A siliceous arms race in pelagic plankton.

Coevolution between predator and prey plays a central role in shaping the pelagic realm and may have significant implications for marine ecosystems and nutrient cycling dynamics. The siliceous diatom frustule is often assumed to have coevolved with the silica-lined teeth of copepods, but empirical evidence of how this relationship drives natural selection and evolution is still lacking. Here, we show that feeding on diatoms causes significant wear and tear on copepod teeth and that this leads to copepods becoming selective feeders. Teeth from copepods feeding on thick-shelled diatoms were more likely to be broken or cracked than those feeding on a dinoflagellate. When fed a large diatom, all analyzed teeth had visible wear. Our results underscore the importance of the predator-prey arms race as a driving force in planktonic evolution and diversity.

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.

Keratinocytes drive the epithelial hyperplasia key to sea lice resistance in coho salmon.

BACKGROUND: Salmonid species have followed markedly divergent evolutionary trajectories in their interactions with sea lice. While sea lice parasitism poses significant economic, environmental, and animal welfare challenges for Atlantic salmon (Salmo salar) aquaculture, coho salmon (Oncorhynchus kisutch) exhibit near-complete resistance to sea lice, achieved through a potent epithelial hyperplasia response leading to rapid louse detachment. The molecular mechanisms underlying these divergent responses to sea lice are unknown. RESULTS: We characterized the cellular and molecular responses of Atlantic salmon and coho salmon to sea lice using single-nuclei RNA sequencing. Juvenile fish were exposed to copepodid sea lice (Lepeophtheirus salmonis), and lice-attached pelvic fin and skin samples were collected 12 h, 24 h, 36 h, 48 h, and 60 h after exposure, along with control samples. Comparative analysis of control and treatment samples revealed an immune and wound-healing response that was common to both species, but attenuated in Atlantic salmon, potentially reflecting greater sea louse immunomodulation. Our results revealed unique but complementary roles of three layers of keratinocytes in the epithelial hyperplasia response leading to rapid sea lice rejection in coho salmon. Our results suggest that basal keratinocytes direct the expansion and mobility of intermediate and, especially, superficial keratinocytes, which eventually encapsulate the parasite. CONCLUSIONS: Our results highlight the key role of keratinocytes in coho salmon's sea lice resistance and the diverged biological response of the two salmonid host species when interacting with this parasite. This study has identified key pathways and candidate genes that could be manipulated using various biotechnological solutions to improve Atlantic salmon sea lice resistance.

Removal of Exogenous Stimuli Reveals a Canalization of Circadian Physiology in a Vertically Migrating Copepod.

Diel rhythms are observed across taxa and are important for maintaining synchrony between the environment and organismal physiology. A striking example of this is the diel vertical migration undertaken by zooplankton, some of which, such as the 5 mm-long copepod Pleuromamma xiphias (P. xiphias), migrate hundreds of meters daily between the surface ocean and deeper waters. Some of the molecular pathways that underlie the expressed phenotype at different stages of this migration are entrained by environmental variables (e.g., day length and food availability), while others are regulated by internal clocks. We identified a series of proteomic biomarkers that vary across ocean DVM and applied them to copepods incubated in 24 h of darkness to assess circadian control. The dark-incubated copepods shared some proteomic similarities to the ocean-caught copepods (i.e., increased abundance of carbohydrate metabolism proteins at night). Shipboard-incubated copepods demonstrated a clearer distinction between night and day proteomic profiles, and more proteins were differentially abundant than in the in situ copepods, even in the absence of the photoperiod and other environmental cues. This pattern suggests that there is a canalization of rhythmic diel physiology in P. xiphias that reflects likely circadian clock control over diverse molecular pathways.

Revealing the profound influence of diapause on gene expression: Insights from the annual transcriptome of the copepod Calanus finmarchicus.

Annual rhythms are observed in living organisms with numerous ecological implications. In the zooplanktonic copepod Calanus finmarchicus, such rhythms are crucial regarding its phenology, body lipid accumulation, and global carbon storage. Climate change drives annual biological rhythms out of phase with the prevailing environmental conditions with yet unknown but potentially catastrophic consequences. However, the molecular dynamics underlying phenology are still poorly described. In a rhythmic analysis of C. finmarchicus annual gene expression, results reveal that more than 90% of the transcriptome shows significant annual rhythms, with abrupt and dramatic upheaval between the active and diapause life cycle states. This work explores the implication of the circadian clock in the annual timing, which may control epigenetic mechanisms to profoundly modulate gene expression in response to calendar time. Results also suggest an increased light sensitivity during diapause that would ensure the photoperiodic entrainment of the endogenous annual clock.

Interactive effects of temperature and salinity on metabolism and activity of the copepod Tigriopus californicus.

In natural environments, two or more abiotic parameters often vary simultaneously, and interactions between co-varying parameters frequently result in unpredictable, non-additive biological responses. To better understand the mechanisms and consequences of interactions between multiple stressors, it is important to study their effects on not only fitness (survival and reproduction) but also performance and intermediary physiological processes. The splash-pool copepod Tigriopus californicus tolerates extremely variable abiotic conditions and exhibits a non-additive, antagonistic interaction resulting in higher survival when simultaneously exposed to high salinity and acute heat stress. Here, we investigated the response of T. californicus in activity and oxygen consumption under simultaneous manipulation of salinity and temperature to identify whether this interaction also arises in these sublethal measures of performance. Oxygen consumption and activity rates decreased with increasing assay salinity. Oxygen consumption also sharply increased in response to acute transfer to lower salinities, an effect that was absent upon transfer to higher salinities. Elevated temperature led to reduced rates of activity overall, resulting in no discernible impact of increased temperature on routine metabolic rates. This suggests that swimming activity has a non-negligible effect on the metabolic rates of copepods and must be accounted for in metabolic studies. Temperature also interacted with assay salinity to affect activity, and with acclimation salinity to affect routine metabolic rates upon acute salinity transfer, implying that the sublethal impacts of these co-varying factors are also not predictable from experiments that study them in isolation.

Environmental influence on the Atlantic salmon transcriptome and methylome during sea lice infestations.

The fish's immune response is affected by different factors, including a wide range of environmental conditions that can also disrupt or promote changes in the host-pathogen interactions. How environmental conditions modulate the salmon genome during parasitism is poorly understood here. This study aimed to explore the environmental influence on the Salmo salar transcriptome and methylome infected with the sea louse Caligus rogercresseyi. Atlantic salmon were experimentally infected with lice at two temperatures (8 and 16 °C) and salinity conditions (32 and 26PSU). Fish tissues were collected from the infected Atlantic salmon for reduced representation bisulfite sequencing (RRBS) and whole transcriptome sequencing (RNA-seq) analysis. The parasitic load was highly divergent in the evaluated environmental conditions, where the lowest lice abundance was observed in fish infected at 8 °C/26PSU. Notably, transcriptome profile differences were statistically associated with the number of alternative splicing events in fish exposed to low temperature/salinity conditions. Furthermore, the temperature significantly affected the methylation level, where high values of differential methylation regions were observed at 16 °C. Also, the association between expression levels of spliced transcripts and their methylation levels was determined, revealing significant correlations with Ferroptosis and TLR KEEG pathways. This study supports the relevance of the environmental conditions during host-parasite interactions in marine ecosystems. The discovery of alternative splicing transcripts associated with DMRs is also discussed as a novel player in fish biology.

Molecular characterization of Lernathropus kroyeri from intensive aquaculture and pathophysiology of infested sea bass.

The copepod Lernathropus kroyeri constitutes one of the major parasites for the Mediterranean aquaculture, infesting the sea bass Dicentrarchus labrax causing thus disruptions of growth performance and occasionally mortalities. Despite the large spread and the high frequency of this parasite in mariculture farms of Eastern Mediterranean, L. kroyeri genetic profile from aquaculture as well as the pathophysiological response of D. labrax have not been studied so far. Keeping this in mind, in the present study we investigated the L. kroyeri infestation on D. labrax from two farms in Greece, examining both healthy and heavy parasitized individuals. Assays included histopathology, phylogenetic reconstruction of the parasite and physiological response of the fish by the means of antioxidant, inflammatory metabolic and stress related gene expression analysis at both mRNA and protein levels. Genetic analysis indicated that L. kroyeri composes a monophyletic group, highly phylogenetically distant from other congeneric groups. Heavy infested D. labrax witnessed a significantly increased immune response that further led to oxidative stress and metabolic alterations. Overall, our results demonstrate the, seasonally independent, high infestation of this parasitic copepods, which continue to affect Mediterranean intensive aquaculture systems.

A comparative analysis of alternative splicing patterns in Atlantic salmon (Salmo salar) in response to Moritella viscosa and sea lice (Lepeophtheirus salmonis) infection.

Moritella viscosa (M. viscosa) and sea lice (Lepeophtheirus salmonis) are severe pathogens that primarily infect the skin of Atlantic salmon (Salmo salar), which cause significant economic losses in the farming industry. However, the pathogenesis and molecular mechanisms underlying the host's immune defence at the post-transcriptional level remain unclear. Alternative splicing (AS) is an evolutionarily conserved post-transcriptional mechanism that can greatly increase the richness of the transcriptome and proteome. In this study, transcriptomic data derived from skin tissues of Atlantic salmon after M. viscosa and sea lice infections were used to examine the AS profiles and their differential expression patterns. In total, we identified 33,044 AS events (involving 13,718 genes) in the control (CON) group, 35,147 AS events (involving 14,340 genes) in the M. viscosa infection (MV) group, and 30,364 AS events (involving 13,142 genes) in the sea lice infection (LC) group, respectively. Among the five types of AS identified in our study (i.e., SE, A5SS, A3SS, MXE, and RI), SE was the most prevalent type in all three groups (i.e., CON, MV, and LC groups). Decreased percent-spliced-in (PSI) levels were observed in SE events under both MV- and LC-infected conditions, suggesting that MV or LC infection elevated exon-skipping isoforms and promoted the selection of shorter transcripts in numerous DAS genes. In addition, most of the differential AS genes were found to be associated with pathways related to mRNA regulation, epithelial or muscle development, and immune response. These findings provide novel insights into the role of AS in host-pathogen interactions and represent the first comparative analysis of AS in response to bacterial and parasitic infections in fish.

Morphological and Molecular Evidence of an Intergeneric Host-Range in Clavisodalis sentifer (Crustacea: Copepoda: Taeniacanthidae) Associated with Diadematid Sea Urchins from the Western Pacific.

Sea urchins have a wide variety of symbionts on their body surfaces and inside their bodies. Copepods of the genus Clavisodalis (Taeniacanthidae) collected from the esophagus of sea urchins of the genera Diadema and Echinothrix in southern Japan were identified based on their morphological characteristics, and molecular analysis was conducted to determine whether genetic variation occurs in copepods from different localities and hosts. Morphological observations identified individuals from southern Japan as Clavisodalis sentifer Dojiri and Humes, 1982, making this the first record of this species in the northern hemisphere and the first record of its genus in Japan. Morphological and molecular analysis suggested that the copepod specimens collected from multiple hosts across two genera would be the same species. Considering the typically observed high level of host specificity among taeniacanthid copepods, the utilization of hosts from two genera by C. sentifer is noteworthy.

Pennella balaenoptera actively select injured cetacean skin as attachment sites, making them potentially useful forensic tags.

Cetaceans harbor multiple epibionts on their external surface, and these attach to particular microhabitats. Understanding what drives the selection of attachment sites is relevant for refining the use of epibionts as indicators of their hosts. We report on about 100 females of the mesoparasitic copepod Pennella balaenoptera attached to a dead Cuvier's beaked whale Ziphius cavirostris stranded in Tunisia (western Mediterranean); the first report of P. balaenoptera in this country. The copepods were exclusively attached to numerous incisive, likely anthropogenic, wounds found on the host's skin. This finding suggests that newly recruited females may actively seek skin areas where physical penetration is facilitated; a factor that may help explain patterns of microhabitat selection by Pennella spp., and perhaps other pennellids, on their hosts. The estimated age of parasitization by P. balaenoptera (supported by age estimations of the co-occurring epibiotic barnacle Conchoderma virgatum) also suggests that the cetacean host likely survived these injuries, at least initially, and the presumed cause of death was starvation due to entanglement in a fishing net.

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.

How host ecology influences the parasite communities of three Australian flathead fishes, Platycephalus spp. (Scorpaeniformes: Platycephalidae).

This study investigated the role of host traits, habitat, and sampling season on the prevalence and intensity of parasites in three species of platycephalid fishes. Three host species sampled were dusky flathead (Platycephalus fuscus, n = 3), blue-spotted flathead (Platycephalus caeruleopunctatus, n = 38), and tiger flathead (Platycephalus richardsoni, n = 59). A total of 14 metazoan parasite species were collected over 15 months, between July 2020 and September 2021. The parasites found included a chondracanthid copepod, adult hemiuroidean trematodes, cestode plerocercoids, larval and adult acanthocephalans, larval and adult nematodes, and an unknown species of helminth. General linear models were used to assess the relationship between host traits and sampling season with parasite infection intensity. The infection intensity of an unidentified plerocercoid species in P. caeruleopunctatus was found to be significantly associated with both sampling season and the interaction of host mass with sex. In P. richardsoni the infection intensity of the acanthocephalan, Corynosoma sp. was found to correlate with sampling season. The highest richness of parasite taxa was recorded from P. richardsoni, which may be due to the wide depth range of P. richardsoni, coupled with its generalist diet.

The impact of dissolved oxygen concentration on the swimming performance and welfare of ballan wrasse (Labrus bergylta Ascanius, 1767).

Sea lice (Lepeophtheirus salmonis [Krøyer, 1838]) are a key issue for salmon aquaculture, contributing to increased mortality for both wild and farmed salmon if no action is taken. Using cleaner fish can be an effective, drug-free treatment method, and ballan wrasse (Labrus bergylta) is a hardy wrasse species that displays cleaning behavior. With concerns about the overharvest of wild ballan wrasse, many companies farm this species, but the optimal ranges of a wide variety of rearing parameters are still unknown. This study investigated the effect of 6-week exposure to four dissolved oxygen (DO) levels (125%, 100%, 85%, and 75% DO saturation as the percentage of air) on ballan wrasse. Survival; growth (specific growth rate, SGR); condition factor (CF); and primary (cortisol), secondary (glucose, lactate, magnesium), and tertiary stress indicators (swimming performance) were investigated. There were no differences in SGR, CF, survival, or cortisol level among the groups at the end of the 6 weeks. There was variation in the magnitude of the cortisol response to an acute stressor at the end of the 6-week period, with the 75% DO treatment exhibiting a 3.3-fold increase in cortisol compared to a 5.2-fold increase in the control group (100%), which could suggest chronic stress. Relative critical swimming speed (RUcrit) was measured to investigate swimming performance once all groups were returned to 100% DO saturation. The 75% RUcrit was lower than the 100% treatment (1.7 ± 0.18 body length [BL]/s compared to 2.5 ± 0.16 BL/s). Overall, these results suggest that DO levels of 75% trigger physiological changes and therefore may negatively affect welfare.

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.

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.

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.

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.

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.

Costs and benefits of predator-induced defence in a toxic diatom.

Phytoplankton employ a variety of defence mechanisms against predation, including production of toxins. Domoic acid (DA) production by the diatom Pseudo-nitzschia spp. is induced by the presence of predators and is considered to provide defence benefits, but the evidence is circumstantial. We exposed eight different strains of P. seriata to chemical cues from copepods and examined the costs and the benefits of toxin production. The magnitude of the induced toxin response was highly variable among strains, while the costs in terms of growth reduction per DA cell quota were similar and the trade-off thus consistent. We found two components of the defence in induced cells: (i) a 'private good' in terms of elevated rejection of captured cells and (ii) a 'public good' facilitated by a reduction in copepod feeding activity. Induced cells were more frequently rejected by copepods and rejections were directly correlated with DA cell quota and independent of access to other food items. By contrast, the public-good effect was diminished by the presence of alternative prey suggesting that it does not play a major role in bloom formation and that its evolution is closely associated with the grazing-deterrent private good.