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1. Research Focus “Urbanistic Studies in Limyra” (M. Seyer) The focus in 2018 should be laid in investigations on the urbanistic development of Limyra mainly during the Hellenistic period. Excavation and research work could be carried out in the frame of the scientific project “The Urbanistic Development of Limyra in the Hellenistic Period”. This project deals with general questions such as the spatial expansion respectively the borders of the settlement, the structural development of the cit...

The mammalian visual system is highly directional and mammals typically employ rapid eye movements to scan their environment. Both sound emission and hearing in echolocating bats are directional but not much is known about how bats use ear movements and possibly movements of the sound-emitting structures to scan space. Here, we investigated in a tightly controlled behavioural experiment how Phyllostomus discolor bats employ their echolocation system while being moved through differently structured environments: we monitored and reconstructed both a close-up of the facial structures in 3D, including the motile noseleaf and outer ears, and the sonar-beam of the bat while it was moved along reflectors. Despite the simple linear movement of the bats in the setup, the bats pointed their beam quite variably in azimuth with a standard deviation of about ±20 deg. This variation arises from yaw-type head rotations. Video analyses show that the bat's noseleaf twitches with every echolocation call. Second, we show that the bat's ears are raised to a rather stereotypical head-centred position with every echolocation call. Surprisingly, P. discolor can adjust the timing and the magnitude of these ear movements to the distance of the reflectors with millisecond precision. Our findings reveal echolocation-specific specialisations as well as general principles of scanning and stabilisation of a directional remote sense. The call-correlated movements of the facial structures may lead to a higher directionality of the echolocation system and may enable the bats to adjust their echo-acoustic gaze to dynamic environments.

This study was supported by the Carlsberg Foundation via a Semper Ardens grant, ONR, N00014-17-1- 2736; AFOSR FA9550-14-1-0398, and NSF NCS-FO:1734744 and a Human Frontiers Science Program Long-Term Fellowship to AS. These experiments were approved by The Danish Council for Experiments on Animals under permit number: 2016-15-0201-00989 and by the Johns Hopkins University Animal Care and Use Committee under protocol number BA17A107. We thank Uwe Firzlaff and Lutz Wiegrebe for their help. 1. To study sensorimotor behaviour in wild animals, it is necessary to synchronously record the sensory inputs available to the animal, and its movements. To do this, we have developed a biologging device that can record the primary sensory information and the associated movements during foraging and navigating in echolocating bats. 2. This 2.6 -gram tag records the sonar calls and echoes from an ultrasonic microphone, while simultaneously sampling fine-scale movement in three dimensions from wideband accelerometers and magnetometers. In this study, we tested the tag on an European noctula (Nyctalus noctula) during target approaches and on four big brown bats (Eptesicus fuscus) during prey interception in a flight room. 3. We show that the tag records both the outgoing calls and echoes returning from objects at biologically relevant distances. Inertial sensor data enables the detection of behavioural events such as flying, turning, and resting. In addition, individual wing-beats can be tracked and synchronized to the bat's sound emissions to study the coordination of different motor events. 4. By recording the primary acoustic flow of bats concomitant with associated behaviours on a very fine time-scale, this type of biologging method will foster a deeper understanding of how sensory inputs guide feeding behaviours in the wild. Postprint

Echolocating bats are known to fly and forage in complete darkness using the echoes of their actively emitted calls to navigate and to detect prey. However, under dim light conditions many bats can also rely on vision. Many flying animals have been shown to navigate by optic flow information, and recently, bats were shown to exploit echo-acoustic flow to navigate through dark habitats. Here we show for the bat Phyllostomus discolor that in lighted habitats where self-motion induced optic flow is strong, optic and echo-acoustic flow interact in their efficiency to guide navigation. Echo-acoustic flow showed a surprisingly strong effect compared to optic flow. We thus demonstrate multimodal interaction between two far-ranging spatial senses, vision and echolocation, available in this combination almost exclusively for bats and toothed whales. Our results highlight the importance of merging information from different sensory systems in a sensory-specialist animal to successfully navigate and hunt under difficult conditions.

Bats are navigation super-performers, flying at high speed through nocturnal forests. Numerous field observations and formal experiments have impressively shown how well bats tackle navigation in 3D with biosonar, i.e., the auditory analysis of self-generated ultrasonic emissions [1-7]. However, unlike in the visual system, where space is explicitly coded at very high resolution in the retinal fovea, the inner ear encodes frequency and time, not space. Spatial attributes of echoes are represented in the space-dependent filtering of the bats' pinnae [8, 9] and binaural computations, like interaural time and level differences [10, 11], as first proposed by Lord Rayleigh [12]. Remarkably, Rayleigh also provided a clear definition of spatial resolution: based on the shape of optical diffraction patterns arising from two closely spaced light sources, Rayleigh defined resolution as the capability to detect a trough in their joint light diffraction patterns [13, 14]. Here, we recruit Rayleigh's classical resolution paradigm to quantify how well bats can resolve multiple simultaneously presented reflectors in space. We show that biosonar spatial resolution in azimuth is no better than about 80° compared to a human visual resolution down to 0.02° [14]. We suggest that bats compensate this effective lack of spatial resolution by sequentially probing their environment in flight. Our data show that low-resolution environment perception is a viable alternative to high-resolution vision to support intelligent behavior in complex environments.

Echolocating bats are known to fly and forage in complete darkness, using the echoes of their actively emitted calls to navigate and to detect prey. However, under dim light conditions many bats can also rely on vision. Many flying animals have been shown to navigate by optic flow information and, recently, bats were shown to exploit echo-acoustic flow to navigate through dark habitats. Here, we show for the bat

>Human hearing is rather insensitive for very low frequencies (i.e. below 100 Hz). Despite this insensitivity, low-frequency sound can cause oscillating changes of cochlear gain in inner ear regions processing even much higher frequencies. These alterations outlast the duration of the low-frequency stimulation by several minutes, for which the term 'bounce phenomenon' has been coined. Previously, we have shown that the bounce can be traced by monitoring frequency and level changes of spontaneous otoacoustic emissions (SOAEs) over time. It has been suggested elsewhere that large receptor potentials elicited by low-frequency stimulation produce a net Ca(2+) influx and associated gain decrease in outer hair cells. The bounce presumably reflects an underdamped, homeostatic readjustment of increased Ca(2+) concentrations and related gain changes after low-frequency sound offset. Here, we test this hypothesis by activating the medial olivocochlear efferent system during presentation of the bounce-evoking low-frequency (LF) sound. The efferent system is known to modulate outer hair cell Ca(2+) concentrations and receptor potentials, and therefore, it should modulate the characteristics of the bounce phenomenon. We show that simultaneous presentation of contralateral broadband noise (100 Hz-8 kHz, 65 and 70 dB SPL, 90 s, activating the efferent system) and ipsilateral low-frequency sound (30 Hz, 120 dB SPL, 90 s, inducing the bounce) affects the characteristics of bouncing SOAEs recorded after low-frequency sound offset. Specifically, the decay time constant of the SOAE level changes is shorter, and the transient SOAE suppression is less pronounced. Moreover, the number of new, transient SOAEs as they are seen during the bounce, are reduced. Taken together, activation of the medial olivocochlear system during induction of the bounce phenomenon with low-frequency sound results in changed characteristics of the bounce phenomenon. Thus, our data provide experimental support for the hypothesis that outer hair cell calcium homeostasis is the source of the bounce phenomenon.

Flying animals need to react fast to rapid changes in their environment. Visually guided animals use optic flow, generated by their movement through structured environments. Nocturnal bats cannot make use of optic flow, but rely mostly on echolocation. Here we show that bats exploit echo-acoustic flow to negotiate flight through narrow passages. Specifically, bats' flight between lateral structures is significantly affected by the echo-acoustic salience of those structures, independent of their physical distance. This is true although echolocation, unlike vision, provides explicit distance cues. Moreover, the bats reduced the echolocation sound levels in stronger flow, likely to compensate for the increased summary target strength of the lateral reflectors. However, bats did not reduce flight velocity under stronger echo-acoustic flow. Our results demonstrate that sensory flow is a ubiquitous principle for flight guidance, independent of the fundamentally different peripheral representation of flow across the senses of vision and echolocation.

Reverberation can dramatically reduce the depth of amplitude modulations which are critical for speech intelligibility. Psychophysical experiments indicate that humans' sensitivity to amplitude modulation in reverberation is better than predicted from the acoustic modulation depth at the receiver position. Electrophysiological studies on reverberation in rabbits highlight the contribution of neurons sensitive to interaural correlation. Here, we use a prepulse-inhibition paradigm to quantify the gerbils' amplitude modulation threshold in both anechoic and reverberant virtual environments. Data show that prepulse inhibition provides a reliable method for determining the gerbils' AM sensitivity. However, we find no evidence for perceptual restoration of amplitude modulation in reverberation. Instead, the deterioration of AM sensitivity in reverberant conditions can be quantitatively explained by the reduced modulation depth at the receiver position. We suggest that the lack of perceptual restoration is related to physical properties of the gerbil's ear input signals and inner-ear processing as opposed to shortcomings of their binaural neural processing.

Research focus “urbanistic studies in Limyra” With the approval of the scientific project “The Urbanistic Development of Limyra in the Hellenistic Period” for three years by the Austrian Science Fund (FWF) (P29027-G25), the research program on urbanism that had already been conducted for several years could be intensified in 2016. Particular focus should be given to the development of Limyra especially in the period under consideration, whereby the extent, the structures, and the urban image ...