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First published online October 17, 2008
Journal of Experimental Biology 211, 3504-3511 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.017848
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Simultaneous color contrast in the foraging swallowtail butterfly, Papilio xuthus

Michiyo Kinoshita1,*, Yuki Takahashi2 and Kentaro Arikawa1

1 Laboratory of Neuroethology, The Graduate University for Advanced Studies (Sokendai), Shonan Village, Hayama 240-0193, Japan
2 Graduate School of Integrated Science, Yokohama City University, 22-2 Seto, Kanazawa-ku, Yokohama 236-0027, Japan


Figure 1
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  		Fig. 1. Illumination and stimuli. (A) Irradiation spectrum of illumination measured as the reflection of MgO-coated surface. (B) Reflectance spectra of colored papers used for the experiments with pale-green (PG)-trained butterflies. (C) Reflectance spectra of colored papers used for the experiments with orange (O)-trained butterflies. (D) Reflectance spectra of background cardboards.

 

Figure 2
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  		Fig. 2. Color loci of color disks and background cardboards on a presumptive three-dimensional color space of Papilio xuthus. Thin lines with numbers show the loci of monochromatic lights. (A) Color loci of two series of color disks and backgrounds, each relating to the training colors; circled letters PG (pale green) and O (orange). Filled circles in the left triangle are colors of disks used for PG training: AB, aqua blue; BG, blue green; FG, forest green; LG, lime green; SG, spring green; YG, yellow green; OG, olive green; GB, grayish blue. Open circles in the left triangle are colors of the background for PG training: B; blue; G, green; MY, mustard yellow; Gr1, gray 1. Filled circles in the right triangle are colors of disks used for O training: Y, yellow; YO, yellow orange; DO, dark orange; R, red; SP, salmon pink; P, pink. Open circles in the left triangle were colors of the background for O training: G, green; V, violet; Gr2, gray 2. (B) Enlarged view of color loci of colors presented with PG for tests 1 and 2. Loci of colors in test 1 are arranged approximately in linear fashion, whereas those in test 2 are distributed around the training color, PG. (C) Enlarged view of color loci of colors presented with O for tests 1 and 2.

 

Figure 3
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  		Fig. 3. Layout patterns of color disks. (A) Training pattern used from post-emergence day 2 to day 5 (1 disk). (B) Five-disk pattern. (C) Seven-disk pattern.

 

Figure 4
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  		Fig. 4. Relative subjective brightness of all color disks and backgrounds, taking the brightness of the black background as 1.0. The dotted lines indicate the brightness of the training colors. (A) Colors related to the pale-green (PG) training. (B) Colors related to the orange (O) training. (C) Action spectrum of foraging behavior of freely flying Papilio [modified from Koshitaka et al. (Koshitaka et al., 2004Go)].

 

Figure 5
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  		Fig. 5. Results of pale-green (PG)-trained Papilio. Colors in test 1 were arranged linearly in the color space, whereas colors in test 2 were arranged radially around PG, the training color (see Fig. 2B). (A) Effect of a blue (B) background. (B) Effect of a mustard yellow (MY) background. The choices were not random in each test. (A) Test 1: Gray (Gr1), H=42.57, P<0.0001; B, H=46.20, P<0.0001. Test 2: Gr1, H=19.46, P<0.01; B, H=20.66, P<0.001. (B) Test 1: Gr1, H=45.54, P<0.0001; MY, H=36.98, P<0.0001. Test 2: Gr1, H=19.46, P<0.01; MY, H=17.43, P<0.001; Kruskal–Wallis test). Each bar shows the mean (±s.e.m.) number of visits. Letters above bars indicate statistical significance. Bars with different letters are significantly different (Tukey's HSD test, P<0.05). Capital letters are used for the results of the control test, and lowercase letters are used for the result of the color contrast tests.

 

Figure 6
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  		Fig. 6. Results of orange (O)-trained Papilio. Colors in test 1 were arranged linearly in the color space, whereas colors in test 2 were arranged radially around O, the training color (see Fig. 2C). (A) Effect of a green (G) background. (B) Effect of a violet (V) background. The choices were not random in each test. (A) Test 1: Gray (Gr2), H=35.64, P<0.0001; G, H=32.73, P<0.0001. Test 2: Gr2, H=21.54, P<0.01; G, H=22.66, P<0.01. (B) Test 1: Gr2, H=44.15, P<0.0001; V; H=29.75, P<0.0001. Test 2: Gr2, H=21.54, P<0.01; V, H=25.31, P<0.0001; Kruskal–Wallis test). Each bar shows the mean (±s.e.m.) number of visits. Letters above bars indicate statistical significance. Bars with different letters are significantly different (Tukey's HSD test, P<0.05). Capital letters are used for the results of the control test, and lowercase letters are used for the result of the color contrast tests.

 

Figure 7
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  		Fig. 7. Prediction of induced colors. (A) Summed spectral sensitivity of the ventral region of the Papilio retina measured by ERG recording [modified from Arikawa et al. (Arikawa et al., 1987Go)]. (B) Difference spectra of receptor excitations between pale green (PG) and the selected color, blue green (BG) with a blue (B) background, and spring green (SG) with a mustard yellow (MY) background. (C) Difference spectra of receptor excitations between orange (O) and the selected color, yellow orange (YO) with a green (G) background, and dark orange (DO) with a violet (V) background.

 

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© The Company of Biologists Ltd 2008