krill Euphausia superba

Feeding experiments were conducted at Palmer Station from December 1985 to February 1986 to examine the potential role of copepod prey as an alternative food source for Euphausia superba. Copepod concentration, copepod size, phytoplankton concentration, the duration of krill starvation and the volume of experimental vessels were altered to determine effects on ingestion and clearance rates. Krill allowed to feed on phytoplankton and copepods in 50-litre tubs showed greatly increased feeding rates relative to animals feeding in the much smaller volumes of water traditionally used for krill-feeding studies. Clearance rates on copepods remained constant over the range of concentrations offered, but clearance rates on phytoplankton increased linearly with phytoplankton concentration. Feeding rates increased when larger copepods were offered and when krill were starved for two weeks prior to experiments. Clearance rates of krill feeding on copepods were higher than, but not correlated with, their clearance rates on phytoplankton in the same vessel. E. superba may have a distinct mechanism for capturing copepods, perhaps through mechanoreception. Although our observed clearance rate of 1055 ml krill^-1 h^-1 indicates that krill can feed very efficiently on copepod prey, such feeding would meet less than 10% of their minimum metabolic requirements at the typical copepod concentrations reported for Antarctic waters. However, substantial energy could be gained if krill fed on the patches of high copepod concentrations occasionally reported during the austral summer, or if krill and copepods were concentrated beneath the sea ice during the winter or spring months. Our results, indicating efficient feeding on zooplankton and higher clearance rates on phytoplankton than previously believed, represent a step towards balancing the energy budget of E. superba in Antarctic waters.     

The fatty acids of antarctic phytoplankton and euphausiids. Fatty acid exchange among trophic levels of the Ross Sea

The fatty acids of 3 samples of Euphausia superba, 7 samples of E. crystallorophias, and 12 samples of phytoplankton collected in the Ross Sea, Antarctica, during Eltanin Cruise 51 were examined. The fatty acid profiles of the samples of E. superba resembled each other closely. The fatty acid profiles of the E. crystallorophias samples were also similar to each other but different quantitatively from those of E. superba. Phytoplankton fatty acid patterns varied with the geographical location and species composition of the samples. The fatty acids of euphausiids were compared to those of the phytoplankton from the corresponding locations. Rather similar fatty acid patterns in phytoplankton and E. superba corroborate the herbivorous nature of this euphausiid. On the other hand, phytoplankton and E. crystallorophias showed quite different fatty acid patterns. The differences were mostly due to the presence of waxes among the lipids of E. crystallorophias. It is not clear whether these waxes are of dietary origin or are synthesized endogenously.     

Assimilation of 210Po by the mussel Mytilus edulis from the alga Isochrysis galbana

Marine invertebrates are thought to accumulate ²¹⁰Po primarily from their food. In this study, a pulse-chase methodology was used to examine the assimilation and depuration of ²¹⁰Po by Mytilus edulis from the common marine alga Isochrysis galbana. The digestion of ²¹⁰Po from I. galbana occurred via a biphasic process, characteristic of a rapid (extracellular) and slow (intracellular) digestion typical of marine bivalves. The mantle/gill and foot have no known digestive role, yet their ²¹⁰Po specific activities increased after 24 h. It is proposed that this increase in ²¹⁰Po specific activity was related to ²¹⁰Po being incorporated into these tissues from ²¹⁰Po assimilated from I. galbana during extracellular digestion. It is proposed that the linear loss of ²¹⁰Po previously accumulated by control mussels was related to the continual state of renewal and replacement of cellular proteins, with ²¹⁰Po turnover and metabolism governed by protein turnover and metabolism. M. edulis' assimilation efficiency of ²¹⁰Po from the ²¹⁰Po-labelled alga was calculated to be 17.2 ± 2.1%, and thus similar to that of Ag, Cd, Co, Se and Zn by bivalves from other marine algae species. It is proposed that the assimilation efficiency of ²¹⁰Po is a function of protein assimilation. Received: 27 August 1998 / Accepted: 3 September 1999     

Effect of Cd on accumulation and loss of 210Po in the mussel Mytilus edulis

As a result of ²¹⁰Po's previously identified association with sulphur-rich proteins, metallothioneins could have a significant effect on the behaviour and fate of ²¹⁰Po in molluscs. Starved control and cadmium-exposed mussels, Mytilus edulis, were fed ²¹⁰Po-labelled algae (Isochrysis galbana) for 5 d and then allowed to depurate in clean sea water. Cadmium-exposed M. edulis accumulated less ²¹⁰Po in the digestive gland and the remainder of the tissue than control mussels, although this was due to a decrease in tissue weight. More than 40% of ²¹⁰Po was identified as being associated with high molecular weight and heat-treated cytosol proteins in M. edulis. Mussels in a starved state are known to recycle as much as 90% of their amino acids. It is proposed that ²¹⁰Po associated with these and other proteins is recycled, explaining why no significant loss of ²¹⁰Po was observed from the remainder of the tissue in either control or Cd-exposed mussels. Cadmium-induced metallothioneins had no effect on the distribution of ²¹⁰Po in M. edulis; <5% associated with the cytosolic fraction was considered to principally contain metallothioneins. It is suggested that ²¹⁰Po's apparent relationship with metallothioneins is coincidental rather than connected with its role in the regulation of metals. Received: 30 March 1998 / Accepted: 3 August 1998     

Krill evolution and the Antarctic ocean currents: evidence of vicariant speciation as inferred by molecular data

The phylogenetic relationships of the Antarctic krill Euphausia superba, the key species in the Antarctic food web, and other Antarctic and sub-Antarctic cuphausiids have been investigated using the 16S ribosomal mitochondrial gene. The phylogenetic reconstructions indicated that the Antarctic species form a monophyletic clade separated by the non-Antarctic species. The results revealed a large genetic divergence between the Antarctic (E. superba and E. crystallorophias) and sub-Antarctic species (E. vallentini). The time of separation between these species, estimated from the molecular data, is around 20 million years ago, which is comparable with the geological time of the formation of a circum-Antarctic water circulation and the Antarctic Polar Frontal Zone. The euphausiid molecular phylogeny therefore represents evidence for vicariant speciation.     

Laboratory and field estimates of the rate of faecal pellet production by Antarctic krill,Euphausia superba

The time course of faecal pellet production (egestion) was monitored in January 1985 for a population of Antarctic krill, Euphausia superba Dana, maintained in flowing seawater aquaria at Palmer Station, Antarctica. Following transfer to filtered seawater, krill produced faecal strings for roughly 40 min, after which time faecal egestion virtually ceased. Similar results were obtained for freshly-trawled krill at sea in February and March 1985. There were wide daily variations in total faecal egestion rate; mean rates varied from 0.54 to 1.66 mg dry wt h^-1 and individual rates from 0.25 to 2.35 mg h^-1 (all data corrected to a standard krill of 600 mg fresh weight). Despite these wide fluctuations in total faecal egestion, the loss of organic matter showed no significant daily variation, with a mean value of 0.13 mgh^-1. The relationship between faecal egestion rate and faecal organic content suggested that feeding rate was governed by food quality; when inorganic load was high, feeding rate increased to ensure sufficient energy intake. The data suggest that superfluous feeding does not occur in krill and that values of gut-clearance time calculated from time intervals greater than about 40 min will not be representative of previous feeding history. the rates of faecal egestion observed in this study indicate that the flux of faecal pellets from krill is substantial. They imply an energy intake in E. superba of 17 to 28% body weight per day, much higher than estimated previously for this species by summing known energy losses, but similar to estimates for other euphausiids.     

Timing of septal formation in two species of Nautilus based on radiometric and aquarium data

Growth rates of four Nautilus scrobiculatus Lightfoot and three N. pompilius Linnaeus captured in Papua New Guinea in 1985 were measured using a chronology based on the naturally occurring radionuclides ²¹⁰Pb and ²¹⁰Po. The age of the las septum in mature specimens of both species was more than one year, whereas the time of septal formation in a submature specimen of N. pompilius was ca 180 d. The ²¹⁰Po/²¹⁰Pb method was also used to determine the growth rates of four immature N. pompilius captured in the Philippines in 1983 before and after introduction into an aquarium. Introduction into the aquarium was marked by a dramatic decrease in the activity of ²¹⁰Pb of the septa, the formation of a black line on the exterior of the shell, and a reduction in the spacing of adjacent septa in two specimens. Based on the number of septa formed in the aquarium, the average time of septal formation ranged from 50 to 80 d. These estimates were consistent with those calculated from the ²¹⁰Po/²¹⁰Pb activity ratios of the septa that formed in the aquarium, thus providing a check on the radiometric method. The average time of septal formation just prior to introduction into the aquarium appears, within the uncertainty of the radioactive measurements, to have been similar to that in the aquarium, indicating no dramatic alteration in the growth rate of these specimens.     

Larval distribution and reproductive ecology of Thysanoessa macrura (Crustacea: Euphausiacea) in the Scotia Sea

In the Scotia Sea, the larvae of Thysanoessa macrura G.O. Sars, 1885 were scarce in spring (September 27 to October 6, 1971) and mainly present at the Calyptopis I stage, while in summer (December 17, 1974 to January 3, 1975) they were much more abundant and much further developed (up to Furcilia VI). The spawning season commences as early as September and, as indicated by the presence of numerous nauplii and metanauplii in summer, is rather long. T. macrura begins to spawn much earlier in West-Wind-Drift waters than in waters of the Weddell Sea. It starts to spawn simultaneously with Euphausia frigida and E. triacantha, but significantly earlier than E. superba. These timings of the breeding process are related to differential biogeographical characteristics of these species. Plankton collections made at depths of 500 or 1000 m have, for T. macrura, revealed a developmental ascent analogous to that of E. superba. Ontogenetical deep-water migrations are, consequently, characteristic not only for E. superba, but also for other Antarctic euphausiids (in particular E. frigida, but possibly not for inshore E. crystallorophias) and, as correctly postulated by several other investigators, for all oceanic euphausiid species.     

Contribution of heterotrophic material to diet and energy budget of Antarctic krill, Euphausia superba

A novel approach was used to estimate the heterotrophic carbon component in the diet of the Antarctic krill Euphausia superba. Over 200 specimens from seven samples collected in the Lazarev Sea (January 1993 and 1995), at the Antarctic Polar Front (January 1993), and around South Georgia (February/March 1994) were dissected, and the total carbon content of their stomachs was estimated with a CHN-analyser. Gut-pigment contents were also measured by the gut-fluorescence technique in specimens collected at the same time, and the equivalent amount of their gut carbon was then subtracted from the total organic carbon content of guts from the same samples. The remaining carbon was assumed to originate entirely from heterotrophic food sources. This heterotrophic component accounted for a substantial proportion of the total food consumed by Antarctic krill, ranging from 17.4 to 98.9% of the mass of the gut contents (mean = 78.8% ± 21.2 SD). The results make an important contribution to the elucidation of the energy budget of krill and its daily carbon ration. With a few exceptions, previous estimates were largely calculated from a solely autotrophic carbon source, and were unable to account for the metabolic requirements of E. superba. Krill plays an important role in Antarctic food webs, as it often constitutes ≃50% of the total biomass of the zooplankton, and produces fast-sinking, dense faecal pellets which are important in the vertical transport of organic carbon from the euphotic layer to the deep ocean. High consumption rates of smaller heterotrophic organisms by krill suggest that this large microphage may be more important than previously believed in re-packaging micro- and mesozooplankton into a longer-lasting and more easily sequestered carbon pool. Received: 26 October 1998 / Accepted: 14 October 1999     

Lead-210 and Beryllium-7 in the Aerosol Particles Around Taiwan Off-Shore Areas

Abstract

Distributions of ²¹⁰Pb and ⁷Be in the aerosol particles of different size fractions were measured around Taiwan offshore areas from October, 1994 to October, 1995. the size distribution and abundance of the aerosol particles showed both spatial and temporal variations. the particle concentrations off northwestern Taiwan were more than twice as those off northeastern Taiwan both in November and March, and the values were much higher in November than in March for both areas. the measured particle concentrations, except for one sample with an unusually high value, ranged from 10 to 83 μOgm ^?3 in the study areas. the ²¹⁰Pb concentrations in the aerosol particles filtered from the air mass varied between 20 × 10^?3 and 72 × 10^?3 dpm m^?3 (1dpm = 0.45 pCi). the ²¹⁰Pb concentrations in the area off southwestern Taiwan appeared somewhat higher than in other areas and were probably contributed by the land air from Taiwan which contains higher ²¹⁰Pb in winter. the ²¹⁰Pb concentrations off northern Taiwan were low in November when the aerosol concentrations were high, but they were high in March when the aerosol concentrations were low.

Based on the ⁷Be monitoring records of 20 years on the aerosol particles of the island by the Taiwan Radiation Monitoring Center, the mean settling rate averaged 0.91, 0.79 and 0.68 cm s^?1, respectively, in northern, central and southern Taiwan. with these values and the ²¹⁰Pb concentrations in the aerosols, the ²¹⁰Pb flux was determined to be between 0.58 and 2.30 dpm cm^?2y^?1, with an average of 1.19 dpm cm^?2y^?1. Excluding the highest value due to its extremely high ²¹⁰Pb and aerosol concentrations, the average is reduced to 1.05 dpm cm^?2y^?1. the activity median diameters (AMD) for ²¹⁰Pb in the study areas were between 0.69 and < 0.49 μM with a mean of 0.5 öm based on ²¹⁰Pb distributions in different particle size fractions. Thus, ²¹⁰Pb was preferentially adsorbed on to the submicron particles. ⁷Be in the study areas showed a good correlation with ²¹⁰Pb in total activity although the two nucides had different sources.