An analysis is presented of the mechanical response to a sound field of the ears of the parasitoid fly Ormia ochracea. This animal shows a remarkable ability to detect the direction of an incident sound stimulus even though its acoustic sensory organs are in very close proximity to each other. This close proximity causes the arrival times of the sound pressures at the two ears to be less than 1 to 2 microseconds depending on the direction of propagation of the sound wave. The small differences in these two pressures must be processed by the animal in order to determine the incident direction of the sound. In this fly, the ears are so close together that they are actually joined by a cuticular structure which couples their motion mechanically and subsequently magnifies interaural differences. The use of a cuticular structure as a means to couple the ears to achieve directional sensitivity is novel and has not been reported in previous studies of directional hearing. An analytical model of the mechanical response of the ear to a sound stimulus is proposed which supports the claim that mechanical interaural coupling is the key to this animal's ability to localize sound sources. Predicted results for sound fields having a range of incident directions are presented and are found to agree very well with measurements.

Insects use several senses to forage, detecting floral cues such as color, shape, pattern, and volatiles. We report a formerly unappreciated sensory modality in bumblebees (Bombus terrestris), detection of floral electric fields. These fields act as floral cues, which are affected by the visit of naturally charged bees. Like visual cues, floral electric fields exhibit variations in pattern and structure, which can be discriminated by bumblebees. We also show that such electric field information contributes to the complex array of floral cues that together improve a pollinator's memory of floral rewards. Because floral electric fields can change within seconds, this sensory modality may facilitate rapid and dynamic communication between flowers and their pollinators.

In humans and other vertebrates, hearing is improved by active contractile properties of hair cells. Comparable active auditory mechanics is now demonstrated in insects. In mosquitoes, Johnston's organ transduces sound-induced vibrations of the antennal £agellum. A non-muscular`motor' activity enhances the sensitivity and tuning of the £agellar mechanical response in physiologically intact animals. This motor is capable of driving the £agellum autonomously, amplifying sound-induced vibrations at speci¢c frequencies and intensities. Motor-related electrical activity of Johnston's organ strongly suggests that mosquito hearing is improved by mechanoreceptor motility.

In insects and vertebrates alike, hearing is assisted by the motility of mechanosensory cells. Much like pushing a swing augments its swing, this cellular motility is thought to actively augment vibrations inside the ear, thus amplifying the ear's mechanical input. Power gain is the hallmark of such active amplification, yet whether and how much energy motile mechanosensory cells contribute within intact auditory systems has remained uncertain. Here, we assess the mechanical energy provided by motile mechanosensory neurons in the antennal hearing organs of Drosophila melanogaster by analyzing the fluctuations of the sound receiver to which these neurons connect. By using dead WT flies and live mutants (tilB 2 , btv 5P1 , and nompA 2 ) with defective neurons as a background, we show that the intact, motile neurons do exhibit power gain. In WT flies, the neurons lift the receiver's mean total energy by 19 zJ, which corresponds to 4.6 times the energy of the receiver's Brownian motion. Larger energy contributions (200 zJ) associate with self-sustained oscillations, suggesting that the neurons adjust their energy expenditure to optimize the receiver's sensitivity to sound. We conclude that motile mechanosensory cells provide active amplification; in Drosophila, mechanical energy contributed by these cells boosts the vibrations that enter the ear.cochlear amplifier ͉ hearing ͉ auditory mechanics ͉ cell mobility ͉ hair cell T he cochlear amplifier is the dominant unifying concept in cochlear mechanics (1). The concept assumes that the cochlea is endowed with a biological energy source that amplifies the ear's input by pumping mechanical energy into the vibrations inside the ear (1-6). The validity of the concept is supported by the mechanics of the cochlea and its mechanosensory cells. Hair cells, the cochlear mechanosensory cells, provide a source of mechanical energy. In addition to transducing mechanical vibrations into electrical responses, some hair cells are equipped with molecular motors that convert metabolic or electrical energy into mechanical energy, resulting in active movements of the cells (1, 3-7). These cellular movements, in turn, exert positive feedback on the cochlear mechanics. By nonlinearly undamping the cochlear resonances as the stimulus intensity declines, this feedback selectively improves the ear's sensitivity to small vibrations induced by faint sound (1, 3-6). Notably, this hair cell-based feedback occasionally becomes unstable, leading to self-sustained feedback oscillations within the cochlear duct. Such self-sustained feedback oscillations may account for the ear's ability to generate spontaneous otoacoustic emissions, i.e., to spontaneously emit sound (8, 9).Collectively, the hair cells' motility, the cochlea's nonlinearity, and the ear's spontaneous otoacoustic emissions document the presence of hair cell-based mechanical feedback inside the cochlear duct. Yet, whether this feedback brings about power gain by expending biological energy, as assumed by the concept of the cochlear ampli...

In mammals, hearing is dependent on three canonical processing stages: (i) an eardrum collecting sound, (ii) a middle ear impedance converter, and (iii) a cochlear frequency analyzer. Here, we show that some insects, such as rainforest katydids, possess equivalent biophysical mechanisms for auditory processing. Although katydid ears are among the smallest in all organisms, these ears perform the crucial stage of air-to-liquid impedance conversion and signal amplification, with the use of a distinct tympanal lever system. Further along the chain of hearing, spectral sound analysis is achieved through dispersive wave propagation across a fluid substrate, as in the mammalian cochlea. Thus, two phylogenetically remote organisms, katydids and mammals, have evolved a series of convergent solutions to common biophysical problems, despite their reliance on very different morphological substrates.