Spatio-temporal variability of polychaete colonization at volcanic CO2 vents indicates high tolerance to ocean acidification

Ocean acidification is predicted to have negative effects on marine biota, resulting in the loss of biodiversity and changes in marine ecosystem structure and function. However, some species and life stages may be capable of thriving in low pH conditions, either due to their natural ability to tolerate stressful low pH–high pCO₂ conditions and/or alteration of species interactions caused by changes in pH profiles, or due to evolutionary trade-offs. A better understanding of which species may be capable of tolerating ocean acidification can guide future research into the mechanisms for physiological and ecological resilience to future carbon dioxide (CO₂) conditions. We investigated the colonization of selected polychaete species along a pH gradient originating from shallow, coastal volcanic CO₂ vents (Ischia, Italy). Colonization was quantified by exposing artificial invertebrate collectors attached to the substratum for 30 days during different periods of the year (late spring, fall and late winter). Three species, Amphiglena mediterranea, Platynereis dumerilii and Syllis prolifera, were present and abundant along the gradient throughout the year. All three species were significantly more abundant in the most acidified areas, confirming their high tolerance and capacity to cope with very low pH. Abundances of all three species were compared to data previously collected via collectors suspended in the water column. More individuals were found in the collectors attached to the substratum, suggesting that abundances may have previously been underestimated. This is likely due to the close proximity of these collectors with the natural rocky substratum. All three species exhibited similar temporal variability, consistent with their life cycle and reproductive biology. Our results demonstrate high tolerance of the species for low and variable pH and corroborate their use as robust models to explore the capacity to cope with low pH–high pCO₂ conditions, both in the natural vent systems and in the laboratory.     

The effect of ocean acidification on early algal colonization stages at natural CO2 vents

Marine algae exhibit different responses to ocean acidification, suggesting that a decrease in pH does not always favour marine photosynthetic organisms. In order to understand the effect of acidification on algal community development, early colonization stages were investigated using carbon dioxide vents around the Castello Aragonese (Ischia, Italy) as a natural laboratory. Settlement tiles were placed in zones with different pH (normal, medium and low), and species composition and coverage measured after 2, 3 and 4 months of deployment. The number of species decreased by 4 and 18 % at medium and low pH zones, respectively (P < 0.05). The structure of the algal assemblage differed between pH zones during the 4 months of the experiment, due to the addition and/or replacement of new species. This leads to a change in the succession of morphological forms as soft crustose algae replaced calcareous species, and turf species were dominant in cover; more complex thalli started to occur only at medium pH. These results support previous findings that ocean acidification will induce changes in benthic algal communities.     

Appropriate pCO2 treatments in ocean acidification experiments

Experiments in which organisms are reared in treatments simulating current and future pCO₂ concentrations are critical for ocean acidification (OA) research. The majority of OA exposure experiments use average atmospheric pCO₂ levels as a baseline treatment. We conducted an ecoregion-scale analysis of global carbon chemistry datasets. For many locales, atmospheric pCO₂ levels are not an appropriate characterization of marine carbon chemistry. We argue that atmospheric pCO₂ should be disregarded when setting baseline treatment conditions and experimental design should rely on measurements of carbon chemistry in a study subject’s habitat. As carbon chemistry conditions vary with space and time, we suggest using a range of pCO₂ values as a control rather than a single value. We illustrate this issue with data on the habitat of Euphausia pacifica, which currently lives in waters with a pCO₂ around 900 μatm, a concentration much higher than the current global atmospheric mean.     

Impacts of ocean acidification on marine shelled molluscs

Over the next century, elevated quantities of atmospheric CO₂ are expected to penetrate into the oceans, causing a reduction in pH (?0.3/?0.4 pH unit in the surface ocean) and in the concentration of carbonate ions (so-called ocean acidification). Of growing concern are the impacts that this will have on marine and estuarine organisms and ecosystems. Marine shelled molluscs, which colonized a large latitudinal gradient and can be found from intertidal to deep-sea habitats, are economically and ecologically important species providing essential ecosystem services including habitat structure for benthic organisms, water purification and a food source for other organisms. The effects of ocean acidification on the growth and shell production by juvenile and adult shelled molluscs are variable among species and even within the same species, precluding the drawing of a general picture. This is, however, not the case for pteropods, with all species tested so far, being negatively impacted by ocean acidification. The blood of shelled molluscs may exhibit lower pH with consequences for several physiological processes (e.g. respiration, excretion, etc.) and, in some cases, increased mortality in the long term. While fertilization may remain unaffected by elevated pCO₂, embryonic and larval development will be highly sensitive with important reductions in size and decreased survival of larvae, increases in the number of abnormal larvae and an increase in the developmental time. There are big gaps in the current understanding of the biological consequences of an acidifying ocean on shelled molluscs. For instance, the natural variability of pH and the interactions of changes in the carbonate chemistry with changes in other environmental stressors such as increased temperature and changing salinity, the effects of species interactions, as well as the capacity of the organisms to acclimate and/or adapt to changing environmental conditions are poorly described.     

Effects of ocean acidification on the early developmental stages of the horned turban,Turbo cornutus

To estimate the impact of CO₂-driven ocean acidification on the early life stages of gastropods, the effects of increased partial pressure of seawater carbon dioxide (pCO₂) (800–2,000 μatm) on the early developmental stages and larval shell length of the commercially important gastropod, the horned turban snail, Turbo cornutus were investigated. Increase in experimental seawater pCO₂ had an increasingly negative impact on the early developmental rate; the proportion of embryos or larvae displaying retarded development increased at higher pCO₂. The proportion of embryos that developed to the 4-cell stage at 2 h after fertilization decreased linearly with increasing pCO₂. At ~1,000 μatm pCO₂, retarded development was observed in ~50 % of larvae. No embryos developed to the 4-cell stage at 2,000 μatm pCO₂ within 2 h of fertilization. A similar trend continued until 24–26 h after fertilization; the proportion of larvae attaining veliger stage by 24–26 h also decreased with increasing pCO₂. The shell length of T. cornutus veligers decreased gradually as seawater pCO₂ increased, but markedly decreased in seawater under nearly unsaturated and unsaturated conditions (≤1.04) of the aragonite saturation state (Ω _aragonite). The results indicate that increased pCO₂ seawater has a progressive and acute effect on embryonic and larval T. cornutus, and imply that the extended early developmental period and/or the downsized larval shell produced by ocean acidification will have a negative impact on survival, settlement and recruitment well into the future.     

凋落物处理对森林地表CO2通量的影响及其调控机理

全球变化和森林演替可以导致森林地表凋落物数量和质量的变化,从而对森林地表 CO2通量产生影响。本实验对亚热带不同演替阶段的3种,即马尾松林（前期）、混交林（中期）和季风林（后期）进行地表凋落物去除、加倍与置换处理,利用静态箱气相色谱法测定地表CO2通量,并同步测定气温、土壤温度和湿度,分析凋落物质量和数量变化对森林地表CO2通量的影响及其调控机理。结果表明,（1）去除凋落物处理显著降低了不同演替阶段的3种森林地表CO2通量,而加倍凋落物处理可以增加森林地表CO2通量,但不同演替阶段增加的幅度不同,依次为：季风林＞马尾松林＞混交林。（2）置换凋落物对不同演替阶段的森林地表 CO2通量的影响不同,在演替后期的季风林中,置换混交林和马尾松林凋落物处理均增加地表CO2通量;在演替中期的混交林中,置换季风林和马尾松林凋落物均降低地表CO2通量。在演替前期的马尾松林中,置换季风林凋落物增加地表CO2通量,而置换混交林凋落物降低了地表CO2通量（3）结合测定的土壤温度和水分数据分析得出,凋落物处理引起森林地表 CO2通量的变化是通过处理凋落物质量和数量后改变森林地表水热条件来实现的。（4）3个林型的各种处理,地表 CO2通量与土壤温度均呈显著的指数相关关系,但不同处理不同地改变了森林地表土壤 CO2通量对温度的敏感性,即Q10值。     

Predicted impact of ocean acidification on a marine invertebrate: elevated CO<Subscript>2</Subscript> alters response to thermal stress in sea urchin larvae

Ocean acidification (OA) and the biological consequences of altered seawater chemistry have emerged as a significant environmental threat to healthy marine ecosystems. Because a more acidic ocean interferes with fixation of calcium carbonate to form shells or calcified skeletons, future ocean chemistry may significantly alter the physiology of calcifying marine organisms. These alterations may manifest themselves directly in the calcification process, or have synergistic effects with other environmental factors such as elevated temperatures. New tools permit us to explore subtle changes in gene expression patterns in response to environmental conditions. We raised sea urchins (Strongylocentrotus franciscanus) under conditions simulating future atmospheric CO₂ levels of 540 and 970 ppm. When larvae raised under elevated CO₂ conditions were subjected to 1-h acute temperature stress, their ability to mount a physiological response (as measured by expression of the molecular chaperone hsp70) was reduced relative to those raised under ambient CO₂ conditions. These results represent the first use of gene expression assays to study the effects of OA on sea urchin development. They highlight the importance of looking at multiple environmental factors simultaneously as this approach may reveal previously unsuspected biological impacts of atmospheric changes.     

Effects of feeding and light intensity on the response of the coral Porites rus to ocean acidification

Recently, it has been suggested that there are conditions under which some coral species appear to be resistant to the effects of ocean acidification. To test if such resistance can be explained by environmental factors such as light and food availability, the present study investigated the effect of 3 feeding regimes crossed with 2 light levels on the response of the coral Porites rus to 2 levels of pCO₂ at 28 °C. After 1, 2, and 3 weeks of incubation under experimental conditions, none of the factors—including pCO₂—significantly affected area-normalized calcification and biomass-normalized calcification. Biomass also was unaffected during the first 2 weeks, but after 3 weeks, corals that were fed had more biomass per unit area than starved corals. These results suggest that P. rus is resistant to short-term exposure to high pCO₂, regardless of food availability and light intensity. P. rus might therefore represent a model system for exploring the genetic basis of tolerance to OA.     

Effects of ocean acidification, temperature and nutrient regimes on the appendicularian Oikopleura dioica: a mesocosm study

Increasing pCO₂ is hypothesized to induce shifts in plankton communities toward smaller cells, reduced carbon export rates and increased roles of gelatinous zooplankton. Appendicularians, among the most numerous pan-global “gelatinous” zooplankton, continuously produce filter-feeding houses, shortcutting marine food webs by ingesting submicron particles, and their discarded houses contribute significantly to carbon fluxes. We present a first mesocosm-scale study on the effects of temperature, pCO₂ and bloom structures on the appendicularian, Oikopleura dioica. There were effects of temperature and nutrients on phytoplankton communities. No shifts in functional phytoplankton groups, nor changes in particle sizes/morphotypes, known to impact appendicularian feeding, were observed under manipulated pCO₂ conditions. However, appendicularian abundance was positively correlated with increased pCO₂, temperature and nutrient levels, consistent with hypotheses concerning gelatinous zooplankton in future oceans. This suggests appendicularians will play more important roles in marine pelagic communities and vertical carbon transport under projected ocean acidification and elevated temperature scenarios.     

Impact of ocean acidification on marine ecosystems: educational challenges and innovations

Population growth and social/technological developments have resulted in the buildup of carbon dioxide (CO₂) in the atmosphere and oceans to the extent that we now see changes in the earth’s climate and ocean chemistry. Ocean acidification is one consequence of these changes, and it is known with certainty that it will continue to increase as we emit more CO₂ into the atmosphere. Ocean acidification is a global issue likely to impact marine organisms, food webs and ecosystems and to be most severely experienced by the people who depend on the goods and services the ocean provides at regional and local levels. However, research is in its infancy and the available data on biological impacts are complex (e.g., species-specific response). Educating future generations on the certainties and uncertainties of the emerging science of ocean acidification and its complex consequences for marine species and ecosystems can provide insights that will help assessing the need to mitigate and/or adapt to future global change. This article aims to present different educational approaches, the different material available and highlight the future challenges of ocean acidification education for both educators and marine biologists.     

Biological impacts of ocean acidification: a postgraduate perspective on research priorities

Research into the effects of ocean acidification (OA) on marine organisms has greatly increased during the past decade, as realization of the potential dramatic impacts has grown. Studies have revealed the multifarious responses of organisms to OA conditions, indicating a high level of intra- and interspecific variation in species’ ability to accommodate these alterations. If we are to provide policy makers with sound, scientific input regarding the expected consequences of OA, we need a broader understanding of these predicted changes. As a group of 20 multi-disciplinary postgraduate students from around the globe, with a study focus on OA, we are a strong representation of ‘next generation’ scientists in this field. In this unique cumulative paper, we review knowledge gaps in terms of assessing the biological impacts of OA, outlining directions for future research.     

Benthic invertebrate larval response to substrate characteristics at settlement: Shelter preferences of the American lobsterHomarus americanus

The factors controlling shelter selection by the first post-larval stage of the American lobsterHomarus americanus were studied in the laboratory in 1988 using an experimental approach. The relative importance of shelter size, light level within shelters, number of shelter entrances, and presence of algal-film cover in and around them were investigated. Post-larvae showed a strong preference for opaque versus transparent shelters. No difference in post-larval selectivity was observed either for shelters of different widths or heights or between shelters with different numbers of entrances. Post-larvae showed no preference for microhabitats colonized by microorganisms compared to others without colonization. The intensity of the response measured suggests a possible hierarchical organization of the different cues involved in shelter selection, the light level within the shelter apparently being the most important. Simple linear regression was used to assess the effect of shelter area on the number of post-larvae occupying a shelter. Up to four post-larvae were observed simultaneously in a shelter of 40 cm² basal area, indicating that they can cohabit at settlement. The minimum shelter area necessary to the first post-larval stage was estimated to be about 3.25 cm². The results are discussed with reference to recent hypotheses concerning the mechanisms controlling larval settlement in other benthic species with relatively weak mobility during the larval and the post-larval phases.Contribution to the programmes of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec) and of the Institut Maurice-Lamontagne     

Health and population-dependent effects of ocean acidification on the marine isopod Idotea balthica

Three populations of the grazing isopod Idotea balthica were exposed to high CO₂ treatment for a period of 20 days to investigate the effect of ocean acidification (OA) on animal health and immunocompetence. The results of the populations from more saline habitats were comparable and showed a 60–80 % decrease in immune response as a result of the high CO₂ treatment. Analysis of protein carbonyls showed no treatment effect, indicating that short-term OA does not increase oxidative protein damage. Meanwhile, the third tested population from the lower saline Baltic Sea had higher background protein carbonyl levels. Ocean acidification in addition to this resulted in 100 % mortality. The results of this study show that OA reduced immunocompetence of this marine isopod. In addition, populations and individuals in poor health are potentially at greater risk to succumb under OA.     

Temporal scales and patterns of invertebrate biodiversity dynamics in boreal lakes recovering from acidification

Despite international policy implementation to reduce atmospheric acid deposition and restore natural resources from cultural acidification, evidence of ecological recovery is equivocal. Failure to meet recovery goals means that acidification still threatens biodiversity in many areas of the world. Managers thus need information to manage biodiversity, especially its components that are sensitive to stress (acid-sensitive taxa). We analyzed 20-year time series (1988-2007) of water quality and littoral invertebrates in acidified and circum-neutral lakes across Sweden to evaluate regional biodiversity dynamics and the extent to which changes in water quality affect these dynamics. We used multivariate time series modeling to (1) test how individual species groups within invertebrate communities track changes in the abiotic environment and (2) reveal congruencies of taxon contributions to species group change across lakes. Chemical recovery in the lakes was equivocal, and increases of pH and alkalinity were observed in subsets of acidified and circum-neutral lakes. Time series analyses revealed two different patterns of species groups for invertebrate communities across lakes; the first species group showed monotonic change over time, while the second group showed fluctuating temporal patterns. These independent species groups correlated distinctly with different sets of environmental variables. Recovery of pH and alkalinity status was associated with species group patterns only in a few lakes, highlighting an overall weak recovery of invertebrate species. The sets of species, including acid-sensitive taxa, composing these species groups differed markedly across lakes, highlighting context-specific responses of invertebrates to environmental variation. These results are encouraging because disparate local-scale dynamics maintain the diversity of sensitive invertebrate species on a regional scale, despite persisting acidification problems. Our study can inform and help refine current acidification-related policy focused on sensitive biodiversity elements.     

Effects of feeding history on the growth characteristics of a reef fish at settlement

A feeding experiment was conducted on the pelagic stages of the tropical goatfish Upeneus tragula (family Mullidae) to examine how food quantity affects growth characteristics and potential success at settlement. Pelagic goatfish were collected from aggregation rafts 3 nautical miles west of Lizard Island on the northern Great Barrier Reef during December 1990. Three tanks in each of four feeding regimes were stocked with 25 pelagic goatfish between 20 and 23 mm standard length (SL). The four feeding regimes were: fed ad lib. (fed): fed once per day (f1pd); starved every second day (stld); starved for 3 d and re-fed (st3d). Fishes were fed 36 to 48 h old Artemia sp. nauplii (Ocean Star strain). Fish were removed from tanks when they underwent metamorphosis, changed pigmentation and settled to the bottom of the tanks. Morphology, muscle development, time taken to settle and biochemical condition were examined. Growth attributes of the treated fish were compared to fish which settled within 24 h of capture (field). All attributes examined were significantly influenced by the feeding treatments. Fish within the fed and st3d treatments were significantly larger and heavier than fish in the less well-fed treatments (f1pd, st1d). Similarly, concentrations of total lipid, carbohydrate and protein in the settled fish were significantly higher in the fed and st3d treatments compared with the st1d and f1pd treatments. For all these morphological and biochemical attributes the st1d and f1pd fish did not significantly differ from the field fishes, but did differ from the fed and st3d fishes. Water content was significantly higher in the f1pd, st1d and field fishes compared with the fed and st3d fish. Furthermore, the average time taken to settle followed the pattern: fed (14d)U. tragula are physiologically well suited to exploiting a patchy food source, and that food availability within the pelagic stages can have a major influence on the growth characteristics of this reef fish at settlement. The ramifications of this finding are discussed in relation to survival and success once fish have recruited to the reef population.