Is Ocean Acidification Really a Threat to Marine Calcifiers? A Systematic Review and Meta-Analysis of 980+ Studies Spanning Two Decades.

Ocean acidification is considered detrimental to marine calcifiers, but mounting contradictory evidence suggests a need to revisit this concept. This systematic review and meta-analysis aim to critically re-evaluate the prevailing paradigm of negative effects of ocean acidification on calcifiers. Based on 5153 observations from 985 studies, many calcifiers (e.g., echinoderms, crustaceans, and cephalopods) are found to be tolerant to near-future ocean acidification (pH ≈ 7.8 by the year 2100), but coccolithophores, calcifying algae, and corals appear to be sensitive. Calcifiers are generally more sensitive at the larval stage than adult stage. Over 70% of the observations in growth and calcification are non-negative, implying the acclimation capacity of many calcifiers to ocean acidification. This capacity can be mediated by phenotypic plasticity (e.g., physiological, mineralogical, structural, and molecular adjustments), transgenerational plasticity, increased food availability, or species interactions. The results suggest that the impacts of ocean acidification on calcifiers are less deleterious than initially thought as their adaptability has been underestimated. Therefore, in the forthcoming era of ocean acidification research, it is advocated that studying how marine organisms persist is as important as studying how they perish, and that future hypotheses and experimental designs are not constrained within the paradigm of negative effects.

Shark teeth can resist ocean acidification.

Ocean acidification can cause dissolution of calcium carbonate minerals in biological structures of many marine organisms, which can be exacerbated by warming. However, it is still unclear whether this also affects organisms that have body parts made of calcium phosphate minerals (e.g. shark teeth), which may also be impacted by the 'corrosive' effect of acidified seawater. Thus, we examined the effect of ocean acidification and warming on the mechanical properties of shark teeth (Port Jackson shark, Heterodontus portusjacksoni), and assessed whether their mineralogical properties can be modified in response to predicted near-future seawater pH (-0.3 units) and temperature (+3°C) changes. We found that warming resulted in the production of more brittle teeth (higher elastic modulus and lower mechanical resilience) that were more vulnerable to physical damage. Yet, when combined with ocean acidification, the durability of teeth increased (i.e. less prone to physical damage due to the production of more elastic teeth) so that they did not differ from those raised under ambient conditions. The teeth were chiefly made of fluorapatite (Ca5 (PO4 )3 F), with increased fluoride content under ocean acidification that was associated with increased crystallinity. The increased precipitation of this highly insoluble mineral under ocean acidification suggests that the sharks could modulate and enhance biomineralization to produce teeth which are more resistant to corrosion. This adaptive mineralogical adjustment could allow some shark species to maintain durability and functionality of their teeth, which underpins a fundamental component of predation and sustenance of the trophic dynamics of future oceans.

Calcifiers can Adjust Shell Building at the Nanoscale to Resist Ocean Acidification.

Ocean acidification is considered detrimental to marine calcifiers based on laboratory studies showing that increased seawater acidity weakens their ability to build calcareous shells needed for growth and protection. In the natural environment, however, the effects of ocean acidification are subject to ecological and evolutionary processes that may allow calcifiers to buffer or reverse these short-term negative effects through adaptive mechanisms. Using marine snails inhabiting a naturally CO2 -enriched environment over multiple generations, it is discovered herein that they build more durable shells (i.e., mechanically more resilient) by adjusting the building blocks of their shells (i.e., calcium carbonate crystals), such as atomic rearrangement to reduce nanotwin thickness and increased incorporation of organic matter. However, these adaptive adjustments to future levels of ocean acidification (year 2100) are eroded at extreme CO2 concentrations, leading to construction of more fragile shells. The discovery of adaptive mechanisms of shell building at the nanoscale provides a new perspective on why some calcifiers may thrive and others collapse in acidifying oceans, and highlights the inherent adaptability that some species possess in adjusting to human-caused environmental change.

Norethindrone alters mating behaviors, ovary histology, hormone production and transcriptional expression of steroidogenic genes in zebrafish (Danio rerio).

The impact of progestins (i.e. synthetic forms of progesterone) on aquatic organisms has drawn increasing attention due to their widespread occurrence in the aquatic environments and potential effects on the endocrine system of fish. In this study, the effects of norethindrone (NET, a progestin) on the reproductive behavior, sex hormone production and transcriptional expressions were evaluated by exposing female zebrafish to NET at 0, 3.1, 36.2 and 398.6 ng L-1 for 60 days. Results showed that NET impaired the mating behaviors of female at 36.2 and 398.6 ng L-1 exhibited by males and increased the frequency of atretic follicular cells in the ovary exposed to NET at 398.6 ng L-1. As for sex hormones, plasma testosterone concentration in zebrafish increased, while estradiol concentration decreased. Up-regulation of genes (Npr, Mpra, Mprß, Fshß, Lß, Tshb, Nis and Dio2) was detected in the brain of fish exposed to NET at 398.6 ng L-1. The transcriptional levels of genes (Esr1, Vtg1, Ar, Cyp19a, Cyp11b and Ptgs2) were generally inhibited in the ovary of zebrafish by NET at 398.6 ng L-1. Moreover, the transcripts of genes (Vtg1, Esr1, Ar and Pgr) in the liver were reduced by NET at 36.2 and 398.6 ng L-1. Our findings suggest that NET can potentially diminish the of fish populations not only by damaging their reproductive organs, but also by altering their mating behavior through the changes in the expressions of genes responsible for the production of sex hormones.

How calorie-rich food could help marine calcifiers in a CO2-rich future.

Increasing carbon emissions not only enrich oceans with CO2 but also make them more acidic. This acidifying process has caused considerable concern because laboratory studies show that ocean acidification impairs calcification (or shell building) and survival of calcifiers by the end of this century. Whether this impairment in shell building also occurs in natural communities remains largely unexplored, but requires re-examination because of the recent counterintuitive finding that populations of calcifiers can be boosted by CO2 enrichment. Using natural CO2 vents, we found that ocean acidification resulted in the production of thicker, more crystalline and more mechanically resilient shells of a herbivorous gastropod, which was associated with the consumption of energy-enriched food (i.e. algae). This discovery suggests that boosted energy transfer may not only compensate for the energetic burden of ocean acidification but also enable calcifiers to build energetically costly shells that are robust to acidified conditions. We unlock a possible mechanism underlying the persistence of calcifiers in acidifying oceans.

Testing for thresholds of ecosystem collapse in seagrass meadows.

Although the public desire for healthy environments is clear-cut, the science and management of ecosystem health has not been as simple. Ecological systems can be dynamic and can shift abruptly from one ecosystem state to another. Such unpredictable shifts result when ecological thresholds are crossed; that is, small cumulative increases in an environmental stressor drive a much greater change than could be predicted from linear effects, suggesting an unforeseen tipping point is crossed. In coastal waters, broad-scale seagrass loss often occurs as a sudden event associated with human-driven nutrient enrichment (eutrophication). We tested whether the response of seagrass ecosystems to coastal nutrient enrichment is subject to a threshold effect. We exposed seagrass plots to different levels of nutrient enrichment (dissolved inorganic nitrogen) for 10 months and measured net production. Seagrass response exhibited a threshold pattern when nutrient enrichment exceeded moderate levels: there was an abrupt and large shift from positive to negative net leaf production (from approximately 0.04 leaf production to 0.02 leaf loss per day). Epiphyte load also increased as nutrient enrichment increased, which may have driven the shift in leaf production. Inadvertently crossing such thresholds, as can occur through ineffective management of land-derived inputs such as wastewater and stormwater runoff along urbanized coasts, may account for the widely observed sudden loss of seagrass meadows. Identification of tipping points may improve not only adaptive-management monitoring that seeks to avoid threshold effects, but also restoration approaches in systems that have crossed them.

Mineralogical Plasticity Acts as a Compensatory Mechanism to the Impacts of Ocean Acidification.

Calcifying organisms are considered particularly susceptible to the future impacts of ocean acidification (OA), but recent evidence suggests that they may be able to maintain calcification and overall fitness. The underlying mechanism remains unclear but may be attributed to mineralogical plasticity, which modifies the energetic cost of calcification. To test the hypothesis that mineralogical plasticity enables the maintenance of shell growth and functionality under OA conditions, we assessed the biological performance of a gastropod (respiration rate, feeding rate, somatic growth, and shell growth of Austrocochlea constricta) and analyzed its shell mechanical and geochemical properties (shell hardness, elastic modulus, amorphous calcium carbonate, calcite to aragonite ratio, and magnesium to calcium ratio). Despite minor metabolic depression and no increase in feeding rate, shell growth was faster under OA conditions, probably due to increased precipitation of calcite and trade-offs against inner shell density. In addition, the resulting shell was functionally suitable for increasingly "corrosive" oceans, i.e., harder and less soluble shells. We conclude that mineralogical plasticity may act as a compensatory mechanism to maintain overall performance of calcifying organisms under OA conditions and could be a cornerstone of calcifying organisms to acclimate to and maintain their ecological functions in acidifying oceans.

Impacts of Near-Future Ocean Acidification and Warming on the Shell Mechanical and Geochemical Properties of Gastropods from Intertidal to Subtidal Zones.

Many marine organisms produce calcareous shells as the key structure for defense, but the functionality of shells may be compromised by ocean acidification and warming. Nevertheless, calcifying organisms may adaptively modify their shell properties in response to these impacts. Here, we examined how reduced pH and elevated temperature affect shell mechanical and geochemical properties of common grazing gastropods from intertidal to subtidal zones. Given the greater environmental fluctuations in the intertidal zone, we hypothesized that intertidal gastropods would exhibit more plastic responses in shell properties than subtidal gastropods. Overall, three out of five subtidal gastropods produced softer shells at elevated temperature, while intertidal gastropods maintained their shell hardness at both elevated pCO2 (i.e., reduced pH) and temperature. Regardless of pH and temperature, degree of crystallization was maintained (except one subtidal gastropod) and carbonate polymorph remained unchanged in all tested species. One intertidal gastropod produced less soluble shells (e.g., higher calcite/aragonite) in response to reduced pH. In contrast, subtidal gastropods produced only aragonite which has higher solubility than calcite. Overall, subtidal gastropods are expected to be more susceptible than intertidal gastropods to shell dissolution and physical damage under future seawater conditions. The increased vulnerability to shell dissolution and predation could have serious repercussions for their survival and ecological contributions in the future subtidal environment.

Ecological risk and pollution history of heavy metals in Nansha mangrove, South China.

Owing to the Industrial Revolution in the late 1970s, heavy metal pollution has been regarded as a serious threat to mangrove ecosystems in the region of the Pearl River Estuary, potentially affecting human health. The present study attempted to characterize the ecological risk of heavy metals (Cd, Cr, Cu, Mn, Ni, Pb and Zn) in Nansha mangrove, South China, by estimating their concentrations in the surface sediment. In addition, the pollution history of heavy metals was examined by determining the concentrations of heavy metals along the depth gradient. The phytoremediation potential of heavy metals by the dominant plants in Nansha mangrove, namely Sonneratia apetala and Cyperus malaccensis, was also studied. Results found that the surface sediment was severely contaminated with heavy metals, probably due to the discharge of industrial sewage into the Pearl River Estuary. Spatial variation of heavy metals was generally unobvious. The ecological risk of heavy metals was very high, largely due to Cd contamination. All heavy metals, except Mn, decreased with depth, indicating that heavy metal pollution has been deteriorating since 1979. Worse still, the dominant plants in Nansha mangrove had limited capability to remove the heavy metals from sediment. Therefore, we propose that immediate actions, such as regulation of discharge standards of industrial sewage, should be taken by the authorities concerned to mitigate the ecological risk posed by heavy metals.

Can PFAS threaten the health of fish consumers? A comprehensive analysis linking fish consumption behaviour and health risk.

Despite being phased out for decades, per- and polyfluoroalkyl substances (PFAS) are still widely detected in the environment and accumulated in many aquaculture organisms for human consumption. Thus, there is growing concern about whether fish consumption can cause PFAS-associated health impacts on humans since fish is a vital protein source for global populations. Here, we assess the potential driving factors of fish consumption by analysing the aquaculture, demographic and socio-economic data across 31 provinces/municipalities in China, followed by estimating the health risk of PFAS via fish consumption. We found that per capita fish consumption was primarily driven by fish production and total area for freshwater aquaculture, while urbanization rate and median age of consumers were also important. The health risk of PFAS was low (hazard quotient <1) in most provinces, while urban consumers were more prone to PFAS than rural consumers across all provinces. Since PFAS have been phased out worldwide, their health risk to humans through fish consumption would be lower than previously thought. To reduce PFAS intake for the high-risk populations, we recommend that fish should be well cooked before consumption, preferably using water-based cooking methods, and that alternative protein sources should be consumed more as the substitute for fish.

Can antimony contamination in soil undermine the ecological contributions of earthworms?

As antimony (Sb) has been increasingly used in manufacturing industries (e.g., alloy, polymer and electronics industries), Sb contamination in the soil environment becomes widely reported and has drawn growing attention due to the toxicity of Sb to living organisms. Whether soil-dwelling organisms can tolerate Sb toxicity and maintain their ecological functions remains poorly understood. Using a cosmopolitan, ecologically important earthworm species (Eisenia fetida) as an ideal model organism, we examine the effects of Sb on the physiological, molecular and behavioural responses of earthworms to different levels of Sb contamination in soil (0, 10, 50, 100, 250 and 500 mg/kg). We found that earthworms could tolerate heavy Sb contamination (100 mg/kg) by boosting their antioxidant defence (POD and GST) and immune systems (ACP) so that their body weight and survival rate were sustained (c.f. control). However, these systems were compromised under extreme Sb contamination (500 mg/kg), leading to mortality. As such, earthworms exhibited avoidance behaviour to escape from the Sb-contaminated soil, implying the loss of their ecological contributions to the environment (e.g., increase in soil aeration and maintenance of soil structure). By measuring various types of biomarkers along a concentration gradient, this study provides a mechanistic understanding of how earthworms resist or succumb to Sb toxicity. Since extreme Sb contamination in soil (>100 mg/kg) is rarely found in nature, we are optimistic that the health and performance of earthworms are not influenced by Sb in most circumstances, but regular monitoring of Sb in soil is recommended to ensure the integrity and functioning of soil environment. Further studies are recommended to evaluate the long-term impact of Sb in the soil ecosystem through bioaccumulation and trophic transfer among soil-dwelling organisms.

Transcriptomic responses reveal impaired physiological performance of the pearl oyster following repeated exposure to marine heatwaves.

Marine heatwaves are predicted to become more intense and frequent in the future, possibly threatening the survival of marine organisms and devastating their communities. While recent evidence reveals the adaptability of marine organisms to heatwaves, substantially overlooked is whether they can also adjust to repeated heatwave exposure, which can occur in nature. By analysing transcriptome, we examined the fitness and recoverability of the pearl oyster (Pinctada maxima) after two consecutive heatwaves (24 °C to 32 °C for 3 days; recovery at 24 °C for 4 days). In the first heatwave, 331 differentially expressed genes (DEGs) were found, such as AGE-RAGE, MAPK, JAK-STAT, FoxO and mTOR. Despite the recovery after the first heatwave, 2511 DEGs related to energy metabolism, body defence, cell proliferation and biomineralization were found, where 1655 of them were downregulated, suggesting a strong negative response to the second heatwave. Our findings imply that some marine organisms can indeed tolerate heatwaves by boosting energy metabolism to support molecular defence, cell proliferation and biomineralization, but this capacity can be overwhelmed by repeated exposure to heatwaves. Since recurrence of heatwaves within a short period of time is predicted to be more prevalent in the future, the functioning of marine ecosystems would be disrupted if marine organisms fail to accommodate repeated extreme thermal stress.

Microplastics can aggravate the impact of ocean acidification on the health of mussels: Insights from physiological performance, immunity and byssus properties.

Ocean acidification may increase the risk of disease outbreaks that would challenge the future persistence of marine organisms if their immune system and capacity to produce vital structures for survival (e.g., byssus threads produced by bivalves) are compromised by acidified seawater. These potential adverse effects may be exacerbated by microplastic pollution, which is forecast to co-occur with ocean acidification in the future. Thus, we evaluated the impact of ocean acidification and microplastics on the health of a mussel species (Mytilus coruscus) by assessing its physiological performance, immunity and byssus properties. We found that ocean acidification and microplastics not only reduced hemocyte concentration and viability due to elevated oxidative stress, but also undermined phagocytic activity of hemocytes due to lowered energy budget of mussels, which was in turn caused by the reduced feeding performance and energy assimilation. Byssus quality (strength and extensibility) and production were also reduced by ocean acidification and microplastics. To increase the chance of survival with these stressors, the mussels prioritized the synthesis of some byssus proteins (Mfp-4 and Mfp-5) to help maintain adhesion to substrata. Nevertheless, our findings suggest that co-occurrence of ocean acidification and microplastic pollution would increase the susceptibility of bivalves to infectious diseases and dislodgement risk, thereby threatening their survival and undermining their ecological contributions to the community.

Microplastic accumulation via trophic transfer: Can a predatory crab counter the adverse effects of microplastics by body defence?

Microplastics are considered detrimental to aquatic organisms due to their potential accumulation along food chains. Thus, it is puzzling why some of them appear unaffected by microplastics. Here, we assessed the contribution of water filtration and food consumption to microplastic accumulation in a predatory marine crab (Charybdis japonica) and examined the associated impacts of microplastics (particle size: 5 µm) following ingestion for one week. Results showed that water filtration and food consumption contributed similarly to the accumulation of microplastics, which were distributed among organs in this order: hepatopancreas > guts > gills > muscles. Yet, biomagnification (i.e. accumulation through consumption of microplastic-contaminated mussels) did not occur possibly due to egestion of microplastics. The crabs upregulated detoxification capacity (EROD) and antioxidant defence (GST) in response to the microplastics accumulated in their tissues. However, these defence mechanisms collapsed when the microplastic concentration in hepatopancreas exceeded ~3 mg g-1, leading to severe hepatic injury (elevated AST and ALT) and impaired neural activity (reduced AChE). Our results suggest that marine organisms have an innate capacity to counter the acute effects of microplastics, but there is a limit beyond which the defence mechanisms fail and hence physiological functions are impaired. As microplastic pollution will deteriorate in the future, the fitness and survival of marine organisms may be undermined by microplastics, affecting the stability and functioning of marine ecosystems.

Agricultural activities compromise ecosystem health and functioning of rivers: Insights from multivariate and multimetric analyses of macroinvertebrate assemblages.

Agricultural activities often lead to nutrient enrichment and habitat modification in rivers, possibly altering macroinvertebrate assemblages and hence ecosystem functioning. For the sake of environmental management and conservation, therefore, assessing the impacts of agricultural activities becomes indispensable, especially when these activities are predicted to be intensified in the future. In this study, the plain river network in the Lake Chaohu Basin was chosen to examine how agricultural activities influence the functioning of rivers by assessing land use, water quality, habitat condition and macroinvertebrate assemblages, followed by calculating the macroinvertebrate-based multimetric index (MMI) to indicate overall ecosystem health of the rivers. We found that agricultural activities lowered the diversity of macroinvertebrates (e.g. total number of taxa and Simpson index) primarily due to elevated ammonium concentrations in water and reduced microhabitat types, thereby impairing the habitat integrity and nutrient cycling of rivers. The macroinvertebrate-based MMI was positively correlated with increasing habitat quality but negatively with increasing nutrient concentrations, suggesting its high reliability for indicating the impacts of agricultural activities, which was further substantiated by classification and regression tree (CART) analysis. We recommend analyzing macroinvertebrate assemblages using both multivariate and multimetric approaches to offer a more comprehensive evaluation of the impacts of agricultural activities on ecosystem health. Some environmental (CODMn, NH4+-N and PO43--P) and biological parameters (total number of taxa), however, can be used as good proxies for MMI, when time and resources for gathering information to develop MMI are limited.

Norethindrone causes cellular and hepatic injury in zebrafish by compromising the metabolic processes associated with antioxidant defence: Insights from metabolomics.

Progestins, such as norethindrone (NET), have been increasingly detected in aquatic environments due to their extensive use for medical applications. While NET is notorious for its endocrine disrupting effects, it has been recently shown to cause cellular damage, suggesting its potential impacts on the body defence of organisms. Hence, we examined the histological features and antioxidant defence of zebrafish (Danio rerio) after exposing to NET (50 ng/L and 500 ng/L) for 72 days, followed by analysing its metabolome to explore whether NET disturbs the metabolic processes responsible for antioxidant defence. While acute mortality was not triggered, we found that antioxidant defence was substantially weakened by NET at 500 ng/L (i.e. reduced SOD and GSH levels) and hence liver injury was inflicted (i.e. elevated ALT and MDA levels), as manifested by vacuolization of liver tissues and reduced number of normal cells in the liver. Metabolomic analysis showed that the metabolic processes responsible for antioxidant defence were disrupted by NET (e.g. upregulation of nervonyl carnitine and chenodeoxycholic acid 3-sulfate; downregulation of homolanthionine and acevaltrate) and these changes can undermine antioxidant defence by suppressing Nrf2-ARE and NF-κB pathways that contribute to the synthesis of SOD and GSH. This study demonstrates how NET can compromise the body defence of aquatic organisms via metabolic disruption, suggesting that the impacts of progestins on their fitness are more detrimental than previously thought.

Long-term thermal acclimation drives adaptive physiological adjustments of a marine gastropod to reduce sensitivity to climate change.

Ocean warming is predicted to challenge the persistence of a variety of marine organisms, especially when combined with ocean acidification. While temperature affects virtually all physiological processes, the extent to which thermal history mediates the adaptive capacity of marine organisms to climate change has been largely overlooked. Using populations of a marine gastropod (Turbo undulatus) with different thermal histories (cool vs. warm), we compared their physiological adjustments following exposure (8-week) to ocean acidification and warming. Compared to cool-acclimated counterparts, we found that warm-acclimated individuals had a higher thermal threshold (i.e. increased CTmax by 2 °C), which was unaffected by the exposure to ocean acidification and warming. Thermal history also strongly mediated physiological effects, where warm-acclimated individuals adjusted to warming by conserving energy, suggested by lower respiration and ingestion rates, energy budget (i.e. scope for growth) and O:N ratio. After exposure to warming, warm-acclimated individuals had higher metabolic rates and greater energy budget due to boosted ingestion rates, but such compensatory feeding disappeared when combined with ocean acidification. Overall, we suggest that thermal history can be a critical mediator of physiological performance under future climatic conditions. Given the relatively gradual rate of global warming, marine organisms may be better able to adaptively adjust their physiology to future climate than what short-term experiments currently convey.

Reproductive potential of mosquitofish is reduced by the masculinizing effect of a synthetic progesterone, gestodene: Evidence from morphology, courtship behaviour, ovary histology, sex hormones and gene expressions.

The ever-increasing use of synthetic hormones, especially progestins, for medical applications has drawn growing concerns due to their potential endocrine disrupting effects that may diminish the reproductive outputs of aquatic organisms. Using mosquitofish (Gambusia affinis) as a model species, we tested whether gestodene (GES), a commonly used progestin, can alter the expressions of genes associated with sex hormone synthesis and cause ensuing changes in morphological features, courtship behaviour and oocyte development. After exposing to GES at environmentally relevant concentrations (2.96, 32.9 and 354 ng L-1) for 40 days, we found that GES, especially at 354 ng L-1, induced masculinization of female fish, indicated by the reduced body weight to length ratio and development of gonopodia (i.e. anal fins of male fish). Thus, the males showed less intimacy and mating interest towards the GES-exposed females, indicated by the reduced time spent on attending, following and mating behaviours. While oocyte development was seemingly unaffected by GES, spermatogonia were developed in the ovary. All the aforementioned masculinizing effects of GES were associated with the increased testosterone level and decreased estradiol level, driven by upregulating androgen receptor genes (Arα and Arß). Overall, our findings suggest that progestins could undermine the reproductive potential of aquatic organisms and hence their persistence in the progestin-contaminated environment.

Stress across life stages: Impacts, responses and consequences for marine organisms.

Population dynamics of marine organisms are strongly driven by their survival in early life stages. As life stages are tightly linked, environmental stress experienced by organisms in the early life stage can worsen their performance in the subsequent life stage (i.e. carry-over effect). However, stressful events can be ephemeral and hence organisms may be able to counter the harmful effects of transient stress. Here, we analysed the published data to examine the relative strength of carry-over effects on the juvenile growth of marine organisms, caused by different stressors (hypoxia, salinity, starvation, ocean acidification and stress-induced delayed metamorphosis) confronted in their larval stage. Based on 31 relevant published studies, we revealed that food limitation had the greatest negative carry-over effect on juvenile growth. In the laboratory, we tested the effects of short-term early starvation and hypoxia on the larval growth and development of a model organism, polychaete Hydroides elegans, and assessed whether the larvae can accommodate the early stress to maintain their performance as juveniles (settlement and juvenile growth). Results showed that early starvation for 3 days (∼50% of normal larval period) retarded larval growth and development, leading to subsequent reduced settlement rate and juvenile growth. When the starvation period decreased to 1 day, however, the larvae could recover from early starvation through compensatory growth and performed normal as juveniles (c.f. control). Early exposure to hypoxia for 3 days did not affect larval growth (body length) and juvenile growth (tube length), but caused malformation of larvae and reduced settlement rate. We conclude that the adverse effects of transient stress can be carried across life stages (e.g. larval to juvenile stage), but depend on the duration of stressful events relative to larval period. As carry-over effects are primarily driven by energy acquisition, how food availability varies over time and space is fundamental to the population dynamics of marine organisms.

Linking energy budget to physiological adaptation: How a calcifying gastropod adjusts or succumbs to ocean acidification and warming.

Accelerating CO2 emissions have driven physico-chemical changes in the world's oceans, such as ocean acidification and warming. How marine organisms adjust or succumb to such environmental changes may be determined by their ability to balance energy intake against expenditure (i.e. energy budget) as energy supports physiological functions, including those with adaptive value. Here, we examined whether energy budget is a driver of physiological adaptability of marine calcifiers to the near-future ocean acidification and warming; i.e. how physiological energetics (respiration rate, feeding rate, energy assimilation and energy budget) relates to adjustments in shell growth and shell properties of a calcifying gastropod (Austrocochlea concamerata). We found that ocean warming boosted the energy budget of gastropods due to increased feeding rate, resulting in faster shell growth and greater shell strength (i.e. more mechanically resilient). When combined with ocean acidification, however, the gastropods had a substantial decrease in energy budget due to reduced feeding rate and energy assimilation, leading to the reduction in shell growth and shell strength. By linking energy budget to the adjustability of shell building, we revealed that energy availability can be critical to determine the physiological adaptability of marine calcifiers to the changing oceanic climate.