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Hausmann FS, Barrett JM, Martin ME, Zhan H, Shepherd GMG (2022) Axonal barcode analysis of pyramidal tract projections from mouse forelimb M1 and M2. Journal of Neuroscience 42:7733-7743.

Barrett JM, Martin ME, Shepherd GMG (2022) Manipulation-specific cortical activity as mice handle food. Current Biology 32:4842-4853.


Shepherd GMG & Yamawaki N (2021) Untangling the cortico-thalamo-cortical loop: cellular pieces of a knotty circuit puzzle. Nature Reviews Neuroscience. 22(7):389-406

Yamawaki N, Raineri Tapies MG, Stults A, Smith GA, Shepherd GMG (2021) Circuit organization of the excitatory sensorimotor loop through hand/forelimb S1 and M1. eLife 10:e66836.


Guo K, Yamawaki N, Barrett JM, Tapies M, Shepherd GMG (2020) Cortico-thalamo-cortical circuits of mouse forelimb S1 are organized primarily as recurrent loops. Journal of Neuroscience 40(14):2849-2858.

Barrett JM, Raineri Tapies MG, Shepherd GMG (2020) Manual dexterity of mice during food-handling involves the thumb and a set of fast basic movements. PLoS One,15(1):e0226774.


Yamawaki N, Li X, Lambot L, Ren LY, Radulovic J, Shepherd GMG (2019) Long-range inhibitory intersection of a retrosplenial thalamocortical circuit by apical tuft-targeting CA1 neurons. Nature Neuroscience, 22(4):618-626.

Jiang Y, Parameswaran R, Li X, Carvalho-de-Souza J, Gao X, Meng L, Bezanilla F, Shepherd GMG, Tian B (2019) Nongenetic optical neuromodulation with silicon-based materials. Nature Protocols, 14(5):1339-1376.

Dura-Bernal S, Suter BA, Gleeson P, Cantarelli M, Quintana A, Rodriguez F, Kedziora DJ, Chadderdon GL, Kerr CC, Neymotin SA, McDougal RA, Hines M, Shepherd GM, Lytton WW (2019) NetPyNE, a tool for data-driven multiscale modeling of brain circuits. eLife, 8. pii: e44494.

Guo K, Yamawaki N, Svoboda K, Shepherd GMG (2018) Anterolateral motor cortex connects with a medial subdivision of ventromedial thalamus through cell-type-specific circuits, forming an excitatory thalamo-cortico-thalamic loop via layer 1 apical tuft dendrites of layer 5B pyramidal tract type neurons. Journal of Neuroscience, 38(41):8787-8797.

Li X, Yamawaki N, Barrett JM, Kording K, Shepherd GMG (2018) Scaling of optogenetically evoked signaling in a higher-order corticocortical pathway in the anesthetized mouse. Frontiers in Systems Neuroscience, 12:16.

Yamawaki N, Corcoran KA, Guedea AL, Shepherd GMG, Radulovic J (2018) Differential contributions of glutamatergic hippocampal→retrosplenial cortical projections to the formation and persistence of context memories. Cerebral Cortex, 29(6):2728-2736.

Jiang Y, Li X, Liu B, Yi J, Fang Y, Shi F, Gao X, Sudzilovsky E, Parameswaran R, Koehler K, Nair V, Yue J, Guo K, Fang Y, Tsai H-M, Freyermuth G, Wong RCS, Kao C-M, Chen C-T, Nicholls AW, Wu X, Shepherd GMG, Tian B (2018) Rational design of silicon structures for optically-controlled multiscale biointerfaces. Nature Biomedical Engineering, 2:508–521.

Neymotin SA, Suter BA, Dura-Bernal S, Shepherd GM, Migliore M, Lytton WW (2017) Optimizing computer models of corticospinal neurons to replicate in vitro dynamics. Journal of Neurophysiology, 117(1):148-162.

​Yamawaki N, Radulovic JR, Shepherd GMG (2016) A corticocortical circuit directly links retrosplenial cortex to M2 in the mouse. Journal of Neuroscience, 36(36):9365–9374.

Yamawaki N, Suter BA, Wickersham IR, Shepherd GM (2016) Combining optogenetics and electrophysiology to analyze projection neuron circuits. Cold Spring Harbor Protocols, 2016(10):pdb.prot090084.

Heuermann RJ, Jaramillo TC, Ying S-W, Suter BA, Lyman KA, Han Y, Lewis AS, Hampton TG, Shepherd GMG, Goldstein PA, Chetkovich DM (2016) Reduction of thalamic and cortical Ih by deletion of TRIP8b produces a mouse model of human absence epilepsy. Neurobiology of Disease, 85:81-92.


Yamawaki N, Shepherd GMG (2015) Synaptic circuit organization of motor corticothalamic neurons. Journal of Neuroscience, 35:2293–2307.


Harris KD, Shepherd GMG (2015) The neocortical circuit: themes and variations. Nature Neuroscience, 18:170-181.


Suter BA, Shepherd GMG (2015) Reciprocal inter-areal connections to corticospinal neurons in mouse M1 and S2. Journal of Neuroscience, 35:2959-2974.


Joshi A, Middleton JW, Anderson CT, Borges K, Suter BA, Shepherd GMG, Tzounopoulos T (2015) Cell-specific, activity-dependent fractionation of layer 2/3→5B excitatory signaling in mouse auditory cortex. Journal of Neuroscience, 35:3112-31123.


Yamawaki N, Borges K, Suter BA, Wokosin D, Harris KD, Shepherd GMG (2014) A genuine layer 4 in motor cortex with prototypical synaptic circuit connectivity. eLife, 3:e05422:1-16.


Shepherd GMG (2014) Diversity and complexity in the PT projectome. Nature Reviews Neuroscience, 15:63.


Brecht M, Hatsopoulos NG, Kaneko T, Shepherd GMG (2014) Motor cortex microcircuits. Frontiers in Neural Circuits, 7:196.


Chadderdon GL, Mohan A, Suter BA, Neymotin SA, Kerr CC, Francis JT, Shepherd GMG, Lytton WW (2014) Motor cortex microcircuit simulation based on brain activity mapping. Neural Computation, 26(7):1239-62.


Suter BA, Yamawaki N, Borges K, Li X, Kiritani T, Hooks BM, Shepherd GMG (2014) Neurophotonics applications to motor cortex research. Neurophotonics, 1:011008(1-5).


Hooks BM, Mao T, Gutnisky D, Yamawaki N, Svoboda K, Shepherd GMG (2013) Organization of cortical and thalamic input to pyramidal neurons in mouse motor cortex. Journal of Neuroscience, 33:748-760.


Shepherd GMG (2013) Corticostriatal connectivity and its role in disease. Nature Reviews Neuroscience, 14:278-291.


Yasvoina M, Genc B, Jara J, Sheets PL, Quinlan K, Milosevic A, Shepherd GMG, Heckman CJ, and Ozdinler PH (2013) eGFP expression under UCHL1 promoter genetically labels corticospinal motor neurons and a subpopulation of degeneration resistant spinal motor neurons in an ALS mouse model. Journal of Neuroscience, 33:7890-7904


Suter B, Migliore M, Shepherd GMG (2013) Intrinsic electrophysiology of mouse corticospinal neurons: a class-specific triad of spike-related properties. Cerebral Cortex, 23:1965-1977.


Kress GJ, Yamawaki N, Wokosin DL, Wickersham IR, Shepherd GMG, Surmeier DJ (2013) Convergent cortical innervation of striatal projection neurons. Nature Neuroscience, 16:665-667.


Kiritani T, Wickersham IR, Seung HS, Shepherd GMG (2012) Hierarchical connectivity and connection-specific dynamics in the corticospinal-corticostriatal microcircuit in mouse motor cortex. Journal of Neuroscience, 32(14):4992–5001.


Apicella A, Wickersham IR, Seung HS, Shepherd GMG (2012) Laminarly orthogonal excitation of fast spiking and low threshold spiking interneurons in mouse motor cortex. Journal of Neuroscience, 32:7021–7033.


Srivastava DP, Woolfrey KM, Jones KA, Anderson CT, Russell TA, Lee H, Yasvoina MV, Wokosin DL, Ozdinler PH, Shepherd GMG, Penzes P (2012) Disease-associated coding variant reveals role for a Ras/Epac2-dependent pathway in controlling basal dendrite maintenance. PLoS Biology, 10:e1001350.


Hooks BM, Hires SA, Zhang Y-X, Huber D, Petreanu LT, Svoboda K, Shepherd GMG (2011) Laminar analysis of excitatory local circuits in vibrissal motor and sensory cortical areas. PLoS Biology, 9(1): e1000572.


Sheets PL, Shepherd GMG (2011) Cortical circuits for motor control. Neuropsychopharmacology, 36:365-366.


Qiu S, Anderson CT, Levitt P, Shepherd GMG (2011) Circuit-specific intracortical hyperconnectivity in mice with deletion of the Met receptor tyrosine kinase. Journal of Neuroscience, 31(15):5855-5864.


Middleton JW, Kiritani T, Pedersen C, Turner J, Shepherd GMG, Tzounopoulos T (2011) Mice with behavioral evidence of tinnitus exhibit dorsal cochlear nucleus hyperactivity due to decreased GABAergic inhibition. Proceedings of the National Academy of Sciences USA, 108:7601-7606.


Sheets PL, Suter BA, Kiritani T, Chan CS, Surmeier DJ, Shepherd GMG (2011) Corticospinal-specific HCN expression in mouse motor cortex: Ih-dependent synaptic integration as a candidate microcircuit mechanism involved in motor control. Journal of Neurophysiology, 106:2216-2231.


Shepherd GMG, Katz DM (2011) Synaptic microcircuit dysfunction in genetic models of neurodevelopmental disorders: focus on Mecp2 and Met. Current Opinion in Neurobiology, 21(6):827-33.


Shepherd GMG (2011) Circuit mapping by UV uncaging of glutamate. In: Optical Imaging in Neuroscience: A Laboratory Manual (Helmchen F, Konnerth A, eds), pp 419-427. Cold Spring Harbor: CSHL Press. Also published online in Cold Spring Harbor Protocols, 2012(9):998-1004.


Wood L, Shepherd GMG (2010) Synaptic circuit abnormalities of motor-frontal layer 2/3 pyramidal neurons in a mutant mouse model of Rett syndrome. Neurobiology of Disease, 38:281-287.


Christie LA, Russell TA, Wood L, Xu J, Shepherd GMG, Contractor A (2010) AMPA receptor desensitization mutation results in severe developmental phenotypes and early postnatal lethality. Proceedings of the National Academy of Sciences USA, 107:9412-9417.


Anderson CT, Sheets PL, Kiritani T, Shepherd GMG (2010) Sublayer-specific microcircuits of corticospinal and corticostriatal neurons in motor cortex. Nature Neuroscience, 13:739-744.


Suter BA, O’Connor T, Iyer V, Petreanu LT, Hooks BM, Kiritani T, Svoboda K, Shepherd GMG (2010) Ephus: multipurpose data acquisition software for neuroscience experiments. Frontiers in Neuroscience Methods, 4:1-13.


Svoboda K, Hooks BM, Shepherd GMG (2010) Barrel cortex. In: Handbook of Microcircuits (Shepherd GM, Grillner S, eds), pp 31-38. New York: Oxford University Press.


Wood L, Gray N, Zhou Z, Greenberg ME, Shepherd GMG (2009) Synaptic circuit abnormalities of MeCP2-deficient layer 2/3 pyramidal neurons in mouse motor-frontal cortex. Journal of Neuroscience, 29:12440-12448.


Shepherd GMG (2009) Intracortical cartography in an agranular area. Frontiers in Neuroscience, 3:337-343.


Bureau I, Shepherd GMG, Svoboda K (2008) Circuit and plasticity defects in the developing somatosensory cortex of Fmr1 knock-out mice. Journal of Neuroscience, 28: 5178-5188.


Yu I, Anderson CT, Sheets PL, Kiritani T, Wokosin DL, Wood L, Shepherd GMG (2008) Local-circuit phenotypes of layer 5 neurons in motor-frontal cortex of YFP-H mice. Frontiers in Neural Circuits, 2:1-8.


Weiler N, Wood L, Yu J, Solla SA, Shepherd GMG (2008) Top-down laminar organization of the excitatory network in motor cortex. Nature Neuroscience, 11:360-366.

Links to more of Gordon M. G. Shepherd’s articles on Pubmed and Google Scholar

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Somatosensory-to-motor transcortical circuits

Cortical circuits - review

Cortico-thalamo-cortical loops - review

Long-range inhibition of thalamocortical excitation

Mouse thumbs and manual dexterity

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