Bone conduction pathways confer directional cues to salamanders.

Sound and vibration are generated by mechanical disturbances within the environment, and the ability to detect and localize these acoustic cues is generally important for survival, as suggested by the early emergence of inherently directional otolithic ears in vertebrate evolutionary history. However, fossil evidence indicates that the water-adapted ear of early terrestrial tetrapods lacked specialized peripheral structures to transduce sound pressure (e.g. tympana). Therefore, early terrestrial hearing should have required nontympanic (or extratympanic) mechanisms for sound detection and localization. Here, we used atympanate salamanders to investigate the efficacy of extratympanic pathways to support directional hearing in air. We assessed peripheral encoding of directional acoustic information using directionally masked auditory brainstem response recordings. We used laser Doppler vibrometry to measure the velocity of sound pressure-induced head vibrations as a key extratympanic mechanism for aerial sound reception in atympanate species. We found that sound generates head vibrations that vary with the angle of the incident sound. This extratympanic pathway for hearing supports a figure-eight pattern of directional auditory sensitivity to airborne sound in the absence of a pressure-transducing tympanic ear.

Seismic sensitivity and bone conduction mechanisms enable extratympanic hearing in salamanders.

The tympanic middle ear is an adaptive sensory novelty that evolved multiple times in all the major terrestrial tetrapod groups to overcome the impedance mismatch generated when aerial sound encounters the air-skin boundary. Many extant tetrapod species have lost their tympanic middle ears, yet they retain the ability to detect airborne sound. In the absence of a functional tympanic ear, extratympanic hearing may occur via the resonant qualities of air-filled body cavities, sensitivity to seismic vibration, and/or bone conduction pathways to transmit sound from the environment to the ear. We used auditory brainstem response recording and laser vibrometry to assess the contributions of these extratympanic pathways for airborne sound in atympanic salamanders. We measured auditory sensitivity thresholds in eight species and observed sensitivity to low-frequency sound and vibration from 0.05-1.2 kHz and 0.02-1.2 kHz, respectively. We determined that sensitivity to airborne sound is not facilitated by the vibrational responsiveness of the lungs or mouth cavity. We further observed that, although seismic sensitivity probably contributes to sound detection under naturalistic scenarios, airborne sound stimuli presented under experimental conditions did not produce vibrations detectable to the salamander ear. Instead, threshold-level sound pressure is sufficient to generate translational movements in the salamander head, and these sound-induced head vibrations are detectable by the acoustic sensors of the inner ear. This extratympanic hearing mechanism mediates low-frequency sensitivity in vertebrate ears that are unspecialized for the detection of aerial sound pressure, and may represent a common mechanism for terrestrial hearing across atympanic tetrapods.

The excitation of solar-like oscillations in a δSct star by efficient envelope convection.

Delta Scuti (δSct) stars are opacity-driven pulsators with masses of 1.5-2.5 M⊙, their pulsations resulting from the varying ionization of helium. In less massive stars such as the Sun, convection transports mass and energy through the outer 30 per cent of the star and excites a rich spectrum of resonant acoustic modes. Based on the solar example, with no firm theoretical basis, models predict that the convective envelope in δSct stars extends only about 1 per cent of the radius, but with sufficient energy to excite solar-like oscillations. This was not observed before the Kepler mission, so the presence of a convective envelope in the models has been questioned. Here we report the detection of solar-like oscillations in the δSct star HD187547, implying that surface convection operates efficiently in stars about twice as massive as the Sun, as the ad hoc models predicted.

How Internally Coupled Ears Generate Temporal and Amplitude Cues for Sound Localization.

In internally coupled ears, displacement of one eardrum creates pressure waves that propagate through air-filled passages in the skull and cause displacement of the opposing eardrum, and conversely. By modeling the membrane, passages, and propagating pressure waves, we show that internally coupled ears generate unique amplitude and temporal cues for sound localization. The magnitudes of both these cues are directionally dependent. The tympanic fundamental frequency segregates a low-frequency regime with constant time-difference magnification from a high-frequency domain with considerable amplitude magnification.

Is human underwater hearing mediated by bone conduction?

In-air and underwater audiograms and directional hearing abilities were measured in humans. The lowest underwater thresholds were 2.8 µW/m2 or 3.6 mPa at a frequency of 500 Hz. The underwater hearing thresholds were 4-26 dB and 40-62 dB higher than in-air hearing thresholds when measured in intensity and pressure units, respectively. This difference is considerably smaller than what has been reported earlier. At frequencies below 1 kHz, when measured in units of particle velocity, the underwater threshold was much lower than published bone conduction thresholds, suggesting that underwater hearing is not always mediated by bone conduction pathways to the inner ear, as previously thought. We suggest it is the resonance of air in the air-filled middle ear that produces the low underwater thresholds, at least at frequencies below 1 kHz. The ability to determine the direction of a 700 Hz underwater sound source while being blindfolded was extremely poor, with submerged test subjects showing only coarse directional hearing abilities at azimuths of less than 50˚. The physical cues to sound direction are different in air and water, and the poor directional hearing abilities indicate that, in spite of low hearing thresholds, humans have no special adaptations to process directional acoustic cues under water.

Seismology of the sun.

Oscillations of the sun make it possible to probe the inside of a star. The frequencies of the oscillations have already provided measures of the sound speed and the rate of rotation throughout much of the solar interior. These quantities are important for understanding the dynamics of the magnetic cycle and have a bearing on testing general relativity by planetary precession. The oscillation frequencies yield a helium abundance that is consistent with cosmology, but they reinforce the severity of the neutrino problem. They should soon provide an important standard by which to calibrate the theory of stellar evolution.

Asteroseismic detection of latitudinal differential rotation in 13 Sun-like stars.

The differentially rotating outer layers of stars are thought to play a role in driving their magnetic activity, but the underlying mechanisms that generate and sustain differential rotation are poorly understood. We report the measurement using asteroseismology of latitudinal differential rotation in the convection zones of 40 Sun-like stars. For the most significant detections, the stars' equators rotate approximately twice as fast as their midlatitudes. The latitudinal shear inferred from asteroseismology is much larger than predictions from numerical simulations.

Hot super-Earths stripped by their host stars.

Simulations predict that hot super-Earth sized exoplanets can have their envelopes stripped by photoevaporation, which would present itself as a lack of these exoplanets. However, this absence in the exoplanet population has escaped a firm detection. Here we demonstrate, using asteroseismology on a sample of exoplanets and exoplanet candidates observed during the Kepler mission that, while there is an abundance of super-Earth sized exoplanets with low incident fluxes, none are found with high incident fluxes. We do not find any exoplanets with radii between 2.2 and 3.8 Earth radii with incident flux above 650 times the incident flux on Earth. This gap in the population of exoplanets is explained by evaporation of volatile elements and thus supports the predictions. The confirmation of a hot-super-Earth desert caused by evaporation will add an important constraint on simulations of planetary systems, since they must be able to reproduce the dearth of close-in super-Earths.

Basic response characteristics of auditory nerve fibers in the grassfrog (Rana temporaria).

Responses to free-field sound of 401 fibers from the VIIIth nerve of the grassfrog, Rana temporaria, are described. The spontaneous activities of the fibers ranged from 0 to 75 spikes/s, showing only weak correlation with frequency or sensitivity of the fibers. The highest spontaneous activities were approximately twice as high as reported previously for frogs. Best frequencies ranged from 100 to 1600 Hz and thresholds ranged from 21 to 80 dB SPL. The median dynamic range was 20 dB and the slopes of the rate-level curves ranged from 5 to 20 spikes/(s-dB). Most of the units showed post-excitatory suppression (PS) of their spontaneous activity. The duration of PS increased with sound level, also in fibers showing a decrease in firing rate at high intensities. Most fibers showing one-tone suppression did not show PS at their best suppression frequencies. Strong suppression was observed also in very phasic cells giving one spike per stimulation. Therefore, the mechanism underlying PS is probably different from that underlying adaptation. The sharpening of the neural encoding of temporal parameters and the strong encoding of sound offset as well as onset caused by PS very likely is biologically important.

In vitro and in vivo responses of saccular and caudal nucleus neurons in the grassfrog (Rana temporaria).

We present results from in vitro and in vivo studies of response properties of neurons in the saccular and caudal nuclei in the frog. In the in vitro studies the saccular nerve of the isolated brain was stimulated with electrical pulses. In the in vivo experiments, the neurons were stimulated by dorso-ventral vibrations of the intact animal. We identified six response types: (1) primary-like cells with short latencies and follow repetition rates up to 100 Hz; (2) phasic cells responding only to the first pulse in a train; (3) bursting cells firing several spikes in response to any stimulation; (4) late responders with very long latencies; (5) integrator cells showing facilitated responses, and (6) inhibitory cells inhibited by saccular nerve stimulation. The cells have comparable sensitivity and frequency characteristics to the primary fibres (BF 10-80 Hz, thresholds from 0.01 cm/s2) and enable a sophisticated analysis of vibrational stimuli.

Kepler detected gravity-mode period spacings in a red giant star.

Stellar interiors are inaccessible through direct observations. For this reason, helioseismologists made use of the Sun's acoustic oscillation modes to tune models of its structure. The quest to detect modes that probe the solar core has been ongoing for decades. We report the detection of mixed modes penetrating all the way to the core of an evolved star from 320 days of observations with the Kepler satellite. The period spacings of these mixed modes are directly dependent on the density gradient between the core region and the convective envelope.

HD 181068: a red giant in a triply eclipsing compact hierarchical triple system.

Hierarchical triple systems comprise a close binary and a more distant component. They are important for testing theories of star formation and of stellar evolution in the presence of nearby companions. We obtained 218 days of Kepler photometry of HD 181068 (magnitude of 7.1), supplemented by ground-based spectroscopy and interferometry, which show it to be a hierarchical triple with two types of mutual eclipses. The primary is a red giant that is in a 45-day orbit with a pair of red dwarfs in a close 0.9-day orbit. The red giant shows evidence for tidally induced oscillations that are driven by the orbital motion of the close pair. HD 181068 is an ideal target for studies of dynamical evolution and testing tidal friction theories in hierarchical triple systems.

Ensemble asteroseismology of solar-type stars with the NASA Kepler mission.

In addition to its search for extrasolar planets, the NASA Kepler mission provides exquisite data on stellar oscillations. We report the detections of oscillations in 500 solar-type stars in the Kepler field of view, an ensemble that is large enough to allow statistical studies of intrinsic stellar properties (such as mass, radius, and age) and to test theories of stellar evolution. We find that the distribution of observed masses of these stars shows intriguing differences to predictions from models of synthetic stellar populations in the Galaxy.

Kepler's optical phase curve of the exoplanet HAT-P-7b.

Ten days of photometric data were obtained during the commissioning phase of the Kepler mission, including data for the previously known giant transiting exoplanet HAT-P-7b. The data for HAT-P-7b show a smooth rise and fall of light from the planet as it orbits its star, punctuated by a drop of 130 +/- 11 parts per million in flux when the planet passes behind its star. We interpret this as the phase variation of the dayside thermal emission plus reflected light from the planet as it orbits its star and is occulted. The depth of the occultation is similar in photometric precision to the detection of a transiting Earth-size planet for which the mission was designed.

Biophysics of underwater hearing in the clawed frog, Xenopus laevis.

Anesthetized clawed frogs (Xenopus laevis) were stimulated with underwater sound and the tympanic disk vibrations were studied using laser vibrometry. The tympanic disk velocities ranged from 0.01 to 0.5 mm/s (at a sound pressure of 2 Pa) in the frequency range of 0.4-4 kHz and were 20-40 dB higher than those of the surrounding tissue. The frequency response of the disk had two peaks, in the range of 0.6-1.1 kHz and 1.6-2.2 kHz, respectively. The first peak corresponded to the peak vibrations of the body wall overlying the lung. The second peak matched model predictions of the pulsations of the air bubble in the middle ear cavity. Filling the middle ear cavity with water lowered the disk vibrations by 10-30 dB in the frequency range of 0.5-3 kHz. Inflating the lungs shifted the low-frequency peak downwards, but did not change the high-frequency peak. Thus, the disk vibrations in the frequency range of the mating call (main energy at 1.7-1.9 kHz) were mainly caused by pulsations of the air in the middle ear cavity; sound transmission via the lungs was more important at low frequencies (below 1 kHz). Furthermore, the low-frequency peak could be reversibly reduced in amplitude by loading the larynx with metal or tissue glue. This shows that the sound-induced vibrations of the lungs are probably coupled to the middle ear cavities via the larynx. Also, anatomical observations show that the two middle ear cavities and the larynx are connected in an air-filled recess in submerged animals.(ABSTRACT TRUNCATED AT 250 WORDS)

One-tone suppression in the frog auditory nerve.

Sixty-seven fibers of a sample of 401 in the auditory nerve of grassfrogs (Rana temporaria) showed one-tone suppression, i.e., their spontaneous activity was suppressed by tones. All fibers were afferents from the amphibian papilla with best frequencies between 100 and 400 Hz. Best suppression frequencies ranged from 700 to 1200 Hz. Spontaneous activities for the fibers showing one-tone suppression ranged from 3 to 75 spikes/s. Spontaneous activities above 40 spikes/s and the phenomenon of one-tone suppression itself has not been reported previously for frogs. The population of fibers showing one-tone suppression comprises 81% of all fibers with best frequencies below 400 Hz and spontaneous activities higher than 3 spikes/s, indicating that the mechanism underlying the suppression is quite general.

The response characteristics of vibration-sensitive saccular fibers in the grassfrog, Rana temporaria.

The response characteristics of saccular nerve fibers in European grassfrogs (Rana temporaria) subjected to dorso-ventral, 10-200 Hz sinusoidal vibrations were studied. Only 4 fibers out of a total of 129 did not respond to the vibrations. 70 fibers had an irregular spontaneous activity of 2-48 spikes/s. These fibers were very vibration-sensitive. The synchronization thresholds at 10-20 Hz varied from below 0.005 to 0.02 cm/s2. In contrast to earlier results, all these fibers had low-pass characteristics (with respect to acceleration) and responded maximally at 10 and 20 Hz. 55 fibers had spontaneous activities from 0-2 spikes/s. These fibers were less sensitive than the fibers with higher spontaneous activity. The spike-rate thresholds varied from about 0.04 to above 1.28 cm/s2, giving a considerable range fractionation. Most of these fibers also had low-pass characteristics with respect to acceleration, but 8 fibers showed band-pass characteristics with maximal synchronizations and spike-rates occurring at 40-80 Hz. At high acceleration levels, most spikes fell within 5-10 degrees of the stimulus cycle. The phase-locking of the saccular fibers is therefore very acute at low frequencies. The phase angles preferred by the fibers at 10 Hz were bimodally distributed with the two peaks about 180 degrees apart. This finding probably reflects the morphological observation that the saccular macula contains two oppositely oriented hair-cell populations. The results also indicate that the actual motion of the otolith relative to the macula is complex. No behavioral role of a vibration receptor has been demonstrated in the grassfrog.(ABSTRACT TRUNCATED AT 250 WORDS)

Sound and vibration sensitivity of VIIIth nerve fibers in the frogs Leptodactylus albilabris and Rana pipiens pipiens.

1. Responses of 73 fibers to dorso-ventral vibration were recorded in the saccular and utricular branchlets of Rana pipiens pipiens using a ventral approach. The saccular branchlet contained nearly exclusively vibration-sensitive fibers (33 out of 36) with best frequencies (BFs) between 10 and 70 Hz, whereas none of the 37 fibers encountered in the utricular branchlet responded to dorso-ventral vibrations. 2. Using a dorsal approach we recorded from the VIIIth nerve near its entry in the brainstem and analyzed responses to both sound and vibration stimuli for 65 fibers in R. pipiens pipiens and 25 fibers in Leptodactylus albilabris. The fibers were classified as amphibian papilla (AP), basilar papilla (BP), saccular or vestibular fibers based on their location in the nerve. Only AP and saccular fibers responded to vibrations. The AP-fibers responded to vibrations from 0.01 cm/s2 and to sound from 40 dB SPL by increasing their spike rate. Best frequencies (BFs) ranged from 60 to 900 Hz, and only fibers with BFs below 500 Hz responded to vibrations. The fibers had identical BF's for sound and vibration. The saccular fibers had BFs ranging from 10 to 80 Hz with 22 fibers having BFs at 40-50 Hz. The fibers responded to sound from 70 dB SPL and to vibrations from 0.01 cm/s2. 3. No differences in sensitivity, tuning or phase-locking were found between the two species, except that most BP-fibers in R. pipiens pipiens had BFs from 1.2 to 1.4 kHz, whereas those in L. albilabris had BFs from 2.0 to 2.2 kHz (matching the energy peak of L. albilabris' mating call). 4. The finding that the low-frequency amphibian papilla fibers are extremely sensitive to vibrations raises questions regarding their function in the behaving animal. They may be substrate vibration receptors, respond to sound-induced vibrations or bone-conducted sound.

Directionality of auditory nerve fiber responses to pure tone stimuli in the grassfrog, Rana temporaria. I. Spike rate responses.

We studied the directionality of spike rate responses of auditory nerve fibers of the grassfrog, Rana temporaria, to pure tone stimuli. All auditory fibers showed spike rate directionality. The strongest directionality was seen at low frequencies (200-400 Hz), where the spike rate could change by up to nearly 200 spikes s-1, with sound direction. At higher frequencies the directional spike rate changes were mostly below 100 spikes s-1. In equivalent dB SPL terms (calculated using the fibers' rate-intensity curves) the maximum directionalities were up to 15 dB at low frequencies and below 10 dB at higher frequencies. Two types of directional patterns were observed. At frequencies below 500 Hz relatively strong responses were evoked by stimuli from the ipsilateral (+90 degrees) and contralateral (-90 degrees) directions while the weakest responses were evoked by stimuli from frontal (0 degree or +30 degrees) or posterior (-135 degrees) directions. At frequencies above 800 Hz the strongest responses were evoked by stimuli from the ipsilateral direction while gradually weaker responses were seen as the sound direction shifted towards the contralateral side. At frequencies between 500 and 800 Hz both directional patterns were seen. The directionality was highly intensity dependent. No special adaptations for localization of conspecific calls were found.

Directionality of auditory nerve fiber responses to pure tone stimuli in the grassfrog, Rana temporaria. II. Spike timing.

We studied the directionality of spike timing in the responses of single auditory nerve fibers of the grass frog, Rana temporaria, to tone burst stimulation. Both the latency of the first spike after stimulus onset and the preferred firing phase during the stimulus were studied. In addition, the directionality of the phase of eardrum vibrations was measured. The response latency showed systematic and statistically significant changes with sound direction at both low and high frequencies. The latency changes were correlated with response strength (spike rate) changes and were probably the result of directional changes in effective stimulus intensity. Systematic changes in the preferred firing phase were seen in all fibers that showed phaselocking (i.e., at frequencies below 500-700 Hz). The mean phase lead for stimulation from the contralateral side was approximately 140 degrees at 200 Hz and decreased to approximately 100 degrees at 700 Hz. These phaseshifts correspond to differences in spike timing of approximately 2 ms and 0.4 ms respectively. The phaseshifts were nearly independent of stimulus intensity. The phase directionality of eardrum vibrations was smaller than that of the nerve fibers. Hence, the strong directional phaseshifts shown by the nerve fibers probably reflect the directional characteristics of extratympanic pathways.