3D intrusions transport active surface microbial assemblages to the dark ocean.

Subtropical oceans contribute significantly to global primary production, but the fate of the picophytoplankton that dominate in these low-nutrient regions is poorly understood. Working in the subtropical Mediterranean, we demonstrate that subduction of water at ocean fronts generates 3D intrusions with uncharacteristically high carbon, chlorophyll, and oxygen that extend below the sunlit photic zone into the dark ocean. These contain fresh picophytoplankton assemblages that resemble the photic-zone regions where the water originated. Intrusions propagate depth-dependent seasonal variations in microbial assemblages into the ocean interior. Strikingly, the intrusions included dominant biomass contributions from nonphotosynthetic bacteria and enrichment of enigmatic heterotrophic bacterial lineages. Thus, the intrusions not only deliver material that differs in composition and nutritional character from sinking detrital particles, but also drive shifts in bacterial community composition, organic matter processing, and interactions between surface and deep communities. Modeling efforts paired with global observations demonstrate that subduction can flux similar magnitudes of particulate organic carbon as sinking export, but is not accounted for in current export estimates and carbon cycle models. Intrusions formed by subduction are a particularly important mechanism for enhancing connectivity between surface and upper mesopelagic ecosystems in stratified subtropical ocean environments that are expanding due to the warming climate.

Deep-pelagic fishes: Demographic instability in a stable environment.

Demographic histories are frequently a product of the environment, as populations expand or contract in response to major environmental changes, often driven by changes in climate. Meso- and bathy-pelagic fishes inhabit some of the most temporally and spatially stable habitats on the planet. The stability of the deep-pelagic could make deep-pelagic fishes resistant to the demographic instability commonly reported in fish species inhabiting other marine habitats, however the demographic histories of deep-pelagic fishes are unknown. We reconstructed the historical demography of 11 species of deep-pelagic fishes using mitochondrial and nuclear DNA sequence data. We uncovered widespread evidence of population expansions in our study species, a counterintuitive result based on the nature of deep-pelagic ecosystems. Frequency-based methods detected potential demographic changes in nine species of fishes, while extended Bayesian skyline plots identified population expansions in four species. These results suggest that despite the relatively stable nature of the deep-pelagic environment, the fishes that reside here have likely been impacted by past changes in climate. Further investigation is necessary to better understand how deep-pelagic fishes, by far Earth's most abundant vertebrates, will respond to future climatic changes.

Incorporating mesopelagic fish into the evaluation of conservation areas for marine living resources under climate change scenarios.

Mesopelagic fish (meso-fish) are central species within the Southern Ocean (SO). However, their ecosystem role and adaptive capacity to climate change are rarely integrated into protected areas assessments. This is a pity given their importance as crucial prey and predators in food webs, coupled with the impacts of climate change. Here, we estimate the habitat distribution of nine meso-fish using an ensemble model approach (MAXENT, random forest, and boosted regression tree). Four climate model simulations were used to project their distribution under two representative concentration pathways (RCP4.5 and RCP8.5) for short-term (2006-2055) and long-term (2050-2099) periods. In addition, we assess the ecological representativeness of protected areas under climate change scenarios using meso-fish as indicator species. Our models show that all species shift poleward in the future. Lanternfishes (family Myctophidae) are predicted to migrate poleward more than other families (Paralepididae, Nototheniidae, Bathylagidae, and Gonostomatidae). In comparison, lanternfishes were projected to increase habitat area in the eastern SO but lose area in the western SO; the opposite was projected for species in other families. Important areas (IAs) of meso-fish are mainly distributed near the Antarctic Peninsula and East Antarctica. Negotiated protected area cover 23% of IAs at present and 38% of IAs in the future (RCP8.5, long-term future). Many IAs of meso-fish still need to be included in protected areas, such as the Prydz Bay and the seas around the Antarctic Peninsula. Our results provide a framework for evaluating protected areas incorporating climate change adaptation strategies for protected areas management. Supplementary Information: The online version contains supplementary material available at 10.1007/s42995-023-00188-9.

A new genus and species of oceanic planktonic Tisbidae (Crustacea, Copepoda, Harpacticoida) with enlarged modified eyes.

Both sexes of a new monotypic genus of Tisbidae (Copepoda, Harpacticoida) are described from the epi- or mesopelagic zone in the Kuroshio region, Japan. Gyoromeguttatumgen. et sp. nov. belongs to a monophyletic lineage of deepwater holoplanktonic genera defined by a suite of characters. Within this clade, Gyoromegen. nov. appears most closely related to Neotisbella Boxshall, 1979. The most distinguishable feature of G.guttatumgen. et sp. nov. is the presence of large, paired, frontal modified eyes, each consisting of a baculiform ocellus, a globular (Gicklhorn's?) organ, and a semi-parabolic plate. The taxonomic position of Tisbespinulosa Bradford & Wells, 1983 is discussed and a key to the six meso- and bathypelagic tisbid species is provided. Confusion surrounding earlier literature reports of supernumerary elements on the caudal ramus in some harpacticoid taxa is clarified. Secondary modifications of ocellar components of the typical naupliar eye in the Harpacticoida are reviewed. It is suggested that the development of specialized eyes in G.guttatumgen. et sp. nov. may provide a means for detecting bioluminescent food particles in oligotrophic pelagic environments. The large, vaulted prosome indicates the species is an opportunistic macrophage that has adopted gorging as a feeding strategy.

Life in the Midwater: The Ecology of Deep Pelagic Animals.

The water column of the deep ocean is dark, cold, low in food, and under crushing pressures, yet it is full of diverse life. Due to its enormous volume, this mesopelagic zone is home to some of the most abundant animals on the planet. Rather than struggling to survive, they thrive-owing to a broad set of adaptations for feeding, behavior, and physiology. Our understanding of these adaptations is constrained by the tools available for exploring the deep sea, but this tool kit is expanding along with technological advances. Each time we apply a new method to the depths, we gain surprising insights about genetics, ecology, behavior, physiology, diversity, and the dynamics of change. These discoveries show structure within the seemingly uniform habitat, limits to the seemingly inexhaustible resources, and vulnerability in the seemingly impervious environment. To understand midwater ecology, we need to reimagine the rules that govern terrestrial ecosystems. By spending more time at depth-with whatever tools are available-we can fill the knowledge gaps and better link ecology to the environment throughout the water column.

Biological information on a rare pelagic fish, black ruff Centrolophus niger, caught in Icelandic waters: Distribution, feeding, and otoliths.

Black ruff (Centrolophus niger) is a rare and poorly studied species found in both the Atlantic and Pacific Oceans and also in the Mediterranean Sea. It is sporadically caught south of Iceland during the annual International Ecosystem Summer Survey of the Nordic Seas. In total, 43 specimens were caught from 2009 to 2021, of which 41 specimens were caught during 2017-2021. All specimens, except one, were caught using a pelagic trawl (cod-end mesh-size: 50 mm) close to the surface (trawl depth: 0-35 m) with in situ temperature ranging from 9 to 13°C. The area south of Iceland is characterized by having warmer temperatures than other areas around the island, which might be indicative of a northern limit for the distribution of black ruff. The fish were primarily in the range of 29-46 cm with a few larger individuals up to 71 cm. Fourteen fish, caught in 2017 and 2021, were dissected to gather biological information on this species. These fish were all juveniles with no obvious sign of gonad development. Correlations between total length, fork length, and standard length are presented. Otoliths were thin and delicate with a length of ~13-16 mm, and otolith size (length, width, and area) was correlated with fish size. Much of the stomach content was at an advanced stage of digestion, but some contents could be identified and consisted of invertebrates, primarily of the orders Amphipoda and Calanoida with some unidentified fish also present.

Reproductive biology of the electric lanternfish Electrona risso (Myctophidae) and the bigscale fishes Melamphaes polylepis and Scopelogadus mizolepis (Melamphaidae).

This study was the first to investigate the key reproductive traits of the electric lantern fish Electrona risso (Myctophidae, n = 918) and the bigscale fishes (Melamphaidae) Melamphaes polylepis (n = 260) and Scopelogadus mizolepis (n = 649). Specimens of these mesopelagic species were collected in March and April 2015 in the eastern Central Atlantic (0-24° N, 20-26° W). Sex ratio was not significantly different from 1:1 in E. risso and M. polylepis but significantly skewed toward female dominance in S. mizolepis. Reproductive phases were determined macroscopically and by histological analyses on selected individuals. Female length at 50% maturity (L50 ) was 55.1 mm standard length (LS ) in E. risso, with an observed female maximum length (Lmax ) of 81.2 mm LS . M. polylepis females had an L50 of 40.2 mm LS and an Lmax of 86.7 mm LS . S. mizolepis had an L50 of 46 mm LS and an Lmax of 97.9 mm LS . The three species show histological features of iteroparity, but the E. risso population appears to occur in two year-classes and experience only one spawning season per lifetime in the study region. All three species are batch-spawners. A batch fecundity of 2668 eggs was estimated from one E. risso individual, with a relative batch fecundity of 369 eggs g-1 gonad-free body mass. M. polylepis had a batch fecundity of 1027 eggs and a relative batch fecundity of 149 eggs g-1 (n = 3). S. polylepis had a batch fecundity of 1545 eggs and a relative batch fecundity of 215 eggs g-1 (n = 21). The median gonado-somatic index during the actively spawning phase of E. risso was 4.5, significantly lower than that of M. polylepis (7.5) and S. mizolepis (7.1). No regressing or regenerating phases were observed in this study. Batch-spawning in all three species is suggested to be advantageous to cope with intra-annual variability in food supply and other risks for offspring survival. With what appears to be in effect a (facultative) semelparous strategy in combination with a short life span in E. risso, interannual differences would have a great effect on population dynamics of this species. Knowledge is still lacking on temporal aspects of reproduction such as the duration of the spawning season and the frequency of spawning, as well as age and growth.

Increased prokaryotic diversity in the Red Sea deep scattering layer.

BACKGROUND: The diel vertical migration (DVM) of fish provides an active transport of labile dissolved organic matter (DOM) to the deep ocean, fueling the metabolism of heterotrophic bacteria and archaea. We studied the impact of DVM on the mesopelagic prokaryotic diversity of the Red Sea focusing on the mesopelagic deep scattering layer (DSL) between 450-600 m. RESULTS: Despite the general consensus of homogeneous conditions in the mesopelagic layer, we observed variability in physico-chemical variables (oxygen, inorganic nutrients, DOC) in the depth profiles. We also identified distinct seasonal indicator prokaryotes inhabiting the DSL, representing between 2% (in spring) to over 10% (in winter) of total 16S rRNA gene sequences. The dominant indicator groups were Alteromonadales in winter, Vibrionales in spring and Microtrichales in summer. Using multidimensional scaling analysis, the DSL samples showed divergence from the surrounding mesopelagic layers and were distributed according to depth (47% of variance explained). We identified the sources of diversity that contribute to the DSL by analyzing the detailed profiles of spring, where 3 depths were sampled in the mesopelagic. On average, 7% was related to the epipelagic, 34% was common among the other mesopelagic waters and 38% was attributable to the DSL, with 21% of species being unique to this layer. CONCLUSIONS: We conclude that the mesopelagic physico-chemical properties shape a rather uniform prokaryotic community, but that the 200 m deep DSL contributes uniquely and in a high proportion to the diversity of the Red Sea mesopelagic.

Metagenome-assembled genomes from particle-associated microbial communities in the mesopelagic zone of the Pacific Ocean.

We report 10 particle-associated metagenome-assembled genomes (MAGs) from the mesopelagic zone of Pacific Ocean seawaters. MAGs comprise members of Flavobacteria Halomonas, Blastomonas, Brevundimonas, Alteromonas, Shingomonas, Sphingopyxis, Tabrizicola, Proteobacteria, and Gammaproteobacteria. Functional annotation suggests that these bacteria are involved in central particulate organic carbon conversion, nitrogen cycling, and phosphorus cycling.

Large sensory volumes enable Southern elephant seals to exploit sparse deep-sea prey.

The ability of echolocating toothed whales to detect and classify prey at long ranges enables efficient searching and stalking of sparse prey in these time-limited dives. However, nonecholocating deep-diving seals such as elephant seals appear to have much less sensory advantage over their prey. Both elephant seals and their prey rely on visual and hydrodynamic cues that may be detectable only at short ranges in the deep ocean, leading us to hypothesize that elephant seals must adopt a less efficient reactive mode of hunting that requires high prey densities. To test that hypothesis, we deployed high-resolution sonar and movement tags on 25 females to record simultaneous predator and prey behavior during foraging interactions. We demonstrate that elephant seals have a sensory advantage over their prey that allows them to potentially detect prey 5 to 10 s before striking. The corresponding prey detection ranges of 7 to 17 m enable stealthy approaches and prey-specific capture tactics. In comparison, prey react at a median range of 0.7 m, close to the neck extension range of striking elephant seals. Estimated search swathes of 150 to 900 m2 explain how elephant seals can locate up to 2,000 prey while swimming more than 100 km per day. This efficient search capability allows elephant seals to subsist on prey densities that are consonant with the deep scattering layer resources estimated by hydroacoustic surveys but which are two orders of magnitude lower than the prey densities needed by a reactive hunter.

The deep dive of organohalogen compounds: Bioaccumulation in the top predators of mesopelagic trophic webs, pygmy and dwarf sperm whales, from the Southwestern Atlantic ocean.

Kogia sima and Kogia breviceps are apex predators of mesopelagic trophic webs being far from most anthropogenic threats. However, chemical pollutants and naturally synthesized compounds may travel long distances. This study aimed to use kogiid whales as sentinels of mesopelagic trophic webs in the Southwestern Atlantic Ocean. Persistent organic pollutants (POPs), e.g., polychlorinated biphenyls (PCBs), dichlorodiphenyltrichloroethane (DDT) and metabolites, mirex, hexachlorobenzene (HCB), polybrominated diphenylethers (PBDEs), pentabromoethylbenzene (PBEB) and hexabromobenzene (HBB), and the naturally produced methoxylated BDE (MeO-BDEs) were determined in the blubber of 16 K. sima and 15 K. breviceps. Among the organochlorine compounds, DDTs were the main group found in K. sima and in K. breviceps (1636.6 and 3983.3 ng g-1 lw, respective medians), followed by PCBs (425.9 and 956.1 ng g-1 lw, respectively), mirex (184.1 and 375.6 ng g-1 lw, respectively), and HCB (132.4 and 340.3 ng g-1 lw, respectively). As for the organobromine, the natural MeO-BDEs were predominant (1676.7 and 501.6 ng g-1 lw, respectively), followed by PBDEs (13.6 and 10.3 ng g-1 lw, respectively) and PBEB (2.2 and 2.9 ng g-1 lw, respectively). In general, POPs concentration was higher in K. breviceps than in K. sima. Conversely, MeO-BDEs concentration was higher in K. sima than in K. breviceps. Differences in concentrations in these sympatric odontocetes were attributed to distinct species, sampling sites, and biological parameters and suggest some level of niche segregation. It is noteworthy the long-range reach and bioaccumulation of these synthetic compounds in an unexplored habitat, that present an increasing economic interest.

The Outsized Role of Salps in Carbon Export in the Subarctic Northeast Pacific Ocean.

Periodic blooms of salps (pelagic tunicates) can result in high export of organic matter, leading to an "outsized" role in the ocean's biological carbon pump (BCP). However, due to their episodic and patchy nature, salp blooms often go undetected and are rarely included in measurements or models of the BCP. We quantified salp-mediated export processes in the northeast subarctic Pacific Ocean in summer of 2018 during a bloom of Salpa aspera. Salps migrated from 300 to 750 m during the day into the upper 100 m at night. Salp fecal pellet production comprised up to 82% of the particulate organic carbon (POC) produced as fecal pellets by the entire epipelagic zooplankton community. Rapid sinking velocities of salp pellets (400-1,200 m d-1) and low microbial respiration rates on pellets (<1% of pellet C respired day-1) led to high salp pellet POC export from the euphotic zone-up to 48% of total sinking POC across the 100 m depth horizon. Salp active transport of carbon by diel vertical migration and carbon export from sinking salp carcasses was usually <10% of the total sinking POC flux. Salp-mediated export markedly increased BCP efficiency, increasing by 1.5-fold the proportion of net primary production exported as POC across the base of the euphotic zone and by 2.6-fold the proportion of this POC flux persisting 100 m below the euphotic zone. Salps have unique and important effects on ocean biogeochemistry and, especially in low flux settings, can dramatically increase BCP efficiency and thus carbon sequestration.

The photic-aphotic divide is a strong ecological and evolutionary force determining the distribution of ciliates (Alveolata, Ciliophora) in the ocean.

The bulk of knowledge on marine ciliates is from shallow and/or sunlit waters. We studied ciliate diversity and distribution across epi- and mesopelagic oceanic waters, using DNA metabarcoding and phylogeny-based metrics. We analyzed sequences of the 18S rRNA gene (V4 region) from 369 samples collected at 12 depths (0-1000 m) at the Bermuda Atlantic Time-series Study site of the Sargasso Sea (North Atlantic) monthly for 3 years. The comprehensive depth and temporal resolutions analyzed led to three main findings. First, there was a gradual but significant decrease in alpha-diversity (based on Faith's phylogenetic diversity index) from surface to 1000-m waters. Second, multivariate analyses of beta-diversity (based on UniFrac distances) indicate that ciliate assemblages change significantly from photic to aphotic waters, with a switch from Oligotrichea to Oligohymenophorea prevalence. Third, phylogenetic placement of sequence variants and clade-level correlations (EPA-ng and GAPPA algorithms) show Oligotrichea, Litostomatea, Prostomatea, and Phyllopharyngea as anti-correlated with depth, while Oligohymenophorea (especially Apostomatia) have a direct relationship with depth. Two enigmatic environmental clades include either prevalent variants widely distributed in aphotic layers (the Oligohymenophorea OLIGO5) or subclades differentially distributed in photic versus aphotic waters (the Discotrichidae NASSO1). These results settle contradictory relationships between ciliate alpha-diversity and depth reported before, suggest functional changes in ciliate assemblages from photic to aphotic waters (with the prevalence of algivory and mixotrophy vs. omnivory and parasitism, respectively), and indicate that contemporary taxon distributions in the vertical profile have been strongly influenced by evolutionary processes. Integration of DNA sequences with organismal data (microscopy, functional experiments) and development of databases that link these sources of information remain as major tasks to better understand ciliate diversity, ecological roles, and evolution in the ocean.

Vertical distribution and acoustic characteristics of deep water micronektonic crustacean in the Bay of Biscay.

Vertical distribution of meso- and bathypelagic crustacean are scarcely known. The logistics involved in their studies hinder an adequate assessment of their role in the deep ecosystems. As a result, the literature on zooplankton scattering models is mainly focused on epipelagic organisms, particularly krill species. This study analyses data of the plankton community classified by family from the surface down to 2000 m taken in the Bay of Biscay, but focusses on the meso- and bathypelagic zone. Photographic data was employed to obtain a micronektonic crustacean shape catalogue. The Distorted Wave Born Approximation (DWBA) model was employed to estimate target strength. Pasiphaeidae, Euphausiidae and Acanthephyridae were mainly distributed above 500 m depth, while Benthesicymidae, Sergestidae and Mysidae were concentrated in the lower mesopelagic to upper bathypelagic area. The most abundant species were Euphausiidae and Benthesicymidae with up to 30 and 40 individuals per cubic meter respectively. Standard length ranged from 8 to 85 mm and was significantly related with height but not with depth. The Pasiphaeidae family presented the largest individuals followed by Acanthephyridae and Sergestidae while Euphausiidae, Benthesicymidae, and Mysidae were shorter. An smooth fluid-like response was estimated for shorter organisms, while individuals of 60 mm or higher present TS oscillations from around 60 kHz. Pasiphaeidae present an almost 10 dB higher TS than Sergestidae, Acanthephyridae and Benthesicymidae while Mysidae followed by Euphausiidae produce the lower TS. Simple models of TS values at broadside versus the logarithm of standard length (SL) that can be employed as an approximation of their scattering are provided for four common frequencies (TS = 58.5*log10(SL)-188.7, TS = 57.03*log10(SL)-174.1, TS = 22.48*log10(SL)-157.14, TS = 17.55*log10(SL)-135 and TS = 10.53*log10(SL)-109 at 18, 38, 70, 120 and 200 kHz respectively). Changes in body density and sound speed contrast may increase by 10 or 2 dB the resulting TS respectively but are constant in phase, while orientation can decrease the TS by up to 20 dB at the higher frequencies and alter the spectra to an almost flat trend. This study provides further insight into the vertical distribution and physical characteristics of the micronektonic crustacean families inhabiting the Bay of Biscay down to 2000 m depth. It also estimates their echo from a real-shape catalogue that can be employed to infer knowledge from acoustic recordings, particularly of the lower mesopelagic and the bathypelagic zones.

Bioactivity Profiling and Untargeted Metabolomics of Microbiota Associated with Mesopelagic Jellyfish Periphylla periphylla.

The marine mesopelagic zone extends from water depths of 200 m to 1000 m and is home to a vast number and diversity of species. It is one of the least understood regions of the marine environment with untapped resources of pharmaceutical relevance. The mesopelagic jellyfish Periphylla periphylla is a well-known and widely distributed species in the mesopelagic zone; however, the diversity or the pharmaceutical potential of its cultivable microbiota has not been explored. In this study, we isolated microorganisms associated with the inner and outer umbrella of P. periphylla collected in Irminger Sea by a culture-dependent approach, and profiled their chemical composition and biological activities. Sixteen mostly gram-negative bacterial isolates were selected and subjected to an OSMAC cultivation regime approach using liquid and solid marine broth (MB) and glucose-yeast-malt (GYM) media. Their ethyl acetate (EtOAc) extracts were assessed for cytotoxicity and antimicrobial activity against fish and human pathogens. All, except one extract, displayed diverse levels of antimicrobial activities. Based on low IC50 values, four most bioactive gram-negative strains; Polaribacter sp. SU124, Shewanella sp. SU126, Psychrobacter sp. SU143 and Psychrobacter sp. SU137, were prioritized for an in-depth comparative and untargeted metabolomics analysis using feature-based molecular networking. Various chemical classes such as diketopiperazines, polyhydroxybutyrates (PHBs), bile acids and other lipids were putatively annotated, highlighting the biotechnological potential in P. periphylla-associated microbiota as well as gram-negative bacteria. This is the first study providing an insight into the cultivable bacterial community associated with the mesopelagic jellyfish P. periphylla and, indeed, the first to mine the metabolome and antimicrobial activities of these microorganisms.

Influencing factors for microplastic intake in abundant deep-sea lanternfishes (Myctophidae).

Plastic debris is ubiquitous in the hydrosphere. Yet, we lack an understanding of contamination among deep-sea species and primarily how each trait can influence microplastic intake. We investigated microplastic contamination in the digestive tract of hyper-abundant mesopelagic lanternfishes (n = 364 individuals) from the Southwestern Tropical Atlantic, captured from 90 to 1000 m depth. Overall, microplastics were detected in most individuals analysed (frequency of occurrence = 68 %). Large microplastics, mostly of a filamentous shape were the most frequent, followed by smaller fragments and foams. Microplastics made of high-density polymers (PET, PVC, PA, SBR rubber) were more prevalent than low-density ones (PE, EVA and PBD rubber), especially under deeper layers. Larger microplastics were detected in lanternfishes captured off the northeastern Brazilian coast (mean 0.88 ± SE 0.06 mm) compared to those from around the Rocas Atoll and Fernando de Noronha Archipelago (0.70 ± 0.07 mm; p≤ 0.05), ∼350 km from the continent. Moreover, lanternfishes that migrate from the upper mesopelagic (200-500 m) to the epipelagic layers (<200 m) had simultaneously the highest intake and the smallest particles (1.65 ± 0.17 particles individual-1 and 0.55 ± 0.07 mm; p≤ 0.05). Biological mediated transport of microplastics from the epipelagic to the mesopelagic waters was evinced, but fishes foraging in shallower layers had the lowest intake (1.11 ± 0.10 part. ind.-1; p≤ 0.05). Furthermore, the jaw length was positively associated with an increment in microplastic intake (Incidence Rate Ratio = 1.1; p≤ 0.05). The lanternfishes that preferably prey upon fish larvae are more prone to microplastic intake than their counterparts, which forage mostly on crustaceans and gelatinous zooplankton (p≤ 0.05).

Particles act as 'specialty centers' with expanded enzymatic function throughout the water column in the western North Atlantic.

Heterotrophic bacteria initiate the degradation of high molecular weight organic matter by producing an array of extracellular enzymes to hydrolyze complex organic matter into sizes that can be taken up into the cell. These bacterial communities differ spatially and temporally in composition, and potentially also in their enzymatic complements. Previous research has shown that particle-associated bacteria can be considerably more active than bacteria in the surrounding bulk water, but most prior studies of particle-associated bacteria have been focused on the upper ocean - there are few measurements of enzymatic activities of particle-associated bacteria in the mesopelagic and bathypelagic ocean, although the bacterial communities in the deep are dependent upon degradation of particulate organic matter to fuel their metabolism. We used a broad suite of substrates to compare the glucosidase, peptidase, and polysaccharide hydrolase activities of particle-associated and unfiltered seawater microbial communities in epipelagic, mesopelagic, and bathypelagic waters across 11 stations in the western North Atlantic. We concurrently determined bacterial community composition of unfiltered seawater and of samples collected via gravity filtration (>3 µm). Overall, particle-associated bacterial communities showed a broader spectrum of enzyme activities compared with unfiltered seawater communities. These differences in enzymatic activities were greater at offshore than at coastal locations, and increased with increasing depth in the ocean. The greater differences in enzymatic function measured on particles with depth coincided with increasing differences in particle-associated community composition, suggesting that particles act as 'specialty centers' that are essential for degradation of organic matter even at bathypelagic depths.

Microbial functional diversity across biogeochemical provinces in the central Pacific Ocean.

Enzymes catalyze key reactions within Earth's life-sustaining biogeochemical cycles. Here, we use metaproteomics to examine the enzymatic capabilities of the microbial community (0.2 to 3 µm) along a 5,000-km-long, 1-km-deep transect in the central Pacific Ocean. Eighty-five percent of total protein abundance was of bacterial origin, with Archaea contributing 1.6%. Over 2,000 functional KEGG Ontology (KO) groups were identified, yet only 25 KO groups contributed over half of the protein abundance, simultaneously indicating abundant key functions and a long tail of diverse functions. Vertical attenuation of individual proteins displayed stratification of nutrient transport, carbon utilization, and environmental stress. The microbial community also varied along horizontal scales, shaped by environmental features specific to the oligotrophic North Pacific Subtropical Gyre, the oxygen-depleted Eastern Tropical North Pacific, and nutrient-rich equatorial upwelling. Some of the most abundant proteins were associated with nitrification and C1 metabolisms, with observed interactions between these pathways. The oxidoreductases nitrite oxidoreductase (NxrAB), nitrite reductase (NirK), ammonia monooxygenase (AmoABC), manganese oxidase (MnxG), formate dehydrogenase (FdoGH and FDH), and carbon monoxide dehydrogenase (CoxLM) displayed distributions indicative of biogeochemical status such as oxidative or nutritional stress, with the potential to be more sensitive than chemical sensors. Enzymes that mediate transformations of atmospheric gases like CO, CO2, NO, methanethiol, and methylamines were most abundant in the upwelling region. We identified hot spots of biochemical transformation in the central Pacific Ocean, highlighted previously understudied metabolic pathways in the environment, and provided rich empirical data for biogeochemical models critical for forecasting ecosystem response to climate change.

Importance of the Lunar Cycle on Mesopelagic Foraging by Atlantic Bluefin Tuna in the Upwelling Area of the Strait of Messina (Central Mediterranean Sea).

The influence of the lunar cycle on bluefin tuna foraging in the upwelling area of the Strait of Messina was investigated by exploring trophic interaction with mesopelagic fish and cephalopod prey. To focus on how the lunar cycle could affect availability of mesopelagic prey for this predator, we tested potential differences in the diet related to each lunar phase. Moreover, we considered two potential impacts of the lunar cycle: the lunar irradiance and the strength of currents. Overall, 2672 prey items were mesopelagic fish and cephalopods, representing 60.7% of overall diet by number. The main mesopelagic fish prey items were lanternfishes and dragonfishes, while Onychoteuthis banksii was the most important cephalopod prey. In summary, the Strait of Messina has highly specific hydrodynamic and biological features which strongly depend on upwelling currents, which in turn are influenced by the lunar cycle (new and full moon with strong currents, quarters with fewer currents). Upwelling causes water mixing, bringing to the surface a large amount of mesopelagic fauna which become more readily available to tuna. Lunar irradiance contributes to the variation of prey composition, increasing the success of visual predation on mesopelagic resources at high light in the water column.

Length-weight relationships of 55 mesopelagic fishes from the eastern tropical North Atlantic: Across- and within-species variation (body shape, growth stanza, condition factor).

We present estimates of length-weight relationships (LWRs) of 55 mesopelagic fish species of 13 taxonomic families based on data collected in the eastern tropical North Atlantic (ETNA) in March/April 2015. Our data include novel records for 19 species, while for 25 species LWRs are based on the most robust sample sizes, and for 21 species they are based on the most representative size ranges available up to now. In 31 species, body lengths were within the maximum range of body lengths recorded in the area, with new records of maximum lengths for 13 species. Most values for b fell between 2.5 and 3.5 with a mean exponent b of 3.08 (median 3.12) and a mean a of 0.0172 (median 0.0113). Body shape as covariate ('elongated', 'fusiform' and 'short-deep') strongly determined the variation in log a as a function of parameter b. For the mesopelagic fish species investigated, the form factor a3.0 indicated a significant increase of median a3.0 from 'elongated' to 'fusiform' to 'short-deep' body shapes. Large variability existed in parameter b between species of the same taxonomic family. Isometric growth was indicated in only nine species, whereas a positive allometry was suggested in 22 species. Using segmented regression analysis, we investigated ontogenetic variation in LWRs in 30 species. Of these, 20 species showed a breakpoint in LWR, whereby nearly equal numbers exhibited an increase or a decrease in slope following the breakpoint. Seven out of nine species showed significant regional variation in the slope of the relationship of the relative condition factor Krel vs. body length between two or more regions of the ETNA [eastern and western part of the oxygen minimum zone (LO-E, LO-W), northern and central equatorial region (EQ-N, EQ-C)]. A conspicuous pattern was an increase in Krel with body size in the LO-E (in six out of eight species), whereas in the LO-W and the equatorial regions the majority of species showed a related decrease. These findings support the idea that growth patterns in mesopelagic fishes in tropical regions show species-specific ecological niche and life-history adaptations that are finely tuned to small-scale regional environmental conditions. Comparison of our data with those of other studies emphasises that, regarding the small adult sizes of many mesopelagic fish species, estimates of LWR parameters are strongly influenced by sampled size distributions.