International audienceAntarctic krill Euphausia superba (hereafter `krill') occur in regions undergoing rapid environmental change, particularly loss of winter sea ice. During recent years, harvesting of krill has in creased, possibly enhancing stress on krill and Antarctic ecosystems. Here we review the overall impact of climate change on krill and Antarctic ecosystems, discuss implications for an ecosystem-based fisheries management approach and identify critical knowledge gaps. Sea ice decline, ocean warming and other environmental stressors act in concert to modify the abundance, distribution and life cycle of krill. Although some of these changes can have positive effects on krill, their cumulative impact is most likely negative. Recruitment, driven largely by the winter survival of larval krill, is probably the population parameter most susceptible to climate change. Predicting changes to krill populations is urgent, because they will seriously impact Antarctic ecosystems. Such predictions, however, are complicated by an intense inter-annual variability in recruitment success and krill abundance. To improve the responsiveness of the ecosystem-based management approach adopted by the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR), critical knowledge gaps need to be filled. In addition to a better understanding of the factors influencing recruitment, management will require a better understanding of the resilience and the genetic plasticity of krill life stages, and a quantitative understanding of under-ice and benthic habitat use. Current precautionary management measures of CCAMLR should be maintained until a better understanding of these processes has been achieved. [GRAPHICS]

No abstract

Considering that swim-flume or chasing methods fail in the estimation of maximum metabolic rate and in the estimation of Aerobic Scope (AS) of sedentary or sluggish aquatic ectotherms, we propose a novel conceptual approach in which high metabolic rates can be obtained through stimulation of organism metabolic activity using high and low non-lethal temperatures that induce high (HMR) and low metabolic rates (LMR), This method was defined as TIMR: Temperature Induced Metabolic Rate, designed to obtain an aerobic power budget based on temperature-induced metabolic scope which may mirror thermal metabolic scope (TMS = HMR—LMR). Prior to use, the researcher should know the critical thermal maximum (CT max) and minimum (CT min) of animals, and calculate temperature TIMR max (at temperatures −5–10% below CT max) and TIMR min (at temperatures +5–10% above CT min), or choose a high and low non-lethal temperature that provoke a higher and lower metabolic rate than observed in routine conditions. Two sets of experiments were carried out. The first compared swim-flume open respirometry and the TIMR protocol using Centropomus undecimalis (snook), an endurance swimmer, acclimated at different temperatures. Results showed that independent of the method used and of the magnitude of the metabolic response, a similar relationship between maximum metabolic budget and acclimation temperature was observed, demonstrating that the TIMR method allows the identification of TMS. The second evaluated the effect of acclimation temperature in snook, semi-sedentary yellow tail (Ocyurus chrysurus), and sedentary clownfish (Amphiprion ocellaris), using TIMR and the chasing method. Both methods produced similar maximum metabolic rates in snook and yellowtail fish, but strong differences became visible in clownfish. In clownfish, the TIMR method led to a significantly higher TMS than the chasing method indicating that chasing may not fully exploit the aerobic power budget in sedentary species. Thus, the TIMR method provides an alternative way to estimate the difference between high and low metabolic activity under different acclimation conditions that, although not equivalent to AS may allow the standardized estimation of TMS that is relevant for sedentary species where measurement of AS via maximal swimming is inappropriate.

Vertical distribution and abundance of three numerically dominant krill species (Nyctiphanes simplex, Nematoscelis difficilis, and Euphausia eximia) were surveyed in the Gulf of California to understand the role of oxidative stress in their daily vertical migration (DVM) and zoogeographic patterns. Superoxide radical production, lipid peroxidation, and antioxidant enzyme activities were analyzed from krill collected with stratified nets from the surface down to 200 m during January, July, and October 2007. The upper boundary of the oxygen minimum zone (OMZ) was significantly shallower during October than during January. N. simplex was always distributed above the hypoxic layers, mostly in coastal upwelling areas. Ne. difficilis and E. eximia were relatively abundant during January, but detected mostly during their ascending migration. N. simplex was the most sensitive species to high temperatures and low oxygen concentrations, showing evidence of oxidative stress during summer (100 times more lipid peroxidation and 30 times more antioxidant enzyme activities than in winter). Ne. difficilis had higher glutathione peroxidase activity than N. simplex, which could facilitate its larger DVM. Low abundance of Ne. difficilis at 100 m during summer suggests that high temperature was also an environmental limiting factor. Oxidative stress indicators could explain the absence of N. simplex and Ne. difficilis in the eastern tropical Pacific and the ability of E. eximia to live in the OMZ and the eastern tropical Pacific. The latter had higher superoxide radical production and smaller lipid peroxidation during October. This suggests that E. eximia antioxidant enzyme activities are enough to avoid oxidative damage when exposed to hypoxic conditions during DVM.

a b s t r a c tVertical and horizontal distributions of the subtropical euphausiid juvenile and adult Nyctiphanes simplex were mapped from samples collected during winter and summer 2007 in the Gulf of California, Mexico. During winter, wide-ranging high densities occurred in most of the Gulf of California. Densities decreased considerably during summer, with only at few locations having high densities. N. simplex made short daily vertical migrations of o 150 m with pronounced vertical seasonal ontogenetic segregation. In winter, most life stages, particularly gravid and ovigerous females, were found in the top 100 m at night, with temperatures o 17 1C throughout the normal, cold, well-mixed water column. During summer, a seasonal thermocline at $ 50 m formed with temperatures between 22-29 1C above the thermocline. Reproductive females and males were found below the thermocline at 450 m, clearly avoiding layers with temperatures 4 20 1C. In both seasons, N. simplex occurred above the low-oxygen layer ( o 1.5 ml l À 1 ), which occurred at 150 m during winter and 90 m during summer. In summer, this layer extended farther north and into shallower water columns than during winter. The low-oxygen layer acts as the bottom limit of vertical distribution and horizontal distribution is limited at the southern part of the gulf to temperatures 423 1C. Seasonal brood size and reproductive effort were estimated for both sides of the Baja California Peninsula under ship board experiments as a proxy of the relative effect of seasonal environmental conditions for euphausiid reproduction. Experiments were done during March, July, and December 2004 at the entrance to Bahía Magdalena and its westward continental shelf and in November 2005 and January and July 2007 in the Gulf of California. Contrary to broadcast-spawning euphausiids, N. simplex, a sac-spawning euphausiid, has a significant association of the brood size as a function of the total length of females. N. simplex produces an average brood of 52 eggs female -1 (range 5-116 eggs female -1 ) with a estimated total fecundity of 936 eggs female -1 in a life span (360-1337 eggs female -1

Sexual maturation and reproduction influence the status of a number of physiological processes and consequently the ecology and behaviour of cephalopods. Using Octopus mimus as a study model, the present work was focused in the changes in biochemical compound and activity that take place during gonadal maturation of females and its consequences in embryo and hatchlings characteristics. To do that, a total of 31 adult females of O. mimus were sampled to follow metabolites (ovaries and digestive gland) and digestive enzyme activities (alkaline and acidic proteases) during physiological and functional maturation. Levels of protein (Prot), triacylglyceride (TG), cholesterol (Chol), glucose (Glu), and glycogen (Gly) were evaluated. Groups of eggs coming from mature females were also sampled along development and after hatching (paralarvae of 1 and 3 days old) to track metabolites (Prot, TG, Glu, Gly, TG, Chol), digestive enzymes activity (Lipase, alkaline proteases, and acidic proteases), and antioxidant/detoxification defence indicators with embryos development. Based on the data obtained, we hypothesized that immature females store Chol in their ovaries, probably from the food they ingested, but switch to TG reserves at the beginning of the maturation processes. At the same time, results suggest that these processes were energetically supported by Glu, obtained probably from Gly breakdown by gluconeogenic pathways. Also, was observed that embryos metabolites and enzyme activities (digestive and antioxidant/detoxification enzymes) where maintained without significant changes and in a low activity during the whole organogenesis, meaning that organogenesis is relatively not energetically costly. In contrast, after organogenesis, a mobilization of nutrients and activation of the metabolic and digestive enzymes was observed, together with increments in consumption of yolk and Gly, and reduction in lipid peroxidation. Derived from our results, we also have the hypothesis that reactive oxygen species (ROS) were produced during the metabolic processes that occurs in ovarian maturation. Those ROS may be in part transferred to the egg provoking a ROS charge to the embryos. The elimination of ROS in embryos started when the activity of the heart and the absorption of the yolk around stages XIV and XV were evident. Altogether, these processes allowed the paralarvae to hatch with buffered levels of ROS and with the antioxidant defence mechanisms ready to support further ROS production derived from paralarvae higher life stage requirements (feeding and metabolic demands).

To understand the adaptation of euphausiid (krill) species to oxygen minimum zones (OMZs), respiratory response and stress experiments combining hypoxia‐reoxygenation exposure with warming were conducted. Experimental krill species were obtained from the Antarctic (South Georgia area), the Humboldt Current System (HCS, Chilean coast) and the Northern California Current System (NCCS, Oregon). Euphausia mucronata from the HCS showed oxyconforming pO2‐dependent respiration below 80% air saturation (18 kPa). Normoxic subsurface oxygenation in winter posed a ‘high oxygen stress’ for this species. The NCCS krill, Euphausia pacifica, and the Antarctic krill, Euphausia superba, maintained respiration rates constant down to low critical pO2 values of 6 kPa (30% air saturation) and 11 kPa (55% air saturation), respectively. Antarctic krill had low antioxidant enzyme activities, but high concentrations of the molecular antioxidant glutathione (GSH) and was not lethally affected by 6 h exposure to moderate hypoxia. The temperate krill species (E. pacifica) had higher superoxide dismutase (SOD) values in winter than in summer, which relates to a higher winter metabolic rate. In all species, antioxidant enzyme activities remained constant during hypoxic exposure at the typical temperature for their habitat. Warming by 7 °C above habitat temperature in summer increased SOD activities and GSH levels in E. mucronata (HCS), but no oxidative damage occurred. In winter, when the NCCS is well mixed and the OMZ is deeper, +4 °C of warming combined with hypoxia represents a lethal condition for E. pacifica. In summer, when the OMZ expands upwards (100 m subsurface), antioxidant defences counteracted hypoxia and reoxygenation effects in E. pacifica, but only at mildly elevated temperature (+2 °C). In this season, experimental warming by +4 °C reduced antioxidant activities and the combination of warming with hypoxia again caused mortality of exposed specimens. We conclude that a climate change scenario combining warming and hypoxia represents a serious threat to E. pacifica and, as a consequence, NCCS food webs.

Histophagous ciliates of the genus Pseudocollinia cause epizootic events that kill adult female krill (Euphausiacea), but their mode of transmission is unknown. We compared 16S rRNA sequences of bacterial strains isolated from stomachs of healthy krill Nyctiphanes simplex specimens with sequences of bacterial isolates and sequences of natural bacterial communities from the hemocoel of N. simplex specimens infected with P. brintoni to determine possible transmission pathways. All P. brintoni endoparasitic life stages and the transmission tomite stage (outside the host) were associated with bacterial assemblages. 16S rRNA sequences from isolated bacterial strains showed that Photobacterium spp. and Pseudoalteromonas spp. were dominant members of the bacterial assemblages during all life phases of P. brintoni and potential pathobionts. They were apparently unaffected by the krill's immune system or the histophagous activity of P. brintoni. However, other bacterial strains were found only in certain P. brintoni life phases, indicating that as the infection progressed, microhabitat conditions and microbial interactions may have become unfavorable for some strains of bacteria. Trophic infection is the most parsimonious explanation for how P. brintoni infects krill. We estimated N. simplex vulnerability to P. brintoni infection during more than three-fourths of their life span, infecting mostly adult females. The ciliates have relatively high prevalence levels (albeit at <10% of sampled stations) and a short life cycle (estimated <7 d). Histophagous ciliate-krill interactions may occur in other krill species, particularly those that form dense swarms and attain high population densities that potentially enhance trophic transmission and allow completion of the Pseudocollinia spp. life cycle.