Ocean acidification and temperature rise: effects on calcification during early development of the cuttlefish Sepia officinalis

This study investigated the effects of seawater pH (i.e., 8.10, 7.85 and 7.60) and temperature (16 and 19 °C) on (a) the abiotic conditions in the fluid surrounding the embryo (viz. the perivitelline fluid), (b) growth, development and (c) cuttlebone calcification of embryonic and juvenile stages of the cephalopod Sepia officinalis. Egg swelling increased in response to acidification or warming, leading to an increase in egg surface while the interactive effects suggested a limited plasticity of the swelling modulation. Embryos experienced elevated pCO₂ conditions in the perivitelline fluid (>3-fold higher pCO₂ than that of ambient seawater), rendering the medium under-saturated even under ambient conditions. The growth of both embryos and juveniles was unaffected by pH, whereas ⁴⁵Ca incorporation in cuttlebone increased significantly with decreasing pH at both temperatures. This phenomenon of hypercalcification is limited to only a number of animals but does not guarantee functional performance and calls for better mechanistic understanding of calcification processes.     

Cuttlebone calcification increases during exposure to elevated seawater pCO2 in the cephalopod Sepia officinalis

Changes in seawater carbonate chemistry that accompany ongoing ocean acidification have been found to affect calcification processes in many marine invertebrates. In contrast to the response of most invertebrates, calcification rates increase in the cephalopod Sepia officinalis during long-term exposure to elevated seawater pCO₂. The present trial investigated structural changes in the cuttlebones of S. officinalis calcified during 6 weeks of exposure to 615 Pa CO₂. Cuttlebone mass increased sevenfold over the course of the growth trail, reaching a mean value of 0.71 ± 0.15 g. Depending on cuttlefish size (mantle lengths 44–56 mm), cuttlebones of CO₂-incubated individuals accreted 22–55% more CaCO₃ compared to controls at 64 Pa CO₂. However, the height of the CO₂-exposed cuttlebones was reduced. A decrease in spacing of the cuttlebone lamellae, from 384 ± 26 to 195 ± 38 μm, accounted for the height reduction The greater CaCO₃ content of the CO₂-incubated cuttlebones can be attributed to an increase in thickness of the lamellar and pillar walls. Particularly, pillar thickness increased from 2.6 ± 0.6 to 4.9 ± 2.2 μm. Interestingly, the incorporation of non-acid-soluble organic matrix (chitin) in the cuttlebones of CO₂-exposed individuals was reduced by 30% on average. The apparent robustness of calcification processes in S. officinalis, and other powerful ion regulators such as decapod cructaceans, during exposure to elevated pCO₂ is predicated to be closely connected to the increased extracellular [HCO₃ ⁻] maintained by these organisms to compensate extracellular pH. The potential negative impact of increased calcification in the cuttlebone of S. officinalis is discussed with regard to its function as a lightweight and highly porous buoyancy regulation device. Further studies working with lower seawater pCO₂ values are necessary to evaluate if the observed phenomenon is of ecological relevance.     

Moderate seawater acidification does not elicit long-term metabolic depression in the blue mussel Mytilus edulis

Marine organisms are exposed to increasingly acidic oceans, as a result of equilibration of surface ocean water with rising atmospheric CO₂ concentrations. In this study, we examined the physiological response of Mytilus edulis from the Baltic Sea, grown for 2 months at 4 seawater pCO₂ levels (39, 113, 243 and 405 Pa/385, 1,120, 2,400 and 4,000 μatm). Shell and somatic growth, calcification, oxygen consumption and \textNH₄ ⁺ {\text{NH}}_{4}^{ + } excretion rates were measured in order to test the hypothesis whether exposure to elevated seawater pCO₂ is causally related to metabolic depression. During the experimental period, mussel shell mass and shell-free dry mass (SFDM) increased at least by a factor of two and three, respectively. However, shell length and shell mass growth decreased linearly with increasing pCO₂ by 6–20 and 10–34%, while SFDM growth was not significantly affected by hypercapnia. We observed a parabolic change in routine metabolic rates with increasing pCO₂ and the highest rates (+60%) at 243 Pa. \textNH₄ ⁺ {\text{NH}}_{4}^{ + } excretion rose linearly with increasing pCO₂. Decreased O:N ratios at the highest seawater pCO₂ indicate enhanced protein metabolism which may contribute to intracellular pH regulation. We suggest that reduced shell growth under severe acidification is not caused by (global) metabolic depression but is potentially due to synergistic effects of increased cellular energy demand and nitrogen loss.     

Blood oxygen-binding characteristics of bigeye tuna (Thunnus obesus), a high-energy-demand teleost that is tolerant of low ambient oxygen

 We found blood from bigeye tuna (Thunnus obesus) to have a significantly higher O₂ affinity than blood from other tunas. Its P₅₀ (partial pressure of oxygen, P_O2 required to reach 50% saturation) was 1.6 to 2.0 kPa (12 to 15 mmHg) when equilibrated with 0.5% CO₂. Previous studies employing similar methodologies found blood from yellowfin tuna (T. albacares), skipjack tuna (Katsuwonus pelamis), and kawakawa (Euthynnus affinis) to have a P₅₀ of 2.8 to 3.1 kPa (21 to 23 mmHg). These observations suggest that bigeye tuna are more tolerant of low ambient oxygen than other tuna species, and support similar conclusions derived from laboratory whole-animal studies, depth-of-capture data, and directly-recorded vertical movements of fish in the open ocean. We also found the O₂ affinity of bigeye tuna blood to be essentially unaffected by a 10 C° open-system temperature change (as is the blood of all tuna species studied to date). The O₂ affinity of bigeye tuna blood was, however, more affected by a 10 C° closed-system temperature change than the blood of any tuna species yet examined. In other words, bigeye tuna blood displayed a significantly enhanced Bohr effect (change in log P₅₀ per unit change in plasma pH at P₅₀) when subjected to the inevitable changes in partial pressure of carbon dioxide (P_CO2) and plasma pH that accompany closed-system temperature shifts, than when subjected to changes in plasma pH accomplished by changing P_CO2 alone. In vivo, the resultant large decrease in O₂ affinity (i.e. the increase in P₅₀) that occurs as the blood of bigeye tuna is warmed during its passage through the vascular counter-current heat exchangers ensures adequate rates of O₂ off-loading in the swimming muscles of this high-energy-demand teleost. Received: 12 March 1999 / Accepted: 18 December 1999     

The hypoxia avoidance behaviour of juvenile Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) depends on the provision and pressure level of an O<Subscript>2</Subscript> refuge

The frequency of low O₂ (hypoxia) has increased in coastal marine areas but how fish avoid deleterious water masses is not yet clear. To assess whether the presence and oxygen pressure (PO₂) level of an O₂ refuge affects the hypoxia avoidance behaviour of fish, individual Atlantic cod (Gadus morhua L.) were exposed to a range of O₂ choices in a 2-way choice chamber at 11.4°C over two different experiments. Cod in the first experiment were allowed access to a fixed O₂ refuge (fully air-saturated seawater) whilst oxygen pressure (PO₂) on the other side was reduced in steps to a critically low level, i.e. 4.3 kPa—a point where cod can no longer regulate O₂ consumption. Under these conditions, cod did not avoid any level of hypoxia and fish swimming speed also remained unchanged. In contrast, strong avoidance reactions were exhibited in a second experiment when fish were again exposed to 4.3 kPa but the safety, i.e. PO₂, of the refuge was reduced. Fish not only spent less time at 4.3 kPa as a result of fewer sampling visits but they also swam at considerably slower speeds. The presence of an avoidance response was thus strongly related to refuge PO₂ and it is unlikely that cod, and possibly other fish species, would enter low O₂ to feed in the wild if a sufficiently safe O₂ refuge was not available. It is therefore hypothesized that the feeding range of fish may be heavily compressed if hypoxia expands and intensifies in future years.     

Effects of ocean acidification on invertebrate settlement at volcanic CO2 vents

We present the first study of the effects of ocean acidification on settlement of benthic invertebrates and microfauna. Artificial collectors were placed for 1 month along pH gradients at CO₂ vents off Ischia (Tyrrhenian Sea, Italy). Seventy-nine taxa were identified from six main taxonomic groups (foraminiferans, nematodes, polychaetes, molluscs, crustaceans and chaetognaths). Calcareous foraminiferans, serpulid polychaetes, gastropods and bivalves showed highly significant reductions in recruitment to the collectors as pCO₂ rose from normal (336–341 ppm, pH 8.09–8.15) to high levels (886–5,148 ppm) causing acidified conditions near the vents (pH 7.08–7.79). Only the syllid polychaete Syllis prolifera had higher abundances at the most acidified station, although a wide range of polychaetes and small crustaceans was able to settle and survive under these conditions. A few taxa (Amphiglena mediterranea, Leptochelia dubia, Caprella acanthifera) were particularly abundant at stations acidified by intermediate amounts of CO₂ (pH 7.41–7.99). These results show that increased levels of CO₂ can profoundly affect the settlement of a wide range of benthic organisms.     

Removal of olive mill waste water phenolics using a crude peroxidase extract from onion by-products

Olive mill waste water (OMWW) originating from a two-phase olive oil-producing plant was treated with a crude peroxidase extract prepared from onion solid by-products. The treatments were based on a 3 × 3 factorial design, employing a series of combinations of pH and H₂O₂, in order to identify optimal operational conditions. The treatment performance was assessed by estimating the removal percentage of total polyphenols. The model established produced a satisfactory fitting of the data (R ² = 0.94, p = 0.0158), while the second-order polynomial equation used to describe the process indicated that peroxidase-catalysed polyphenol removal in diluted OMWW is facilitated at relatively low pH and intermediate H₂O₂ values. A predicted value of 50.7 ± 9.5% removal was calculated under optimal operational conditions (pH 2.76, [H₂O₂] = 3.56 mM). Analysis of an untreated and an optimally treated sample by high performance liquid chromatography revealed that enzyme treatment brought about alteration in the original OMWW polyphenolic profile. The use of peroxidase from onion solid by-products is proposed as an alternative means that could have a prospect in bioremediation applications.     

Future ocean acidification will be amplified by hypoxia in coastal habitats

Ocean acidification is elicited by anthropogenic carbon dioxide emissions and resulting oceanic uptake of excess CO₂ and might constitute an abiotic stressor powerful enough to alter marine ecosystem structures. For surface waters in gas-exchange equilibrium with the atmosphere, models suggest increases in CO₂ partial pressure (pCO₂) from current values of ca. 390 μatm to ca. 700–1,000 μatm by the end of the century. However, in typically unequilibrated coastal hypoxic regions, much higher pCO₂ values can be expected, as heterotrophic degradation of organic material is necessarily related to the production of CO₂ (i.e., dissolved inorganic carbon). Here, we provide data and estimates that, even under current conditions, maximum pCO₂ values of 1,700–3,200 μatm can easily be reached when all oxygen is consumed at salinities between 35 and 20, respectively. Due to the nonlinear nature of the carbonate system, the approximate doubling of seawater pCO₂ in surface waters due to ocean acidification will most strongly affect coastal hypoxic zones as pCO₂ during hypoxia will increase proportionally: we calculate maximum pCO₂ values of ca. 4,500 μatm at a salinity of 20 (T = 10 °C) and ca. 3,400 μatm at a salinity of 35 (T = 10 °C) when all oxygen is consumed. Upwelling processes can bring these CO₂-enriched waters in contact with shallow water ecosystems and may then affect species performance there as well. We conclude that (1) combined stressor experiments (pCO₂ and pO₂) are largely missing at the moment and that (2) coastal ocean acidification experimental designs need to be closely adjusted to carbonate system variability within the specific habitat. In general, the worldwide spread of coastal hypoxic zones also simultaneously is a spread of CO₂-enriched zones. The magnitude of expected changes in pCO₂ in these regions indicates that coastal systems may be more endangered by future global climate change than previously thought.     

Photosynthetic performance of giant clams, Tridacna maxima and T. squamosa, Red Sea

Two species of giant clams, Tridacna maxima and T. squamosa, coexist in the Red Sea, but exhibit distinctly different depth distributions: T. maxima mostly occurs in shallow waters (reef flat and edge), while T. squamosa may occur down to the lower fore-reef slope. Giant clams have been described as mixotrophic, capable of both filter-feeding and photosynthesis due to algal symbionts (zooxanthellae), therefore, observed depth preferences were investigated in relation to possible differences in autotrophy vs. heterotrophy. This study was conducted from April to June 2004, at the reef near the Marine Science Station, Aqaba, Gulf of Aqaba, Red Sea, and in May 2007, at a reef near Dahab, Sinai Peninsula, Egypt. In situ measurements using a submersible pulse amplitude modulated fluorometer (Diving PAM), revealed no significant differences in effective PSII quantum yield (ΔF/Fm′) and relative electron transport rates (ETR) between the two species; but rapid light curves (ETR vs. light, photosynthetically active irradiance, PAR) showed significant differences in maximum photosynthetic rates (ETR_max), with 20% higher values in T. maxima. Chamber incubations displayed higher net and gross oxygen production by T. maxima (88.0 and 120.3 μmol O₂ cm⁻² mantle area day⁻¹) than T. squamosa (56.7 and 84.8 μmol O₂ cm⁻² mantle area day⁻¹); even under shading conditions (simulated depth of 20 m) T. maxima still achieved 93% of the surface gross O₂ production, whereas T. squamosa reached only 44%. A correlation was found between ETR and net photosynthesis measured as oxygen production (T. maxima: R ² = 0.53; T. squamosa: R ² = 0.61). Calculated compensation depth (CD) (gross photosynthesis equals respiration) in T. maxima (16 m) matches the maximum depth of occurrence in this study (17 m). By contrast, the CD of T. squamosa (9 m) was much shallower than the maximum vertical range (42 m). Findings suggest T. maxima is a strict functional photoautotroph limited by light, whereas T. squamosa is a mixotroph whose photoautotrophic range is extended by heterotrophy. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Carbon-dioxide production and metabolic parameters in the ophiurid Ophiothrix fragilis

As part of the evaluation of fluxes between the water column and a rich benthic community of the Dover Strait (Eastern English Channel), laboratory measurements of oxygen consumption were carried out on a common ophiurid, Ophiothrix fragilis (Abildgaard), from February 1993 to February 1995. The mean O₂-consumption rate was evaluated at 0.31 mg O₂ g⁻¹ h⁻¹ (ash-free dry weight). Simultaneous measurements of O₂ consumption and CO₂ production using the pH-alkalinity method revealed an average respiratory quotient of 0.69 proved suitable for converting oxygen demand to carbon flux. A seasonal trend in respiration data was demonstrated by sinusoidal curves fitted to O₂-uptake and CO₂-release data as a function of time. The influence on respiration rate of two seasonal parameters (temperature and food availability) is discussed; linear regression indicated a highly significant relationship between O₂ consumption (or CO₂ production) and temperature; both O₂-consumption and CO₂-production rates decreased with starvation. The average O:N ratio was estimated at 8.46, close to the theoretical value when proteins constitute the catabolic substrate. The annual carbon respired by the O. fragilis community examined and the estimated annual primary production by phytoplankton indicate that the respiration of the O. fragilis community could supply 35% of phytoplankton carbon requirements. Received: 1 August 1996 / Accepted: 4 September 1996     

Microsensor studies of photosynthesis and respiration in larger symbiotic foraminifera. I The physico-chemical microenvironment of Marginopora vertebralis, Amphistegina lobifera and Amphisorus hemprichii

 The physico-chemical microenvironment of larger benthic foraminifera was studied with microsensors for O₂, CO₂, pH, Ca²⁺ and scalar irradiance. Under saturating light conditions, the photosynthetic activity of the endosymbiotic algae increased the O₂ up to 183% air saturation and a pH of up to 8.6 was measured at the foraminiferal shell surface. The photosynthetic CO₂ fixation decreased the CO₂ at the shell down to 4.7 μM. In the dark, the respiration of host and symbionts decreased the O₂ level to 91% air saturation and the CO₂ concentration reached up to 12 μM. pH was lowered relative to the ambient seawater pH of 8.2. The endosymbionts responded immediately to changing light conditions, resulting in dynamic changes of O₂, CO₂ and pH at the foraminiferal shell surface during experimentally imposed light–dark cycles. The dynamic concentration changes demonstrated for the first time a fast exchange of metabolic gases through the perforate, hyaline shell of Amphistegina lobifera. A diffusive boundary layer (DBL) limited the solute exchange between the foraminifera and the surrounding water. The DBL reached a thickness of 400–700 μm in stagnant water and was reduced to 100–300 μm under flow conditions. Gross photosynthesis rates were significantly higher under flow conditions (4.7 nmol O₂ cm⁻³ s⁻¹) than in stagnant water (1.6 nmol O₂ cm ⁻³ s⁻¹), whereas net photosynthesis rates were unaffected by flow conditions. The Ca²⁺ microprofiles demonstrated a spatial variation in sites of calcium uptake over the foraminiferal shells. Ca²⁺ gradients at the shell surface showed total Ca²⁺ uptake rates of 0.6 to 4.2 nmol cm⁻² h⁻¹ in A. lobifera and 1.7 to 3.6 nmol cm⁻² h⁻¹ in Marginopora vertebralis. The scattering and reflection of the foraminiferal calcite shell increased the scalar irradiance at the surface up to 205% of the incident irradiance. Transmittance measurements across the calcite shell suggest that the symbionts are shielded from higher light levels, receiving approximately 30% of the incident light for photosynthesis. Received: 6 July 1999 / Accepted: 28 April 2000     

The burrows and physiological adaptations to a burrowing lifestyle of Natatolana borealis (Isopoda: Cirolanidae)

The isopod Natatolana borealis Lilljeborg constructs U-shaped burrows in soft mud, the bore of which closely approximates the width of the occupant. Within artificial burrows, the isopods are largely quiescent and often adopt a position close to one of the burrow openings. Conditions within burrows constructed in the laboratory are moderately hypoxic [11.7 to 14.9 kPa (88 to 112 torr)], with isopods showing discontinuous irrigation behaviour (pleopod beating). Rates of oxygen consumption (measured at 10°C) are maintained approximately constant over a wide range of oxygen partial pressure (P_{O 2}) due, in part, to a pronounced increase in pleopod beat rate. Values for the critical partial pressure of oxygen (P_c), the P_{O 2} at which can no longer be maintained independent of P_{O 2}, were 2.0 to 3.3 kPa (15 to 25 torr). N. borealis can survive lengthy periods (65 h at 5°C) of anoxia, during which there is a significant reduction in the carbohydrate concentration and an increase in the l-lactate concentration of the tissues. The oxygencarrying capacity of the haemolymph of N. borealis was low. The haemocyanin showed a relatively high oxygen affinity [P₅₀=0.39 kPa (2.99 torr) at 10°C at the in vivo pH of 7.80] and a pronounced Bohr effect (-1.22). These characteristics may be advantageous to a burrowing mode of life and also for the conditions likely to be encountered in fish carcasses into which they burrow en masse to feed.     

Acid-base responses to lethal aquatic hypercapnia in three marine fishes

Ocean sequestration of CO₂ is proposed as a possible measure to mitigate environmental changes due to the increasing atmospheric concentration of the gas. However, toxic effects of CO₂ on marine organisms are poorly understood. We therefore studied acid–base responses and mortality during exposure to fatal levels of CO₂ in three marine fishes (Japanese flounder, Paralichthys olivaceus; yellowtail, Seriola quinqueradiata; and starspotted dogfish, Mustelus manazo). The teleosts died during exposure to seawater equilibrated with a gas mixture containing 5% CO₂ (water PCO₂ 4.95 kPa); 100% mortality occurred within 8 h for yellowtail and within 48 h for flounder. Only 20% mortality was recorded at 72 h for the dogfish during exposure to 7% CO₂ (water PCO₂ 6.96 kPa). Arterial pH (pH_a) initially decreased, but completely recovered within 1–24 h for the teleosts at 1% and 3%, although the recovery was slower and complete only at 1% (water PCO₂ 0.99 kPa) for the dogfish. During exposure to 5%, the flounder died after the pH_a had been completely restored, suggesting that the mortality was not due to plasma acidosis. During exposure to 1% hypercapnia, plasma [Cl^–] appeared to be the main counter ion to balance increases in plasma [HCO₃^-]. There was a 1:1 stoichiometry for the rise in [HCO₃^-] and the fall in [Cl^–] for the teleosts, whereas the ratio was 1:0.4 for the dogfish at 1% CO₂. At the higher levels of hypercapnia, the rise in [HCO₃^-] consistently exceeded the fall in [Cl^–], and plasma [Na⁺] significantly increased.These results do not agree with the generally accepted model for acid–base regulation in marine fish in which Na⁺/H⁺ exchangers are assumed to play a predominant role, and indicate that an acid–base regulatory mechanism differs between teleost and elasmobranch fishes, as well as the intensity of acidic stress.Communicated by T. Ikeda, Hakodate     

Comparing the effect of elevated pCO2 and temperature on the fertilization and early development of two species of oysters

This study compared the synergistic effects of elevated pCO₂ and temperature on the early life history stages of two ecologically and economically important oysters: the Sydney rock oyster, Saccostrea glomerata and the Pacific oyster, Crassostrea gigas. Gametes, embryos, larvae and spat were exposed to four pCO₂ (375, 600, 750, 1,000 μatm) and four temperature (18, 22, 26, 30°C) levels. At elevated pCO₂ and suboptimal temperatures, there was a reduction in the fertilization success of gametes, a reduction in the development of embryos and size of larvae and spat and an increase in abnormal morphology of larvae. These effects varied between species and fertilization treatments with S. glomerata having greater sensitivity than C. gigas. In the absence of adaptation, C. gigas may become the more dominant species along the south-eastern coast of Australia, recruiting into estuaries currently dominated by the native S. glomerata.     

Wet oxidation of recalcitrant lignin water solutions: experimental and reaction kinetics

The purpose of the research was to study the behavior of lignin degradation under different conditions (T 110–190°C, pO₂ 0.5–1.5 MPa, pH 5, 9 and 12) and to develop a predictive model. Temperature increase improved lignin removal from 75% at 110°C to 100% at 190°C (experimental). Increasing the pH enhanced the lignin removal efficiency from 30 to 97% (experimental). The developed model predicted the lignin degradation and changes in COD, BOD and TOC. The model agreed well with the experimental data (R ² = 0.93 at pH 5 and 12).     

Effects of low oxygen concentrations on the hatching and viability of eggs of marine calanoid copepods

Hatching of the eggs of four species of copepods, Acartia tonsa, Labidocera aestiva, Tortanus discaudatus, and Centropages hamatus was suppressed at oxygen concentrations less than 0.02 ml O₂/l. When such eggs were subsequently exposed to normal oxygen concentrations, hatching varied among the species, indicating variability in the capacity of eggs to survive exposure to low oxygen. Incubation of eggs at low oxygen concentrations resulted in an increase in the duration of embryonic development. Experiments were conducted in closed systems and oxygen concentration was determined by thermal conductivity using gas chromatography.     

Effects of O2, pH and DIC on photosynthetic net-O2 evolution by marine macroalgae

Net photosynthetic O₂ evolution by five marine macroalgae:Ulva lactuca L.,Enteromorpha sp.,Ceramium strictum Harvey,Fucus serratus L., andF. vesiculosus L., collected from Danish waters in the summer of 1983 was followed at increasing O₂ and with pH either fixed close to pH 7, 8 or 9, or drifting upwards during photosynthesis in a closed chamber to determine the effects of changing O₂, pH and DIC (dissolved inorganic carbon) on photosynthesis. Increasing O₂, increasing pH and decreasing DIC together limited O₂ evolution. Raising the O₂ concentration with pH and DIC held constant resulted in less inhibition of net-O₂ evolution than when all three factors acted together. The O₂ inhibition of photosynthesis was similar to the reported O₂ inhibition of ribulose 1,5-bisphosphate carboxylase isolated from lower and higher plants. Net-O₂ evolution as a function of the molar ratio of O₂ to HCO^– ₃ + CO₂ in solution provided a general, linear relationship (r ² = 0.72 to 0.84), predicting inhibition of photosynthesis based on O₂ pH and DIC changing together. Slopes of this relationship, representing competition between O₂ and carbon based on external concentrations, were similar for the five taxonomically different algae, suggesting that similar processes act to reduce net-O₂ evolution.     

Effect of salinity on hypoxia tolerance of resting green crabs, Carcinus maenas, after feeding

Mechanisms that can influence the tolerance of hypoxia in brackish waters were studied in resting and fed crabs, Carcinus maenas, at 15?°C. Mortality, blood oxygenation, acid-base status and lactate concentration were analysed in fed crabs held in full-strength normoxic seawater (32.5‰?S) and then transferred for 24?h to a partial pressure of oxygen (P_o2) of 3?kPa (1.4?mg?l^?1) and various salinities (17, 12.5, 10, 8‰?S). At salinity levels >10‰, fed crabs tolerated P_o2 values as low as 3?kPa in the ambient water and 0.5?kPa in their arterial blood for 24?h without switching to anaerobic metabolism. Only below 10‰?S did their blood-lactate content rise, leading to their death despite the fact that their blood O₂-content was twice the control value measured in full-strength normoxic seawater and their blood P_o2 did not decrease below values recorded at higher salinity levels. Addition of CO₂ to 8‰?S water (CO₂ partial pressure increasing from 0.1 to 0.3?kPa) decreased blood-lactate production and mortality, suggesting that at 10‰?S impairment of the O₂ supply is limited by an excessive blood O₂-affinity. The results are discussed in terms of the distribution (?10‰?S) of C. maenas along salinity gradients in estuaries and bays.     

Prolonged exposure to low dissolved oxygen affects early development and swimming behaviour in the gastropod Nassarius festivus (Nassariidae)

Effects of low dissolved oxygen on early development and swimming behaviour of veliger larvae of the scavenging gastropod Nassarius festivus were studied. Embryonic development was significantly delayed when dissolved oxygen level was reduced to 3.0 mg O₂ l⁻¹ and no embryo hatched successfully at 0.5 mg O₂ l⁻¹. Veliger larvae hatched at 4.5 mg O₂ l⁻¹ had significantly smaller velar lobe, shell length and shell width. Median 48-h LC₅₀ value of the veliger larvae was estimated at 1.25 mg O₂ l⁻¹ with lower swimming speed (swimming velocity and dispersal velocity) being recorded for the survivors exposed to reduced oxygen levels. The percentage of veliger larvae that developed into crawling juveniles was significantly reduced and metamorphosis was delayed at 4.5 mg O₂ l⁻¹ whereas all larvae at 3.5 mg O₂ l⁻¹ died before they underwent metamorphosis. Juveniles developed at 4.5 mg O₂ l⁻¹ were also smaller than those at 6.0 mg O₂ l⁻¹. Results indicated that dissolved oxygen levels well above hypoxia levels (2.8 mg O₂ l⁻¹) have already had significant impact on the hatching success and larval development in gastropods, which may lead to long-term decreases in population growth.     

PIXGRO: A model for simulating the ecosystem CO2 exchange and growth of spring barley

A model, PIXGRO, developed by coupling a canopy flux sub-model (PROXEL_NEE; PROcess-based piXEL Net Ecosystem CO₂ Exchange) to a vegetation structure submodel (CGRO), for simulating both net ecosystem CO₂ exchange (NEE) and growth of spring barley is described. PIXGRO is an extension of the stand-level CO₂ and H₂O-flux model PROXEL_NEE, that simulates the NEE on a process basis, but goes further to include the dry matter production, partitioning, and crop development for spring barley. Dry matter partitioned to the leaf was converted to leaf area index (LAI) using relationships for the specific leaf area (SLA). The canopy flux component, PROXEL_NEE was calibrated using information from the literature on C3 plants and was tested using CO₂ flux data from an eddy-covariance (EC) method in Finland with long-term observations. The growth component (CGRO) was calibrated using data from the literature on spring barley as well as data from the Finland site. It was then validated against field data from two sites in Germany and partly via the use of MODIS remotely sensed LAI from the Finland site.Both the diurnal and the seasonal patterns of gross CO₂ uptake were very well simulated (R² = 0.92). A slight seasonal bias may be attributed to leaf ageing. Crop growth was also well simulated; simulated dry matter agreed with field observed data from Germany (R² = 0.90). For LAI, the agreement between the simulated and observed was good (R² = 0.80), giving an indication that functions describing the conversion of fixed CO₂ to dry matter and the subsequent partitioning leaf dry matter and LAI simulation were robust and provided reliable estimates.The MODIS LAI at a resolution of 1000 m agreed poorly (R² = 0.45) with the PIXGRO simulated LAI and the observed LAI at the Finland site in 2001. We attributed this to the coarse resolution of the image and/or the small size of the barley field (about 17 ha or 0.25 km²) at the Finland site. By deriving a regression relation between the observed LAI and NDVI from a higher resolution MODIS (500 m resolution), the MODIS-recalculated LAI agreed better with the PIXGRO-simulated LAI (R² = 0.86).PIXGRO provides a prototype model bridging the disciplines of plant physiology, crop modeling and remote sensing, for use in a spatial context in evaluating carbon balances and plant growth at stand level, landscape, regional, and with some care, continental scales. Since almost 50% of the European land surface is covered by crops, such a model is needed for the dynamic estimation of LAI and NEE of croplands.