1 3
J Comp Physiol A
DOI 10.1007/s00359-014-0914-2
ORIGINAL PAPER
Mechanical basis of otoacoustic emissions in tympanal hearing
organs
Doreen Möckel · Manuela Nowotny · Manfred Kössl
Received: 29 January 2014 / Revised: 14 April 2014 / Accepted: 16 April 2014
© Springer-Verlag Berlin Heidelberg 2014
Keywords DPOAE · Laser Doppler vibrometry · Locust ·
Tympanum · Insect
Abbreviations
DPOAE Distortion–product otoacoustic emission
LDV Laser Doppler vibrometry
SPL Sound pressure level
Introduction
Insects with tympanal organs produce sensitive distortion–
product otoacoustic emissions (DPOAEs), as shown in
locusts (Kössl and Boyan 1998; Möckel et al. 2007, 2012),
moths (Kössl and Coro 2006; Kössl et al. 2007) and bush-
crickets (Möckel et al. 2011), that in mammals are used as
indicator for nonlinear cochlear amplification. Character-
istics of insect DPOAEs are largely comparable to those
reported from vertebrate ears. They appear during simul-
taneous stimulation with two pure tones (f
1
< f
2
) as addi-
tional spectral peaks at frequencies of nf
1
- (n - 1)f
2
and
nf
2
- (n - 1)f
1
. The 2f
1
- f
2
emission is most prominent,
as it is evoked by lowest stimulus levels, with the largest
amplitude reached at frequencies of high auditory sensitiv-
ity. In moths and locusts, the 2f
1
- f
2
DPOAEs are elicited
already at stimuli levels near the species-specific auditory
threshold (Kössl et al. 2008). Insect DPOAEs are highly
vulnerable to manipulations that interfere with the animal’s
physiological state, such as hypoxia (Kössl and Boyan
1998) or ethyl ether (Kössl et al. 2007). First evidence that
the auditory mechanoreceptors and specifically the dynein–
tubulin system within could be relevant for DPOAE gen-
eration came from selective exclusion of mechanoreceptors
via mechanical lesions (on locusts, Möckel et al. 2007),
the effects of a neuroactive insecticide (on bushcrickets,
Abstract Tympanal hearing organs of insects emit dis-
tortion–product otoacoustic emissions (DPOAEs), which
in mammals are used as indicator for nonlinear cochlear
amplification, and which are highly vulnerable to manipu-
lations interfering with the animal’s physiological state.
Although in previous studies, evidence was provided for
the involvement of auditory mechanoreceptors, the source
of DPOAE generation and possible active mechanisms in
tympanal organs remained unknown. Using laser Dop-
pler vibrometry in the locust ear, we show that DPOAEs
mechanically emerge at the tympanum region where the
auditory mechanoreceptors are attached. Those emission-
coupled vibrations differed remarkably from tympanum
waves evoked by external pure tones of the same frequency,
in terms of wave propagation, energy distribution, and loca-
tion of amplitude maxima. Selective inactivation of the
auditory receptor cells by mechanical lesions did not affect
the tympanum’s response to external pure tones, but abol-
ished the emission’s displacement amplitude peak. These
findings provide evidence that tympanal auditory recep-
tors, comparable to the situation in mammals, comprise the
required nonlinear response characteristics, which during
two-tone stimulation lead to additional, highly localized
deflections of the tympanum.
Electronic supplementary material The online version of this
article (doi:10.1007/s00359-014-0914-2) contains supplementary
material, which is available to authorized users.
D. Möckel (*) · M. Nowotny · M. Kössl
Institut für Zellbiologie und Neurowissenschaft,
J. W. Goethe-Universität, Biologicum A,
Max-von-Laue-Straße 13, 60438 Frankfurt am Main, Germany
e-mail: moeckel@bio.uni-frankfurt.de