First published online October 31, 2008
Journal of Experimental Biology 211, 3536-3543 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.022277
Reflected polarization guides chironomid females to oviposition sites
Amit Lerner1,*,
Nikolay Meltser2,
Nir Sapir3,
Carynelisa Erlick1,
Nadav Shashar4 and
Meir Broza2
1 Department of Atmospheric Sciences, The Hebrew University of Jerusalem,
Jerusalem 91904, Israel
2 Faculty of Science and Science Education, University of Haifa at Oranim, Tivon
36006, Israel
3 Department of Evolution, Systematics and Ecology, The Hebrew University of
Jerusalem, Jerusalem 91904, Israel
4 Department of Life Sciences, Eilat Campus, Ben-Gurion University, Hatmarim
St., Elat 88000, Israel

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Fig. 1. An egg trap used in experiment 1. The trap consisted of a 30 cmx30
cmx30 cm wooden box, on the bottom of which was a 10 W bulb. Above the
bulb there was a 10 cmx10 cm open hatch with a 15 cmx15 cm filter
(black rectangle). The filter included diffusers and a plane polarizer that
provided polarized/unpolarized illumination at two levels of intensity. A 15
cmx15 cmx10 cm glass aquarium filled with 2 cm (450
cm3) of tap water was placed on the filter.
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Fig. 2. Intensity (solid lines) and percentage polarization (dashed lines) of
radiation measured in the range 490—510 nm reflected from the egg traps
in experiment 1. The light source was a 10 W bulb. PLIL, polarization low
(unpolarized) intensity low [1 polarizing sheet (1p) + 15 diffusing sheets
(15d)] ordered from light source to viewer; PLIH, polarization low intensity
high (1p + 3d); PHIL, polarization high (polarized) intensity low (15d + 1p);
PHIH, polarization high intensity high (3d + 1p).
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Fig. 3. The spectrum of intensity (dashed grey line, measured without a polarizing
filter), percentage polarization (black solid line) and e-vector orientation
(grey solid line) reflected from the pond's surface, measured on 17 October
2007 at 16.23 h local time (UTC+2 h). The sun was obscured by the Carmel Ridge
from 16.00 h and sunset time was 17.05 h. The sensor was directed 45 deg. from
the nadir and 230 deg. from the north. The intensity, percentage polarization
and e-vector orientation calculated from the readings integrated over the
range 490—510 nm were 2.8x10—3
µEcm—2 s—1, 71.5%, and 4.4 deg.,
respectively (where 0 deg. represents horizontal alignment).
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Fig. 4. Mean ± s.d. of the intensity (A) and percentage polarization (B),
reflected from the four tubs in experiment 2, as a function of wavelength in
the range 490—510 nm. WT, black tub painted white up to water level
filled with tap water (N=10; 10 tubs, one measurement from each tub);
WP, black tub painted white up to water level filled with pond water
(N=10); BT, black tub filled with tap water (N=18); and BP,
black tub filled with pond water (N=18). For the reflected intensity,
only WT differed significantly from the other tubs (Kruskal—Wallis
non-parametric ANOVA; P<0.05). For the reflected percentage
polarization, only BT and BP did not differ significantly from each other
(pair-wise comparisons, one-way ANOVA, P<0.001). None of the
e-vector orientations reflected from the tubs differed significantly from 0
deg. (horizontal).
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Fig. 5. Intensity (triangles), percentage polarization (circles), and e-vector
orientation (diamonds) of the radiance reflected from the pond water surface
during sunset, measured on December 10, 2007. Values were measured for the
wavelength range 490—510 nm. The sun was obscured by the Carmel Ridge at
15.57 h local time (black vertical bar) and sunset time was 16.36 h (grey
vertical bar). Complete darkness occurred at 17.01 h. The e-vector orientation
did not differ from horizontal (0±4 deg.). Note that the light
intensity decreased by 96%, while the percentage polarization remained stable
at around 60%.
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Fig. 6. Chironomid egg batch (EB) types and their relative contribution (%) to the
total eggs laid in experiment 1. (A) An unidentified genus of chironomids
(0.1%), (B) Kiefferulus sp. (6.3%), (C) Chironomus
transvaalensis (83.5%), and (D) Polypedilum nubifer (10.1%).
Note that in experiment 2 conducted outdoors, only C. transvaalensis
eggs were found.
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Fig. 7. Mean ± s.d. of percentages of egg batches (EB) laid in artificial
egg traps in experiment 1 (total EB=10592; N=10 repeats). Planned
comparison of means of PLIL vs PLIH, PLIL vs PHIL, PHIL
vs PHIH, and PLIH vs PHIH were found significant in all four
pairs (P<0.005). PLIL, polarization low (unpolarized) intensity
low; PLIH, polarization low intensity high; PHIL, polarization high
(polarized) intensity low; PHIH, polarization high intensity high.
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Fig. 8. Mean ± s.d. of percentages of egg batches (EB) laid in the tubs in
experiment 2 (total EB=3543; N=16 repeats). A Bonferroni
post-hoc test revealed a significant difference between all
treatments (P<0.05). Note that EB laid in WT (black tub painted
white up to water level and filled with tap water) was not significantly
different from 0%. WP, black tub painted white up to water level filled with
pond water; BT, black tub filled with tap water; BP, black tub filled with
pond water.
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Fig. 9. Intensity (triangles), percentage polarization (circles), and e-vector
orientation (diamonds) of the radiance reflected from a 70 l white barrel
filled with 60 l of pond water versus the total organic carbon (TOC)
concentration measured in the water. Values were measured for the wavelength
range 490—510 nm. The e-vector orientation did not differ significantly
from horizontal (—6±17 deg.). A value of TOC=0 mg
l—1 was excluded, since the corresponding reflected radiance
was almost unpolarized (percentage polarization=2.6%).
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Fig. 10. (A) Transmission electron micrograph (TEM) of a transverse section of an
ommatidium from the ventral part (white arrow in B) of the Chironomus
transvaalensis female eye. The double-headed black arrows in A indicate
the general alignment of the microvilli of rhabdomeres R1—R7. Note that
the microvilli of R4 and R7 are orthogonal to the microvilli of R1 and R2, and
both are aligned at 45 deg. with respect to R5,R6 and R3, providing the
anatomical basis for linear polarization detection.
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© The Company of Biologists Ltd 2008