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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. 

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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. 

Deoxygenation worldwide is increasing in aquatic systems with implications for organisms' biology, communities and ecosystems. Eastern Baltic cod has experienced a strong decline in mean body condition (i.e. weight at a specific length) over the past 20 years with effects on the fishery relying on this resource. The decrease in cod condition has been tentatively linked in the literature to increased hypoxic areas potentially affecting habitat range, but also to benthic prey and/or cod physiology directly. To date, no studies have been performed to test these mechanisms. Using otolith trace element microchemistry and hypoxia-responding metrics based on manganese (Mn) and magnesium (Mg), we investigated the relationship between fish body condition at capture and exposure to hypoxia. Cod individuals collected after 2000 with low body condition had a higher level of Mn/Mg in the last year of life, indicating higher exposure to hypoxic waters than cod with high body condition. Moreover, lifetime exposure to hypoxia was even more strongly correlated to body condition, suggesting that condition may reflect long-term hypoxia status. These results were irrespective of fish age or sex. This implies that as Baltic cod visit poor-oxygen waters, perhaps searching for benthic food, they compromise their own performance. This study specifically sheds light on the mechanisms leading to the low condition of cod and generally points to the impact of deoxygenation on ecosystems and fisheries. 

Accurate age data are essential for reliable fish stock assessment. Yet many stocks suffer from inconsistencies in age interpretation. A new approach to obtain age makes use of the chemical composition of otoliths. This study validates the periodicity of recurrent patterns in 25 Mg, 31 P, 34 K, 55 Mn, 63 Cu, 64 Zn, 66 Zn, 85 Rb, 88 Sr, 138 Ba, and 208 Pb in Baltic cod (Gadus morhua) otoliths from tag–recapture and known-age samples. Otolith P concentrations showed the highest consistency in seasonality over the years, with minima co-occurring with otolith winter zones in the known-age otoliths and in late winter – early spring when water temperatures are coldest in tagged cod . The timing of minima differs between stocks, occurring around February in western Baltic cod and 1 month later in eastern Baltic cod; seasonal maxima are also stock-specific, occurring in August and October, respectively. The amplitude in P is larger in faster-growing western compared with eastern Baltic cod. Seasonal patterns with minima in winter – late spring were also evident in Mg and Mn, but less consistent over time and fish size than P. Chronological patterns in P, and to a lesser extent Mg and Mn, may have the potential to supplement traditional age estimation or to guide the visual identification of translucent and opaque otolith patterns used in traditional age estimation. 

Otolith chemistry has gained increasing attention as a tool for analyzing various aspects of fish biology, such as stock dynamics, migration patterns, hypoxia and pollution exposure, and connectivity between habitats. While these studies often assume otolith elemental concentrations reflect environmental conditions, physiological processes are increasingly recognized as a modulating and/or controlling factor. In particular, biomineralization—the complex, enzyme-regulated construction of CaCO3 crystals scaffolded by proteins—is believed to play a critical role in governing otolith chemical patterns. This review aims to summarize the knowledge on otolith composition and biophysical drivers of biomineralization, present hypotheses on how biomineralization should affect element incorporation, and test the validity thereof with selected case studies. Tracers of environmental history are assumed to be dominated by elements that substitute for Ca during crystal growth or that occur randomly trapped within the crystal lattice. Strontium (Sr) and barium (Ba) largely comply with the biomineralization-based hypotheses that otolith element patterns reflect environmental concentrations, without additional effects of salinity, but can be influenced by physiological processes, typically exhibiting decreasing incorporation with increasing growth. Conversely, tracers of physiology are assumed to be elements under physiological control and primarily occur protein-bound in the otolith’s organic matrix. Physiological tracers are hypothesized to reflect feeding rate and/or growth, decrease with fish age, and exhibit minimal influence of environmental concentration. The candidate elements phosphorus (P), copper (Cu) and zinc (Zn) confirm these hypotheses. Magnesium (Mg) is believed to be randomly trapped in the crystal structure and hence a candidate for environmental reconstruction, but the response to all examined drivers suggest Mg to be coupled to growth. Manganese (Mn) substitutes for Ca, but is also a co-factor in matrix proteins, and therefore exhibits otolith patterns reflecting both environmental (concentration and salinity) and physiological (ontogeny and growth) histories. A consistent temperature response was not evident across studies for either environmental or physiological tracers, presumably attributable to variable relationships between temperature and fish behavior and physiology (e.g., feeding rate, reproduction). Biomineralization thus has a controlling effect on otolith element concentrations for elements that are linked with somatic growth, but not for elements that substitute for Ca in the crystal lattice. Interpretation of the ecological significance of patterns from field samples therefore needs to consider the impact of the underlying biomineralization processes of the element in question as well as physiological processes regulating the availability of ions for inclusion in the growing crystal lattice. Such understanding will enhance the utility of this technique to address fisheries management questions.