The timing and magnitude of an escape reaction is often the determining factor governing a copepod’s success at avoiding predation. Copepods initiate rapid and directed escapes in response to fluid signals created by predators; however little is known about how copepods modulate their behavior in response to additional sensory input. This study investigates the effect of light level on the escape behavior of Calanus finmarchicus. A siphon flow was used to generate a consistent fluid signal and the behavioral threshold and magnitude of the escape response was quantified in the dark and in the light. The results show that C. finmarchicus initiated their escape reaction further from the siphon and traveled with greater speed in the light than in the dark. However, no difference was found in the escape distance. These results suggest that copepods use information derived from multiple sensory inputs to modulate the sensitivity and strength of the escape in response to an increase risk of predation. Population and IBM models that predict optimal vertical distributions of copepods in response to visual predators need to consider changes in the copepod’s behavioral thresholds when predicting predation risk within the water column.
Carotenoids are protective pigments present in many aquatic organisms that reduce the photooxidative stress induced by short-wavelenght solar radiation, yet increase their susceptibility to predators. Arctodiaptomus spinosus, a calanoid copepod typically found in many fishless shallow soda lakes, shows large between-lake differences in pigmentation. Here, we attribute these differences to the environmental state of these ecosystems, namely, ‘dark water’ lakes with submersed vegetation and turbid ‘white’ lakes lacking macrophytes. Copepod carotenoid concentration in the turbid ‘white’ lakes was significantly (about 20-fold) higher than in the ‘dark water’ ones, although the latter systems were characterized by higher transparency. In addition, males had on a dry weight basis around three times higher carotenoid concentrations than females. Mycosporine-like amino acids (direct UV screening substances) were found in all cases, but in low concentration. The environmental conditions in these ecosystems were largely shaped by the presence/absence of submersed macrophytes Thus, in the turbid lakes, the strong wind-driven mixis allows for copepods to be brought to the surface and being exposed to solar radiation, whereas in ‘dark water’ ones, macrophytes reduce water turbulence and additionally provide shelter. Our results explain the counter-intuitive notion of strong red pigmentation in copepods from a turbid ecosystem and suggest that factors other than high UV transparency favor carotenoid accumulation in zooplankton.
The dynamics of the protozoan parasite Marteilia refringens was studied in Thau lagoon, an important French shellfish site, for 1 year in three potential hosts: the Mediterranean mussel Mytilus galloprovincialis (Mytiliidae) the grooved carpet shell Ruditapes decussatus (Veneriidae) and the copepod Paracartia grani (Acartiidae). Parasite DNA was detected by PCR in R. decussatus. In situ hybridization (ISH) showed necrotic cells of M. refringens in the digestive epithelia of some R. decussatus suggesting the non-involvement of this species in the parasite life cycle. In contrast, the detection of M. refringens in mussels using PCR appeared bimodal with two peaks in spring and autumn. Histological observations of PCR-positive mussels revealed the presence of different parasite stages including mature sporangia in spring and autumn. These results suggest that the parasite has two cycles per year in the Thau lagoon and that mussels release parasites into the water column during these two periods. Moreover, PCR detection of the parasite in the copepodid stages of P. grani between June and November supports the hypothesis of the transmission of the parasite from mussels to copepods and conversely. In situ hybridization performed on copepodites showed labeling in some sections. Unusual M. refringens cells were observed in the digestive tract and the gonad from the third copepodid stage, suggesting that the parasite could infect a copepod by ingestion and be released through the gonad. This hypothesis is supported by the PCR detection of parasite DNA in copepod eggs from PCR-positive females, which suggests that eggs could contribute to the parasite spreading in the water and could allow overwintering of M. refringens. Finally, in order to understand the interactions between mussels and copepods, mussel retention efficiency (number of copepods retained by a mussel) was measured for all P. grani developmental stages. Results showed that all copepod stages could contribute to the transmission of the parasite, especially eggs and nauplii which were retained by up to 90%.
Many studies have focused on natural stress factors that shape the spatial and temporal distribution of calanoid copepods, but bioassays have shown that copepods are also sensitive to a broad range of contaminants. Although both anthropogenic and natural stress factors are obviously at play in natural copepod communities, most studies consider only one or the other. In the present investigation, we modelled the combined impact of both anthropogenic and natural stress factors on copepod populations. The model was applied to estimate Eurytemora affinis densities in the contaminated Scheldt estuary and the relatively uncontaminated Darß-Zingst estuary in relation to temperature, salinity, chlorophyll a and sediment concentrations of cadmium, copper and zinc. The results indicated that temperature was largely responsible for seasonal fluctuations of E. affinis densities. Our model results further suggested that exposure to zinc and copper was largely responsible for the reduced population densities in the contaminated estuary. The model provides a consistent framework for integrating and quantifying the impacts of multiple anthropogenic and natural stress factors on copepod populations. It facilitates the extrapolation of laboratory experiments to ecologically relevant endpoints pertaining to population viability.
In the Gulf of Maine, the copepod Calanus finmarchicus co-occurs with the neurotoxin-producing dinoflagellate, Alexandrium fundyense. The copepod is resistant to this toxic alga, but little is known about other effects. Gene expression profiles were used to investigate the physiological response of females feeding for two and five days on a control diet or a diet containing either a low or a high dose of A. fundyense. The physiological responses to the two experimental diets were similar, but changed between the time points. At 5-days the response was characterized by down-regulated genes involved in energy metabolism. Detoxification was not a major component of the response. Instead, genes involved in digestion were consistently regulated, suggesting that food assimilation may have been affected. Thus, predicted increases in the frequency of blooms of A. fundyense could affect C. finmarchicus populations by changing the individuals' energy budget and reducing their ability to build lipid reserves.
DNA metabarcoding is a promising approach for rapidly surveying biodiversity and is likely to become an important tool for measuring ecosystem responses to environmental change. Metabarcoding markers need sufficient taxonomic coverage to detect groups of interest, sufficient sequence divergence to resolve species, and will ideally indicate relative abundance of taxa present. We characterized zooplankton assemblages with three different metabarcoding markers (nuclear 18S rDNA, mitochondrial COI, and mitochondrial 16S rDNA) to compare their performance in terms of taxonomic coverage, taxonomic resolution, and correspondence between morphology- and DNA-based identification. COI amplicons sequenced on separate runs showed that operational taxonomic units representing >0.1% of reads per sample were highly reproducible, although slightly more taxa were detected using a lower annealing temperature. Mitochondrial COI and nuclear 18S showed similar taxonomic coverage across zooplankton phyla. However, mitochondrial COI resolved up to threefold more taxa to species compared to 18S. All markers revealed similar patterns of beta-diversity, although different taxa were identified as the greatest contributors to these patterns for 18S. For calanoid copepod families, all markers displayed a positive relationship between biomass and sequence reads, although the relationship was typically strongest for 18S. The use of COI for metabarcoding has been questioned due to lack of conserved primer-binding sites. However, our results show the taxonomic coverage and resolution provided by degenerate COI primers, combined with a comparatively well-developed reference sequence database, make them valuable metabarcoding markers for biodiversity assessment.
- Proceedings. Biological sciences / The Royal Society
- Published almost 3 years ago
Understanding interactions between harmful algal bloom (HAB) species and their grazers is essential for determining mechanisms of bloom proliferation and termination. We exposed the common calanoid copepod, Temora longicornis to the HAB species Alexandrium fundyense and examined effects on copepod survival, ingestion, egg production and swimming behaviour. A. fundyense was readily ingested by T. longicornis and significantly altered copepod swimming behaviour without affecting copepod survival or fitness. A. fundyense caused T. longicornis to increase their swimming speed, and the straightness of their path long after the copepods had been removed from the A. fundyense treatment. Models suggest that these changes could lead to a 25-56% increase in encounter frequency between copepods and their predators. This work highlights the need to determine how ingesting HAB species alters grazer behaviour as this can have significant impacts on the fate of HAB toxins in marine systems.
For the first time, the phylogenetic relationships between representatives of all 10 copepod orders have been investigated using 28S and 18S rRNA, Histone H3 protein and COI mtDNA. The monophyly of Copepoda (including Platycopioida Fosshagen, 1985) is demonstrated for the first time using molecular data. Maxillopoda is rejected, as it is a polyphyletic group. The monophyly of the major subgroups of Copepoda, including Progymnoplea Lang, 1948 (=Platycopioida); Neocopepoda Huys and Boxshall, 1991; Gymnoplea Giesbrecht, 1892 (=Calanoida Sars, 1903); and Podoplea Giesbrecht, 1892, are supported in this study. Seven copepod orders are monophyletic, including Platycopioida, Calanoida, Misophrioida Gurney, 1933; Monstrilloida Sars, 1901; Siphonostomatoida Burmeister, 1834; Gelyelloida Huys, 1988; and Mormonilloida Boxshall, 1979. Misophrioida (=Propodoplea Lang, 1948) is the most basal Podoplean order. The order Cyclopoida Burmeister, 1835, is paraphyletic and now encompasses Poecilostomatoida Thorell, 1859, as a sister to the family Schminkepinellidae Martinez Arbizu, 2006. Within Harpacticoida Sars, 1903, both sections, Polyarthra Lang, 1948, and Oligoarthra Lang, 1948, are monophyletic, but not sister groups. The order Canuelloida is proposed while maintaining the order Harpacticoida s. str. (Oligoarthra). Cyclopoida, Harpacticoida and Cyclopinidae are redefined, while Canuelloida ordo. nov., Smirnovipinidae fam. nov. and Cyclopicinidae fam. nov are proposed as new taxa.
Background. Climate change concurrent with anthropogenic disturbances can initiate serial changes that reverberate up the food chain with repercussions for fisheries. To date, there is no information available concerning the combined effects of global warming and human impacts on tropical marine food webs. While temperate copepods respond differently to warming and environmental stressors, the extent to which tropical copepods can adapt to rising temperature of already warm waters remains unknown. We hypothesize that sea warming and other anthropogenic disturbances over the long term will have the greatest impact on the copepod community in nearshore waters where their effects are accentuated, and therefore vulnerable and resilient species could be identified. Methods. Zooplankton samples were collected during two time periods (1985-86 and 2014-15) interposed by marked anthropogenic disturbances, and at the same five stations located progressively from inshore to offshore in Klang Strait, Malaysia, following the asymmetrical before-after-control-impact (BACI) design. Copepods were identified to species, and results were interpreted by univariate (ANOVA) and multivariate (PERMANOVA, PCO) analyses of the computed species abundance and diversity measures. Results. Copepod total abundance was not significantly different among stations but higher after disturbance than before disturbance. However, changes in the abundance of particular species and the community structure between time periods were dramatic. Coastal large-bodied calanoid species (e.g., Acartia spinicauda, Calanopia thompsoni, Pseudodiaptomus bowmani and Tortanus forcipatus) were the most vulnerable group to disturbance. This however favored the opportunistic species (e.g., Oithona simplex, O. attenuata, Hemicyclops sp., Pseudomacrochiron sp. and Microsetella norvegica). Small-bodied copepods (e.g., Paracalanus sp., Parvocalanus crassirostris and Euterpina acutifrons) were unaffected. Centropages tenuiremis was likely an introduced species. There was no significant loss in species richness of copepods despite the dramatic changes in community structure. Discussion. Sea warming and other human-induced effects such as eutrophication, acidification and coastal habitat degradation are likely the main factors that have altered copepod community structure. The large-bodied estuarine and coastal calanoid copepods are surmised to be vulnerable to eutrophication and hypoxia, while both resilient and opportunistic species are largely unaffected by, or adaptable to, degraded coastal environments and observed sea surface temperature (SST) rise. It is forecasted that SST rise with unmitigated anthropogenic impacts will further reduce large-bodied copepod species the favoured food for fish larvae with dire consequences for coastal fish production.
Assessing the impact of global warming on the food web of the North Atlantic will require difficult-to-obtain physiological data on a key copepod crustacean, Calanus finmarchicus. The de novo transcriptome presented here represents a new resource for acquiring such data. It was produced from multiplexed gene libraries using RNA collected from six developmental stages: embryo, early nauplius (NI-II), late nauplius (NV-VI), early copepodite (CI-II), late copepodite (CV) and adult (CVI) female. Over 400,000,000 paired-end reads (100 base-pairs long) were sequenced on an Illumina instrument, and assembled into 206,041 contigs using Trinity software. Coverage was estimated to be at least 65%. A reference transcriptome comprising 96,090 unique components (“comps”) was annotated using Blast2GO. 40% of the comps had significant blast hits. 11% of the comps were successfully annotated with gene ontology (GO) terms. Expression of many comps was found to be near zero in one or more developmental stages suggesting that 35 to 48% of the transcriptome is “silent” at any given life stage. Transcripts involved in lipid biosynthesis pathways, critical for the C. finmarchicus life cycle, were identified and their expression pattern during development was examined. Relative expression of three transcripts suggests wax ester biosynthesis in late copepodites, but triacylglyceride biosynthesis in adult females. Two of these transcripts may be involved in the preparatory phase of diapause. A key environmental challenge for C. finmarchicus is the seasonal exposure to the dinoflagellate Alexandrium fundyense with high concentrations of saxitoxins, neurotoxins that block voltage-gated sodium channels. Multiple contigs encoding putative voltage-gated sodium channels were identified. They appeared to be the result of both alternate splicing and gene duplication. This is the first report of multiple NaV1 genes in a protostome. These data provide new insights into the transcriptome and physiology of this environmentally important zooplankter.