Resonance of the tympanoperiotic complex of fin whales with implications for their low frequency hearing.

The tympanoperiotic complex (TPC) bones of the fin whale skull were studied using experimental measurements and simulation modeling to provide insight into the low frequency hearing of these animals. The study focused on measuring the sounds emitted by the left and right TPC bones when the bones were tapped at designated locations. Radiated sound was recorded by eight microphones arranged around the tympanic bulla. A finite element model was also created to simulate the natural mode vibrations of the TPC and ossicular chain, using a 3D mesh generated from a CT scan. The simulations produced mode shapes and frequencies for various Young's modulus and density values. The recorded sound amplitudes were compared with the normal component of the simulated displacement and it was found that the modes identified in the experiment most closely resembled those found with Young's modulus for stiff and flexible bone set to 25 and 5 GPa, respectively. The first twelve modes of vibration of the TPC had resonance frequencies between 100Hz and 6kHz. Many vibrational modes focused energy at the sigmoidal process, and therefore the ossicular chain. The resonance frequencies of the left and right TPC were offset, suggesting a mechanism for the animals to have improved hearing at a range of frequencies as well as a mechanism for directionality in their perception of sounds.

Cholla cactus frames as lightweight and torsionally tough biological materials.

Biological materials tested in compression, tension, and impact inspire designs for strong and tough materials, but torsion is a relatively neglected loading mode. The wood skeletons of cholla cacti, subject to spartan desert conditions and hurricane force winds, provide a new template for torsionally resilient biological materials. Novel mesostructural characterization methods of laser-scanning and photogrammetry are used alongside traditional optical microscopy, scanning electron microscopy, and micro-computed tomography to identify mechanisms responsible for torsional resistance. These methods, in combination with finite element analysis reveal how cholla meso and macro-porosity and fibril orientation contribute to highly density-efficient mechanical behavior. Selective lignification and macroscopic tubercle pore geometry contribute to density-efficient shear stiffness, while mesoscopic wood fiber straightening, delamination, pore collapse, and fiber pullout provide extrinsic toughening mechanisms. These energy absorbing mechanisms are enabled by the hydrated material level properties. Together, these hierarchical behaviors allow the cholla to far exceed bamboo and trabecular bone in its ability to combine specific torsional stiffness, strength, and toughness. STATEMENT OF SIGNIFICANCE: The Cholla cactus experiences, due to the high velocity desert winds, high torsional loads. Our study has revealed the amazingly ingenious strategy by which the tubular structure containing arrays of voids intermeshed with wood fibers resists these high loads. Deformation is governed by compressive and tensile stresses which are greatest at 45 degrees to the cross section. It proceeds by stretching, sliding, and bending of the wood fibers which are coupled with the pore collapse, resulting in delayed failure and a high torsional toughness.

Auditory chain reaction: Effects of sound pressure and particle motion on auditory structures in fishes.

Despite the diversity in fish auditory structures, it remains elusive how otolith morphology and swim bladder-inner ear (= otophysic) connections affect otolith motion and inner ear stimulation. A recent study visualized sound-induced otolith motion; but tank acoustics revealed a complex mixture of sound pressure and particle motion. To separate sound pressure and sound-induced particle motion, we constructed a transparent standing wave tube-like tank equipped with an inertial shaker at each end while using X-ray phase contrast imaging. Driving the shakers in phase resulted in maximised sound pressure at the tank centre, whereas particle motion was maximised when shakers were driven out of phase (180°). We studied the effects of two types of otophysic connections-i.e. the Weberian apparatus (Carassius auratus) and anterior swim bladder extensions contacting the inner ears (Etroplus canarensis)-on otolith motion when fish were subjected to a 200 Hz stimulus. Saccular otolith motion was more pronounced when the swim bladder walls oscillated under the maximised sound pressure condition. The otolith motion patterns mainly matched the orientation patterns of ciliary bundles on the sensory epithelia. Our setup enabled the characterization of the interplay between the auditory structures and provided first experimental evidence of how different types of otophysic connections affect otolith motion.

Coupled finite element/boundary element formulation for scattering from axially-symmetric objects in three dimensions.

The fluid-structure interaction technique provides a paradigm for solving scattering from elastic structures embedded in an environment characterized by a Green's function, by a combination of finite and boundary element methods. In this technique, the finite element method is used to discretize the equations of motion for the structure and the Helmholtz-Kirchhoff integral with the appropriate Green's function is used to produce the discrete pressure field in the exterior medium. The two systems of equations are coupled at the surface of the structure by imposing the continuity of pressure and normal particle velocity. The present method condenses the finite element model so that finally only the boundary element problem needs to be solved. This results in a significant reduction in the number of unknowns and hence a much lower cost. In this paper, the fluid-structure interaction method is specialized to axially-symmetric objects for non-axially-symmetric loading in free space using a circumferential Fourier expansion of the fields. The specialization of the method to axially-symmetric objects results in even further significant reductions in computation. The method is validated using well-known benchmark solutions. A derivation of the method for an arbitrarily-shaped elastic structure embedded in an arbitrary environment characterized by a Green's function is given in the Appendix.

Functional Morphology and Symmetry in the Odontocete Ear Complex.

Odontocete ear complexes or tympanoperiotic complexes (TPCs) were compared for asymmetry. Left and right TPCs were collected from one long-beaked common dolphin (Delphinus capensis) and one Amazon River dolphin (Inia geoffrensis). Asymmetry was assessed by volumetric comparisons of left and right TPCs and by visual comparison of superimposed models of the right TPC to a reflected mirror image of the left TPC. Kolmogorov-Smirnov tests were performed to compare the resonant frequencies of the TPCs as calculated by vibrational analysis. All analyses found slight differences between TPCs from the same specimen in contrast to the directional asymmetry in the nasal region of odontocete skulls.

Directional Hearing and Head-Related Transfer Function in Odontocete Cetaceans.

The head-related transfer function (HRTF) is an important descriptor of spatial sound field reception by the listener. In this study, we computed the HRTF of the common dolphin Delphinus delphis. The received sound pressure level at various locations within the acoustic fats of the internal pinna near the surface of the tympanoperiotic complex (TPC) was calculated for planar incident waves directed toward the animal. The relative amplitude of the received pressure versus the incident pressure was the representation of the HRTF from the point of view of the animal. It is of interest that (1) different locations on the surface of the TPC resulted in different HRTFs, (2) the HRTFs for the left and right ears were slightly asymmetric, and (3) the locations of the peaks of the HRTF depended on the frequency of the incident wave.

Sound Transmission Validation and Sensitivity Studies in Numerical Models.

In 1974, Norris and Harvey published an experimental study of sound transmission into the head of the bottlenose dolphin. We used this rare source of data to validate our Vibroacoustic Toolkit, an array of numerical modeling simulation tools. Norris and Harvey provided measurements of received sound pressure in various locations within the dolphin's head from a sound source that was moved around the outside of the head. Our toolkit was used to predict the curves of pressure with the best-guess input data (material properties, transducer and hydrophone locations, and geometry of the animal's head). In addition, we performed a series of sensitivity analyses (SAs). SA is concerned with understanding how input changes to the model influence the outputs. SA can enhance understanding of a complex model by finding and analyzing unexpected model behavior, discriminating which inputs have a dominant effect on particular outputs, exploring how inputs combine to affect outputs, and gaining insight as to what additional information improves the model's ability to predict. Even when a computational model does not adequately reproduce the behavior of a physical system, its sensitivities may be useful for developing inferences about key features of the physical system. Our findings may become a valuable source of information for modeling the interactions between sound and anatomy.

Fin whale sound reception mechanisms: skull vibration enables low-frequency hearing.

Hearing mechanisms in baleen whales (Mysticeti) are essentially unknown but their vocalization frequencies overlap with anthropogenic sound sources. Synthetic audiograms were generated for a fin whale by applying finite element modeling tools to X-ray computed tomography (CT) scans. We CT scanned the head of a small fin whale (Balaenoptera physalus) in a scanner designed for solid-fuel rocket motors. Our computer (finite element) modeling toolkit allowed us to visualize what occurs when sounds interact with the anatomic geometry of the whale's head. Simulations reveal two mechanisms that excite both bony ear complexes, (1) the skull-vibration enabled bone conduction mechanism and (2) a pressure mechanism transmitted through soft tissues. Bone conduction is the predominant mechanism. The mass density of the bony ear complexes and their firmly embedded attachments to the skull are universal across the Mysticeti, suggesting that sound reception mechanisms are similar in all baleen whales. Interactions between incident sound waves and the skull cause deformations that induce motion in each bony ear complex, resulting in best hearing sensitivity for low-frequency sounds. This predominant low-frequency sensitivity has significant implications for assessing mysticete exposure levels to anthropogenic sounds. The din of man-made ocean noise has increased steadily over the past half century. Our results provide valuable data for U.S. regulatory agencies and concerned large-scale industrial users of the ocean environment. This study transforms our understanding of baleen whale hearing and provides a means to predict auditory sensitivity across a broad spectrum of sound frequencies.

Angular oscillation of solid scatterers in response to progressive planar acoustic waves: do fish otoliths rock?

Fish can sense a wide variety of sounds by means of the otolith organs of the inner ear. Among the incompletely understood components of this process are the patterns of movement of the otoliths vis-à-vis fish head or whole-body movement. How complex are the motions? How does the otolith organ respond to sounds from different directions and frequencies? In the present work we examine the responses of a dense rigid scatterer (representing the otolith) suspended in an acoustic fluid to low-frequency planar progressive acoustic waves. A simple mechanical model, which predicts both translational and angular oscillation, is formulated. The responses of simple shapes (sphere and hemisphere) are analyzed with an acoustic finite element model. The hemispherical scatterer is found to oscillate both in the direction of the propagation of the progressive waves and also in the plane of the wavefront as a result of angular motion. The models predict that this characteristic will be shared by other irregularly-shaped scatterers, including fish otoliths, which could provide the fish hearing mechanisms with an additional component of oscillation and therefore one more source of acoustical cues.

A new acoustic portal into the odontocete ear and vibrational analysis of the tympanoperiotic complex.

Global concern over the possible deleterious effects of noise on marine organisms was catalyzed when toothed whales stranded and died in the presence of high intensity sound. The lack of knowledge about mechanisms of hearing in toothed whales prompted our group to study the anatomy and build a finite element model to simulate sound reception in odontocetes. The primary auditory pathway in toothed whales is an evolutionary novelty, compensating for the impedance mismatch experienced by whale ancestors as they moved from hearing in air to hearing in water. The mechanism by which high-frequency vibrations pass from the low density fats of the lower jaw into the dense bones of the auditory apparatus is a key to understanding odontocete hearing. Here we identify a new acoustic portal into the ear complex, the tympanoperiotic complex (TPC) and a plausible mechanism by which sound is transduced into the bony components. We reveal the intact anatomic geometry using CT scanning, and test functional preconceptions using finite element modeling and vibrational analysis. We show that the mandibular fat bodies bifurcate posteriorly, attaching to the TPC in two distinct locations. The smaller branch is an inconspicuous, previously undescribed channel, a cone-shaped fat body that fits into a thin-walled bony funnel just anterior to the sigmoid process of the TPC. The TPC also contains regions of thin translucent bone that define zones of differential flexibility, enabling the TPC to bend in response to sound pressure, thus providing a mechanism for vibrations to pass through the ossicular chain. The techniques used to discover the new acoustic portal in toothed whales, provide a means to decipher auditory filtering, beam formation, impedance matching, and transduction. These tools can also be used to address concerns about the potential deleterious effects of high-intensity sound in a broad spectrum of marine organisms, from whales to fish.

A theoretical and experimental analysis of radiofrequency ablation with a multielectrode, phased, duty-cycled system.

BACKGROUND: The development of a unique radiofrequency (RF) cardiac ablation system, for the treatment of cardiac arrhythmias, is driven by the clinical need to safely create large uniform lesions while controlling lesion depth. Computational analysis of a finite element model of a three-dimensional, multielectrode, cardiac ablation catheter, powered by a temperature-controlled, multiphase, duty-cycled RF generator, is presented. METHODS: The computational model for each of the five operating modes offered by the generator is compared to independent tissue temperature measurements taken during in vitro ablation experiments performed on bovine myocardium. RESULTS: The results of the model agree with experimental temperature measurements very closely-the average values for mean error, root mean square difference, and correlation coefficient were 1.9°C, 13.3%, and 0.97, respectively. Lesions are shown to be contiguous and no significant edge effects are observed. CONCLUSIONS: Both the in vitro and computational model results demonstrate that lesion depth decreases consistently as the bipolar-to-unipolar ratio increases-suggesting a clinical application to potentially control lesion depth with higher fidelity than is currently available. The effect of variable design parameters and clinical conditions on RF ablation can now be expeditiously studied with this validated model.

Dynamic assessment of the vaginal high-pressure zone using high-definition manometery, 3-dimensional ultrasound, and magnetic resonance imaging of the pelvic floor muscles.

OBJECTIVE: We used a novel technique, high-definition manometry (HDM) that utilizes 256 tactile sensitive microtransducers to define the characteristics of vaginal high-pressure zone. STUDY DESIGN: Sixteen nullipara asymptomatic women were studied using HDM, transperineal 2-dimensional dynamic ultrasound and dynamic magnetic resonance (MR) imaging. RESULTS: Vaginal high-pressure zone revealed higher contact pressures in anterior and posterior directions compared with lateral directions, both at rest and squeeze. At rest, anterior pressure cluster is located 10 mm cephalad to posterior pressure cluster; with squeeze the latter moves in the cranial direction by 7 mm. Ultrasound and MR images revealed that the anorectal angle moves cephalad and ventrally during squeeze. Cephalad movement of posterior pressure cluster during squeeze is similar to the cranial movement of anorectal angle. CONCLUSION: We propose that the vaginal high-pressure zone represents the constrictor function and cranial movement of the posterior pressure cluster represents the elevator function of pelvic floor. HDM may be used to measure the constrictor and elevator functions of pelvic floor muscles.

Quantity and distribution of levator ani stretch during simulated vaginal childbirth.

OBJECTIVE: The objective of the study was to develop a model of the female pelvic floor to study levator stretch during simulated childbirth. STUDY DESIGN: Magnetic resonance data from an asymptomatic nulligravida were segmented into pelvic muscles and bones to create a simulation model. Stiffness estimates of lateral and anteroposterior levator attachments were varied to estimate the impact on levator stretch. A 9 cm sphere was passed through the pelvis, along the path of the vagina, simulating childbirth. Levator response was interpreted at 4 positions of the sphere, simulating fetal head descent. The levator was color mapped to display the stretch experienced. RESULTS: A maximum stretch ratio of 3.5 to 1 was seen in the posteriomedial puborectalis. Maximum stretch increased with increasing stiffness of lateral levator attachments. CONCLUSION: Although preliminary, this work may help explain epidemiologic data regarding the pelvic floor impact of a first delivery. The models and simulation technique need refinement, but they may help study the effect of labor parameters on the pelvic floor.

Acoustic pathways revealed: simulated sound transmission and reception in Cuvier's beaked whale (Ziphius cavirostris).

The finite element modeling (FEM) space reported here contains the head of a simulated whale based on CT data sets as well as physical measurements of sound-propagation characteristics of actual tissue samples. Simulated sound sources placed inside and outside of an adult male Cuvier's beaked whale (Ziphius cavirostris) reveal likely sound propagation pathways into and out of the head. Two separate virtual sound sources that were located at the left and right phonic lips produced beams that converged just outside the head. This result supports the notion that dual sound sources can interfere constructively to form a biologically useful and, in fact, excellent sonar beam in front of the animal. The most intriguing FEM results concern pathways by which sounds reach the ears. The simulations reveal a previously undescribed 'gular pathway' for sound reception in Ziphius. Propagated sound pressure waves enter the head from below and between the lower jaws, pass through an opening created by the absence of the medial bony wall of the posterior mandibles, and continue toward the bony ear complexes through the internal mandibular fat bodies. This new pathway has implications for understanding the evolution of underwater hearing in odontocetes. Our model also provides evidence for receive beam directionality, off-axis acoustic shadowing and a plausible mechanism for the long-standing orthodox sound reception pathway in odontocetes. The techniques developed for this study can be used to study acoustic perturbation in a wide variety of marine organisms.

Anatomic geometry of sound transmission and reception in Cuvier's beaked whale (Ziphius cavirostris).

This study uses remote imaging technology to quantify, compare, and contrast the cephalic anatomy between a neonate female and a young adult male Cuvier's beaked whale. Primary results reveal details of anatomic geometry with implications for acoustic function and diving. Specifically, we describe the juxtaposition of the large pterygoid sinuses, a fibrous venous plexus, and a lipid-rich pathway that connects the acoustic environment to the bony ear complex. We surmise that the large pterygoid air sinuses are essential adaptations for maintaining acoustic isolation and auditory acuity of the ears at depth. In the adult male, an acoustic waveguide lined with pachyosteosclerotic bones is apparently part of a novel transmission pathway for outgoing biosonar signals. Substitution of dense tissue boundaries where we normally find air sacs in delphinoids appears to be a recurring theme in deep-diving beaked whales and sperm whales. The anatomic configuration of the adult male Ziphius forehead resembles an upside-down sperm whale nose and may be its functional equivalent, but the homologous relationships between forehead structures are equivocal.