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ABSTRACT Neuronal dendrites are electrically excitable: they can generate regenerative events such as dendritic spikes in response to sufficiently strong synaptic input1,2,3. Although such

events have been observed in many neuronal types4,5,6,7,8,9, it is not well understood how active dendrites contribute to the tuning of neuronal output _in vivo_. Here we show that dendritic

spikes increase the selectivity of neuronal responses to the orientation of a visual stimulus (orientation tuning). We performed direct patch-clamp recordings from the dendrites of

pyramidal neurons in the primary visual cortex of lightly anaesthetized and awake mice, during sensory processing. Visual stimulation triggered regenerative local dendritic spikes that were

distinct from back-propagating action potentials. These events were orientation tuned and were suppressed by either hyperpolarization of membrane potential or intracellular blockade of NMDA

(_N_-methyl-d-aspartate) receptors. Both of these manipulations also decreased the selectivity of subthreshold orientation tuning measured at the soma, thus linking dendritic regenerative

events to somatic orientation tuning. Together, our results suggest that dendritic spikes that are triggered by visual input contribute to a fundamental cortical computation: enhancing

orientation selectivity in the visual cortex. Thus, dendritic excitability is an essential component of behaviourally relevant computations in neurons. Access through your institution Buy or

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EMERGENCE OF LOCAL AND GLOBAL SYNAPTIC ORGANIZATION ON CORTICAL DENDRITES Article Open access 28 June 2021 FUNCTIONAL ARCHITECTURE OF INTRACELLULAR OSCILLATIONS IN HIPPOCAMPAL DENDRITES

Article Open access 26 July 2024 DENDRITIC EXCITATIONS GOVERN BACK-PROPAGATION VIA A SPIKE-RATE ACCELEROMETER Article Open access 04 February 2025 CHANGE HISTORY * _ 06 NOVEMBER 2012

Reference 13 has been replaced. _ REFERENCES * Johnston, D. & Narayanan, R. Active dendrites: colorful wings of the mysterious butterflies. _Trends Neurosci._ 31, 309–316 (2008) Article

  CAS  Google Scholar  * London, M. & Häusser, M. Dendritic computation. _Annu. Rev. Neurosci._ 28, 503–532 (2005) Article  CAS  Google Scholar  * Spruston, N. Pyramidal neurons:

dendritic structure and synaptic integration. _Nature Rev. Neurosci._ 9, 206–221 (2008) Article  CAS  Google Scholar  * Larkum, M. E., Zhu, J. J. & Sakmann, B. A new cellular mechanism

for coupling inputs arriving at different cortical layers. _Nature_ 398, 338–341 (1999) Article  ADS  CAS  Google Scholar  * Schiller, J., Major, G., Koester, H. J. & Schiller, Y. NMDA

spikes in basal dendrites of cortical pyramidal neurons. _Nature_ 404, 285–289 (2000) Article  ADS  CAS  Google Scholar  * Helmchen, F., Svoboda, K., Denk, W. & Tank, D. W. _In vivo_

dendritic calcium dynamics in deep-layer cortical pyramidal neurons. _Nature Neurosci._ 2, 989–996 (1999) Article  CAS  Google Scholar  * Llinas, R., Nicholson, C., Freeman, J. A. &

Hillman, D. E. Dendritic spikes and their inhibition in alligator Purkinje cells. _Science_ 160, 1132–1135 (1968) Article  ADS  CAS  Google Scholar  * Kamondi, A., Acsady, L. & Buzsaki,

G. Dendritic spikes are enhanced by cooperative network activity in the intact hippocampus. _J. Neurosci._ 18, 3919–3928 (1998) Article  CAS  Google Scholar  * Yuste, R., Gutnick, M. J.,

Saar, D., Delaney, K. R. & Tank, D. W. Ca2+ accumulations in dendrites of neocortical pyramidal neurons: an apical band and evidence for two functional compartments. _Neuron_ 13, 23–43

(1994) Article  CAS  Google Scholar  * Branco, T., Clark, B. A. & Häusser, M. Dendritic discrimination of temporal input sequences in cortical neurons. _Science_ 329, 1671–1675 (2010)

Article  ADS  CAS  Google Scholar  * Ferster, D. & Jagadeesh, B. EPSP-IPSP interactions in cat visual cortex studied with _in vivo_ whole-cell patch recording. _J. Neurosci._ 12,

1262–1274 (1992) Article  CAS  Google Scholar  * Volgushev, M., Pei, X., Vidyasagar, T. R. & Creutzfeldt, O. D. Postsynaptic potentials in cat visual cortex: dependence on polarization.

_Neuroreport_ 3, 679–682 (1992) Article  CAS  Google Scholar  * Hirsch, J. A., Alonso, J. M. & Reid, R. A. Visually evoked calcium action potentials in cat striate cortex. _Nature_ 378,

612–616 (1995) Article  ADS  CAS  Google Scholar  * Hubel, D. H. & Wiesel, T. N. Receptive fields of single neurones in the cat’s striate cortex. _J. Physiol. (Lond.)_ 148, 574–591

(1959) Article  CAS  Google Scholar  * Larkum, M. E., Waters, J., Sakmann, B. & Helmchen, F. Dendritic spikes in apical dendrites of neocortical layer 2/3 pyramidal neurons. _J.

Neurosci._ 27, 8999–9008 (2007) Article  CAS  Google Scholar  * Waters, J. & Helmchen, F. Background synaptic activity is sparse in neocortex. _J. Neurosci._ 26, 8267–8277 (2006) Article

  CAS  Google Scholar  * Niell, C. M. & Stryker, M. P. Highly selective receptive fields in mouse visual cortex. _J. Neurosci._ 28, 7520–7536 (2008) Article  CAS  Google Scholar  * Yu,

Y., Shu, Y. & McCormick, D. A. Cortical action potential backpropagation explains spike threshold variability and rapid-onset kinetics. _J. Neurosci._ 28, 7260–7272 (2008) Article  CAS 

Google Scholar  * Svoboda, K., Helmchen, F., Denk, W. & Tank, D. W. Spread of dendritic excitation in layer 2/3 pyramidal neurons in rat barrel cortex _in vivo_. _Nature Neurosci._ 2,

65–73 (1999) Article  CAS  Google Scholar  * Tan, A. Y., Brown, B. D., Scholl, B., Mohanty, D. & Priebe, N. J. Orientation selectivity of synaptic input to neurons in mouse and cat

primary visual cortex. _J. Neurosci._ 31, 12339–12350 (2011) Article  CAS  Google Scholar  * Jia, H., Rochefort, N. L., Chen, X. & Konnerth, A. Dendritic organization of sensory input to

cortical neurons _in vivo_. _Nature_ 464, 1307–1312 (2010) Article  ADS  CAS  Google Scholar  * Polsky, A., Mel, B. W. & Schiller, J. Computational subunits in thin dendrites of

pyramidal cells. _Nature Neurosci._ 7, 621–627 (2004) Article  CAS  Google Scholar  * Lavzin, M., Rapoport, S., Polsky, A., Garion, L. & Schiller, J. Nonlinear dendritic processing

determines angular tuning of barrel cortex neurons _in vivo_. _Nature_ 490, 397–401 (2012) Article  ADS  CAS  Google Scholar  * Mel, B. W. Synaptic integration in an excitable dendritic

tree. _J. Neurophysiol._ 70, 1086–1101 (1993) Article  CAS  Google Scholar  * Smith, S. L. & Häusser, M. Parallel processing of visual space by neighboring neurons in mouse visual

cortex. _Nature Neurosci._ 13, 1144–1149 (2010) Article  CAS  Google Scholar  * Ohiorhenuan, I. E. et al. Sparse coding and high-order correlations in fine-scale cortical networks. _Nature_

466, 617–621 (2010) Article  ADS  CAS  Google Scholar  * London, M., Roth, A., Beeren, L., Häusser, M. & Latham, P. E. Sensitivity to perturbations _in vivo_ implies high noise and

suggests rate coding in cortex. _Nature_ 466, 123–127 (2010) Article  ADS  CAS  Google Scholar  * Xu, N. L. et al. Nonlinear dendritic integration of sensory and motor input during an active

sensing task. _Nature_ 492, 247–251 (2012) Article  ADS  CAS  Google Scholar  * Gentet, L. J. et al. Unique functional properties of somatostatin-expressing GABAergic neurons in mouse

barrel cortex. _Nature Neurosci._ 15, 607–612 (2012) Article  CAS  Google Scholar  * Jiang, X., Wang, G., Lee, A. J., Stornetta, R. L. & Zhu, J. J. The organization of two new cortical

interneuronal circuits. _Nature Neurosci._ 16, 210–218 (2013) Article  CAS  Google Scholar  * Brainard, D. H. The psychophysics toolbox. _Spat. Vis._ 10, 433–436 (1997) Article  CAS  Google

Scholar  * Pelli, D. G. The VideoToolbox software for visual psychophysics: transforming numbers into movies. _Spat. Vis._ 10, 437–442 (1997) Article  CAS  Google Scholar  * Kitamura, K.,

Judkewitz, B., Kano, M., Denk, W. & Häusser, M. Targeted patch-clamp recordings and single-cell electroporation of unlabeled neurons _in vivo_. _Nature Methods_ 5, 61–67 (2008) Article 

CAS  Google Scholar  * Jahr, C. E. & Stevens, C. F. Voltage dependence of NMDA-activated macroscopic conductances predicted by single-channel kinetics. _J. Neurosci._ 10, 3178–3182

(1990) Article  CAS  Google Scholar  * Nevian, T., Larkum, M. E., Polsky, A. & Schiller, J. Properties of basal dendrites of layer 5 pyramidal neurons: a direct patch-clamp recording

study. _Nature Neurosci._ 10, 206–214 (2007) Article  CAS  Google Scholar  * Spruston, N., Jonas, P. & Sakmann, B. Dendritic glutamate receptor channels in rat hippocampal CA3 and CA1

pyramidal neurons. _J. Physiol. (Lond.)_ 482, 325–352 (1995) Article  CAS  Google Scholar  * Pologruto, T. A., Sabatini, B. L. & Svoboda, K. ScanImage: flexible software for operating

laser scanning microscopes. _Biomed. Eng. Online_ 2, 13 (2003) Article  Google Scholar  * Fasano, G. & Franceschini, A. A multidimensional version of the Kolomogorov–Smirnov test. _Mon.

Not. R. Astron. Soc._ 225, 155–170 (1987) Article  ADS  Google Scholar  Download references ACKNOWLEDGEMENTS We are grateful to B. Clark, P. Latham, M. London, D. Ringach, A. Roth, C.

Schmidt-Hieber and C. Wilms for discussions and comments on the manuscript. This work was supported by the following: a Long-Term Fellowship and a Career Development Award from the Human

Frontier Science Program and a Klingenstein Fellowship (S.L.S.); a Helen Lyng White Fellowship (I.T.S.); a Wellcome Trust and Royal Society Fellowship and MRC Programme Leader Track (T.B.);

and by grants from the Wellcome Trust, ERC and Gatsby Charitable Foundation (M.H.). AUTHOR INFORMATION AUTHORS AND AFFILIATIONS * Wolfson Institute for Biomedical Research and Department of

Neuroscience, Physiology and Pharmacology, University College London, Gower Street, London WC1E 6BT, UK, Spencer L. Smith, Ikuko T. Smith, Tiago Branco & Michael Häusser * Department of

Cell Biology and Physiology and Neuroscience Center, University of North Carolina School of Medicine, Chapel Hill, 27599, North Carolina, USA Spencer L. Smith & Ikuko T. Smith *

Laboratory of Molecular Biology, Medical Research Council, Cambridge CB2 0QH, UK, Tiago Branco Authors * Spencer L. Smith View author publications You can also search for this author

inPubMed Google Scholar * Ikuko T. Smith View author publications You can also search for this author inPubMed Google Scholar * Tiago Branco View author publications You can also search for

this author inPubMed Google Scholar * Michael Häusser View author publications You can also search for this author inPubMed Google Scholar CONTRIBUTIONS S.L.S. and M.H. conceived and

designed the experiments. S.L.S. and I.T.S. performed the experiments. S.L.S. analysed the data. T.B. designed and carried out the compartmental modelling. S.L.S., I.T.S., T.B. and M.H.

interpreted the data and wrote the paper. CORRESPONDING AUTHORS Correspondence to Spencer L. Smith or Michael Häusser. ETHICS DECLARATIONS COMPETING INTERESTS The authors declare no

competing financial interests. EXTENDED DATA FIGURES AND TABLES EXTENDED DATA FIGURE 1 ELECTROPHYSIOLOGICAL FEATURES OF LAYER 2/3 DENDRITES _IN VIVO_. A, The input resistance of distal

dendrites was typically 100–300 MΩ but sometimes larger (up to 600 MΩ). The input resistance increased as function of dendritic distance from the soma, approximately doubling every 300 µm.

The grey point indicates the input resistance measured in somatic patch-clamp recordings (mean ± s.e.m.). B, During a dendritic recording at 150 µm from the soma, hyperpolarizing current

steps did not reveal a voltage sag; thus, there is probably little to no hyperpolarization-activated cation current, _I_h, in the dendrites of layer 2/3 pyramidal neurons _in vivo_. C, The

peak voltage response plotted against the hyperpolarizing current step amplitude in an _I_–_V_ plot was well fit by a linear function, confirming the lack of _I_h. D, Representative

dendritic bursts evoked by visual stimulation at the optimal orientation in nine different dendritic recordings at progressively increasing distances from the soma. All right-hand scale bars

are 20 mV. E, F, Compared with action potentials recorded at the soma, bAPs had a lower amplitude (E) and were prolonged in time (F), and both of these trends were more pronounced with

increasing dendritic distance from the soma (error bars, s.d.). Both the amplitude and width were significantly different among the three groups (_P_ < 0.01, unpaired _t_-tests with the

Bonferroni correction for multiple comparisons). EXTENDED DATA FIGURE 2 ORIENTATION TUNING CURVES OF DENDRITIC BURSTS COMPARED WITH BAPS. Tuning curves for dendritic spike bursts and bAPs

recorded at distal dendritic locations (>75 µm from soma) are shown. The tuning curves for dendritic bursts match the tuning curves for isolated bAPs. The statistical significance of

dendritic burst tuning curves was tested by randomly shuffling responses (details in the Methods) and was found to be significant (_P_ < 0.05) for 7 out of 9 cells (dendritic burst tuning

in cells 6 and 9 was not significant). The curves were normalized to the maximal values, which are shown at the bottom right of each polar plot. The small qualitative differences may be due

to dendrites that are topologically distant from the dendritic recording site exhibiting slightly different tuning curves. The grating drift direction that elicited the largest response is

indicated with an arrow. The difference between these directions is indicated at the bottom of each polar plot. The cross correlation between dendritic bursts and isolated bAPs was highly

significant: Pearson’s _R_ = 0.54, _P_ = 0.000013, paired _t_-test; _n_ = 9. When only the spikes in the bursts with rise times in the slowest quartile of the distribution were considered to

be dendritic in origin, the preferred orientation of bAPs and the slowest quartile were still matched within individual dendritic recordings (difference in preferred orientation, 41.5 ± 

58.1°; _P_ = 0.49, paired _t_-test; _n_ = 9). EXTENDED DATA FIGURE 3 DENDRITIC RECORDINGS IN AWAKE MICE EXHIBIT DENDRITIC BURSTS. A, Awake, head-fixed mice viewed drifting gratings during

electrophysiological recordings. B, A two-photon image of the patched dendrite (117 µm from the soma) of a layer 2/3 pyramidal neuron in the mouse visual cortex, after filling with Alexa

Fluor 594 with the dendritic patch-clamp pipette. C, Dendritic bursts were observed when the preferred orientation was presented. D, Tuning curves for the isolated bAPs and dendritic bursts.

E, Example bursts from three different distal dendritic recordings in awake mice. Calibration bars, 25 mV. EXTENDED DATA FIGURE 4 THE DIVERSITY OF ONSET DYNAMICS VERSUS MEMBRANE POTENTIAL.

A, Spikes from each distal dendritic recording (both isolated bAPs (black) and spikes in dendritic burst events (red)) were normalized such that isolated bAPs had a mean phase slope of 1.

The mean baseline membrane potential (_V_m) of isolated bAPs was subtracted from the mean baseline _V_m of all spikes. Although many spikes in bursts had a depolarized baseline _V_m relative

to isolated spikes, there was overlap between the two populations around ±3 mV. B, Magnification of panel A to show spikes at ±3 mV relative to the mean baseline _V_m of isolated bAPs. C,

Histograms of the two populations reveal a tendency towards lower phase slope values for spikes in bursts (_P_ = 0.041, Kolmogorov–Smirnov test ; _n_ = 211 bAPs, 80 spikes in bursts). D, An

example of bAPs and a spike in a burst (both from the same distal dendritic recording): although the bAP has a more depolarized baseline _V_m, it still exhibits a steeper phase slope (a kink

at the foot of the voltage waveform), indicative of a propagated action potential. EXTENDED DATA FIGURE 5 CA2+ IMAGING AT THE SITE OF DENDRITIC RECORDING REVEALS THAT GLOBAL CA2+ SIGNALS

ARE ASSOCIATED WITH FASTER ONSET SPIKES. A, During dendritic recordings, Ca2+ signals were simultaneously imaged at the site of the recording and at nearby dendrites. B, In dendritic bursts

with global Ca2+ signals that were simultaneously observed in all regions of interest (ROIs), the spikes recorded at the dendrite exhibited steep onsets, indicating that they were probably

bAPs. C, In local Ca2+ signals that were observed only in the ROI at the site of recording, the dendritic spikes exhibited slower onsets, indicating that they were probably locally

generated. D, The maximum phase slope of spikes occurring during global Ca2+ events was higher than for spikes occurring during local Ca2+ events (_P_ = 0.0069, _t_-test). E, F, When global

Ca2+ signals occurred during ongoing local Ca2+ signals, the initiation was associated with a steep onset spike. Two examples are shown. EXTENDED DATA FIGURE 6 NON-FIRING CELLS EXHIBIT

SUBTHRESHOLD ORIENTATION TUNING. A, Raw data for an example cell in which subthreshold orientation tuning was observed, although no spikes were fired during stimulus presentations. B, In

this case, the tuning width of the subthreshold membrane potential was quite sharp and was confined to two directions. EXTENDED DATA FIGURE 7 TUNING OF ACTION POTENTIALS AND SUBTHRESHOLD

MEMBRANE POTENTIAL. A, In individual cells, the orientation tuning of spikes and the membrane potential were highly correlated, indicating that the tuning of the subthreshold responses was

not spurious (mean difference in preferred orientation, 14.8 ± 5.3°). B, In individual cells, the orientation selectivity index based on the membrane potential response (_V_mOSI) was highly

correlated with the conventional spiking-based OSI. C, In individual cells, the preferred orientation of the control subthreshold response was correlated with the preferred orientation of

the subthreshold response during hyperpolarization. D, The black curve is the fitted subthreshold orientation tuning curve (the black circles are raw data points), and the red curve is the

subthreshold tuning curve during hyperpolarization (the red circles are raw data points). The _V_mOSI values for the control and hyperpolarized conditions are shown next to each plot. The

radial axes are linear and start at 0. The maximal radial axis range is shown below each polar plot. The differences in _V_mOSI are quantified in Fig. 5g. EXTENDED DATA FIGURE 8 CHANGES IN

THE DRIVING FORCE FOR CL−DO NOT ACCOUNT FOR THE EFFECTS OF HYPERPOLARIZATION ON _V_MOSI. A, When the pipette solution contained 10 mM Cl−, the reversal potential for chloride, _E_Cl, was

estimated to be −71 mV (based on the assumption that natural cerebrospinal fluid contains a similar amount of Cl− to the artificial cerebrospinal fluid used). In this situation,

hyperpolarization decreased the orientation selectivity index. B, Even with a low Cl− concentration (4 mM; estimated _E_Cl, −95 mV), the result was the same. C, There was no significant

correlation between the driving force for Cl− and _V_mOSI. D, The data were resampled (15 of the data points in C were selected at random, and the _R_ and _P_ values for that set of data

points were calculated; this process was repeated 10,000 times), and this process confirmed that the result from the correlational analysis in C was not biased by a small subset of the data

points (the mean _R_ and _P_ values from the resampling analysis match the values for the full data set in C well). EXTENDED DATA FIGURE 9 THE EFFECT OF INTRACELLULAR MK-801 ON UP AND DOWN

STATES AND DENDRITIC SPIKES. A, To determine whether MK-801 that might have leaked out of the pipette during patching affects network circuitry, we examined the dynamics of up and down

states in recordings in which MK-801 was in the pipette and in control recordings (no MK-801). B, Although, in general, the membrane potential drifted up slightly (<5 mV on average) and

the time spent in the up state increased over long recordings (possibly due to the anaesthesia wearing off), these trends were identical with or without MK-801 in the patch pipette. C, When

1 µM MK-801 was included in the recording pipette, the visually evoked responses contained fewer bAPs and bursts. This trend was clear in individual cells (A) and across the population (D).

This reduction in spiking confirms that dendritic bursts do not occur when NMDA receptors are blocked. Because the low firing rate in MK-801 recordings prevented a reliable measurement of

orientation tuning in the MK-801 dendritic patch-clamp recordings, we averaged over all of the stimulus presentations for both conditions, resulting in a lower average firing rate for bAPs

and dendritic bursts. EXTENDED DATA FIGURE 10 COMPARTMENTAL MODELLING OF DENDRITIC EVENTS. A, A detailed reconstruction of a layer 2/3 pyramidal cell was used in the simulations. Light green

circles over the dendritic tree represent background synapses, and dark green circles represent signal synapses (the model had 1,100 synapses; not all are illustrated). Voltage was recorded

at the soma and at all dendritic branches simultaneously. B, Activation of signal synapses at 5 Hz produced high-frequency dendritic bursts, composed of local dendritic spikes and bAPs.

These bursts were always accompanied by dendritic Ca2+ transients. The timing of the activation of excitatory synapses on the recorded dendritic branch is illustrated. Note how the local

excitatory postsynaptic potentials (EPSPs) are clearly smaller than the dendritic spikes. C, Examples of specific features consistently observed in the model. Isolated bAPs were associated

with global Ca2+ transients and had kinked onsets. Dendritic spikes often preceded somatic action potentials, had smooth onsets and Ca2+ transients that were localized to the branches where

the spikes were recorded, and clearly started before the global transients associated with bAPs. Local dendritic spikes initiated in the dendrite could often be recorded in multiple

electrotonically close dendritic branches. Pairs of local spikes and bAPs reached very high frequencies; the example shows a pair at >400 Hz. When NMDA receptors were removed from the

simulations, no dendritic spikes were observed, and the soma failed to reach the threshold for action potential firing. This also occurred when there were no dendritic voltage-activated Na+

channels, indicating that the generation of dendritic spikes is required for producing axonal output. D, Quantification of spike onset for local dendritic spikes and bAPs in the model

reproduced the experimentally observed effect reported in Extended Data Fig. 5d. E, Example trial showing the somatic voltage and recordings for two dendrites indicated in A. For each

dendrite, the local voltage, the Na+ channel conductance (gNa, expressed as a fraction of the maximum conductance) and the timing of activation of excitatory synapses on the recorded

dendrite are shown. The gNa traces show that there is significant local Na+ channel inactivation after the first spike and that subsequent spikes are associated with varying degrees of Na+

channel conductance. Asterisks denote extreme cases when a bAP followed a local dendritic spike at very high frequency and did not recruit any local gNa, thereby indicating that the

propagation into the recorded branch was passive. SUPPLEMENTARY INFORMATION SUPPLEMENTARY INFORMATION This file contains a Supplementary Note and additional references. _This file was

replaced on 6 November 2013._ (PDF 206 kb) THE PROPAGATION OF VOLTAGE IN THE COMPARTMENTAL MODEL DURING SYNAPTIC ACTIVATION Background input was continuously active, and signal synapses were

activated at 8 Hz for 200 ms (100 ms after the start of the trace). The traces show the local voltage at the indicated dendrites, together with the somatic voltage. During the period of

high synaptic input local dendritic spikes are elicited in multiple regions of the dendritic tree, and eventually lead to a somatic action potential the backpropagates globally. Note how all

of the global spikes are preceded by at least one dendritic spike in a region of the dendritic tree. (MOV 5410 kb) SLOW MOTION VIDEO OF THE SAME TRIAL SHOWN IN VIDEO 1, HIGHLIGHTING THE

ACTIVITY IN DENDRITE 3 Note that the first two global spikes are immediately preceded by local spikes in this dendrite, and that while the last dendritic spike does not directly trigger a

somatic action potential it leads to a significant charge build up in the soma that facilitates the subsequent somatic firing. (MOV 3761 kb) SLOW MOTION VIDEO OF THE SAME TRIAL SHOWN IN

VIDEO 1, HIGHLIGHTING THE ACTIVITY IN DENDRITE 1 Note that dendrite 1 fires bursts of spikes at frequencies >50 Hz. (MOV 3793 kb) POWERPOINT SLIDES POWERPOINT SLIDE FOR FIG. 1 POWERPOINT

SLIDE FOR FIG. 2 POWERPOINT SLIDE FOR FIG. 3 POWERPOINT SLIDE FOR FIG. 4 POWERPOINT SLIDE FOR FIG. 5 RIGHTS AND PERMISSIONS Reprints and permissions ABOUT THIS ARTICLE CITE THIS ARTICLE

Smith, S., Smith, I., Branco, T. _et al._ Dendritic spikes enhance stimulus selectivity in cortical neurons _in vivo _. _Nature_ 503, 115–120 (2013). https://doi.org/10.1038/nature12600

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