Presence of competitors influences photosynthesis, but not growth, of the hard coral Porites cylindrica at elevated seawater CO2

Changes in environmental conditions, such as those caused by elevated carbon dioxide (CO2), potentially alter the outcome of competitive interactions between species. This study aimed to understand how elevated CO2 could influence competitive interactions between hard and soft corals, by investigating growth and photosynthetic activity of Porites cylindrica (a hard coral) under elevated CO2 and in the presence of another hard coral and two soft coral competitors. Corals were collected from reefs around Orpheus and Pelorus Islands on the Great Barrier Reef, Australia. They were then exposed to elevated pCO2 for 4 weeks with two CO2 treatments: intermediate (pCO2 648) and high (pCO2 1003) compared with a control (unmanipulated seawater) treatment (pCO2 358). Porites cylindrica growth did not vary among pCO2 treatments, regardless of the presence and type of competitors, nor was the growth of another hard coral species, Acropora cerealis, affected by pCO2 treatment. Photosynthetic rates of P. cylindrica were sensitive to variations in pCO2, and varied between the side of the fragment facing the competitors vs. the side facing away from the competitor. However, variation in photosynthetic rates depended on pCO2 treatment, competitor identity, and whether the photosynthetic yields were measured as maximum or effective photosynthetic yield. This study suggests that elevated CO2 may impair photosynthetic activity, but not growth, of a hard coral under competition and confirms the hypothesis that soft corals are generally resistant to elevated CO2. Overall, our results indicate that shifts in the species composition in coral communities as a result of elevated CO2 could be more strongly related to the individual tolerance of different species rather than a result of competitive interactions between species.

Food selectivity and processing by the cold-water coral Lophelia pertusa

Cold-water corals form prominent reef ecosystems along ocean margins that depend on suspended resources produced in surface waters. In this study, we investigated food processing of 13C and 15N labelled bacteria and algae by the cold-water coral Lophelia pertusa. Coral respiration, tissue incorporation of C and N and metabolic-derived C incorporation into the skeleton were traced following the additions of different food concentrations (100, 300, 1300 µg C/l) and two ratios of suspended bacterial and algal biomass (1:1, 3:1). Respiration and tissue incorporation by L. pertusa increased markedly following exposure to higher food concentrations. The net growth efficiency of L. pertusa was low (0.08±0.03), which is consistent with their slow growth rates. The contribution of algae and bacteria to total coral assimilation was proportional to the food mixture in the two lowest food concentrations, but algae were preferred over bacteria as food source at the highest food concentration. Similarly, the stoichiometric uptake of C and N was coupled in the low and medium food treatment, but was uncoupled in the high food treatment and indicated a comparatively higher uptake or retention of bacterial carbon as compared to algal nitrogen. We argue that behavioural responses for these small-sized food particles, such as tentacle behaviour, mucus trapping and physiological processing, are more likely to explain the observed food selectivity as compared to physical-mechanical considerations. A comparison of the experimental food conditions to natural organic carbon concentrations above CWC reefs suggests that L. pertusa is well adapted to exploit temporal pulses of high organic matter concentrations in the bottom water caused by internal waves and down-welling events.

Seawater carbonate chemistry and net ecosystem calcification and production in a coral community Kaneohe Bay, Oahu, Hawaii, 2011

Net ecosystem calcification rates (NEC) and net photosynthesis (NP) were determined from CO2 seawater parameters on the barrier coral reef of Kaneohe Bay, Oahu, Hawaii. Autosamplers were deployed to collect samples on the barrier reef every 2 hours for six 48-hour deployments, two each in June 2008, August 2009, and January/February 2010. NEC on the Kaneohe Bay barrier reef increased throughout the day and decreased at night. Net calcification continued at low rates at night except for six time periods when net dissolution was measured. The barrier reef was generally net photosynthetic (positive NP) during the day and net respiring (negative NP) at night. NP controlled the diel cycles of the partial pressure of CO2 (pCO2) and aragonite saturation state resulting in high daytime aragonite saturation state levels when calcification rates were at their peak. However, the NEC and NP diel cycles can become decoupled for short periods of time (several hours) without affecting calcification rates. On a net daily basis, net ecosystem production (NEP) of the barrier reef was found to be sometimes net photosynthetic and sometimes net respiring and ranged from -378 to 80 mmol m-2 d-1 when calculated using simple box models. Daily NEC of the barrier reef was positive (net calcification) for all deployments and ranged from 174 to 331 mmol CaCO3 m-2 d-1. Daily NEC was strongly negatively correlated with average daily pCO2 (R2 = 0.76) which ranged from 431 to 622 µatm. Daily NEC of the Kaneohe Bay barrier reef is similar to or higher than daily NEC measured on other coral reefs even though aragonite saturation state levels (mean aragonite saturation state = 2.85) are some of the lowest measured in coral reef ecosystems. It appears that while calcification rate and ?arag are correlated within a single coral reef ecosystem, this relationship does not necessarily hold between different coral reef systems. It can be expected that ocean acidification will not affect coral reefs uniformly and that some may be more sensitive to increasing pCO2 levels than others.

Seawater carbonate chemistry, Mg/Ca, Sr/Ca, oxygen production and calcification rate during experiments with coral Pocillopora damicornis, 2011

The Sr/Ca of aragonitic coral skeletons is a commonly used palaeothermometer. However skeletal Sr/Ca is typically dominated by weekly-monthly oscillations which do not reflect temperature or seawater composition and the origins of which are currently unknown. To test the impact of transcellular Ca2+ transport processes on skeletal Sr/Ca, colonies of the branching coral, Pocillopora damicornis, were cultured in the presence of inhibitors of Ca-ATPase (ruthenium red) and Ca channels (verapamil hydrochloride). The photosynthesis, respiration and calcification rates of the colonies were monitored throughout the experiment. The skeleton deposited in the presence of the inhibitors was identified (by 42Ca spike) and analysed for Sr/Ca and Mg/Ca by secondary ion mass spectrometry. The Sr/Ca of the aragonite deposited in the presence of either of the inhibitors was not significantly different from that of the solvent (dimethyl sulfoxide) control, although the coral calcification rate was reduced by up to 66% and 73% in the ruthenium red and verapamil treatments, respectively. The typical precision (95% confidence limits) of mean Sr/Ca determinations within any treatment was <±1% and differences in skeletal Sr/Ca between treatments were correspondingly small. Either Ca-ATPase and Ca channels transport Sr2+ and Ca2+ in virtually the same ratio in which they are present in seawater or transcellular processes contribute little Ca2+ to the skeleton and most Ca is derived from seawater transported directly to the calcification site. Variations in the activities of Ca-ATPase and Ca-channels are not responsible for the weekly-monthly Sr/Ca oscillations observed in skeletal chronologies, assuming that the specificities of Ca transcellular transport processes are similar between coral genera.

Seawater carbonate chemistry, photosynthesis, respiration and calcification during experiments with scleractinian coral Stylophora pistillata, 2003

We show here that CO2 partial pressure (pCO2) and temperature significantly interact on coral physiology. The effects of increased pCO2 and temperature on photosynthesis, respiration and calcification rates were investigated in the scleractinian coral Stylophora pistillata. Cuttings were exposed to temperatures of 25°C or 28°C and to pCO2 values of ca. 460 or 760 muatm for 5 weeks. The contents of chlorophyll c2 and protein remained constant throughout the experiment, while the chlorophyll a content was significantly affected by temperature, and was higher under the 'high-temperature-high-pCO2' condition. The cell-specific density was higher at 'high pCO2' than at 'normal pCO2' (1.7 vs. 1.4). The net photosynthesis normalized per unit protein was affected by both temperature and pCO2, whereas respiration was not affected by the treatments. Calcification decreased by 50% when temperature and pCO2 were both elevated. Calcification under normal temperature did not change in response to an increased pCO2. This is not in agreement with numerous published papers that describe a negative relationship between marine calcification and CO2. The confounding effect of temperature has the potential to explain a large portion of the variability of the relationship between calcification and pCO2 reported in the literature, and warrants a re-evaluation of the projected decrease of marine calcification by the year 2100.

Coral and mollusc resistance to ocean acidification adversely affected by warming, 2011

Increasing atmospheric carbon dioxide (CO2) concentrations are expectedto decrease surface ocean pH by 0.3-0.5 units by 2100, lowering the carbonate ion concentration of surfacewaters. This rapid acidification is predicted to dramatically decrease calcification in many marine organisms. Reduced skeletal growth under increased CO2 levels has already been shown for corals, molluscs and many other marine organisms. The impact of acidification on the ability of individual species to calcify has remained elusive, however, as measuring net calcification fails to disentangle the relative contributions of gross calcification and dissolution rates on growth. Here, we show that corals and molluscs transplanted along gradients of carbonate saturation state at Mediterranean CO2 vents are able to calcify and grow at even faster than normal rates when exposed to the high CO2 levels projected for the next 300 years. Calcifiers remain at risk, however, owing to the dissolution of exposed shells and skeletons that occurs as pH levels fall. Our results show that tissues and external organic layers play a major role in protecting shells and skeletons from corrosive sea water, limiting dissolution and allowing organisms to calcify. Our combined field and laboratory results demonstrate that the adverse effects of global warming are exacerbated when high temperatures coincide with acidification.

Seawater carbonate chemistry, calcification and dissolution of a coral reef in the northern Red Sea, 2007

In this study we investigated the relations between community calcification of an entire coral reef in the northern Red Sea and annual changes in temperature, aragonite saturation and nutrient loading over a two year period. Summer (April-October) and winter (November-March) average calcification rates varied between 60 ± 20 and 30 ± 20 mmol·m-2·d-1, respectively. In general, calcification increased with temperature and aragonite saturation state of reef water with an apparent effect of nutrients, which is in agreement with most laboratory studies and in situ measurements of single coral growth rates. The calcification rates we measured in the reef correlated remarkably well with precipitation rates of inorganic aragonite calculated for the same temperature and degree of saturation ranges using empirical equations from the literature. This is a very significant finding considering that only a minute portion of reef calcification is inorganic. Hence, these relations could be used to predict the response of coral reefs to ocean acidification and warming.

Seawater carbonate chemistry and community calcification during Reunion Island coral reef community study, 2007

To assess the contribution of soft-bottoms to the carbon cycle in coral reefs, the net community production (p) was measured in winter at 3 stations on La Saline inner reef flat (Reunion Island). Changes in pH and total alkalinity at different irradiances (I) were assessed using benthic chambers (0.2 m2) during a 1-h incubation. Mean grain size, the silt and clay load and chlorophyll a content of the sediments were analysed in each chamber. Daily community production (P), gross community production (Pg) and community respiration (R) were estimated from p-I curves and daily irradiance variations (PAR, 400-700 nm). Sediment characteristics and chlorophyll a contents did not differ between the three sites, except for the silt and clay fraction at one station. R being higher than Pg (84.88 ± 7.36 and -62.29 ± 3.34 mmolC m-2 d-1 respectively), P value reached 22.59 ± 5.66 mmolC m-2 d-1. The sediments were therefore heterotrophic with a mean Pg/R lower than 1 (0.74 ± 0.05) and appear to be a carbon source. Our data suggested the importance of the degradation process in the functioning of near-reef sediments.

Seawater carbonate chemistry and community metabolism during an experiment with a coral reef, 2003

The effect of elevated pCO2 on the metabolism of a coral reef community dominated by macroalgae has been investigated utilizing the large 2650 m3 coral reef mesocosm at the Biosphere-2 facility near Tucson, Arizona. The carbonate chemistry of the water was manipulated to simulate present-day and a doubled CO2 future condition. Each experiment consisted of a 1-2 month preconditioning period followed by a 7-9 day observational period. The pCO2 was 404 ± 63 ?atm during the present-day pCO2 experiment and 658 ± 59 ?atm during the elevated pCO2 experiment. Nutrient levels were low and typical of natural reefs waters (NO3? 0.5-0.9 ?M, NH4+ 0.4 ?M, PO43? 0.07-0.09 ?M). The temperature and salinity of the water were held constant at 26.5 ± 0.2°C and 34.4 ± 0.2 ppt. Photosynthetically available irradiance was 10 ± 2 during the present-day experiment and 7.4 ± 0.5 mol photons m?2 d?1 during the elevated pCO2 experiment. The primary producer biomass in the mesocosm was dominated by four species of macroalgae; Haptilon cubense, Amphiroa fragillisima, Gelidiopsis intricata and Chondria dasyphylla. Algal biomass was 10.4 mol C m?2 during the present-day and 8.7 mol C m?2 and during the elevated pCO2 experiments. As previously observed, the increase in pCO2 resulted in a decrease in calcification from 0.041 ± 0.007 to 0.006 ± 0.003 mol CaCO3 m?2 d?1. Net community production (NCP) and dark respiration did not change in response to elevated pCO2. Light respiration measured by a new radiocarbon isotope dilution method exceeded dark respiration by a factor of 1.2 ± 0.3 to 2.1 ± 0.4 on a daily basis and by 2.2 ± 0.6 to 3.9 ± 0.8 on an hourly basis. The 1.8-fold increase with increasing pCO2 indicates that the enhanced respiration in the light was not due to photorespiration. Gross production (GPP) computed as the sum of NCP plus daily respiration (light + dark) increased significantly (0.24 ± 0.03 vs. 0.32 ± 0.04 mol C m?2 d?1). However, the conventional calculation of GPP based on the assumption that respiration in the light proceeds at the same rate as the dark underestimated the true rate of GPP by 41-100% and completely missed the increased rate of carbon cycling due to elevated pCO2. We conclude that under natural, undisturbed, nutrient-limited conditions elevated CO2 depresses calcification, stimulates the rate of turnover of organic carbon, particularly in the light, but has no effect on net organic production. The hypothesis that an increase pCO2 would produce an increase in net production that would counterbalance the effect of decreasing saturation state on calcification is not supported by these data.

Seawater carbonate chemistry and calcification rate during mesocosm experiments with coral Montipora capitata, 2008

A long-term (10 months) controlled experiment was conducted to test the impact of increased partial pressure of carbon dioxide (pCO2) on common calcifying coral reef organisms. The experiment was conducted in replicate continuous flow coral reef mesocosms flushed with unfiltered sea water from Kaneohe Bay, Oahu, Hawaii. Mesocosms were located in full sunlight and experienced diurnal and seasonal fluctuations in temperature and sea water chemistry characteristic of the adjacent reef flat. Treatment mesocosms were manipulated to simulate an increase in pCO2 to levels expected in this century [midday pCO2 levels exceeding control mesocosms by 365 ± 130 µatm (mean ± sd)]. Acidification had a profound impact on the development and growth of crustose coralline algae (CCA) populations. During the experiment, CCA developed 25% cover in the control mesocosms and only 4% in the acidified mesocosms, representing an 86% relative reduction. Free-living associations of CCA known as rhodoliths living in the control mesocosms grew at a rate of 0.6 g buoyant weight per year while those in the acidified experimental treatment decreased in weight at a rate of 0.9 g buoyant weight per year, representing a 250% difference. CCA play an important role in the growth and stabilization of carbonate reefs, so future changes of this magnitude could greatly impact coral reefs throughout the world. Coral calcification decreased between 15% and 20% under acidified conditions. Linear extension decreased by 14% under acidified conditions in one experiment. Larvae of the coral Pocillopora damicornis were able to recruit under the acidified conditions. In addition, there was no significant difference in production of gametes by the coral Montipora capitata after 6 months of exposure to the treatments.

Cape Lookout cold-water coral area, coral ages and environmental parameters for the last glacial cycle

Climatic and oceanographic changes, as occurring at a glacial-interglacial scale, may alter the environmental conditions needed for the development of prolific cold-water coral reefs and mounds. Studies constraining the temporal distribution of cold-water corals in the NE Atlantic suggested the cyclic changes of the Atlantic Meridional Overturning Circulation as the main driver for the development and dispersal of cold-water coral ecosystems. However, conclusions were hindered by lack of data from the NW Atlantic. Aiming to overcome this lack of data, the temporal occurrence of cold-water corals in the Cape Lookout area along the southeastern US margin was explored by U-series dating. Furthermore, the local influence of the regional water masses, namely the Gulf Stream, on cold-water coral proliferation and occurrence since the Last Glacial Maximum was examined. Results suggest that the occurrence of cold-water corals in the Cape Lookout area is restricted to interglacial periods, with corals being present during the last ~7 kyr and also during the Eemian (~125 ka). The reconstructed local environmental conditions suggest an offshore displacement of the Gulf Stream and increased influence from the Mid-Atlantic Bight shelf waters during the last glacial period. During the deglacial sea level rise, the Gulf Stream moved coastward providing present-day-like conditions to the surface waters. Nevertheless, present-day conditions at the ocean sea floor were not established before 7.5 cal ka BP once the ultimate demise of the Laurentide ice-sheet caused the final sea level rise and the displacement of the Gulf Stream to its present location. Occasional presence of the Gulf Stream over the site during the Mid- to Late Holocene coincides with enhanced bottom current strength and a slightly higher bottom water temperature, which are environmental conditions that are favorable for cold-water coral growth.