Krill body size drives particulate organic carbon export in West Antarctica.

The export of carbon from the ocean surface and storage in the ocean interior is important in the modulation of global climate1-4. The West Antarctic Peninsula experiences some of the largest summer particulate organic carbon (POC) export rates, and one of the fastest warming rates, in the world5,6. To understand how warming may alter carbon storage, it is necessary to first determine the patterns and ecological drivers of POC export7,8. Here we show that Antarctic krill (Euphausia superba) body size and life-history cycle, as opposed to their overall biomass or regional environmental factors, exert the dominant control on the POC flux. We measured POC fluxes over 21 years, the longest record in the Southern Ocean, and found a significant 5-year periodicity in the annual POC flux, which oscillated in synchrony with krill body size, peaking when the krill population was composed predominately of large individuals. Krill body size alters the POC flux through the production and export of size-varying faecal pellets9, which dominate the total flux. Decreases in winter sea ice10, an essential habitat for krill, are causing shifts in the krill population11, which may alter these export patterns of faecal pellets, leading to changes in ocean carbon storage.

Relationship of morphometrics, total carotenoids, and total lipids with activity and sexual and spatial features in Euphausia superba.

Morphological differences associated with sex or stage, together with total lipids and carotenoids, were studied in Euphausia superba as possible indicators of physiological condition. E. superba displays sexual dimorphism during growth. A group of mature males, called Males II herein, has a greater abdominal length, suggesting that they are faster swimmers, a feature implying higher metabolic rates and a higher demand for protecting pigments like carotenoids. Mature Males II have proportionally lower lipids but higher total lipid-soluble carotenoids, a counterintuitive finding. Males II also have bigger eyes. Significant regressions with carotenoids were found for wet weight, abdominal length, and eye diameter. On a spatial analysis, population composition reflects reproductive activity. Males II would be in search of females for fecundation and, thus, are dominant in some areas. The PCA analysis of 10 allometric and biochemical variables show a distinct Males II group differing in morphology, carotenoids, and lipid contents. The carotenoid:lipid ratio was highest for Males II, supporting the hypothesis of the role of carotenoids in the activity of the species. Mature males may experience physiological stress during reproduction and probably die shortly afterwards. A relationship between activity, morphometrics, and carotenoid content seems evident, deserving further investigation.

Quantifying the Escape Mortality of Trawl Caught Antarctic Krill (Euphausia superba).

Antarctic krill (Euphausia superba) is an abundant fishery resource, the harvest levels of which are expected to increase. However, many of the length classes of krill can escape through commonly used commercial trawl mesh sizes. A vital component of the overall management of a fishery is to estimate the total fishing mortality and quantify the mortality rate of individuals that escape from fishing gear. The methods for determining fishing mortality in krill are still poorly developed. We used a covered codend sampling technique followed by onboard observations made in holding tanks to monitor mortality rates of escaped krill. Haul duration, hydrological conditions, maximum fishing depth and catch composition all had no significant effect on mortality of krill escaping 16 mm mesh size nets, nor was any further mortality associated with the holding tank conditions. A non- parametric Kaplan-Meier analysis was used to model the relationship between mortality rates of escapees and time. There was a weak tendency, though not significant, for smaller individuals to suffer higher mortality than larger individuals. The mortality of krill escaping the trawl nets in our study was 4.4 ± 4.4%, suggesting that krill are fairly tolerant of the capture-and-escape process in trawls.

Naupliar and Metanaupliar Development of Thysanoessa raschii (Malacostraca, Euphausiacea) from Godthåbsfjord, Greenland, with a Reinstatement of the Ancestral Status of the Free-Living Nauplius in Malacostracan Evolution.

The presence of a characteristic crustacean larval type, the nauplius, in many crustacean taxa has often been considered one of the few uniting characters of the Crustacea. Within Malacostraca, the largest crustacean group, nauplii are only present in two taxa, Euphauciacea (krill) and Decapoda Dendrobranchiata. The presence of nauplii in these two taxa has traditionally been considered a retained primitive characteristic, but free-living nauplii have also been suggested to have reappeared a couple of times from direct developing ancestors during malacostracan evolution. Based on a re-study of Thysanoessa raschii (Euphausiacea) using preserved material collected in Greenland, we readdress this important controversy in crustacean evolution, and, in the process, redescribe the naupliar and metanaupliar development of T. raschii. In contrast to most previous studies of euphausiid development, we recognize three (not two) naupliar (= ortho-naupliar) stages (N1-N3) followed by a metanauplius (MN). While there are many morphological changes between nauplius 1 and 2 (e.g., appearance of long caudal setae), the changes between nauplius 2 and 3 are few but distinct. They involve the size of some caudal spines (largest in N3) and the setation of the antennal endopod (an extra seta in N3). A wider comparison between free-living nauplii of both Malacostraca and non-Malacostraca revealed similarities between nauplii in many taxa both at the general level (e.g., the gradual development and number of appendages) and at the more detailed level (e.g., unclear segmentation of naupliar appendages, caudal setation, presence of frontal filaments). We recognize these similarities as homologies and therefore suggest that free-living nauplii were part of the ancestral malacostracan type of development. The derived morphology (e.g., lack of feeding structures, no fully formed gut, high content of yolk) of both euphausiid and dendrobranchiate nauplii is evidently related to their non-feeding (lecithotrophic) status.

The application of optical coherence tomography to image subsurface tissue structure of Antarctic krill Euphausia superba.

Many small open ocean animals, such as Antarctic krill, are an important part of marine ecosystems. To discover what will happen to animals such as krill in a changing ocean, experiments are run in aquaria where conditions can be controlled to simulate water characteristics predicted to occur in the future. The response of individual animals to changing water conditions can be hard to observe, and with current observation techniques it is very difficult to follow the progress of an individual animal through its life. Optical coherence tomography (OCT) is an optical imaging technique that allows images at high resolution to be obtained from depths up to a few millimeters inside biological specimens. It is compatible with in vivo imaging and can be used repeatedly on the same specimens. In this work, we show how OCT may be applied to post mortem krill samples and how important physiological data such as shell thickness and estimates of organ volume can be obtained. Using OCT we find an average value for the thickness of krill exoskeleton to be (30±4) µm along a 1 cm length of the animal body. We also show that the technique may be used to provide detailed imagery of the internal structure of a pleopod joint and provide an estimate for the heart volume of (0.73±0.03) mm3.

Comparative morphology of the hemolymph vascular system in krill (Euphausiacea; Crustacea).

The phylogenetic position of Euphausiacea within Malacostraca is still under debate. Either they are seen as sister group to a taxon comprising Pancarida and Peracarida or closer related to Decapoda. Both hypotheses can be supported by characters of the circulatory system. Therefore, a comparative re-evaluation of the circulatory system seems to be feasible. Here we present the first three-dimensional data of the circulatory system of three euphausiacean species based on semi-thin sections and micro computer tomography in combination with corrosion casting. We were also able to study for the first time representatives of Bentheuphausia amblyops, the suggested sister taxon to all other euphausiaceans. The main pumping structure in the open circulatory system of Euphausiacea is the globular heart in the rear thoracic segments. From the anterior and posterior end of the heart two unpaired and four pairs of arteries emanate. The unpaired anterior aorta runs below the carapace from the anterior part of the heart into the anterior cephalothorax, where it supplies the first antennae, the brain and the eyes. The paired posterior aortae run into the pleon supplying the pleopods, uropods and the telson. The four pairs of cardiac arteries supply appendages in the cephalic region and viscera in the trunk. The unpaired descending artery connects to the subneural vessel supplying the thoracopods. A myoarterial formation of the anterior aorta is described in Bentheuphausia amblyops. The observed pattern of a globular heart situated in the posterior cephalothorax and comprising a meshwork of muscular strands (also running through its lumen) is highly likely to be homologous in Euphausiacea and Decapoda. The data are compared with the scarce literature present to date.

Material properties of euphausiids and other zooplankton from the Bering Sea.

Acoustic assessment of Bering Sea euphausiids and their predators can provide useful data for ecosystem studies if the acoustic scattering characteristics of these animals are known. The amount of acoustic energy that is scattered by different marine zooplankton taxa is strongly affected by the contrast of the animal's density (g) and sound speed (h) with the surrounding seawater. Density and sound speed contrast were measured in the Bering Sea during the summer of 2008 for several different zooplankton and nekton taxa including: euphausiids (Thysanoessa inermis, Thysanoessa raschii, and Thysanoessa spinifera), copepods, amphipods, chaetognaths, gastropods, fish larvae, jellyfish, and squid. Density contrast values varied between different taxa as well as between individual animals within the same species. Sound speed contrast was measured for monospecific groups of animals and differences were found among taxa. The range, mean, and standard deviation of g and h for all euphausiid species were: g = 1.001-1.041; 1.018 ± 0.009 and h = 0.990-1.017; 1.006 ± 0.008. Changes in the relationship between euphausiid material properties and animal length, seawater temperature, seawater density, and geographic location were also evaluated. Results suggest that environmental conditions at different sample locations led to significant differences in animal density and material properties.

Coordination of multiple appendages in drag-based swimming.

Krill are aquatic crustaceans that engage in long distance migrations, either vertically in the water column or horizontally for 10 km (over 200,000 body lengths) per day. Hence efficient locomotory performance is crucial for their survival. We study the swimming kinematics of krill using a combination of experiment and analysis. We quantify the propulsor kinematics for tethered and freely swimming krill in experiments, and find kinematics that are very nearly metachronal. We then formulate a drag coefficient model which compares metachronal, synchronous and intermediate motions for a freely swimming body with two legs. With fixed leg velocity amplitude, metachronal kinematics give the highest average body speed for both linear and quadratic drag laws. The same result holds for five legs with the quadratic drag law. When metachronal kinematics is perturbed towards synchronous kinematics, an analysis shows that the velocity increase on the power stroke is outweighed by the velocity decrease on the recovery stroke. With fixed time-averaged work done by the legs, metachronal kinematics again gives the highest average body speed, although the advantage over synchronous kinematics is reduced.

Systematic sequencing of mRNA from the Antarctic krill (Euphausia superba) and first tissue specific transcriptional signature.

BACKGROUND: Little is known about the genome sequences of Euphausiacea (krill) although these crustaceans are abundant components of the pelagic ecosystems in all oceans and used for aquaculture and pharmaceutical industry. This study reports the results of an expressed sequence tag (EST) sequencing project from different tissues of Euphausia superba (the Antarctic krill). RESULTS: We have constructed and sequenced five cDNA libraries from different Antarctic krill tissues: head, abdomen, thoracopods and photophores. We have identified 1.770 high-quality ESTs which were assembled into 216 overlapping clusters and 801 singletons resulting in a total of 1.017 non-redundant sequences. Quantitative RT-PCR analysis was performed to quantify and validate the expression levels of ten genes presenting different EST countings in krill tissues. In addition, bioinformatic screening of the non-redundant E. superba sequences identified 69 microsatellite containing ESTs. Clusters, consensuses and related similarity and gene ontology searches were organized in a dedicated E. superba database http://krill.cribi.unipd.it. CONCLUSION: We defined the first tissue transcriptional signatures of E. superba based on functional categorization among the examined tissues. The analyses of annotated transcripts showed a higher similarity with genes from insects with respect to Malacostraca possibly as an effect of the limited number of Malacostraca sequences in the public databases. Our catalogue provides for the first time a genomic tool to investigate the biology of the Antarctic krill.

Computational determination of refractive index distribution in the crystalline cones of the compound eye of Antarctic krill (Euphausia superba).

In order to understand how a compound eye channels light to the retina and forms an image, one needs to know the refractive index distribution in the crystalline cones. Direct measurements of the refractive indices require sections of fresh, unfixed tissue and the use of an interference microscope, but frequently neither is available. Using the eye of the Antarctic krill Euphausia superba (the main food of baleen whales) we developed a computational method to predict a likely refractive index distribution non-invasively from sections of fixed material without the need of an interference microscope. We used a computer model of the eye and calculated the most realistic spatial distribution of the refractive index gradient in the crystalline cone that would enable the eye to produce a sharp image on the retina. The animals are known to see well and on the basis of our computations we predict that for the eyes of the adult a maximum refractive index of 1.45-1.50 in the centre of the cone yields a better angular sensitivity and light absorption in a target receptor of the retina than if N(max) were 1.55. In juveniles with a narrower spatial separation between dioptric structures and retina, however, an N(max) of 1.50-1.55 gives a superior result. Our method to determine the most likely refractive index distribution in the cone without the need of fresh material and an interference microscope could be useful in the study of other invertebrate eyes that are known to possess good resolving power, but for a variety of reasons are not suitable for or will not permit direct refractive index measurements of their dioptric tissues to be taken.