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• W2005126983 abstract "The daily vertical migration of pelagic animals to and from the upper ocean layers is the most widespread and coordinated movement of biomass on Earth. The behaviour is a major pathway for carbon to reach the ocean's interior as organisms eat at the surface and release their waste at depth. Estimates of carbon sequestration via this ‘active flux’ of pelagic animals assume that the behaviour is performed just once every 24 hours [1Longhurst A.R. Bedo A.W. Harrison W.G. Head E.J.H. Sameoto D.D. Vertical flux of respiratory carbon by oceanic diel migrant biota.Deep Sea Res. A. 1990; 37: 685-694Crossref Scopus (203) Google Scholar], but there is indirect evidence that there may be a continuous movement between the upper and lower layers, with individuals ascending when hungry and descending once satiated [2Frost B.W. Variability and possible adaptive significance of diel vertical migration in Calanus pacificus, a planktonic marine copepod.Bull. Mar. Sci. 1988; 43: 675-694Google Scholar, 3Pearre S. Vertical migration and feeding in Sagitta elegans (Verrill).Ecology. 1973; 54: 300-314Crossref Google Scholar]. We found that Antarctic krill, one of the most abundant pelagic animals found in a region known to be an important carbon sink [4Schlitzer R. Carbon export fluxes in the Southern Ocean: results from inverse modeling and comparison with satellite-based estimates.Deep-sea Res. II. 2002; 49: 1623-1644Crossref Scopus (226) Google Scholar], exhibited a sinking response when replete that could contribute significantly to carbon sequestration. Many animals perform daily vertical migration, indicating that it confers selective advantages. The behaviour minimises visibility to predators whilst satisfying the need to exploit the food-rich surface layers. However, daily vertical migration patterns are not always up at night and down in the day, because there is evidence that many populations divide into deep and shallow layers at the same time [2Frost B.W. Variability and possible adaptive significance of diel vertical migration in Calanus pacificus, a planktonic marine copepod.Bull. Mar. Sci. 1988; 43: 675-694Google Scholar]. Individuals found in deeper layers can contain food types that could only have been eaten at the surface [3Pearre S. Vertical migration and feeding in Sagitta elegans (Verrill).Ecology. 1973; 54: 300-314Crossref Google Scholar]. This suggests a pattern of swimming to the surface, sinking when full and migrating up again when digestion is complete. Such behaviour has yet to be directly observed in individual pelagic animals. Amongst pelagic animals, Antarctic krill has a considerable biomass and profoundly influences the ecology of the Southern Ocean. Krill remain pelagic for their entire life cycle with 73% of their total daily metabolism being used to fuel swimming [5Kils U. Swimming behaviour, swimming performance and energy balance of Antarctic krill Euphausia superba.BIOMASS Sci. Ser. 1981; 3: 1-121Google Scholar]. These animals must overcome their negative buoyancy through the continuous generation of upward thrust from specialised swimming legs on the abdomen, called pleopods. Krill sink immediately when inactive, although the rate of descent may be controlled by fanning out the pleopods in a horizontal plane to form a parachute [6Kils U. Marschall H.-P. Antarctic Krill (Euphausia superba) feeding and swimming performance: New insights with new methods.in: Hempel I. Hempel G. Biologie der Polarmeere. Fischer-Jena, Stuttgart1995: 201-210Google Scholar] (Figure 1). To measure krill swimming characteristics in laboratory experiments, it is necessary to use some sort of tethering device to prevent them from hitting container walls. Such studies have found that gender, size and moult-stage affect maximum power output [7Thomasson M.A. Johnson M.L. Stromberg J.O. Gaten E. Swimming capacity and pleopod beat rate as a function of sex, size and moult stage in Northern krill Meganyctiphanes norvegica.Mar. Ecol. Prog. Ser. 2003; 250: 205-213Crossref Scopus (27) Google Scholar]. The use of free-falling cameras in situ have revealed that krill adopt a variety of swimming patterns, from the continuous beating of pleopods to regular switching between beating and parachuting [6Kils U. Marschall H.-P. Antarctic Krill (Euphausia superba) feeding and swimming performance: New insights with new methods.in: Hempel I. Hempel G. Biologie der Polarmeere. Fischer-Jena, Stuttgart1995: 201-210Google Scholar]. No studies of krill have yet been able to relate swimming behaviours to internal states, such as levels of hunger and satiation. We examined swimming activity patterns in 113 krill through tethering them to a MLT0015 rotational displacement transducer (see Supplemental Experimental Procedures in the Supplemental Data published online). Measurements were made during a research campaign aboard the RRS James Clark Ross (December 2004–January 2005) in the vicinity of South Georgia (54.5°S, 37°W). Krill caught with a pelagic net (RMT8) were placed in incubation facilities continuously supplied with water from the sea surface. This meant that food availability depended on whether or not the ship had recently passed through any food (phytoplankton) patches. Accordingly, krill used for swimming experiments contained stomachs that ranged from full to empty. Krill exhibited continuous swimming, cyclic swimming (alternating phases of strong and weak pleopod strokes) and regular switching between swimming and parachuting during our observations (∼20 min per krill). Most exhibited just one or two of these behavioural modes, although around 10% exhibited all three. The swim–parachute mode was exhibited for around 27% of total experimental time. We found, however, that the level of stomach fullness affected its prevalence. Krill with full to half-full stomachs spent a significantly greater amount of time in the swim–parachute mode than those with empty stomachs (Mann-Whitney U=1940.5, N1=73, N2= 40, P=0.042; Figure 2). Hungry krill were more likely to beat their pleopods continuously. Krill in swim–parachute mode sink at a rate of between 0.2 cm s−1 and 0.8 cm s−1 (see Supplemental Data). In the 90min it would take a krill to digest its stomach contents [8Atkinson A. Snyder R. Krill–copepod interactions at South Georgia, Antarctica, I. Omnivory by Euphausia superba.Mar. Ecol. Prog. Ser. 1997; 160: 63-76Crossref Scopus (129) Google Scholar], it would descend between 9 m and 43 m. Krill mainly feed in the mid to lower reaches of the surface mixed layer [9Godlewska M. Vertical migrations of krill (Euphausia superba Dana).Pol. Arch. Hydrobiol. 1996; 43: 9-63Google Scholar], which is usually 40 m deep in the Southern Ocean [10Meredith M.P. Brandon M.A. Murphy E.J. Trathan P.N. Thorpe S.E. Bone D.G. Chernyshkov P.P. Sushin V.A. Variability in hydrographic conditions to the east and northwest of South Georgia, 1996–2001.J. Mar. Sys. 2005; 53: 143-167Crossref Scopus (46) Google Scholar]. Satiation-induced descent would result in krill migrating below this layer, so releasing faeces into the ocean’s interior where it becomes sequestered. During the productive summer period, it is possible that individuals go through the cycle of feeding, sinking and re-ascending three times each night. We estimate that krill in the Southern Ocean sequester 2.3×1013 grams carbon each year (see Supplemental Data) through releasing faeces below the mixed layer, adding 8% to previous estimates of global active flux [1Longhurst A.R. Bedo A.W. Harrison W.G. Head E.J.H. Sameoto D.D. Vertical flux of respiratory carbon by oceanic diel migrant biota.Deep Sea Res. A. 1990; 37: 685-694Crossref Scopus (203) Google Scholar]. Egestion via hunger–satiation-driven vertical migration has not been considered by other studies of active flux. This study highlights the need to examine the prevalence of this behaviour worldwide. Download .pdf (.05 MB) Help with pdf files Document S1. Supplemental Experimental Procedures and Supplemental Analysis" @default.
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• W2005126983 title "Satiation gives krill that sinking feeling" @default.
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