ABSTRACT
Although gigantic body size and obligate filter feeding mechanisms have evolved in multiple vertebrate lineages (mammals and fishes), intermittent ram (lunge) filter feeding is unique to a specific family of baleen whales: rorquals. Lunge feeding is a high cost, high benefit feeding mechanism that requires the integration of unsteady locomotion (i.e., accelerations and maneuvers); the impact of scale on the biomechanics and energetics of this foraging mode continues to be the subject of intense study. The goal of our investigation was to use a combination of multi-sensor tags paired with UAS footage to determine the impact of morphometrics such as body size on kinematic lunging parameters such as fluking timing, maximum lunging speed, and deceleration during the engulfment period for a range of species from minke to blue whales. Our results show that, in the case of krill-feeding lunges and regardless of size, animals exhibit a skewed gradient between powered and fully unpowered engulfment, with fluking generally ending at the point of both the maximum lunging speed and mouth opening. In all cases, the small amounts of propulsive thrust generated by the tail were unable to overcome the high drag forces experienced during engulfment. Assuming this thrust to be minimal, we predicted the minimum speed of lunging across scale. To minimize the energetic cost of lunge feeding, hydrodynamic theory predicts slower lunge feeding speeds regardless of body size, with a lower boundary set by the ability of the prey to avoid capture. We used empirical data to test this theory and instead found that maximum foraging speeds remain constant and high (â¼4 m s-1) across body size, even as higher speeds result in lower foraging efficiency. Regardless, we found an increasing relationship between body size and this foraging efficiency, estimated as the ratio of energetic gain from prey to energetic cost. This trend held across timescales ranging from a single lunge to a single day and suggests that larger whales are capturing more prey-and more energy-at a lower cost.
ABSTRACT
We consider in this work a bicriteria scheduling problem on two different parallel machines with a periodic preventive maintenance policy. The two objectives considered involve minimization of job rejection costs and weighted sum of completion times. They are handled through a lexicographic approach, due to a natural hierarchy among the two objectives in the applications considered. The main contributions of this paper are first to present a new problem relevant to practice, second, to develop a mixed-integer-linear-program model for the problem, and third, to introduce two generalizable tabu-search metaheuristics relying on different neighborhood structures and solution spaces. Computational results for 120 instances (generated from a real case) are reported to empirically demonstrate the effectiveness of the proposed metaheuristics.
ABSTRACT
Despite their enormous size, whales make their living as voracious predators. To catch their much smaller, more maneuverable prey, they have developed several unique locomotor strategies that require high energetic input, high mechanical power output and a surprising degree of agility. To better understand how body size affects maneuverability at the largest scale, we used bio-logging data, aerial photogrammetry and a high-throughput approach to quantify the maneuvering performance of seven species of free-swimming baleen whale. We found that as body size increases, absolute maneuvering performance decreases: larger whales use lower accelerations and perform slower pitch-changes, rolls and turns than smaller species. We also found that baleen whales exhibit positive allometry of maneuvering performance: relative to their body size, larger whales use higher accelerations, and perform faster pitch-changes, rolls and certain types of turns than smaller species. However, not all maneuvers were impacted by body size in the same way, and we found that larger whales behaviorally adjust for their decreased agility by using turns that they can perform more effectively. The positive allometry of maneuvering performance suggests that large whales have compensated for their increased body size by evolving more effective control surfaces and by preferentially selecting maneuvers that play to their strengths.
Subject(s)
Motivation , Whales , Animals , Body Size , SwimmingABSTRACT
High efficiency lunate-tail swimming with high-aspect-ratio lifting surfaces has evolved in many vertebrate lineages, from fish to cetaceans. Baleen whales (Mysticeti) are the largest swimming animals that exhibit this locomotor strategy, and present an ideal study system to examine how morphology and the kinematics of swimming scale to the largest body sizes. We used data from whale-borne inertial sensors coupled with morphometric measurements from aerial drones to calculate the hydrodynamic performance of oscillatory swimming in six baleen whale species ranging in body length from 5 to 25â m (fin whale, Balaenoptera physalus; Bryde's whale, Balaenoptera edeni; sei whale, Balaenoptera borealis; Antarctic minke whale, Balaenoptera bonaerensis; humpback whale, Megaptera novaeangliae; and blue whale, Balaenoptera musculus). We found that mass-specific thrust increased with both swimming speed and body size. Froude efficiency, defined as the ratio of useful power output to the rate of energy input ( Sloop, 1978), generally increased with swimming speed but decreased on average with increasing body size. This finding is contrary to previous results in smaller animals, where Froude efficiency increased with body size. Although our empirically parameterized estimates for swimming baleen whale drag were higher than those of a simple gliding model, oscillatory locomotion at this scale exhibits generally high Froude efficiency as in other adept swimmers. Our results quantify the fine-scale kinematics and estimate the hydrodynamics of routine and energetically expensive swimming modes at the largest scale.
Subject(s)
Balaenoptera , Fin Whale , Animals , Antarctic Regions , Biomechanical Phenomena , SwimmingABSTRACT
BACKGROUND: As a result of the COVID-19 pandemic first wave, reductions in ST-elevation myocardial infarction (STEMI) invasive care, ranging from 23% to 76%, have been reported from various countries. Whether this change had any impact on coronary angiography (CA) volume or on mechanical support device use for STEMI and post-STEMI mechanical complications in Canada is unknown. METHODS: We administered a Canada-wide survey to all cardiac catheterization laboratory directors, seeking the volume of CA use for STEMI performed during the period from March 1 2020 to May 31, 2020 (pandemic period), and during 2 control periods (March 1, 2019 to May 31, 2019 and March 1, 2018 to May 31, 2018). The number of left ventricular support devices used, as well as the number of ventricular septal defects and papillary muscle rupture cases diagnosed, was also recorded. We also assessed whether the number of COVID-19 cases recorded in each province was associated with STEMI-related CA volume. RESULTS: A total of 41 of 42 Canadian catheterization laboratories (98%) provided data. There was a modest but statistically significant 16% reduction (incidence rate ratio [IRR] 0.84; 95% confidence interval 0.80-0.87) in CA for STEMI during the first wave of the pandemic, compared to control periods. IRR was not associated with provincial COVID-19 caseload. We observed a 26% reduction (IRR 0.74; 95% confidence interval 0.61-0.89) in the use of intra-aortic balloon pump use for STEMI. Use of an Impella pump and mechanical complications from STEMI were exceedingly rare. CONCLUSIONS: We observed a modest 16% decrease in use of CA for STEMI during the pandemic first wave in Canada, lower than the level reported in other countries. Provincial COVID-19 caseload did not influence this reduction.
INTRODUCTION: Après la première vague de la pandémie de COVID-19, de nombreux pays ont déclaré une réduction de 23 % à 76 % des soins invasifs de l'infarctus du myocarde avec élévation du segment ST (STEMI). On ignore si ce changement a entraîné des répercussions sur le volume d'angiographies coronariennes (AC) ou sur l'utilisation des dispositifs d'assistance mécanique lors de STEMI et des complications mécaniques post-STEMI au Canada. MÉTHODES: Nous avons réalisé un sondage pancanadien auprès de tous les directeurs de laboratoire de cathétérisme cardiaque pour obtenir le volume d'utilisation des AC lors des STEMI réalisées durant la période du 1er mars 2020 au 31 mai 2020 (période de pandémie) et durant 2 périodes témoins (1er mars 2019 au 31 mai 2019 et 1er mars 2018 au 31 mai 2018). Le nombre de dispositifs d'assistance ventriculaire gauche utilisés et le nombre de cas de communications interventriculaires et de ruptures du muscle papillaire diagnostiqués ont également été enregistrés. Nous avons aussi évalué si le nombre de cas de COVID-19 enregistrés dans chaque province était associé au volume d'AC liées aux STEMI. RÉSULTATS: Au total, 41 des 42 laboratoires canadiens de cathétérisme (98 %) ont fourni des données. Lors de la comparaison de la première vague de la pandémie aux périodes témoins, nous avons noté une réduction modeste, mais significative, sur le plan statistique de 16 % (ratio du taux d'incidence [RTI] 0,84; intervalle de confiance à 95 % 0,80-0,87) des AC lors de STEMI. Le RTI n'était pas associé au nombre provincial de cas de COVID-19. Nous avons observé une réduction de 26 % (RTI 0,74; intervalle de confiance à 95 % 0,61-0,89) de l'utilisation de pompes à ballonnet intra-aortique lors de STEMI. L'utilisation d'une pompe Impella et les complications mécaniques après les STEMI étaient extrêmement rares. CONCLUSIONS: Nous avons observé une diminution modeste de 16 % de l'utilisation des AC lors de STEMI durant la première vague de la pandémie au Canada, soit une diminution plus faible que ce que les autres pays ont signalé. Le nombre provincial de cas de COVID-19 n'a pas influencé cette réduction.
ABSTRACT
The considerable power needed for large whales to leap out of the water may represent the single most expensive burst maneuver found in nature. However, the mechanics and energetic costs associated with the breaching behaviors of large whales remain poorly understood. In this study we deployed whale-borne tags to measure the kinematics of breaching to test the hypothesis that these spectacular aerial displays are metabolically expensive. We found that breaching whales use variable underwater trajectories, and that high-emergence breaches are faster and require more energy than predatory lunges. The most expensive breaches approach the upper limits of vertebrate muscle performance, and the energetic cost of breaching is high enough that repeated breaching events may serve as honest signaling of body condition. Furthermore, the confluence of muscle contractile properties, hydrodynamics, and the high speeds required likely impose an upper limit to the body size and effectiveness of breaching whales.
Subject(s)
Body Size , Energy Metabolism/physiology , Whales/anatomy & histology , Whales/physiology , Animals , Biomechanical Phenomena/physiology , Feeding Behavior , Species Specificity , Whales/classificationABSTRACT
The unique engulfment filtration strategy of microphagous rorqual whales has evolved relatively recently (<5 Ma) and exploits extreme predator/prey size ratios to overcome the maneuverability advantages of swarms of small prey, such as krill. Forage fish, in contrast, have been engaged in evolutionary arms races with their predators for more than 100 million years and have performance capabilities that suggest they should easily evade whale-sized predators, yet they are regularly hunted by some species of rorqual whales. To explore this phenomenon, we determined, in a laboratory setting, when individual anchovies initiated escape from virtually approaching whales, then used these results along with in situ humpback whale attack data to model how predator speed and engulfment timing affected capture rates. Anchovies were found to respond to approaching visual looming stimuli at expansion rates that give ample chance to escape from a sea lion-sized predator, but humpback whales could capture as much as 30-60% of a school at once because the increase in their apparent (visual) size does not cross their prey's response threshold until after rapid jaw expansion. Humpback whales are, thus, incentivized to delay engulfment until they are very close to a prey school, even if this results in higher hydrodynamic drag. This potential exaptation of a microphagous filter feeding strategy for fish foraging enables humpback whales to achieve 7× the energetic efficiency (per lunge) of krill foraging, allowing for flexible foraging strategies that may underlie their ecological success in fluctuating oceanic conditions.
Subject(s)
Escape Reaction/physiology , Feeding Behavior , Fishes/physiology , Humpback Whale/physiology , Predatory Behavior/physiology , Animals , Biological Evolution , Euphausiacea/physiology , Filtration , Humpback Whale/anatomy & histology , Hydrodynamics , Jaw/anatomy & histology , Locomotion/physiology , Models, Biological , Organ Size/physiology , Time FactorsABSTRACT
The largest animals are baleen filter feeders that exploit large aggregations of small-bodied plankton. Although this feeding mechanism has evolved multiple times in marine vertebrates, rorqual whales exhibit a distinct lunge filter feeding mode that requires extreme physiological adaptations-most of which remain poorly understood. Here, we review the biomechanics of the lunge feeding mechanism in rorqual whales that underlies their extraordinary foraging performance and gigantic body size.
Subject(s)
Feeding Behavior/physiology , Whales/physiology , Adaptation, Physiological/physiology , Animals , Biomechanical Phenomena/physiology , Energy Metabolism/physiologyABSTRACT
The scale dependence of locomotor factors has long been studied in comparative biomechanics, but remains poorly understood for animals at the upper extremes of body size. Rorqual baleen whales include the largest animals, but we lack basic kinematic data about their movements and behavior below the ocean surface. Here, we combined morphometrics from aerial drone photogrammetry, whale-borne inertial sensing tag data and hydrodynamic modeling to study the locomotion of five rorqual species. We quantified changes in tail oscillatory frequency and cruising speed for individual whales spanning a threefold variation in body length, corresponding to an order of magnitude variation in estimated body mass. Our results showed that oscillatory frequency decreases with body length (âlength-0.53) while cruising speed remains roughly invariant (âlength0.08) at 2â mâ s-1 We compared these measured results for oscillatory frequency against simplified models of an oscillating cantilever beam (âlength-1) and an optimized oscillating Strouhal vortex generator (âlength-1). The difference between our length-scaling exponent and the simplified models suggests that animals are often swimming non-optimally in order to feed or perform other routine behaviors. Cruising speed aligned more closely with an estimate of the optimal speed required to minimize the energetic cost of swimming (âlength0.07). Our results are among the first to elucidate the relationships between both oscillatory frequency and cruising speed and body size for free-swimming animals at the largest scale.
Subject(s)
Swimming/physiology , Whales/physiology , Animals , Regression Analysis , Species Specificity , Whales/anatomy & histologyABSTRACT
Buoyancy control is a fundamental aspect of aquatic life that has major implications for locomotor performance and ecological niche. Unlike terrestrial animals, the densities of aquatic animals are similar to the supporting fluid, thus even small changes in body density may have profound effects on locomotion. Here, we analysed the body composition (lipid versus lean tissue) of 32 shark species to study the evolution of buoyancy. Our comparative phylogenetic analyses indicate that although lean tissue displays minor positive allometry, liver volume exhibits pronounced positive allometry, suggesting that larger sharks evolved bulkier body compositions by adding lipid tissue to lean tissue rather than substituting lean for lipid tissue, particularly in the liver. We revealed a continuum of buoyancy control strategies that ranged from more buoyant sharks with larger livers in deeper ecosystems to relatively denser sharks with small livers in epipelagic habitats. Across this eco-morphological spectrum, our hydrodynamic modelling suggests that neutral buoyancy yields lower drag and more efficient steady swimming, whereas negative buoyancy may be more efficient during accelerated movements. The evolution of buoyancy control in sharks suggests that ecological and physiological factors mediate the selective pressures acting on these traits along two major gradients, body size and habitat depth.
Subject(s)
Biological Evolution , Body Composition , Sharks/anatomy & histology , Sharks/physiology , Swimming , Animal Distribution , Animals , Hydrodynamics , Models, Theoretical , Phylogeny , Species SpecificityABSTRACT
Balaenid whales feed on large aggregates of small and slow-moving prey (predominantly copepods) through a filtration process enabled by baleen. These whales exhibit continuous filtration, namely, with the mouth kept partially opened and the baleen exposed to oncoming prey-laden waters while fluking. The process is an example of crossflow filtration (CFF) in which most of the particulates (prey) are separated from the substrate (water) without ever coming into contact with the filtering surface (baleen). This paper discusses the simulation of baleen filtration hydrodynamics based on a type of hydraulic circuit modeling commonly used in microfluidics, but adapted to the much higher Reynolds number flows typical of whale hydrodynamics. This so-called Baleen Hydraulic Circuit (BHC) model uses as input the basic characteristics of the flows moving through a section of baleen observed in a previous flume study by the authors. The model has low-spatial resolution but incorporates the effects of fluid viscosity, which doubles or more a whale's total body drag in comparison to non-feeding travel. Modeling viscous friction is crucial here since exposing the baleen system to the open ocean ends up tripling a whale's total wetted surface area. Among other findings, the BHC shows how CFF is enhanced by a large filtration surface and hence large body size; how it is carried out via the establishment of rapid anteroposterior flows transporting most of the prey-water slurry towards the oropharyngeal wall; how slower intra-baleen flows manage to transfer most of the substrate out of the mouth, all the while contributing only a fraction to overall oral cavity drag; and how these anteroposterior and intra-baleen flows lose speed as they approach the oropharyngeal wall.
Subject(s)
Balaenoptera/physiology , Feeding Behavior , Hydrodynamics , Mouth/physiology , AnimalsABSTRACT
Maneuverability is one of the most important and least understood aspects of animal locomotion. The hydrofoil-like flippers of cetaceans are thought to function as control surfaces that effect maneuvers, but quantitative tests of this hypothesis have been lacking. Here, we constructed a simple hydrodynamic model to predict the longitudinal-axis roll performance of fin whales, and we tested its predictions against kinematic data recorded by on-board movement sensors from 27 free-swimming fin whales. We found that for a given swimming speed and roll excursion, the roll velocity of fin whales calculated from our field data agrees well with that predicted by our hydrodynamic model. Although fluke and body torsion may further influence performance, our results indicate that lift generated by the flippers is sufficient to drive most of the longitudinal-axis rolls used by fin whales for feeding and maneuvering.
Subject(s)
Animal Structures/physiology , Fin Whale/anatomy & histology , Fin Whale/physiology , Hydrodynamics , Animals , Biomechanical Phenomena , Models, Biological , Swimming/physiologyABSTRACT
Forces due to propulsion should approximate forces due to hydrodynamic drag for animals horizontally swimming at a constant speed with negligible buoyancy forces. Propulsive forces should also correlate with energy expenditures associated with locomotion-an important cost of foraging. As such, biologging tags containing accelerometers are being used to generate proxies for animal energy expenditures despite being unable to distinguish rotational movements from linear movements. However, recent miniaturizations of gyroscopes offer the possibility of resolving this shortcoming and obtaining better estimates of body accelerations of swimming animals. We derived accelerations using gyroscope data for swimming Steller sea lions (Eumetopias jubatus), and determined how well the measured accelerations correlated with actual swimming speeds and with theoretical drag. We also compared dive averaged dynamic body acceleration estimates that incorporate gyroscope data, with the widely used Overall Dynamic Body Acceleration (ODBA) metric, which does not use gyroscope data. Four Steller sea lions equipped with biologging tags were trained to swim alongside a boat cruising at steady speeds in the range of 4 to 10 kph. At each speed, and for each dive, we computed a measure called Gyro-Informed Dynamic Acceleration (GIDA) using a method incorporating gyroscope data with accelerometer data. We derived a new metric-Averaged Propulsive Body Acceleration (APBA), which is the average gain in speed per flipper stroke divided by mean stroke cycle duration. Our results show that the gyro-based measure (APBA) is a better predictor of speed than ODBA. We also found that APBA can estimate average thrust production during a single stroke-glide cycle, and can be used to estimate energy expended during swimming. The gyroscope-derived methods we describe should be generally applicable in swimming animals where propulsive accelerations can be clearly identified in the signal-and they should also prove useful for dead-reckoning and improving estimates of energy expenditures from locomotion.
Subject(s)
Sea Lions/physiology , Acceleration , Animals , Diving , Energy Metabolism , Female , Hydrodynamics , SwimmingABSTRACT
The traditional view of mysticete feeding involves static baleen directly sieving particles from seawater using a simple, dead-end flow-through filtration mechanism. Flow tank experiments on bowhead (Balaena mysticetus) baleen indicate the long-standing model of dead-end filtration, at least in balaenid (bowhead and right) whales, is not merely simplistic but wrong. To recreate continuous intraoral flow, sections of baleen were tested in a flume through which water and buoyant particles circulated with variable flow velocity. Kinematic sequences were analyzed to investigate movement and capture of particles by baleen plates and fringes. Results indicate that very few particles flow directly through the baleen rack; instead much water flows anteroposteriorly along the interior (lingual) side of the rack, allowing items to be carried posteriorly and accumulate at the posterior of the mouth where they might readily be swallowed. Since water flows mainly parallel to rather than directly through the filter, the cross-flow mechanism significantly reduces entrapment and tangling of minute items in baleen fringes, obviating the need to clean the filter. The absence of copepods or other prey found trapped in the baleen of necropsied right and bowhead whales supports this hypothesis. Reduced through-baleen flow was observed with and without boundaries modeling the tongue and lips, indicating that baleen itself is the main if not sole agent of crossflow. Preliminary investigation of baleen from balaenopterid whales that use intermittent filter feeding suggests that although the biomechanics and hydrodynamics of oral flow differ, cross-flow filtration may occur to some degree in all mysticetes.
Subject(s)
Bowhead Whale/physiology , Feeding Behavior , Animal Structures/anatomy & histology , Animal Structures/physiology , Animals , Biomechanical Phenomena , Bowhead Whale/anatomy & histology , Hydrodynamics , Models, Biological , Mouth/anatomy & histology , Mouth/physiologyABSTRACT
Major transitions between marine and freshwater habitats are relatively infrequent, primarily as a result of major physiological and ecological challenges. Few species of cartilaginous fish have evolved to occupy freshwater habitats. Current thought suggests that the metabolic physiology of sharks has remained a barrier to the diversification of this taxon in freshwater ecosystems. Here, we demonstrate that the physical properties of water provide an additional constraint for this species-rich group to occupy freshwater systems. Using hydromechanical modeling, we show that occurrence in fresh water results in a two- to three-fold increase in negative buoyancy for sharks and rays. This carries the energetic cost of lift production and results in increased buoyancy-dependent mechanical power requirements for swimming and increased optimal swim speeds. The primary source of buoyancy, the lipid-rich liver, offers only limited compensation for increased negative buoyancy as a result of decreasing water density; maintaining the same submerged weight would involve increasing the liver volume by very large amounts: 3- to 4-fold in scenarios where liver density is also reduced to currently observed minimal levels and 8-fold without any changes in liver density. The first data on body density from two species of elasmobranch occurring in freshwater (the bull shark Carcharhinus leucas, Müller and Henle 1839, and the largetooth sawfish Pristis pristis, Linnaeus 1758) support this hypothesis, showing similar liver sizes as marine forms but lower liver densities, but the greatest negative buoyancies of any elasmobranch studied to date. Our data suggest that the mechanical challenges associated with buoyancy control may have hampered the invasion of freshwater habitats in elasmobranchs, highlighting an additional key factor that may govern the predisposition of marine organisms to successfully establish in freshwater habitats.
Subject(s)
Elasmobranchii/anatomy & histology , Elasmobranchii/physiology , Liver/anatomy & histology , Animals , Biomechanical Phenomena , Ecosystem , Fresh Water , Models, Biological , SwimmingABSTRACT
The role of endomyocardial biopsy (EMB) in the diagnosis and treatment of adults presenting a newly unexplained cardiomyopathy remains controversial and the clinical indication varies among different centers. Furthermore, the choice of the ventricular site for EMB is still under debate. The right ventricular EMB is commonly used due to safety features and left ventricle (LV) EMB has not yet gained acceptance because of concerns about possible complications. Of note, when LV-EMB is performed, the femoral artery is usually the access site. There is a large body of evidence supporting the safety and feasibility of transradial approach in a broad spectrum of patients and settings in the catheterization laboratory. Therefore, the aim of the present manuscript is to report the feasibility of performing a LV-EMB by the transradial approach using a sheathless guiding catheter as an ad hoc procedure.
Subject(s)
Amyloidosis/pathology , Biopsy/instrumentation , Cardiac Catheterization/instrumentation , Cardiomyopathies/pathology , Endocardium/pathology , Heart Failure/pathology , Heart Ventricles/pathology , Hypertrophy, Left Ventricular/pathology , Myocardium/pathology , Adult , Biopsy/methods , Cardiac Catheterization/methods , Equipment Design , Female , Humans , Middle Aged , Radial ArteryABSTRACT
A 24-year-old healthy man consulted to our center because of typical on-and-off chest-pain and an electrocardiogram showing ST-segment elevation in inferior leads. An urgent coronary angiography showed angiographically normal coronary arteries. Cardiovascular magnetic resonance imaging confirmed acute myocarditis. Although acute myocarditis triggering coronary spasm is an uncommon association, it is important to recognize it, particularly for the management for those patients presenting with ST-segment elevation and suspect myocardial infarction and angiographically normal coronary arteries. The present report highlights the role of cardiovascular magnetic resonance imaging to identify acute myocarditis as the underlying cause.
ABSTRACT
Muscle serves a wide variety of mechanical functions during animal feeding and locomotion, but the performance of this tissue is limited by how far it can be extended. In rorqual whales, feeding and locomotion are integrated in a dynamic process called lunge feeding, where an enormous volume of prey-laden water is engulfed into a capacious ventral oropharyngeal cavity that is bounded superficially by skeletal muscle and ventral groove blubber (VGB). The great expansion of the cavity wall presents a mechanical challenge for the physiological limits of skeletal muscle, yet its role is considered fundamental in controlling the flux of water into the mouth. Our analyses of the functional properties and mechanical behaviour of VGB muscles revealed a crimped microstructure in an unstrained, non-feeding state that is arranged in parallel with dense and straight elastin fibres. This allows the muscles to accommodate large tissue deformations of the VGB yet still operate within the known strain limits of vertebrate skeletal muscle. VGB transverse strains in routine-feeding rorquals were substantially less than those observed in dead ones, where decomposition gas stretched the VGB to its elastic limit, evidence supporting the idea that eccentric muscle contraction modulates the rate of expansion and ultimate size of the ventral cavity during engulfment.
Subject(s)
Connective Tissue/anatomy & histology , Feeding Behavior/physiology , Locomotion/physiology , Models, Biological , Muscle, Skeletal/anatomy & histology , Oropharynx/anatomy & histology , Whales/physiology , Animals , Biomechanical Phenomena/physiology , Connective Tissue/physiology , Muscle, Skeletal/physiology , Oropharynx/physiologyABSTRACT
Bulk-filter feeding is an energetically efficient strategy for resource acquisition and assimilation, and facilitates the maintenance of extreme body size as exemplified by baleen whales (Mysticeti) and multiple lineages of bony and cartilaginous fishes. Among mysticetes, rorqual whales (Balaenopteridae) exhibit an intermittent ram filter feeding mode, lunge feeding, which requires the abandonment of body-streamlining in favor of a high-drag, mouth-open configuration aimed at engulfing a very large amount of prey-laden water. Particularly while lunge feeding on krill (the most widespread prey preference among rorquals), the effort required during engulfment involve short bouts of high-intensity muscle activity that demand high metabolic output. We used computational modeling together with morphological and kinematic data on humpback (Megaptera noveaangliae), fin (Balaenoptera physalus), blue (Balaenoptera musculus) and minke (Balaenoptera acutorostrata) whales to estimate engulfment power output in comparison with standard metrics of metabolic rate. The simulations reveal that engulfment metabolism increases across the full body size of the larger rorqual species to nearly 50 times the basal metabolic rate of terrestrial mammals of the same body mass. Moreover, they suggest that the metabolism of the largest body sizes runs with significant oxygen deficits during mouth opening, namely, 20% over maximum VO2 at the size of the largest blue whales, thus requiring significant contributions from anaerobic catabolism during a lunge and significant recovery after a lunge. Our analyses show that engulfment metabolism is also significantly lower for smaller adults, typically one-tenth to one-half VO2|max. These results not only point to a physiological limit on maximum body size in this lineage, but also have major implications for the ontogeny of extant rorquals as well as the evolutionary pathways used by ancestral toothed whales to transition from hunting individual prey items to filter feeding on prey aggregations.
Subject(s)
Biological Evolution , Body Size , Energy Metabolism , Feeding Behavior , Whales/physiology , Animals , Biomechanical Phenomena , Computer Simulation , Hydrodynamics , Models, Biological , MovementABSTRACT
Rorqual whales (Balaenopteridae) represent not only some of the largest animals of all time, but also exhibit a wide range in intraspecific and interspecific body size. Balaenopterids are characterized by their extreme lunge-feeding behaviour, a dynamic process that involves the engulfment of a large volume of prey-laden water at a high energetic cost. To investigate the consequences of scale and morphology on lunge-feeding performance, we determined allometric equations for fin whale body dimensions and engulfment capacity. Our analysis demonstrates that larger fin whales have larger skulls and larger buccal cavities relative to body size. Together, these data suggest that engulfment volume is also allometric, increasing with body length as L(3.5)(body). The positive allometry of the skull is accompanied by negative allometry in the tail region. The relative shortening of the tail may represent a trade-off for investing all growth-related resources in the anterior region of the body. Although enhanced engulfment volume will increase foraging efficiency, the work (energy) required to accelerate the engulfed water mass during engulfment will be relatively higher in larger rorquals. If the mass-specific energetic cost of a lunge increases with body size, it will have major consequences for rorqual foraging ecology and evolution.