Discrimination between Inhibitory and Excitatory Neurons of Mouse Hippocampus Based on the Shape of Extracellular Spike Waveforms
Abstract
Purpose: Inhibitory and excitatory neurons play an essential role in brain function, and we aim to introduce an automatic method to discriminate these two populations based on features of the shape of their spikes. Consequently, we will explain the spike extraction from raw data of a single shank electrode and determine the best features of spike waveforms for the classification of neurons. It is noteworthy that, to the best of our knowledge, classification of inhibitory and excitatory neurons using the shape features extracted from their spike waveforms has not been done before.
Materials and Methods: In this paper, we use a dataset of mouse hippocampus neurons in which the neuron types (inhibitory or excitatory) have been verified optogenetically. For the classification of mouse hippocampus neurons, we extracted eight shape features of their spike waveforms in addition to their firing rates and used three types of classifiers: K-Nearest Neighbors (KNN), Linear Discriminant Analysis (LDA), and Support Vector Machine (SVM) to analyze the discriminatory power of features based on the accuracy of the classifications.
Results: We showed that Spike asymmetry, Peak-to-trough ratio, Recovery slope, and Duration between peaks were four shape features of spike waveforms participated in the optimum feature subsets that resulted in maximum classification accuracy. Moreover, the SVM classifier with RBF kernel resulted in maximum accuracy of %96.91 ± %13.03 and was identified as the best classifier.
Conclusion: In this study, we found that shape features of spike waveforms can accurately classify inhibitory and excitatory neurons of mouse hippocampus. Also, we found an optimum subset of shape features of spike waveforms that resulted in better classification performance than previously proposed subsets of features used for clustering of neurons. Our findings open a promising way toward a functional classification of neurons automatically.
2-Petilla Interneuron Nomenclature Group (PING), "Petilla terminology: nomenclature of features of GABAergic interneurons of the cerebral cortex," Nature reviews. Neuroscience, vol. 9, no. 7, pp. 557–568, 2008.
3- D. F. English, S. Mckenzie, T. Evans, K. Kim, E. Yoon, and G. Buzsáki, "Pyramidal cell-interneuron circuit architecture and dynamics in hippocampal networks," Neuron, vol. 96, no. 2, pp. 505–520, 2018.
4- M. Pachitariu, Marius, Nicholas Steinmetz, Shabnam Kadir, and Matteo Carandin, N. Steinmetz, S. Kadir, M. Carandini, and K. D. Harris, "Kilosort: realtime spike-sorting for extracellular electrophysiology with hundreds of channels," BioRxiv , p. 061481, 2016.
5- Nowak, Lionel G., Rony Azouz, Maria V. Sanchez-Vives, Charles M. Gray, and David A. McCormick, "Electrophysiological Classes of Cat Primary Visual Cortical Neurons In Vivo as Revealed by Quantitative Analyses," ournal of neurophysiology, vol. 89, no. 3, pp. 1541–1566, 2003.
6- M. Avermann, C. Tomm, C. Mateo, W. Gerstner, and C. C. H. Petersen, "Microcircuits of excitatory and inhibitory neurons in layer 2 / 3 of mouse barrel cortex," J. Neurophisiology, vol. 107, pp. 3116–3134, 2012.
7- A. Sirota, S. Montgomery, S. Fujisawa, Y. Isomura, and G. Buzsáki, "Entrainment of neocortical neurons and gamma oscillations by the hippocampal theta rhythm," Neuron, vol. 60, no. 4, pp. 683–697, 2009.
8- Jia, Xiaoxuan, Joshua H. Siegle, Corbett Bennett, Samuel D. Gale, Daniel J. Denman, Christof Koch, and Shawn R. Olse, "High-density extracellular probes reveal dendritic backpropagation and facilitate neuron classification," journal of neurophysiology, vol. 121, no. 5, pp. 1831-1847, 2019.
9- H. Cheng, D. J. Garrick, and R. L. Fernando, "Efficient strategies for leave-one-out cross validation for genomic best linear unbiased prediction," Anim. Sci. Biotechnol., vol. 8, no. 38, pp. 1-5, 2017.
10- K. Fukunaga, in Introduction to statistical pattern recognition, San Diego: Academic Press Professional, pp. 132–153.
11- R. Duda and P. Hart, "Pattern Classification," New York: Wiley-Interscience, 2000, pp. 174–188.
12- S. Theodoridis and K. Koutroumbas, in Pattern Recognition, California: Academic Press, 2009.
13- Becchetti, Andrea, Francesca Gullo, Giuseppe Bruno, Elena Dossi, Marzia Lecchi, and Enzo Wanke, "Exact distinction of excitatory and inhibitory neurons in neural networks: a study with GFP-GAD67 neurons optically and electrophysiologically recognized on multielectrode arrays," Frontiers in neural circuits, vol. 6, p. 63, 2012.
14- D. A. Mccormick, B. W. Connors, J. W. Lighthall, and D. A. Prince, "Comparative Electrophysiology of Pyramidal and Sparsely Spiny Stellate Neurons of the Neocortex," J. neurophisiology, vol. 54, no. 4, October 1985.
15- J. A. Cardin, L. A. Palmer, and D. Contreras, "Stimulus Feature Selectivity in Excitatory and Inhibitory Neurons in Primary Visual Cortex," J. Neurosci, vol. 27, no. 39, pp. 10333–10344, 2007.
16- Henze, Darrell A., Zsolt Borhegyi, Jozsef Csicsvari, Akira Mamiya, Kenneth D. Harris, and Gyorgy Buzsaki, "ntracellular Features Predicted by Extracellular Recordings in the Hippocampus In Vivo," Journal of neurophysiology, vol. 84, no. 1, pp. 390-400, 2000.
17- S. Y. Shin and M. A. Sommer, "Activity of Neurons in Monkey Globus Pallidus During Oculomotor Behavior Compared With That in Substantia Nigra Pars Reticulata," Neurophysiology, vol. 103, no. 4, pp. 1874–1887, 2010.
18- J. F. Mitchell, K. A. Sundberg, and J. H. Reynolds, "Differential Attention-Dependent Response Modulation across Cell Classes in Macaque Visual Area V4," Neuron, vol. 55, no. 1, pp. 131–141, 2007.
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Issue | Vol 8 No 3 (2021) | |
Section | Original Article(s) | |
DOI | https://doi.org/10.18502/fbt.v8i3.7110 | |
Keywords | ||
Inhibitory and Excitatory Neurons Classification Mouse Hippocampus Shape Features Spike Waveform Extracellular Recordings |
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