Working with living krill – the people and the places

The fascination of Antarctic scientists with Antarctic krill and their capabilities has a long and varied history, and prompted many scientists to maintain and manipulate krill under laboratory conditions. Starting in the Discovery era with Mackintosh at the King Edward Point labs on South Georgia, 1930, scientists have collected krill from sailing vessels, small boats, inflatable zodiacs and large ice-breaking vessels. Krill have been maintained in small and large jars, deep rectangular tanks, large round tanks and in flow-through and recycling systems. They have been maintained both on board research vessels and in laboratories, in flowing seawater systems at ambient conditions and in temperature-controlled environmental rooms. A few researchers have transported living krill back to their home laboratories, for example tropical laboratories in Japan (Murano) and Australia (Ikeda), temperate laboratories (Nicol) in Australia, a northern European laboratory in Germany (Marschall) and a sunny maritime laboratory in California (Ross and Quetin). The goals have been varied: short-term experiments to understand in situ physiological rates, long-term experiments to test the effects of manipulations or controlled changes in environmental conditions, and behavioral responses. We take you on a brief historical tour as we trace the lineage of modern day research on living Antarctic krill.     

Krill maintenance and experimentation at the australian antarctic division

Live Antarctic krill, Euphausia superba, have been maintained for experimental purposes at the Australian Antarctic Division since 1981. This population has been replenished on an annual basis with animals taken from the wild. Techniques used to capture and maintain live krill are discussed here, with particular reference given to the development of systems for their maintenance. Details are also provided for specific experimental systems that have been used to conduct research into the behaviour and physiology of krill both at-sea and in shore-based laboratories.     

Krill maintenance and experimentation at the australian antarctic division

Live Antarctic krill, Euphausia superba, have been maintained for experimental purposes at the Australian Antarctic Division since 1981. This population has been replenished on an annual basis with animals taken from the wild. Techniques used to capture and maintain live krill are discussed here, with particular reference given to the development of systems for their maintenance. Details are also provided for specific experimental systems that have been used to conduct research into the behaviour and physiology of krill both at-sea and in shore-based laboratories.     

A photographic documentation of the development of Antarctic krill (Euphausia superba) from egg to early juvenile

Antarctic krill (Euphausia superba) is a key species in Antarctic marine ecosystems, as well as an important species in the Southern Ocean fishery. Here, we provide the first detailed photographic documentation of embryonic and larval development of Antarctic krill over a 5-month developmental period under controlled laboratory conditions. Developing embryos and larvae were photographed every 3 h and every 5 days, respectively. Our results indicated a developmental time of approximately 6 days for embryos and 138 days for larvae (0.5 °C). This study provided baseline biometry information for future investigations of Antarctic krill development under changing environmental conditions.     

British biological research in the Antarctic

A tradition of biological research in the Antarctic was established by Cook 200 years ago. This tradition has been built on by other British expeditions, notably the 'Discovery' Investigations. The British Antarctic Survey, which arose from Operation Tabarin and the Falkland Islands Dependencies Survey, now carries out a programme of coordinated and continuous biological research. The Atlantic sector of the Antarctic, in which the Survey operates, is of key importance biologically. The Antarctic provides a striking biological contrast between a species-poor and very barren terrestrial ecosystem and the species-rich and productive ocean which surrounds it. Severe climatic conditions and great isolation (a contrast to the Arctic) characterize the Antarctic environment. Work at the Survey's biological research stations is designed to study the distribution and interactions of organisms and communities, how they have adapted to Antarctic conditions, and which by their abundance may be deemed successful. Research is done into terrestrial, fresh-water and marine systems. Additionally, there is a major research programme into the biologv, environment and principal predators of krill, Euphausia superba. The Antarctic is a laboratory where opportunities exist for natural experiments to test theories and elucidate basic biological problems.     

Protection of the antarctic ecosystem: A study in international environmental law

Abstract

The Antarctic region constitutes a fragile eco‐system closely related to the unique features of the physical environment of that continent. The Antarctic Ocean is central to the region's living systems, with krill as the ecological basis of life in the ocean and on land. So far, man's impact upon the Antarctic environment has been negligible, but there is concern that overharvesting of krill and possible marine pollution resulting from any future offshore oil exploration may undermine the krill basis of the Antarctic ecosystem. The legal status of Antarctica is largely determined by the 1959 Antarctic Treaty, and especially by the inner circle of the currently fourteen “consultative”; status signatories. These states have given high priority to ecological considerations by enacting a series of environmental and conservationist regulations, as well as two conventions, one protecting the Antarctic seals and the other marine living resources in general. Environmental regulations will form an important part of the legal regime for the exploration and exploitation of the Antarctic mineral resources, primarily oil and gas. The Antarctic Treaty framework does not legally bind nonsignatory states, but under general international law all states are bound to refrain from inflicting damage upon the planet's environment. Also, some marine pollution conventions apply to the Antarctic waters, and the 1982 U.N. Convention on the Law of the Sea comprehensively covers the protection and preservation of the marine environment of all oceans and seas. The preservation of the Antarctic environment will remain a high priority irrespective of what legal regime will govern Antarctica after 1991 when the Antarctic Treaty may, and most probably will, be subject to review.     

Laboratory observations of moulting,growth and maturation in Antarctic krill (Euphausia superba Dana)

Summary Observations of intermoult period, growth and maturation were made on krill which were transported from Antarctic waters and maintained in the laboratory in Australia over a three year period. The mean intermoult period (IP) for each of 10 specimens, with initial body lengths of 24.7=46.8 mm, kept at -0.5° C varied from 22.0 to 29.8 days (overall mean = 26.6 days). These measurements of IP are significantly longer than those obtained in some previous studies. Differences in experimental temperatures, light, body sizes and growth patterns of the specimens between studies are unlikely to be causes of these dissimilar results. The pattern of changes in body length (BL) varies from one individual to the next. The greatest increase in BL over a series of 4–5 moults ranged from 0.024 to 0.070 mm/day, which is equivalent to 0.0020 to 0.0086/day in body weight, assuming exponential growth. This maximum growth rate is about half the rate predicted from the growth scheme of Mauchline (1980) for wild krill. Comparison of growth data for other euphausiids suggests that Mauchline's scheme produces anomalous growth rate. The slower growth rate observed in the present study would extend the estimated life span of krill from 3–4 years, as calculated by Mauchline (1980), to 4–7 years. If krill undergo body shrinkage during the Antarctic winter the estimated life span might be even longer. Examination of the external sexual characters of moults showed both progression and regression of maturity stage in association with changes in BL.     

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.     

Relative changes in krill abundance inferred from Antarctic fur seal

Antarctic krill Euphausia superba is a predominant species in the Southern Ocean, it is very sensitive to climate change, and it supports large stocks of fishes, seabirds, seals and whales in Antarctic marine ecosystems. Modern krill stocks have been estimated directly by net hauls and acoustic surveys; the historical krill density especially the long-term one in the Southern Ocean, however, is unknown. Here we inferred the relative krill population changes along the West Antarctic Peninsula (WAP) over the 20th century from the trophic level change of Antarctic fur seal Arctocephalus gazella using stable carbon (δ(13)C) and nitrogen (δ(15)N) isotopes of archival seal hairs. Since Antarctic fur seals feed preferentially on krill, the variation of δ(15)N in seal hair indicates a change in the proportion of krill in the seal's diets and thus the krill availability in local seawater. For the past century, enriching fur seal δ(15)N values indicated decreasing krill availability. This is agreement with direct observation for the past ～30 years and suggests that the recently documented decline in krill populations began in the early parts of the 20th century. This novel method makes it possible to infer past krill population changes from ancient tissues of krill predators.     

Environmental response of upper trophic-level predators reveals a system change in an Antarctic marine ecosystem

Long-term changes in the physical environment in the Antarctic Peninsula region have significant potential for affecting populations of Antarctic krill (Euphausia superba), a keystone food web species. In order to investigate this, we analysed data on krill-eating predators at South Georgia from 1980 to 2000. Indices of population size and reproductive performance showed declines in all species and an increase in the frequency of years of low reproductive output. Changes in the population structure of krill and its relationship with reproductive performance suggested that the biomass of krill within the largest size class was sufficient to support predator demand in the 1980s but not in the 1990s. We suggest that the effects of underlying changes in the system on the krill population structure have been amplified by predator-induced mortality, resulting in breeding predators now regularly operating close to the limit of krill availability. Understanding how krill demography is affected by changes in physical environmental factors and by predator consumption and how, in turn, this influences predator performance and survival, is one of the keys to predicting future change in Antarctic marine ecosystems.     

The krill maturity cycle: a conceptual model of the seasonal cycle in Antarctic krill

A long-term study on the maturity cycle of Antarctic krill was conducted in a research aquarium. Antarctic krill were either kept individually or in groups for 8 months under different temperature and food conditions, and the succession of female maturity stages and intermoult periods were observed. In all cases regression and re-maturation of external sexual characteristics were observed, but there were differences in length of the cycle and intermoult periods between the experimental conditions. Based on these results, and information available from previous studies, we suggest a conceptual model describing seasonal cycle of krill physiology which provides a framework for future studies and highlight the importance of its link to the timings of the environmental conditions.     

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.     

Dietary segregation of krill-eating South Georgia seabirds

The diets of six of the main seabird species (two petrels, two albatrosses, two penguins) breeding at Bird Island, South Georgia were studied simultaneously during the chick-rearing period in 1986. For five species, Antarctic krill Euphausia superba was the main food (39–98% by mass); grey-headed albatrosses took mainly the ommastrephid squid Martialia hyadesi (71%) and only 16% krill. The size of the krill taken was similar between seabird species, although there were small but significant differences between penguins and the other species. Sex and reproductive status of krill, however, was different between all seabird species, reflecting some combination of differences in foraging ranges, selectivity by predators, or differences in escape responses of krill. For the krill-eating species, the rest of the diet varied substantially between species, comprising Martialia and nototheniid fish (blackbrowed albatross and, along with lanternfish, white-chinned petrel), lanternfish and amphipods (Antarctic prion and macaroni penguin), and icefish (gentoo penguin). Long-term data on breeding success and information on diet in 5–10 other years suggest that in 1986 seabird diet and reproductive performance was indicative of a year of good availability of krill around South Georgia. In such circumstances, ecological segregation between krill-eating species appears to be maintained chiefly by differences in foraging range and feeding methods, which are reviewed. This situation is rather different from the few studies of seabird communities elsewhere, where prey type and size are believed to be the main mechanisms of dietary segregation.     

Antarctic krill population genomics: apparent panmixia,but genome complexity and large population size muddy the water

Antarctic krill (Euphausia superba; hereafter krill) are an incredibly abundant pelagic crustacean which has a wide, but patchy, distribution in the Southern Ocean. Several studies have examined the potential for population genetic structuring in krill, but DNA‐based analyses have focused on a limited number of markers and have covered only part of their circum‐Antarctic range. We used mitochondrial DNA and restriction site‐associated DNA sequencing (RAD‐seq) to investigate genetic differences between krill from five sites, including two from East Antarctica. Our mtDNA results show no discernible genetic structuring between sites separated by thousands of kilometres, which is consistent with previous studies. Using standard RAD‐seq methodology, we obtained over a billion sequences from >140 krill, and thousands of variable nucleotides were identified at hundreds of loci. However, downstream analysis found that markers with sufficient coverage were primarily from multicopy genomic regions. Careful examination of these data highlights the complexity of the RAD‐seq approach in organisms with very large genomes. To characterize the multicopy markers, we recorded sequence counts from variable nucleotide sites rather than the derived genotypes; we also examined a small number of manually curated genotypes. Although these analyses effectively fingerprinted individuals, and uncovered a minor laboratory batch effect, no population structuring was observed. Overall, our results are consistent with panmixia of krill throughout their distribution. This result may indicate ongoing gene flow. However, krill's enormous population size creates substantial panmictic inertia, so genetic differentiation may not occur on an ecologically relevant timescale even if demographically separate populations exist.     

Mesozooplankton biomass and indices of grazing and metabolic activity in Antarctic waters

Mesozooplankton abundance, body area spectrum, biomass, gut fluorescence and electron transfer system (ETS) activity were studied in the Antarctic Peninsula during the post-bloom scenario in these waters. Values of abundance and biomass were rather low and decreased sharply from the slope waters to the coastal area. In contrast, specific gut fluorescence and ETS activity were high in the coastal area and decreased through the shelf-break. Large copepods were very scarce, similarly to the post-bloom conditions in phytoplankton where large cells are not abundant and small cells such as flagellates dominate the water column. The vertical distribution showed two well defined layers by day, one at the surface which corresponded to krill organisms and a second at depth (>300 m) formed mainly by the large copepod Metridia gerlachei. During the short night, this layer ascended at the time that krill at the surface migrated to deeper waters as observed from acoustics and net sampling. This observation and the absence of large copepods over the shelf suggest that krill consumption on large phytoplankton cells during the bloom is followed by an increase in predation upon mesozooplankton. It also suggests that krill decrease the abundance and biomass of mesozooplankton over the shelf and continues their predation upon mesopelagic copepods during the post-bloom in Antarctic waters. This behaviour could explain the long ago described impoverishment in mesozooplankton south of the Antarctic Circumpolar Current.     

Histopathology of Antarctic krill, Euphausia superba, bearing black spots

Small black spots have been noticed on the cephalothorax of Antarctic krill, Euphausia superba, since January, 2001. To study the nature of the black spots, the krill were sampled in the winter of 2003, 2006, and 2007 in the South Georgia region, the Antarctic Ocean. Histological observations revealed that the black spots were melanized nodules that were composed of hemocytes surrounding either bacteria or amorphous material. In the 2007 samples, 42% of the krill had melanized nodules. Most of the nodules had an opening on the body surface of the krill. A single melanized nodule often contained more than one type of morphologically distinct bacterial cell. Three bacteria were isolated from these black spots, and classified into either Psychrobacter or Pseudoalteromonas based on the sequences of 16S rRNA genes. More than three bacterial species or strains were also confirmed by in situ hybridization for 16S rRNA. The melanized nodules were almost always accompanied by a mass of atypical, large heteromorphic cells, which were not observed in apparently healthy krill. Unidentified parasites were observed in some of the krill that had melanized nodules. These parasites were directly surrounded by the large heteromorphic cells. Histological observations suggested that these heteromorphic cells were attacking the parasites. These results suggest the possibility that the krill had been initially affected by parasite infections, and the parasitized spots were secondary infected by environmental bacteria after the parasites had escaped from the host body.     

Modelling individual growth of the Antarctic krill Euphausia superba Dana

Summary Growth of the Antarctic krill, Euphausia superba, is not easily determined from net catches nor from laboratory experiments. Therefore, in support of these methods, a phenomenological model was constructed which in its present state describes the growth of a single krill specimen under periodically limiting food conditions with summer seasons of variable lengths. Published data of krill body length vs. age and of the annual cycle of primary production of algae in the Drake Passage were used to formulate equations and to calculate growth curves. At 1,000 days after hatching, the model predicts a body length of 63 mm, growth being delayed by 380 days compared with constant, optimal feeding conditions. Final length, weight and time delay are related to the amount of food supplied and compared with published population growth curves.     

Energetic demands of larval krill, Euphausia superba, in winter

Metabolic rates of larval and juvenile krill, Euphausia superba, were measured on board ship during three winter cruises west of the Antarctic Peninsula (June-July 1987, June 1993, and June 1994), and also under different temperature regimes and feeding conditions during long-term maintenance in the laboratory (Palmer Station, winter 1993). A mean oxygen consumption and nitrogen excretion ratio of 31.1 measured on board ship at ambient ocean temperatures suggested that larval and juvenile krill from ice-covered waters were primarily herbivorous. Results from both shipboard and laboratory experiments demonstrated that oxygen consumption increased with temperature, but that larvae subjected to acute temperature increases exhibited higher rates. Experiments conducted at near ambient water temperatures for winter were also conducted to test the effect of habitat on the energy requirements of larval and juvenile krill. A comparison of the field and laboratory studies conducted at −1.5 to −1.8 °C showed that larvae from ice-covered waters and fed larvae in the laboratory had oxygen consumption rates significantly higher than those of larvae collected from open, i.e. ice-free, water and those starved in the laboratory. Results of the comparison lend support to the concept that in winter, larval and juvenile krill are better fed in ice-covered waters than in open water, and to the hypothesis that ice biota in the pack ice are an important food resource in winter for larval and juvenile krill.     

2018年夏秋季南设得兰群岛周边水域南极磷虾集群类型及其影响因素

南极磷虾是一种典型的集群性海洋生物,其集群特征为行为生态学研究领域的重要内容之一。南极磷虾在南设得兰群岛周围高度密集分布,然而磷虾集群形状和大小的机制解释仍存在较大的争议。基于南设得兰群岛周边水域收集的Simrad EK80声学数据,本研究利用Echoview V6.16软件,对声学数据进行了分析,并对磷虾集群特征进行了划分。通过主坐标分析(PCoA)检验了环境因素(表温和海况)以及时空因素对各类型磷虾集群产生的影响。结果表明:海况对磷虾集群影响较大,光照强度次之;块状小型集群的时空分布较广,夜间与白天均占有较高比例(>30%);小型集群更易出现在白天,而大型集群则更多出现在深夜; 2月,磷虾集群与海况及纬度显著相关; 3月,集群与时段显著相关; 4月,集群与时段及海况显著相关。