Antarctic Rockcod Research Articles

Description of Pseudobenedenia coriicepsi sp. n. (Monogenea: Capsalidae) from the Antarctic Black Rockcod, Notothenia coriiceps Richardson in Coastal Waters of West Antarctica Using Novel SEM Images, Energy Dispersive X-Ray Analysis and Molecular Analysis

Morphological and metrical analyses of sclerotized parts of haptor and body morphology and anatomy of monogeneans from Antarctic rockcod Notothenia coriiceps Richardson revealed the presence of a new species, Pseudobenedenia coriicepsi sp. n., supported with molecular studies, EDXA and scanning electron microscopy. New species differs from Pseudobenedenia nototheniae Johnston, 1931 by characteristic shape of anterior hamulus, size of sclerotized structures of haptor, measurements of the body, measurements ratios, v-shaped consistent commissure in vitellaria field, relative proportions of body and haptor, unlobed testes, and by host species. Published drawings of sclerotized parts of haptor and measurements of the species of the genus Pseudobenedenia Johnston, 1931 are provided for comparison and discussed. Status of P. nototheniae as species complex and host-specificity in this group are discussed. Molecular data of the P. coriicepsi sp. n. (partial 18S and 28S) were generated to provide the first molecular analysis for any species of Pseudobenedenia to date. Phylogenetic analyses inferred from the two molecular markers indicate that P. coriicepsi sp. n. is closely related to the species of the genus Benedenia and formed a sister clade to it as positioned under the family Capsalidae.

Helminth Assemblages of the Antarctic Black Rockcod, Notothenia coriiceps (Actinopterygii: Nototheniidae) in Coastal Waters near Galindez Island (Argentine Islands, West Antarctic): Temporal Changes in the Endoparasite Community.

Analysis and comparison of the helminth assemblages in Antarctic rockcod Notothenia coriiceps collected near the UAS "Akademik Vernadsky" (Argentine Islands, West Antarctica) in 2002 and 2014-2015 were performed to characterise the parasite community and investigate the temporal changes in helminth assemblages and infection parameters. All specimens of N. coriiceps (n = 194) were caught at depths of 10-30m. Parasites (22,856 helminth specimens and 15,057 cysts) were collected manually and identified based on their morphology. Statistical analysis of the quantitative data was performed using the Quantitative Parasitology 3.0 (QP 3.0), Paleontological Statistics (PAST v. 3.1), and PRIMER 6 software. Twenty-seven species of four taxonomic groups were recorded: trematodes (8 species), cestodes (4), nematodes (5), and acanthocephalans (10). Helminth samples collected in 2002 and 2014-2015 showed a rather high similarity in species composition. The species richness was higher in the sample collected in 2014-2015, while the evenness and diversity in the two samples were similar. The dissimilarity between helminth infracommunities in the two samples appeared to be statistically significant. Larval cestodes Diphyllobotrium sp., the acanthocephalan Metacanthocephalus rennicki, and the trematode Neoleoburia antarctica were found to make the most significant impact on the dissimilarity. The analysis of the composition and structure of helminth community in N. coriiceps revealed the changes that have happened during the last decade. At least some of the changes are attributed to the changes in marine ecosystems in Western Antarctica.

Helminths of antarctic rockcod Notothenia coriiceps (Perciformes, Nototheniidae) from the akademik Vernadsky station area (Argentine Islands, West Antarctica): new data on the parasite community

The aim of our study was to update the information on taxonomic diversity and structure of the parasite community of Antarctic rockcod, Notothenia coriiceps in the coastal water off the Argentine Islands (Wilhelm Archipelago, Graham Land, West Antarctica). The material for this study was collected in 2014–2015 at the Ukrainian Antarctic station “Akademik Vernadsky”. More than 8,500 specimens of parasites from 106 host specimens were collected and identified. All fishes were found to be infected with helminths; 25 helminth species were identified. Acanthocephalans were recorded in 93.4 % of hosts; eight species of acanthocephalans (Metacanthocephalus rennicki, M. johnstoni, M. campbelli, M. dalmori, Aspersentis megarhynchus, Corynosoma hammani, C. pseudohammani, and C. evae) were found. Nematodes were found in 96.2 % of fishes; larval stages of Pseudoterranova sp., Contraceacum sp., Anisakis sp. and adults Ascarophis nototheniae and Dichelyne fraseri were identified. Trematodes were found in 94.3 % of fishes; seven species (Macvicaria georgiana, Neolebouria antarctica, Lepidapedon garrardi, Genolinea bowersi, Elytrophalloides oatesi, Lecithaster macrocotyle, Derogenes johnstoni) were identified. Larval stages of cestodes (Diphyllobothrium sp. and tetraphyllidean metacestodes) and the monogenean species Pseudobenedenia nototheniae were found. Our studies revealed significant changes in the structure of the parasite community of N. coriiceps during the last decade.

Application of a quantitative histological health index for Antarctic rock cod (Trematomus bernacchii) from Davis Station, East Antarctica

A quantitative Histological Health Index (HHI) was applied to Antarctic rock cod (Trematomus bernacchii) using gill, liver, spleen, kidney and gonad to assess the impact of wastewater effluent from Davis Station, East Antarctica. A total of 120 fish were collected from 6 sites in the Prydz Bay region of East Antarctica at varying distances from the wastewater outfall. The HHI revealed a greater severity of alteration in fish at the wastewater outfall, which decreased stepwise with distance. Gill and liver displayed the greatest severity of alteration in fish occurring in close proximity to the wastewater outfall, showing severe and pronounced alteration respectively. Findings of the HHI add to a growing weight of evidence indicating that the current level of wastewater treatment at Davis Station is insufficient to prevent impact to the surrounding environment. The HHI for T. bernacchii developed in this study is recommended as a useful risk assessment tool for assessing in situ, sub-lethal impacts from station-derived contamination in coastal regions throughout Antarctica.

Novel flame retardants (N-FRs), polybrominated diphenyl ethers (PBDEs) and dioxin-like polychlorinated biphenyls (DL-PCBs) in fish, penguin, and skua from King George Island, Antarctica

Persistent organic pollutants (POPs), including polychlorinated biphenyls (PCBs) and polybrominated diphenyl ethers (PBDEs), are frequently detected in biota from Antarctica, whereas no data are available for their replacements, such as novel flame retardants (N-FRs). This study presented the occurrence of several N-FRs, PBDEs, and PCBs in tissue samples of an Antarctic rock cod (Trematomus bernacchii), a young gentoo penguin (Pygoscelis papua), and a brown skua (Stercorarius antarcticus) collected from King George Island. The total concentrations of N-FRs (ΣN-FRs; mean: 931pg/g dry weight (dw)) were comparable to PBDEs (Σ8PBDEs; 681pg/gdw), which were much lower than PCBs (ΣDL-PCBs; 12,800pg/gdw). Overall, skua contained two to three orders of magnitude higher contamination than penguin and fish. In the future, more attention should be focused on the fate of N-FRs in Antarctica, where usages have increased since PBDEs were banned. To our knowledge, this is the first report of N-FRs in biota from Antarctica.

Tracking spatial distribution of human-derived wastewater from Davis Station, East Antarctica, using δ15N and δ13C stable isotopes

Stable isotope ratios, δ15N and δ13C were effectively used to determine the geographical dispersion of human derived sewage from Davis Station, East Antarctica, using Antarctic rock cod (Trematomus bernacchii). Fish within 0–4km downstream of the outfall exhibited higher δ15N and δ13C values relative to reference sites. Nitrogen in particular showed a stepped decrease in δ15N with increasing distance from the discharge point by 1–2‰. Stable isotopes were better able to detect the extent of wastewater contamination than other techniques including faecal coliform and sterol measures. Uptake and assimilation of δ15N and δ13C up to 4km from the outfall adds to growing evidence indicating the current level of wastewater treatment at Davis Station is not sufficient to avoid impact to the surrounding environment. Isotopic assimilation in T. bernacchii is a viable biomarker for investigation of initial sewage exposure and longer term monitoring in the future.

Direct evidence of histopathological impacts of wastewater discharge on resident Antarctic fish (Trematomus bernacchii) at Davis Station, East Antarctica

During the 2009/2010 summer, a comprehensive environmental impact assessment (EIA) of the wastewater discharge at Davis Station, East Antarctica was completed. As part of this, histological alteration of gill and liver tissue in Antarctic Rock-cod (Trematomus bernacchii) from four sites along a spatial gradient from the wastewater outfall were assessed. All fish within 800m of the outfall exhibited significant histological changes in both tissues. Common pathologies observed in fish closest to the outfall include proliferation of epithelial cells with associated secondary lamellar fusion in the gills and multifocal granulomata with inflammation and necrosis as well as cysts in the liver. Fish from sites >800m from the outfall also exhibited alterations but to a lesser degree, with prevalence and severity decreasing with increasing distance from the outfall. This study highlights the value of histopathological investigations as part of EIAs and provides the first evidence of sub-lethal alteration associated with wastewater discharge in East Antarctica.

Interactions between trace metals (Cu, Hg, Ni, Pb) and 2,3,7,8‐tetrachlorodibenzo‐p‐dioxin in the antarctic fish Trematomus bernacchii: Oxidative effects on biotransformation pathway

Biological effects of chemical mixtures are an emerging issue when using biomarkers in field conditions, because synergistic or inhibitory interactions, cascade, and indirect mechanisms can both enhance or suppress responses to specific classes of pollutants. The Antarctic rock cod (Trematomus bernacchii) was exposed to various trace metals (Cu, Hg, Ni, and Pb) and to 2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD), dosed alone or in combination, to generate hypotheses regarding the mechanisms of action at the cellular level. Concentrations of metals significantly increased in liver of T. bernacchii without significant differences between organisms exposed only to the elements or cotreated with TCDD. The marked induction of cytochrome P450 (CYP) by TCDD was greatly suppressed by cotreatment with Cu at both catalytic and protein expression levels, whereas no significant effects were caused by coexposures with Hg, Ni, and Pb. The oxidative status was measured by integrating individual antioxidants with the total oxyradical scavenging capacity toward peroxyl radicals (ROO*) and hydroxyl radicals (*OH). Synergistic oxidative effects appeared in fish coexposed to Cu and TCDD, suggesting that biotransformation efficiency is down-regulated by higher levels of hydrogen peroxide, hydroxyl radicals, and/or mechanisms limiting availability of heme groups. Major oxidative perturbations also were observed in organisms coexposed to TCDD and Hg; in these organisms, however, the marked increase of glutathione content could explain the absence of oxidative inhibition of the CYP system. More limited oxidative effects were caused by Ni and Pb, and the contemporary absence of inhibitory effects on CYP would further confirm the importance of pro-oxidant mechanisms in modulating the biotransformation pathway. The overall results indicate that a wide battery of biomarkers is necessary to assess the impact of chemical mixtures in field conditions.

Purification and characterization of pepsins A1 and A2 from the Antarctic rock cod Trematomus bernacchii

The Antarctic notothenioid Trematomus bernacchii (rock cod) lives at a constant mean temperature of -1.9 degrees C. Gastric digestion under these conditions relies on the proteolytic activity of aspartic proteases such as pepsin. To understand the molecular mechanisms of Antarctic fish pepsins, T. bernacchii pepsins A1 and A2 were cloned, overexpressed in Escherichia coli, purified and characterized with a number of biochemical and biophysical methods. The properties of these two Antarctic isoenzymes were compared to those of porcine pepsin and found to be unique in a number of ways. Fish pepsins were found to be more temperature sensitive, generally less active at lower pH and more sensitive to inhibition by pepstatin than their mesophilic counterparts. The specificity of Antarctic fish pepsins was similar but not identical to that of pig pepsin, probably owing to changes in the sequence of fish enzymes near the active site. Gene duplication of Antarctic rock cod pepsins is the likely mechanism for adaptation to the harsh temperature environment in which these enzymes must function.

Interactions between metabolism of trace metals and xenobiotic agonists of the aryl hydrocarbon receptor in the antarctic fish Trematomus bernacchii: Environmental perspectives

Although Antarctica is a pristine environment, organisms are challenged with contaminants either released locally or transported from industrialized regions through atmospheric circulation and marine food webs. Organisms from Terra Nova Bay also are exposed to a natural enrichment of cadmium, but to our knowledge, whether such environmental conditions influence biological responses to anthropogenic pollutants has never been considered. In the present study, the Antarctic rock cod (Trematomus bernacchii) was exposed to model chemicals, including polycyclic aromatic hydrocarbons (benzo[a]pyrene), persistent organic pollutants (2,3,7,8-tetrachlorodibenzo-p-dioxin [TCDD]), cadmium, and a combination of cadmium and TCDD. Analyzed parameters included chemical bioaccumulation, activity, and levels of biotransformation enzymes (cytochrome P4501A); metallothioneins and the efficiency of the antioxidant system measured as individual defenses (catalase, glutathione, glutathione reductase, glutathione S-transferases, and glutathione peroxidases); and total scavenging capacity toward peroxyl and hydroxyl radicals. Reciprocal interactions between metabolism of inorganic and organic pollutants were demonstrated. Dioxin enhanced the accumulation of cadmium, probably stored within proliferating endoplasmic reticulum, and cadmium suppressed the inducibility of cytochrome P4501A, allowing us to hypothesize a posttranscriptional mechanism as the depletion of heme group availability. Clear evidence of oxidative perturbation was provided by the inhibition of antioxidants and enhanced sensitivity to oxyradical toxicity in fish exposed to organic chemicals. Exposure to cadmium revealed counteracting responses of glutathione metabolism; however, these responses did not prevent a certain loss of antioxidant capacity toward peroxyl radicals. The pattern of antioxidant responses exhibited by fish coexposed to cadmium and TCDD was more similar to that observed for cadmium than to that observed for TCDD. The overall results suggest that elevated natural levels of cadmium in Antarctic organisms from Terra Nova Bay can limit biotransformation capability of polycyclic (halogenated) hydrocarbons, thus influencing the bioaccumulation and biological effects of these chemicals in key sentinel species.

Bloodthirsty, an RBCC/TRIM gene required for erythropoiesis in zebrafish

The Antarctic icefishes (family Channichthyidae, suborder Notothenioidei) constitute the only vertebrate taxon that fails to produce red blood cells. These fishes can be paired with closely related, but erythrocyte-producing, notothenioids to discover erythropoietic genes via representational difference analysis. Using a B30.2-domain-encoding DNA probe so derived from the hematopoietic kidney (pronephros) of a red-blooded Antarctic rockcod, Notothenia coriiceps, we discovered a related, novel gene, bloodthirsty ( bty), that encoded a 547-residue protein that contains sequential RING finger, B Box, coiled-coil, and B30.2 domains. bty mRNA was expressed by the pronephric kidney of N. coriiceps at a steady-state level 10-fold greater than that found in the kidney of the icefish Chaenocephalus aceratus. To test the function of bty, we cloned the orthologous zebrafish gene from a kidney cDNA library. Whole-mount in situ hybridization of zebrafish embryos showed that bty mRNA was present throughout development and, after the mid-blastula transition, was expressed in the head and in or near the site of primitive erythropoiesis in the tail just prior to red cell production. One- to four-cell embryos injected with two distinct antisense morpholino oligonucleotides (MOs) targeted to the 5′-end of the bty mRNA failed to develop red cells, whereas embryos injected with 4- and 5-bp mismatch control MOs produced wild-type quantities of erythrocytes. The morphant phenotype was rescued by co-injection of synthetic bty mRNA containing an artificial 5′-untranslated region (UTR) with the antisense MO that bound the 5′-UTR of the wild-type bty transcript. Furthermore, the expression of genes that mark terminal erythroid differentiation was greatly reduced in the antisense-MO-treated embryos. We conclude that bty is likely to play a role in differentiation of the committed red cell progenitor.

Posttranslational modification of brain tubulins from the Antarctic fish Notothenia coriiceps: reduced C-terminal glutamylation correlates with efficient microtubule assembly at low temperature.

We have shown previously that the tubulins of Antarctic fish assemble into microtubules efficiently at low temperatures (-2 to +2 degrees C) due to adaptations intrinsic to the tubulin subunits. To determine whether changes in posttranslational glutamylation of the fish tubulins may contribute to cold adaptation of microtubule assembly, we have characterized C-terminal peptides from alpha- and beta-tubulin chains from brains of adult specimens of the Antarctic rockcod Notothenia coriiceps by MALDI-TOF mass spectrometry and by Edman degradation amino acid sequencing. Of the four fish beta-tubulin isotypes, nonglutamylated isoforms were more abundant than glutamylated isoforms. In addition, maximal glutamyl side-chain length was shorter than that observed for mammalian brain beta tubulins. For the nine fish alpha-tubulin isotypes, nonglutamylated isoforms were also generally more abundant than glutamylated isoforms. When glutamylated, however, the maximal side-chain lengths of the fish alpha tubulins were generally longer than those of adult rat brain alpha chains. Thus, Antarctic fish adult brain tubulins are glutamylated differently than mammalian brain tubulins, resulting in a more heterogeneous population of alpha isoforms and a reduction in the number of beta isoforms. By contrast, neonatal rat brain tubulin possesses low levels of glutamylation that are similar to that of the adult fish brain tubulins. We suggest that unique residue substitutions in the primary structures of Antarctic fish tubulin isotypes and quantitative changes in isoform glutamylation act synergistically to adapt microtubule assembly to low temperatures.

Adaptive Evolution of Gene Expression in Antarctic Fishes: Divergent Transcription of the 5′-to-5′ Linked Adult α1- and β-Globin Genes of the Antarctic TeleostNotothenia coriicepsis Controlled by Dual Promoters and Intergenic Enhancers1

Unlike temperate fishes, Antarctic fishes of the notothenioid suborder, whose body temperatures (−2 to 1°C) conform to the Southern Ocean, must express their genomes in an extremely cold thermal regime. To determine whether these fishes have evolved compensatory adjustments that maintain efficient gene transcription at low temperatures, we have initiated studies of the cis-acting regulatory elements that control globin gene expression in the Antarctic rockcod Notothenia coriiceps and in its close relative, the temperate New Zealand black cod N. angustata (habitat temperature = 6 to 15°C). The genes encoding the major α1 and β globins of these fishes are tightly linked in head-to-head (5′ to 5′) orientation. The intergenic regions separating the globin genes in the two fishes, ∼4.3 kb in N. coriiceps and ∼3.2 kb in N. angustata, are highly similar in sequence, the major difference being the absence of a 1.1-kb, repeat-containing segment in the latter. To assess the promoter and enhancer activities of the intergenic regions, each was cloned into the luciferase-reporter vector pGL3-Basic, and the constructs were transfected into MEL cells. Upon DMSO induction of MEL cell differentiation, each of the α/β-intergenic regions functioned in both orientations as erythroid-responsive transcriptional regulators. However, expression of luciferase mediated by the N. coriiceps intergene was 6-fold greater in the α orientation than that for the N. angustata intergene and 2-fold greater for the β. The greater transcription-stimulating activity of the N. coriiceps intergene can be attributed to two enhancers composed of combinations of CAC/Sp1 and GATA motifs and located in direct repeat elements. N. angustata, which lacked repetitive structure in its intergene, contained a single copy of the enhancer. We propose that cold adaptation of globin gene expression in N. coriiceps evolved in part through duplication and refinement of critical cis-acting regulatory elements as the Southern Ocean cooled during the past 25 million years.

Adaptive Evolution of Gene Expression in Antarctic Fishes: Divergent Transcription of the 5′-to-5′ Linked Adult α1- and β-Globin Genes of the Antarctic TeleostNotothenia coriicepsis Controlled by Dual Promoters and Intergenic Enhancers

Unlike temperate fishes, Antarctic fishes of the notothenioid suborder, whose body temperatures (−2 to +1°C) conform to the Southern Ocean, must express their genomes in an extremely cold thermal regime. To determine whether these fishes have evolved compensatory adjustments that maintain efficient gene transcription at low temperatures, we have initiated studies of the cis-acting regulatory elements that control globin gene expression in the Antarctic rockcod Notothenia coriiceps and in its close relative, the temperate New Zealand black cod N. angustata (habitat temperature = +6 to +15°C). The genes encoding the major α1 and β globins of these fishes are tightly linked in head-to-head (5′ to 5′) orientation. The intergenic regions separating the globin genes in the two fishes, ∼4.3 kb in N. coriiceps and ∼3.2 kb in N. angustata, are highly similar in sequence, the major difference being the absence of a 1.1-kb, repeat-containing segment in the latter. To assess the promoter and enhancer activities of the intergenic regions, each was cloned into the luciferase-reporter vector pGL3-Basic, and the constructs were transfected into MEL cells. Upon DMSO induction of MEL cell differentiation, each of the α/β-intergenic regions functioned in both orientations as erythroid-responsive transcriptional regulators. However, expression of luciferase mediated by the N. coriiceps intergene was 6-fold greater in the α orientation than that for the N. angustata intergene and 2-fold greater for the β. The greater transcription-stimulating activity of the N. coriiceps intergene can be attributed to two enhancers composed of combinations of CAC/Sp1 and GATA motifs and located in direct repeat elements. N. angustata, which lacked repetitive structure in its intergene, contained a single copy of the enhancer. We propose that cold adaptation of globin gene expression in N. coriiceps evolved in part through duplication and refinement of critical cis-acting regulatory elements as the Southern Ocean cooled during the past 25 million years.

Identification of betaIII- and betaIV-tubulin isotypes in cold-adapted microtubules from Atlantic cod (Gadus morhua): antibody mapping and cDNA sequencing.

Isolated microtubule proteins from the Atlantic cod (Gadus morhua) assemble at temperatures between 8 and 30 degrees C. The cold-adaptation is an intrinsic property of the tubulin molecules, but the reason for it is unknown. To increase our knowledge of tubulin diversity and its role in cold-adaptation we have further characterized cod tubulins using alpha- and beta-tubulin site-directed antibodies and antibodies towards posttranslationally modified tubulin. In addition, one cod brain beta-tubulin isotype has been sequenced. In mammals there are five beta-tubulins (betaI, betaII, betaIII, betaIVa and betaIVb) expressed in brain. A cod betaIII-tubulin was identified by its electrophoretic mobility after reduction and carboxymethylation. The betaIII-like tubulin accounted for more than 30% of total brain beta-tubulins, the highest yield yet observed in any animal. This tubulin corresponds most probably with an additional band, designated beta(x), which was found between alpha- and beta-tubulins on SDS-polyacrylamide gels. It was found to be phosphorylated and neurospecific, and constituted about 30% of total cod beta-tubulin isoforms. The sequenced cod tubulin was identified as a betaIV-tubulin, and a betaIV-isotype was stained by a C-terminal specific antibody. The amount of staining indicates that this isotype, as in mammals, only accounts for a minor part of the total brain beta-tubulin. Based on the estimated amounts of betaIII- and betaIV-tubulins in cod brain, our results indicate that cod has at least one additional beta-tubulin isotype and that beta-tubulin diversity evolved early during fish evolution. The sequenced cod betaIV-tubulin had four unique amino acid substitutions when compared to beta-tubulin sequences from other animals, while one substitution was in common with Antarctic rockcod beta-tubulin. Residues 221, Thr to Ser, and 283, Ala to Ser, correspond in the bovine tubulin dimer structure to loops that most probably interact with other tubulin molecules within the microtubule, and might contribute to cold-adaptation of microtubules.

Muscle power output during escape responses in an Antarctic fish

Escape responses (C-shaped fast-starts) were filmed at 500 frames s-1 in the Antarctic rock cod (Notothenia coriiceps) at 0 °C. The activation and strain patterns of the superficial fast myotomal muscle were measured simultaneously using electromyography and sonomicrometry respectively. In order to bend the body into the initial C-shape, the muscle fibres in the rostral myotomes (at 0.35L, where L is total length) shortened by up to 13 % of their resting length at a maximum velocity of 1.68 fibre lengths s-1. During the contralateral contraction, muscle fibres were stretched (by 5 % and 7 % at 0.35L and 0.65L, respectively) and were activated prior to the end of lengthening, before shortening by up to 12 % of resting fibre length (peak-to-peak strain). Representative strain records were digitised to create cyclical events corresponding to the C-bend and contralateral contraction. Isolated fibres were subjected to the abstracted strain cycles and stimulated at the same point and for the same duration as occurs in vivo. During the early phase of shortening, muscle shortening velocity (V) increased dramatically whilst the load was relatively constant and represented a substantial fraction of the maximum isometric stress. Pre-stretch of active muscle was associated with significant force enhancement. For the contralateral contraction, V exceeded that predicted by the steady-state force­velocity relationship for considerable periods during each tailbeat, contributing to relatively high maximum instantaneous power outputs of up to 290 W kg-1 wet muscle mass. In vitro experiments, involving adjusting strain, cycle duration and stimulation parameters, indicated that in vivo muscle fibres produce close to their maximum power. During escape responses, the maximum velocity and acceleration recorded from the centre of gravity of the fish were 0.71±0.03 m s-1 and 17.1±1.4 m s-2, respectively (mean ± s.e.m., N=7 fish). Muscle performance was sufficient to produce maximum velocities and accelerations that were within the lower end of the range reported for temperate-zone fish.