John I. Spicer

Ocean acidification may increase calcification rates, but at a cost

Ocean acidification is the lowering of pH in the oceans as a result of increasing uptake of atmospheric carbon dioxide. Carbon dioxide is entering the oceans at a greater rate than ever before, reducing the oceans natural buffering capacity and lowering pH. Previous work on the biological consequences of ocean acidification has suggested that calcification and metabolic processes are compromised in acidified seawater. By contrast, here we show, using the ophiuroid brittlestar Amphiura filiformis as a model calcifying organism, that some organisms can increase the rates of many of their biological processes (in this case, metabolism and the ability to calcify to compensate for increased seawater acidity). However, this upregulation of metabolism and calcification, potentially ameliorating some of the effects of increased acidity comes at a substantial cost (muscle wastage) and is therefore unlikely to be sustainable in the long term.

Macrophysiology: A Conceptual Reunification

Widespread recognition of the importance of biological studies at large spatial and temporal scales, particularly in the face of many of the most pressing issues facing humanity, has fueled the argument that there is a need to reinvigorate such studies in physiological ecology through the establishment of a macrophysiology. Following a period when the fields of ecology and physiological ecology had been regarded as largely synonymous, studies of this kind were relatively commonplace in the first half of the twentieth century. However, such large‐scale work subsequently became rather scarce as physiological studies concentrated on the biochemical and molecular mechanisms underlying the capacities and tolerances of species. In some sense, macrophysiology is thus an attempt at a conceptual reunification. In this article, we provide a conceptual framework for the continued development of macrophysiology. We subdivide this framework into three major components: the establishment of macrophysiological patterns, determining the form of those patterns (the very general ways in which they are shaped), and understanding the mechanisms that give rise to them. We suggest ways in which each of these components could be developed usefully.

Ocean acidification disrupts induced defences in the intertidal gastropod Littorina littorea

Carbon dioxide-induced ocean acidification is predicted to have major implications for marine life, but the research focus to date has been on direct effects. We demonstrate that acidified seawater can have indirect biological effects by disrupting the capability of organisms to express induced defences, hence, increasing their vulnerability to predation. The intertidal gastropod Littorina littorea produced thicker shells in the presence of predation (crab) cues but this response was disrupted at low seawater pH. This response was accompanied by a marked depression in metabolic rate (hypometabolism) under the joint stress of high predation risk and reduced pH. However, snails in this treatment apparently compensated for a lack of morphological defence, by increasing their avoidance behaviour, which, in turn, could affect their interactions with other organisms. Together, these findings suggest that biological effects from ocean acidification may be complex and extend beyond simple direct effects.

Oxygen supply in aquatic ectotherms: Partial pressure and solubility together explain biodiversity and size patterns

Aquatic ectotherms face the continuous challenge of capturing sufficient oxygen from their environment as the diffusion rate of oxygen in water is 3 x 10(5) times lower than in air. Despite the recognized importance of oxygen in shaping aquatic communities, consensus on what drives environmental oxygen availability is lacking. Physiologists emphasize oxygen partial pressure, while ecologists emphasize oxygen solubility, traditionally expressing oxygen in terms of concentrations. To resolve the question of whether partial pressure or solubility limits oxygen supply in nature, we return to first principles and derive an index of oxygen supply from Ficks classic first law of diffusion. This oxygen supply index (OSI) incorporates both partial pressure and solubility. Our OSI successfully explains published patterns in body size and species across environmental clines linked to differences in oxygen partial pressure (altitude, organic pollution) or oxygen solubility (temperature and salinity). Moreover, the OSI was more accurately and consistently related to these ecological patterns than other measures of oxygen (oxygen saturation, dissolved oxygen concentration, biochemical oxygen demand concentrations) and similarly outperformed temperature and altitude, which covaried with these environmental clines. Intriguingly, by incorporating gas diffusion rates, it becomes clear that actually more oxygen is available to an organism in warmer habitats where lower oxygen concentrations would suggest the reverse. Under our model, the observed reductions in aerobic performance in warmer habitats do not arise from lower oxygen concentrations, but instead through organismal oxygen demand exceeding supply. This reappraisal of how organismal thermal physiology and oxygen demands together shape aerobic performance in aquatic ectotherms and the new insight of how these components change with temperature have broad implications for predicting the responses of aquatic communities to ongoing global climate shifts.

Adaptation and acclimatization to ocean acidification in marine ectotherms: an in situ transplant experiment with polychaetes at a shallow CO2 vent system

Metabolic rate determines the physiological and life-history performances of ectotherms. Thus, the extent to which such rates are sensitive and plastic to environmental perturbation is central to an organisms ability to function in a changing environment. Little is known of long-term metabolic plasticity and potential for metabolic adaptation in marine ectotherms exposed to elevated pCO2. Consequently, we carried out a series of in situ transplant experiments using a number of tolerant and sensitive polychaete species living around a natural CO2 vent system. Here, we show that a marine metazoan (i.e. Platynereis dumerilii) was able to adapt to chronic and elevated levels of pCO2. The vent population of P. dumerilii was physiologically and genetically different from nearby populations that experience low pCO2, as well as smaller in body size. By contrast, different populations of Amphiglena mediterranea showed marked physiological plasticity indicating that adaptation or acclimatization are both viable strategies for the successful colonization of elevated pCO2 environments. In addition, sensitive species showed either a reduced or increased metabolism when exposed acutely to elevated pCO2. Our findings may help explain, from a metabolic perspective, the occurrence of past mass extinction, as well as shed light on alternative pathways of resilience in species facing ongoing ocean acidification.

Immunological function in marine invertebrates: Responses to environmental perturbation

The inception of ecological immunology has led to an increase in the number of studies investigating the impact of environmental stressors on host immune defence mechanisms. This in turn has led to an increased understanding of the importance of invertebrate groups for immunological research. This review discusses the advances made within marine invertebrate ecological immunology over the past decade. By demonstrating the environmental stressors tested, the immune parameters typically investigated, and the species that have received the greatest level of investigation, this review provides a critical assessment of the field of marine invertebrate ecological immunology. In highlighting the methodologies employed within this field, our current inability to understand the true ecological significance of any immune dysfunction caused by environmental stressors is outlined. Additionally, a number of examples are provided in which studies successfully demonstrate a measure of immunocompetence through alterations in disease resistance and organism survival to a realized pathogenic threat. Consequently, this review highlights the potential to advance our current understanding of the ecological and evolutionary significance of environmental stressor related immune dysfunction. Furthermore, the potential for the advancement of our understanding of the immune system of marine invertebrates, through the incorporation of newly emerging and novel molecular techniques, is emphasized.

Student perceptions of a virtual field trip to replace a real field trip

This study examines student perceptions on the use of virtual field trips (VFT) as part of their university experience and in particular the extent to which they could replace real field trips. While students were extremely positive about the potential of VFT to provide valuable learning experiences (and in particular a VFT constructed by the authors of this paper) nearly all of the students were insistent that it could not, and should not, replace real field trips. Furthermore when the same students were re-approached after having been on a real field trip, these perceptions were strengthened and they thought VFT could be most effective in preparing for, or revising after, a real field trip.

Metabolic responses of the prawns Palaemon elegans and P. serratus (Crustacea: Decapoda) to acute hypoxia and anoxia

Using prawns, Palaemon elegans (Rathke) from intertidal pools on the Isle of Cumbrae, Scotland, and P. serratus (Pennant) from the subtidal at Plymouth, England, some metabolic responses to hypoxia and anoxia have been studied. P. elegans was found to have a greater tolerance of severe hypoxia than P. serratus. Tolerance of totally anoxic conditions, however, was limited to only 4 h in P. elegans and to approximately 1 h in P. serratus. exposure to moderate hypoxia (30 torr) resulted in little change in the concentration of L-lactate in the blood or in the tissues of either P. elegans or P. serratus. When exposed to extreme hypoxia (10 or 5 torr for P. elegans), however, there was a progressive increase in the concentration of L-lactate in the blood and in the tissues of both species. After normoxic conditions had been restored, the concentration of L-lactate in the blood and in the tissues returned to normal resting levels more rapidly in P. elegans than in P. serratus. Under hypoxic conditions, both P. elegans and P. serratus showed an increase in the concentration of blood glucose and a slight reduction in the glycogen content of the tissues. The concentrations of blood glucose and of tissue glycogen returned to normal levels within 6 h of the prawns being returned to normoxic conditions. The results of an in situ study in April and August 1986 to examine the metabolic responses of P. elegans to the hypoxic conditions normally experienced in high-shore rock pools are also presented. The ecological significance of the differing abilities of these species to survive hypoxic exposure is discussed.

The Physiological Ecology of Land Invasion by the Talitridae (Crustacea: Amphipoda)

Bousfield’s phylogenetic–systematic scheme (Bernice P. Bishop Mus.spec. Publ. no. 72 (1984)) for the family Talitridae has given renewed impetus to comparative physiological studies on representatives of his different morphological groupings within this, the only amphipod family with truly terrestrial constituents. Our comparative review of talitrid physiology presented here reveals the ecological adaptation of extant species. This has been set against Bousfield’s view of the evolutionary history of the group in an attempt to produce a comprehensive and realistic organismic biology. The beachflea and sandhopper genera are highly modified for life in the supralittoral zone. The former group has given rise to euterrestrial amphipods which are to a certain extent physiologically pre-adapted for more rigorous terrestrial environments. Their success, however, compared with a more ancient landhopper group that invaded land directly (via the leaf litter of newly established angiosperm rainforests in the Cretaceous) may have been limited not solely by desiccation stress but also by more severe osmo- or iono-regulatory constraints.

Comparing the impact of high CO2 on calcium carbonate structures in different marine organisms

Abstract Coastal seas are critical components of the global carbon cycle, yet little research has been conducted on the impact of ocean acidification on coastal benthic organisms. Calcifying marine organisms are predicted to be most vulnerable to a decline in oceanic pH (ocean acidification) based on the assumption that calcification will decrease as a result of changes in seawater carbonate chemistry, particularly reduced carbonate ion concentration (and associated saturation states). Net calcium carbonate production is dependent on an organisms ability to increase calcification sufficiently to counteract an increase in dissolution. Here, a critical appraisal of calcification in five benthic species showed, contrary to popular predictions, the deposition of calcium carbonate can be maintained or even increased in acidified seawater. This study measured changes in the concentration of calcium ions seen in shells taken from living animals exposed to acidified seawater. These data were compared with data from isolated shells that were not associated with living material to determine a species’ ability to maintain the physiological process of calcification under high carbon dioxide (CO2) conditions and characterize the importance of dissolution and abiotic influences associated with decreasing pH. Comparison with palaeoecological studies of past high CO2 events presents a similar picture. This conclusion implies that calcification may not be the physiological process that suffers most from ocean acidification; particularly as all species investigated displayed physiological trade-offs including increased metabolism, reduced health, and changes in behavioural responses in association with this calcification upregulation, which poses as great a threat to survival as an inability to calcify.