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Smaller body size under warming is not due to gill-oxygen limitation in a coldwater salmonid

Declining body size in fishes and other aquatic ectotherms associated with anthropogenic climate warming has significant implications for future fisheries yields, stock assessments and aquatic ecosystem stability. One proposed mechanism seeking to explain such body-size reductions, known as the gill oxygen limitation (GOL) hypothesis, has recently been used to model future impacts of climate warming on fisheries but has not been robustly empirically tested. We used brook trout ( Salvelinus fontinalis ), a fast-growing, cold-water salmonid species of broad economic, conservation and ecological value, to examine the GOL hypothesis in a long-term experiment quantifying effects of temperature on growth, resting metabolic rate (RMR), maximum metabolic rate (MMR) and gill surface area (GSA). Despite significantly reduced growth and body size at an elevated temperature, allometric slopes of GSA were not significantly different than 1.0 and were above those for RMR and MMR at both temperature treatments (15°C and 20°C), contrary to GOL expectations. We also found that the effect of temperature on RMR was time-dependent, contradicting the prediction that heightened temperatures increase metabolic rates and reinforcing the importance of longer-term exposures (e.g. >6 months) to fully understand the influence of acclimation on temperature–metabolic rate relationships. Our results indicate that although oxygen limitation may be important in some aspects of temperature–body size relationships and constraints on metabolic supply may contribute to reduced growth in some cases, it is unlikely that GOL is a universal mechanism explaining temperature–body size relationships in aquatic ectotherms. We suggest future research focus on alternative mechanisms underlying temperature–body size relationships, and that projections of climate change impacts on fisheries yields using models based on GOL assumptions be interpreted with caution.

Journal of Experimental Biology

Male lake char release taurocholic acid as part of a mating pheromone

The evolutionary origins of sexual preferences for chemical signals remain poorly understood, due, in part, to scant information on the molecules involved. In the current study, we identified a male pheromone in lake char ( Salvelinus namaycush ) to evaluate the hypothesis that it exploits a non-sexual preference for juvenile odour. In anadromous char species, the odour of stream-resident juveniles guides migratory adults into spawning streams. Lake char are also attracted to juvenile odour but have lost the anadromous phenotype and spawn on nearshore reefs, where juvenile odour does not persist long enough to act as a cue for spawning site selection by adults. Previous behavioural data raised the possibility that males release a pheromone that includes components of juvenile odour. Using metabolomics, we found that the most abundant molecule released by males was also released by juveniles but not females. Tandem mass spectrometry and nuclear magnetic resonance were used to identify the molecule as taurocholic acid (TCA), which was previously implicated as a component of juvenile odour. Additional chemical analyses revealed that males release TCA at high rates via their urine during the spawning season. Finally, picomolar concentrations of TCA attracted pre-spawning and spawning females but not males. Taken together, our results indicate that male lake char release TCA as a mating pheromone and support the hypothesis that the pheromone is a partial match of juvenile odour.

Journal of Experimental Biology

Glucocorticoid and glycemic responses to immune challenge in a viviparous snake afflicted with an emerging mycosis

Disease may be both a cause and consequence of stress, and physiological responses to infectious disease may involve stress coping mechanisms that have important fitness consequences. For example, glucocorticoid and glycemic responses may affect host fitness by altering resource allocation and use in hosts, and these responses may be affected by competing stressors. To better understand the factors that affect host responses to infection, we challenged the immune system of field acclimatized pygmy rattlesnakes, Sistrurus miliarius , with a sterile antigen, lipopolysaccharide (LPS), and measured the glucocorticoid and glycemic response in healthy non-reproductive snakes, snakes afflicted with an emerging mycosis (ophidiomycosis), and pregnant snakes. We hypothesized that LPS challenge would result in a glucocorticoid and glycemic response typical of the vertebrate acute phase response (APR), and therefore predicted that LPS challenge would result in an acute increase in plasma corticosterone (CORT) and a decline in plasma glucose in all individuals. Additionally, we hypothesized that the APR would be attenuated in individuals simultaneously coping with additional challenges to homeostasis (i.e., disease or reproduction). As predicted, immune challenge elicited an acute increase in plasma CORT and a decrease in plasma glucose. Snakes coping with ophidiomycosis and pregnant snakes were able to mount a robust glucocorticoid and hypoglycemic response to LPS challenge, which was contrary to our hypothesis. Our findings clarify directions of causality linking infection, glucocorticoids, and glucose, and emphasize the importance of future research examining the fitness consequences of interactions between stress and disease in wildlife threatened by emerging pathogens.

Journal of Experimental Biology

Not all who meander are lost: Migrating sea lamprey follow river thalwegs to facilitate safe and efficient passage upstream

Efficient navigation is crucial for the reproductive success of many migratory species, often driven by competing pressures to conserve energy and reduce predation risk. Little is known about how non-homing species achieve this balance. We show that sea lamprey ( Petromyzon marinus ), an ancient extant vertebrate, uses persistent patterns in hydro-geomorphology to quickly and efficiently navigate through complex ecosystems. Hydrodynamic flow models coupled with bathymetric mapping and fine-scale acoustic telemetry revealed movement paths that tracked thalweg scour channels, which are often the deepest and fastest-flowing sections of a river. These paths allow rapid and efficient upstream migration and suggest the existence of a bathymetric highway system. Near-substrate swimming along this path resulted in a median of 5.8% energy savings while also promoting improved safety from nocturnally active predators. We hypothesize sea lampreys use hydrostatic pressure-guided rheotaxis to achieve this navigation. It is likely this tactic relies on sensory information from the animal's primitive lateral line and perhaps the inner ear. Insights from this study can be used to redesign conservation practices to achieve improved control where the animal is invasive and improved fish passage within its native range.

Michigan

A Cftr-independent, Ano1-rich seawater-adaptive ionocyte in sea lamprey gills

All ionoregulating marine fishes examined to date utilize seawater-type ionocytes expressing the apical Cl- channel, cystic fibrosis transmembrane conductance regulator (Cftr) to secrete Cl − . We performed transcriptomic, molecular, and functional studies to identify Cl − transporters in the seawater-type ionocytes of sea lamprey ( Petromyzon marinus ). Gill cftr expression was minimal or undetectable in larvae and post-metamorphic juveniles. We identified other Cl − transporters highly expressed in the gills and/or upregulated following metamorphosis and further investigated two candidates that stood out in our analysis, a Ca 2+ -activated Cl − channel, anoctamin 1 ( ano1 ), and the Clc chloride channel family member 2 ( clcn2 ). Of these, ano1 was expressed 10-100 times more than clcn2 in the gills; moreover, ano1 was upregulated during seawater acclimation, while clcn2 was not. Using an antibody raised against sea lamprey Ano1, we did not detect Ano1 in the gills of larvae, found elevated levels in juveniles and observed a 4-fold increase in juveniles after seawater acclimation. Ano1 was localized to seawater-type branchial ionocytes but, surprisingly, was localized to the basolateral membrane. In vivo pharmacological inhibition experiments demonstrated that a DIDS-sensitive mechanism was critical to the maintenance of osmoregulatory homeostasis in seawater- but not freshwater-acclimated sea lamprey. Taken together, our results provide evidence of a Cftr-independent mechanism for branchial Cl − secretion in sea lamprey that leverages Ano1-expressing ionocytes. Once further characterized, the Cftr-independent, Ano1-rich ionocytes of sea lamprey could reveal novel strategies for branchial Cl − secretion, whether by Ano1 or some other Cl − transporter, not previously known in ionoregulating marine organisms.

Journal of Experimental Biology

Fifty-fifth Christmas Bird Count. 159. Ocean City, Md

The distribution of dynamic pressure behind a Harris' hawk's wing was sampled using a wake rake consisting of 15 pitot tubes and one static tube. The hawk was holding on to a perch, but at an air speed and gliding angle at which it was capable of gliding. The perch was instrumented, so that the lift developed by the wing was known and the lift coefficient could be calculated. The mean of 92 estimates of profile drag coefficient was 0.0207, with standard deviation 0.0079. Lift coefficients ranged from 0.51 to 1.08. Reynolds numbers were nearly all in the range 143000-194000. The estimates of profile drag coefficient were reconcilable with previous estimates of the wing profile drag of the same bird, obtained by the subtractive method, and also with values predicted by the `Airfoil-ii? program for designing aerofoils, based on a digitized wing profile from the ulnar region of the wing. The thickness of the wake suggested that the boundary layer was mostly or fully turbulent in most observations and separated in some, possibly as an active means of creating drag for control purposes. It appears that the bird could momentarily either increase or decrease the profile drag of specific parts of the wing, by active changes of shape, and it appeared to use the carpo-metacarpal region especially for such control movements. Further investigation in a low turbulence wind tunnel would help to resolve doubts about the possible influence of airstream turbulence on the behaviour of the boundary layer.

Journal of Experimental Biology

Biorecovery of gold

Recovery of ionic and metallic gold (Au) from a wide variety of solutions by selected species of bacteria, yeasts, fungi, algae, and higher plants is documented. Gold accumulations were up to 7.0 g/kg dry weight (DW) in various species of bacteria, 25.0 g/kg DW in freshwater algae, 84.0 g/kg DW in peat, and 100.0 g/kg DW in dried fungus mixed with keratinous material. Mechanisms of accumulation include oxidation, dissolution, reduction, leaching, and sorption. Uptake patterns are significantly modified by the physicochemical milieu. Crab exoskeletons accumulate up to 4.9 g Au/kg DW; however, gold accumulations in various tissues of living teleosts, decapod crustaceans, and bivalve molluscs are negligible.

Indian Journal of Experimental Biology

The colonial ascidian Didemnum sp. A: current distribution, basic biology and potential threat to marine communities of the northeast and west coasts of North America

Didemnum sp. A is a colonial ascidian with rapidly expanding populations on the east and west coasts of North America. The origin of Didemum sp. A is unknown. Populations were first observed on the northeast coast of the U.S. in the late 1980s and on the west coast during the 1990s. It is currently undergoing a massive population explosion and is now a dominant member of many subtidal communities on both coasts. To determine Didemnum sp. A's current distribution, we conducted surveys from Maine to Virginia on the east coast and from British Columbia to southern California on the west coast of the U.S. between 1998 and 2005. In nearshore locations Didemnum sp. A currently ranges from Eastport, Maine to Shinnecock Bay, New York on the east coast. On the west coast it has been recorded from Humboldt Bay to Port San Luis in California, several sites in Puget Sound, Washington, including a heavily fouled mussel culture facility, and several sites in southwestern British Columbia on and adjacent to oyster and mussel farms. The species also occurs at deeper subtidal sites (up to 81 m) off New England, including Georges, Stellwagen and Tillies Banks. On Georges Bank numerous sites within a 230 km2 area are 50–90% covered by Didemnum sp. A; large colonies cement the pebble gravel into nearly solid mats that may smother infaunal organisms. These observations suggest that Didemnum sp. A has the potential to alter marine communities and affect economically important activities such as fishing and aquaculture.

British Columbia, California, Connecticut, Maine,

Larval settlement into marine soft-sediment systems: Interactions with the meiofauna

The distribution and abundance of organisms exhibiting complex life cycles, like marine invertebrates, reflect ecological processes operating at all stages in the life history. Few studies have yet considered the mortality patterns of larvae and juveniles. To test whether the permanent meiofauna play a role in macrofaunal larval site-selection, and/or if permanent meiofaunal predators or competitors and disturbers are a significant source of mortality for newly settled macrofaunal juveniles, a series of manipulations of meiofauna were performed. Densities of (1) predatory turbellarians alone, and (2) other meiofauna, were increased in small boxes placed on a holding table in a shallow subtidal habitat. Natural settlement into meiofauna manipulations was examined approximately monthly for a year. Other experiments were conducted to examine the relative importance of site-selection and post-settlement mortality and to determine if settlement was density-dependent or density-independent. Densities of spionid, cirratulid, terebellid, capitellid, and maldanid polychaetes, oligochaetes, bivalves, and amphipods were all reduced by high densities of turbellarians, and sometimes by other meiofauna. Turbellarian predation was responsible for significant post-settlement mortality, but spionids, cirratulids, terebellids, and maldanids also appeared to avoid dense turbellarian treatments. Bivalve densities were reduced by high densities of other meiofauna, probably due to their sedimentary disturbance. Syllid polychaete densities were higher in turbellarian treatments, suggesting that they select for areas with high densities of their prey (small turbellarians). The effects of meiofauna on settlement and survival were greatest in spring and summer. The results of these experiments suggest larval settlement patterns and early juvenile survival should be considered in developing general models of life history and community structure for soft-sediment systems.

Journal of Experimental Marine Biology and Ecology

Effects of kelp forest removal on associated fish assemblages in central California

Visual surveys along subtidal belt transects were used to compare fish assemblages on an experimental and a control site before and after the removal of a canopy-forming kelp forest. The giant kelp Macrocystis pyrifera (L.) C.A. Agardh was removed at the holdfast from ≈ 1 ha of high relief structurally complex rock substratum. The abundance of seven species of fish, of which five were considered midwater species, significantly declined after the kelp was removed. No increases in abundance were noted within the experimental site. On the control site, two declines in abundance were observed, while two species increased in abundance. These results indicate that the presence of a giant kelp forest may increase the abundance and species diversity of the fish assemblages over a high relief rocky reef in central California, U.S.A.

Journal of Experimental Marine Biology and Ecology

Food availability controls seasonal cycle of growth in Macoma balthica (L.) in San Francisco Bay, California

A 2-yr field study of growth in the bivalve Macoma balthica (L.) at four locations in San Francisco Bay, California, U.S., showed that the timing and rate of growth (increase in shell length) were related to food supply. This clam feeds on both planktonic and benthic microalgae, depending on availability. Growth was apparently food-limited during some months, during one year more than the other, and at some locations more than others. Tissue-weight changes were also related to food availability. The heaviest animals were found in that year and at those locations with the highest chl a concentrations. Tissue-weight gains usually coincided with increased shell-growth rate or with reproductive development, although some large animals showed weight gain independent of both of these factors during periods with mid-range chl a levels. Weight losses coincided with spawning or periods of low growth rate, except at one station where, during a period when most animals were reproductively ripe, food concentrations were high, and shell growth was rapid, animals lost weight. This study failed to show a relation between salinity and the timing or rate of change of either shell length or tissue weight. The mild temperatures (10-23 ??C water temperature) of the area studied resulted in no growth inhibition due to low temperature, but there was some indication that the high air temperatures found in these intertidal areas limited growth rates. ?? 1988.

Journal of Experimental Marine Biology and Ecology

Effects of experimental overgrowth on survival and change in the turf assemblage of a giant kelp forest

Crustose coralline algae were the prevalent cover among sessile organisms that paved or grew near the substratum, and also the most commonly overgrown species in a giant kelp Macrocystis pyrifera (L.) C.A. Agardh forest located off San Nicolas Island, California. Giant kelp was the largest and most conspicuous species that overgrew large patches of the substrata; overgrowth among turf organisms also appeared common. To determine the effects of giant kelp holdfasts on crustose coralline algae and other turf organisms, “artificial holdfasts” were placed on 0.125-m 2 plots for 5, 8 and 12 months. In these treatments, 50–57% of the crustose coralline algae survived. Because these algae also recruited while covered, the total cover (survivorship plus recruitment) differed by only 7–26% from that sampled at the start of the study. The decline of these algae in control plots was similar to that in the treatment plots mostly because of overgrowth by sessile invertebrates. Bryozoans increased markedly on the control plots, whereas 0–12% survived in the treatment plots. Bryozoans and sponges also recruited under the artificial holdfasts. Some arborescent turf algae survived in the 5- and 8-month treatments; articulated coralline algae survived better than did foliose algae. High survival recruitment of crustose coralline algae while overgrown contributed to their prevalence in benthic communities.

Journal of Experimental Marine Biology and Ecology

Can otolith microchemistry chart patterns of migration and habitat utilization in anadromous fishes?

Seasonal and ontogenetic patterns in estuarine and coastal migrations of anadromous fish species have important consequences to their survival, growth, recruitment, and reproduction. We tested the hypothesis that otolith (sagitta) microchemistry can document the environmental history of individual fish across an estuarine salinity gradient. Juvenile striped bass, Morone saxatilis (Walbaum), (80 days posthatch) were reared for 3 wk in aquaria at two temperatures and six salinities. The ratio of strontium/calcium ( Sr Ca "> SrCa ) deposited in the sagittal otoliths of reared juveniles was positively related to salinity. Temperature and growth rate had relatively minor, but significant effects on the Sr Ca "> SrCa ratio. In a second experiment, juveniles (80 days posthatch) were exposed to increasing salinity (0 ppt to 25 ppt) and then decreasing salinity (25 ppt to 0 ppt) over a 20-wk period. Electron microprobe examination of the otoliths from these juveniles showed a gradual rise and decline in Sr Ca "> SrCa during the experimental period which corresponded directly with experimental changes in salinity. Field data on subadult and adult striped bass corroborated the laboratory analyses and indicated a logistic relationship between ambient salinity and otolith Sr Ca "> SrCa ratio. Verification studies support the use of otolith microchemistry to measure migratory schedules and habitat utilization patterns in anadromous striped bass populations.

Journal of Experimental Marine Biology and Ecology

Resilience of predators to fishing pressure on coral patch reefs

Numbers and biomass of piscivorous fish and their predation on other fish may often be high in undisturbed coral reef communities. The effects of such predation have sometimes been studied by removal of piscivores (either experimentally or by fishermen). Such perturbations have usually involved removal of large, highly vulnerable, mobile piscivores that are often actively sought in fisheries. The effects of fishing on smaller, demersal, semi-resident piscivores have been little studied. We studied such effects on the fish communities of patch reefs at Midway atoll by experimentally removing major resident, demersal, piscivorous fishes. First, four control reefs and four experimental reefs were selected, their dimensions and habitats mapped, and their visible fish communities censused repeatedly over 1 year. Census of all control and experimental reefs was continued for the following 39 months, during which known piscivores were collected repeatedly by hand spearing. Records were kept of catch and effort to calculate CPUE as an index of predator density. Spearfishing on the experimental reefs removed 2504 piscivorous fish from 12 families and 43 taxa (mostly species). The species richness of the catch did not show an overall change over the duration of the experiment. Spearman rank correlation analysis showed some unexpected positive correlations for density in numbers and biomass of major fished piscivorous groups (especially lizardfish) over the experiment. Only two relatively minor fished piscivorous taxa declined in abundance over the experiment, while the overall abundance of piscivores increased. Visual censuses of fish on the experimental reefs also failed to show reduction of total piscivores over the full experimental period. No significant trend in the abundance of lizardfish censused over the full period was apparent on any of the control reefs. The high resilience of piscivores on these experimental reefs to relatively intense fishing pressure could result from their protracted recruitment seasons, high immigration rates, cryptic habits, or naturally high abundances. A major factor was the high immigration rates of lizardfish, replacing lizardfish and other less mobile piscivores removed from the reefs by spearing. On the fished reefs, the removed lizardfish population replaced itself >20 times during the experiment; other piscivorous taxa replaced themselves only 5 times.

Journal of Experimental Marine Biology and Ecology

Biomechanical factors contributing to self-organization in seagrass landscapes

Field observations have revealed that when water flow is consistently from one direction, seagrass shoots align in rows perpendicular to the primary axis of flow direction. In this study, live Zostera marina shoots were arranged either randomly or in rows perpendicular to the flow direction and tested in a seawater flume under unidirectional flow and waves to determine if shoot arrangement: a) influenced flow-induced force on individual shoots, b) differentially altered water flow through the canopy, and c) influenced light interception by the canopy. In addition, blade breaking strength was compared with flow-induced force to determine if changes in shoot arrangement might reduce the potential for damage to shoots. Under unidirectional flow, both current velocity in the canopy and force on shoots were significantly decreased when shoots were arranged in rows as compared to randomly. However, force on shoots was nearly constant with downstream distance, arising from the trade-off of shoot bending and in-canopy flow reduction. The coefficient of drag was higher for randomly-arranged shoots at low velocities (< 30 cm s- 1) but converged rapidly among the two shoot arrangements at higher velocities. Shoots arranged in rows tended to intercept slightly more light than those arranged randomly. Effects of shoot arrangement under waves were less clear, potentially because we did not achieve the proper plant size?row spacing ratio. At this point, we may only suggest that water motion, as opposed to light capture, is the dominant physical mechanism responsible for these shoot arrangements. Following a computation of the Environmental Stress Factor, we concluded that even photosynthetically active blades may be damaged or broken under frequently encountered storm conditions, irrespective of shoot arrangement. We hypothesize that when flow is generally from one direction, seagrass bed patterns over multiple scales of consideration may arise as a cumulative effect of individual shoot self-organization driven by reduced force and drag on the shoots and somewhat improved light capture.

Journal of Experimental Marine Biology and Ecology

Ecological observations on the colonial ascidian Didemnum sp. in a New England tide pool habitat

The colonial ascidian Didemnum sp. has colonized northwestern Atlantic coastal habitats from southern Long Island, New York, to Eastport, Maine. It is also present in offshore habitats of the Georges Bank fishing grounds. It threatens to alter fisheries habitats and shellfish aquacultures. Observations in a tide pool at Sandwich, MA from December 2003 to February 2006 show that Didemnum sp. tolerates water temperatures ranging from ≤ 1 to > 24 °C, with daily changes of up to 11 °C. It attaches to pebbles, cobbles, and boulders, and it overgrows other tunicates, seaweeds, sponges, and bivalves. From May to mid July, colonies appear as small patches on the bottoms of rocks. Colonies grow rapidly from July to September, with some growth into December, and they range in color from pink to pale yellow to pale orange. Colony health declines from October through April, presumably in response to changes in water temperatures, and this degenerative process is manifested by color changes, by the appearance of small dark brown spots that represent clumps of fecal pellets in the colony, by scavenging by periwinkles, and by a peeling-away of colonies from the sides of cobbles and boulders. At Sandwich, colonies died that were exposed to air at low tide. The species does not exhibit this seasonal cycle of growth and decline in subtidal habitats (40–65 m) on the Georges Bank fishing grounds where the daily climate is relatively stable and annual water temperatures range from 4 to 15 °C. Experiments in the tide pool with small colony fragments (5 to 9 cm 2 ) show they re-attach and grow rapidly by asexual budding, increasing in size 6- to 11-fold in the first 15 days. Didemnum sp. at Sandwich has no known predators except for common periwinkles ( Littorina littorea ) that graze on degenerating colonies in the October to April time period and whenever colonies are stressed by desiccation. The tendencies of the ascidian (1) to attach to firm substrates, (2) to rapidly overgrow other species, (3) to tolerate a wide temperature range, (4) to be free from predation, and (5) to spread by colony fragmentation combine to make it a potential threat to benthic marine habitats and aquacultures. Didemnum sp. is known to overgrow mussels, oysters, and sea scallops, and it likely envelops other bivalves too.

Massachusetts

The occurrence of the colonial ascidian Didemnum sp. on Georges Bank gravel habitat: ecological observations and potential effects on groundfish and scallop fisheries

The colonial ascidian Didemnum sp. is present on the Georges Bank fishing grounds in a gravel habitat where the benthic invertebrate fauna has been monitored annually since 1994. The species was not noted before 2002 when large colonies were first observed; and by 2003 and 2004 it covered large areas of the seabed at some locations. The latest survey in 2005 documented the tunicate's presence in two gravel areas that total more than 67 nm 2 (230 km 2 ). The affected area is located on the Northern Edge of the bank in United States waters near the U.S./Canada boundary ( Fig. 1). This is the first documented offshore occurrence of a species that has colonized eastern U.S. coastal waters from New York to Maine during the past 15–20 years ( U.S. Geological Survey, 2006). Video imagery shows colonies coalescing to form large mats that cover more than 50% of the seabed along some video/photo transects. The affected area is an immobile pebble and cobble pavement that lies at water depths of 40 to 65 m where strong semidiurnal tidal currents reach speeds of 1 to 2 kt (50–100 cm/s). The water column is mixed year round, ensuring a constant supply of nutrients to the seabed. Annual temperatures range from 4 to 15 °C ( Mountain and Holzwarth, 1989). The gravel areas are bounded by sand ridges whose mobile surfaces are moved daily by the strong tidal currents. Studies commenced here in 1994 to characterize the gravel habitat and to document the effects of fishing disturbance on it ( Collie et al., 2005).

Massachusetts

Effects of intermittent flow and irradiance level on back reef Porites corals at elevated seawater temperatures

Corals inhabiting shallow back reef habitats are often simultaneously exposed to elevated seawater temperatures and high irradiance levels, conditions known to cause coral bleaching. Water flow in many tropical back reef systems is tidally influenced, resulting in semi-diurnal or diurnal flow patterns. Controlled experiments were conducted to test effects of semi-diurnally intermittent water flow on photoinhibition and bleaching of the corals Porites lobata and P. cylindrica kept at elevated seawater temperatures and different irradiance levels. All coral colonies were collected from a shallow back reef pool on Ofu Island, American Samoa. In the high irradiance experiments, photoinhibition and bleaching were less for both species in the intermittent high-low flow treatment than in the constant low flow treatment. In the low irradiance experiments, there were no differences in photoinhibition or bleaching for either species between the flow treatments, despite continuously elevated seawater temperatures. These results suggest that intermittent flow associated with semi-diurnal tides, and low irradiances caused by turbidity or shading, may reduce photoinhibition and bleaching of back reef corals during warming events. ?? 2006 Elsevier B.V. All rights reserved.

Journal of Experimental Marine Biology and Ecology