Contrasting responses of Thermocyclops crassus and T. oithonoides (Crustacea, Copepoda) to thermal stress.

Thermal tolerance is a critical factor influencing the survival of living organisms. This study focuses on the thermal resistance of copepod species, Thermocyclops crassus (Fischer, 1853) and T. oithonoides (Sars G.O., 1863), with overlapping distribution ranges in Europe. Short-term heat shock experiments were conducted to assess the thermal resistance of these copepods, considering various temperature increments and exposure durations. Additionally, the study explored the influence of heat shock on egg sac shedding, a vital indicator of population dynamics. Results indicate that widely distributed T. crassus exhibits higher thermal tolerance compared to narrowly distributed T. oithonoides, with survival rates varying under different heat shock conditions. Furthermore, T. crassus demonstrated a quicker response in dropping egg sacs in response to thermal stress, suggesting a potential adaptive mechanism for the survival of adults. However, rapid egg sac droppings pose high risks for eggs facing unfavorable conditions. T. crassus, inhabiting environments with greater temperature fluctuations such as the littoral and pelagial zones, exhibited better survival mechanisms compared to T. oithonoides, which predominantly resides in the pelagic zone. The findings have implications for understanding copepod responses to global warming and thermal pollution. This research contributes insights into the adaptive strategies of thermophilic copepod species and their ecological consequences.

Environmental factors controlling seasonal and spatial variability of zooplankton in thermokarst lakes along a permafrost gradient of Western Siberia.

Humic thermokarst lakes of permafrost peatlands in Western Siberia Lowland (WSL) are major environmental controllers of carbon and nutrient storage in inland waters and greenhouse gases emissions to the atmosphere in the subarctic. In contrast to sizable former research devoted to hydrochemical and hydrobiological (phytoplankton) composition, zooplankton communities of these thermokarst lakes and thaw ponds remain poorly understood, especially along the latitudinal gradient, which is a perfect predictor of permafrost zones. To fill this gap, 69 thermokarst lakes of the WSL were sampled using unprecedented spatial coverage, from continuous to sporadic permafrost zone, in order to assess zooplankton (Cladocera, Copepoda, Rotifera) diversity and abundance across three main open water physiological seasons (spring, summer and autumn). We aimed at assessing the relationship of environmental factors (water column hydrochemistry, nutrients, and phytoplankton parameters) with the abundance and diversity of zooplankton. A total of 74 zooplankton species and taxa were detected, with an average eight taxa per lake/pond. Species richness increased towards the north and reached the maximum in the continuous permafrost zone with 13 species found in this zone only. In contrast, the number of species per waterbody decreased towards the north, which was mainly associated with a decrease in the number of cladocerans. Abundance and diversity of specific zooplankton groups strongly varied across the seasons and permafrost zones. Among the main environmental controllers, Redundancy Analysis revealed that water temperature, lake area, depth, pH, Dissolved Inorganic and Organic Carbon and CO2 concentrations were closely related to zooplankton abundance. Cladocerans were positively related to water temperature during all seasons. Copepods were positively related to depth and lake water pH in all seasons. Rotifers were related to different factors in each season, but were most strongly associated with DOC, depth, CH4, phytoplankton and cladoceran abundance. Under climate warming scenario, considering water temperature increase and permafrost boundary shift northward, one can expect an increase in the diversity and abundance of cladocerans towards the north which can lead to partial disappearance of copepods, especially rare calanoid species.

Escape performance in the cyclopoid copepod Oithona davisae.

Escaping a predator is one of the keys to success for any living creature. The performance of adults (males, females, and ovigerous females) of the cyclopoid copepod Oithona davisae exposed to an electrical stimulus is analysed as a function of temperature by measuring characteristic parameters associated with the escape movement (distance covered, duration of the appendage movement, mean and maximum escape speeds, Reynolds number). In addition, as a proxy for the efficiency of the motion, the Strouhal number was calculated. The escape performance showed temperature-dependent relationships within each adult state, as well as differences between sexes; additionally, changes owing to the presence of the egg sac were recorded in females. In a broader perspective, the results collected reveal the occurrence of different behavioural adaptations in males and females, adding to the comprehension of the mechanisms by which O. davisae interacts with its environment and shedding new light on the in situ population dynamics of this species.

Lufenuron treatment temporarily represses gene expression and affects the SUMO pathway in liver of Atlantic salmon.

Lufenuron is a benzoylurea insecticide currently in use to combat sea lice infestation in salmon aquaculture in Chile. With pending approval in Norway, the aim of this work was to study the uptake and toxicity of lufenuron in liver tissue of Atlantic salmon. Juvenile salmon weighing 40 g were given a standard 7-day oral dose, and bioaccumulation and transcriptional responses in the liver were examined 1 day after the end-of-treatment (day 8) and after 1 week of elimination (day 14). Bioaccumulation levels of lufenuron were 29 ± 3 mg/kg at day 8 and 14 ± 1 mg/kg at day 14, indicating relatively rapid clearance. However, residues of lufenuron were still present in the liver after 513 days of depuration. The exposure gave a transient inhibition of transcription in the liver at day 8 (2437 significant DEGs, p-adj < .05), followed by a weaker compensatory response at day 14 (169 significant DEGs). Pathways associated with RNA metabolism such as the sumoylation pathway were most strongly affected at day 8, while the apelin pathway was most profoundly affected at day 14. In conclusion, this study shows that lufenuron easily bioaccumulates and that a standard 7-day oral dose induces a transient inhibition of transcription in liver of salmon.

Transcriptome analysis revealed immune responses in the kidney of Atlantic salmon (Salmo salar) co-infected with sea lice (Lepeophtheirus salmonis) and infectious salmon anemia virus.

Sea lice (Lepeophtheirus salmonis) and infectious salmon anemia virus (ISAv) are two of the most important pathogens in Atlantic salmon (Salmo salar) farming and typically cause substantial economic losses to the industry. However, the immune interactions between hosts and these pathogens are still unclear, especially in the scenario of co-infection. In this study, we artificially infected Atlantic salmon with sea lice and ISAv, and investigated the gene expression patterns of Atlantic salmon head kidneys in response to both lice only and co-infection with lice and ISAv by transcriptomic analysis. The challenge experiment indicated that co-infection resulted in a cumulative mortality rate of 47.8 %, while no mortality was observed in the lice alone infection. We identified 240 differentially expressed genes (DEGs) under the lice alone infection, of which 185 were down-regulated and 55 were up-regulated, while a total of 994 DEGs were identified in the co-infection, of which 206 were down-regulated and 788 were significantly up-regulated. The pathway enrichment analysis revealed that single-infection significantly suppressed the innate immune system (e.g., the complement system), whereas co-infection induced a strong immune response, leading to the activation of immune-related signaling pathways such as Toll-like receptors and NOD-like receptors pathways, as well as significant upregulation of genes related to the activation of interferon and MH class I protein complex. Our results provide the first global transcriptomic study of gene expression in the Atlantic salmon head kidney in response to co-infection with sea lice and ISAv, and provided the baseline knowledge for understanding the immune responses during co-infection.

A modeling study of the impact of treatment policies on the evolution of resistance in sea lice on salmon farms.

Salmonid aquaculture is an important source of nutritious food with more than 2 million tonnes of fish produced each year (Food and Agriculture Organisation of the United Nations, 2019). In most salmon producing countries, sea lice represent a major barrier to the sustainability of salmonid aquaculture. This issue is exacerbated by widespread resistance to chemical treatments on both sides of the Atlantic. Regulation for sea lice management mostly involves reporting lice counts and treatment thresholds, which depending on interpretation may encourage preemptive treatments. We have developed a stochastic simulation model of sea lice infestation including the lice life-cycle, genetic resistance to treatment, a wildlife reservoir, salmon growth and stocking practices in the context of infestation, and coordination of treatment between farms. Farms report infestation levels to a central organisation, and may then cooperate or not when coordinated treatment is triggered. Treatment practice then impacts the level of resistance in the surrounding sea lice population. Our simulation finds that treatment drives selection for resistance and coordination between managers is key. We also find that position in the hydrologically-derived network of farms can impact individual farm infestation levels and the topology of this network can impact overall infestation and resistance. We show how coordination and triggering of treatment alongside varying hydrological topology of farm connections affects the evolution of lice resistance, and thus optimise salmon quality within socio-economic and environmental constraints. Network topology drives infestation levels in cages, treatments, and hence treatment-driven resistance. Thus farmer behaviour may be highly dependent on hydrologically position and local level of infestation.

The response of zooplankton network indicators to winter water warming using shallow artificial reservoirs as model case study.

To predict the most likely scenarios, the consequences of the rise in water surface temperature have been studied using various methods. We tested the hypothesis that winter water warming significantly alters the importance and nature of the relationships in zooplankton communities in shallow reservoirs. These relationships were investigated using network graph analysis for three thermal variants: warm winters (WW), moderate winters (MW) and cold winters (CW). The CW network was the most cohesive and was controlled by eutrophic Rotifera and Copepoda, with a corresponding number of positive and negative interspecific relationships. An increase in water temperature in winter led to a decrease in the centrality of MW and WW networks, and an increase in the importance of species that communicated with the highest number of species in the subnetworks. The WW network was the least cohesive, controlled by psammophilous and phytophilous rotifers, and littoral cladocerans. Adult copepods were not identified in the network and the importance of antagonistic relationships decreased, indicating that the WW network structure was weak and unstable. This study can serve as a model for generalisations of zooplankton community response to the disappearance of long winter periods of low temperatures, as predicted in global climate change projections.

Joint species distribution modeling reveals a changing prey landscape for North Pacific right whales on the Bering shelf.

The eastern North Pacific right whale (NPRW) is the most endangered population of whale and has been observed north of its core feeding ground in recent years with low sea ice extent. Sea ice and water temperature are important drivers for zooplankton dynamics within the whale's core feeding ground in the southeastern Bering Sea, seasonally forming stable fronts along the shelf that give rise to distinct zooplankton communities. A northward shift in NPRW distribution driven by changing distribution of prey resources could put this species at increased risk of entanglement and vessel strikes. We modeled the abundance of NPRW prey, Calanus glacialis, Neocalanus, and Thysanoessa species, using a dynamic biophysical food web model of nine zooplankton guilds in the Bering shelf zooplankton community during a period of warming (2006-2016). This model is unique among prior zooplankton studies from the region in that it includes density dependence, thereby allowing us to ask whether species interactions influence zooplankton dynamics. Modeling confirmed the importance of sea ice and ocean temperature to zooplankton dynamics in the region. Density-independent growth drove community dynamics, while dependent factors were comparatively minimal. Overall, Calanus responded to environment terms, with the strength and direction of response driven by copepodite stage. Neocalanus and Thysanoessa responses were weaker, likely due to their primary occurrence on the outer shelf. We also modeled the steady-state (equilibrium) abundance of Calanus in conditions with and without wind gusts to test whether advection of outer shelf species might disrupt the steady-state dynamics of Calanus abundance; the results did not support disruption. Given the annual fall sampling design, we interpret our results as follows: low-ice-extent winters induced stronger spring winds and weakened fronts on the shelf, thereby advecting some outer shelf species into the study region; increased development rates in these warm conditions influenced the proportion of C. glacialis copepodite stages over the season. Residual correlation suggests missing drivers, possibly predators, and phytoplankton bloom composition. Given the continued loss of sea ice in the region and projected continued warming, our findings suggest that C. glacialis will move northward, and thus, whales may move northward to continue targeting them.

Immune response induced by coinfection of the sea louse Caligus rogercresseyi and the intracellular bacteria Piscirickettsia salmonis in vaccinated Atlantic salmon.

Recently, we showed that Atlantic salmon vaccinated against Piscirickettsia salmonis lose their protection upon coinfection with Caligus rogercresseyi (sea lice). However, the causes of the overriding effect of C. rogercresseyi infection have not been elucidated, and the molecular basis of the cellular and humoral immune responses upon C. rogercresseyi infection has not been described for vaccinated salmon. Therefore, we studied changes in the transcription of immune genes in vaccinated Atlantic salmon that were experimentally challenged by co-infecting them with C. rogercresseyi and P. salmonis. In general, coinfection treatments showed immune gene expression similar to treatments with a single P. salmonis infection, showing a decreased cellular response. However, a high variance was found between individual fish in the case of crucial cellular immune genes, with a few fish reacting overwhelmingly highly compared to the majority. This supports our previous findings on vaccination response variation and reinforces the idea that vaccination failures in the field might be caused by an overwhelming amount of vaccinated fish that display a deficient immune response to the infection.

Repeated exposure affects susceptibility and responses of Atlantic salmon (Salmo salar) towards the ectoparasitic salmon lice (Lepeophtheirus salmonis).

Atlantic salmon (Salmo salar) is repeatedly exposed to and infected with ectoparasitic salmon lice (Lepeophtheirus salmonis) both in farms and in nature. However, this is not reflected in laboratory experiments where fish typically are infected only once. To investigate if a previous lice infection affects host response to subsequent infections, fish received 4 different experimental treatments; including 2 groups of fish that had previously been infected either with adult or infective salmon lice larvae (copepodids). Thereafter, fish in all treatment groups were infected with either a double or a single dose of copepodids originating from the same cohort. Fish were sampled when lice had developed into the chalimus, the pre-adult and the adult stage, respectively. Both the specific growth rate and cortisol levels (i.e. a proxy for stress) of the fish differed between treatments. Lice success (i.e. ability to infect and survive on the host) was higher in naïve than in previously infected fish (pre-adult stage). The expression of immune and wound healing transcripts in the skin also differed between treatments, and most noticeable was a higher upregulation early in the infection in the group previously infected with copepodids. However, later in the infection, the least upregulation was observed in this group, suggesting that previous exposure to salmon lice affects the response of Atlantic salmon towards subsequent lice infections.

Rejection of Lepeophtheirus salmonis driven in part by chitin sensing is not impacted by seawater acclimitization in Coho salmon (Oncorhynchus kisutch).

There is tremendous variation in life-history strategies among anadromous salmonids. Species that enter the ocean environment at small sizes (< 20 g) are likely under more physiological pressure from pathogens; however, little data is available on responses at these early stages. With this in mind, we performed salmon louse challenges with Coho salmon either immediately after seawater entry (SW; ca. 10 g) or after 30 days in SW (ca. 20 g). Irrespective of size or time in SW, parasites were rapidly rejected by the host, with > 90% of all parasites lost by 16 days post-infection (dpi). Rejection was concomitant with host epithelial granulomatous infiltrations that initially targeted the embedded frontal filament (4 dpi) and the entire parasite by 10 dpi. Illumina sequencing, followed by functional enrichment analysis, revealed a concerted defense response in the fin within 1 dpi that included multiple innate and adaptive immunity components. Strikingly, early indications of an allergic-type inflammatory response were associated with chitin sensing pathways orchestrated by early overexpression of the IgE-receptor, fcer1g. Additionally, there was profound overexpression of several classes of c-type lectin receptors, including dectin-2, mincle, and dc-sign at 1 dpi onward. These profiles and upregulation of cellular effector markers were corroborated by histopathological evaluation, revealing the simultaneous presence of mast cell/eosinophilic granular cells, sacciform cells, macrophages/histiocytes, and granulocytes in fin. At 10 dpi and concurrent with parasite expulsion, there was evidence of immunoregulation in addition to tissue remodelling pathways. At 16 dpi, the response was effectively abrogated. Simultaneous profiling of the parasite transcriptome revealed early induction of chitin metabolism and immunomodulation, toxin production and ECM degradation; however, after 7 dpi, these were replaced with overexpression of stress and immune defense genes. These data present the first evidence for Coho salmon demonstrating chitin- and sugar moiety-sensing as key drivers of salmon louse rejection.

Decadal decline of dominant copepod species in the North Sea is associated with ocean warming: Importance of marine heatwaves.

Time-series are crucial to understand the status of zooplankton communities and to anticipate changes that might affect the entire food web. Long-term time series allow us to understand impacts of multiple environmental and anthropogenic stressors, such as chemical pollution and ocean warming, on the marine ecosystems. Here, a recent time series (2018-2022) of abundance data of four dominant calanoid and one harpacticoid copepod species from the Belgian Part of the North Sea was combined with previously collected (2009-2010, 2015-2016) datasets for the same study area. The time series reveals a significant decrease (up to two orders of magnitude) in calanoid copepod abundance (Temora longicornis, Acartia clausi, Centropages spp., Calanus helgolandicus), while this was not the case for the harpacticoid Euterpina acutifrons. We applied generalized additive models to quantify the relative contribution of temperature, nutrients, salinity, primary production, turbidity and pollution (anthropogenic chemicals, i.e., polychlorinated biphenyls and polycyclic aromatic hydrocarbons) to the population dynamics of these species. Temperature, turbidity and chlorophyll a concentrations were the only variables consistently showing a relative high contribution in all models predicting the abundances of the selected species. The observed heat waves which occurred during the summer periods of the investigated years coincided with population collapses (versus population densities in non-heatwave years) and are considered the most likely cause for the observed copepod abundance decreases. Moreover, the recorded water temperatures during these heatwaves correspond to the physiological thermal limit of some of the studied species. As far as we know, this is the first study to observe ocean warming and marine heat waves having such a dramatic impact (population collapse) on the dominant zooplankton species in shallow coastal areas.

High and diurnally fluctuating carbon dioxide exposure produces lower mercury toxicity in a marine copepod.

Coastal waters have experienced fluctuations in partial pressure of carbon dioxide (pCO2) and mercury (Hg) pollution, yet little is known concerning how natural pCO2 fluctuations affect Hg biotoxicity. Here, a marine copepod Tigriopus japonicus was interactively exposed to different seawater pCO2 (ambient 400, steady elevated 1000, and fluctuating elevated 1000 ± 600 µatm) scenarios and Hg (control, 2 µg/L) treatments for 7 d. The results showed that elevated pCO2 decreased Hg bioaccumulation, and it was even more under fluctuating elevated pCO2 condition. We found energy depletion and oxidative stress under Hg-treated copepods, while combined exposure initiated compensatory response to alleviate Hg toxicity. Intriguingly, fluctuating acidification presented more immune defense related genes/processes in Hg-treated copepods when compared to steady acidification, probably linking with the greater decrease in Hg bioaccumulation. Collectively, understanding how fluctuating acidification interacts with Hg contaminant will become more crucial in predicting their risks to coastal biota and ecosystems.

Global distribution patterns of Caligus Müller, 1785 (Copepoda: Caligidae) associated to teleost fishes, with physiological and histopathological data and description of treatment strategies.

This review surveyed information on Caligus Müller, 1785 to identify global infestation patterns and geographic distribution in teleost fishes, as well as physiological and histopathological data and description of treatment strategies. A total 990 samples of Caligus spp. (N = 212 species) obtained of 233 scientific papers on farmed and wild teleost species from 99 families and 30 orders were used, and the highest number of occurrences was on Carangidae. Caligus spp. was predominantly found in marine environments, and only Caligus lacustris and Caligus epidemicus were found in teleost fish of freshwater environments. There was a high prevalence of Caligus spp. on hosts and infestation occurred predominantly in both the tegument and the gills. Caligus species are distributed across different countries and some particularities were identified and discussed. Caligus elongatus and Caligus bonito bonito had the broadest geographic distribution. Histomorphological and hematological disorders caused by infestation by Caligus spp. were reported and discussed, as well as chemotherapeutic products used for controlling and treating the infestations. Variation in the distribution and geographic patterns of Caligus spp. were little evident in many ecosystems and due to the limited data on the infestation of these sea lice on teleost populations in different regions.

Ecophysiological response of marine copepods to dietary elemental imbalances.

We assessed the effects of nutrient imbalanced diets on the feeding, reproduction and gross-growth efficiency of egg production of the copepod Paracartia grani. The cryptophyte Rhodomonas salina, cultivated under balanced (f/2) and imbalanced growth conditions (N and P limitation), served as prey. Copepod C:N and C:P ratios increased in the imbalanced treatments, particularly under P limitation. Feeding and egg production rates did not differ between the balanced and N-limited treatments but decreased under P limitation. We found no evidence of compensatory feeding in P. grani. C gross-growth efficiency averaged 0.34 in the balanced treatment and declined to values of 0.23 and 0.14 for the N- and P-limited treatments, respectively. Under N limitation, N gross-growth efficiency increased significantly to a mean value of 0.69, likely as a result of increasing the nutrient absorption efficiency. P gross-growth efficiency reached values > 1 under P limitation, involving the depletion of body P. Hatching success was >80%, with no differences among diets. Hatched nauplii, however, had lower size and slower development when the progenitor was fed a P-limited diet. This study highlights the effects of P limitation in copepods, which are more constraining than N, and the presence of maternal effects driven by prey nutritional composition that ultimately may affect population fitness.

Modeling present and future distribution of plankton populations in a coastal upwelling zone: the copepod Calanus chilensis as a study case.

Predicting species distribution in the ocean has become a crucial task to assess marine ecosystem responses to ongoing climate change. In the Humboldt Current System (HCS), the endemic copepod Calanus chilensis is one of the key species bioindicator of productivity and water masses. Here we modeled the geographic distribution of Calanus chilensis for two bathymetric ranges, 0-200 and 200-400 m. For the 0-200 m layer, we used the Bayesian Additive Regression Trees (BART) method, whereas, for the 200-400 m layer, we used the Ensembles of Small Models (ESMs) method and then projected the models into two future scenarios to assess changes in geographic distribution patterns. The models were evaluated using the multi-metric approach. We identified that chlorophyll-a (0.34), Mixed Layer Depth (0.302) and salinity (0.36) explained the distribution of C. chilensis. The geographic prediction of the BART model revealed a continuous distribution from Ecuador to the southernmost area of South America for the 0-200 m depth range, whereas the ESM model indicated a discontinuous distribution with greater suitability for the coast of Chile for the 200-400 m depth range. A reduction of the distribution range of C. chilensis is projected in the future. Our study suggests that the distribution of C. chilensis is conditioned by productivity and mesoscale processes, with both processes closely related to upwelling intensity. These models serve as a tool for proposing indicators of changes in the ocean. We further propose that the species C. chilensis is a high productivity and low salinity indicator at the HCS. We recommend further examining multiple spatial and temporal scales for stronger inference.

Nanoplastics induce epigenetic signatures of transgenerational impairments associated with reproduction in copepods under ocean acidification.

Ocean acidification (OA) is one of many major global climate changes that pose a variety of risks to marine ecosystems in different ways. Meanwhile, there is growing concern about how nanoplastics (NPs) affect marine ecosystems. Combined exposure of marine organisms to OA and NPs is inevitable, but their interactive effects remain poorly understood. In this study, we investigated the multi- and transgenerational toxicity of NPs on copepods under OA conditions for ten generations. The findings revealed that OA and NPs have a synergistic negative effect on copepod reproduction across generations. In particular, the transgenerational groups showed reproductive impairments in the F1 and F2 generations (F1T and F2T), even though they were never exposed to NPs. Moreover, our epigenetic examinations demonstrated that the observed intergenerational reproductive impairments are associated with differential methylation patterns of specific genes, suggesting that the interaction of OA and NPs can pose a significant threat to the sustainability of copepod populations through epigenetic modifications. Overall, our findings provide valuable insight into the intergenerational toxicity and underlying molecular mechanisms of responses to NPs under OA conditions.

Towards better survival: Modeling drivers for daily mortality in Norwegian Atlantic salmon farming.

Mortality in the production of farmed salmonids is a major constraint to the sustainability of this form of animal husbandry. We have developed a model for the daily mortality in salmon farming over a full production cycle from stocking to harvest, considering different environmental and production factors. These factors included sea temperature, salinity, day within year, fish weight at stocking, stocking day, four types of lice treatments and the possible occurrence of pancreas disease (PD). We considered a generalized additive model following full production cycles, allowing for non-linear descriptions of how relevant factors relate to the daily mortality. We saw a high overall mortality rate immediately after stocking, which decreased the first three months in the cycle and thereafter increased. We found that the total mortality could be reduced by 21% if avoiding all lice treatments, and similarly reduced by 20% if no PD infections occurred. If avoiding jointly PD and all lice treatments, the accumulated mortality could be reduced by 34%. A single thermal or hydrogen peroxide treatment was associated with a mortality of around 1.6% and 1.3%, respectively. This modeling approach gave a unique opportunity to model how different factors interact on the overall global mortality and can easily be extended by other factors, such as additional fish diseases.

Experimental evolution reveals the synergistic genomic mechanisms of adaptation to ocean warming and acidification in a marine copepod.

Metazoan adaptation to global change relies on selection of standing genetic variation. Determining the extent to which this variation exists in natural populations, particularly for responses to simultaneous stressors, is essential to make accurate predictions for persistence in future conditions. Here, we identified the genetic variation enabling the copepod Acartia tonsa to adapt to experimental ocean warming, acidification, and combined ocean warming and acidification (OWA) over 25 generations of continual selection. Replicate populations showed a consistent polygenic response to each condition, targeting an array of adaptive mechanisms including cellular homeostasis, development, and stress response. We used a genome-wide covariance approach to partition the allelic changes into three categories: selection, drift and replicate-specific selection, and laboratory adaptation responses. The majority of allele frequency change in warming (57%) and OWA (63%) was driven by shared selection pressures across replicates, but this effect was weaker under acidification alone (20%). OWA and warming shared 37% of their response to selection but OWA and acidification shared just 1%, indicating that warming is the dominant driver of selection in OWA. Despite the dominance of warming, the interaction with acidification was still critical as the OWA selection response was highly synergistic with 47% of the allelic selection response unique from either individual treatment. These results disentangle how genomic targets of selection differ between single and multiple stressors and demonstrate the complexity that nonadditive multiple stressors will contribute to predictions of adaptation to complex environmental shifts caused by global change.

Swimming and respiration in cyclopoid copepods Thermocyclops oithonoides and Oithona davisae and calanoid copepod Paracalanus parvus.

Cyclopoid and calanoid copepods differ in how they move. Cyclopoid copepods use the thoracic legs for cruise and escape swimming while most calanoid copepods use the cephalic appendages for cruise swimming and the thoracic legs for escape reactions. Apart from this gross difference, little is known on the comparative aspects of the locomotor function of copepod appendages. This study investigated the main kinematic patterns of cruise and escape swimming of two small cyclopoid copepods, Thermocyclops oithonoides and Oithona davisae, and a small calanoid copepod, Paracalanus parvus, by video filming at a frame rate of up to 1200 frames/s. During escape swimming, O. davisae and the twice as large P. parvus showed similar movement, jumping at a frequency of 150 Hz and moving at 12 cm s-1 ; however, at a lower jump frequency (∼100 Hz), the cyclopoid T. oithonoides showed an almost two times faster escape swimming than that of P. parvus which has the same body size. This higher speed can be linked to the greater role of the longer abdomen for the flopping strokes in T. oithonoides. In accordance with the Arrhenius law, the kinematic parameters of cruise and escape swimming of T. oithonoides showed temperature dependence in the range of 6.5-27°Ð¡. At a temperature of about 20°C, the respiration rate of O. davisae and P. parvus was 1.6 times higher (i.e., ∼1.5 µg O2 mg-1 h-1 ) than in T. oithonoides during normal swimming; however, in the swarming state, the respiration rate of T. oithonoides increased 3.4 times to 3.0 µg O2 mg-1 h-1 , which was nine times higher than the respiratory rate of anesthetized individuals of this species. Based on the speed and duration of locomotor acts, the cyclopoid T. oithonoides consumed about the same amount of respiratory energy as the calanoid P. parvus, but the mechanical energy required for movement in jumps mode was 1.5 times higher.