Neurophysiology goes wild: from exploring sensory coding in sound proof rooms to natural environments.

To perform adaptive behaviours, animals have to establish a representation of the physical "outside" world. How these representations are created by sensory systems is a central issue in sensory physiology. This review addresses the history of experimental approaches toward ideas about sensory coding, using the relatively simple auditory system of acoustic insects. I will discuss the empirical evidence in support of Barlow's "efficient coding hypothesis", which argues that the coding properties of neurons undergo specific adaptations that allow insects to detect biologically important acoustic stimuli. This hypothesis opposes the view that the sensory systems of receivers are biased as a result of their phylogeny, which finally determine whether a sound stimulus elicits a behavioural response. Acoustic signals are often transmitted over considerable distances in complex physical environments with high noise levels, resulting in degradation of the temporal pattern of stimuli, unpredictable attenuation, reduced signal-to-noise levels, and degradation of cues used for sound localisation. Thus, a more naturalistic view of sensory coding must be taken, since the signals as broadcast by signallers are rarely equivalent to the effective stimuli encoded by the sensory system of receivers. The consequences of the environmental conditions for sensory coding are discussed.

Spatial orientation in the bushcricket Leptophyes punctatissima (Phaneropterinae; Orthoptera): II. Phonotaxis to elevated sound sources on a walking compensator.

The ability of the bushcricket Leptophyes punctatissima to orient to elevated sound sources was investigated. Males were placed on a walking compensator and oriented in response to a synthetic female reply, which was broadcast via one of five loudspeakers placed at elevations of 0 degrees , 30 degrees , 60 degrees , 75 degrees and 90 degrees . Forward and backward movements were compensated, so that males remained at the same distance and elevation to the sound source. With increasing loudspeaker elevation, the males meandered more, and the ratio of the ideal path length to the actual path length decreased. The same was true for the correlation between stimulus angle and turn angle, and there were more turns to the wrong side with increasing loudspeaker elevation. Most males performed phonotaxis with a high acuity up to an elevation of 60 degrees . Individuals varied strongly in their performance especially at a source elevation of 75 degrees , where some were still very accurate in their approach, whereas the acuity of others decreased rapidly. We also describe a behaviour where males tilt their body axis to more anterior and sideward positions, both during walking and while calling on the spot. This behaviour is interpreted as a kind of directional scanning in order to actively induce changes in binaural cues.

Spatial orientation in the bushcricket Leptophyes punctatissima (Phaneropterinae; Orthoptera): I. Phonotaxis to elevated and depressed sound sources.

Many species of acoustically interacting insects live in a complex, arboreal or semi-arboreal habitat. Thus mate finding by phonotaxis requires sound localization in the horizontal and vertical plane. Here we investigated the ability of the duetting bushcricket Leptophyes punctatissima to orient to one of three speakers, positioned at different levels in an artificial grid system, where each point in space could be reached by the male with almost equal probability. The system was designed analogous to a spherical calotte model of bismuth, where, once the male arrived at any nodal point had to decide between only three directions: either up or down and/or left and right. This design does not favour any phonotactic path of the males. All 12 males tested reached the three speaker positions (one in the horizontal plane, one elevated by 45 degrees , one depressed by 45 degrees relative to the starting position) with only little deviation from the shortest possible path. There was no significant difference with respect to the whole phonotactic time needed, the number of segments passed, or the number of stimuli received for the different speaker positions. This remarkable spatial orientation is achieved although the insects have no specialized external ear structures such as mammals, or some owls.

Auditory lateralization in bushcrickets: a new dichotic paradigm.

Pair formation in the bushcricket Gampsocleis gratiosa is achieved through acoustic signalling by the male and phonotactic approaches of the female towards the calling song. On a walking belt in the free sound field, females tracked the position of the speaker broadcasting the male calling song with a remarkable precision, deviating by no more than 10 cm in either direction from the ideal course. Starting with stimulus angles of 6-10 degrees the females significantly turned to the correct side, and with stimulus angles greater than 25 degrees no incorrect turns were made. Using neurophysiological data on the directionality of the ear we calculated that with such stimulus angles the available binaural intensity difference is in the order of 1-2 dB. We developed a dichotic ear stimulation device for freely moving females with a cross-talk barrier of about 50 dB, which allowed to precisely apply small binaural intensity differences. In such a dichotic stimulation paradigm, females on average turned to the tronger stimulated side starting with a 1 dB difference between both ears. The significance of such a reliable lateralization behaviour with small interaural intensity differences for phonotactic behaviour under natural conditions is discussed.