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First published online June 29, 2006
Journal of Experimental Biology 209, 2794-2803 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02307
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Evidence for a role of orcokinin-related peptides in the circadian clock controlling locomotor activity of the cockroach Leucophaea maderae

Sabine Hofer and Uwe Homberg*

Fachbereich Biologie, Tierphysiologie, Philipps Universität Marburg, D-35032 Marburg, Germany


Figure 1
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  		Fig. 1. Confocal laser images obtained from vibratome sections of the accessory medulla. Orcokinin immunoreactivity is shown in green (Ai,Bi,Ci,Di) and Texas Red dextran (TRed-D) fluorescence in red (Aii,Bii,Cii,Dii). (Aiii,Biii,Ciii,Diii) Colocalization of orcokinin immunoreactivity and TRed-D fluorescence in yellow. Stacks of 7-11 optical sections (z-distance between single sections=2 µm). (Ai-iii) The TRed-D-injected accessory medulla (outlined by the dotted line). Arrowheads point to the injection site. (Bi-iii) The contralateral accessory medulla (AMe) after injection of TRed-D into the opposite AMe revealed one colabeled orcokinin/TRed-D fluorescent ventral neuron (VNe; arrowheads). (Ci-iii) Section posterior to B. Near the AMe, one ventromedian neuron (VMNe, arrowheads) shows anti-orcokinin/TRed-D colabeling. Colabeled fibers from the injected AMe project via the lobula valley tract (arrows) into the anterior and internodular neuropil of the contralateral AMe. (Di-iii) Section posterior to C. Two additional VMNe (arrowheads) and fibers in the lobula valley tract (arrows) show colabeled anti-orcokinin/TRed-D fluorescence. Scale bars, 50 µm.

 

Figure 2
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  		Fig. 2. Confocal laser images obtained from vibratome sections of (A) the anterior- and (B) posterior optic commissure. Green shows orcokinin immunoreactivity (Ai,Bi), red shows Texas Red dextran (TRed-D) fluorescence (Aii,Bii), after injection of TRed-D into one AMe (see Fig. 1A). The right column (Aiii,Biii) shows colocalization of orcokinin immunoreactivity and TRed-D fluorescence in yellow. Stacks of 10 optical sections (z-distance between single sections=1 µm). (Ai-iii) Orcokinin-ir and TRed-D fluorescent fibers project in parallel via the anterior optic commissure (arrowheads), but do not show colabeling. (Bi-iii) Colabeled orcokinin/TRed-D fluorescent fibers in the posterior optic commissure (arrowheads). Scale bars, 50 µm.

 

Figure 3
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  		Fig. 3. Three-dimensional model of the cockroach brain showing orcokinin-ir (blue, green) and PDF-ir (red) connections between both AMae. Three ventromedian neurons (VMNe, blue, p) project via the posterior optic commissure (POC) into the contralateral AMe and arborize in several median layers of the medulla. One ventral neuron (VNe, green) projects via an unidentified commissure. One PDF-ir VNe (red, p) projects via the POC and two PDF-ir VNe (a) via the anterior optic commissure (AOC) to the contralateral AMe and into the distalmost layer of the medulla (Reischig et al., 2004Go). The 3D model was provided by T. Reischig and modified. Scale bar, 200 µm.

 

Figure 4
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  		Fig. 4. Records of circadian wheel-running activity (A,C) and plots of activity onsets of cockroaches kept in constant darkness (B,D). (A,B) After injection of 150 fmol of Asn13-orcokinin in 2 nl saline at CT14 of day 10 (arrow in A), regression analysis through consecutive activity onsets (B) revealed a phase delay {Delta}{varphi} of 3.7 circadian hours (hct). (C,D) After injection at CT18 of day 10 (arrow in C), the regression analysis through consecutive activity onsets (D) revealed a phase advance {Delta}{varphi} of 2.2 hct.

 

Figure 5
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  		Fig. 5. Orcokinin- and saline-dependent phase shifts at different circadian times (CT; hct). (A) Orcokinin injections (150 fmol in 2 nl saline with blue food dye, N=50) caused maximal phase delays during the early subjective night (up to -3.8 hct at CT13) and maximal phase advances during the middle of the subjective night (up to 2.2 hct at CT18). (B) Control injections (2 nl saline with blue food dye, N=46) caused small and non-significant phase shifts in both directions except for a significant phase delay at CT22. (C) Phase-response curves obtained in response to 150 fmol Asn13-orcokinin and control injections. Data were merged into 2 h bins. Orcokinin-dependent phase shifts (black) and phase shifts following control injections (gray) are plotted. Black asterisks indicate orcokinin-dependent phase shifts that were significantly different (P<0.05) from control injections at the same CT. The grey asterisk indicates control injections that produced phase shifts that were significantly different from zero.

 

Figure 6
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  		Fig. 6. Dose dependency of Asn13-orcokinin-induced phase shifts between CT13 and CT15. Bars show phase shifts resulting from injections of saline (N=6), 1.5x10-6 fmol orcokinin (N=6), 1.5x10-2 fmol orcokinin (N=8), and 150 fmol of orcokinin (N=5) in 2 nl saline. Asterisks indicate orcokinin doses that induced phase shifts significantly different from control injections (P<0.05).

 

Figure 7
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  		Fig. 7. Smoothed phase-response curves for injections of 150 fmol orcokinin (black) compared with the phase-response curve for 6 h light-pulses [blue (Page and Barrett, 1989Go)], GABA (red) and allatotropin (dotted line) injections (Petri et al., 2002Go).

 

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© The Company of Biologists Ltd 2006