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ArticleResponse to ‘On the importance of understanding physiology when estimating energetics in cetaceans’(Company of Biologists, 2017-02-15) Fahlman, Andreas ; van der Hoop, Julie ; Moore, Michael J. ; Levine, Gregg ; Rocho-Levine, Julie ; Brodsky, MicahWe are grateful for the interest in our paper by two eminent physiologists and hope this response to their comments will clarify the objectives of our paper. The analysis in Fahlman et al. (2016) was not intended to provide an accurate method to estimate field metabolic rate (FMR) in large mysticetes; the objective was to measure the dynamic changes in physiology associated with recovery from exercise and show that they are important to consider when estimating FMR. While static averages can provide useful estimates of FMR for a variety of situations, these need to be appropriately selected. For example, we illustrate that it is not possible to use selected average values chosen from excised tissues or resting animals (as in Blix and Folkow, 1995) to provide meaningful estimates of FMR for animals at different activities (i.e. the dolphins in our study). Our study highlights the importance of temporal variation in physiological models: the Blix and Folkow (1995) estimates rely on the assumption that only breathing frequency (fR) changes with activity, while we argue that both the tidal volume (VT) and mixed lung O2 content also vary with activity and recovery from a dive (Ridgway et al., 1969). Including this variation in all three parameters reduces temporal uncertainty in the same conceptual model (see Eqn. 1 in Fahlman et al., 2016).
ArticleEstimating energetics in cetaceans from respiratory frequency : why we need to understand physiology(The Company of Biologists, 2016-04-15) Fahlman, Andreas ; van der Hoop, Julie ; Moore, Michael J. ; Levine, Gregg ; Rocho-Levine, Julie ; Brodsky, MicahThe accurate estimation of field metabolic rates (FMR) in wild animals is a key component of bioenergetic models, and is important for understanding the routine limitations for survival as well as individual responses to disturbances or environmental changes. Several methods have been used to estimate FMR, including accelerometer-derived activity budgets, isotope dilution techniques, and proxies from heart rate. Counting the number of breaths is another method used to assess FMR in cetaceans, which is attractive in its simplicity and the ability to measure respiration frequency from visual cues or data loggers. This method hinges on the assumption that over time a constant tidal volume (VT) and O2 exchange fraction (ΔO2) can be used to predict FMR. To test whether this method of estimating FMR is valid, we measured breath-by-breath tidal volumes and expired O2 levels of bottlenose dolphins, and computed the O2 consumption rate (V̇O2) before and after a pre-determined duration of exercise. The measured V̇O2 was compared with three methods to estimate FMR. Each method to estimate V̇O2 included variable VT and/or ΔO2. Two assumption-based methods overestimated V̇O2 by 216-501%. Once the temporal changes in cardio-respiratory physiology, such as variation in VT and ΔO2, were taken into account, pre-exercise resting V̇O2 was predicted to within 2%, and post-exercise V̇O2 was overestimated by 12%. Our data show that a better understanding of cardiorespiratory physiology significantly improves the ability to estimate metabolic rate from respiratory frequency, and further emphasizes the importance of eco-physiology for conservation management efforts.
ArticleTag-based estimates of bottlenose dolphin swimming behavior and energetics(The Company of Biologists, 2022-11-30) Gabaldon, Joaquin T ; Zhang, Ding ; Rocho-Levine, Julie ; Moore, Michael J ; van der Hoop, Julie ; Barton, Kira ; Shorter, K AlexCurrent estimates of marine mammal hydrodynamic forces tend to be made using camera-based kinematic data for a limited number of fluke strokes during a prescribed swimming task. In contrast, biologging tag data yield kinematic measurements from thousands of strokes, enabling new insights into swimming behavior and mechanics. However, there have been limited tag-based estimates of mechanical work and power. In this work, we investigated bottlenose dolphin (Tursiops truncatus) swimming behavior using tag-measured kinematics and a hydrodynamic model to estimate propulsive power, work and cost of transport. Movement data were collected from six animals during prescribed straight-line swimming trials to investigate swimming mechanics over a range of sustained speeds (1.9-6.1 m s-1). Propulsive power ranged from 66 W to 3.8 kW over 282 total trials. During the lap trials, the dolphins swam at depths that mitigated wave drag, reducing overall drag throughout these mid- to high-speed tasks. Data were also collected from four individuals during undirected daytime (08:30-18:00 h) swimming to examine how self-selected movement strategies are used to modulate energetic efficiency and effort. Overall, self-selected swimming speeds (individual means ranging from 1.0 to 1.96 m s-1) tended to minimize cost of transport, and were on the lower range of animal-preferred speeds reported in literature. The results indicate that these dolphins moderate propulsive effort and efficiency through a combination of speed and depth regulation. This work provides new insights into dolphin swimming behavior in both prescribed tasks and self-selected swimming, and presents a path forward for continuous estimates of mechanical work and power from wild animals.
PreprintBottlenose dolphins modify behavior to reduce metabolic effect of tag attachment( 2014-10) van der Hoop, Julie ; Fahlman, Andreas ; Hurst, Thomas P. ; Rocho-Levine, Julie ; Shorter, K. Alex ; Petrov, Victor ; Moore, Michael J.Attaching bio-telemetry or -logging devices (‘tags’) to marine animals for research and monitoring adds drag to streamlined bodies, thus affecting posture, swimming gaits and energy balance. These costs have never been measured in free-swimming cetaceans. To examine the effect of drag from a tag on metabolic rate, cost of transport and swimming behavior, four captive male dolphins (Tursiops truncatus) were trained to swim a set course, either non-tagged (n=7) or fitted with a tag (DTAG2; n=12), and surface exclusively in a flow-through respirometer in which oxygen consumption (Graphic) and carbon dioxide production (Graphic; ml kg−1 min−1) rates were measured and respiratory exchange ratio (Graphic/Graphic) was calculated. Tags did not significantly affect individual mass-specific oxygen consumption, physical activity ratios (exercise Graphic/resting Graphic), total or net cost of transport (COT; J m−1 kg−1) or locomotor costs during swimming or two-minute recovery phases. However, individuals swam significantly slower when tagged (by ~11%; mean ± s.d., 3.31±0.35 m s−1) than when non-tagged (3.73±0.41 m s−1). A combined theoretical and computational fluid dynamics model estimating drag forces and power exertion during swimming suggests that drag loading and energy consumption are reduced at lower swimming speeds. Bottlenose dolphins in the specific swimming task in this experiment slowed to the point where the tag yielded no increases in drag or power, while showing no difference in metabolic parameters when instrumented with a DTAG2. These results, and our observations, suggest that animals modify their behavior to maintain metabolic output and energy expenditure when faced with tag-induced drag.
ArticleSwimming energy economy in bottlenose dolphins under variable drag loading(Frontiers Media, 2018-12-11) van der Hoop, Julie ; Fahlman, Andreas ; Shorter, K. Alex ; Gabaldon, Joaquin ; Rocho-Levine, Julie ; Petrov, Victor ; Moore, Michael J.Instrumenting animals with tags contributes additional resistive forces (weight, buoyancy, lift, and drag) that may result in increased energetic costs; however, additional metabolic expense can be moderated by adjusting behavior to maintain power output. We sought to increase hydrodynamic drag for near-surface swimming bottlenose dolphins, to investigate the metabolic effect of instrumentation. In this experiment, we investigate whether (1) metabolic rate increases systematically with hydrodynamic drag loading from tags of different sizes or (2) whether tagged individuals modulate speed, swimming distance, and/or fluking motions under increased drag loading. We detected no significant difference in oxygen consumption rates when four male dolphins performed a repeated swimming task, but measured swimming speeds that were 34% (>1 m s-1) slower in the highest drag condition. To further investigate this observed response, we incrementally decreased and then increased drag in six loading conditions. When drag was reduced, dolphins increased swimming speed (+1.4 m s-1; +45%) and fluking frequency (+0.28 Hz; +16%). As drag was increased, swimming speed (-0.96 m s-1; -23%) and fluking frequency (-14 Hz; 7%) decreased again. Results from computational fluid dynamics simulations indicate that the experimentally observed changes in swimming speed would have maintained the level of external drag forces experienced by the animals. Together, these results indicate that dolphins may adjust swimming speed to modulate the drag force opposing their motion during swimming, adapting their behavior to maintain a level of energy economy during locomotion.