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First published online July 14, 2008
Journal of Experimental Biology 211, 2388-2396 (2008)
Published by The Company of Biologists 2008
doi: 10.1242/jeb.015040
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Nitric oxide induces aspects of egg-laying behavior in Aplysia

Nimrod Miller1, Ayelet Katzoff1,2 and Abraham J. Susswein1,2,*

1 Leslie and Susan Gonda (Goldschmied) Multidisciplinary Brain Research Center, Bar Ilan University, Ramat Gan, 52900, Israel
2 Mina and Everard Goodman Faculty of Life Sciences, Bar Ilan University, Ramat Gan, 52900, Israel


Figure 1
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  		Fig. 1. Examples of egg cordons deposited after treatment. (A) An egg cordon produced in response to the NO donor SNAP (S-nitroso-N-acetyl-penicillamine) that began with empty egg capsules and ended with capsules filled with eggs. The eggs are filled with yolk, which is yellow. The portions of the egg cordon deposited earlier are white, reflecting the lack of yolk and eggs in the capsules. Note that both the white and the yellow portions of the cordon are knotted and raveled, reflecting identical tamping and weaving behaviors that accompany egg deposition. (B) A section of an egg cordon produced in response to BC homogenate, showing capsules filled with numerous eggs. (C–E) Sections of egg cordons laid in response to NO. (C) The cordons contain empty capsules; this is the most common appearance of the cordons. (D) At the start of egg-laying, cordons sometimes lacked capsules. (E) As deposition of a cordon similar to C progressed, eggs sometimes began to appear in some capsules. Scale bar in D is for B–E.

 

Figure 2
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  		Fig. 2. Latency and duration of egg-laying in response to a number of stimuli. (A) Latency to the start of egg-laying in response to the NO donor, and in response to large hermaphroditic duct (LHD) homogenate, bag cell (BC) homogenate, and egg-laying hormone (ELH). Data for these treatments were combined. Latency in response to the NO donor was significantly shorter than in response to LHD homogenate, BC homogenate and ELH. (B) The duration of egg laying in response to the NO donor versus the other treatments. The duration of egg-laying in response to the NO donor is significantly shorter than in response to the BC homogenate. (C) Application of the NOS (nitric oxide synthase) inhibitor L-NAME (N{omega}-nitro-L-arginine methyl ester) significantly increased the duration of egg-laying in response to the LHD homogenate.

 

Figure 3
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  		Fig. 3. (A) Diagram of the reproductive system of Aplysia showing the sites tested for localization of the effect of NO. (B) Analysis of the effect of the NO donor application to the abdominal ganglion. (i) Effect of application of the NO donor on the activity recorded extracellularly using a suction electrode placed on the cut pleural-abdominal connective, close to the bag cells. The two traces are continuous. The stimulus did not elicit a bag cell after-discharge. (ii) Electrical stimulation of the connective (in the example shown, 7 V, 7 Hz, 5 ms pulse duration, maintained for 5 s) induced an after-discharge. The three traces are continuous. The recordings began a few seconds after the stimulus, since the same suction electrode was used for stimulation and recording. The stimulating electrode was removed from the tubing and the recording electrode was then inserted.

 

Figure 4
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  		Fig. 4. Appetitive and consummatory behaviors elicited by artificial sea water (ASW), and by the NO donor, in A. californica during the winter months, when most of them are immature. Data are shown separately for animals that responded to the NO donor by depositing an egg cordon (mature), and for animals in which no egg cordon was produced (immature). (A) The NO donor elicited a large increase in appetitive behaviors, with respect to that observed in response to ASW. In mature animals the appetitive behaviors stopped abruptly at the start of the consummatory behaviors, whereas in immature animals appetitive behaviors gradually declined over the hour of observation. (B) Consummatory behaviors were observed in mature animals.

 

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© The Company of Biologists Ltd 2008