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.

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.

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.

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.

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.

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.

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.

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.

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.

Helioseismic measurement of solar torsional oscillations.

Bands of slower and faster rotation, the so-called torsional oscillations, are observed at the Sun's surface to migrate in latitude over the 11-year solar cycle. Here, we report on the temporal variations of the Sun's internal rotation from solar p-mode frequencies obtained over nearly 6 years by the Michelson Doppler Imager (MDI) instrument on board the Solar and Heliospheric Observatory (SOHO) satellite. The entire solar convective envelope appears to be involved in the torsional oscillations, with phase propagating poleward and equatorward from midlatitudes at all depths throughout the convective envelope.

Deeply Penetrating Banded Zonal Flows in the Solar Convection Zone.

Helioseismic observations have detected small temporal variations of the rotation rate below the solar surface that correspond to the so-called "torsional oscillations" known from Doppler measurements of the surface. These appear as bands of slower- and faster-than-average rotation moving equatorward. Here we establish, using complementary helioseismic observations over 4 yr from the GONG network and from the MDI instrument on board SOHO, that the banded flows are not merely a near-surface phenomenon: rather, they extend downward at least 60 Mm (some 8% of the total solar radius) and thus are evident over a significant fraction of the nearly 200 Mm depth of the solar convection zone.

Dynamic variations at the base of the solar convection zone

We have detected changes in the rotation of the sun near the base of its convective envelope, including a prominent variation with a period of 1.3 years at low latitudes. Such helioseismic probing of the deep solar interior has been enabled by nearly continuous observation of its oscillation modes with two complementary experiments. Inversion of the global-mode frequency splittings reveals that the largest temporal changes in the angular velocity Omega are of the order of 6 nanohertz and occur above and below the tachocline that separates the sun's differentially rotating convection zone (outer 30% by radius) from the nearly uniformly rotating deeper radiative interior beneath. Such changes are most pronounced near the equator and at high latitudes and are a substantial fraction of the average 30-nanohertz difference in Omega with radius across the tachocline at the equator. The results indicate variations of rotation close to the presumed site of the solar dynamo, which may generate the 22-year cycles of magnetic activity.

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.

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.

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.

Sound and vibration sensitivity of VIIIth nerve fibers in the grassfrog, Rana temporaria.

We have studied the sound and vibration sensitivity of 164 amphibian papilla fibers in the VIIIth nerve of the grassfrog, Rana temporaria. The VIIIth nerve was exposed using a dorsal approach. The frogs were placed in a natural sitting posture and stimulated by free-field sound. Furthermore, the animals were stimulated with dorso-ventral vibrations, and the sound-induced vertical vibrations in the setup could be canceled by emitting vibrations in antiphase from the vibration exciter. All low-frequency fibers responded to both sound and vibration with sound thresholds from 23 dB SPL and vibration thresholds from 0.02 cm/s2. The sound and vibration sensitivity was compared for each fiber using the offset between the rate-level curves for sound and vibration stimulation as a measure of relative vibration sensitivity. When measured in this way relative vibration sensitivity decreases with frequency from 42 dB at 100 Hz to 25 dB at 400 Hz. Since sound thresholds decrease from 72 dB SPL at 100 Hz to 50 dB SPL at 400 Hz the decrease in relative vibration sensitivity reflects an increase in sound sensitivity with frequency, probably due to enhanced tympanic sensitivity at higher frequencies. In contrast, absolute vibration sensitivity is constant in most of the frequency range studied. Only small effects result from the cancellation of sound-induced vibrations. The reason for this probably is that the maximal induced vibrations in the present setup are 6-10 dB below the fibers' vibration threshold at the threshold for sound. However, these results are only valid for the present physical configuration of the setup and the high vibration-sensitivities of the fibers warrant caution whenever the auditory fibers are stimulated with free-field sound. Thus, the experiments suggest that the low-frequency sound sensitivity is not caused by sound-induced vertical vibrations. Instead, the low-frequency sound sensitivity is either tympanic or mediated through bone conduction or sound-induced pulsations of the lungs.

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.