First published online September 19, 2006
Journal of Experimental Biology 209, 3758-3765 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02431
The spectral sensitivity of the lens eyes of a box jellyfish, Tripedalia cystophora (Conant)
Melissa M. Coates1,*,
Anders Garm2,
Jamie C. Theobald2,
Stuart H. Thompson1 and
Dan-Eric Nilsson2
1 Hopkins Marine Station, Department of Biological Sciences, Stanford
University, Oceanview Boulevard, Pacific Grove, California, 93950,
USA
2 Department of Cell and Organism Biology, Lund University, Zoology
Building, Helgonavägen 3, S-223 62 Lund, Sweden

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Fig. 1. (A) Photograph of Tripedalia cystophora. There are four rhopalia
located on the sides of the bell (arrow), which alternate with the four groups
of tentacles at the corners. The lens eyes point inward, toward the center of
the bell. Scale bar, 1 cm. (B) A photograph of a single rhopalium inside the
rhopalial niche. Arrows indicate a pit eye and a slit eye. The slit and pit
eyes are both identically matched on the other side of the rhopalium. (C) A
photograph of an isolated rhopalium. Arrows indicate the upper and lower lens
eyes. Scale bar, 200 µm.
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Fig. 2. A sample ERG trace showing the response of an isolated rhopalia to a flash
of white light, 40 ms in duration, at 5.47x103 W
m-2 sr-1 (ND 0.5). Peak response is taken as first peak
(a), regardless of polarity. It is assumed that the first response is from
photoreceptors and subsequent peaks (b) may be from downstream events.
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Fig. 3. A single V-logI curve from a lower lens eye. Data points (open
circles) show how the response changed with changing light intensity (I; W
m-2 sr-1), while the model fit (solid line) shows the
sigmoid shape of this response. For each preparation the
V-logI-based model fit was used to calculate spectral
sensitivity for that preparation (see text for details).
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Fig. 4. Response characteristics change with intensity of the stimulus. Here, an
ERG trace showing the response to a flash of white light at
5.47x103 W m-2 sr-1 (ND 0.5, black
trace) is noticeably biphasic. However, when the same preparation was
stimulated by a much lower intensity (1.73x101 W
m-2 sr-1, ND 3.0), the response appears more monophasic
and is of smaller magnitude (gray trace). Inset shows onset (1.5 s) and
duration (40 ms) of flash.
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Fig. 5. Normalized response versus stimulus intensity (W m-2
sr-1) for the lower and upper eyes (V-logI). Values are
means ± 1 s.e.m. Solid line, lower eye (N=14); broken line,
upper eye (N=8).
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Fig. 6. Mean spectral sensitivity curves for both the lower (solid line, open
circles, N=14) and the upper (broken line, open triangles,
N=8) lens eyes (values are means ± 1 s.e.m.). When plotted on
the same graph the similarity, in both the peak sensitivity (500 nm) and the
shape of the curves, between the two eye types is apparent.
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Fig. 8. Removing the ß-peak from the GFRKD visual pigment template
optimization allows for a much better fit in both eye types (gray lines;
correlations=0.984, lower eye; 0.968, upper eye). With invertebrate levels of
self-screening (k=0.0067; A,B, black lines) the curves are
indistinguishable from the no screening case (A,B, gray lines). However, these
fits can be improved further by adding vertebrate level self-screening to the
optimization (C,D, black lines); where k= 0.035 µm-1,
and l=50 µm for the lower eye photoreceptors and l=35
µm for the upper eye photoreceptors. Here, self-screening has the effect of
broadening the absorption curve. For the lower eye (C) the half-width
increases to 112.1 nm with the addition of screening, compared to 82.0 nm
without screening. For the upper eye (D) the half-width increases to 111.2 nm
from 82.8 nm. Correspondingly the correlation values increase to 0.991 and
0.980 for the lower and upper eyes, respectively.
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© The Company of Biologists Ltd 2006