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This study investigated the biogeography, the presence and diversity of potentially harmful taxa harbored, and potential interactions between and within bacterial and eukaryotic domains of life on plastic debris in the Mediterranean. Using a combination of high-throughput DNA sequencing (HTS), Causal Network Analysis, and Scanning Electron Microscopy (SEM), we show regional differences and gradients in the Mediterranean microbial communities associated with marine litter, positive causal effects between microbes including between and within domains of life, and how these might impact the marine ecosystems surrounding them. Adjacent seas within the Mediterranean region showed a gradient in the microbial communities on plastic with non-overlapping endpoints (Adriatic and Ligurian Seas). The largest predicted inter-domain effects included positive effects of a novel red-algal Plastisphere member on its potential microbiome community. Freshwater and marine samples housed a diversity of fungi including some related to disease-causing microbes. Algal species related to those responsible for Harmful Blooms (HABs) were also observed on plastic pieces including members of genera not previously reported on Plastic Marine Debris (PMD).

Summary Microalgae play a major role as primary producers in aquatic ecosystems. Cell signalling regulates their interactions with the environment and other organisms, yet this process in phytoplankton is poorly defined. Using the marine planktonic diatom Pseudo‐nitzschia multistriata, we investigated the cell response to cues released during sexual reproduction, an event that demands strong regulatory mechanisms and impacts on population dynamics.We sequenced the genome of P. multistriata and performed phylogenomic and transcriptomic analyses, which allowed the definition of gene gains and losses, horizontal gene transfers, conservation and evolutionary rate of sex‐related genes. We also identified a small number of conserved noncoding elements.Sexual reproduction impacted on cell cycle progression and induced an asymmetric response of the opposite mating types. G protein‐coupled receptors and cyclic guanosine monophosphate (cGMP) are implicated in the response to sexual cues, which overall entails a modulation of cell cycle, meiosis‐related and nutrient transporter genes, suggesting a fine control of nutrient uptake even under nutrient‐replete conditions.The controllable life cycle and the genome sequence of P. multistriata allow the reconstruction of changes occurring in diatoms in a key phase of their life cycle, providing hints on the evolution and putative function of their genes and empowering studies on sexual reproduction.

The Gulf of Naples is a dynamical area with intense exchanges between offshore oligotrophic and coastal eutrophic waters with frequent freshwater inputs. The Sarno River, one of the most polluted rivers in Europe, strongly contributes to the pollution of the area, discharging high amounts of heavy metals and organic wastes from heavily cultivated and industrial areas. This paper reports on the diversity and community structure of the marine residential Bacteria and Archaea of the Gulf of Naples in an area close to the river Sarno plume and investigates their small-scale taxonomic diversity and expression patterns as a proxy of potential metabolic activity using metagenomics and metatranscriptomics. Bacteria and Archaea were mainly represented by marine clades, with only minor contributors from freshwater ones. The community was dominated by Alpha- and Gammaproteobacteria, of which Rhodospirillales, Pelagibacteriales, and Oceanospirilalles were most represented. However, Alteromonadales and Rhodobacterales were the most active, despite their relative lower abundance, suggesting that they are important for overall ecosystem functioning and nutrient cycling. Nitrification and a reversed form of dissimilatory sulfate reduction were the major metabolic processes found in the metatrascriptomes and were mainly associated to Nitrosopumilales and Pelagibacter, respectively. No clear indication of transcripts related to stress induced by heavy metals or organic pollutants was found. In general, despite the high loads of pollutants discharged continuously by the Sarno River, the microbial community did not show marks of stress-induced changes neither structural nor functional, thus suggesting that this river has little or no effect on the planktonic bacterial community of the Gulf of Naples.

High-throughput sequencing of Prasinovirus DNA polymerase and host green algal (Mamiellophyceae) ribosomal RNA genes was used to analyse the diversity and distribution of these taxa over a ∼10 000 km latitudinal section of the Indian Ocean. New viral and host groups were identified among the different trophic conditions observed, and highlighted that although unknown prasinoviruses are diverse, the cosmopolitan algal genera Bathycoccus, Micromonas and Ostreococcus represent a large proportion of the host diversity. While Prasinovirus communities were correlated to both the geography and the environment, host communities were not, perhaps because the genetic marker used lacked sufficient resolution. Nevertheless, analysis of single environmental variables showed that eutrophic conditions strongly influence the distributions of both hosts and viruses. Moreover, these communities were not correlated, in their composition or specific richness. These observations could result from antagonistic dynamics, such as that illustrated in a prey–predator model, and/or because hosts might be under a complex set of selective pressures. Both of these reasons must be considered to interpret environmental surveys of viruses and hosts, because covariation does not always imply interaction.

Bacterioplankton are fundamental components of marine ecosystems and influence the entire biosphere by contributing to the global biogeochemical cycles of key elements. Yet, there is a significant gap in knowledge about their diversity and specific activities, as well as environmental factors that shape their community composition and function. Here, the distribution and diversity of surface bacterioplankton along the coastline of the Gulf of Naples (GON; Italy) were investigated using flow cytometry coupled with high-throughput sequencing of the 16S rRNA gene. Heterotrophic bacteria numerically dominated the bacterioplankton and comprised mainly Alphaproteobacteria , Gammaproteobacteria , and Bacteroidetes . Distinct communities occupied river-influenced, coastal, and offshore sites, as indicated by Bray-Curtis dissimilarity, distance metric (UniFrac), linear discriminant analysis effect size (LEfSe), and multivariate analyses. The heterogeneity in diversity and community composition was mainly due to salinity and changes in environmental conditions across sites, as defined by nutrient and chlorophyll a concentrations. Bacterioplankton communities were composed of a few dominant taxa and a large proportion (92%) of rare taxa (here defined as operational taxonomic units [OTUs] accounting for IMPORTANCE The study of bacterioplankton in coastal zones is of critical importance, considering that these areas are highly productive and anthropogenically impacted. Their richness and evenness, as well as their potential activity, are very important to assess ecosystem health and functioning. Here, we investigated bacterial distribution, community composition, and potential metabolic activity in the GON, which is an ideal test site due to its heterogeneous environment characterized by a complex hydrodynamics and terrestrial inputs of varied quantities and quality. Our study demonstrates that bacterioplankton communities in this region are highly diverse and strongly regulated by a combination of different environmental factors leading to their heterogeneous distribution, with the rare taxa contributing to a major proportion of diversity and shifts in community composition and potentially holding a key role in ecosystem functioning.

This paper presents a winter carbon budget for the northern Adriatic Sea, obtained through direct measurements during two multidisciplinary cruises and literature data. A box model approach was adopted to integrate estimates of stocks and fluxes of carbon species over the total area. The oligotrophy at the basin scale and the start of primary productivity well before the onset of spring stratification were observed. In winter, the system underwent a complete reset, as the mixing of water masses erased any signal of previous hypoxia or anoxia episodes. The northern Adriatic Sea was phosphorus depleted with respect to C and N availability. This fact confirms the importance of mixing with deep-sea water for P supply to biological processes on the whole. Despite the abundant prokaryotic biomass, the microbial food web was less efficient in organic C production than phytoplankton. In the upper layer, the carbon produced by primary production exceeded the fraction respired by planktonic community smaller than 200 µm. On the contrary, respiration processes prevailed in the water column below the pycnocline. The carbon budget also proved that the northern Adriatic Sea can be an effective sink for atmospheric CO2 throughout the entire winter season.

Egg production rates and/or hatching success in the copepods Acartia clausi, Calanus helgolandicus and Temora longicornis were negatively affected by a late spring (May-June 2003) phytoplankton bloom in the North Adriatic Sea, dominated mainly by the large diatom Cerataulina pelagica. Highest total concentrations of 3.3- 104 cells-ml(-1) were located in the vicinity of the Po River, which also corresponded to the area where the highest numbers of phaeophorbides were measured (0.779, 0.528 and 0.419 mu g.l(-1), respectively, compared to an average of the remaining stations of 0.183 +/- 0.049 SD), suggesting some grazing on the bloom. Phytoplankton biomass in terms of carbon was dominated by diatoms, representing on average 42% of total phytoplankton carbon and more than 80% at several stations. Cerataulina pelagica, Cyclotella spp., Chaetoceros spp. and small unidentified centric diatoms dominated the diatom community numerically but C. pelagica was by far the dominant diatom in terms of carbon due to its large cell size. This species represented more than 60% of the diatom biomass at nine of the 14 stations sampled, and was absent only at one station, which was the most offshore station sampled during the cruise. Although polyunsaturated aldehydes (PUAs) were not detected, other oxylipins which are hydroxy and keto derivatives of eicosapentaenoic and docosahexaenoic acids that affect copepod reproduction were found in these samples. Hence, we can attribute the negative impact of diatoms not only to PUAs, as previously believed, but also to these compounds. This is the first direct evidence of the presence of oxylipins other than PUAs in marine blooms dominated by diatoms.

Photophysiological parameters provide useful insights into the effects of environmental forcings on phytoplankton physiology. We present data on the short-term photoacclimative responses of phytoplankton throughout the water column during a diel sampling (every 1.5–3 h for 33 h) at a coastal station in the Gulf of Naples (Italy) in November 1996. Liposoluble pigments (high performance liquid chromatography), variable fluorescence (Prim-Prod probe) and picoplankton cell counts and autofluorescence (flow cytometry) were investigated every 1.5–3 h over a period of 33 h. The phytoplankton was phased to the alternation of light and dark and also showed acclimation to the different light intensities. Photoprotective pigments were synthesized during the day at the surface (0–20 m) and were significantly correlated with light intensity changes, as well as with the effective quantum yield of fluorescence. At night, recovery of photosystems from excess light was observed as was redistribution of nutrients and algae due to vertical convective motions caused by thermal dissipation. Equations linking photobiological parameters and time or light evolution were inferred to obtain kinetic coefficients. These were then used as biological tracers of vertical mixing whose velocity in the surface layer was estimated to be ,0.05 cm s 21 .

Phytoplankton pigment diversity and photoacclimation during the natural day-night cycle was investigated at a fixed location in the Strait of Sicily in July 1997 using HPLC pigment analysis on fractionated samples ( 3 µm) and flow cytometry. Picophytoplankton dominated phytoplankton biomass in terms of chl a with an average value of 57% and was mainly represented by prokaryotes, prymnesiophytes and pelagophytes. Prochlorococcus and picoeukaryotes con- tributed equally to the picophytoplankton in terms of chl a, but Prochlorococcus were numerically more abundant and were represented by 2 ecotypes, one replacing the other according to depth. Larger phytoplankton were dominated by prymnesiophytes and diatoms. Photoacclimation was evi- dent from changes in pigment content and strongly increased with depth. The deep chlorophyll max- imum (DCM), present between 75 and 90 m, showed a diverse and rich phytoplankton community with the 2 size classes almost equally represented. Growth rates of Prochlorococcus and Synechococ- cus, estimated from cell cycle measurements, were 0.67 and 0.41 d -1 , respectively, at 75 m. Only Prochlorococcus was able to sustain a good growth rate of 0.43 d -1 at the base of the DCM (90 m) where only 0.5% of incident light was available. Light-shift experiments using onboard incubated natural seawater showed much faster kinetic coefficients for acclimation in picophytoplankton than in larger algae. In general, the data describe the dynamics of picophytoplankton and its light adapta- tion through the water column and in the DCM, and can be considered representative of stable sum- mer conditions in the Mediterranean Sea.

Most diatom species produce at least two classes of oxylipins, polyunsaturated aldehydes (PUAs) and non-volatile oxylipins (NVOs). These compounds can act as grazer deterrents by reducing the reproductive success of copepod grazers. The aim of the present study was to better understand the impact of late-winter diatom blooms and of diatom-derived PUAs and NVOs on the reproduction of two dominant copepod species in the northern Adriatic Sea, in two consecutive years. Higher diatom abundance in 2005 compared to 2004 coincided with higher egg-production rates in both Acartia clausi and Calanus helgolandicus. The increased contribution of diatoms to the phytoplankton assemblage in 2005 also led to a higher phytoplankton production of NVOs, whereas PUA levels were low in both years. These three-times-higher oxylipin levels correlated with a decrease in average hatching success in both A. clausi and C helgolandicus from similar to 80% in 2004 to similar to 60% in 2005, indicating that the observed negative effects on copepod recruitment in the latter year were related to the greater production of diatom NVOs. Our findings reveal the importance of these metabolites and offer an explanation as to why diatom blooms may be more toxic for copepods in some years compared to others. (C) 2015 Elsevier B.V. All rights reserved.

Aspects of phytoplankton photophysiology were investigated in a coastal area of the Mediterranean Sea (Gulf of Naples) using high performance liquid chromatography pigment analysis, flow cytometric cellular fluorescence information and in vivo pump and probe measurements. Discrete sampling, continuous profiles and on-board kinetic experiments were used to estimate photoacclimation kinetics and to identify the time scales of these reactions in phytoplankton. Based on the relationship between the depth of the euphotic zone and the mean chlorophyll concentration, the area could be classified as Case 1 water, although, oligotrophy was not reflected in phytoplankton photosynthetic efficiencies, suggesting that it represented a transient feature induced by an eddy, originating offshore. Fronts at the edge of the eddy exhibited different photophysiological features, in which the phytoplankton showed greater physiological stress, brought about by the convergent motion within the front. Kinetic experiments allowed identification of time scales for photoacclimation. Using photoprotective pigments as tracers we estimated vertical mixing velocities in the upper layer of 0.07 cm s � 1 and from this we inferred kinetic coefficients for different photophysiological processes in the mixed layer as well as significant threshold values for photoacclimation rates to be 4% h � 1 . From the threshold value, the vertical eddy diffusivity in the area at the time of sampling was estimated as 1.75 � 10 � 2 m 2 s � 1 , a high value for vertical mixing.

Diatoms are able to react to biotic and abiotic stress, such as competition, predation and unfavorable growth conditions, by producing bioactive compounds including polyunsaturated aldehydes (PUAs). PUAs have been shown to act against grazers and either enhance or inhibit the growth of different phytoplankton and bacteria both in culture and in the field. Presence of nanomolar concentrations of dissolved PUAs in seawater has been reported in the North Adriatic Sea (Mediterranean), suggesting that these compounds are released in seawater following diatom cell lysis. However, the origin of the PUAs and their effects on natural phytoplankton assemblages remain unclear. Here we present data from four oceanographic cruises that took place during diatom blooms in the northern Adriatic Sea where concentrations of particulate and dissolved PUAs were monitored along with phytoplankton cell lysis. Cell lysis was positively correlated with both concentrations of particulate and dissolved PUAs (R = 0.69 and R = 0.77, respectively), supporting the hypothesis that these compounds are released by cell lysis. However, the highest concentration of dissolved PUAs (2.53 nM) was measured when cell lysis was high (0.24 d(-1)) but no known PUA-producing diatoms were detected, suggesting either that other organisms can produce PUAs or that PUA-producing enzymes retain activity extracellularly after diatom cells have lysed. Although in situ concentrations of dissolved PUAs were one to three orders of magnitude lower than those typically used in laboratory culture experiments, we argue that concentrations produced in the field could induce similar effects to those observed in culture and therefore may help shape plankton community composition and function in the oceans.

The effect of light on the synchronization of cell cycling was investigated in several strains of the oceanic photosynthetic prokaryote Prochlorococcus using flow cytometry. When exposed to a light-dark (L-D) cycle with an irradiance of 25 μmol of quanta · m −2 s −1 , the low-light-adapted strain SS 120 appeared to be better synchronized than the high-light-adapted strain PCC 9511. Submitting L-D-entrained populations to shifts (advances or delays) in the timing of the “light on” signal translated to corresponding shifts in the initiation of the S phase, suggesting that this signal is a key parameter for the synchronization of population cell cycles. Cultures that were shifted from an L-D cycle to continuous irradiance showed persistent diel oscillations of flow-cytometric signals (light scatter and chlorophyll fluorescence) but with significantly reduced amplitudes and a phase shift. Complete darkness arrested most of the cells in the G 1 phase of the cell cycle, indicating that light is required to trigger the initiation of DNA replication and cell division. However, some cells also arrested in the S phase, suggesting that cell cycle controls in Prochlorococcus spp. are not as strict as in marine Synechococcus spp. Shifting Prochlorococcus cells from low to high irradiance translated quasi-instantaneously into an increase of cells in both the S and G 2 phases of the cell cycle and then into faster growth, whereas the inverse shift induced rapid slowing of the population growth rate. These data suggest a close coupling between irradiance levels and cell cycling in Prochlorococcus spp.

The effects of different water masses and their physical structures on phytoplankton and bacteria were investigated in a coastal area of the Mediterranean Sea, the Gulf of Naples (Italy) on 9 and 10 November 1995. A small eddy (8 km in diameter), generated from offshore waters. occupied the inner part of the Gulf at surface, while the Modified Atlantic Water (MAW) intruded between 60 and 80 m depth. Phytoplankton communities as identified by HPLC pigment analysis (total) and flow cytometry (ultraphytoplankton) showed peculiarities in the different subsystems. Cryptophytes domlnated by pigment biomass were everywhere, while diatoms were more abundant at the coast and nanoflagellates at offshore stations. In the MAW, a peculiar phytoplankton community, different to that in the other water masses, occurred, and 2 populations of prochlorophytes were distinguished based on scatter and red fluorescence observations. Total bacterial numbers were significantly correlated to chlorophyll a (chl a) concentrations. Different populations of heterotrophic bacteria were identified based on apparent DN.4 content. Percentages of 2 such populations were significantly correlated to total chl a. Therefore, phytoplankton appears to regulate not only bacterial quantity but also composition. Although our study area was characterized by a marked oligotrophy, heterotrophic bacterial biomass never exceeded autotrophic biomass, as derived using constant conversion factors of bacteria and chl a to C. Therefore, no inverted pyramid was observed and bacterial biomass was still controlled by phytoplankton biomass. Mesoscale variability is of great relevance for energy and matter budgrts of coastal areas and its study is therefore fundamental. For example, eddies can be very important in coastal areas, influencing distribution of nutrients and organisms. Our approach, couphng HPLC and flow cytometry, represents a valuable tool for obtaining information on both phytoplankton coniposition and ecophysiology on a large number of samples, necessary for a valid representation of the space scales involved.

Phytoplankton abundance, species composition and primary production of the Gulf of Naples were investigated during an autumn bloom in November 1985. Hydrographic data and surface phytoplankton samples were collected during a 3 day cruise, whereas investigations on in situ primary production and phytoplankton vertical distribution were conducted from a second boat on three different dates. A coast-offshore gradient was recorded for most of the chemical and biological parameters analysed. At stations closer to the coast, which were affected by land run-off, phytoplankton populations attained concentrations of 2.5 x lO* cells P'. with a marked dominance of diatoms belonging to the genera Thalassiosira and Chaetoceros. The most striking character of the system was a remarkably high carbon assimilation rate (up to 1.04 g C m'2 day"1) at stations closer to the coast. The causative mechanism for this bloom appeared to be land-derived nutrient enrichment, possibly enhanced by autumn rains, followed by a period of favourable meteorological conditions, which occurs recurrently in the region for a brief period around November, known locally as "St Martin's Summer'. We hypothesize that a similar mechanism can stimulate phytoplankton growth more than once every year, since high-stability periods following meteoro- logical perturbations can occur several times in temperate regions of the northern hemisphere in autumn.

The present paper reviews the literature related to the life cycle of the prymnesiophyte Phaeocystis and its controlling factors and proposes novel hypotheses based on unpublished observations in culture and in the field. We chiefly refer to P. globosa Scherffel as most of the observations concern this species. P. globosa exhibits a complex alternation between several types of free-living cells (non-motile, flagellates, microzoopores and possibly macrozoospores) and colonies for which neither forms nor pathways have been completely identified and described. The different types of Phaeocystis cells were reappraised on the basis of existing microscopic descriptions complemented by unpublished flow cytometric investigations. This analysis revealed the existence of at least three different types of free-living cells identified on the basis of a combination of size, motility and ploidy characteristics: non-motile cells, flagellates and microzoospores. Their respective function within Phaeocystis life cycle, and in particular their involvement in colony formation is not completely understood. Observational evidence shows that Phaeocystis colonies are initiated at the early stage of their bloom each by one free-living cell. The mechanisms controlling this cellular transformation are still uncertain due to the lack of information on the overwintering Phaeocystis fomms and on the cell type responsible for colony induction. The existence of haploid microzoospores released from senescent colonies gives however some support to sexuality involvement at some stages of colony formation. Once colonies are formed, at least two mechanisms were identified as responsible of the spreading of colony form: colony multiplication by colonial division or budding and induction of new colony from colonial cells released in the external medium after colony disruption. The latter mechanism was clearly identified, involving at least two successive cell differentiations in the following sequence: motility development, subsequent flagella loss and settlement to a surface, mucus secretion and colony formation, colonial cell division and colony growth. Aggregate formation, cell motility development and subsequent emigration from the colonies, release of non-motile cells after colony lysis on the other hand, were identified as characteristics for termination of Phaeocystis colony development. These pathways were shown to occur similarly in natural environments. In the early stages of the bloom however, many recently-formed colonies were found on the setae of Chaetoceros spp, suggesting this diatom could play a key-role in Phaeocystis bloom inception. Analysis of the possible environmental factors regulating the transition between the different phases of the life cycle, suggested that nutrient status and requirement of a substrate for attachment of free-living cells would be essential for initiation of the colonial form. Physical constraints obviously would be important in determining colony shape and fragmentation although autogenic factors cannot be excluded. Some evidence exists that nutrients regulate colony division, while temperature and nutrient stress would stimulate cell emigration from the colonies.

Eric Karsenti et al.-- 5 pages, 2 figures, With biology becoming quantitative, systems-level studies can now be performed at spatial scales ranging from molecules to ecosystems. Biological data generated consistently across scales can be integrated with physico-chemical contextual data for a truly holistic approach, with a profound impact on our understanding of life [1]–[5]. Marine ecosystems are crucial in the regulation of Earth's biogeochemical cycles and climate [6],[7]. Yet their organization, evolution, and dynamics remain poorly understood [8],[9]. The Tara Oceans project was launched in September 2009 for a 3-year study of the global ocean ecosystem aboard the ship Tara. A unique sampling programme encompassing optical and genomic methods to describe viruses, bacteria, archaea, protists, and metazoans in their physico-chemical environment has been implemented. Starting as a grassroots initiative of a few scientists, the project has grown into a global consortium of over 100 specialists from diverse disciplines, including oceanography, microbial ecology, genomics, molecular, cellular, and systems biology, taxonomy, bioinformatics, data management, and ecosystem modeling. This multidisciplinary community aims to generate systematic, open access datasets usable for probing the morphological and molecular makeup, diversity, evolution, ecology, and global impacts of plankton on the Earth system, We are keen to thank the commitment of the following people and sponsors who made this singular expedition possible: CNRS, EMBL, Genoscope/CEA, VIB, Stazione Zoologica Anton Dohrn, UNIMIB, ANR (projects POSEIDON, BIOMARKS, PROMETHEUS, and TARA-GIRUS), FWO, BIO5, Biosphere 2, agnès b., the Veolia Environment Foundation, Region Bretagne, World Courier, Illumina, Cap L'Orient, the EDF Foundation EDF Diversiterre, FRB, the Prince Albert II de Monaco Foundation, Etienne Bourgois, the Tara schooner and its captain and crew. Tara Oceans would not exist without the continuous support of the participating 23 institutes (see http://oceans.taraexpeditions.org). Special thanks also goes to Fabrice Not and Noan Le Bescot for creating the figures of this manuscript. This is contribution number 0001 of the Tara Oceans Expedition 2009–2012

Marine chemical ecology comprises the study of the production and interaction of bioactive molecules affecting organism behavior and function. Here we focus on bioactive compounds and interactions associated with phytoplankton, particularly bloom-forming diatoms, prymnesiophytes and dinoflagellates. Planktonic bioactive metabolites are structurally and functionally diverse and some may have multiple simultaneous functions including roles in chemical defense (antipredator, allelopathic and antibacterial compounds), and/or cell-to-cell signaling (e. g., polyunsaturated aldehydes (PUAs) of diatoms). Among inducible chemical defenses in response to grazing, there is high species-specific variability in the effects on grazers, ranging from severe physical incapacitation and/or death to no apparent physiological response, depending on predator susceptibility and detoxification capability. Most bioactive compounds are present in very low concentrations, in both the producing organism and the surrounding aqueous medium. Furthermore, bioactivity may be subject to synergistic interactions with other natural and anthropogenic environmental toxicants. Most, if not all phycotoxins are classic secondary metabolites, but many other bioactive metabolites are simple molecules derived from primary metabolism (e. g., PUAs in diatoms, dimethylsulfoniopropionate (DMSP) in prymnesiophytes). Producing cells do not seem to suffer physiological impact due to their synthesis. Functional genome sequence data and gene expression analysis will provide insights into regulatory and metabolic pathways in producer organisms, as well as identification of mechanisms of action in target organisms. Understanding chemical ecological responses to environmental triggers and chemically-mediated species interactions will help define crucial chemical and molecular processes that help maintain biodiversity and ecosystem functionality.

Diatoms have evolved a silicified cell wall that provides an efficient barrier against herbivores. These microalgae also produce chemical compounds such as polyunsaturated aldehydes (PUAs) that can potentially impair recruitment and cause malformations in the offspring of such grazers. We measured silica content as an indication of cell wall thickness, organic nutrient cell quotas, PUAs and polyunsaturated fatty acid cell content in Skeletonema marinoi grown under N-, P- and Si-limitation in continuous cultures. A 7.5 fold increase in PUA production was observed (27.5 fmol cell(-1)) in Si-limited cells with respect to the controls, while Si content decreased by 50%. PUA production decreased in nitrogen-limited cells to 0.14 fmol cell(-1) and increased 3 times in phosphorus-limited cells (7.53 fmol cell(-1)), while silica content increased by 20% in both cases. The substrate to product ratio suggested that production of the PUAs heptadienal and octadienal was limited by the amount of substrate under P- and Si-limitation, and by enzyme activity under N-limitation. Octatrienal production was likely limited by enzyme activity in all growth conditions. The high PUA levels produced under Si-limitation, when cells have a thinner cell wall, suggests a compensatory alternation of mechanical and chemical defense mechanisms.

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This essay is the outcome of a colloquium convened in November 2005 at the Benthos Laboratory of the Stazione Zoologica Anton Dohrn in Ischia, Italy, on chemical ecology and the role of secondary metabolites in the structuring and functioning of marine biodiversity. The participants of the workshop are part of the European Network of Excellence MarBEF (Marine Biodiversity and Ecosystem Function), a consortium of 56 European marine institutes to integrate and disseminate knowledge and expertise on marine biodiversity. Here we review some of the new trends and emerging topics in marine chemical ecology. The first section deals with microbial chemical interactions. Microbes communicate with each other using diffusible molecules such as N-acylhomoserine lactones (AHL). These are regulators in cell-density-dependent gene regulation (quorum sensing) controlling microbial processes. In chemical interactions with higher organisms, microbes can act either as harmful pathogens that are repelled by the host’s chemical defense or as beneficial symbionts. These symbionts are sometimes the true producers of the host’s secondary metabolites that have defensive and protective functions for their hosts. We also describe how allelochemicals can shape phytoplankton communities by regulating competition for available resources, and also interactions among individuals of the same species. Compounds such as the diatom-derived unsaturated aldehydes have been demonstrated to act as info chemicals, and they possibly function as a diffusible bloom-termination signal that triggers an active cell death and bloom termination at sea. The same molecules have also been shown to interfere with the reproductive capacity of grazing animals deterring future generations of potential predators. Such compounds differ from those that act as feeding deterrents since they do not target the predator but its offspring. Many of the neurotoxins produced by dinoflagellates act as feeding deterrents, and laboratory experiments have shown that ingestion of these algae by some microzooplankton and macrozooplankton can cause acute, responses such as death, incapacitation, altered swimming behavior, and reduced fecundity and egg-hatching success. These effects may rarely occur in nature because of low individual grazing rates on dinoflagellate cells and grazing on other food sources such as microflagellates and diatoms. We also consider the nutritional component of marine plant-herbivore interactions, especially in the plankton, and the information available on the effects of growing conditions of algae on the production of toxic metabolites. Species producing saxitoxins seem to consistently produce the highest amounts of toxins (on a per cell basis) in the exponential phase of growth, and there is a decrease in their production under nitrogen, but not under phosphorus stress, where the production actually increases. We try to explain the circumstances under which organisms defend themselves chemically and argue that the most likely explanatory model for the production of secondary metabolites used for defense in planktonic organisms is the carbon nutrient balance hypothesis, which predicts that most algae produce their toxins mainly under conditions where carbon is in excess and nitrogen (or other nutrients) is limiting. We also discuss chemically mediated macroalgal-herbivore interactions in the benthos and the large variation in concentration of seaweed defense metabolites at different spatial and temporal scales. Seaweeds have been shown to produce a large variety of secondary metabolites with highly variable chemical structures such as terpenoids, acetogenins, amino acid derivates, and polyphenols. Many of these compounds probably have multiple simultaneous functions for the seaweeds and can act as allelopathic, antimicrobial, and antifouling or ultraviolet-screening agents, as well as herbivore deterrents. We also provide examples of interactions between marine benthic invertebrates, especially sponges, molluscs, and cnidarians, that are mediated by specific secondary metabolites and discuss the role of these in shaping benthic communities.

Diatoms are an important group of eukaryotic phytoplankton, responsible for about 20% of global primary productivity. Study of the functional role of chemical signaling within phytoplankton assemblages is still in its infancy although recent reports in diatoms suggest the existence of chemical-based defense strategies. Here, we demonstrate how the accurate perception of diatom-derived reactive aldehydes can determine cell fate in diatoms. In particular, the aldehyde (2E, 4E/Z)-decadienal (DD) can trigger intracellular calcium transients and the generation of nitric oxide ( NO) by a calcium-dependent NO synthase-like activity, which results in cell death. However, pretreatment of cells with sublethal doses of aldehyde can induce resistance to subsequent lethal doses, which is reflected in an altered calcium signature and kinetics of NO production. We also present evidence for a DD-derived NO-based intercellular signaling system for the perception of stressed bystander cells. Based on these findings, we propose the existence of a sophisticated stress surveillance system in diatoms, which has important implications for understanding the cellular mechanisms responsible for acclimation versus death during phytoplankton bloom successions.

Paffenhöfer, G. A. ... et al.-- 13 pages, 1 table From 3 to 6 November 2002, a colloquium was convened at the Benthos Laboratory of the Stazione Zoologica Anton Dohrn on Ischia, Italy, with the goal of evaluating the present status of the effects of diatoms on their main consumers, planktonic copepods, and to develop future research strategies to enhance our understanding of such interactions. These included toxic effects of diatom metabolites on copepods, particularly reproduction, and nutritional effects of diatoms on juvenile to adult copepods. Key issues involved in the impact of diatoms on the dynamics of natural plankton communities in situ were also addressed. During the plenary session, the most recent advances on this topic were presented. The plenary session was followed by 3 working groups on (1) production of aldehydes by phytoplankton, (2) toxic and nutritional effects of diatoms on zooplankton, and (3) the chemistry of diatom defense, as well as of their nutritional quality. These working groups focused on suggesting future research needs for the different topics. As a result, several recommendations were outlined, including experimental studies. It became evident that interdisciplinary efforts are needed, involving chemists, oceanographers and experimentalists, since many of the biological observations under controlled conditions and in situ require an integrated approach, including chemical causation. Extensive field observations based on common protocols are also recommended for investigation of the intrinsic variability of such effects and their environmental controls. Laboratory experiments are seen to be essential for the full understanding of environmentally occurring processes

Ecophysiology Laboratory, Stazione Zoologica A. Dohrn, Villa Comunale, 80121 Naples, Italy. ianora@szn.it The growth cycle in nutrient-rich, aquatic environments starts with a diatom bloom that ends in mass sinking of ungrazed cells and phytodetritus. The low grazing pressure on these blooms has been attributed to the inability of overwintering copepod populations to track them temporally. We tested an alternative explanation: that dominant diatom species impair the reproductive success of their grazers. We compared larval development of a common overwintering copepod fed on a ubiquitous, early-blooming diatom species with its development when fed on a typical post-bloom dinoflagellate. Development was arrested in all larvae in which both mothers and their larvae were fed the diatom diet. Mortality remained high even if larvae were switched to the dinoflagellate diet. Aldehydes, cleaved from a fatty acid precursor by enzymes activated within seconds after crushing of the cell, elicit the teratogenic effect. This insidious mechanism, which does not deter the herbivore from feeding but impairs its recruitment, will restrain the cohort size of the next generation of early-rising overwinterers. Such a transgenerational plant-herbivore interaction could explain the recurringly inefficient use of a predictable, potentially valuable food resource--the spring diatom bloom--by marine zooplankton.

[1] The relationship between phytoplankton distribution and dynamics, and the resident water masses in the Ionian Sea was investigated in the spring of 1999 by flow cytometry, HPLC, microscopy, 14C incorporation, and serial dilutions. More than 50% of total biomass was represented by ultraphytoplankton, with proportions increasing with depth (up to 80%) and eastward (up to 60%). Synechococcus sp. cyanobacteria dominated ultraphytoplankton numbers and biomass. Hydrological stability explained the relative vertical distribution of Synechococcus sp. and Prochlorococcus sp. cyanobacteria and their degree of photoacclimation. The northwestern area (“Italian Side”) was in a transition from bloom to oligotrophic conditions, reflected in high biological instability in terms of phytoplankton composition, photoacclimative properties, and photosynthetic efficiency (P/B of 11.40 mg C mg chl−1 h−1). The influence of the Adriatic Surface Water, carrying a trace of the spring bloom was visible from satellite imagery (SeaWiFS). The eastern part of the sampled area (“Greek Side,” GS) was hydrologically more stable (low estimated vertical velocities), resulting in higher photosynthetic efficiency (P/B of 20.47 mg C mg chl−1 h−1) and phytoplankton photoacclimative properties. The Atlantic Water (AW) was the most oligotrophic in terms of nutrient concentration, chla, and productivity (208.0 mg C m−2 d−1), but also the most variable in terms of mesoscale features. Growth of Synechococcus sp. and Prochlorococcus sp. was slow (0.36 d−1 at max), while picoeukaryotes grew well in the deep chlorophyll maximum (DCM) of the GS (0.67 d−1) Picoeukaryote growth was tightly coupled with grazing (0.80 d−1), indicating efficient biomass recycling. Biological characterization and DCM dynamics in the three areas are discussed in relation to physical features present at the time of sampling.