Tiny spies: mosquito antennae are sensitive sensors for eavesdropping on frog calls.

Most mosquito and midge species use hearing during acoustic mating behaviors. For frog-biting species, however, hearing plays an important role beyond mating as females rely on anuran calls to obtain blood meals. Despite the extensive work examining hearing in mosquito species that use sound in mating contexts, our understanding of how mosquitoes hear frog calls is limited. Here, we directly investigated the mechanisms underlying detection of frog calls by a mosquito species specialized on eavesdropping on anuran mating signals: Uranotaenia lowii. Behavioral, biomechanical and neurophysiological analyses revealed that the antenna of this frog-biting species can detect frog calls by relying on neural and mechanical responses comparable to those of non-frog-biting species. Our findings show that in Ur. lowii, contrary to most species, males do not use sound for mating, but females use hearing to locate their anuran host. We also show that the response of the antennae of this frog-biting species resembles that of the antenna of species that use hearing for mating. Finally, we discuss our data considering how mosquitoes may have evolved the ability to tap into the communication system of frogs.

Outsourced hearing in an orb-weaving spider that uses its web as an auditory sensor.

SignificanceThe sense of hearing in all known animals relies on possessing auditory organs that are made up of cellular tissues and constrained by body sizes. We show that hearing in the orb-weaving spider is functionally outsourced to its extended phenotype, the proteinaceous self-manufactured web, and hence processes behavioral controllability. This finding opens new perspectives on animal extended cognition and hearing-the outsourcing and supersizing of auditory function in spiders. This study calls for reinvestigation of the remarkable evolutionary ecology and sensory ecology in spiders-one of the oldest land animals. The sensory modality of outsourced hearing provides a unique model for studying extended and regenerative sensing and presents new design features for inspiring novel acoustic flow detectors.

The Long and Short of Hearing in the Mosquito Aedes aegypti.

Mating behavior in Aedes aegypti mosquitoes occurs mid-air and involves the exchange of auditory signals at close range (millimeters to centimeters) [1-6]. It is widely assumed that this intimate signaling distance reflects short-range auditory sensitivity of their antennal hearing organs to faint flight tones [7, 8]. To the contrary, we show here that male mosquitoes can hear the female's flight tone at surprisingly long distances-from several meters to up to 10 m-and that unrestrained, resting Ae. aegypti males leap off their perches and take flight when they hear female flight tones. Moreover, auditory sensitivity tests of Ae. aegypti's hearing organ, made from neurophysiological recordings of the auditory nerve in response to pure-tone stimuli played from a loudspeaker, support the behavioral experiments. This demonstration of long-range hearing in mosquitoes overturns the common assumption that the thread-like antennal hearing organs of tiny insects are strictly close-range ears. The effective range of a hearing organ depends ultimately on its sensitivity [9-13]. Here, a mosquito's antennal ear is shown to be sensitive to sound levels down to 31 dB sound pressure level (SPL), translating to air particle velocity at nanometer dimensions. We note that the peak of energy of the first formant of the vowels of the human speech spectrum range from about 200-1,000 Hz and is typically spoken at 45-70 dB SPL; together, they lie in the sweet spot of mosquito hearing. VIDEO ABSTRACT.

Sensing fluctuating airflow with spider silk.

The ultimate aim of flow sensing is to represent the perturbations of the medium perfectly. Hundreds of millions of years of evolution resulted in hair-based flow sensors in terrestrial arthropods that stand out among the most sensitive biological sensors known, even better than photoreceptors which can detect a single photon (10-18-10-19 J) of visible light. These tiny sensory hairs can move with a velocity close to that of the surrounding air at frequencies near their mechanical resonance, despite the low viscosity and low density of air. No man-made technology to date demonstrates comparable efficiency. Here we show that nanodimensional spider silk captures fluctuating airflow with maximum physical efficiency (Vsilk/Vair ∼ 1) from 1 Hz to 50 kHz, providing an effective means for miniaturized flow sensing. Our mathematical model shows excellent agreement with experimental results for silk with various diameters: 500 nm, 1.6 µm, and 3 µm. When a fiber is sufficiently thin, it can move with the medium flow perfectly due to the domination of forces applied to it by the medium over those associated with its mechanical properties. These results suggest that the aerodynamic property of silk can provide an airborne acoustic signal to a spider directly, in addition to the well-known substrate-borne information. By modifying a spider silk to be conductive and transducing its motion using electromagnetic induction, we demonstrate a miniature, directional, broadband, passive, low-cost approach to detect airflow with full fidelity over a frequency bandwidth that easily spans the full range of human hearing, as well as that of many other mammals.

Motor and mechanical bases of the courtship call of the male treehopper Umbonia crassicornis.

This study is a physiological, anatomical and biophysical analysis of how plant-borne vibrational signals are produced by the treehopper Umbonia crassicornis During courtship, males and females engage in a vibrational duet, with each producing a characteristic call. For males, this consists of a frequency-modulated tonal signal which is accompanied by rhythmic broad-band clicks. Although previous studies have described these complex signals in detail, little is known about how they are produced. By combining video recordings, electromyograms, dissections and mechanical modeling, we describe the mechanism by which the male produces his courtship signal. High-speed videos show that the tonal portion of the call is produced by periodic dorso-ventral movements of the abdomen, with a relatively large amplitude oscillation alternating with a smaller oscillation. Electromyograms from the muscles we identified that produce this motion reveal that they fire at half the frequency of the abdominal oscillation, throughout the frequency modulation of the tonal signal. Adding weight to the abdomen of a calling male reduces the frequency of motion, demonstrating that the abdominal motion is strongly influenced by its mechanical resonance. A mathematical model accounting for this resonance provides excellent qualitative agreement with measurements of both the muscle firing rate recorded electrophysiologically and the oscillatory motion of the abdomen as recorded in the high-speed video. The model, electromyograms and analysis of video recordings further suggest that the frequency modulation of the abdominal response is due to a simultaneous modulation in the muscle firing rate and a fluctuation in stiffness of the abdominal attachment.

Airborne Acoustic Perception by a Jumping Spider.

Jumping spiders (Salticidae) are famous for their visually driven behaviors [1]. Here, however, we present behavioral and neurophysiological evidence that these animals also perceive and respond to airborne acoustic stimuli, even when the distance between the animal and the sound source is relatively large (∼3 m) and with stimulus amplitudes at the position of the spider of ∼65 dB sound pressure level (SPL). Behavioral experiments with the jumping spider Phidippus audax reveal that these animals respond to low-frequency sounds (80 Hz; 65 dB SPL) by freezing-a common anti-predatory behavior characteristic of an acoustic startle response. Neurophysiological recordings from auditory-sensitive neural units in the brains of these jumping spiders showed responses to low-frequency tones (80 Hz at ∼65 dB SPL)-recordings that also represent the first record of acoustically responsive neural units in the jumping spider brain. Responses persisted even when the distances between spider and stimulus source exceeded 3 m and under anechoic conditions. Thus, these spiders appear able to detect airborne sound at distances in the acoustic far-field region, beyond the near-field range often thought to bound acoustic perception in arthropods that lack tympanic ears (e.g., spiders) [2]. Furthermore, direct mechanical stimulation of hairs on the patella of the foreleg was sufficient to generate responses in neural units that also responded to airborne acoustic stimuli-evidence that these hairs likely play a role in the detection of acoustic cues. We suggest that these auditory responses enable the detection of predators and facilitate an acoustic startle response. VIDEO ABSTRACT.

The analysis of plane discontinuities in offset cylindrical waveguides.

This paper applies the re-expansion method for analyzing the effects on the sound field due to planar discontinuities at the junction of two offset circular acoustic waveguides. The normal modes in the two waveguides are expanded at the junction plane into a system of functions accounting for velocity singularities at the corner points. As the new expansion has a high convergence order, only a few terms have to be considered for obtaining the solution of most practical problems. This paper gives the equivalent impedance accounting for nonplanar waves into a plane-wave analysis. The last section of the paper applies the re-expansion technique to the case of two offset pipes (step discontinuity) and to that of an offset aperture in a cylindrical pipe (diaphragm-type discontinuity). The plots of the discontinuity inductance, characterized by Karal's factor, are quite similar but the values in the second case are 1.5-2 times larger, showing that the diaphragm-type discontinuity excites much more nonplanar evanescent modes than the step discontinuity.

Viscous damping and spring force calculation of regularly perforated MEMS microstructures in the Stokes' approximation.

There are a number of applications for microstructure devices consisting of a regular pattern of perforations, and many of these utilize fluid damping. For the analysis of viscous damping and for calculating the spring force in some cases, it is possible to take advantage of the regular hole pattern by assuming periodicity. Here a model is developed to determine these quantities based on the solution of the Stokes' equations for the air flow. Viscous damping is directly related to thermal-mechanical noise. As a result, the design of perforated microstructures with minimal viscous damping is of real practical importance. A method is developed to calculate the damping coefficient in microstructures with periodic perforations. The result can be used to minimize squeeze film damping. Since micromachined devices have finite dimensions, the periodic model for the perforated microstructure has to be associated with the calculation of some frame (edge) corrections. Analysis of the edge corrections has also been performed. Results from analytical formulas and numerical simulations match very well with published measured data.

Re-expansion method for circular waveguide discontinuities: application to concentric expansion chambers.

The paper applies the re-expansion method for analyzing planar discontinuities at the junction of two axi-symmetrical circular waveguides. The normal modes in the two waveguides are expanded at the junction plane into a system of functions accounting for velocity singularities at the corner points. As the new expansion has a high convergence order, only a few terms have to be considered for obtaining the solution of most practical problems. This paper gives the equivalent impedance accounting for nonplanar waves into a plane-wave analysis and also the scattering matrix describing the coupling of arbitrary modes at each side of the discontinuity valid in the case of many propagating modes in both sides of the duct. The last section applies the re-expansion technique to some concentric expansion chambers providing an explicit formula for the transmission loss coefficient.

An analytical-numerical method for determining the mechanical response of a condenser microphone.

The paper is based on determining the reaction pressure on the diaphragm of a condenser microphone by integrating numerically the frequency domain Stokes system describing the velocity and the pressure in the air domain beneath the diaphragm. Afterwards, the membrane displacement can be obtained analytically or numerically. The method is general and can be applied to any geometry of the backplate holes, slits, and backchamber. As examples, the method is applied to the Bruel & Kjaer (B&K) 4134 1/2-inch microphone determining the mechanical sensitivity and the mechano-thermal noise for a domain of frequencies and also the displacement field of the membrane for two specified frequencies. These elements compare well with the measured values published in the literature. Also a new design, completely micromachined (including the backvolume) of the B&K micro-electro-mechanical systems (MEM) 1/4-inch measurement microphone is proposed. It is shown that its mechanical performances are very similar to those of the B&K MEMS measurement microphone.

A re-expansion method for determining the acoustical impedance and the scattering matrix for the waveguide discontinuity problem.

The paper gives a new method for analyzing planar discontinuities in rectangular waveguides. The method consists of a re-expansion of the normal modes in the two ducts at the junction plane into a system of functions accounting for the velocity singularities at the corner points. As the new expansion has an exponential convergence, only a few terms have to be considered for obtaining the solution of most practical problems. To see how the method works some closed form solutions, obtained by the conformal mapping method, are used to discuss the convergence of the re-expanded series when the number of retained terms increases. The equivalent impedance accounting for nonplanar waves into a plane-wave analysis is determined. Finally, the paper yields the scattering matrix which describes the coupling of arbitrary modes at each side of the discontinuity valid in the case of many propagating modes in both parts of the duct.

Viscous damping and spring force in periodic perforated planar microstructures when the Reynolds' equation cannot be applied.

A model of squeeze-film behavior is developed based on Stokes' equations for viscous, compressible isothermal flows. The flow domain is an axisymmetrical, unit cell approximation of a planar, periodic, perforated microstructure. The model is developed for cases when the lubrication approximation cannot be applied. The complex force generated by vibrations of the diaphragm driving the flow has two components: the damping force and the spring force. While for large frequencies the spring force dominates, at low (acoustical) frequencies the damping force is the most important part. The analytical approach developed here yields an explicit formula for both forces. In addition, using a finite element software package, the damping force is also obtained numerically. A comparison is made between the analytic result, numerical solution, and some experimental data found in the literature, which validates the analytic formula and provides compelling arguments about its value in designing microelectomechanical devices.

Mechanical peeling of free-standing single-walled carbon-nanotube bundles.

An in situ electron microscopy study is presented of adhesion interactions between single-walled carbon nanotubes (SWNTs) by mechanically peeling thin free-standing SWNT bundles using in situ nanomanipulation techniques inside a high-resolution scanning electron microscope. The in situ measurements clearly reveal the process of delaminating one SWNT bundle from its originally bound SWNT bundle in a controlled-displacement manner and capture the deformation curvature of the delaminated SWNT bundle during the peeling process. A theoretical model based on nonlinear elastica theory is employed to interpret the measured deformation curvatures of the SWNTs and to quantitatively evaluate the peeling force and the adhesion strength between bundled SWNTs. The estimated adhesion energy per unit length for each pair of neighboring tubes in the peeling interface based on our peeling experiments agrees reasonably well with the theoretical value. This in situ peeling technique provides a potential new method for separating bundled SWNTs without compromising their material properties. The combined peeling experiments and modeling presented in this paper will be very useful to the study of the adhesion interactions between SWNTs and their nonlinear mechanical behaviors in the large-displacement regime.

Analytical model for viscous damping and the spring force for perforated planar microstructures acting at both audible and ultrasonic frequencies.

The paper presents a model for the squeezed film damping, the resistance of the holes, and the corresponding spring forces for a periodic perforated microstructure including the effects of compressibility, inertia, and rarefied gas. The viscous damping and spring forces are obtained by using the continuity equation. The analytical formula for the squeezed film damping is applied to analyze the response of an ultrasonic transducer. The inclusion of these effects in a model significantly improves the agreement with measured results. Finally, it is shown that the frequency dependence of the total damping and total spring force for a cell are very similar to those corresponding to a rectangular open microstructure without holes. A separate analysis reveals the importance of each particular correction. The most important is the compressibility correction; the inertia has to be considered only for determining the spring force and the damping force for sufficiently high frequencies.

Influence of viscosity on the reflection and transmission of an acoustic wave by a periodic array of screens. The general 3-D problem.

An analysis is presented of the diffraction of a pressure wave by a periodic grating including the influence of the air viscosity. The direction of the incoming pressure wave is arbitrary. As opposed to the classical nonviscous case, the problem cannot be reduced to a plane problem having a definite 3-D character. The system of partial differential equations used for solving the problem consists of the compressible Navier-Stokes equations associated with no-slip boundary conditions on solid surfaces. The problem is reduced to a system of two hypersingular integral equations for determining the velocity components in the slits' plane and a hypersingular integral equation for the normal component of velocity. These equations are solved by using Galerkin's method with some special trial functions. The results can be applied in designing protective screens for miniature microphones realized in MEMS technology. In this case, the physical dimensions of the device are on the order of the viscous boundary layer so that the viscosity cannot be neglected. The analysis indicates that the openings in the screen should be on the order of 10 microns in order to avoid excessive attenuation of the signal. This paper also provides the variation of the transmission coefficient with frequency in the acoustical domain.

Viscous microstructural dampers with aligned holes: design procedure including the edge correction.

The paper is a continuation of the works "Modelling of viscous damping of perforated planar micromechanical structures. Applications in acoustics" [Homentcovschi and Miles, J. Acoust. Soc. Am. 116, 2939-2947 (2004)] and "Viscous Damping of Perforated Planar Micromechanical Structures" [Homentcovschi and Miles, Sensors Actuators, A119, 544-552 (2005)] where design formulas for the case of an offset (staggered) system of holes was provided. The present work contains design formulas for perforated planar microstructures used in MEMS devices (such as proof-masses in accelerometers, backplates in microphones, micromechanical switches, resonators, tunable microoptical interferometers, etc.) in the case of aligned (nonstaggered) holes of circular and square section. The given formulas assure a minimum total damping coefficient (including the squeeze film damping and the direct and indirect resistance of the holes) for an assigned open area. The paper also gives a simple edge correction, making it possible to consider real (finite) perforated planar microstructures. The proposed edge correction is validated by comparison with the results obtained by FEM simulations: the relative error is found to be smaller than 0.04%. By putting together the design formulas with the edge correction a simple integrated design procedure for obtaining viscous perforated dampers with assigned properties is obtained.

A boundary integral approach to analyze the viscous scattering of a pressure wave by a rigid body.

The paper provides boundary integral equations for solving the problem of viscous scattering of a pressure wave by a rigid body. By using this mathematical tool uniqueness and existence theorems are proved. Since the boundary conditions are written in terms of velocities, vector boundary integral equations are obtained for solving the problem. The paper introduces single-layer viscous potentials and also a stress tensor. Correspondingly, a viscous double-layer potential is defined. The properties of all these potentials are investigated.By representing the scattered field as a combination of a single-layer viscous potential and a double-layer viscous potential the problem is reduced to the solution of a singular vectorial integral equation of Fredholm type of the second kind.In the case where the stress vector on the boundary is the main quantity of interest the corresponding boundary singular integral equation is proved to have a unique solution.

A method for two-dimensional characterization of animal vibrational signals transmitted along plant stems.

Conventional approaches to measuring animal vibrational signals on plant stems use a single transducer to measure the amplitude of vibrations. Such an approach, however, will often underestimate the amplitude of bending waves traveling along the stem. This occurs because vibration transducers are maximally sensitive along a single axis, which may not correspond to the major axis of stem motion. Furthermore, stem motion may be more complex than that of a bending wave propagating along a single axis, and such motion cannot be described using a single transducer. Here, we describe a method for characterizing stem motion in two dimensions by processing the signals from two orthogonally positioned transducers. Viewed relative to a cross-sectional plane, a point on the stem surface moves in an ellipse at any one frequency, with the ellipse's major axis corresponding to the maximum amplitude of vibration. The method outlined here measures the ellipse's major and minor axes, and its angle of rotation relative to one of the transducers. We illustrate this method with measurements of stem motion during insect vibrational communication. It is likely the two-dimensional nature of stem motion is relevant to insect vibration perception, making this method a promising avenue for studies of plant-borne transmission.

A two-dimensional model of a directional microphone: calculation of the normal force and moment on the diaphragm.

It has been shown that the parasitoid fly Ormia Ochracea exhibits exceptional sound localization ability achieved through the mechanical coupling of its eardrums [R. N. Miles et al., J. Acoust. Soc. Am. 98, 3059-3070 (1995)]. Based on this biological system a new directional microphone has been designed, having as a basic element a special diaphragm undergoing a rocking motion. This paper considers a 2D model of the microphone in which the diaphragm is considered as a 2D plate having slits on the sides. The slits lead to a backing volume limited by an infinite rigid wall parallel to the diaphragm in its neutral position. The reflection and diffraction of an incoming plane wave by this system are studied to determine the resultant force and resultant moment of pressure upon the diaphragm. The results show that such a microphone will be driven better in the case of narrow slits and deep cavities.

Viscous scattering of a pressure wave: calculation of the fluid tractions on a biomimetic acoustic velocity sensor.

In the paper we give a method for calculating the tractions (local forces) of the fluid motion determined by an incoming plane pressure wave on an artificial hair cell transducer structure. The sensing element of the transducer is a standing high aspect ratio cilium in the shape of a narrow thin curved beam (tape-like), which can be easily fabricated in micro-/nanotechnology. The method is based on considering the system of partial differential equations describing the motion of the compressible viscous fluid in an acoustic linearized approximation, and representation of the velocity field as a viscous acoustic single-layer potential. The boundary conditions, stating the cancellation of the velocity components on the solid beam, yield a two-dimensional (2-D) system of three integral equations over the beam's surface for the traction components. In the case of a narrow cilium, the system of integral equations furnishes a system of two 1-D integral equations over the symmetry curve of the structure for obtaining the tangential and normal components of the traction. This system is solved numerically by a finite (boundary) element method. The numerical code written for solving the problem was applied to some particular structures. The last structure is similar to the trichobothrium of a spider Cupiennius salei. The results obtained show that the curvature of the hair is enhancing sensitivity to flows directed normal to the main shaft of the hair confirming the assertion of Barth et al. [Philos. Trans. R. Soc. London, Ser. B 340, 445-461 (1993)].