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Research Article
Cuvier's beaked whale (Ziphius cavirostris) head tissues: physical properties and CT imaging
Melissa S. Soldevilla, Megan F. McKenna, Sean M. Wiggins, Robert E. Shadwick, Ted W. Cranford, John A. Hildebrand
Journal of Experimental Biology 2005 208: 2319-2332; doi: 10.1242/jeb.01624
Melissa S. Soldevilla
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Megan F. McKenna
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Sean M. Wiggins
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Robert E. Shadwick
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Ted W. Cranford
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John A. Hildebrand
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Figures

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  • Fig. 1.
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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.

  • Fig. 2.
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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.

  • Fig. 3.
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    Fig. 3.

    (A) Head of a neonate Z. cavirostris. Arrowheads point to transverse slices (A–O) 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.

  • Fig. 4.
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    Fig. 4.

    Sound speed versus temperature from Ziphius cavirostris head tissues.

  • Fig. 5.
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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 s–1), density (1026 kg m–3) and HU (0), respectively, of seawater at 15°C and 101 kPa.

  • Fig. 6.
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    Fig. 6.

    Sound speed through 13 transverse slices (A–M) 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 s–1).

  • Fig. 7.
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    Fig. 7.

    Stress–strain 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.

  • Fig. 8.
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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 (anterior–posterior, dorsal–ventral, lateral).

  • Fig. 9.
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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.

  • Fig. 10.
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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.

  • Fig. 11.
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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.

Tables

  • Figures
  • Table 1.

    List of samples used in the physical properties analysis of a Ziphius cavirostris neonate head

    Sample nameTissue categorySlice nameSamples/slice
    Left mandible acoustic blubberBlubberA–J1
    Left mandible external acoustic fatAcoustic fatA–L1
    Left mandible internal acoustic fatAcoustic fatA1
    Left mandible internal acoustic fatAcoustic fatB2
    Left mandible internal acoustic fatAcoustic fatC2
    Left mandible internal acoustic fatAcoustic fatD2
    Left mandible internal acoustic fatAcoustic fatE4
    Left mandible internal acoustic fatAcoustic fatF5
    Left mandible internal acoustic fatAcoustic fatG5
    Left mandible internal acoustic fatAcoustic fatH6
    Left mandible internal acoustic fatAcoustic fatI6
    Left mandible internal acoustic fatAcoustic fatJ5
    Left mandible internal acoustic fatAcoustic fatK7
    ForeheadVariableA3
    ForeheadVariableB6
    ForeheadVariableC9
    ForeheadVariableD11
    ForeheadVariableE13
    ForeheadVariableF17
    ForeheadVariableG17
    ForeheadVariableH20
    ForeheadVariableI23
    ForeheadVariableJ16
    ForeheadVariableK31
    ForeheadVariableL20
    ForeheadVariableM14
    ForeheadVariableN/O3
  • Table 2.

    Results of sound speed–temperature analysis for various tissue types from the head of a Ziphius cavirostris neonate

    Acoustic fat: foreheadAcoustic fat: mandibleBlubber: foreheadBlubber: mandibleConnective tissueMuscle
    Slope (m)-4.4225-7.113-4.1655-4.80810.4842-0.116
    r 2 0.9790.9460.9320.953–0.080
    • Slopes and r2 values of the best-fit line are presented.

  • Table 3.

    Physical property measurements of Ziphius cavirostris forehead and mandibular tissues by tissue type

    Forehead samples Mandible samples
    Blubber (N=36)Acoustic fats (N=66)Muscle (N=13)Connective tissue (N=30)Blubber (N=11)Acoustic fats (N=44)
    Sound speed at 37°C (m s-1)
       Mean140113821517162813761350
       s.d.7.82346.848.71310.6
       Min.138213411460153813451331
       Max.142214091592170813941378
    Sound speed at 21°C (m s-1)
       Mean146514501522162014531460
       s.d.8.317.647.248.78.810.6
       Min.144614121461153114411442
       Max.148914831594170014681488
    Density (kg m-3)
       Mean9359379931087919890
       s.d.251758411323
       Min.8829089091007901854
       Max.98698710661175937941
    Hounsfield unit
       Mean-84-81-3345-102-78
       s.d.15122345327
       Min.-106-96-61-36-106-106
       Max.-46-472109-95-65
    Hysteresis (%)
       Mean28.432.5–53.521.729.4
       s.d.11.29.2–16.07.814.1
       Min.0.615.5–11.211.65.6
       Max.46.951.4–77.133.088.8
  • Table 4.

    Mean elastic modulus (kPa) at selected stresses in the various tissue types of the forehead and mandible of a Ziphius cavirostris neonate

    Forehead tissues Mandible tissues
    Stress (kPa)BlubberAcoustic fatConnective tissueBlubberAcoustic fat
    2.5597512415878
    5208153149220215
    10411212279400303
    20712412618650420
    3010066261030913587
    4012007391295991726
    50141091015101174934
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Research Article
Cuvier's beaked whale (Ziphius cavirostris) head tissues: physical properties and CT imaging
Melissa S. Soldevilla, Megan F. McKenna, Sean M. Wiggins, Robert E. Shadwick, Ted W. Cranford, John A. Hildebrand
Journal of Experimental Biology 2005 208: 2319-2332; doi: 10.1242/jeb.01624
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Research Article
Cuvier's beaked whale (Ziphius cavirostris) head tissues: physical properties and CT imaging
Melissa S. Soldevilla, Megan F. McKenna, Sean M. Wiggins, Robert E. Shadwick, Ted W. Cranford, John A. Hildebrand
Journal of Experimental Biology 2005 208: 2319-2332; doi: 10.1242/jeb.01624

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