Direct Effects Of Ocean Acidification Research Articles

Are physiological and ecosystem-level tipping points caused by ocean acidification? A critical evaluation

Abstract. Ocean acidification (OA) is predicted to cause profound shifts in many marine ecosystems by impairing the ability of calcareous taxa to calcify and grow and by influencing the physiology of many others. In both calcifying and non-calcifying taxa, ocean acidification could further impair the ability of marine life to regulate internal pH and thus metabolic function and/or behaviour. Identifying tipping points at which these effects will occur for different taxa due to the direct impacts of ocean acidification on organism physiology is difficult because they have not adequately been determined for most taxa nor for ecosystems at higher levels. This is due to the presence of both resistant and sensitive species within most taxa. However, calcifying taxa such as coralline algae, corals, molluscs, and sea urchins appear to be most sensitive to ocean acidification. Conversely, non-calcareous seaweeds, seagrasses, diatoms, cephalopods, and fish tend to be more resistant or even benefit from the direct effects of ocean acidification, though the effects of ocean acidification are more subtle for these taxa. While physiological tipping points of the effects of ocean acidification either do not exist or are not well defined, their direct effects on organism physiology will have flow-on indirect effects. These indirect effects will cause ecological tipping points in the future through changes in competition, herbivory, and predation. Evidence for indirect effects and ecological change is mostly taken from benthic ecosystems in warm temperate–tropical locations in situ that have elevated CO2. Species abundances at these locations indicate a shift away from calcifying taxa and towards non-calcareous taxa at high-CO2 concentrations. For example, lower abundance of corals and coralline algae and higher covers of non-calcareous macroalgae, often turfing species, are often found at elevated CO2. However, there are some locations where only minor changes or no detectable changes occur. Where ecological tipping points do occur, it is usually at locations with naturally elevated mean pCO2 concentrations of 500 µatm or more, which also corresponds to just under that concentration where the direct physiological impacts of ocean acidification are detectable in the most sensitive taxa in laboratory research (coralline algae and corals). Collectively, the available data support the concern that ocean acidification will most likely cause ecological change in the near future in most benthic marine ecosystems, with tipping points in some ecosystems as low as 500 µatm pCO2. However, further research is required to more adequately quantify and model the extent of these impacts in order to accurately project future marine ecosystem tipping points under ocean acidification.

Meta-analysis reveals an extreme "decline effect" in the impacts of ocean acidification on fish behavior.

Ocean acidification-decreasing oceanic pH resulting from the uptake of excess atmospheric CO2-has the potential to affect marine life in the future. Among the possible consequences, a series of studies on coral reef fish suggested that the direct effects of acidification on fish behavior may be extreme and have broad ecological ramifications. Recent studies documenting a lack of effect of experimental ocean acidification on fish behavior, however, call this prediction into question. Indeed, the phenomenon of decreasing effect sizes over time is not uncommon and is typically referred to as the "decline effect." Here, we explore the consistency and robustness of scientific evidence over the past decade regarding direct effects of ocean acidification on fish behavior. Using a systematic review and meta-analysis of 91 studies empirically testing effects of ocean acidification on fish behavior, we provide quantitative evidence that the research to date on this topic is characterized by a decline effect, where large effects in initial studies have all but disappeared in subsequent studies over a decade. The decline effect in this field cannot be explained by 3 likely biological explanations, including increasing proportions of studies examining (1) cold-water species; (2) nonolfactory-associated behaviors; and (3) nonlarval life stages. Furthermore, the vast majority of studies with large effect sizes in this field tend to be characterized by low sample sizes, yet are published in high-impact journals and have a disproportionate influence on the field in terms of citations. We contend that ocean acidification has a negligible direct impact on fish behavior, and we advocate for improved approaches to minimize the potential for a decline effect in future avenues of research.

Ocean acidification increases domoic acid contents during a spring to summer succession of coastal phytoplankton

Enrichment of the oceans with CO2 may be beneficial for some marine phytoplankton, including harmful algae. Numerous laboratory experiments provided valuable insights into the effects of elevated pCO2 on the growth and physiology of harmful algal species, including the production of phycotoxins. Experiments close to natural conditions are the next step to improve predictions, as they consider the complex interplay between biotic and abiotic factors that can confound the direct effects of ocean acidification. We therefore investigated the effect of ocean acidification on the occurrence and abundance of phycotoxins in bulk plankton samples during a long-term mesocosm experiment in the Gullmar Fjord, Sweden, an area frequently experiencing harmful algal blooms. During the experimental period, a total of seven phycotoxin-producing harmful algal genera were identified in the fjord, and in accordance, six toxin classes were detected. However, within the mesocosms, only domoic acid and the corresponding producer Pseudo-nitzschia spp. was observed. Despite high variation within treatments, significantly higher particulate domoic acid contents were measured in the mesocosms with elevated pCO2. Higher particulate domoic acid contents were additionally associated with macronutrient limitation. The risks associated with potentially higher phycotoxin levels in the future ocean warrants attention and should be considered in prospective monitoring strategies for coastal marine waters.

Early life stages of Calanus pacificus are neither exposed nor sensitive to low pH waters

Abstract We characterized the vertical distribution of Calanus pacificus eggs and larvae and the carbonate chemistry that they are exposed to in Puget Sound, WA. We found that, under stratified conditions, more than 90% of eggs and nauplii stages 1–4 were distributed above the pycnocline, in seawater with pH higher than 7.7. In addition, eggs and larvae from 101 females were reared for 5 days under a range of pH conditions (7.2–8.0) to investigate how pH sensitivity varies among individuals. We observed a slight increase in naupliar survival at pH 7.3 in Individual Brood experiments, while in Mixed Brood experiments, exposure to pH 7.3 led to a small decline in hatching success. In a Split Brood experiment, inter-individual variability among different females’ broods masked pH effects. These results indicate that C. pacificus early life stages are generally tolerant to short-term direct effects of ocean acidification.

No maternal or direct effects of ocean acidification on egg hatching in the Arctic copepod Calanus glacialis.

Widespread ocean acidification (OA) is transforming the chemistry of the global ocean and the Arctic is recognised as the region where this transformation will occur at the fastest rate. Moreover, many Arctic species are considered less capable of tolerating OA due to their lower capacity for acid-base regulation. This inability may put severe restraints on many fundamental functions, such as growth and reproductive investments, which ultimately may result in reduced fitness. However, maternal effects may alleviate severe effects on the offspring rendering them more tolerant to OA. In a highly replicated experiment we studied maternal and direct effects of OA predicted for the Arctic shelf seas on egg hatching time and success in the keystone copepod species Calanus glacialis. We incubated females at present day conditions (pHT 8.0) and year 2100 extreme conditions (pHT 7.5) during oogenesis and subsequently reciprocally transplanted laid eggs between these two conditions. Statistical tests showed no effects of maternal or direct exposure to OA at this level. We hypothesise that C. glacialis may be physiologically adapted to egg production at low pH since oogenesis can also take place at conditions of potentially low haemolymph pH of the mother during hibernation in the deep.

The effects of pH and pCO2 on photosynthesis and respiration in the diatom Thalassiosira weissflogii.

The response of marine phytoplankton to the ongoing increase in atmospheric pCO2 reflects the consequences of both increased CO2 concentration and decreased pH in surface seawater. In the model diatom Thalassiosira weissflogii, we explored the effects of varying pCO2 and pH, independently and in concert, on photosynthesis and respiration by incubating samples in water enriched in H218O. In long-term experiments (~6-h) at saturating light intensity, we observed no effects of pH or pCO2 on growth rate, photosynthesis or respiration. This absence of a measurable response reflects the very small change in energy used by the carbon concentrating mechanism (CCM) compared to the energy used in carbon fixation. In short-term experiments (~3min), we also observed no effects of pCO2 or pH, even under limiting light intensity. We surmise that in T. weissflogii, it is the photosynthetic production of NADPH and ATP, rather than the CO2-saturation of Rubisco that controls the rate of photosynthesis at low irradiance. In short-term experiments, we observed a slightly higher respiration rate at low pH at the onset of the dark period, possibly reflecting the energy used for exporting H+ and maintaining pH homeostasis. Based on what is known of the biochemistry of marine phytoplankton, our results are likely generalizable to other diatoms and a number of other eukaryotic species. The direct effects of ocean acidification on growth, photosynthesis and respiration in these organisms should be small over the range of atmospheric pCO2 predicted for the twenty-first century.

Lost at sea: ocean acidification undermines larval fish orientation via altered hearing and marine soundscape modification.

The dispersal of larvae and their settlement to suitable habitat is fundamental to the replenishment of marine populations and the communities in which they live. Sound plays an important role in this process because for larvae of various species, it acts as an orientational cue towards suitable settlement habitat. Because marine sounds are largely of biological origin, they not only carry information about the location of potential habitat, but also information about the quality of habitat. While ocean acidification is known to affect a wide range of marine organisms and processes, its effect on marine soundscapes and its reception by navigating oceanic larvae remains unknown. Here, we show that ocean acidification causes a switch in role of present-day soundscapes from attractor to repellent in the auditory preferences in a temperate larval fish. Using natural CO2 vents as analogues of future ocean conditions, we further reveal that ocean acidification can impact marine soundscapes by profoundly diminishing their biological sound production. An altered soundscape poorer in biological cues indirectly penalizes oceanic larvae at settlement stage because both control and CO2-treated fish larvae showed lack of any response to such future soundscapes. These indirect and direct effects of ocean acidification put at risk the complex processes of larval dispersal and settlement.

Oceanographic patterns and carbonate chemistry in the vicinity of cold-water coral reefs in the Gulf of Mexico: Implications for resilience in a changing ocean

To accurately assess the threat that global climate change poses to marine systems, a detailed baseline of the current carbonate chemistry and other oceanographic conditions is required. Despite the heightened vulnerability of deep-sea communities to ocean acidification, there have been relatively few studies investigating the carbonate chemistry immediately above cold-water coral reefs. Here, we present data collected during five cruises from 2010 to 2014 in the northern Gulf of Mexico and quantify the carbonate system and other oceanographic parameters in offshore surface-waters, the water column, and at deep benthic sites. Benthic sites containing the scleractinian cold-water coral L. pertusa occurred in waters with a relatively wide temperature range (6.8–13.6°C), low potential density (σθ = 26.9 ± 0.3 kg m−3), low dissolved oxygen concentration (111.3 ± 2.0 μmol kg−1), low pHT (7.87 ± 0.04), low ΩARAG (1.31 ± 0.14), and a low availability of carbonate ions (94.4 ± 9.2 μmol kg−1) compared with L. pertusa habitats in other regions. Based on previous modelling and experimental results, these values place L. pertusa at the edge of its viable niche in the deep Gulf of Mexico. However, significantly elevated total alkalinity (+39–44 μmol kg−1) was detected above large L. pertusa mounds, suggesting that carbonate dissolution within the mounds may be partially ameliorating the direct effects of ocean acidification. Together, these results provide an important baseline for assessing future oceanographic changes in the Gulf of Mexico and for predicting the resilience of cold-water coral reefs to global climate and ocean change.

Marine capture fisheries in the Arctic: winners or losers under climate change and ocean acidification?

AbstractClimate change, ocean acidification (OA) and the subsequent changes in marine productivity may affect fisheries and eventually the whole economy in the Arctic. We analysed how changes in climate and ocean pH under scenarios of anthropogenic CO2 emissions are likely to affect the economics of marine fisheries in the Arctic. We applied a Dynamic Bioclimate Envelope Model (DBEM) and outputs from four different Earth System Models (ESMs) to project future changes in the distribution and maximum catch potential of exploited marine fishes and invertebrates. We projected that total fisheries revenue in the Arctic region may increase by 39% (14–59%) by 2050 relative to 2000 under the Special Reports on Emission Scenario (SRES) A2. Simultaneously, total fishing costs, fishers’ incomes, household incomes and economy‐wide impacts in the Arctic are also projected to increase. Climate change with OA is expected to reduce the potential increases in catch and the economic indicators studied herein. Although the projections suggest that Arctic countries are likely to be ‘winners’ under climate change in comparison with tropical developing countries, the effects of OA will lower the expected future benefits in the Arctic. The predicted impacts are likely to be conservative as we consider only the direct effects of OA on fishes and calcifiers, of which there are only a few in the Arctic. Results of this study would be useful for designing effective adaptation strategies to climate change and measures to mitigate the potential negative impacts of OA in the Arctic.

Elevated carbon dioxide alters the plasma composition and behaviour of a shark.

Increased carbon emissions from fossil fuels are increasing the pCO2 of the ocean surface waters in a process called ocean acidification. Elevated water pCO2 can induce physiological and behavioural effects in teleost fishes, although there appear to be large differences in sensitivity between species. There is currently no information available on the possible responses to future ocean acidification in elasmobranch fishes. We exposed small-spotted catsharks (Scyliorhinus canicula) to either control conditions or a year 2100 scenario of 990 μatm pCO2 for four weeks. We did not detect treatment effects on growth, resting metabolic rate, aerobic scope, skin denticle ultrastructure or skin denticle morphology. However, we found that the elevated pCO2 group buffered internal acidosis via [Formula: see text] accumulation with an associated increase in Na(+), indicating that the blood chemistry remained altered despite the long acclimation period. The elevated pCO2 group also exhibited a shift in their nocturnal swimming pattern from a pattern of many starts and stops to more continuous swimming. Although CO2-exposed teleost fishes can display reduced behavioural asymmetry (lateralization), the CO2-exposed sharks showed increased lateralization. These behavioural effects may suggest that elasmobranch neurophysiology is affected by CO2, as in some teleosts, or that the sharks detect CO2 as a constant stressor, which leads to altered behaviour. The potential direct effects of ocean acidification should henceforth be considered when assessing future anthropogenic effects on sharks.

Effects of Elevated Carbon Dioxide (CO2) Concentrations on Early Developmental Stages of the Marine Copepod Calanus finmarchicus Gunnerus (Copepoda: Calanoidae)

Ocean acidification poses an ongoing threat to marine organisms, and early life stages are believed to be particularly sensitive. The boreal calanoid copepod Calanus finmarchicus seasonally dominates the standing stock of zooplankton in the northern North Sea and North Atlantic, and due to its size and abundance is considered an ecological key species linking energy from primary producers to higher trophic levels. To examine whether the early stages of C. finmarchicus are particularly vulnerable to elevated levels of CO2, eggs and nauplii were subjected to different levels of CO2-acidified seawater for 1 wk. The first experiment, with eggs as the starting point, revealed no marked effect on hatching success, but a significant reduction in nauplii survival during incubation at 8800 ppm CO2. In addition, a significant decrease in ontogenetic development rate during incubation at 8800 ppm CO2 was observed in this experiment. In the second experiment, where third-stage nauplii represented the starting point, no significant effects on ontogenetic development and survival following exposure to pCO2 ≥ 7700 ppm were observed. Data suggest that the two first nauplii stages, which are fed endogenously, may be more vulnerable and therefore likely to represent the “bottleneck” for this species in a more acidic ocean. However, the absence of significant effects in the most sensitive stages during exposure to 2800 ppm CO2, a level that is well above worst-case scenario predictions for year 2300 (approximately 2000 ppm CO2), suggests that this species may be generally robust to direct effects of ocean acidification.

High tolerance of microzooplankton to ocean acidification in an Arctic coastal plankton community

Abstract. Impacts of ocean acidification (OA) on marine biota have been observed in a wide range of marine systems. We used a mesocosm approach to study the response of a high Arctic coastal microzooplankton community during the post-bloom period in Kongsfjorden (Svalbard) to direct and indirect effects of high pCO2/low pH. We found almost no direct effects of OA on microzooplankton composition and diversity. Both the relative shares of ciliates and heterotrophic dinoflagellates as well as the taxonomic composition of microzooplankton remained unaffected by changes in pCO2/pH. Although the different pCO2 treatments affected food availability and phytoplankton composition, no indirect effects (e.g. on the total carrying capacity and phenology of microzooplankton) could be observed. Our data point to a high tolerance of this Arctic microzooplankton community to changes in pCO2/pH. Future studies on the impact of OA on plankton communities should include microzooplankton in order to test whether the observed low sensitivity to OA is typical for coastal communities where changes in seawater pH occur frequently.

Using functional genomics to explore the effects of ocean acidification on calcifying marine organisms

As the research community attempts to forecast the effects of ocean acidification on marine ecosystems, a critical element is a clear understanding of the effects of ocean acidification on an individual organism’s physiology. This article explores how the use of genomics-based tools that measure gene expression — DNA microarrays and quantitative PCR — can assist in this effort and reveal aspects of how calcifying marine organisms will respond to ocean acidification. More specifically, what stands to be gained from this approach is an understanding of the direct effects of ocean acidification and whether organisms have sufficient physiological plasticity to adapt to the altered CO2 conditions. We provide a brief overview of biomineralization processes in corals and sea urchin larvae, and then link these pathways to ways in which gene expression analysis can reveal physiological responses and mechanisms, and further, can define new testable hypotheses. In addition, we review the resources available and strategies that might be taken for each of 2 study organisms, stony corals and sea urchins. Finally, we suggest strategies for gene expression profiling in organisms that differ in availability of genomic resources.