ISSN 0006-3509, Biophysics, 2008, Vol. 53, No. 3, pp. 237–239. © Pleiades Publishing, Inc., 2008.

Original Russian Text © E.S. Babushina, M.A. Polyakov, 2008, published in Biofizika, 2008, Vol. 53, No. 3, pp. 499–503.

COMPLEX SYSTEMS
BIOPHYSICS

Vertical Auditory Acuity of the Bottlenose Dolphin

E. S. Babushina and M. A. Polyakov
Karadag Natural Reserve, National Academy of Sciences of Ukraine, Kurortnoe, Crimea, 98188 Ukraine
E-mail: poza@dolphin.crimea.com
Received June 27, 2007; in final form, November 22, 2007

Abstract—The accuracy in locating underwater sounds in the vertical median plane was determined for the
Black Sea bottlenose dolphin Tursiops truncatus trained by operant conditioning with food reinforcement. The
minimal perceived angles for 1-s tone signals were 2.5°at 5 or 20 kHz and 2.0° at 120 kHz; for 1-s trains of
clicks centered at 120 kHz the acuity was still better, ~1.5°. Dolphins may locate different sounds using differ-
ent yet equally efficient mechanisms, and they are the best in analyzing the acoustic space among marine mam-
mals studied.

Key words: odontocetes, Tursiops truncatus, acoustic signal, vertical localization

DOI: 10.1134/S0006350908030123

INTRODUCTION The dolphin was trained to take a start position just

Adequate orientation in space is important for any
organism. Dolphins with their unique auditory analyzer
superbly tackle this task.
A special role in 3D hearing is played by locating
the sound source in the vertical direction. In humans,
this is associated with directional filtration in a certain
frequency range by sound-conducting structures in the
head; filtration is determined by the type of acoustic
signal [1, 2]. Successful monaural localization of a
composite sound depends on comparison of at least
three frequency components and their relative ampli-
tudes [3–5]. The process is optimized in binaural per-
ception [6, 7]. It is reasonable to suppose that the basic
physical principles of sound location should be true for
other mammals.
It has been experimentally established that aquatic
animals with a highly specialized auditory analyzer use
echolocation and of course can detect sources of sound
[8–15]. Quite a few works have been devoted to assess-
ing the localization accuracy and the mechanisms of
directional hearing in odontocetes, but mainly in the
horizontal plane [16–21].
Here we examined the acuity of vertical localization
of underwater sound by a Black Sea bottlenose dolphin
(Tursiops truncatus) as dependent on the signal param-
eters.
EXPERIMENTAL
Experiments were performed in a muck-bottom sea
bay within a 9 × 6 × 4.5 m pile-net pen, using behav-
ioral response techniques (operant conditioning with
food reinforcement).
below the water surface and keep still until delivery of
a signal, rostrum in touch with the start manipulator (a
ceramic knob of 2 cm base diameter) suspended at a
depth of 10 cm. The central manipulator was the same
knob 6 m away; under it were the emitters: one at a
depth of 1 m (which was taken as the zero angle) and
the second was lower and its position could be varied.
In the start position the dolphin had its rostrum directed
onto the upper emitter. If a signal came from this direc-
tion, the dolphin went to the central manipulator,
pushed it and got fish. If a signal came at a larger angle,
the dolphin stayed at the start. The angular distance
between the emitters was gradually decreased from 20
to 1 or 0.5° in 5–1, 0.5° steps. A signal was delivered
from either of the emitters, interchanged at random. At
the moment the dolphin started to move, the signal was
switched off. The minimal angle between the emitters
distinguished by the dolphin corresponded to 75%-cor-
rect response.
The signal intensity well exceeded the dolphin hear-
ing threshold.
Use was made of a barium titanate piezoceramic trans-
ducer 14 mm in diameter, tone signal (5 and 20 kHz)
transducers 50 mm in diameter, and standard radio-
measuring equipment.
RESULTS AND DISCUSSION
We assessed the vertical angular acuity of the dol-
phin with tone signals (5, 20, 120 kHz) and with trains
of clicks (centered on 120 kHz, pulse duration 20 µs,
exponential fall, repetition rate 300 s–1); in all cases the
signal duration was 1 s.
The table shows that dolphin’s performance appre-
ciably improves with increasing tone frequency, the

237
238 BABUSHINA, POLYAKOV

Efficacy of vertical discrimination of sound sources by the Thus, it is clear that dolphins can determine the
bottlenose dolphin direction of the sound source with high accuracy both
in the horizontal and the vertical plane. These vital abil-
Correct
Frequency, Angle between Number of
responses, ities are due to the sophisticated complex of sound-con-
kHz emitters, deg trials ducting structures.
%
5 1.0 48 65 ± 6.3 The earlier works on sound conduction in marine
1.5 44 59 ± 14.6 mammals have been reviewed in this Journal [22]; here we
2.0 194 69 ± 6.5
should mention the more recent publications [23–27].
Experimental proof has been obtained for the concurrent
3.0 18 95 ± 4.8 operation of two pathways: via the acoustic meatuses,
4.5 15 100 which dominate in the lower frequency range, and the
20 1.0 21 57 ± 3.5 lower jaw, which is the main conductor above 30 kHz.
2.0 117 72 ± 8.1 Together with the surrounding tissues, the auditory pas-
3.0 163 76 ± 6.5 sage and the lower jaw largely ensure the directionality
of auditory perception [28]. Scanning head movements
4.5 41 88 ± 10
favor more detailed analysis of the differences in inten-
9.0 19 100 sity and spectral composition at the two receivers [29].
120 2.0 243 75 ± 5.5 Composite sounds “spread out” over the conducting
3.0 42 83 ± 10 pathways and interact with them to create a specific pat-
3.5 84 80 ± 2.5 tern in the auditory centers depending on the coordi-
5.0 20 100 nates of the sound source.
10 60 100 For the dolphin, the directionality diagram in either
15 56 100 plane has been shown to change with increasing fre-
20 380 100 quency and decreasing duration of the signal, becoming
Clicks* 1.0 44 61 ± 14 narrower and steeper [30, 31]. Particularly, the upper
slope of the vertical diagram is steeper than that of the
1.5 88 72 ± 9.4 horizontal one [31], suggesting more favorable condi-
2.0 82 83 ± 8.0 tions for source localization by intensity difference. It
* Centered on 120 kHz, pulse duration 20 µs, exponential fall, has been supposed [30] that the dolphin can scan the
repetition rate 300 s–1; 26 dB re 1 Pa. signal by head movements and adjust the characteris-
tics of the receiver to optimize spatial–frequency filtra-
tion.
minimal distinguished angle being about 2.5° at 5 and
20 kHz and 2.0° at 120 kHz, the latter value equaling Collation of the data on vertical acuity for tone sig-
the acuity obtained in the horizontal plane [21]. The nals suggests the involvement of different but equally
acuity for clicks (at an intensity of 26 dB re 1 Pa at the efficient mechanisms of localization at different fre-
start site) was nearly 1.5° (figure). quencies. This is in line with our earlier data on hori-
zontal accuracy for two dolphins [21] and the results of
other authors [10, 11, 19]. Especially good agreement
P, % is seen with the performance of an Atlantic bottlenose
1 * 3 [19] with similar kinds of signal, despite the substantial
100
* differences in the experimental setup (dolphin’s head
90 2 was immobile, and the “vertical” arrangement was
examined by training the animal to lie on its side). The
80 4 acuity for simulated echolocation clicks at ~64 kHz
70 was even better than in our case (within 1° in both
* planes). However, the authors [19] noted that in the
*
60 * pseudovertical localization the upper jaw was directed
towards one transducer and the lower jaw towards the
50 other. Therefore, the dolphin could use e.g. the differ-
40 ence in response evoked in the inferior colliculus by
sounds arriving through the lower jaw and through the
30 dorsal portion of the rostrum [32]. Hence, these data
1 2 4 6 8 10 θ, deg
may pertain to intensity discrimination rather than true
vertical localization. Again, an unrestrained animal (as
Graphical representation of the data on vertical angular acu-
ity of the dolphin (table): tones of (1) 5, (2) 20, and (3) 120 in our experiments) can locate the sound source by
kHz, and (4) trains of clicks; dashed line marks the 75%- memorizing the signal and comparing it with the one
correct threshold. received at another head position.

BIOPHYSICS Vol. 53 No. 3 2008

 VERTICAL AUDITORY ACUITY OF THE BOTTLENOSE DOLPHIN
Thus, the auditory acuity of dolphins exceeds that of
semiaquatic mammals (pinnipeds) by a factor of 1.6–
1.8 in the horizontal and 5–9 in the vertical plane (at
frequencies optimal for each species) [13–15]. Obvi-
ously, vertical localization requires a set of high-fre-
quency components, and the task is best solved by the
dolphin with a sophisticated auditory apparatus that has
largely evolved with echolocation. Dolphins are supe-
rior to all marine mammals studied in analyzing the
acoustic space.
ACKNOWLEDGMENTS
The authors are grateful to N.G. Bibikov for discus-
sions and valuable advice.
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