First published online June 6, 2005
Journal of Experimental Biology 208, 2319-2332 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01624
Cuvier's beaked whale (Ziphius cavirostris) head tissues: physical properties and CT imaging
Melissa S. Soldevilla1,*,
Megan F. McKenna2,
Sean M. Wiggins1,
Robert E. Shadwick1,
Ted W. Cranford2 and
John A. Hildebrand1
1 Scripps Institution of Oceanography, University of California, San Diego,
9500 Gilman Drive, La Jolla, CA 92093-0205, USA
2 San Diego State University, 5500 Campanile Drive, San Diego, CA 92182,
USA

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Fig. 1. (A) Reconstructed computer image of a neonate Z. cavirostris head.
Dark grey areas show the outer skin layer. The light blue region is bone, i.e.
the skull and mandible. The white area is the connective tissue theca that
encompasses the melon, seen in yellow. Mandibular fat bodies are also shown in
yellow. Colorized CT scans of transverse slices through (B) a posterior region
and (D) an anterior region of the head. C and E represent line drawings that
diagram the body parts seen in scans B and D, respectively. Abbreviations: b,
blubber; ct, connective tissue; ma, maxilla; me, melon; mf, mandibular fat;
mn, mandibular bones; mu, muscle. Only interior mandibular fat is represented
in these images.
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Fig. 2. Ziphius cavirostris skull. The concave skull basin where the melon
is located is readily apparent. The hollowed mandibles, which house the
acoustic fats important to sound reception, are also noted.
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Fig. 3. (A) Head of a neonate Z. cavirostris. Arrowheads point to
transverse slices (AO) that were taken through the melon. (B) Anterior
side of a transverse slice through the melon, which corresponds to slice K,
from A. Skin, blubber, connective tissue, muscle and melon tissues are noted.
(C) Transverse melon slice K, showing grid-like sampling of tissues into
numbered cubes. (D) Transverse CT scan image of slice K. Maxillary and
mandibular bones are visible as bright white areas. Between these is the
tongue. A blubber sheath can be seen surrounding the animal. The connective
tissue, muscle and melon fats can be discerned in the upper portion of the
figure. Interior and exterior mandibular fats can be seen inside of and around
the mandibular bones. Throat grooves are also visible.
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Fig. 4. Sound speed versus temperature from Ziphius cavirostris
head tissues.
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Fig. 5. Box and whisker plots of (A) sound speed corrected to 37°C, (B) mass
density and (C) CT scan Hounsfield units for various tissue types in the
forehead and mandible. The lines of the boxes represent the lower-quartile,
median and upper-quartile values. The whiskers extending from each end of the
box show the extent of the remaining data. Outliers are represented by + signs
beyond the whiskers. The dark blue horizontal lines represent the sound speed
(1507 m s1), density (1026 kg m3) and HU
(0), respectively, of seawater at 15°C and 101 kPa.
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Fig. 6. Sound speed through 13 transverse slices (AM) of Ziphius
cavirostris melon. Slices N and O are not shown in this analysis since
they encompass the nasal plugs. The figure shows the anterior face with dorsal
side up. The animal's right (R) and left (L) sides are as shown in the figure.
The color bar scale represents sound speed values (m
s1).
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Fig. 7. Stressstrain curve for a sample of Ziphius cavirostris
forehead blubber. The 10 low-stress pulses are visible at the low end of the
curve, while the larger curve represents the single high-stress pulse.
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Fig. 8. Elastic modulus versus stress for tissues from a Ziphius
cavirostris forehead and mandible. Blubber, acoustic fats and connective
tissues are plotted for each location. Note that values have been averaged
across three orientations (anteriorposterior, dorsalventral,
lateral).
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Fig. 9. Results of a principal components analysis showing the loadings of each
variable for the first two principal components. Density, Hounsfield unit and
sound speed are all described primarily by the 1st principal component, while
elastic modulus is described by the 2nd principal component.
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Fig. 10. Scatter plot of 1st and 2nd principal component loadings for each
Ziphius cavirostris tissue sample. Lipids, muscle and connective
tissue can be distinguished from each other by the first component, while
acoustic fats are distinguished from blubber by the second component.
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Fig. 11. Regression analysis of Hounsfield units versus (A) sound speed and
(B) density for Ziphius cavirostris forehead tissues. Best-fit lines
are represented by the following equations: sound speed=1.6889HU+1535.9;
density=0.001HU+1.0265.
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© The Company of Biologists Ltd 2005