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Groundwater structures fish growth and production across a riverscape

Landscapes are composed of habitat patches and conditions that vary across space and time. While habitat variability and complexity can support important ecological processes and ecosystem services, the dynamic nature of habitats can also constrain organismal growth and production as optimal conditions are fleeting. In riverine ecosystems, groundwater discharge to streams stabilises water temperature and flow regimes, thus mediating how habitat complexity is expressed. Yet, how stable habitats structure growth and production within the broader landscape matrix is not well understood. In this study, we explored the effects of groundwater on spatiotemporal variation in growth and production for juvenile Yellowstone cutthroat trout ( Oncorhynchus virginalis bouvieri ) across the upper Snake River catchment, Wyoming, USA. We combined machine learning techniques and remotely sensed landscape data to estimate groundwater availability across the river network, which we linked to stream temperature regimes and conspecific density. We then used Bayesian hierarchical models to quantify the effects of temperature, density and groundwater on spatiotemporal variation in fish growth and production in 52 focal reaches. Finally, we predicted body size trajectories and trends in total production continuously over both space and time to understand the effect of groundwater at the riverscape scale. Groundwater discharged to streams where topography changes abruptly in valley-bottom areas underlain by coarse glacial deposits. Groundwater stabilised temperature regimes and was associated with high trout densities. Temperature and density, in turn, interacted to influence growth rates: growth increased strongly with temperature, but this effect was reduced when density was high. Accordingly, variation in groundwater availability among stream reaches diversified growth and production regimes. In reaches with low groundwater availability, growth and production declined over time from summer maxima. In contrast, in reaches with high groundwater availability, temporal trends in growth and production were hump-shaped—peaking in autumn—and mean production was greater. At the riverscape scale, temporal asynchrony in growth rates generated convergent spatial variation in growth capacity, but—when combined with density—led to the formation of distinct hotspots of production. Our results demonstrate how groundwater, an important driver of aquatic ecosystem heterogeneity, structures trout growth and production across space and time. Importantly, rare, but stable habitats may disproportionately affect ecological processes and serve as key sources of population diversity at larger spatial scales.

Wyoming

Integrating Sr isotopes, microchemistry, and genetics to reconstruct Salmonidae species and life history

Recent approaches to fisheries research emphasize the importance of the coproduction of knowledge in building resilient and culturally mindful fisheries management frameworks. Despite widespread recognition of the need for Indigenous knowledge and historical reference points as baseline data, archaeological data are rarely included in conservation biology research designs. Here we propose a novel multiproxy method to learn from former fisheries stewards by generating archaeological data on past salmonid population parameters. We used a newly developed, high throughput qPCR (HT-qPCR) chip, originally designed for environmental DNA (eDNA), for species identification of archaeological salmonid vertebrae. We combine this with the laser ablation split-stream (LASS) approach to identify ocean-migration versus freshwater residency. We test this multidisciplinary approach using both contemporary and archaeological salmonid samples and new radiocarbon dates from the Tronsdal Site on the Skagit River, Washington State, USA. This is a useful approach for extracting information about Salmonidae species and life history diversity from archaeological remains to reconstruct historic baselines for several population parameters in anadromous species with long periods of freshwater residency. The approach outlined in this paper may be particularly useful for research investigating past fisheries dynamics, offering hundreds to thousands of years of temporal depth for modern fisheries management, harvest policies, restoration ecology, and conservation biology.

Idaho, Oregon, Washington

Expression of corticoid-regulatory genes in the gills of Atlantic salmon (Salmo salar) parr and smolt and during salinity acclimation

In teleost fishes, cortisol is the major corticoid and has both glucocorticoid and mineralocorticoid actions. However, how fish tissues discriminate between these distinct corticosteroid actions is unclear. In mammals, the major factors responsible for intracellular corticosteroid regulation are glucocorticoid receptors ( gr s) and the mineralocorticoid receptor ( mr ), but their role in osmoregulation of fish is unclear. 11β-hydroxysteroid dehydrogenases ( hsd11b s) control the levels of intracellular corticosteroids by converting from bioactive forms to inert forms. To investigate how Atlantic salmon ( Salmo salar ) respond to cortisol in different physiological or environmental conditions, we performed comparisons of parr and smolt, and osmotic challenge experiments to examine the physiological responses and gill transcript levels of genes underlying cortisol-signalling, including gr1 , gr2 , mr , hsd11b2 and hsd11b3 . Because cortisol may interact with growth hormone and prolactin during salinity changes, transcript levels encoding growth hormone receptors ( ghr1 , ghr2 ) and the prolactin receptor ( prlr ) were also examined. Hsd11b2 transcript levels in seawater-acclimated fish were consistently lower compared to fish acclimated to fresh water. After transfer to seawater, prlr transcript levels in fish significantly decreased and transcript levels of ghr1 , ghr2 and hsd11b3 showed no change or were slightly higher than those of freshwater control groups. Gr1 , gr2 and mr transcript levels were slightly but consistently higher in fish acclimated to fresh water relative to seawater. Our results indicate that changes in corticosteroid receptor and hsd11b2 transcript levels in the gills may be important mechanisms that regulate corticoid signals to achieve ion homeostasis in Atlantic salmon.

Journal of Fish Biology

Variable pH habitats could help prepare stone crabs for coastal acidification

Coastal acidification is being exacerbated by terrestrial organic inputs, especially after high precipitation events. Florida’s rainy season coincides with stone crab ( Menippe mercenaria) reproduction, and pH extremes could limit future harvests by reducing reproductive output. Populations that experience pH variability can serve as “natural laboratories” for estimating the tolerance of a species to coastal acidification. Here, we conducted a series of experiments to determine if ovigerous stone crabs conditioned in more variable pH habitats (seagrass) would result in faster embryonic development, greater hatching success, and higher larval survival relative to crabs conditioned in habitats with lower pH variability (sandy habitats). After field conditioning, crabs were transported to the laboratory and randomly acclimated to either a control pH (pH = ~ 7.90) or a reduced pH condition (pH = ~ 7.60) until larval release. The rate of embryo development was slower in the laboratory reduced pH treatment, however, there were no observable field condition effects on embryo development rate. Crabs conditioned in the more pH variable seagrass habitat did have greater hatching success and higher larval survival than crabs in less pH variable sandy habitats; however, larval survival was low across all treatments. These results suggest that the pH variability experienced in seagrass habitats during brooding may serve as a mechanism for stone crabs to acclimatize to extremes in seawater pH.

Marine Biology

Light-dependent activity of deepwater sculpin (Myoxocephalus thompsonii) across substrates with and without predation risk

Light availability strongly influences predator–prey interactions in deepwater ecosystems, where visual constraints shape both foraging success and prey behavior. The behavioral response of deepwater sculpin ( Myoxocephalus thompsonii ) to siscowet lake trout ( Salvelinus namaycush siscowet ) was studied under ecologically relevant light intensities spanning several orders of magnitude typical of daytime downwelling light in the 20–100 m depth range of Lake Superior. Trials were conducted over varying substrates (gravel, sand, and black fabric). Deepwater sculpin showed a significant preference for gravel over sand and black fabric. In the absence of siscowet, sculpin movement frequency increased as light intensity decreased. Sculpin reaction distance to siscowet was influenced by both light and substrate. Reaction distance was shortest at low light intensities, peaked at intermediate light intensities (3.05 × 10⁹ to < 6.0 × 10⁹ photons m⁻2 s⁻1), and declined again at the highest light intensities tested. In the presence of siscowet, sculpin activity was suppressed at the upper end of the light range (≥ 6.0 × 10⁹ photons m⁻2 s⁻1). The greatest increase in movement occurred between 6.0 × 10⁹ and 3.05 × 10⁹ photons m⁻2 s⁻1, which corresponded to the range where siscowet prey capture declined, suggesting sculpin exploit this low-light window to move with reduced risk. Reduced activity in the presence of predators is common among cryptic species, and our findings suggest that sculpin restrict movement at higher light levels to avoid detection by siscowet.

Wisconsin

Navigating uncertainty and competing objectives: Spring chinook salmon recovery in the upper Willamette river

Globally, anadromous fish populations are threatened with extinction due to multiple factors, including river impoundments that block migration, widespread alteration of physical habitat, temperature, and flow regimes, commercial and recreational harvest, changes in biological communities, and long-term climate trends. In this synthesis, we describe how physical and biological factors, policy and law, and public resource allocation affect management of spring-run Chinook Salmon ( Oncorhynchus tshawytscha ) in the Upper Willamette River (UWR), Oregon, USA. Efforts to restore salmon populations often involve trade-offs, for example, water management decisions must account for flood control, agricultural and municipal water use, and management of other species. UWR Chinook Salmon recovery efforts are also governed by multiple laws, policies, and action agencies that in some cases have incongruent objectives. Competing and conflicting objectives, combined with uncertainty regarding the outcomes of proposed management actions, create significant challenges for decision makers. Resource limitations, regulatory constraints, and incongruent objectives have created challenges for managers in the UWR, but opportunities exist for improved adaptive management and collaboration, which could build trust and engagement among stakeholders and ultimately support the successful implementation of decision tools. A suite of decision support tools that have been used in the Willamette River basin and elsewhere across the range of Pacific salmon have the potential to enhance decision processes. These tools can be effective when decision makers and stakeholders commit to long-term, collaborative utilization of the tools for decision making, and when available resources and regulatory environments are conducive to implementation and adaptive refinement of both the decision-support tools and management plans. The general biological and institutional principles discussed for UWR Chinook Salmon in this work apply more generally to populations of anadromous fishes across their range, especially to populations in highly regulated river systems.

Oregon

Unprecedented burning in tropical peatlands during the 20th century compared to the previous two millennia

Tropical peatland wildfire incidence has risen in recent decades, driven by drainage for land use and intensified by severe droughts with global climate change. These disturbances have altered vegetation structure, disrupted ecosystem functioning, and increased carbon emissions, particularly in Southeast Asia. However, the long-term history and characteristics of wildfires in tropical peatlands remain largely unknown. Here, we compiled fifty-eight macro-charcoal records from peatlands across the tropics, ranging from lowland forested to montane peatlands, to assess millennia-scale changes and controlling factors of tropical peatland burning. We divided the datasets into four main sub-regions: Neotropical, Afrotropical, Indomalayan and Australasian ecoregions to explore regional variability. Tropical peatlands had high burning levels between 0 and 850 ce , followed by a relatively low and stable period until a marked increase during the 20th century. The general trend in tropical peatland burning follows changes in global temperature, and climate variables that control the length and severity of drought events have a notable influence on peat burning before 1900 ce . During the 20th century, regional differences were observed, with declining fire trends in the Neotropical and Afrotropical regions and increasing fire trends in the Indomalayan and Australasian regions. This difference is likely attributable to human activities, and such intervention is also evident in palm swamps and hardwood swamps under similar wet, weakly seasonal climates. With the increase in anthropogenic pressures on peatlands and greater climate variability, future wildfires in peatlands are likely to become more frequent and widespread across all tropical ecoregions. Conservation and sustainable land-use practices could be used to mitigate and control peatland burning and protect these carbon-rich sinks.

Global Change Biology

Comparative assessment of STIC sensors, streamflow and rain gauges for quantifying river connectivity in intermittent systems

In intermittent stream systems, including those occurring in Texas, USA, the severity of low-flow conditions, duration of seasonal disconnection, and frequency of no-flow events have been amplified by drought. Documentation of these no-flow events is necessary to evaluate ecosystem health. However, many intermittent reaches remain un-gauged given that perennial river sec-tions are often prioritized for gauge placement. Our objectives were to 1) document stream flow using Stream Temperature, Intermittency, and Conductivity (STIC) loggers to determine the frequency and duration of no-flow events in intermittent tributaries of the Colorado River, Texas and 2) compare logger data to publicly available data from streamflow discharge and precipitation gauge networks to understand differences among these data types for drying event characterization. We use these comparisons to summarize benefits and limitations of the application of in-stream data loggers. STIC loggers were deployed at 19 sites, one in each pool and riffle habitat of a stream reach. STIC loggers recorded a measurement of relative conductance every six hours from June 2022 to March 2024, which was used to determine the presence or absence of flow connectivity in a reach. No-flow duration among intermittent reaches varied between 37 and 270 days across tributaries during an ongoing drought in the study area. Overall, logger data was more precise than discharge data for characterizing no-flow events or precipitation data when documenting presence of water in the stream channel due to runoff. Lack of discharge gauges in intermittent tributaries left large sections of stream reaches undocumented and resulted in mischaracterization of flow patterns. Drought severity across the tributaries did not follow longitudinal patterns that would be expected by the climatic precipitation gradient of the study area. More research is needed to determine if factors such as population size affect severity. Likewise, precipitation data did not correlate well with logger water presence data, lacking consideration for groundwater recharge, soil hydrophobicity, and surface compaction. This study shows that to monitor no-flow events, detailed spatial datasets are necessary and that STIC loggers are useful tools that provide data to fill spatial information gaps and facilitate more accurate flow characterization and water presence data in intermittent systems.

Texas

Ice sheet dynamics drive pronounced changes in the subsurface freshwater-saltwater interface

Saltwater is migrating into freshwater aquifers globally with water quality and biogeochemical implications, yet saltwater intrusion in glaciated regions is sparsely investigated. Field observations suggest that groundwater head in glaciated systems is influenced by ice sheet forcings and provides evidence that seawater infiltrated into offshore aquifers during past deglaciation events. To understand links between ice sheet dynamics, groundwater head, and saltwater intrusion, we use numerical models to explore the effects of deglaciation on nearshore head and salinity distributions. We find that ice sheet thinning diminishes groundwater head, and the resulting shift in subsurface pressure gradients drives rapid landward movement of the subsurface freshwater-saltwater interface up to 4.0 km or 1.3 m per m ice sheet loss. Results highlight an overlooked saltwater intrusion mechanism that aligns with field observations and affects glaciated coastlines undergoing ice sheet retreat, underscoring the need to consider this mechanism in studies of contemporary coastal water quality.

Geophysical Research Letters

Method for estimating water use and interbasin transfers of freshwater and wastewater in an urbanized basin

Techniques for management of drainage basins that use water budgets to balance available water resources with actual or anticipated water use require accurate and precise estimates of basin withdrawals, interbasin transfers of freshwater, unaccounted-for use, water use, consumptive use, inflow and infiltration, basin return flow, and interbasin transfers of wastewater. Frequently, interbasin transfers of freshwater and wastewater are not included in basin water budgets because they occur within public water-delivery and wastewater-collection systems. A new 10-step method was developed to improve estimates of inflow and infiltration and interbasin transfers using readily available statewide data. The accuracy and precision of water-use estimates determined by this method are improved through careful application of coefficients for small users and the use of metered values for large users. The method was developed and tested with data for the Ten Mile River Basin in southeastern Massachusetts. This report uses examples from the basin to illustrate each step of the method.

Massachusetts

Transit time modeling framework for predicting freshwater salinization in urban catchments

The salinity of inland freshwaters is rising globally, particularly in urban watersheds where winter road deicers are widely applied. Attributing stream salinity dynamics to specific sources and transport pathways remains challenging due to episodic salt inputs, engineered drainage, and strong coupling between hydrology and subsurface storage. We present a modeling framework that couples climate-driven deicer build-up and wash-off with transient transit time distribution theory to simulate salt transport through drainage, interflow, and groundwater pathways. Applied to an urban watershed in Northern Virginia (USA), the model reproduces ten years of high-frequency stream salinity measurements across daily-to-decadal timescales. The calibrated model implies an average deicer application of 206 tonnes Cl −1 yr −1 , or roughly one 20 kg bag of rock salt person −1 yr −1 when normalized by the 20,000 people living in the watershed. In winter months, higher infiltration routes a large fraction of snowmelt and deicers into shallow subsurface pathways, enhancing vadose-zone and interflow contributions to stream salinity. Limited subsurface storage capacity and seasonal hydrologic turnover flush excess chloride from the vadose zone and groundwater during subsequent summer storms. By linking climate-driven deicer inputs, hydrologic connectivity, and stream water age, the framework provides a transferable basis for diagnosing and managing freshwater salinization in urban watersheds.

Virginia

Dietary bioavailability of uranium to a model freshwater invertebrate

Uranium (U) mining increases environmental exposures. Understanding how U is taken up by organisms can aid in evaluating the potential for bioaccumulation and toxicity. Although the importance of aqueous geochemical speciation is well recognized for U bioavailability after dissolved exposures, far less is known about the processes controlling U bioavailability after dietary exposures. This study characterizes the biogeochemical drivers of dietary U uptake in the freshwater snail Lymnaea stagnalis in laboratory experiments. Solids tested included benthic diatoms pre-exposed to dissolved U(VI), soils from contaminated U mine sites, and colloidal hydrous ferric oxide (HFO) synthesized in the presence of dissolved U(VI) or with U complexed by natural organic matter (NOM). Results showed that U was bioavailable from all solids. Uranium assimilation efficiency (AE), a proxy for dietary U bioavailability, varied among solids. AE was lowest for the U-contaminated soils (25 ± 17%) and highest for the U-laden diatoms (71 ± 13%). AE varied slightly among HFO preparations, suggesting modest influences of NOM and iron on U bioavailability. Increases in dietary U exposures reduced feeding rates, and the extent of feeding inhibition appeared inversely related to U bioavailability. The high U assimilation and range of bioavailability have implications for toxicity risks inferred without considering dietary uptake.

Environmental Science and Technology

The effect of temperature, flow, density and disease on smallmouth bass (Micropterus dolomieu) early life growth rate

Individual growth rates of fish are influenced by gradually changing environmental conditions and rapid-onset events. However, there is a paucity of information on drivers of spatiotemporal variability in growth rates of fishes across large spatial extents. Using a 36-year dataset of smallmouth bass ( Micropterus dolomieu ) length-at-age data across 10 river reaches in Pennsylvania, USA ( n = 54,068 individuals), we estimated annual early life growth rates (mm year −1 ) and assessed the effects of relative abundance, summer temperature, summer streamflow rate and a disease outbreak on growth rate. We found that temperature had a positive effect on growth rate, varied spatially, and had a stronger effect in diseased reaches. There was also evidence for an impact of disease on early life growth through density-dependent mechanisms, with growth rates increasing from ∼85 to 101 mm·year −1 following disease-related decreases in abundance. This study adds to our understanding of the factors that shape early life growth, including rapid-onset events like disease which can have immediate and lasting effects on growth rate trajectory.

Pennsylvania

Disturbance is the primary determinant of food chain length when the top predator is constant

Food chain length (FCL) is a primary determinant of food web structure and is hypothesized to be influenced by habitat size, productivity, and disturbance. Understanding the environmental characteristics that determine food chain length can assist in understanding how food webs may be impacted due to changes in habitats and environmental characteristics. This study examines the impact of hydrologic disturbance on stream food webs when the top predator is constant. We analyzed FCL in less disturbed groundwater flashy streams and more disturbed runoff flashy streams using stable isotopes. Despite no difference in species richness or fish density, food chains in more disturbed streams had a lower FCL compared to food chains in more stable streams. Assemblage analysis showed that flow regime and drainage area significantly impacted individual species abundances. The more disturbed runoff flashy streams had higher proportions of primary consumer fish, such as the algivorous Campostoma sp. (Stonerollers), which likely drives the reduced FCL. Drainage area and land cover had non-significant relationships with FCL. Shifting community structure due to hydrologic variability likely leads to differences in diet of Micropterus dolomieu (Smallmouth Bass), and thus a difference in FCL.

Arkansas, Missouri, Oklahoma

Beyond the mangroves: A global synthesis of tidal forested wetland types, drivers and future information opportunities

There is increasing awareness of the global diversity of tidal forested wetlands (TFWs) and their significance in the provision of ecosystem services. These ecosystems, including mangrove forests, tidal freshwater forested wetlands, supratidal forests and transitional forests together span multiple climatic zones, geomorphic settings, and inundation and salinity regimes. We utilise case studies across five continents to demonstrate the state of knowledge among TFWs. Intertidal mangroves are the best-defined of the TFWs thanks to decades of research on their geomorphology, hydrology and ecology across their broad distribution. Non-mangrove forest settings, however, demonstrate more diverse hydrological, biochemical and vegetation conditions. In many cases, non-mangrove forests are situated at upper intertidal or supratidal elevations, where surface waters and groundwater are subject to interactions between tides freshwater inputs. Salinity datasets show variations ranging from tidal freshwater forested wetlands and ‘low-salinity mangroves’ to mesohaline or marine salinities, often with high temporal variability. While the floristic composition of non-mangrove forests vary among biogeographic regions, locally dominant TFW species are commonly distributed beyond the tidal niche into non-tidal wetland and upland forests. This presents challenges for traditional remote sensing approaches to ecosystem mapping, which are mostly lacking for non-mangrove forests. Geomorphic approaches and developments in machine learning offer opportunities to address this.

Preprint

Structural evolution of iron coordination in proteins across Earth’s oxygenation history

Protein metal-binding sites support essential biological functions shaped by protein fold, subunit interactions, and cofactor chemistry. Because these sites encode both biochemical function and environmental constraint, they offer a route to connect protein evolution with changes in Earth’s surface environment through time. Of particular interest is iron (Fe), the most widely used metal in biology and a cofactor central to both anaerobic and aerobic metabolism. Here, we systematically compare the immediate chemical environments of functionally essential Fe-binding sites in three-dimensional protein structures to test whether Fe coordination spheres differ across oxygen contexts. Using a curated dataset of experimentally determined structures, we identify a clear shift in the local chemistry of Fe-binding environments from anaerobic to aerobic proteins. Aerobic Fe sites are significantly more hydrophilic than anaerobic ones, and amino-acid composition analyses show reduced cysteine use in aerobic Fe-binding neighborhoods. These patterns suggest that as Earth’s surface environments became more oxygenated, proteins retained Fe as a core redox metal while reconfiguring local coordination chemistry in ways less vulnerable to oxidative damage. More broadly, this study introduces and applies the Coordination Sphere Analysis and Comparison (CSAC) workflow, an open and archived Python workflow for extracting local metal-binding environments from structure datasets, providing a framework for linking metalloprotein structure to evolutionary and geobiological transitions across Earth history.

Discover Life

Decoding the oxidative digestion mechanism for polystyrene nanoplastic detection in the Great Lakes using a customizable Raman spectral processing algorithm

Despite the concerns surging around nanoplastics (NPs) regarding their prevalence and bioavailability in freshwater systems, robust detection of NPs in complex environmental matrices is hindered by the lack of standardized sample pretreatment and a mechanistic understanding of oxidative digestion. Here, we systematically investigate the interaction between hydrogen peroxide (H 2 O 2 ) and polystyrene (PS) NPs during digestion in deionized (DI) water and four environmental matrices from in and around the Great Lakes Basin. To facilitate high-throughput analysis, we develop Pre_peak, a customizable Raman spectral processing algorithm that achieves >99% accuracy for both NP identification and interference rejection, allowing reliable NP quantification via pixel counting and systematic decoding of the oxidative digestion mechanisms. In DI water, varying H 2 O 2 doses from 0 to 30% has negligible effects on the recovery and Raman signal intensity of PS NPs over 24 hours of digestion. However, morphological changes and aggregation of PS NPs are observed when the H 2 O 2 dose exceeds 20%. Prolonged digestion further leads to progressive NP loss. In natural waters, the optimal dosage and digestion duration depend on matrix characteristics, including dissolved organic matter (DOM) and ion composition. This study provides mechanistic insights into NP–oxidant interactions and underscores the need for matrix-tailored digestion protocols to advance standardized NP detection in freshwater environments.

Great Lakes

Diverging mineral chemistry of iron and nickel throughout Earth’s changing redox conditions reveals foundation for their evolution as protein cofactors

Iron (Fe) and nickel (Ni) were both foundational to early metabolism, yet their biological trajectories diverged as Earth’s surface redox state changed. Here, we integrate mineral chemistry network analysis, protein metal-site coordination-sphere analysis, and curated redox comparisons to test how geochemistry and metalloprotein architecture co-evolved. Mineral network analyses show broader electronegativity variation and network diversity for Fe-bearing minerals through time relative to Ni-bearing minerals. In structural analyses of protein metal centers in a combined Fe/Ni protein structure set, it is shown that Fe- and Ni-associated environments differ in amino-acid composition, hydropathy structure, and cysteine representation. The greater chemical diversity and electronegativity variation in Fe minerals mirror the higher redox and structural versatility of Fe-binding proteins. The presence of Fe in a broader range of mineral and protein environments demonstrates the chemical adaptability of the metal, from the anoxic Archean to oxidative Earth surface conditions following the Great Oxidation Event. Iron, with its broad redox potential range in Fe-oxidoreductases, has a central role in both anaerobic and aerobic metabolisms. Nickel, by contrast, is less widespread in biology. Today, Ni is predominantly employed in deeply branching anaerobic pathways and by proteins with narrower redox potential ranges. Our results show that evolutionary processes, constrained by metal chemistry, habitually utilize Fe as a redox generalist while retaining Ni in specialized roles. The divergent paths of Ni and Fe, from rocks to proteins, demonstrate the intimate relationship between planetary geochemistry and metabolic origins on Earth and suggest that Fe/Ni geochemistry may inform habitability assessments in extraterrestrial environments when interpreted within specific planetary environmental contexts.

Life