Wenyu Lin

Old World frog and bird vocalizations contain prominent ultrasonic harmonics

Several groups of mammals such as bats, dolphins and whales are known to produce ultrasonic signals which are used for navigation and hunting by means of echolocation, as well as for communication. In contrast, frogs and birds produce sounds during night- and day-time hours that are audible to humans; their sounds are so pervasive that together with those of insects, they are considered the primary sounds of nature. Here we show that an Old World frog (Amolops tormotus) and an oscine songbird (Abroscopus albogularis) living near noisy streams reliably produce acoustic signals that contain prominent ultrasonic harmonics. Our findings provide the first evidence that anurans and passerines are capable of generating tonal ultrasonic call components and should stimulate the quest for additional ultrasonic species.

Temporal selectivity for complex signals by single neurons in the torus semicircularis of Pleurodema thaul (Amphibia:Leptodactylidae).

Abstract Responses of auditory neurons in the torus semicircularis (TS) of Pleurodema thaul, a leptodactylid from Chile, to synthetic stimuli having diverse temporal patterns and to digitized advertisement calls of P. thaul and three sympatric species, were recorded to investigate their temporal response selectivities. The advertisement call of this species consists of a long sequence of sound pulses (a pulse-amplitude-modulated, or PAM, signal) having a dominant frequency of about 2000 Hz. Each of the sound pulses contains intra-pulse sinusoidal-amplitude-modulations (SAMs). Synthetic stimuli consisted of six series in which the following acoustic parameters were systematically modified, one at a time: PAM rate, pulse duration, number of pulses, and intra-pulse SAM rate. The carrier frequency of these stimuli was set at the characteristic frequency (CF) of the isolated units (n = 47). Response patterns of TS units to synthetic call variants reveal different degrees of selectivities for each of the temporal variables, with populations of neurons responding maximally to specific values found in the advertisement call of this species. These selectivities are mainly shaped by neuronal responsiveness to the overall sound energy of the stimulus and by the inability of neurons to discharge to short inter-pulse gaps.

Detection of gaps in sinusoids by frog auditory nerve fibers: importance in AM coding

Physiological studies were carried out in the frog (Rana pipiens pipiens) eighth nerve to determine: (i) whether the modulation rate or the silent gap was the salient feature that set the upper limit of time-locking to pulsed amplitude-modulated (PAM) stimuli, (ii) the gap detection capacity of individual eighth nerve fibers. Time-locked responses of 79 eighth nerve fibers to PAM stimuli (at the fibers characteristic frequency) showed that the synchronization coefficient was a low-pass function of the modulation rate. In response to PAM stimuli having different pulse durations, a fiber gave rise to non-overlapping modulation transfer functions. The upper cut-off frequency of time locking was higher when tonepulses in PAM stimuli had shorter duration. The fact that the cut-off frequency was different for the different PAM series suggested that the AM rate was neither the sole, nor the main, determinant for the decay in time-locking at high AM rates. Gap detection capacity was determined for 69 eighth nerve fibers by assessing fibers spiking activities to paired tone-pulses during an OFF-window and an ON-window. It was found that the minimum detectable gap of eighth nerve fibers ranged from 0.5 to 10 ms with an average of 1.23–2.16 ms depending on the duration of paired tone pulses. For each fiber, the minimum detectable gap was longer when the duration of tone pulses comprising the twin-pulse stimuli was more than four times longer. When the synchronization coefficient was plotted against the silent gap between tones pulses in the PAM stimuli, the gap response functions of a fiber as derived from multiple PAM series were equivalent to gap response functions deriving from twin-pulse series suggesting that it was the silent gap which primarily determined the upper limit of time-locking to PAM stimuli.

NEURONAL ARCHITECTURE OF THE DORSAL NUCLEUS (COCHLEAR NUCLEUS) OF THE FROG, RANA PIPIENS PIPIENS

The neuronal architecture of the dorsal nucleus of the Northern leopard frog (Rana pipiens pipiens), which is a homolog of the cochlear nucleus of mammals and birds, was investigated. Our study showed that the frog dorsal nucleus contains a number of morphologically distinct cell types that are discernible in terms of the cellular architecture as derived from Nissl‐stained material and in terms of the dendritic profile as revealed by horseradish peroxidase‐filled single neurons. These cell types are bushy cells, bipolar (or fusiform) cells, octopus cells, stellate cells, giant cells, radiate (or round) cells, and a variety of small cells. The different cell types occupy different regions of the nucleus. Therefore, our results suggest that the dorsal nucleus should no longer be considered to be a uniform nucleus containing a homogeneous population of neurons. Homologies of these cell types with those described in other vertebrate species, including mammals, are proposed.

Wide-ranging frequency preferences of auditory midbrain neurons: Roles of membrane time constant and synaptic properties

Periodicity is a fundamental sound attribute. Its coding has been the subject of intensive research, most of which has focused on investigating how the periodicity of sounds is processed through the synaptic machinery in the brain. The extent to which the intrinsic properties of cells play in periodicity coding, particularly in the creation of selectivity to periodic signals, is not well understood. We performed in vitro whole‐cell patch recordings in the frog torus semicircularis to investigate each neuron’s intrinsic membrane properties as well as responses to sinusoidal current injected through the electrode and periodic stimulation of the ascending afferent. We found that: (i) toral neurons were heterogeneous, showing diverse biophysical phenotypes having distinct membrane characteristics, including membrane time constants (τ) and ionic channel compositions (Ih, Ikir, Ikv and INaP); (ii) a neuron’s τ was tightly correlated with its current‐evoked frequency preference (FP; range: 0.05–50 Hz); (iii) application of blockers for Ih, Ikir and Ikv (but not INaP) shifted the τ as well as the cell’s current‐evoked FP, suggesting that these ion channels contribute to the cell’s FP through regulation of τ; (iv) a neuron’s τ was also correlated with its afferent‐evoked FP (range: 10–300 pulses/s); and (v) the range of afferent‐evoked FP was approximately one order higher than the range of current‐evoked FPs, suggesting that both the cell’s intrinsic membrane and synaptic properties contribute to determining the afferent‐evoked cell‐specific FP (whose range matched those of cell‐specific responses to sound stimulation, e.g. selectivity to amplitude modulation rate).

Voices of the dead: Complex nonlinear vocal signals from the larynx of an ultrasonic frog

Most anurans are highly vocal but their vocalizations are stereotyped and simple with limited repertoire sizes compared to other vocal vertebrates, due presumably to the limited mechanisms for fine vocal motor control. We recently reported that the call of the concave‐eared torrent frog (Amolops tormotus) is an exception in its seemingly endless variety, musical warbling quality, extension of call frequency into the ultrasonic range, and the prominence of nonlinear features such as period doubling. We now show that the major spectral features of its calls, responsible for this frog’s vocal diversity, can be generated by forcing pressurized air through the larynx of euthanized males. Laryngeal specializations for ultrasound appear to include very thin portions of the medial vocal ligaments and the reverse sexual size dimorphism of the larynx being smaller in males than in females. The intricate morphology of the vocal cords, which changes along their length, suggests that nonlinear phenomena likely arise f...