ALBERT

Description: Future ocean acidification has the potential to adversely affect many marine organisms. A growing body of evidence suggests that many species could suffer from reduced fertilization success, decreases in larval- and adult growth rates, reduced calcification rates, and even mortality when being exposed to near-future levels (year 2100 scenarios) of ocean acidification. Little research focus is currently placed on those organisms/taxa that might be less vulnerable to the anticipated changes in ocean chemistry; this is unfortunate, as the comparison of more vulnerable to more tolerant physiotypes could provide us with those physiological traits that are crucial for ecological success in a future ocean. Here, we attempt to summarize some ontogenetic and lifestyle traits that lead to an increased tolerance towards high environmental pCO2. In general, marine ectothermic metazoans with an extensive extracellular fluid volume may be less vulnerable to future acidification as their cells are already exposed to much higher pCO2 values (0.1 to 0.4 kPa, ca. 1000 to 3900 μatm) than those of unicellular organisms and gametes, for which the ocean (0.04 kPa, ca. 400 μatm) is the extracellular space. A doubling in environmental pCO2 therefore only represents a 10% change in extracellular pCO2 in some marine teleosts. High extracellular pCO2 values are to some degree related to high metabolic rates, as diffusion gradients need to be high in order to excrete an amount of CO2 that is directly proportional to the amount of O2 consumed. In active metazoans, such as teleost fish, cephalopods and many brachyuran crustaceans, exercise induced increases in metabolic rate require an efficient ion-regulatory machinery for CO2 excretion and acid-base regulation, especially when anaerobic metabolism is involved and metabolic protons leak into the extracellular space. These ion-transport systems, which are located in highly developed gill epithelia, form the basis for efficient compensation of pH disturbances during exposure to elevated environmental pCO2. Compensation of extracellular acid-base status in turn may be important in avoiding metabolic depression. So far, maintained "performance" at higher seawater pCO2 (〉0.3 to 0.6 kPa) has only been observed in adults/juveniles of active, high metabolic species with a powerful ion regulatory apparatus. However, while some of these taxa are adapted to cope with elevated pCO2 during their regular embryonic development, gametes, zygotes and early embryonic stages, which lack specialized ion-regulatory epithelia, may be the true bottleneck for ecological success – even of the more tolerant taxa. Our current understanding of which marine animal taxa will be affected adversely in their physiological and ecological fitness by projected scenarios of anthropogenic ocean acidification is quite incomplete. While a growing amount of empirical evidence from CO2 perturbation experiments suggests that several taxa might react quite sensitively to ocean acidification, others seem to be surprisingly tolerant. However, there is little mechanistic understanding on what physiological traits are responsible for the observed differential sensitivities (see reviews of Seibel and Walsh, 2003; Pörtner et al., 2004; Fabry et al., 2008; Pörtner, 2008). This leads us to the first very basic question of how to define general CO2 tolerance on the species level.

Description: Future scenarios project combined developments of elevated CO2 concentrations and global warming and their impact on marine ecosystems. The synergistic impact of both factors was addressed by studying the effect of CO2 accumulation on thermal tolerance of the cold-eurythermal spider crab Hyas araneus. Animals were exposed to present day normocapnia (380 ppm CO2), CO2 levels expected towards 2100 (710 ppm) and beyond (3000 ppm). Heart rate and haemolymph PO2 (PeO2) were measured during progressive short term cooling from 10 to 0°C and during warming from 10 to 25°C. An increase of PeO2 occurred during cooling with highest values reached at 0°C under all three CO2 levels. Heart rate increased during warming until a critical temperature (Tc) was reached. The putative Tc under normocapnia was presumably 〉25°C, from where it fell to 23.5°C under 710 ppm and then 21.1°C under 3000 ppm. At the same time, thermal sensitivity, as seen in the Q10 values of heart rate, rose with increasing CO2 concentration in the warmth. Our results suggest a narrowing of the thermal window of Hyas araneus under moderate increases in CO2 levels by exacerbation of the heat or cold induced oxygen and capacity limitation of thermal tolerance.

Description: Future ocean acidification has the potential to adversely affect many marine organisms. A growing body of evidence suggests that many species could suffer from reduced fertilization success, decreases in larval- and adult growth rates, reduced calcification rates, metabolic depression and even mortality when being exposed to near-future levels (year 2100 scenarios) of ocean acidification. Little research focus is currently placed on those organisms/taxa that might be less vulnerable to the anticipated changes in ocean chemistry; this is unfortunate, as the comparison of more vulnerable to more tolerant physiotypes could provide us with those physiological traits that are crucial for ecological success in a future ocean. Here, we attempt to summarize some ontogenetic and lifestyle traits that lead to an increased tolerance towards high environmental pCO2. In general, marine ectothermic metazoans with an extensive extracellular fluid volume may be less vulnerable to future acidification as their cells are already exposed to much higher pCO2 values (0.1 to 0.4 kPa, 1000 to 4000 μatm) than those of unicellular organisms and gametes, for which the ocean (0.04 kPa, 400 μatm) is the extracellular space. A doubling in environmental pCO2 therefore only represents a 10% change in extracellular CO2 in some marine teleosts. High extracellular pCO2 values are to some degree related to high metabolic rates, as diffusion gradients need to be high in order to excrete an amount of CO2 that is directly proportional to the amount of O2 consumed. In active metazoans, such as teleost fish, cephalopods and many brachyuran crustaceans, exercise induced increases in metabolic rate require an efficient ion-regulatory machinery for CO2 excretion and acid-base regulation, especially when anaerobic metabolism is involved and metabolic protons leak into the extracellular space. These ion-transport systems, which are located in highly developed gill epithelia, form the basis for efficient compensation of pH disturbances during exposure to elevated environmental pCO2. Compensation of extracellular acid-base status in turn may be extremely important in avoiding metabolic depression. So far, maintained "performance" at higher seawater pCO2 (〉0.3 to 0.6 kPa) has only been observed in adults/juveniles of active, high metabolic species with a powerful ion regulatory apparatus. However, while some of these taxa are adapted to cope with elevated pCO2 during their regular embryonic development, unicellular gametes, which lack specialized ion-regulatory epithelia, may be the true bottleneck for ecological success – even of the more tolerant taxa.

Description: Dissolved oxygen (DO) concentration in the water column is an environmental parameter that is crucial for the successful development of many pelagic organisms. Hypoxia tolerance and threshold values are species- and stage-specific and can vary enormously. While some fish species may suffer from oxygen values of less than 3 ml L−1 and show impact on growth, development and behaviour, other organisms such as euphausiids may survive DO levels as low as 0.1 ml L−1. A change in the average or the minimum or maximum DO in an area may have significant impacts on the survival of certain species and hence on the species composition in the ecosystem with consequent changes in trophic pathways and productivity. Evidence of the deleterious effects of oxygen depletion on species of the pelagic realm is scarce, particularly in terms of the effect of low oxygen on development, recruitment and patterns of migration and distribution. While planktonic organisms have to cope with different DOs and find adaptive mechanisms, nektonic species may avoid areas of inconvenient DO and develop adapted migrational strategies. Planktonic organisms may only be able to escape vertically, above or beneath the Oxygen Minimum Zone (OMZ). In shallow areas only the surface layer can serve as a refuge, in deep waters many organisms have developed vertical migration strategies to use, pass and cope with the OMZ. This paper elucidates the role of DO for different taxa in the pelagic realm and the consequences of low oxygen for foodweb structure and system productivity.

Description: Future scenarios for the oceans project combined developments of CO2 accumulation and global warming and their impact on marine ecosystems. The synergistic impact of both factors was addressed by studying the effect of elevated CO2 concentrations on thermal tolerance of the cold-eurythermal spider crab Hyas araneus from the population around Helgoland. Here ambient temperatures characterize the southernmost distribution limit of this species. Animals were exposed to present day normocapnia (380 ppm CO2), CO2 levels expected towards 2100 (710 ppm) and beyond (3000 ppm). Heart rate and haemolymph PO2 (PeO2) were measured during progressive short term cooling from 10 to 0°C and during warming from 10 to 25°C. An increase of PeO2 occurred during cooling, the highest values being reached at 0°C under all three CO2 levels. Heart rate increased during warming until a critical temperature (Tc) was reached. The putative Tc under normocapnia was presumably 〉25°C, from where it fell to 23.5°C under 710 ppm and then 21.1°C under 3000 ppm. At the same time, thermal sensitivity, as seen in the Q10 values of heart rate, rose with increasing CO2 concentration in the warmth. Our results suggest a narrowing of the thermal window of Hyas araneus under moderate increases in CO2 levels by exacerbation of the heat or cold induced oxygen and capacity limitation of thermal tolerance.

Description: Dissolved oxygen (DO) concentration in the water column is an environmental parameter that is crucial for the successful development of many pelagic organisms. Hypoxia tolerance and threshold values are species- and stage-specific and can vary enormously. While some fish species may suffer from oxygen values of less than 3 mL O2 L−1 through impacted growth, development and behaviour, other organisms such as euphausiids may survive DO levels as low as 0.1 mL O2 L−1. A change in the average or the range of DO may have significant impacts on the survival of certain species and hence on the species composition in the ecosystem with consequent changes in trophic pathways and productivity. Evidence for the deleterious effects of oxygen depletion on pelagic species is scarce, particularly in terms of the effect of low oxygen on development, recruitment and patterns of migration and distribution. While planktonic organisms have to cope with variable DOs and exploit adaptive mechanisms, nektonic species may avoid areas of unfavourable DO and develop adapted migration strategies. Planktonic organisms may only be able to escape vertically, above or beneath the Oxygen Minimum Zone (OMZ). In shallow areas only the surface layer can serve as a refuge, but in deep waters many organisms have developed vertical migration strategies to use, pass through and cope with the OMZ. This paper elucidates the role of DO for different taxa in the pelagic realm and the consequences of low oxygen for foodweb structure and system productivity. We describe processes in two contrasting systems, the semi-enclosed Baltic Sea and the coastal upwelling system of the Benguela Current to demonstrate the consequences of increasing hypoxia on ecosystem functioning and services.