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UC San Diego UC San Diego Electronic Theses and Dissertations UC San Diego UC San Diego Electronic Theses and Dissertations Title Sensory Tuning of Thalamic and Intracortical Excitation in Primary Visual Cortex and Novel Methods for Circuit Analysis Permalink https://escholarship.org/uc/item/0p0306wv Author Lien, Anthony D. Publication Date 2013 Peer reviewed|Thesis/dissertation eScholarship.org Powered by the California Digital Library University of California UNIVERSITY OF CALIFORNIA, SAN DIEGO Sensory Tuning of Thalamic and Intracortical Excitation in Primary Visual Cortex and Novel Methods for Circuit Analysis A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Neurosciences by Anthony D. Lien Committee in charge: Professor Massimo Scanziani, Chair Professor Edward M. Callaway Professor E.J. Chichilnisky Professor Timothy Q. Gentner Professor Jeffry S. Isaacson Professor Takaki Komiyama 2013 Copyright Anthony D. Lien, 2013 All rights reserved. The Dissertation of Anthony D. Lien is approved and it is acceptable in quality and form for publication on microfilm and electronically: Chair University of California, San Diego 2013 iii Table of Contents Signature Page .............................................................................................iii Table of Contents......................................................................................... iv List of Figures .............................................................................................. vi Acknowledgements.....................................................................................vii Vita................................................................................................................ ix Abstract ......................................................................................................... x Chapter 1. Thalamic and intracortical excitation and sensory tuning in the visual cortex ............................................................................................ 1 Introduction..................................................................................................................... 1 Tuning properties of V1 neurons.................................................................................... 2 Anatomical and synaptic properties of thalamic excitation............................................ 4 Tuning properties of thalamic neurons ........................................................................... 6 Circuit mechanisms for orientation tuned thalamic excitation ....................................... 7 Experimental evidence for orientation tuning of thalamic excitation............................. 8 Silencing the cortex to isolate thalamic excitation ....................................................... 11 Results on thalamic excitation ...................................................................................... 13 What are the tuning properties of cortical inputs?........................................................ 15 Results on cortical excitation........................................................................................ 18 Conclusion .................................................................................................................... 20 The origin of direction selectivity............................................................................. 20 Contribution of thalamic and intracortical excitation to tuning properties in other cortical layers............................................................................................................ 23 References..................................................................................................................... 33 Chapter 2. Tuned thalamic excitation is amplified by visual cortical circuits ......................................................................................................... 38 Abstract......................................................................................................................... 38 Introduction................................................................................................................... 38 Results........................................................................................................................... 41 Discussion..................................................................................................................... 49 Methods ........................................................................................................................ 56 Acknowledgements....................................................................................................... 84 References..................................................................................................................... 84 Chapter 3. In vivo labeling of constellations of functionally identified neurons for targeted in vitro recordings................................................... 88 iv Preface .......................................................................................................................... 88 Abstract......................................................................................................................... 89 Introduction................................................................................................................... 90 Results........................................................................................................................... 91 Discussion..................................................................................................................... 96 Methods ........................................................................................................................ 98 Acknowledgments ...................................................................................................... 107 References................................................................................................................... 107 v List of Figures Chapter 1 Figure 1.1 Tuning properties of cortical neurons.................................................. 26 Figure 1.2 Tuning properties of thalamic neurons................................................ 27 Figure 1.3 Possible circuits for orientation selective thalamic excitation ............ 28 Figure 1.4 Experimental evidence for oriented spatial arrangement of thalamic inputs onto a cortical neuron................................................................................. 29 Figure 1.5 Experimental evidence for tuning of thalamic excitation.................... 30 Figure 1.6 Predicted orientation tuning of thalamic excitation in response to drifting gratings..................................................................................................... 31 Figure 1.7 Proposed intracortical and thalamic excitatory connectivity scheme.. 32 Chapter 2 Figure 2.1 Isolating thalamic excitation................................................................ 67 Figure 2.2 Receptive field structure of thalamic excitation.................................. 69 Figure 2.3 Orientation tuning of thalamic excitation............................................ 71 Figure 2.4 Separation of ON and OFF thalamic subfields predicts preferred orientation of thalamic excitation ......................................................................... 72 Figure 2.5 Tuning of non-thalamic excitatory charge .......................................... 74 Figure 2.6 Tuning of non-thalamic excitatory F1 modulation.............................. 75 Figure 2.7 Co-tuning and phase relationship between thalamic and non-thalamic excitation............................................................................................................... 76 Supplementary Figure 2.1 Suppression of the local field potential is correlated with the degree of cortical silencing ..................................................................... 77 Supplementary Figure 2.2 Cortical depth of recorded neurons ............................ 78 Supplementary Figure 2.3 Tuning of isolated dLGN units................................... 79 Supplementary Figure 2.4 Blocking GABAB receptors does not affect grating- evoked local field potential during cortical silencing........................................... 81 Supplementary Figure 2.5 Additional measures of orientation tuning................. 82 Chapter 3 Figure 3.1 In vivo photo-labeling of functionally identified cortical neurons followed by in vitro targeted recording. ............................................................. 105 Figure 3.2 Matching in vivo and in vitro identities of photo-labeled neurons to correlate visual response properties with intrinsic electrophysiological properties. ............................................................................................................................. 106 vi Acknowledgements I thoroughly enjoyed my time in graduate school. This would not be possible without the kind support of my mentors, friends, and family. My most sincere gratitude goes to: Massimo Scanziani for his elegant approach to science, for sharing his enthusiasm and joy of discovery, for his outstanding guidance and mentorship, for his willingness to step with me into the unknown, for his refined aesthetic sense, and for his humor and wit. Shaul Hestrin for patiently teaching me when I knew nothing and inspiring me to pursue science. Bassam Atallah for his infectious passion for science and life and for
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