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First published online July 2, 2004
Journal of Experimental Biology 207, 2803-2810 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.01078

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A unique visual pigment expressed in green, red and deep-red receptors in the eye of the small white butterfly, Pieris rapae crucivora

Motohiro Wakakuwa¹, Doekele G. Stavenga², Masumi Kurasawa¹ and Kentaro Arikawa^1,*

¹ Graduate School of Integrated Science, Yokohama City University, Yokohama 236-0027, Japan
² Department of Neurobiophysics, University of Groningen, Groningen, The Netherlands

View larger version (72K): [in a new window]   Fig. 1. Nucleotide and deduced amino acid sequence of the PrL opsin of Pieris rapae crucivora. Single- or double-headed arrows indicate the regions corresponding to primer attachment: the sequences of degenerate primers are TTGAAGCTTCARTTYCCNCCNATGAAYCC (L01F, forward primer for QFPPLNP), TTGAAGCTTAYGTNCCIGARGGNAAYATGAC (L02F, forward primer for PEGNMTA), CGAATTCGTCATRTTNCCYTCIGGNACRTA (L02R, reverse primer for PEGNMTA) and CGAATTCGGRTTRTANACIGCRTTNGCYTT (L03R, reverse primer for KANAVYNP). Forward and reverse primers have a HindIII site and EcoRI site (underlined), respectively.

View larger version (36K): [in a new window]   Fig. 2. Phylogenetic relationship of insect visual pigment opsins calculated using the neighbour joining method with octopus opsin as the outgroup. PrL is grouped in the long-wavelength absorbing type (arrow). Data accession numbers: Apis mellifera AMBLOP=AF004168; Apis mellifera AMLOP=AF091732; Apis mellifera AMUVOP=AF004169; Camponotus abdominalis LW=U32502; Camponotus abdominalis UV=AF042788; Cataglyphis bombycina LW=U32501; Cataglyphis bombycinus UV=AF042787; Drosophila melanogaster Rh1=K02315; Drosophila melanogaster Rh2=M12896; Drosophila melanogaster Rh3=M17718; Drosophila melanogaster Rh4=P08255; Drosophila melanogaster Rh5=U67905; Drosophila melanogaster Rh6=Y00043; Manduca sexta Manop1=L78080; Manduca sexta Manop2=L78081; Manduca sexta Manop3=AD001674; Papilio glaucus PglRh1=AF077189; Papilio glaucus PglRh2=AF077190; Papilio glaucus PglRh3=AF067080; Papilio glaucus PglRh3=AF098283; Papilio glaucus PglRh4=AF077193; Papilio glaucus PglRh5=AF077191; Papilio glaucus PglRh6=AF077192; Papilio xuthus PxB=AB028217; Papilio xuthus PxUV=AB028218; Papilio xuthus PxL1=AB007423; Papilio xuthus PxL2=AB007424; Papilio xuthus PxL3=AB007425; Schistocerca gregaria Lo1=X80071; Schistocerca gregaria Lo2=X80072; Sphodromantis sp. LW=X71665.

View larger version (86K): [in a new window]   Fig. 3. In situ hybrization of the PrL mRNA of Pieris rapae crucivora. (A) Diagrammatical sketch of a Pieris ommatidium. Arrowheads indicate the planes where the transverse sections B,D and C,E were obtained. (B) Non-stained transverse section through the distal tier. Roman numbers indicate the ommatidial type I, II and III. (C) Unstained transverse section of the identical region of the eye shown in B but through the proximal tier. (D) In situ hybridization of PrL mRNA on a transverse section through the distal tier. R3 and R4 photoreceptors in all ommatidia were labelled. (E) PrL mRNA in situ hybridization in the proximal tier. R5–8 photoreceptors in all ommatidia were labelled. Scale bars, 10 µm.

View larger version (17K): [in a new window]   Fig. 4. Spectral sensitivities of Pieris photoreceptors and absorbance spectra of screening pigments. (A) Log spectral sensitivity data (error bars are S.E.M.) of green sensitive photoreceptors determined by intracellular recordings of R3,4 photoreceptors of type I and III ommatidia (open symbols; L560) and type II ommatidia (filled symbols; L560-II); from Qiu and Arikawa (2003a). The continuous green line gives the absorption spectrum of a visual pigment with peak absorption at 563 nm. (B) Log spectral sensitivity data (error bars are S.E.M.) determined by intracellular recordings on R5–8 photoreceptors of type I and III ommatidia (open symbols; L620) and type II ommatidia (filled symbols; L640); from Qiu and Arikawa (2003b). The dashed and dotted lines give log(1–Tp) and log(1+Tp), respectively, where Tp is the transmittance of the proximal rhabdom tier, length Lp=150 µm, diameter Dr=2.0 µm, with a visual pigment R563, with maximum absorption coefficient max=0.006 µm–1, in all R5–8 photoreceptors. The dot/dash green line gives the absorbance due to the visual pigment in photoreceptors R3 and R4 of the distal rhabdom tier, Adv=–log(Tdv), where Tdv is the transmittance. The summed volume fraction of R3 and R4 is assumed to be f3,4=0.3 (see Materials and methods). The used spectral sensitivity data are normalized to the peak value, meaning that the log sensitivity data have a zero peak value. The log sensitivity data of L620 and L640 were shifted 0.66 and 1.1 log unit, respectively, to fit the log (1–Tp) spectrum at the longest four (L620) or three (L640) wavelength values. (C) Absorbance spectra of screening pigments in the pale-red (PR) ommatidia and the deep-red (DR) ommatidia derived by subtracting the log spectral sensitivities of L620 and L640 from the log (1–Tp) spectrum; the error bars are taken to be identical to those in B.