Fast-moving bat ears create informative Doppler shifts.

Many animals have evolved adept sensory systems that enable dexterous mobility in complex environments. Echolocating bats hunting in dense vegetation represent an extreme case of this, where all necessary information about the environment must pass through a parsimonious channel of pulsed, 1D echo signals. We have investigated whether certain bats (rhinolophids and hipposiderids) actively create Doppler shifts with their pinnae to encode additional sensory information. Our results show that the bats' active pinna motions are a source of Doppler shifts that have all attributes required for a functional relevance: (i) the Doppler shifts produced were several times larger than the reported perception threshold; (ii) the motions of the fastest moving pinna portions were oriented to maximize the Doppler shifts for echoes returning from the emission direction, indicating a possible evolutionary optimization; (iii) pinna motions coincided with echo reception; (iv) Doppler-shifted signals from the fast-moving pinna portion entered the ear canal of a biomimetic pinna model; and (v) the time-frequency Doppler shift signatures were found to encode target direction in an orderly fashion. These results indicate that instead of avoiding or suppressing all self-produced Doppler shifts, rhinolophid and hipposiderid bats actively create Doppler shifts with their own pinnae. These bats could hence make use of a previously unknown nonlinear mechanism for the encoding of sensory information, based on Doppler signatures. Such a mechanism could be a source for the discovery of sensing principles not only in sensory physiology but also in the engineering of sensory systems.

Facial Blood Flow Responses to Dynamic Exercise.

We reported previously that a static handgrip exercise evoked regional differences in the facial blood flow. The present study examined whether regional differences in facial blood flow are also evoked during dynamic exercise. Facial blood flow was measured by laser speckle flowgraphy during 15 min of cycling exercise at heart rates of 120 bpm, 140 bpm and 160 bpm in 12 subjects. The facial vascular conductance index was calculated from the blood flow and mean arterial pressure. The regional blood flow and conductance index values were determined in the forehead, eyelid, nose, cheek, ear and lip. One-way ANOVA and Tukey's post-hoc test were used to examine effects of exercise intensity and target regions. The blood flow and conductance index in skin areas increased significantly with the exercise intensity. The blood flow and conductance index in the lip increased significantly at 120 bpm and 140 bpm compared to the control, while the values in the lip at 160 bpm did not change from the control values. These results suggest that the blood flow in facial skin areas, not in the lip, responds similarly to dynamic exercise, in contrast to the responses to static exercise.

The Auditory Mechanics of the Outer Ear of the Bush Cricket: A Numerical Approach.

Bush crickets have tympanal ears located in the forelegs. Their ears are elaborate, as they have outer-, middle-, and inner-ear components. The outer ear comprises an air-filled tube derived from the respiratory trachea, the acoustic trachea (AT), which transfers sound from the mesothoracic acoustic spiracle to the internal side of the ear drums in the legs. A key feature of the AT is its capacity to reduce the velocity of sound propagation and alter the acoustic driving forces of the tympanum (the ear drum), producing differences in sound pressure and time between the left and right sides, therefore aiding the directional hearing of the animal. It has been demonstrated experimentally that the tracheal sound transmission generates a gain of ∼15 dB and a propagation velocity of 255 ms-1, an approximately 25% reduction from free-field propagation. However, the mechanism responsible for this change in sound pressure level and velocity remains elusive. In this study, we investigate the mechanical processes behind the sound pressure gain in the AT by numerically modeling the tracheal acoustic behavior using the finite-element method and real three-dimensional geometries of the tracheae of the bush cricket Copiphora gorgonensis. Taking into account the thermoviscous acoustic-shell interaction on the propagation of sound, we analyze the effects of the horn-shaped domain, material properties of the tracheal wall, and the thermal processes on the change in sound pressure level in the AT. Through the numerical results obtained, it is discerned that the tracheal geometry is the main factor contributing to the observed pressure gain.

Genetic correlations between cartilage regeneration and degeneration reveal an inverse relationship.

OBJECTIVE: The etiology of osteoarthritis (OA) is unknown, however, there appears to be a significant contribution from genetics. We have identified recombinant inbred strains of mice derived from LG/J (large) and SM/J (small) strains that vary significantly in their ability to repair articular cartilage and susceptibility to post-traumatic OA due to their genetic composition. Here, we report cartilage repair phenotypes in the same strains of mice in which OA susceptibility was analyzed previously, and determine the genetic correlations between phenotypes. DESIGN: We used 12 recombinant inbred strains, including the parental strains, to test three phenotypes: ear-wound healing (n = 263), knee articular cartilage repair (n = 131), and post-traumatic OA (n = 53) induced by the surgical destabilization of the medial meniscus (DMM). Genetic correlations between various traits were calculated as Pearson's correlation coefficients of strain means. RESULTS: We found a significant positive correlation between ear-wound healing and articular cartilage regeneration (r = 0.71; P = 0.005). We observed a strong inverse correlation between articular cartilage regeneration and susceptibility to OA based on maximum (r = -0.54; P = 0.036) and summed Osteoarthritis Research Society International (OARSI) scores (r = -0.56; P = 0.028). Synovitis was not significantly correlated with articular cartilage regeneration but was significantly positively correlated with maximum (r = 0.63; P = 0.014) and summed (r = 0.70; P = 0.005) OARSI scores. Ectopic calcification was significantly positively correlated with articular cartilage regeneration (r = 0.59; P = 0.021). CONCLUSIONS: Using recombinant inbred strains, our study allows, for the first time, the measurement of genetic correlations of regeneration phenotypes with degeneration phenotypes, characteristic of OA (cartilage degeneration, synovitis). We demonstrate that OA is positively correlated with synovitis and inversely correlated with the ability to repair cartilage. These results suggest an addition to the risk paradigm for OA from a focus on degeneration to regeneration.

Ethyl Alcohol Versus Botulinum Toxin A: A Comparative Study of the Visual and Histopathological Outcomes in the Rabbit Anterior Auricular Muscle Model.

BACKGROUND: Botulinum toxin has long been known for its paralytic effects at the neuromuscular junction. Although it has been widely used for vascular and nervous tissues, there has been no study of the aesthetic effects of the application of ethanol to muscle tissues to date. OBJECTIVE: The authors aimed to demonstrate the effects of the application of ethanol to muscle tissues after an intramuscular injection and to compare the effects of botulinum toxin A (BTA) and ethanol. METHODS AND MATERIALS: A total of 28 rabbits were divided into 4 groups (n = 7 each). Botulinum toxin A (5 units) and different concentrations of ethanol (5 cc) were injected into the left and right anterior auricular muscles of all rabbits, respectively. Ear ptosis was assessed, and histopathological examination was performed after all rabbits were euthanized in the eighth week. RESULTS: Muscle function was affected earlier in ethanol-treated ears than in botulinum-treated ears; however, the ptotic effect lasted for a significantly shorter duration in ethanol-injected ears than in BTA-applied ears. CONCLUSION: Ethanol can block muscle function reversibly and can serve as an alternative to BTA, particularly when rapid results are desirable.

Dynamic relationship between noseleaf and pinnae in echolocating hipposiderid bats.

Old World leaf-nosed bats (family Hipposideridae) can deform the shapes of their 'noseleaves' (i.e. ultrasonic emission baffles) and outer ears during echolocation behaviors. Prior work has shown that deformations on the emission as well as on the reception side can have an impact on the properties of the emitted/received sonar signals. The occurrence of the deformations on the emission and reception sides raises the question of whether the bats coordinate these two dynamic biosonar features to achieve synergistic effects. To address this question, simultaneous three-dimensional reconstructions of the trajectories of landmarks on the dynamic noseleaf and pinna geometries have been obtained in great roundleaf bats (Hipposideros pratti). These joint kinematics data on the noseleaf and pinnae have shown both qualitative and quantitative relationships between the noseleaf and pinna motions: large noseleaf deformations (opening or closing) tended to be associated with non-rigid pinna motions. Furthermore, closing deformations of the noseleaves tended to co-occur with closing motions of the pinna. Finally, a canonical correlation analysis of the motion trajectories has revealed a tight correlation between the motions of the landmarks on the noseleaf and both pinnae. These results demonstrate that the biosonar system of hipposiderid bats includes coordinated emission and reception dynamics.

Limitations of present models of blast-induced sound power conduction through the external and middle ear.

The use of models to predict the effect of blast-like impulses on hearing function is an ongoing topic of investigation relevant to hearing protection and hearing-loss prevention in the modern military. The first steps in the hearing process are the collection of sound power from the environment and its conduction through the external and middle ear into the inner ear. Present efforts to quantify the conduction of high-intensity sound power through the auditory periphery depend heavily on modeling. This paper reviews and elaborates on several existing models of the conduction of high-level sound from the environment into the inner ear and discusses the shortcomings of these models. A case is made that any attempt to more accurately define the workings of the middle ear during high-level sound stimulation needs to be based on additional data, some of which has been recently gathered.

Skin shedding and tissue regeneration in African spiny mice (Acomys).

Evolutionary modification has produced a spectrum of animal defence traits to escape predation, including the ability to autotomize body parts to elude capture. After autotomy, the missing part is either replaced through regeneration (for example, in urodeles, lizards, arthropods and crustaceans) or permanently lost (such as in mammals). Although most autotomy involves the loss of appendages (legs, chelipeds, antennae or tails, for example), skin autotomy can occur in certain taxa of scincid and gekkonid lizards. Here we report the first demonstration of skin autotomy in Mammalia (African spiny mice, Acomys). Mechanical testing showed a propensity for skin to tear under very low tension and the absence of a fracture plane. After skin loss, rapid wound contraction was followed by hair follicle regeneration in dorsal skin wounds. Notably, we found that regenerative capacity in Acomys was extended to ear holes, where the mice exhibited complete regeneration of hair follicles, sebaceous glands, dermis and cartilage. Salamanders capable of limb regeneration form a blastema (a mass of lineage-restricted progenitor cells) after limb loss, and our findings suggest that ear tissue regeneration in Acomys may proceed through the assembly of a similar structure. This study underscores the importance of investigating regenerative phenomena outside of conventional model organisms, and suggests that mammals may retain a higher capacity for regeneration than was previously believed. As re-emergent interest in regenerative medicine seeks to isolate molecular pathways controlling tissue regeneration in mammals, Acomys may prove useful in identifying mechanisms to promote regeneration in lieu of fibrosis and scarring.

Sweep frequency impedance measures in young infants: developmental characteristics from birth to 6 months.

OBJECTIVE: International Journal of Audiology To study the developmental characteristics of sweep frequency impedance (SFI) measures in healthy infants from birth to 6 months. DESIGN: All infants were assessed using high-frequency tympanometry (HFT), distortion product otoacoustic emission (DPOAE) and SFI tests. SFI measures consisted of measurement of resonance frequency (RF) and mobility (ΔSPL) of the outer and middle ear. A mixed model analysis of variance was applied to the SFI data to examine the effect of age on RF and ΔSPL. STUDY SAMPLE: Study included 117 ears from 83 infants of different age groups from birth to 6 months. RESULTS: The mean RF of the outer ear increased from 279 Hz at birth to 545 Hz at 4 months, whereas mean ΔSPL of the outer ear decreased from 7.9 dB at birth to 3.7 dB at 4 months of age. In contrast, the mean RF and ΔSPL of the middle ear did not change significantly with age up to 6 months. CONCLUSIONS: Developmental characteristics should be considered when evaluating the function of the outer and middle ear of young infants (≤6 months) using the SFI. The preliminary normative SFI data established in this study may be used to assist with the evaluation.

Vertical normal modes of human ears: Individual variation and frequency estimation from pinna anthropometry.

Beyond the first peak of head-related transfer functions or pinna-related transfer functions (PRTFs) human pinnae are known to have two normal modes with "vertical" resonance patterns, involving two or three pressure anti-nodes in cavum, cymba, and fossa. However, little is known about individual variations in these modes, and there is no established model for estimating their center-frequencies from anthropometry. Here, with geometries of 38 pinnae measured, PRTFs were calculated and vertical modes visualized by numerical simulation. Most pinnae were found to have both Cavum-Fossa and Cavum-Cymba modes, with opposite-phase anti-nodes in cavum and either fossa or cymba, respectively. Nevertheless in both modes, fossa involvement varied substantially across pinnae, dependent on scaphoid fossa depth and cymba shallowness. Linear regression models were evaluated in mode frequency estimation, with 3322 measures derived from 31 pinna landmarks. The Cavum-Fossa normal mode frequency was best estimated [correlation coefficient r = 0.89, mean absolute error (MAE) = 257 Hz or 4.4%] by the distance from canal entrance to helix rim, and cymba horizontal depth. The Cavum-Cymba normal mode frequency was best estimated (r = 0.92, MAE = 247 Hz or 3.2%) by the sagittal-plane distance from concha floor to cymba anterior wall, and cavum horizontal depth.

Otoplasty Outcomes With Different Suture Materials in a Rabbit Model.

Otoplasty is a commonly performed procedure to correct prominent ears. Many different otoplasty techniques have been described but there is no gold standard technique. As well, many different suture materials are used in otoplasty but studies directly comparing different sutures materials are lacking. An otoplasty outcome study with Nylon and Mersilene (2 of the most commonly used sutures in otoplasty) sutures was conducted using a rabbit model. Each rabbit ear was randomized to receive a Mustardé-type horizontal mattress suture with either 4-0 clear Nylon (N = 12 ears) or 4-0 Mersilene sutures (N = 12 ears). Two weeks after surgery, the auricular bend angle was measured with a finger goniometer and histologic analysis with hematoxylin and eosin staining was performed on the rabbit auricular cartilage. Overall, there was no significant difference in the mean bend angle between the 2 groups (Nylon: 135.8°, SD = 22.7° and Mersilene: 143.2°, SD = 19.7°; P = 0.559). Also, no qualitative difference was observed on histologic analysis between the 2 suture groups. In the current rabbit model study, both Nylon and Mersilene sutures performed well and no significant differences were noted.

Modeling and estimating acoustic transfer functions of external ears with or without headphones.

The acoustic transfer functions of external ears with or without headphones affect the features of perceived sounds and vary considerably with listeners and headphones. A method for estimating the frequency responses of external-ear transfer functions from the sound at the entrance of a blocked ear canal (or from the input of a headphone) to the sound at the eardrum for different listeners and headphones is developed based on an acoustic signal model of external ears. The model allows for applying realistic data about individual external ears and headphones and is advantageous over current standard ear simulators with fixed structures limited to simulating average ear canals and eardrum impedances below 10 kHz. Given different eardrum impedances, ear canal shapes, lengths, and headphones, the frequency responses of external-ear transfer functions are estimated and presented. In addition, a method of determining the Norton equivalent volume velocity or Thevenien equivalent sound pressure sources of a headphone from sound pressure signals in an acoustic tube is presented. These methods are validated via direct measurements and expected to have applications in headphone sound reproduction, headphone and hearing aid design, and audiometric and psychoacoustic measurements to produce desired sounds at the eardrums of different listeners.

[Aural Stimulation with Capsaicin Ointment Improved the Swallowing Function in Patients with Dysphagia: Evaluation by the SMRC Scale].

Cough and swallowing reflexes are important airway-protective mechanisms against aspiration. Angiotensin-converting enzyme (ACE) inhibitors, one of the side effects of which is cough, have been reported to reduce the incidence of aspiration pneumonia in hypertensive patients with stroke. ACE inhibitors have also been reported to improve the swallowing function in post-stroke patients. On the other hand, stimulation of the Arnold nerve, the auricular branch of the vagus, triggers the cough reflex (Arnold's ear-cough reflex). Capsaicin, an agonist of Transient Receptor Potential Vanilloid 1 (TRPV1), has been shown to activate the peripheral sensory C-fibers. Stimulation of the sensory branches of the vagus in the laryngotracheal mucosa with capsaicin induces the cough reflex and has been reported to improve the swallowing function in patients with dysphagia. In our previous study, we showed that aural stimulation of the Arnold nerve with 0.025% capsaicin ointment improved the swallowing function, as evaluated by the endoscopic swallowing score, in 26 patients with dysphagia. In the present study, the video images of swallowing recorded in the previous study were re-evaluated using the SMRC scale by an independent otolaryngologist who was blinded to the information about the patients and the endoscopic swallowing score. The SMRC scale is used to evaluate four aspects of the swallowing function: 1) Sensory: the initiation of the swallowing reflex as assessed by the white-out timing; 2) Motion: the ability to hold blue-dyed water in the oral cavity and induce laryngeal elevation; 3) Reflex: glottal closure and the cough reflex induced by touching the epiglottis or arytenoid with the endoscope; 4) Clearance: pharyngeal clearance of the blue-dyed water after swallowing. Accordingly, we demonstrated that a single application of capsaicin ointment to the external auditory canal of patients with dysphagia significantly improved the R, but not the S, M or C scores, and this effect lasted for 60 min. After repeated aural stimulation with the ointment for 7 days, the R score improved significantly in patients with severe dysphagia. The present findings suggest that stimulation of the Arnold's branch of the vagus in the external auditory canal with capsaicin improves the glottal closure and cough reflex in patients with dysphagia. Thus, aural stimulation with capsaicin represents a novel treatment for dysphagia. It is also suggested that repeated alternative aural stimulation with capsaicin for a week, rather than a single application, is needed to improve the swallowing function in patients with severe dysphagia. By the same mechanism as that underlying the effect of ACE inhibitors, aural stimulation with capsaicin may reduce the incidence of aspiration pneumonia in patients with dysphagia.

Cell cycle regulation and regeneration.

Regeneration of ear punch holes in the MRL mouse and amputated limbs of the axolotl show a number of similarities. A large proportion of the fibroblasts of the uninjured MRL mouse ear are arrested in G2 of the cell cycle, and enter nerve-dependent mitosis after injury to form a ring-shaped blastema that regenerates the ear tissue. Multiple cell types contribute to the establishment of the regeneration blastema of the urodele limb by dedifferentiation, and there is substantial reason to believe that the cells of this early blastema are also arrested in G2, and enter mitosis under the influence of nerve-dependent factors supplied by the apical epidermal cap. Molecular analysis reveals other parallels, such as; (1) the upregulation of Evi5, a centrosomal protein that prevents mitosis by stabilizing Emi1, a protein that inhibits the degradation of cyclins by the anaphase promoting complex and (2) the expression of sodium channels by the epidermis. A central feature in the entry into the cell cycle by MRL ear fibroblasts is a natural downregulation of p21, and knockout of p21 in wild-type mice confers regenerative capacity on non-regenerating ear tissue. Whether the same is true for entry into the cell cycle in regenerating urodele limbs is presently unknown.

Fine-mapping quantitative trait loci affecting murine external ear tissue regeneration in the LG/J by SM/J advanced intercross line.

External ear hole closure in LG/J mice represents a model of regenerative response. It is accompanied by the formation of a blastema-like structure and the re-growth of multiple tissues, including cartilage. The ability to regenerate tissue is heritable. An F34 advanced intercross line of mice (Wustl:LG,SM-G34) was generated to identify genomic loci involved in ear hole closure over a 30-day healing period. We mapped 19 quantitative trait loci (QTL) for ear hole closure. Individual gene effects are relatively small (0.08 mm), and most loci have co-dominant effects with phenotypically intermediate heterozygotes. QTL support regions were limited to a median size of 2 Mb containing a median of 19 genes. Positional candidate genes were evaluated using differential transcript expression between LG/J and SM/J healing tissue, function analysis and bioinformatic analysis of single-nucleotide polymorphisms in and around positional candidate genes of interest. Analysis of the set of 34 positional candidate genes and those displaying expression differences revealed over-representation of genes involved in cell cycle regulation/DNA damage, cell migration and adhesion, developmentally related genes and metabolism. This indicates that the healing phenotype in LG/J mice involves multiple physiological mechanisms.

Spatial release from simultaneous echo masking in bat sonar.

Big brown bats (Eptesicus fuscus) use biosonar to navigate and locate objects in their surroundings. During natural foraging, they often encounter echoes returned by a target of interest located to the front while other, often stronger, clutter echoes are returned from objects, such as vegetation, located to the sides or above. Nevertheless, bats behave as if they do not suffer interference from this clutter. Using a two-choice delay discrimination procedure, bats were tested for the masking effectiveness of clutter echoes on target echoes when the target echoes were delivered from the bat's front while clutter echoes were delivered from 90° overhead, a direction of lowpass filtering by the external ears. When clutter echoes are presented from the front at the same delay as target echoes, detection performance declines and clutter masking occurs. When the clutter echoes are presented at the same delay but from overhead, discrimination performance is unaffected and no masking occurs. Thus there is masking release for simultaneous off-axis lowpass clutter compared to masking by simultaneous clutter from the front. The bat's performance for simultaneous target and clutter echoes indicates a new role for the mechanism that separates overlapping echoes by decomposing the bat's auditory time-frequency representation.

External and middle ear sound pressure distribution and acoustic coupling to the tympanic membrane.

Sound energy is conveyed to the inner ear by the diaphanous, cone-shaped tympanic membrane (TM). The TM moves in a complex manner and transmits sound signals to the inner ear with high fidelity, pressure gain, and a short delay. Miniaturized sensors allowing high spatial resolution in small spaces and sensitivity to high frequencies were used to explore how pressure drives the TM. Salient findings are: (1) A substantial pressure drop exists across the TM, and varies in frequency from ∼10 to 30 dB. It thus appears reasonable to approximate the drive to the TM as being defined solely by the pressure in the ear canal (EC) close to the TM. (2) Within the middle ear cavity (MEC), spatial variations in sound pressure could vary by more than 20 dB, and the MEC pressure at certain locations/frequencies was as large as in the EC. (3) Spatial variations in pressure along the TM surface on the EC-side were typically less than 5 dB up to 50 kHz. Larger surface variations were observed on the MEC-side.

Three-dimensional finite element modeling of the human external ear: simulation study of the bone conduction occlusion effect.

A linear three-dimensional (3D) elasto-acoustic finite element model was used to simulate the occlusion effect following mechanical vibration at the mastoid process. The ear canal and the surrounding soft and bony tissues were reconstructed using images of a female cadaver head (Visible Human Project(®)). The geometrical model was coupled to a 3D earplug model and imported into comsol Multiphysics (COMSOL(®), Sweden). The software was used to solve for the sound pressure at the eardrum. Finite element modeling of the human external ear and of the occlusion effect has several qualities that can complement existing measuring and modeling techniques. First, geometrically complex structures such as the external ear can be reconstructed. Second, various material behavioral laws and complex loading can be accounted for. Last, 3D analyses of external ear substructures are possible allowing for the computation of a broad range of acoustic indicators. The model simulates consistent occlusion effects (e.g., insertion depth variability). Comparison with an experimental dataset, kindly provided by Stenfelt and Reinfeldt [Int. J. Audiol. 46, 595-608 (2007)], further demonstrates the model's accuracy. Power balances were used to analyze occlusion effect differences obtained for a silicone earplug and to examine the increase in sound energy when the ear canal is occluded (e.g., high-pass filter removal).

[Mechanisms of sound reception and conduction in a dolphin].

Morphology of a lower jaw, model and behavioral experiments are discussed with the aim of exploring the mechanisms of sound reception and conduction to the dolphin's lower jaw canals taking into account known concepts of acoustics and a theory of group antennas. It is shown that the left and right row of mental foramens with the respective mandibular canal and tissues of the canals are forming the new outer ear and the new external auditory canal by which the sound (in frequency band of 0.1-160 kHz) is transmitted into the middle ear, in contrast to the dolphin's non-functional outer ear. This new external ear is created by nature as a receiving array of the traveling wave antenna located in the throat of the acoustical horn (a corresponding mandibular canal). The results give reason to assume the presence of similar new outer ear in Odontoceti.

[The threshold levels of the stapedius reflex depending on the relationship between the gas pressure in the external auditory meatus and the middle ear].

The objective of the present work was to detect and elucidate effects of a single factor (gas pressure) responsible for the threshold levels of stapedius reflex. It was shown that the threshold reflexes were recorded at 105 dB SPL when the gas pressure in the external auditory meatus decreased by 150-200 pPa against the atmospheric pressure and when it increased by 100-150 pPa. These effects can be accounted for by the reduced compliance of the tympanic membrane and different changes in the state of the auditory ossicles undergoing compression and tension. When entering the most comfortable stapedial reflex levels into the patient's tuning chart, an account should be taken of the maximum value of tympanic membrane compliance.