ISSN 0006-3509, Biophysics, 2008, Vol. 53, No. 3, pp. 235–236. © Pleiades Publishing, Inc., 2008.

Original Russian Text © E.S. Babushina, M.A. Polyakov, 2008, published in Biofizika, 2008, Vol. 53, No. 3, pp. 495–498.

COMPLEX SYSTEMS
BIOPHYSICS

Efficiency of Body Conduction of Underwater Sounds

in 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 December 1, 2006; in final form, November 12, 2007

Abstract—Hearing thresholds of the Black Sea bottlenose dolphin Tursiops truncatus for tonal and composite
underwater sounds within 50 kHz were ascertained in fully or partly submerged (head out of water) animals
(trained by operant conditioning with food reinforcement). Perception of sounds conducted through the body
deteriorated in all cases (thresholds for 75%-correct response rose typically by 6–24 dB); the least changes were
observed for 10 and 20 kHz tones. The aggregate data also suggested significant individual variations.

Key words: odontocetes, Tursiops truncatus, hearing threshold, sound transmission

DOI: 10.1134/S0006350908030111

INTRODUCTION 9.5 × 4.5 m, inside a hall 30 m long, 15 m wide, and 8 m

The acoustic orientation of marine mammals such
as dolphins and pinnipeds is determined by their anat-
omy, morphology, and ecology, by the functional prop-
erties of the conduction pathways as well as by the
parameters of the acoustic signals. We have previously
assessed sound conduction in the bottlenose dolphin,
the Caspian seal, and the northern fur seal [1–3] by
comparing the perception thresholds for underwater
sounds in animals submerged completely, with the head
under water; and partially, with the head out of water.
This comparison provided an estimate for the fraction
of the sound energy transmitted to the inner ear through
body tissues. Dolphins and true seals were found to
hear underwater sounds well even when the animal’s
head was out of water. In the sea-bear, sound transmis-
sion through the body was substantially hampered in
the main range of sound reception because of the muf-
fling properties of its air-containing fur. Works on
sound conduction through dolphin’s head tissues [4–6]
and our data on masking thresholds [3, 7] suggested
that the sound-transmitting channels of aquatic and
semiaquatic mammals interact with the signal to alter
the spectral structure of the composite sound. Being
aware of the possible individual differences, here we
continue our studies on body conduction and percep-
tion of various sounds by the bottlenose dolphin.
EXPERIMENTAL
Main experiments were performed with an adult
male dolphin (Tursiops truncatus) using behavioral
response techniques (operant conditioning with food
reinforcement) in a concrete pool measuring 27.5 ×
high.
At the start, the animal was either in a horizontal
position just below the water surface (complete sub-
mergence) or in a vertical position, with the head above
the water surface (partial submergence). In the latter
position, the sound was conducted through body tis-
sues. Waiting for a signal, the animal kept its nose in
touch with a start manipulator suspended at a height (H)
that we could vary. The waiting time was varied at ran-
dom from 1 to 7 s. Hearing a signal, the animal
approached its source. Either a pause or a signal was
presented. In the absence of a signal (that is, during the
pause), the animal had to stay at the start. Its approach
to the generator in this case was recorded as a false
response. The signal intensity was varied discretely, in
decrements of 3–6 dB, from a level several times
greater than the threshold to a subthreshold level; 10–
20 trials were made at each intensity. Usually the dol-
phin responded to sound within fraction of a second,
not waiting for the signal to end. In doubtful cases, near
the perception threshold, the dolphin preferred not to
go; sometimes the dolphin stopped halfway and
returned to check whether there was any sound.
The threshold was defined as the stimulus intensity
corresponding to 75% correct responses. The procedure
of gradual attenuation continued until the stimulus elic-
ited only 5060% correct responses. We repeated this
procedure several times a day; therefore, the animal
was presented with a signal of a given intensity at least
20 times (up to several tens of times) throughout one
experiment, and tens or hundreds of times over all
experiments. Presented below are the relative estimates
for the thresholds, allowing their comparison in fully
and partially submerged animals at various frequencies.

235
236 BABUSHINA, POLYAKOV

Auditory sensitivity (decibels relative to 1 W/cm2) of a bottlenose dolphin (24-years-old male)

Broadband noise @@@ Frequency of signal, kHz

H, cm
(0–50 kHz) (37–50 kHz) 5 10 20 40
Tone, H = 0
–38 ± 1 –41 ± 1 –44 + 3, –4 –43 ± 1
0 –34 + 2, –3 –36 ± 0
Narrowband, H = 0
–22 ± 1 –40 + 2, –3 –46 ± 0 –39 ± 1
Tone, H = 50
0±1 –35 + 2 –30 + 2, –3 –16 ± 1
50 –10 ± 2 –18 + 1, –2
Narrowband, H = 50
1±1 –28 ± 1 –26 + 4, –9 –15 ± 2
Note: H is the height of suspension of the start manipulator (centimeters): H = 0 corresponds to full submergence of the animal (head in
water), and H = 50 cm corresponds to partial submergence (head out of water).

The hearing thresholds for underwater acoustic sig-
nals were determined in the 5–40 kHz range, using tone
pulses (100-ms duration, repetition time 200 ms, 0.2–
5.0 ms rise, 5–25 ms fall) of 5, 10, 20, and 40 kHz or
corresponding narrowband (10% of the central fre-
quency) continuous noise-like signals. Also tested were
broadband Gaussian signals encompassing this fre-
quency range.
The generator was a cylindrical magnetostriction
transducer 35 mm in diameter and 120 mm in height
with a working frequency range of 0.5100.0 kHz. The
signal amplitude was maximal at the center of the range
and decreased to its edges by 28 and 13 dB. The emitter
was placed 0.6 m below the water surface, 2.4 m from
the start position.
RESULTS AND DISCUSSION
The results are shown in the table and figure. For all
kinds of signal, false responses at near-threshold levels
P, dB re 1 µbar
0
on average accounted for 1.3% of all approaches, and
there were none at higher levels.
It is clear that the hearing thresholds appreciably
increased for all signals throughout the frequency range
tested if the animal raised its head out of water and the
sound was conducted through the body. The differences
were minimal for tone and narrowband signals at
10 kHz and increased with frequency. Quite unex-
pected was the great drop of through-body perception
at 5 kHz, especially for the tone signal (38 dB); this was
most probably associated with some individual alter-
ations, because another dolphin under the same condi-
tions showed a difference of only 12 dB (working log
data). Note that in an earlier work [7] we found marked
differences in sensitivity only above 60 kHz. With the
5 kHz exception, tone and narrowband signals were
perceived quite similarly (figure). In the natural milieu
(fully under water) the dolphin was equally sensitive to
signals of any structure throughout the frequency range
examined, which is in accord with previous data [3, 7].
Clearly, further work should be both in-depth (regard-
ing signal and noise parameters) and extensive (to take
account of individual peculiarities).

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