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ABSTRACT ObjectiveThe purpose of this study was to understand the possible provenance of a unique population of Atlantic Cod Gadus morhua that is found in the low-salinity Åland Sea region of the northern Baltic Sea. This population consists of large, healthy individuals, in contrast to the Atlantic Cod in the rest of the Baltic Sea. MethodsWe used laser ablation inductively coupled plasma mass spectrometry to measure levels of boron (as B:Ca) in the otoliths of Atlantic Cod in regions throughout the Baltic Sea. We examined both lifetime chronologies and concentrations in the core region that corresponds to birth and early life. ResultsWe found that B:Ca concentrations were 31 to 348 times higher in the otoliths of cod that occupy the Åland Sea, including in the core region. These concentrations were much higher than expected given that boron is linearly, positively proportional to salinity, which is higher in the southern Baltic Sea, and other populations displayed very low concentrations by comparison. ConclusionsBased on the otolith B:Ca as a unique marker, we suggest that the cod that are sampled in the Åland Sea may be a separate population from those that inhabit the rest of the Baltic Sea. This would not prevent it from mixing with other populations but could point to a separate spawning area. The source of the elevated boron is currently unknown, but the widespread occurrence in cod otoliths from the Åland Sea indicates an extensive nonpoint source.more » « lessFree, publicly-accessible full text available July 25, 2026
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ABSTRACT Fjords provide valuable research opportunities for marine scientists. They are excellent natural infrastructure for climate impact studies associated with hypoxic episodes and consequences for mesopelagic and deep‐sea ecosystems involving oceanographic circulation processes and basin water renewals. Repeated sampling from the same populations is possible, making fjords excellent systems for developing time series of data for climate impact studies. We provide an overview of the 14 years of data from Norwegian West Coast fjords, focusing on Masfjorden, and report major findings from Oslofjorden in Eastern Norway, exhibiting recurrent hypoxia in the basin waters. We document that the oxygen levels in Masfjorden decreased rapidly by over 60% at 450 m depth in < 8 years, which is much faster than the average rate of deoxygenation in the global ocean. We also discuss the increase in the deep‐sea and low‐light‐adapted coronate jellyfishPeriphylla periphyllain view of altered optical conditions of the basin water potentially related to deoxygenation. We argue that fjords like Masfjorden and Oslofjorden are not only macrocosms for ecological processes but also are likely an accelerated version of deep oceans with respect to climate impacts.more » « less
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Abstract Anthropogenic deoxygenation of the Baltic Sea caused major declines in demersal and benthic habitat quality with consequent impacts on biodiversity and ecosystem services. Using Baltic cod otolith chemical proxies of hypoxia, salinity, and fish metabolic status and growth, we tracked changes from baseline conditions in the late Neolithic (4500 BP) and early twentieth century to the present, in order to understand how recent, accelerating climate change has affected this key species. Otolith hypoxia proxies (Mn:Mg) increased with expanding anoxic water volumes, but decreased with increasing salinity indexed by otolith Sr:Ca. Metabolic status proxied by otolith Mg:Ca and reconstructed growth were positively related to dissolved oxygen percent saturation, with particularly severe declines since 2010. This long-term record of otolith indicators provides further evidence of a profound state change in oxygen for the worse, in one of the world’s largest inland seas. Spreading hypoxia due to climate warming will likely impair fish populations globally and evidence can be tracked with otolith chemical biomarkers.more » « less
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Abstract Chemical analysis of calcified structures continues to flourish, as analytical and technological advances enable researchers to tap into trace elements and isotopes taken up in otoliths and other archival tissues at ever greater resolution. Increasingly, these tracers are applied to refine age estimation and interpretation, and to chronicle responses to environmental stressors, linking these to ecological, physiological, and life-history processes. Here, we review emerging approaches and innovative research directions in otolith chemistry, as well as in the chemistry of other archival tissues, outlining their value for fisheries and ecosystem-based management, turning the spotlight on areas where such biomarkers can support decision making. We summarise recent milestones and the challenges that lie ahead to using otoliths and archival tissues as biomarkers, grouped into seven, rapidly expanding and application-oriented research areas that apply chemical analysis in a variety of contexts, namely: (1) supporting fish age estimation; (2) evaluating environmental stress, ecophysiology and individual performance; (3) confirming seafood provenance; (4) resolving connectivity and movement pathways; (5) characterising food webs and trophic interactions; (6) reconstructing reproductive life histories; and (7) tracing stock enhancement efforts. Emerging research directions that apply hard part chemistry to combat seafood fraud, quantify past food webs, as well as to reconcile growth, movement, thermal, metabolic, stress and reproductive life-histories provide opportunities to examine how harvesting and global change impact fish health and fisheries productivity. Ultimately, improved appreciation of the many practical benefits of archival tissue chemistry to fisheries and ecosystem-based management will support their increased implementation into routine monitoring. Graphical abstractmore » « less
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Abstract Fish otoliths' chronometric properties make them useful for age and growth rate estimation in fisheries management. For the Eastern Baltic Sea cod stock (Gadus morhua), unclear seasonal growth zones in otoliths have resulted in unreliable age and growth information. Here, a new age estimation method based on seasonal patterns in trace elemental otolith incorporation was tested for the first time and compared with the traditional method of visually counting growth zones, using otoliths from the Baltic and North seas. Various trace elemental ratios, linked to fish metabolic activity (higher in summer) or external environment (migration to colder, deeper habitats with higher salinity in winter), were tested for age estimation based on assessing their seasonal variations in concentration. Mg:Ca and P:Ca, both proxies for growth and metabolic activity, showed greatest seasonality and therefore have the best potential to be used as chemical clocks. Otolith image readability was significantly lower in the Baltic than in the North Sea. The chemical (novel) method had an overall greater precision and percentage agreement among readers (11.2%, 74.0%) than the visual (traditional) method (23.1%, 51.0%). Visual readers generally selected more highly contrasting zones as annuli whereas the chemical readers identified brighter regions within the first two annuli and darker zones thereafter. Visual estimates produced significantly higher, more variable ages than did the chemical ones. Based on the analyses in our study, we suggest that otolith microchemistry is a promising alternative ageing method for fish populations difficult to age, such as the Eastern Baltic cod.more » « less
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Typical analyses of otolith microchemistry use calcium, a major constituent, as an internal standard, setting its value as a constant and ignoring any potential variations. In fact, patterns do occur in otolith Ca deposition, as can be observed either by repeating the analysis, by creating two-dimensional maps of Ca, or both. Here we present evidence of Ca variations in fish otoliths from analyses using synchrotron-based scanning X-ray fluorescence microscopy, electron microprobe analysis, and laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS). 2-D maps of otoliths created with LA-ICP-MS indicate that Ca is elevated where especially Zn and P are low, and vice versa, suggesting that spatial variations in protein deposition may affect concentrations of Ca. We encourage others to examine Ca concentrations in their biomineralized samples to check for variations, using LA-ICP-MS and other methods.more » « lessFree, publicly-accessible full text available March 1, 2026
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Chronological records of elemental concentrations in fish otoliths are a widely used tool to infer the environmental conditions experienced by individual fish. To interpret elemental signals within the otolith, it is important to understand how both external and internal factors impact ion uptake, transport and incorporation. In this study, we have combined chronological records from otoliths and archival data storage tags to quantify the influence of internal (sex, size, age, growth) and external (temperature, depth, salinity) conditions on otolith elemental chemistry of cod (Gadus morhua) in natural settings of the Baltic Sea. This study focused on elements primarily under physiological control: Phosphorus (P), magnesium (Mg) and zinc (Zn); and elements under environmental control: Strontium (Sr), barium (Ba) and manganese (Mn). Based on known spatial and temporal patterns in environmental conditions and fish size, growth, and maturity, we posed a series of hypotheses of expected otolith element patterns. Partial effects of internal and external drivers on element concentration were analyzed using a Linear Mixed Model approach with random variables (fish and year). Predicted effects of otolith concentrations of all elements under physiological control (P, Mg, Zn) showed similar trends, with distinct seasonal patterns (lowest concentration in late spring, highest concentrations in winter), and a positive correlation with water temperature, in addition to higher Zn and lower P in spawning individuals. Predicted effects of otolith concentrations of elements expected to be predominantly under environmental control showed the predicted geographic and depth-related trends based on ambient salinity (Ba) and coastal hypoxia (Mn). However, contrary to expectation, Sr was unrelated to salinity. Predicted otolith Ba, Sr and Mn concentrations also exhibited pronounced seasonal patterns that were out of phase with each other but appeared to be partly explained by spawning/feeding migrations. While performing laboratory validation studies for adult fish is typically not possible, these results highlight the importance of assessing local water chemistry and freshwater endmembers in one’s study system before otolith elemental chemistry can be reliably used to reconstruct fish habitat use and environmental histories.more » « less
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Abstract. Anthropogenic warming and nutrient over-enrichment of our oceans have resulted in significant, and often catastrophic, reductions in dissolved oxygen (deoxygenation). Stress on water-breathing animals from this deoxygenation has been shown to occur at all levels of biological organization: cellular, organ, individual, species, population, community, and ecosystem. Most climate forecasts predict increases in ocean deoxygenation; thus, it is essential to develop reliable biological indicators of low-oxygen stress that can be used by regional and global oxygen monitoring efforts to detect and assess the impacts of deoxygenation on ocean life. This review focuses on responses to low-oxygen stress that are manifest at different levels of biological organization and at a variety of spatial and temporal scales. We compare particular attributes of these biological indicators to the dissolved oxygen threshold of response, timescales of response, sensitive life stages and taxa, and the ability to scale the response to oxygen stress across levels of organization. Where there is available evidence, we discuss the interactions of other biological and abiotic stressors on the biological indicators of low-oxygen stress. We address the utility, confounding effects, and implementation of the biological indicators of oxygen stress for research and societal applications. Our hope is that further refinement and dissemination of these oxygen stress indicators will provide more direct support for environmental managers, fisheries and mariculture scientists, conservation professionals, and policymakers to confront the challenges of ocean deoxygenation. An improved understanding of the sensitivity of different ocean species, communities, and ecosystems to low-oxygen stress will empower efforts to design monitoring programs, assess ecosystem health, develop management guidelines, track conditions, and detect low-oxygen events.more » « less
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Marine hypoxia has had major consequences for both economically and ecologically critical fish species around the world. As hypoxic regions continue to grow in severity and extent, we must deepen our understanding of mechanisms driving population and community responses to major stressors. It has been shown that food availability and habitat use are the most critical components of impacts on individual fish leading to observed outcomes at higher levels of organization. However, differences within and among species in partitioning available energy for metabolic demands – or metabolic prioritization – in response to stressors are often ignored. Here, I use both a multispecies size spectrum model and a meta-analysis to explore evidence in favor of metabolic prioritization in a community of commercially important fish species in the Baltic Sea. Modeling results suggest that metabolic prioritization is an important component of the individual response to hypoxia, that it interacts with other components to produce realistic community dynamics, and that different species may prioritize differently. It is thus suggested that declines in feeding activity, assimilation efficiency, and successful reproduction – in addition to low food availability and changing habitat use – are all important drivers of the community response to hypoxia. Meta-analysis results also provide evidence that the dominant predator in the study system prioritizes among metabolic demands, and that these priorities may change as oxygen declines. Going forward, experiments and models should explore how differences in priorities within and among communities drive responses to environmental degradation. This will help management efforts to tailor recovery programs to the physiological needs of species within a given system.more » « less
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