First published online December 15, 2004
Journal of Experimental Biology 208, 169-177 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01361
Burrow air phase maintenance and respiration by the mudskipper Scartelaos histophorus (Gobiidae: Oxudercinae)
Heather J. Lee1,*,
Callie A. Martinez2,
Katherine J. Hertzberg3,
Amy L. Hamilton4 and
Jeffrey B. Graham1
1 Center for Marine Biotechnology and Biomedicine and Marine Biology
Research Division, Scripps Institution of Oceanography, University of
California, San Diego, La Jolla, CA 92093-0204, USA
2 Department of Integrative Biology, 3060 VLSB, University of California,
Berkeley, Berkeley, CA 94720-3140, USA
3 Biological Oceanography Division, Scripps Institution of Oceanography,
University of California, San Diego, La Jolla, CA 92093-0236, USA
4 MEC Analytical Systems Inc., 2433 Impala Drive, Carlsbad, CA 92008,
USA

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Fig. 1. Laboratory burrow system with the O2 electrode circulation loop
for continuous measurement of water-phase PO2.
a, mudflat with shallow water; b, connecting tube; c, burrow chamber filled
with water; d, temperature-controlled water bath; e, water-phase outlet; f,
and inlet; g, air phase; h, oscillating pump; i, O2 electrode in
housing; j, N2 gas-equilibration chamber; k, stopcock for air-phase
sampling. Arrows indicate direction of water flow.
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Fig. 2. Record of burrow air-deposition behavior by one S. histophorus
(5.1 g male) over an 8 day period. Diagonal lines show increased air-phase
volume deposited by the fish over time. Vertical lines indicate when all
deposited air was removed by the experimenter. Zero air-phase volume occurred
when the fish was first placed in the system and immediately after each air
phase was withdrawn. Numbers above air-deposition events indicate
air-deposition rate (trips h-1). (A) The three air phases added
during the first 9 h period (burrow-water PO2
indicated in boxes). (B) Entire observation record for the fish showing 14
separate air-phase depositions. (Boxed section is the compressed record shown
in A.)
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Fig. 3. Mean air-deposition rates of S. histophorus over different
treatments (the time intervals over which the fish was left undisturbed)
(N=8 individuals + 2 pairs) based on the volume of air accumulated,
fish buccal chamber volume and the time interval between withdrawals. Values
are means ± S.E.M.
*Significant difference from the 1.5-8 h and >8 h intervals.
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Fig. 4. Hypothesized daily pattern of burrow and surface respiration by S.
histophorus. Horizontal bar shows times of high tide when the burrow
entrance is covered by water (black) and when the burrow is uncovered at low
tide (white). Dashed line shows tidal variation in burrow-water
PO2, which is highest at the time the tide
rises to cover the burrow and lowest when the tide recedes. The fish remains
in the burrow during high tide. Solid black line shows fish aerial
VO2
(Milward, 1974 ); when
burrow-water PO2 is at or below air-breathing
threshold PO2 (horizontal dotted line), the
fish requires air-phase breathing. When burrow-water
PO2 is above threshold, aquatic respiration
would occur. During low tide, the fish is up on the mudflat where it breathes
air. Figure compiled with laboratory data on threshold
PO2 and burrow-water
PO2 decline, and field data for burrow position
in the intertidal zone, low-tide burrow PO2,
and tide tables for the area of study.
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© The Company of Biologists Ltd 2005