Adult acclimation to combined temperature and pH stressors significantly enhances reproductive outcomes compared to short-term exposures

This study examined the effects of long-term culture under altered conditions on the Antarctic sea urchin, Sterechinus neumayeri. Sterechinus neumayeri was cultured under the combined environmental stressors of lowered pH (-0.3 and -0.5 pH units) and increased temperature (+2 °C) for 2 years. This time-scale covered two full reproductive cycles in this species and analyses included studies on both adult metabolism and larval development. Adults took at least 6-8 months to acclimate to the altered conditions, but beyond this, there was no detectable effect of temperature or pH. Animals were spawned after 6 and 17 months exposure to altered conditions, with markedly different outcomes. At 6 months, the percentage hatching and larval survival rates were greatest in the animals kept at 0 °C under current pH conditions, whilst those under lowered pH and +2 °C performed significantly less well. After 17 months, performance was not significantly different across treatments, including controls. However, under the altered conditions urchins produced larger eggs compared with control animals. These data show that under long-term culture adult S. neumayeri appear to acclimate their metabolic and reproductive physiology to the combined stressors of altered pH and increased temperature, with relatively little measureable effect. They also emphasize the importance of long-term studies in evaluating effects of altered pH, particularly in slow developing marine species with long gonad maturation times, as the effects of altered conditions cannot be accurately evaluated unless gonads have fully matured under the new conditions.

Size matters: plasticity in metabolic scaling shows body-size may modulate responses to climate change

Variability in metabolic scaling in animals, the relationship between metabolic rate ( R) and body mass ( M), has been a source of debate and controversy for decades. R is proportional to Mb, the precise value of b much debated, but historically considered equal in all organisms. Recent metabolic theory, however, predicts b to vary among species with ecology and metabolic level, and may also vary within species under different abiotic conditions. Under climate change, most species will experience increased temperatures, and marine organisms will experience the additional stressor of decreased seawater pH ('ocean acidification'). Responses to these environmental changes are modulated by myriad species-specific factors. Body-size is a fundamental biological parameter, but its modulating role is relatively unexplored. Here, we show that changes to metabolic scaling reveal asymmetric responses to stressors across body-size ranges; b is systematically decreased under increasing temperature in three grazing molluscs, indicating smaller individuals were more responsive to warming. Larger individuals were, however, more responsive to reduced seawater pH in low temperatures. These alterations to the allometry of metabolism highlight abiotic control of metabolic scaling, and indicate that responses to climate warming and ocean acidification may be modulated by body-size.

Sea urchins in a high CO2 world: partitioned effects of body-size, ocean warming and acidification on metabolic rate

Body-size and temperature are the major factors explaining metabolic rate, and the additional factor of pH is a major driver at the biochemical level. These three factors have frequently been found to interact, complicating the formulation of broad models predicting metabolic rates and hence ecological functioning. In this first study of the effects of warming and ocean acidification, and their potential interaction, on metabolic rate across a broad body-size range (two-to-three orders of magnitude difference in body mass) we addressed the impact of climate change on the sea urchin Heliocidaris erythrogramma in context with climate projections for east Australia, an ocean warming hotspot. Urchins were gradually introduced to two temperatures (18 and 23 °C) and two pH (7.5 and 8.0), and maintained for two months. That a new physiological steady-state had been reached, otherwise know as acclimation, was validated through identical experimental trials separated by several weeks. The relationship between body-size, temperature and acidification on the metabolic rate of H. erythrogramma was strikingly stable. Both stressors caused increases in metabolic rate; 20% for temperature and 19% for pH. Combined effects were additive; a 44% increase in metabolism. Body-size had a highly stable relationship with metabolic rate regardless of temperature or pH. None of these diverse drivers of metabolism interacted or modulated the effects of the others, highlighting the partitioned nature of how each influences metabolic rate, and the importance of achieving a full acclimation state. Despite these increases in energetic demand there was very limited capacity for compensatory modulating of feeding rate; food consumption increased only in the very smallest specimens, and only in response to temperature, and not pH. Our data show that warming, acidification and body-size all substantially affect metabolism and are highly consistent and partitioned in their effects, and for H. erythrogramma near-future climate change will incur a substantial energetic cost.

Paleontological and sedimentological investigations of a sediment profile from northeast Greenland

The Kap Mackenzie area on the outer coast of northeast Greenland was glaciated during the last glacial stage, and pre-Holocene shell material was brought to the area. Dating of marine shells indicates that deglaciation occurred in the earliest Holocene, before 10 800 cal. a BP. The marine limit is around 53 m a.s.l. In the wake of the deglaciation, a glaciomarine fauna characterized the area, but after c. one millennium a more species-rich marine fauna took over. This fauna included Mytilus edulis and Mysella sovaliki, which do not live in the region at present; the latter is new to the Holocene fauna of northeast Greenland. The oldest M. edulis sample is dated to c. 9500 cal. a BP, which is the earliest date for the species from the region and indicates that the Holocene thermal maximum began earlier in the region than previously documented. This is supported by driftwood dated to c. 9650 cal. a BP, which is the earliest driftwood date so far from northeastern Greenland and implies that the coastal area was at least partly free of sea ice in summer. As indicated by former studies, the Storegga tsunami hit the Kap Mackenzie area at c. 8100 cal. a BP. Loon Lake, at 18 m a.s.l., was isolated from the sea at c. 6200 cal. a BP, which is distinctly later than expected from existing relative sea-level curves for the region.

Food choice in Siganus fuscescens: influence of macrophyte nutrient content and availability

Using multiple-choice feeding experiments, the selection of six species of macrophytes by the herbivorous rabbitfish Siganus fuscescens was examined. The rabbitfish showed distinct food choice in the laboratory; however, selection of macrophytes by S. fuscescens was not related to their absolute nutrient content (nitrogen, carbon, energy and ash free dry mass). Nutrient assimilation estimates showed that the macrophytes which were most preferred were those that S. fuscescens assimilated best. In S. fuscescens, the macrophytes that were preferred passed through the gut significantly faster than the less preferred species. Gut transit time had a significant effect on the absolute value of a food item in terms of net nutrient gain per unit time. This study showed that food value could be inferred from the absolute nutrient content of the macrophytes. Thus both the ability to assimilate nutrients as well as the absolute nutrient content of macrophytes must be quantified when assessing food value. (C) 2004 The Fisheries society of the British Isles.

Lack of an osmotic constraint on intake rate of the eastern curlew Numenius madagascariensis

Rates of food intake in animals consuming abundant prey can be constrained by the rates of digestion or excretion of ingested substances, such as salt, particularly so in the animals that regularly migrate between freshwater and saltwater environments. We tested this hypothesis in a long-distance migrant shorebird, the eastern curlew Numenius madagascariensis (suborder Charadrii), foraging on intertidal decapods in eastern Australia. We predicted that if food intake rates are constrained osmotically, individuals with access to freshwater and less saline prey (FW group) would have higher rates of food and water intake than individuals with seawater-only access (SW group). Food intake rates did not differ between the FW and SW groups (0.14 g ash-free dry mass min(-1)), nor did the water influx rates (0.75 g min(-1)). Salt intake rates were lower at FW sites (19.3 versus 23.3 mg NaCl min(-1)) and overall they were similar to those of marine birds. Food intake rate in the eastern curlew appeared limited by digestive rather than by osmoregulatory capacity.

Intake rates and the functional response in shorebirds (Charadriiformes) eating macro-invertebrates

As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type 11 ('disc equation') formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching. A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (< 150/m(-2)). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote. A multivariate analysis of 468 'spot' estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81 % of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3 %), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6%, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93 % of cases. We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested. As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shorebird ecology and behaviour to estimate intake rate without the need for conventional time-consuming field studies, including species for which it has not yet proved possible to measure intake rate in the field.

Fast pyrolysis of biomass in a circulating fluidised bed reactor

The objective of this work was to design, construct, test and operate a novel circulating fluid bed fast pyrolysis reactor system for production of liquids from biomass. The novelty lies in incorporating an integral char combustor to provide autothermal operation. A reactor design methodology was devised which correlated input parameters to process variables, namely temperature, heat transfer and gas/vapour residence time, for both the char combustor and biomass pyrolyser. From this methodology a CFB reactor was designed with integral char combustion for 10 kg/h biomass throughput. A full-scale cold model of the CFB unit was constructed and tested to derive suitable hydrodynamic relationships and performance constraints. Early difficulties encountered with poor solids circulation and inefficient product recovery were overcome by a series of modifications. A total of 11 runs in a pyrolysis mode were carried out with a maximum total liquids yield of 61.50% wt on a maf biomass basis, obtained at 500°C and with 0.46 s gas/vapour residence time. This could be improved by improved vapour recovery by direct quenching up to an anticipated 75 % wt on a moisture-and-ash-free biomass basis. The reactor provides a very high specific throughput of 1.12 - 1.48 kg/hm2 and the lowest gas-to-feed ratio of 1.3 - 1.9 kg gas/kg feed compared to other fast pyrolysis processes based on pneumatic reactors and has a good scale-up potential. These features should provide significant capital cost reduction. Results to date suggest that the process is limited by the extent of char combustion. Future work will address resizing of the char combustor to increase overall system capacity, improvement in solid separation and substantially better liquid recovery. Extended testing will provide better evaluation of steady state operation and provide data for process simulation and reactor modeling.

Effect of the temperature on the composition of lignin pyrolysis products

The temperature dependence of the pyrolysis products of two types of lignin (Alcell lignin and Asian lignin) was investigated using pyrolysis-gas chromatography-mass spectrometry (PyGC-MS). About 50 compounds were identified and quantified for each type of lignin over a temperature range of 400-800C. The maximum yield of phenolic compounds was obtained at 600C for both lignins, which was 17.2% for Alcell lignin and 15.5% for Asian lignin. Most of the phenolic compounds had an individual yield of less than 1%; however, for Alcell lignin, 5-hydroxyvanillin was the highest yield at 4.29 wt%on dry ash-free lignin, and for Asian lignin, 2-methoxy-4-vinylphenol was the highest yield at 4.15 wt % on dry ash-free lignin.

Physiological energetics of the thick shell mussel Mytilus coruscus exposed to seawater acidification and thermal stress

Anthropogenic CO2 emissions have caused seawater temperature elevation and ocean acidification. In view of both phenomena are occurring simultaneously, their combined effects on marine species must be experimentally evaluated. The purpose of this study was to estimate the combined effects of seawater acidification and temperature increase on the energy budget of the thick shell mussel Mytilus coruscus. Juvenile mussels were exposed to six combined treatments with three pH levels (8.1, 7.7 and 7.3) * two temperatures (25 °C and 30 °C) for 14 d. We found that clearance rates (CRs), food absorption efficiencies (AEs), respiration rates (RRs), ammonium excretion rates (ER), scope for growth (SFG) and O:N ratios were significantly reduced by elevated temperature sometimes during the whole experiments. Low pH showed significant negative effects on RR and ER, and significantly increased O:N ratios, but showed almost no effects on CR, AE and SFG of M. coruscus. Nevertheless, their interactive effects were observed in RR, ER and O:N ratios. PCA revealed positive relationships among most physiological indicators, especially between SFG and CR under normal temperatures compared to high temperatures. PCA also showed that the high RR was closely correlated to an increasing ER with increasing pH levels. These results suggest that physiological energetics of juvenile M. coruscus are able to acclimate to CO2 acidification with a little physiological effect, but not increased temperatures. Therefore, the negative effects of a temperature increase could potentially impact the ecophysiological responses of M. coruscus and have significant ecological consequences, mainly in those habitats where this species is dominant in terms of abundance and biomass.

The distribution of meiofauna in surface sediments of the Fladen Ground, North Sea

A general study of structure, biomass, and dynamic estimates on meiofauna was carried out during PREFLEX (1975) and FLEX (1976), in 117- 141 m water depth. On the basis of these data an attempt was made to estimate meiofauna production, and this is discussed in relation to the energy input from the spring phytoplankton bloom. Sampling was performed at five stations, but only the stations 1, 4, and 5 were covered by a complete series from August 1975 to July 1976. At each station, from four replicate box core samples, two were withdrawn to study the abundance, distribution, and biomass of meiofauna, the content of chloroplastic pigment equivalents (CPE), and chemical and grain size analyses. At all stations grain size fell in the range of fine sand having median diameters (MD) of < 125 µm. From station 1 to 5 an increase in MD was observed. Highest values of CPE (7.81 µg m l**-1) and organic matter (4.7 %) were obtained in June and July (1976)/ August (1975), respectively. Meiofauna abundance was fairly uniform at all stations examined. Station 1 displayed maximal numbers during the whole investigation period. The abundance per 100 cm**2 varied between 15,550 and 34,900 organisms. All meiofauna studied both in total and as separate taxa showed annual cycles of abundance. Low abundance values were recorded during early summer, and maximum values during winter. High numbers of Foraminifera were obtained for August 1975 (9,460 per 100 cm**2) and July 1976 (9,710 per 100 cm**2). From December to June the values decreased from 3,280 to 1,030 per 100 cm**2. At station 1 maximum values of meiofauna biomass were recorded ranging from 1.5 to 2.7 g DWT m**-2. The mean meiofauna dry weight amounted to 2.1 g DWT m**-2. Based on minimum production, the P/B ratio for the area of station 1 might have a mean of 1.4. Taking into consideration generation times we believe that a turnover ratio of 2 is a conservative value for the Fladen Ground meiofauna. The annual production would amount to 4.2 g DWT m**-2 yr**-1. This is 27.5 % of the energy supply during the spring phytoplankton bloom, which is channelled into the meiofauna. The hypothesis is put forward that the energetic strategy of deep offshore meiofauna differs distinctively from that of shallow inshore meiofauna. While the shallow inshore meiofauna show a relatively fast response to organic matter input, the deep offshore meiofauna reacts much more slowly, the food energy consumption seems to be spread out over a longer period.