Vision and Cortical Map Development

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Vision and Cortical Map Development Leonard E. White, David Fitzpatrick  Neuron  Volume 56, Issue 2, Pages 327-338 (October 2007) DOI: 10.1016/j.neuron.2007.10.011 Copyright © 2007 Elsevier Inc. Terms and Conditions

Figure 1 Summary of Changes that Ensue following Eye Opening in Ferret Visual Cortex and Lateral Geniculate Nucleus Black curve represents average volumetric synaptic density in cortical layer 2/3 (from Erisir and Harris, 2003). This marked increase in synaptogenesis in supragranular layers is accompanied by an outgrowth of long-range horizontal connections, which accelerates after eye opening. The axonal reconstructions above were traced from tangential sections containing small clusters of neurons (located in vacant space) and their supragranular axons labeled with biocytin (White et al., 2001; see also Durack and Katz, 1996; Ruthazer and Stryker, 1996); arrows indicate age of animals (scale bar = 1 mm). The maturation of NMDA-receptor currents in visual cortex follows a similar developmental trajectory, as exemplified by the increasing expression of the NR2A subunit (green line) (after Erisir and Harris, 2003; see also Roberts and Ramoa, 1999). Likewise, the expression of parvalbumin immunoreactivity (solid blue line) and calbindin immunoreactive (dashed blue line) in cortical inhibitory neurons also increases (after Gao et al., 2000). Finally, the response latency of neurons in the lateral geniculate nucleus decreases (orange line), suggesting a coordination of maturation in the thalamus and cortex (after Tavazoie and Reid, 2000). Curves are representations of functions fit to published values normalized to each parameter's peak value in this phase of cortical maturation. Neuron 2007 56, 327-338DOI: (10.1016/j.neuron.2007.10.011) Copyright © 2007 Elsevier Inc. Terms and Conditions

Figure 2 Development of Orientation and Direction Maps in Ferret Visual Cortex Assessed by Means of Intrinsic Signal Optical Imaging (after Li et al., 2006) Graph plots normalized orientation (blue squares) and direction (orange circles) selectivity indices obtained from the same animals (dashed vertical lines); representative difference images for four animals are shown above (arrows indicate age; scale bar = 1 mm). Note the maturation of orientation maps after eye opening and the relative time-lag in the development of direction maps, which become detectable after about 1 week of visual experience. Neuron 2007 56, 327-338DOI: (10.1016/j.neuron.2007.10.011) Copyright © 2007 Elsevier Inc. Terms and Conditions

Figure 3 Representative Maps of Orientation Preference from Age-Matched, Juvenile Normal, Dark-Reared, and Binocularly Lid-Sutured Ferrets (after White et al., 2001) The upper two rows show single-condition images (stimulated condition minus blank image) evoked by horizontal (first row) and vertical (second row) grating stimuli; the third row shows difference images generated by subtracting horizontal and vertical single-condition images (scale bar = 1 mm). The bottom row shows polar-magnitude maps generated by vector summation, where color represents orientation preference (legend below middle column) and brightness represents the magnitude of selectivity. Note the presence of well-formed but attenuated orientation maps in dark-reared ferrets (effects quantified in Figure 4). In contrast, very little columnar structure was evident in lid-sutured ferrets, making maps of orientation preference nearly undetectable in this group. Thus, the impact of abnormal visual experience engendered by lid-suture was far more devastating for the development of the map of orientation preference than was the absence of visual experience. Neuron 2007 56, 327-338DOI: (10.1016/j.neuron.2007.10.011) Copyright © 2007 Elsevier Inc. Terms and Conditions

Figure 4 Early Vision Is Necessary for the Development of Direction Selectivity in Ferret Visual Cortex (after Li et al., 2006) Upper panels: difference images of orientation selectivity (upper row) generated by subtracting horizontal and vertical single-condition images and difference images of direction selectivity (bottom row) for the same animals generated by subtracting responses to upward- and downward-moving grating stimuli (scale bar = 1 mm). The attenuation of orientation selectivity induced by dark-rearing can be normalized by provision of 2–3 weeks of normal visual experience beginning a few days after eye opening (early vision) or about 3 weeks after eye opening (late vision). In contrast, the formation of the map of direction preference is uniquely dependent on early vision. Unlike orientation columns, direction columns are only detected in ferrets that were provided with vision in the first 2–3 weeks after eye opening (normal and early-vision groups). Lower graph: quantification of orientation and direction selectivity indices from the four groups of experimental animals (blue squares, orientation selectivity index; orange circles, direction selectivity index; horizontal lines, median values). Neuron 2007 56, 327-338DOI: (10.1016/j.neuron.2007.10.011) Copyright © 2007 Elsevier Inc. Terms and Conditions