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During polar nights of 2012 and 2017, bioluminescence (BL) potential surveys demonstrated high emissions at depths around and below 100 m at offshore stations to the north of a Svalbard fjord (Rijpfjorden). We demonstrated that the highest bioluminescent emissions for offshore stations are located at depths below depths of modelled/ambient light intensities corresponding to the reported irradiance thresholds for the behavioral light sensitivity of krill and copepods, and suggest that behavioral light sensitivity is one of the reasons for high values of BL potential observed below 100 m at offshore stations. In order to understand sources of bioluminescent taxa responsible for the observed high values of BL potential in offshore waters, we have investigated the origin and pathways of water masses circulating to the north, offshore of the fjord Rijpfjorden by using a hydrodynamic model. For both 2012 and 2017, the model water masses mostly originate from the west, where the Atlantic water is flowing northward, then along the shelf and shelf slope of northern Svalbard, and to the offshore of the fjord. This indicates that the advection of zooplankton by North Atlantic Water is one possible source of bioluminescent organisms offshore of northern Svalbard. In 2012, water masses also originated from the inflow through Hinlopen trench and strait, while, in 2017, the offshore water was advected and upwelled into the fjord on time scales less than 10 days, and after that there was a recirculation back from the fjord to offshore on time scales larger than 10 days. This recirculation from the fjord might be another source of bioluminescent organisms in the offshore waters.

Larval stomatopods have generally been described as having a typical larval crustacean compound eye, which lacks the visual pigment diversity and morphological specializations of the well-studied stomatopod adult eye. However, recent work has suggested that larval stomatopod eyes are more complex than previously described. In this study, we provide physiological and behavioral evidence of at least three distinct photoreceptor classes in three species of larval stomatopods: Gonodactylellus n. sp., Gonodactylaceus falcatus and Pullosquilla n. sp. First, electroretinogram recordings were used to measure the spectral sensitivity of each species. Evidence for at least three spectral classes were identified in each: an ultraviolet, peaking at 340–376 nm; a short-wavelength blue, peaking at 455–464 nm; and a long-wavelength orange, peaking at 576–602 nm. Next, the behavioral response to light was investigated. We found that each species demonstrated positive phototactic responses to monochromatic stimuli across the UV–visible spectrum. In wavelength preference trials, distinct preferences among species were identified when different colored light stimuli were presented simultaneously. All species displayed a strong response to the UV stimulus, as well as responses to blue and orange stimuli, although at different response strengths, but no response to green. The results of this study demonstrate that larval stomatopods not only have multiple physiologically active spectral classes but they also display clear and distinct responses to wavelengths across the spectrum. We propose that the spectral classes demonstrated in each are related to visually guided ecological tasks of the larvae, which may differ between species.

During polar nights in January 2012 and 2017, significantly higher bioluminescence (BL) potential emissions in the upper 50 m were observed in the fjord Rijpfjorden (Svalbard, Norway) in comparison to offshore stations (located on the shelf-break, shelf-slope areas and in the deeper water). The objective of this paper is to better understand why, during two polar nights (separated by 5 years), the values of BL potential in the northern Svalbard fjord are higher than at offshore stations, and what the role of advection is in observed elevated BL potential values in the top 50 m of the fjord. To address the above objective, we applied the same BL potential modeling approach and strategies during polar nights for both 2012 and 2017. For both years, advection of BL potential from offshore (including upwelling along the shelf, shelf-slope) produced an increase of BL potential in the fjord area, in spite of the introduction of mortality in bioluminescent organisms. Observations of BL potential indicated high emissions at depths below 100 m at offshore stations for both polar nights. Our modeling studies demonstrated that these high values of BL potential below 100 m are upwelled and advected to the top 50 m of the fjord. We demonstrated that upwelling and advection of these deep high BL potential values (and therefore, upwelling and advection of corresponding bioluminescent taxa) from offshore areas are dominant factors in observed BL potential dynamics in the top 50 m in the fjord.

Bioluminescence (BL) potential observations registered high BL potential emissions during the polar night of January 2012 at the mouth of a fjord Rijpfjorden (northern Svalbard, Norway). Notably, observations of BL potential at this location were significantly higher in the upper 50 m than observed BL potential at offshore stations located on the shelf-slope areas and in the deeper water off northern Svalbard. In the present paper, we address questions as to why the values of BL potential in the fjord are higher than at offshore stations and what the role of advection is in the observed elevation of BL potential values in the fjord in comparison to offshore stations. We utilized the ensemble approach when changes in BL potential are modeled with the advection-diffusion-source (ADS) model, and the focus is on modeling and predictions of a large ensemble of averaged values of BL potential over a specific domain of interest at the fjord mouth. Results of the modeling have demonstrated increases in ensemble members with BL potential values larger than or equal to the specific threshold in comparison to corresponding ensemble members in the initial distribution. Even when we introduced mortality rates in our simulations (values from 0.1 to 0.3 day−1), increases in ensemble members with higher than the threshold value were more than 3.7 times larger in comparison to ensemble members in the initial distribution. The interpretation of our results is that during the polar night, the fjord represents an area where bioluminescent organisms from offshore aggregate through advection and mixing.

Background The Bethesda System for Reporting Thyroid Cytopathology (TBSRTC) comprises 6 categories used for the diagnosis of thyroid fine-needle aspiration biopsy (FNAB). Each category has an associated risk of malignancy, which is important in the management of a thyroid nodule. More accurate predictions of malignancy may help to reduce unnecessary surgery. A machine learning algorithm (MLA) was developed to evaluate thyroid FNAB via whole slide images (WSIs) to predict malignancy. Methods Files were searched for all thyroidectomy specimens with preceding FNAB over 8 years. All cytologic and surgical pathology diagnoses were recorded and correlated for each nodule. One representative slide from each case was scanned to create a WSI. An MLA was designed to identify follicular cells and predict the malignancy of the final pathology. The test set comprised cases blindly reviewed by a cytopathologist who assigned a TBSRTC category. The area under the receiver operating characteristic curve was used to assess the MLA performance. Results Nine hundred eight FNABs met the criteria. The MLA predicted malignancy with a sensitivity and specificity of 92.0% and 90.5%, respectively. The areas under the curve for the prediction of malignancy by the cytopathologist and the MLA were 0.931 and 0.932, respectively. Conclusions The performance of the MLA in predicting thyroid malignancy from FNAB WSIs is comparable to the performance of an expert cytopathologist. When the MLA and electronic medical record diagnoses are combined, the performance is superior to the performance of either alone. An MLA may be used as an adjunct to FNAB to assist in refining the indeterminate categories.

The ArcLight observatory provides hourly continuous time series of light regime data (intensity, spectral composition, and photoperiod) from the Arctic, Svalbard at 79° N. Until now, no complete annual time series of biologically relevant light has been provided from the high Arctic due to insufficient sensitivity of commercial light sensors during the Polar Night. We describe a camera system providing all-sky images and the corresponding integrated spectral irradiance ( E P A R ) in energy or quanta units, throughout a complete annual cycle. We present hourly–diel–annual dynamics from 2017 to 2020 of irradiance and its relation to weather conditions, sun and moon trajectories.

Microplastic (MP) in estuarine and coastal environments remains poorly characterized, despite the importance of these physically dynamic regions as a buffer between land, freshwater environments, and the open ocean where plastic debris accumulates. We sampled MP particles to determine concentration, size, and type in Delaware Bay and numerically simulated transport and distribution at a high spatiotemporal resolution of positively buoyant particles, representing common MP types. Baywide MP concentrations averaged between 0.19 and 1.24 pieces m-3 depending on size fraction (300-1000 and 1000-5000 μm) and sampling month (April and June 2017). Upper bay stations, which are located in or near the estuarine turbidity maximum, had higher MP concentrations than lower bay and New Jersey shore stations. Fragments were predominately polyethylene, and filaments predominately polypropylene. Model results suggest that buoyant particles quickly (i.e., within hours) organize in patchy, highly inhomogeneous distributions, creating "hot spots" of MP. In the presence of variable currents driven by buoyancy, wind, and tides, we predict high spatial and temporal variability of MP distributions in Delaware Bay; MP concentrations could vary by a factor of 1000 within a tidal cycle at our sample locations. Collectively, these observations and simulations provide a baseline of MP concentrations in Delaware Bay along with broader, contextual understanding for how measurements reflect MP concentrations in a dynamic estuarine system.

Stomatopod crustaceans have among the most complex eyes in the animal kingdom, with up to twelve different color detection channels. The capabilities of these unique eyes include photoreception of ultraviolet (UV) wavelengths (

In situ observations of pelagic fish and zooplankton with optical instruments usually rely on external light sources. However, artificial light may attract or repulse marine organisms, which results in biased measurements. It is often assumed that most pelagic organisms do not perceive the red part of the visible spectrum and that red light can be used for underwater optical measurements of biological processes. Using hull-mounted echosounders above an acoustic probe or a baited video camera, each equipped with light sources of different colours (white, blue and red), we demonstrate that pelagic organisms in Arctic and temperate regions strongly avoid artificial light, including visible red light (575–700 nm), from instruments lowered in the water column. The density of organisms decreased by up to 99% when exposed to artificial light and the distance of avoidance varied from 23 to 94 m from the light source, depending on colours, irradiance levels and, possibly, species communities. We conclude that observations from optical and acoustic instruments, including baited cameras, using light sources with broad spectral composition in the 400–700 nm wavelengths do not capture the real state of the ecosystem and that they cannot be used alone for reliable abundance estimates or behavioural studies.

The predation risk of many aquatic taxa is dominated by visually searching predators, commonly a function of ambient light. Several studies propose that changes in visual predation will become a major climate-change impact on polar marine ecosystems. The High Arctic experiences extreme seasonality in the light environment, from 24 h light to 24 h darkness, and therefore provides a natural laboratory for studying light and predation risk over diel to seasonal timescales. Here, we show that zooplankton (observed using acoustics) in an Arctic fjord position themselves vertically in relation to light. A single isolume (depth-varying line of constant light intensity, the value of which is set at the lower limit of photobehaviour reponses of Calanus spp. and krill) forms a ceiling on zooplankton distribution. The vertical distribution is structured by light across timescales, from the deepening of zooplankton populations at midday as the sun rises in spring, to the depth to which zooplankton ascend to feed during diel vertical migration. These results suggest that zooplankton might already follow a foraging strategy that will keep visual predation risk roughly constant under changing light conditions, such as those caused by the reduction of sea ice, but likely with energetic costs such as lost feeding opportunities as a result of altered habitat use.

Observing multiple size classes of organisms, along with oceanographic properties and water mass origins, can improve our understanding of the drivers of aggregations, yet acquiring these measurements remains a fundamental challenge in biological oceanography. By deploying multiple biological sampling systems, from conventional bottle and net sampling to in situ imaging and acoustics, we describe the spatial patterns of different size classes of marine organisms (several microns to ∼10 cm) in relation to local and regional (m to km) physical oceanographic conditions on the Delaware continental shelf. The imaging and acoustic systems deployed included (in ascending order of target organism size) an imaging flow cytometer (CytoSense), a digital holographic imaging system (HOLOCAM), an In Situ Ichthyoplankton Imaging System (ISIIS, 2 cameras with different pixel resolutions), and multi-frequency acoustics (SIMRAD, 18 and 38 kHz). Spatial patterns generated by the different systems showed size-dependent aggregations and differing connections to horizontal and vertical salinity and temperature gradients that would not have been detected with traditional station-based sampling (∼9-km resolution). A direct comparison of the two ISIIS cameras showed composition and spatial patchiness changes that depended on the organism size, morphology, and camera pixel resolution. Large zooplankton near the surface, primarily composed of appendicularians and gelatinous organisms, tended to be more abundant offshore near the shelf break. This region was also associated with high phytoplankton biomass and higher overall organism abundances in the ISIIS, acoustics, and targeted net sampling. In contrast, the inshore region was dominated by hard-bodied zooplankton and had relatively low acoustic backscatter. The nets showed a community dominated by copepods, but they also showed high relative abundances of soft-bodied organisms in the offshore region where these organisms were quantified by the ISIIS. The HOLOCAM detected dense patches of ciliates that were too small to be captured in the nets or ISIIS imagery. This near-simultaneous deployment of different systems enables the description of the spatial patterns of different organism size classes, their spatial relation to potential prey and predators, and their association with specific oceanographic conditions. These datasets can also be used to evaluate the efficacy of sampling techniques, ultimately aiding in the design of efficient, hypothesis-driven sampling programs that incorporate these complementary technologies.

Objective: Lateral neck dissection (LND) is important in managing papillary thyroid carcinomas (PTCs). This study aimed to evaluate the relationship between lymph node yield (LNY) for LND and patient outcomes, specifically postoperative serum thyroglobulin levels (sTG) and radioiodine uptake on thyroid scan, and to estimate a threshold LNY to signify adequate LND. Methods: Patients diagnosed with PTC who underwent LND from 2006 to 2015 at a single institution were included. Linear regression with restricted cubic splines was used to characterize the association of LNY with outcomes. Outcomes were log-transformed to achieve a more symmetric distribution prior to regression. For nonlinear associations, a Monte Carlo Markov Chain procedure was used to estimate a threshold LNY associated with postoperative outcome. This threshold was then used to define high LNY versus low LNY in the subsequent analyses. Results: In total, 107 adult patients were included. There was a significant relationship between LNY and postoperative sTG level ( P = .004), but not radioiodine uptake ( P = .64). An LNY of 42.96 was identified, which was associated with the maximum change in sTG level. No association was found between LNY groups (LNY ≥ 43 vs < 43) and radioiodine uptake, risk of complication, or longer operative times (all P > .05). High LNY was associated with a decrease in log(sTG) (estimate = −1.855, P = .03), indicating that adequacy of LND is associated with an 84.4% decrease in sTG. Conclusion: These results suggest an association between LNY and postoperative sTG level, with an estimated threshold of 43 nodes. This has implications for adequate therapeutic LND; additional work is needed to validate thresholds for clinical practice.

Early life history in marine benthic crustaceans often includes externally brooded eggs that hatch into free-swimming planktonic larvae. These larvae are relatively strong swimmers, and movement in the vertical plane provides a number of advantages, including modulation of horizontal transport and assurance of favorable predator–prey interactions. Swimming behavior in larval crustaceans is regulated by predictable external cues in the water column, primarily light, gravity, and hydrostatic pressure. Light-regulated behavior depends upon the optical physics of seawater and the physiology of light-detecting sensory structures in the larvae, which overall vary little with ontogeny. Swimming in response to light contributes to ecologically significant behaviors in planktonic crustacean larvae, including shadow responses, depth regulation, and diel vertical migration. Moreover, the photoresponses themselves, and in turn the evoked behaviors, change with the needs of larvae as development progresses. Regarding other sensory modalities, crustacean embryos and larvae respond to chemical cues using bimodal sensilla (chemosensory and mechanosensory) as contact receptors, and aesthetascs for detection of water-soluble cues. Processes and behaviors are stimulated by larval detection of chemical cues throughout ontogeny, including egg-hatching, avoidance of predators during free-swimming stages, and, ultimately, settlement and metamorphosis in juvenile habitats. The latter process can also involve tactile cues. The sensory-mediated behaviors described here for crustacean larvae have parallels in numerous arthropod and nonarthropod taxa. Emerging directions for future research on sensory aspects of behavior in crustacean larvae include multimodal sensory integration and behavioral responses to changing environmental stressors.

For organisms that remain active in one of the last undisturbed and pristine dark environments on the planet—the Arctic Polar Night—the moon, stars and aurora borealis may provide important cues to guide distribution and behaviours, including predator-prey interactions. With a changing climate and increased human activities in the Arctic, such natural light sources will in many places be masked by the much stronger illumination from artificial light. Here we show that normal working-light from a ship may disrupt fish and zooplankton behaviour down to at least 200 m depth across an area of >0.125 km2 around the ship. Both the quantitative and qualitative nature of the disturbance differed between the examined regions. We conclude that biological surveys in the dark from illuminated ships may introduce biases on biological sampling, bioacoustic surveys, and possibly stock assessments of commercial and non-commercial species., Berge et al. find that the normal working-light from a ship impacts on the vertical distribution of macrozooplankton and pelagic fish communities around the ship at three stations during the Arctic Polar Night. These data suggest bias from such effects should be taken into account when performing surveys and stock assessments in the Arctic.

The currently unsurpassed diversity of photoreceptors found in the eyes of stomatopods, or mantis shrimps, is achieved through a variety of opsin-based visual pigments and optical filters. However, the presence of extraocular photoreceptors in these crustaceans is undescribed. Opsins have been found in extraocular tissues across animal taxa, but their functions are often unknown. Here, we show that the mantis shrimp Neogonodactylus oerstedii has functional cerebral photoreceptors, which expands the suite of mechanisms by which mantis shrimp sense light. Illumination of extraocular photoreceptors elicits behaviors akin to common arthropod escape responses, which persist in blinded individuals. The anterior central nervous system, which is illuminated when a mantis shrimp’s cephalothorax protrudes from its burrow to search for predators, prey, or mates, appears to be photosensitive and to feature two types of opsin-based, potentially histaminergic photoreceptors. A pigmented ventral eye that may be capable of color discrimination extends from the cerebral ganglion, or brain, against the transparent outer carapace, and exhibits a rapid electrical response when illuminated. Additionally, opsins and histamine are expressed in several locations of the eyestalks and cerebral ganglion, where any photoresponses could contribute to shelter-seeking behaviors and other functions.

Specialized defense strategies are induced in zooplankton upon detection of predator chemical cues or kairomones. These defenses are well-described for freshwater zooplankters, with morphological defenses being particularly striking, but few studies have reported kairomone-induced morphological defenses in marine zooplankton. Here, we compare morphological responses to kairomones in the larvae of two marine crab species, estuarine mud crabs (Rhithropanopeus harrisii) and Asian shore crabs (Hemigrapsus sanguineus). When reared in the presence of fish kairomones, spine length increased by 2–3% in larval R. harrisii, while no morphological changes were identified in H. sanguineus. In subsequent feeding assays with a co-occurring fish predator (Atlantic silversides, Menidia menidia), consumption of R. harrisii was lower on larvae that had been reared with kairomones. In addition, we found that broods with smaller larvae are more likely to exhibit increases in spine length after kairomone exposure. Hence, the observed morphological response is likely influenced by larval size. This article is protected by copyright. All rights reserved.

The Polar Night has long been regarded as a period of no biological activity. As a logical consequence, environmental management has mainly been neglected in this period. We will use the Northguider accident in December 2018 as a case study for the need to prioritize operative habitat mapping and monitoring to provide a sufficiently knowledge-based environmental operative management. After the ship ran ashore in a remote location in the northern part of Svalbard, the entire crew was safely rescued and airlifted back to Longyearbyen within hours. The ship, however, remained, potentially posing a threat to wildlife – not only wildlife that is expected to return in spring but also for the many organisms that we now know are present and active during the Polar Night. There is, however, still a strong need to provide necessary understanding of the dynamic marine biodiversity at the sea surface, water column, and seafloor during the Polar Night, focusing on the threats for ecosystems and habitats.

Fish odor induces predator avoidance behaviors in zooplankton, like vertical migration, by making zooplankton more responsive to light. Odor cues that alter behavior in marine crustacean zooplankton in the laboratory include sulfated glycosaminoglycans (sGAGs) derived from fish body mucus. Few studies quantify these cues in estuarine/marine environments or assess whether laboratory studies reflect natural scenarios. We collected fish and water samples weekly in Broadkill River, Delaware, USA. We used field-collected water in colorimetric assays to determine the concentration of sGAG-equivalent molecules and in behavioral assays with a zooplankton model, brine shrimp (Artemia spp.) nauplii, which only descend in response to downwelling light after fish odor exposure. Fish quantity was positively related to sGAG-equivalents and zooplankton photosensitivity, indicated by descent responses at lower light levels and across a broad intensity range. Our results support that fish odor concentrations used in previous laboratory assays are consistent with levels found in an estuary.

How much light isa vailable for biological processes during Polar Night? This question appears simple enough. But the reality is that conventional light sen- sors for measuring visible light (~350 to ~700 nm) have not been sensitive enough to answer it. Beyond this technical challenge, “light” is a general term that must be qualified in terms of “light climate” before it has meaning for biological systems. In this chapter, we provide an answer to the question posed above and explore aspects of light climate during Polar Night with relevance to biology, specifically, how Polar Night is defined by solar elevation, atmospheric light in Polar Night and its propaga- tion underwater, bioluminescence in Polar Night and the concept of Polar Night as a deep-sea analogue, light pollution, and future perspectives. This chapter focuses on the quantity and quality of light present during Polar Night, while subsequent chapters in this volume focus on specific biological effects of this light for algae (Chap. “Marine Micro- and Macroalgae in the Polar Night”), zooplankton (Chaps.“Zooplankton in the Polar Night” and “Biological Clocks and Rhythms in Polar Organisms”), and fish (Chap. “Fish Ecology in the Polar Night”).

Larval development in brachyuran crabs includes a number of zoeal stages followed by a single megalopal stage. Zoeae and megalopae are relatively strong swimmers, and movement up or down in the water column takes advantage of vertical shear in ambient currents with consequent transport in the horizontal dimension. For some species, this process is important in controlling the dispersal of early-stage larvae away from spawning sites and in maintaining the supply of late-stage larvae to juvenile nursery habitat. For other species it enables retention near spawning sites throughout the larval period. Vertical position in the water column also modulates predator-prey interactions, which impact growth and survival of larvae. Swimming behavior in larval crabs is regulated by both external cues detected in the water column and endogenous rhythms entrained by external oscillators. Gravity, hydrostatic pressure, and light are the primary external cues because of their predictability in the environment. Light is also the most common external oscillator entraining swimming rhythms. Secondary cues include salinity, temperature, turbulent kinetic energy, and feeding state. Crab larvae also respond to chemical and tactile cues that facilitate settlement in juvenile habitat. This paper presents a review of the physical and chemical characteristics of these cues, the behavioral responses of crab larvae to the cues, and the patterns of larval transport that emanate from these responses.

Many species of marine larvae use exogenous cues, such as gravity and pressure, for orientation and dispersal, but it is unclear if parasitic larvae exhibit comparable geotactic and barokinetic behaviors. Using two species of marine cercariae, Euhaplorchis sp. and Probolocoryphe lanceolata , which utilize the same first intermediate host, the mangrove snail Cerithidea scalariformis , the hypothesis that they use gravity and pressure to position themselves in the water column to increase the probability of contacting their respective second intermediate hosts was tested. In a test chamber under dark laboratory conditions, Euhaplorchis sp. cercariae ascended, and therefore displayed negative geotaxis, increasing the probability of finding their second intermediate host, salt marsh killifish ( Fundulus spp.). In contrast, P. lanceoloata cercariae swam downwards towards the bottom of the chamber, and this positive geotactic behavior would place them in a benthic microhabitat consistent with their second intermediate host, fiddler crabs ( Uca spp.). In a subsequent experiment testing swimming activity in response to step increases in pressure, no effect of pressure changes over the range tested (0.3–30 kPa) was observed for either species. Collectively, these results suggest that while cercariae are unresponsive to changes in pressure, they are able to utilize gravity for orientation, placing them in the preferred microhabitat of the second intermediate host. This finding is consistent with published photobehavioral studies in which Euhaplorchis swam upwards towards light and P . lanceolata swam downward towards the bottom upon light stimulation. These complementary phototactic and geotactic behaviors optimize host contact, and thus completion of their complex life cycle.

We consider machine-learning-based thyroid-malignancy prediction from cytopathology whole-slide images (WSI). Multiple instance learning (MIL) approaches, typically used for the analysis of WSIs, divide the image (bag) into patches (instances), which are used to predict a single bag-level label. These approaches perform poorly in cytopathology slides due to a unique bag structure: sparsely located informative instances with varying characteristics of abnormality. We address these challenges by considering multiple types of labels: bag-level malignancy and ordered diagnostic scores, as well as instance-level informativeness and abnormality labels. We study their contribution beyond the MIL setting by proposing a maximum likelihood estimation (MLE) framework, from which we derive a two-stage deep-learning-based algorithm. The algorithm identifies informative instances and assigns them local malignancy scores that are incorporated into a global malignancy prediction. We derive a lower bound of the MLE, leading to an improved training strategy based on weak supervision, that we motivate through statistical analysis. The lower bound further allows us to extend the proposed algorithm to simultaneously predict multiple bag and instance-level labels from a single output of a neural network. Experimental results demonstrate that the proposed algorithm provides competitive performance compared to several competing methods, achieves (expert) human-level performance, and allows augmentation of human decisions.

Diel vertical migration involves vertical migration over the 24-h day. The three patterns are nocturnal, twilight, and reverse, and they function as strategies for feeding and predator avoidance. Zooplankton either follow a preferred light level during migration or are cued to migration by the rate and direction of change in light at twilight. During selective tidal-stream transport, animals migrate vertically relative to tides. During flood-tide transport, animals move up estuaries by swimming in the water column during flood tide and descending to the bottom during ebb tides. The opposite occurs during ebb tide transport. The underlying mechanisms are tidal activity rhythms and responses to environmental factors.

Due to its Arctic location at 79°N, Kongsfjorden in Svalbard experiences strong seasonality in light climate, changing from polar night to midnight sun. Sea ice conditions and the optical properties of seawater further modify the amount and the spectral composition of solar radiation penetrating into the water column, thus defining the underwater light climate in Kongsfjorden. Light represents one of the major shaping factors for the entire marine ecosystem. A number of studies focusing on implications of the underwater light for marine organisms have been conducted in Kongsfjorden, generating diverse datasets on seawater optical properties, scattered over time and space. This review synthesizes the fragmentary information available from the literature as well as presenting some unpublished data, and discusses the underwater light climate and its main controlling factors in Kongsfjorden. Furthermore, we provide a short synopsis about the relevance of light for different components of an Arctic marine ecosystem, exemplified by studies carried out in Kongsfjorden. Due to its year-round accessibility and its high-Arctic location, Kongsfjorden has become a prime fjord for studying how the strong seasonal changes in light availability, ranging from polar night to midnight sun, affect marine life with respect to primary production, behavioural aspects and synchronization of growth and reproduction.

Characterizing the thermal niche of harmful algae is crucial for understanding and projecting the effects of future climate change on harmful algal blooms. The effects of 6 different temperatures (18–32 °C) on the growth, photophysiology, and toxicity were examined in the dinoflagellate Karlodinium veneficum, and the raphidophytes, Heterosigma akashiwo and Chattonella subsalsa from the Delaware Inland Bays (DIB). K. veneficum and H. akashiwo had skewed unimodal growth patterns, with temperature optima (Topt) at 28.6 and 27.3 °C respectively and an upper thermal niche limit of 32 °C. In contrast, C. subsalsa growth increased linearly with temperature, suggesting Topt and upper thermal boundaries >32 °C. K. veneficum photosystem II (PSII) photochemical efficiency remained stable across all temperatures, while H. akashiwo PSII efficiency declined at higher temperature and C. subsalsa was susceptible to low temperature (~18 °C) photoinactivation. Cell toxicity thermal response was species-specific such that K. veneficum toxicity increased with temperature above Topt. Raphidophyte toxicity peaked at 25–28 °C and was in close agreement with Topt for growth in H. akashiwo but below C. subsalsa maximal growth. The mode of toxicity was markedly different between the dinoflagellate and the raphidophytes such that K. veneficum had greater hemolytic activity while the raphidophytes had pronounced fish gill cell toxicity. These results and patterns of natural abundance for these algae in the DIB suggest that continued ocean warming may contribute to C. subsalsa bloom formation while possibly promoting highly toxic blooms of K. veneficum.

Several predator avoidance strategies in zooplankton rely on the use of light to control vertical position in the water column. Although light is the primary cue for such photobehavior, predator chemical cues or kairomones increase swimming responses to light. We currently lack a mechanistic understanding for how zooplankton integrate visual and chemical cues to mediate phenotypic plasticity in defensive photobehavior. In marine systems, kairomones are thought to be amino sugar degradation products of fish body mucus. Here, we demonstrate that increasing concentrations of fish kairomones heightened sensitivity of light-mediated swimming behavior for two larval crab species (Rhithropanopeus harrisii and Hemigrapsus sanguineus). Consistent with these behavioral results, we report increased visual sensitivity at the retinal level in larval crab eyes directly following acute (1–3 h) kairomone exposure, as evidenced electrophysiologically from V–log I curves and morphologically from wider, shorter rhabdoms. The observed increases in visual sensitivity do not correspond with a decline in temporal resolution, because latency in electrophysiological responses actually increased after kairomone exposure. Collectively, these data suggest that phenotypic plasticity in larval crab photobehavior is achieved, at least in part, through rapid changes in photoreceptor structure and function.

We established the lethal levels for water-accommodated fractions of Corexit ® 9500A chemical dispersant, crude oil (WAF), and dispersed crude oil (CEWAF) for the ctenophore Mne- miopsis leidyi at both 15 and 23°C. This gelatinous zooplankter was sensitive to dispersant at both temperatures, as well as to oil solutions, with some increase in toxicity of CEWAF as compared to WAF. Subsequent sublethal assays for routine respiration rate, bioluminescence, and glutathione- s-transferase (GST) activity were conducted on individuals surviving 24 h exposures to test solu- tions at both 15 and 23°C. GST activity increased significantly in 2.5 and 5 mg l �1 dispersant solu- tions at 15°C, suggesting a metabolic detoxification response to the dispersant-containing solutions, but no effect of any solution type on routine respiration rate was observed. Light emis- sion through mechanically stimulated bioluminescence and photocyte lysis decreased with exposure to crude oil WAF and CEWAF at both temperatures and to dispersant exposure at 23°C. Collectively, these results demonstrate that M. leidyi exhibits both lethal and sublethal effects from acute crude oil exposure, with an elevation of some sublethal responses upon addition of chemical dispersant. Sublethal effects of oil and dispersants in pelagic species, most notably impairment of luminescence, should be considered when evaluating oil spill response strategies.

Animals that have true color vision possess several spectral classes of photoreceptors. Pancrustaceans (Hexapoda + Crustacea) that integrate spectral information about their reconstructed visual world do so from photoreceptor terminals supplying their second optic neuropils, with subsequent participation of the third (lobula) and deeper centers (optic foci). Here we describe experiments and correlative neural arrangements underlying convergent visual pathways in two species of branchiopod crustaceans that have to cope with a broad range of spectral ambience and illuminance in ephemeral pools, yet possess just two optic neuropils, the lamina and optic tectum. Electroretinographic recordings and multimodel inference based on modeled spectral absorptance were used to identify the most likely number of spectral photoreceptor classes in their compound eyes. Recordings from the retina provide support for four color channels. Neuroanatomical observations resolve arrangements in their laminas that suggest signal summation at low light intensities, incorporating chromatic channels. Neuroanatomical observations demonstrate that spatial summation in the lamina of the two species are mediated by quite different mechanisms, both of which allow signals from several ommatidia to be pooled at single lamina monopolar cells. We propose that such summation provides sufficient signal for vision at intensities equivalent to those experienced by insects in terrestrial habitats under dim starlight. Our findings suggest that despite the absence of optic lobe neuropils necessary for spectral discrimination utilized by true color vision, four spectral photoreceptor classes have been maintained in Branchiopoda for vision at very low light intensities at variable ambient wavelengths that typify conditions in ephemeral fresh water habitats.

Using new enabling technologies, we document behavioral patterns and susceptibility to light pollution never previously seen., Light is a major cue for nearly all life on Earth. However, most of our knowledge concerning the importance of light is based on organisms’ response to light during daytime, including the dusk and dawn phase. When it is dark, light is most often considered as pollution, with increasing appreciation of its negative ecological effects. Using an Autonomous Surface Vehicle fitted with a hyperspectral irradiance sensor and an acoustic profiler, we detected and quantified the behavior of zooplankton in an unpolluted light environment in the high Arctic polar night and compared the results with that from a light-polluted environment close to our research vessels. First, in environments free of light pollution, the zooplankton community is intimately connected to the ambient light regime and performs synchronized diel vertical migrations in the upper 30 m despite the sun never rising above the horizon. Second, the vast majority of the pelagic community exhibits a strong light-escape response in the presence of artificial light, observed down to 100 m. We conclude that artificial light from traditional sampling platforms affects the zooplankton community to a degree where it is impossible to examine its abundance and natural rhythms within the upper 100 m. This study underscores the need to adjust sampling platforms, particularly in dim-light conditions, to capture relevant physical and biological data for ecological studies. It also highlights a previously unchartered susceptibility to light pollution in a region destined to see significant changes in light climate due to a reduced ice cover and an increased anthropogenic activity.

Many estuarine crab species are exported from this habitat at spawning and are physically transported to coastal waters where they undergo zoeal development. Later, they recruit back to the adult habitat as megalopae. Larval behavior within the context of wind-driven coastal circulation contributes to this process. The present study tested whether the crab Hemigrapsus sanguineus , an invasive species to the mid-Atlantic coast of the United States as well as in Europe, possesses behavioral responses to exogenous cues (gravity and light) and endogenous rhythms that would facilitate this scenario. Throughout larval development, H. sanguineus zoeae were negatively buoyant. In a laboratory test chamber, early stage (Z1) larvae displayed a strong negative geotaxis in darkness which resulted in surface swimming that was not offset by behavioral responses to light exposure. These Z1 stage larvae also possessed an endogenous tidal rhythm in swimming activity that was in phase with ebb tides in the spawning habitat and was independent of light exposure. These behaviors are consistent with export of larvae from estuarine to coastal waters where larval development occurs. Intermediate (Z3) and late (Z5) stage larvae exhibited a predominant negative geotaxis and surface swimming, with some zoeae showing positive geotaxis and/or sinking in the Z5 stage. Visual spectral sensitivity of eyes in Z3 larvae peaked between 450 and 510 nm, with a secondary peak evident at short-wavelengths (~ 370 nm). In both the Z3 and Z5 stages, larvae descended in response to light exposure, as expected under a negative feedback model of depth regulation common among crab larvae. Photoresponse thresholds of 5.8 × 10 12 and 1.2 × 10 14 photons m − 2 s − 1 for Z3 and Z5 stage larvae, respectively, predict both stages to be in the lower water column during the day. After molting to megalopae, H. sanguineus display positive geotaxis and/or sinking in darkness and lack a phototactic response, consistent with settlement in the adult benthic habitat. Thus, larvae of the invasive crab H. sanguineus possess endogenous rhythms and behavioral responses to environmental cues that are similar to those observed in successful native species with a similar coastal larval development pattern.

Chemical cues from fish, or kairomones, often impact the behavior of zooplankton. These behavioral changes are thought to improve predator avoidance. For example, marine and estuarine crustacean zooplankton become more sensitive to light after kairomone exposure, which likely deepens their vertical distribution into darker waters during the day and thereby reduces their visibility to fish predators. Here, we show that kairomones from an estuarine fish induce similar behavioral responses in adult brine shrimp (Artemia franciscana) from an endorheic, hypersaline lake, Great Salt Lake, Utah, USA. Given downwelling light stimuli, kairomone-exposed A. franciscana induce a descent response upon dimmer light flashes than they do in the absence of kairomones. Using extracellular electroretinogram (ERG) recordings, we also find that kairomones induce physiological changes in the retina that may lead to increased visual sensitivity, suggesting that kairomone-induced changes to photobehavior are mediated at the photoreceptor level. However, kairomones did not induce structural changes within the eye. Although A. franciscana inhabit endorheic environments that are too saline for most fish, kairomones from an estuarine fish amplify photobehavior in these branchiopod crustaceans. The mechanism for this behavioral change has both similarities to and differences from that described in marine malacostracan crustaceans.

Light is a primary environmental factor used by aquatic invertebrates for depth selection behavior. Many branchiopod crustaceans live in ephemeral aquatic habitats. All branchiopod crustaceans studied to date express four or more visual opsins in their compound eyes. We asked whether two branchiopods, Triops longicaudatus, and Streptocephalus mackini, use multiple spectral channels to regulate their position in the water column. At the lowest intensities that elicited photonegative behavior, both species had broad spectral bandwidths, suggesting they use multiple spectral photoreceptor classes. Male Streptocephalus mackini were more likely to maintain a vertical position 8.0-12.0 cm below the surface than females, independently of whether females were present. Male photopositive behavior at low intensity was restricted to narrow bandwidth centered at 532nm, suggesting a single photoreceptor class is used to maintain position above females. We compared ephemeral pools from two regions in Arizona and found that diffuse light attenuation coefficients were two orders of magnitude greater than the most heavily attenuating coastal waters. At less than a meter of depth, pools were often dimmer than terrestrial habitats under starlight. Soil particle size distribution in each region affected spectral light environments, and behavioral responses of field-caught shrimp were adapted to the spectral properties of their region. The results suggest that branchiopods predominantly use luminance vision summed from multiple spectral photoreceptor classes for depth selection in dim, spectrally variable environments. The neuroanatomical basis for summation is described in a companion paper.

The subtidal crab Rhithropanopeus harrisii has a circadian rhythm in larval release; under constant condi- tions eggs hatch in the 2-3-h interval after the time of sunset in nontidal estuaries. Eggs that are removed from the female hatch rhythmically, indicating that the circadian rhythm resides in the embryos. The model for entrainment is that mature embryos have functional sensory systems that detect and entrain to environmental cycles. This model was reex- amined by confirming that the visual system of advanced embryos responds to light and thus could mediate entrain- ment to the light/dark cycle. We then determined whether the hatching rhythm of mature embryos that are removed from the female can be entrained to new light/dark cycles. Contrary to expectations, these embryos did not entrain to new cycles. Instead, they remained entrained to the light/ dark cycle to which they were exposed when still attached to the female, suggesting that the female entrains the rhythm. Indeed, hatching by embryos collected from the field when they had not yet developed eye pigments, kept in constant conditions attached to their mother, exhibited the circadian hatching rhythm. They could also be entrained to a new photoperiod in the laboratory. The role of the female is further supported by experiments showing that the hatch- ing rhythm in embryos carried by females lacking one but not both eyes can be entrained to a new cycle in the laboratory. Thus, the revised model is that the female per- ceives the light/dark cycle and entrains the circadian rhythm in the embryos.

Knowledge of lethal and sublethal effects of crude oil and dispersants on mesozooplankton are important to understanding ecosystem impacts of oil spills in marine environments. Here we (1) establish median lethal concentrations for water accommodated fractions of Corexit EC9500A dispersant, MC-252 crude oil (WAF), and dispersed crude oil (CEWAF) for the coastal copepod Labidocera aestiva, and (2) assess acute effects on L. aestiva swimming activity. Mortality assays with L. aestiva support that copepods are more sensitive than other zooplankton taxa to dispersant toxicity, while WAF and CEWAF are generally similar in their toxicity to this copepod species and other zooplankton. Acute effects on L. aestiva activity included impaired swimming upon WAF and CEWAF exposure. These results highlight that copepods are particularly sensitive to dispersant exposure, with acute effects on survival most evident with dispersant alone, and on swimming behavior when dispersant is mixed with crude oil.

Talitrid amphipods use many cues for orientation during forays between temporary burrows and feeding areas, and for locating beaches when submerged, with visual cues being particularly important. Little evidence exists for polarized light among these visual cues despite extensive orientation by celestial and underwater polarized light in other crustaceans and in insects. We used electroretinography to assess spectral sensitivity in the eye of the beach flea Platorchestia platensis, and behavioral studies to test whether linearly polarized light serves as an orientation cue. Two spectral classes were present in the P. platensis eye with maxima at 431 and 520 nm. Non-uniform orientation of amphipods in the laboratory arena required either light/dark or polarized cues. Scototactic movements depended on arena conditions (day/night, wet/dry), while orientation under linearly polarized light was wavelength-dependent and parallel to the e-vector. Subsequent tests presented conflicting and additive scototactic and polarotactic cues to differentiate among these responses. In dry conditions, orientation parallel to the polarization e-vector overcame a dominant negative scototaxis, confirming that polarotaxis and scototaxis are separate orientation responses in this species. These behavioral results demonstrate talitrid amphipods can perceive and orient to linearly polarized light, and may use it to orient toward preferred zones on beaches.

Exposure to high pCO2 or low pH alters sensation and behaviour in many marine animals. We show that crab larvae lose their ability to detect and/or process predator kairomones after exposure to low pH over a time scale relevant to diel pH cycles in coastal environments. Previous work suggests that acidification affects sensation and behaviour through altered neural function, specifically the action of γ-aminobutyric acid (GABA), because a GABA antagonist, gabazine, restores the original behaviour. Here, however, gabazine resulted in a loss of kairomone detection/processing, regardless of pH. Our results also suggest that GABAergic signalling is necessary for kairomone identification in these larvae. Hence, the mechanism for the observed pH effect varies from the original GABA hypothesis. Furthermore, we suggest that this pH effect is adaptive under diel-cycling pH.

Diel vertical migration (DVM) is a characteristic behavioural pattern performed by zooplankton in which their vertical distribution changes over the 24-h day. Here the proximate control of zooplankton DVM is reviewed. Light has emerged as the major proximate cue controlling DVM behaviour and the understanding of zooplankton visual physiology and the light-mediated behaviour underlying DVM is expanding. Field and laboratory evidence exist to support each of the three major hypotheses for the exogenous role of light in DVM: (1) preferendum or isolume, (2) absolute intensity threshold, and (3) relative rate of change. Light may also play an endogenous role in DVM by entraining circadian rhythms in vertical movement or activity. This appreciation of the role of light has improved modelling efforts into the causes and consequences of DVM. The most important recent advance in the study of DVM is the recognition that this behaviour is a pheno typic response in many species and is most commonly activated by chemical cues (kairomones) from sh predators. High levels of kairomones signal high levels of predation pressure, and DVM-related photobehaviours, such as swimming responses on relative rates of irradiance change, are altered such that migration occurs and zooplankton achieve a refuge from visual predators.