Ted W.

Cranford, Biology Department, San Diego State University,

tcranfor@mail.sdsu.edu
Petr Krysl, Structural Engineering, University of California at San Diego,
pkrysl@ucsd.edu
John A. Hildebrand, Marine Physical Lab, University of California San Diego,
jhildebrand@ucsd.edu

This lay-language submission is based on two presentations:

“Rivers” of sound in Cuvier’s beaked whale (Ziphius cavirostris): implications for the
evolution of sound reception in odontocetes.
4aAB5 - http://asa.aip.org/web2/asa/abstracts/search.may09/asa782.html
Vibration characteristics of the tympanoperiotic complex in the bottlenose dolphin,
Tursiops truncatus.
4aAB4 - http://asa.aip.org/web2/asa/abstracts/search.may09/asa781.html

THURSDAY MORNING, 21 MAY 2009 GALLERIA NORTH, 8:25 TO 11:50 A.M.

Session 4aAB
Animal Bioacoustics: General Topics in Animal Bioacoustics I

Investigating the physiology of sound reception in the largest animals in the world is
problematic. Their gargantuan size and marine habitat often makes them difficult to find
or access, impossible to handle alive; quite simply intractable subjects for study. The
rapid advance of blazing fast computer hardware, giant industrial x-ray CT scanners, and
modern mathematical and engineering approaches to building computer models, have
allowed us to pierce the seeming impenetrable veil around the bioacoustics of study large
whales bioacoustics.

The bioacoustics of living marine resources has become an important topic in the last
decade, initiated by stranded whales associated with Navy sonar, heightened public
concern, and at least one court case reaching all of the way to the United States Supreme
Court. Recently, evidence has surfaced that suggests human-generated sound can have
detrimental effects on fish hearing, reproductive habits, and stress levels.

We have developed a suite of techniques that combine to produce a “finite element

model” of a whale’s head. This model allows us to simulate what happens when the
anatomy of the whale interacts with sound. We are now capable of deciphering the
physics and physiology of sound production and sound reception in the head of an adult
male Cuvier’s beaked whale (Ziphius cavirostris).

The first step in the process was to develop a method for collecting a representation of in
situ anatomic structure, the anatomic geometry, from large whales. Currently, we are the
only group in the world that has surmounted this daunting challenge. The central key to
the solution is rocket science! More specifically, the technology used to inspect solid
fuel rockets for flaws, an industrial x-ray CT scanner, can be used to scan a dead whale.

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Not just any dead whale, we wait for an opportunity when we know a whale has recently
beached itself alive and then subsequently died. If we can get to the recently deceased
behemoth and cover it with ice before it gets too far along in the decomposition process,
then we can salvage the specimen in good enough condition to scan it and eventually
build a computer model from its carcass.

Industrial x-ray CT scanner at Hill Air Force Base is normally used to scan solid fuel rocket motors. In this
instance, it will scan a frozen whale head contained within a sarcophagus built specifically for this purpose.

After scanning the head of a Cuvier’s Beaked Whale, we published a report about the
structure of its biosonar anatomy. (Cranford TW, et al., (2008), Anatomic Geometry of
Sound Transmission and Reception in Cuvier's Beaked Whale (Ziphius cavirostris). Anat
Rec 291:353-378). That paper was the first quantitative description of biosonar anatomy
for any toothed whale.

After the scanning process we dissect the specimen, take it apart systematically, and
measure the elastic properties of the various tissues. These values for tissue elasticity and
the tissue density values given by the CT scans are the two primary building blocks for
the computer model. The toolbox that takes the building blocks and constructs the model
is the computer software. Our finite element modeling software, the Vibroacoustic
Toolkit, was custom built for this purpose.

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Computer simulations led to another published paper that reported the discovery a new
channel for sound reaching the ears in Cuvier’s Beaked Whale. (Cranford TW, Krysl P,
Hildebrand JA. (2008). Sound pathways revealed: Simulated sound transmission and
reception in Cuvier’s beaked whale (Ziphius cavirostris). Bioinsp. Biomim. 3:1, 1-10.).

The computer simulations show that most sounds arriving from in front of the animal,
enter the head from underneath the tongue region, passes through the throat and an
opening in the posterior part of hollow lower jaw, and propagate along a fat body to the
ear.

These views of the jaws (magenta) from a Cuvier’s Beaked Whale are reconstructed from x-ray CT scans.
The image on the left shows that the rear portion of the jaws is hollow and there is no bony wall on the
inside. This is even more evident in the image on the right, because the fat body (yellow) is clearly seen
because as the bony wall is missing. The ear complex (red) is also shown.

In order for this sound reception pathway to function, the bony wall on the inside of the
lower jaws must be absent; we say, “The door must be open.” As it turns out, all living
toothed whales have this open door. And some of the earliest fossils also show the same
hollow jaw with the open door. This suggests that this sound reception pathway
developed early in the evolution of whales.

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Ancient whales are known as archaeocetes. The fossil whale shown in the above image is Basilosaurus
cetoides. It lived approximately 35-40 million years ago. Notice that it has the same hollow (asterisk) in
the rear portion of the lower jaw, and perhaps more importantly, the same missing wall of bone.

The computer model allows us to visualize the pathway or “river” of sound as it “flows”
from the front of the head to the ear complex.

This image shows a view from the underside of the head. The arrows show the generalized pathway
(rivers) for sound passing from in front of Cuvier’s Beaked Whale, underneath the jaws (magenta), through
the fat body (yellow), and to the ears (red).

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These images show the anatomy of the bony ear complex (tympanoperiotic complex or TPC) and
predictions for how it vibrates. Panel A shows a left lateral view of the TPC, the periotic bone (salmon),
the tympanic bone (cyan), and the middle ear bones or ossicles (malleus=yellow, incus=magenta,
stapes=green). Panel C shows the TPC upside down with one side removed to facilitate visualizing the
ossicles (same color map). Panels B & D show the TPC subdivided into finite elements whose properties
are known. The analysis predicts the vibrational patterns for the entire TPC at 69 kHz. The lighter colors
indicate higher amplitude motions of elements.

Once the river of sound reaches the ear complex, the computer model can allow us to
predict how the bony ear complex will vibrate as a result of being stimulated by the
incoming sound. This is known as vibrational analysis. A series calculation It describes
how all of the elements stiffness and mass features interact to produce the collective
motion of the entire structure based on elementary physical principalsprinciples.

The TPC collects sound from the gular pathway (river) and transmits it to the cochlea
(inner ear). Analysis suggests that Navy sonar sounds reach the ears with reduced
amplitude but the frequencies used by these animals to catch prey are amplified.

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This composite image tells us about the received sound pressure over the surface of the TPC with respect to
the sound incident on the head. The image on the right shows the TPC of Cuvier’s Beaked Whale
reconstructed from CT scans. The twenty four subpanels on the left show where the relative pressure
“pushes” on the TPC after flowing through the river. Relative pressures are color coded (Green =1=
pressure is equal to the sound pressure incident on the head. Blue = 0 = sound pressure is below incident
pressure. 2=red=sound pressure is twice the incident pressure). Interestingly, the first two subpanels
(outlined with a red line) indicate that the frequencies in the range produced by U.S. Navy’s Mid Frequency
Active Sonar are largely filtered out (i.e. blue indicates less sound pressure than incident) before they reach
the ear complex! The other subpanels (7.5 kHz to 42.5 kHz) show that those frequencies and locations that
are particularly effective at driving the vibrations of the bony ear complex (TPC). These frequencies are
also those that are produced by these animals in the wild for the purposes of biosonar.

Computer models are powerful tools for investigation and discovery of bioacoustic
physiology. This is particularly valuable because it provides a means to simulate across a
broad range of scales and taxonomies, from whales to fish. Virtual experiments can also
determine potential for, and mechanism(s) of physical injury. Our techniques also
provide potential for evaluating and directing mitigation efforts.

We acknowledge the continued support of Frank Stone and Ernie Young at the Chief of
Naval Operations N45; Curt Collins from the Naval Postgraduate school, as well as
support from Mike Wiese and Bob Gisiner at the Office of Naval Research.