Jacob L. Johansen

Finding the best estimates of metabolic rates in a coral reef fish

SUMMARY Metabolic rates of aquatic organisms are estimated from measurements of oxygen consumption rates () through swimming and resting respirometry. These distinct approaches are increasingly used in ecophysiology and conservation physiology studies; however, few studies have tested whether they yield comparable results. We examined whether two fundamental measures, standard metabolic rate (SMR) and maximum metabolic rate (MMR), vary based on the method employed. Ten bridled monocle bream (Scolopsis bilineata) were exercised using (1) a critical swimming speed (Ucrit) protocol, (2) a 15 min exhaustive chase protocol and (3) a 3 min exhaustive chase protocol followed by brief (1 min) air exposure. Protocol 1 was performed in a swimming respirometer whereas protocols 2 and 3 were followed by resting respirometry. SMR estimates in swimming respirometry were similar to those in resting respirometry when a three-parameter exponential or power function was used to extrapolate the swimming speed– relationship to zero swimming speed. In contrast, MMR using the Ucrit protocol was 36% higher than MMR derived from the 15 min chase protocol and 23% higher than MMR using the 3 min chase/1 min air exposure protocol. For strong steady (endurance) swimmers, such as S. bilineata, swimming respirometry can produce more accurate MMR estimates than exhaustive chase protocols because oxygen consumption is measured during exertion. However, when swimming respirometry is impractical, exhaustive chase protocols should be supplemented with brief air exposure to improve measurement accuracy. Caution is warranted when comparing MMR estimates obtained with different respirometry methods unless they are cross-validated on a species-specific basis.

Increasing ocean temperatures reduce activity patterns of a large commercially important coral reef fish

Large-bodied fish are critical for sustaining coral reef fisheries, but little is known about the vulnerability of these fish to global warming. This study examined the effects of elevated temperatures on the movement and activity patterns of the common coral trout Plectropomus leopardus (Serranidae), which is an important fishery species in tropical Australia and throughout the Indo West-Pacific. Adult fish were collected from two locations on Australias Great Barrier Reef (23°S and 14°S) and maintained at one of four temperatures (24, 27, 30, 33 °C). Following >4 weeks acclimation, the spontaneous swimming speeds and activity patterns of individuals were recorded over a period of 12 days. At 24-27 °C, spontaneous swimming speeds of common coral trout were 0.43-0.45 body lengths per second (bls(-1)), but dropped sharply to 0.29 bls(-1) at 30 °C and 0.25 bls(-1) at 33 °C. Concurrently, individuals spent 9.3-10.6% of their time resting motionless on the bottom at 24-27 °C, but this behaviour increased to 14.0% at 30 °C and 20.0% of the time at 33 °C (mean ± SE). The impact of temperature was greatest for smaller individuals (<45 cm TL), showing significant changes to swimming speeds across every temperature tested, while medium (45-55 cm TL) and large individuals (>55 cm TL) were first affected by 30 °C and 33 °C, respectively. Importantly, there was some indication that populations can adapt to elevated temperature if presented with adequate time, as the high-latitude population decreased significantly in swimming speeds at both 30 °C and 33 °C, while the low-latitude population only showed significant reductions at 33 °C. Given that movement and activity patterns of large mobile species are directly related to prey encounter rates, ability to capture prey and avoid predators, any reductions in activity patterns are likely to reduce overall foraging and energy intake, limit the energy available for growth and reproduction, and affect the fitness and survival of individuals and populations.

Living in a risky world: the onset and ontogeny of an integrated antipredator phenotype in a coral reef fish

Prey individuals with complex life-histories often cannot predict the type of risk environment to which they will be exposed at each of their life stages. Because the level of investment in defences should match local risk conditions, we predict that these individuals should have the ability to modulate the expression of an integrated defensive phenotype, but this switch in expression should occur at key life-history transitions. We manipulated background level of risk in juvenile damselfish for four days following settlement (a key life-history transition) or 10 days post-settlement, and measured a suite of physiological and behavioural variables over 2 weeks. We found that settlement-stage fish exposed to high-risk conditions displayed behavioural and physiological alterations consistent with high-risk phenotypes, which gave them a survival advantage when exposed to predators. These changes were maintained for at least 2 weeks. The same exposure in post-settlement fish failed to elicit a change in some traits, while the expression of other traits disappeared within a week. Our results are consistent with those expected from phenotypic resonance. Expression of antipredator traits may be masked if individuals are not exposed to certain conditions at key ontogenetic stages.

Large predatory coral trout species unlikely to meet increasing energetic demands in a warming ocean.

Increased ocean temperature due to climate change is raising metabolic demands and energy requirements of marine ectotherms. If productivity of marine systems and fisheries are to persist, individual species must compensate for this demand through increasing energy acquisition or decreasing energy expenditure. Here we reveal that the most important coral reef fishery species in the Indo-west Pacific, the large predatory coral trout Plectropomus leopardus (Serranidae), can behaviourally adjust food intake to maintain body-condition under elevated temperatures, and acclimate over time to consume larger meals. However, these increased energetic demands are unlikely to be met by adequate production at lower trophic levels, as smaller prey species are often the first to decline in response to climate-induced loss of live coral and structural complexity. Consequently, ubiquitous increases in energy consumption due to climate change will increase top-down competition for a dwindling biomass of prey, potentially distorting entire food webs and associated fisheries.

Sediment‐induced turbidity impairs foraging performance and prey choice of planktivorous coral reef fishes

Sedimentation is a substantial threat to aquatic ecosystems and a primary cause of habitat degradation on near-shore coral reefs. Although numerous studies have demonstrated major impacts of sedimentation and turbidity on corals, virtually nothing is known of the sensitivity of reef fishes. Planktivorous fishes are an important trophic group that funnels pelagic energy sources into reef ecosystems. These fishes are visual predators whose foraging is likely to be impaired by turbidity, but the threshold for such effects and their magnitude are unknown. This study examined the effect of sediment-induced turbidity on foraging in four species of planktivorous damselfishes (Pomacentridae) of the Great Barrier Reef, including inshore and offshore species that potentially differ in tolerance for turbidity. An experimental flow tunnel was used to quantify their ability to catch mobile and immobile planktonic prey under different levels of turbidity and velocity in the range encountered on natural and disturbed reefs. Turbidity of just 4 NTU (nephelometric turbidity units) reduced average attack success by up to 56%, with higher effect sizes for species with offshore distributions. Only the inshore species (Neopomacentrus bankieri), which frequently encounters this turbidity on coastal reefs, could maintain high prey capture success. At elevated turbidity similar to that found on disturbed reefs (8 NTU), attack success was reduced in all species examined by up to 69%. These reductions in attack success led to a 21-24% decrease in foraging rates for all mid to outer-shelf species, in spite of increasing attack rates at high turbidity. Although effects of turbidity varied among species, it always depended heavily on prey mobility and ambient velocity. Attack success was up to 14 times lower on mobile prey, leaving species relatively incapable of foraging on anything but immobile prey at high turbidity. Effects of turbidity were particularly prominent at higher velocities, as attack success was overall 20-fold lower and foraging rates 3.3-fold lower at flow velocities > 30 cm/s relative to < or = 10 cm/s. Given that many planktivorous reef fishes predominantly occupy exposed, high-flow habitats, these results provide a reasonable explanation for the lack of planktivores on inshore coral reefs and warn that the performance of visual predators could be impaired at turbidity levels of only 4 NTU.

Unsteady flow affects swimming energetics in a labriform fish (Cymatogaster aggregata)

Unsteady water flows are common in nature, yet the swimming performance of fishes is typically evaluated at constant, steady speeds in the laboratory. We examined how cyclic changes in water flow velocity affect the swimming performance and energetics of a labriform swimmer, the shiner surfperch, Cymatogaster aggregata, during station holding. Using intermittent-flow respirometry, we measured critical swimming speed (Ucrit), oxygen consumption rates (ṀO2) and pectoral fin use in steady flow versus unsteady flows with either low- [0.5 body lengths (BL) s−1] or high-amplitude (1.0 BL s−1) velocity fluctuations, with a 5 s period. Individuals in low-amplitude unsteady flow performed as well as fish in steady flow. However, swimming costs in high-amplitude unsteady flow were on average 25.3% higher than in steady flow and 14.2% higher than estimated values obtained from simulations based on the non-linear relationship between swimming speed and oxygen consumption rate in steady flow. Time-averaged pectoral fin use (fin-beat frequency measured over 300 s) was similar among treatments. However, measures of instantaneous fin use (fin-beat period) and body movement in high-amplitude unsteady flow indicate that individuals with greater variation in the duration of their fin beats were better at holding station and consumed less oxygen than fish with low variation in fin-beat period. These results suggest that the costs of swimming in unsteady flows are context dependent in labriform swimmers, and may be influenced by individual differences in the ability of fishes to adjust their fin beats to the flow environment.

Energetic extremes in aquatic locomotion by coral reef fishes

Underwater locomotion is challenging due to the high friction and resistance imposed on a body moving through water and energy lost in the wake during undulatory propulsion. While aquatic organisms have evolved streamlined shapes to overcome such resistance, underwater locomotion has long been considered a costly exercise. Recent evidence for a range of swimming vertebrates, however, has suggested that flapping paired appendages around a rigid body may be an extremely efficient means of aquatic locomotion. Using intermittent flow-through respirometry, we found exceptional energetic performance in the Bluelined wrasse Stethojulis bandanensis, which maintains tuna-like optimum cruising speeds (up to 1 metre s−1) while using 40% less energy than expected for their body size. Displaying an exceptional aerobic scope (22-fold above resting), streamlined rigid-body posture, and wing-like fins that generate lift-based thrust, S. bandanensis literally flies underwater to efficiently maintain high optimum swimming speeds. Extreme energetic performance may be key to the colonization of highly variable environments, such as the wave-swept habitats where S. bandanensis and other wing-finned species tend to occur. Challenging preconceived notions of how best to power aquatic locomotion, biomimicry of such lift-based fin movements could yield dramatic reductions in the power needed to propel underwater vehicles at high speed.

Quantifying water flow within aquatic ecosystems using load cell sensors: a profile of currents experienced by coral reef organisms around Lizard Island, Great Barrier Reef, Australia.

Current velocity in aquatic environments has major implications for the diversity, abundance and ecology of aquatic organisms, but quantifying these currents has proven difficult. This study utilises a simple and inexpensive instrument (<

Methods matter: considering locomotory mode and respirometry technique when estimating metabolic rates of fishes

150) to provide a detailed current velocity profile of the coral-reef system around Lizard Island (Great Barrier Reef, Australia) at a spatial and temporal scale relevant to the ecology of individual benthos and fish. The instrument uses load-cell sensors to provide a correlation between sensor output and ambient current velocity of 99%. Each instrument is able to continuously record current velocities to >500 cms−1 and wave frequency to >100 Hz over several weeks. Sensor data are registered and processed at 16 MHz and 10 bit resolution, with a measuring precision of 0.06±0.04%, and accuracy of 0.51±0.65% (mean ±S.D.). Each instrument is also pressure rated to 120 m and shear stresses ≤20 kNm−2 allowing deployment in harsh environments. The instrument was deployed across 27 coral reef sites covering the crest (3 m), mid-slope (6 m) and deep-slope (9 m depth) of habitats directly exposed, oblique or sheltered from prevailing winds. Measurements demonstrate that currents over the reef slope and crest varies immensely depending on depth and exposure: Currents differ up to 9-fold within habitats only separated by 3 m depth and 15-fold between exposed, oblique and sheltered habitats. Comparisons to ambient weather conditions reveal that currents around Lizard Island are largely wind driven. Zero to 22.5 knot winds correspond directly to currents of 0 to >82 cms−1, while tidal currents rarely exceed 5.5 cms−1. Rather, current velocity increases exponentially as a function of wave height (0 to 1.6 m) and frequency (0.54 to 0.20 Hz), emphasizing the enormous effect of wind and waves on organisms in these shallow coral reef habitats.

Fast-starting after a breath: air-breathing motions are kinematically similar to escape responses in the catfish Hoplosternum littorale

We estimated maximum metabolic rates (MMR) in four steady-swimming coral reef fishes - two body-caudal and two median-paired fin swimmers - using three different techniques. Differences between MMR estimates were not due to locomotory mode, but rather the method used to induce maximum performance, with swimming respirometry resulting in the highest estimates.