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The ipsilaterally projecting axons from the treated eye are unlikely to have been killed by the drug because, when treated with binocular epibatidine injections, axons from the treated eye grow exuberantly. Because many ganglion cell axons at this age also have sustaining collaterals to the superior colliculus [A. Ramoa et al., Proc. Natl. Acad. Sci. U.S.A. 86, 2061 (1989)], we consider it more likely that the axons withdrew into the optic tract.
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Visual inspection of the projection contralateral to the epibatidine-treated eye in Fig. 2D (red) indicates that some contralateral axons from the treated eye remain within the binocular zone, rather than being entirely excluded by the projection from the active eye. This may result from slight differences in the time of onset of effective blockade or in the exact age, in hours postpartum, of the animal (it is possible that activity blockade more rapidly influences axonal extention than axonal retraction). This scattering of remaining axons from the epibatidine-treated eye accounts for the lack of quantitative difference in area occupied by the contralateral projection when compared to that occupied by the contralateral projection from the saline-injected eye (Fig. 3) (34), because the threshold used for measurements was chosen to give a maximal estimate of the area filled by the projection [see also (34)].
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3H]leucine-labeled retinogeniculate projections were examined under light- or dark-field illumination, respectively, and the distribution of label was recorded digitally with a charge-coupled-device camera mounted on a Nikon SMU-Z microscope attached to a Macintosh computer running NIH Image. The mid-section of each LGN (where, in normal animals, the eye-specific layers are most distinct) was selected for quantitation. The scanned gray-scale images were filtered to reduce noise; threshold images were made (for brightfield images, 30% above background; for darkfield images, 70%); and the labeled areas were calculated and compared to the total LGN area in that section. Statistical significance was calculated with a Student's t test (of two populations).
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Epibatidine-treated retinas sustained waves either after incubation in fura-2AM (n = 9 retinas 18 to 48 hours after injection) or after additional washout in ACSF (n = 1 retina 48 hours after a single injection; n = 2 retinas after a full series of injections from P0 to P9). Recovery of the retinal waves after fura-2AM loading of the retina was expected if the retina was healthy, because the vitreous humor (containing the epibatidine-treated beads) had been removed for imaging and the fura-2AM incubation volume was relatively large (2 ml).
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In several experiments (n = 3 epibatidine-treated retinas; n = 3 saline-treated retinas), the eyes that received intraocular injections were removed after perfusion and stored in 4% paraformaldehyde in 0.1 M sodium phosphate buffer (PFA in PBS). The eyes were opened, the lens and vitreous humor were removed, tension-relieving slits were made, and the retina was flattened between two glass slides and placed in PFA. The retina was examined under a stereomicroscope for any visible damage and was stained with 0.01% bisbenzimide (Hoechst) in 0.1 M PBS to reveal the overall density of cells in the ganglion cell layer.
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Immediately after the removal of the eyes for imaging, the brain was removed from the skull with the optic nerves attached (n = 2 brains; in each animal, one eye was epibatidine-treated and the other eye was saline-treated). The brain was placed in oxygenated chilled ACSF (12) and sectioned coronally on a vibratome to expose the optic tract. The preparation was perfused with oxygenated ACSF in a recording chamber. A glass electrode filled with ACSF was placed in the optic tract, and the nerves were stimulated with suction electrodes. Recordings were obtained with an extracellular amplifier (Walsh Electronics, Pasadena, CA) and digitized (Axon Instruments).
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3H]leucine) were the same as those used for the eye injections. The location of the injection was confirmed by the presence of the fluorescent latex microspheres located throughout the lateral ventricle (Fig. 4D, top panel).
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We thank G. Grant (Washington University, St. Louis) for providing recombinant κ-bungarotoxin, B. Olivera and J. M. McIntosh (University of Utah, Salt Lake City) for conotoxins, R. Loring (Northeastern University, Boston) for nereistoxin, R Corriveau for advice on acetylcholine receptors, and H. Aaron and C. Cowdry for technical assistance. Supported by NIH grant MH 98108 to C.J.S., NIH MSTP Training Grant to A.A.P., and U. C. Berkeley Miller Fellowship to M.B.F. A.A.P. was in the Neurosciences Program at Stanford University. C.J.S. is an investigator of the Howard Huges Medical Institute.
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