Iannetti, GD

Novembre, G

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Novembre,	G.,	&	Iannetti,	G.	D.	(2018).	Tagging	the	musical	beat:	Neural	entrainment	or	event-related	potentials?.	Proceedings	of	the	National	Academy	of	Sciences,	115(47),	E11002-E11003.		Tagging	the	musical	beat:	Neural	Entrainment	or	Event-Related	Potentials?			Giacomo	Novembre1,2,*,	Gian	Domenico	Iannetti1,2,*		1	Department	of	Neuroscience,	Physiology	and	Pharmacology,	University	College	London,	UK	2	Neuroscience	and	behaviour	laboratory,	Istituto	Italiano	di	Tecnologia,	Rome,	Italy	*	Correspondence	to:	g.novembre@ucl.ac.uk;	g.iannetti@ucl.ac.uk			In	 a	 recent	 issue	 of	 PNAS,	 Lenc	 et	 al.	 (1)	 report	 that	 Steady-State	 Responses	(SSRs)	induced	by	rhythmic	auditory	stimuli	are	enhanced	when	listeners	attend	to	 low	 frequency	 sounds,	 particularly	 when	 these	 sounds	 reflect	 salient	periodicities	like	the	beat	or	the	meter.	Lenc	et	al.	(1)	interpret	this	finding	as	an	entrainment	of	sensory-motor	networks	important	for	planning	and	controlling	movements,	 underlying	 humans’	 spontaneous	 tendency	 to	move	 to	music,	 and	particularly	 to	 instruments	 producing	 low-frequency	 sounds,	 such	 as	 bass	 or	drums	(1,	2).	We	 propose	 a	 more	 parsimonious	 interpretation	 of	 these	 findings:	 the	 EEG	periodicities	contributing	to	the	enhanced	SSRs	may	not	reflect	an	actual	neural	entrainment	 but,	 instead,	 might	 be	 driven	 by	 a	 sequence	 of	 auditory	 event-related	potentials	(ERPs).	In	an	earlier	study,	Nozaradan	et	al.	(3)	dismissed	this	interpretation,	by	 simply	 reporting	 that	only	 the	 first	 stimulus	of	 the	 sequence	elicited	an	ERP,	while	the	following	stimuli	did	not	elicit	“measurable”	ERPs.	The	latter	 point,	 however,	was	merely	 qualitative,	 and	 not	 supported	 by	 statistical	evidence.	 Relevant	 to	 this	 discourse	 is	 a	 recent	 study	 that	 disclosed	 that,	although	habituated	over	 repetition,	 small	 amplitude	ERPs	 are	 reliably	 elicited	even	following	tens	of	consecutive	stimuli	presented	at	1Hz	(4).	These	preserved	ERPs	included	the	supramodal	biphasic	negative-positive	vertex	wave	(N-P),	and	the	 contingent	 negative	 variation	 (CNV),	 a	 slow	 rising	 negativity	 anticipating	predictable	 events.	 Notably,	 these	 ERPs	 are	 commonly	 evoked	 by	 auditory	stimuli	(5,	6).	A	modulation	of	either	of	these	ERPs	could	explain	the	results	reported	by	Lenc	et	al.	(1),	because	both	the	N-P	vertex	wave	and	the	CNV	are	modulated	by	both	temporal	 attention	 and	 anticipation	 (5–7).	 If	 a	 participant	 allocates	 more	Novembre,	G.,	&	Iannetti,	G.	D.	(2018).	Tagging	the	musical	beat:	Neural	entrainment	or	event-related	potentials?.	Proceedings	of	the	National	Academy	of	Sciences,	115(47),	E11002-E11003.	attentional	resources	to	the	periodic	events	marking	the	beat	or	the	meter	(or	to	any	 predictable	 or	 salient	 event	 within	 a	 pattern),	 then	 the	 ERP	 amplitude	associated	 with	 these	 events	 becomes	 larger.	 It	 follows	 that	 such	 time-locked	modulations	would	enhance	the	power	at	beat-	and	meter-relevant	frequencies	(Fig.	 1).	 Supporting	 this	 alternative	 interpretation,	 we	 highlight	 that	 (i)	 the	reported	 SSRs	had	 a	 scalp	 topography	 similar	 to	 that	 of	 both	 the	CNV	and	 the	negative	vertex	wave	(auditory	N1);	 (ii)	 the	SSRs	enhancement	was	contingent	on	attentional	allocation	(5,	6);	(iii)	remarkably,	the	N-P	vertex	wave	has	larger	amplitude	when	 elicited	 by	 low-frequency	 sounds	 (8);	 and	 (iv)	 the	N-P	 vertex	wave	can	also	be	evoked	by	imagery	of	transient	auditory	events	that	are	typical	of	syncopated	rhythms	(9).	This	alternative	interpretation	could	be	easily	tested	by	analysing	the	EEG	data	in	 the	 time	 domain,	 and	 statistically	 assessing	 the	 presence	 of	 ERPs	 in	 the	waveforms	 time-locked	 to	 each	 stimulus	 onset	 (4).	 Should	 this	 analysis	 reveal	the	presence	of	ERPs,	 two	 fundamental	 implications	of	 the	 results	 reported	by	Lenc	 et	 al.	 (1)	 follow.	 First,	 the	 notion	 that	 these	 SSRs	 tag	 neural	 entrainment	should	 be	 reconsidered:	 they	 might	 not	 reflect	 stimulus-induced	 sustained	oscillatory	 activity,	 but	 rather	 repeated	 stimulus-evoked	 transient	 ERPs	 (10).	Second,	 the	 cortical	 network	 underlying	 the	 results	 of	 Lenc	 et	 al.	 (1)	 and	Nozaradan	 et	 al.	 (3)	 would	 subserve	 attention	 to	 and	 anticipation	 of	 salient	events	(5–7).			Novembre,	G.,	&	Iannetti,	G.	D.	(2018).	Tagging	the	musical	beat:	Neural	entrainment	or	event-related	potentials?.	Proceedings	of	the	National	Academy	of	Sciences,	115(47),	E11002-E11003.	Figure.	 Two	 possible	 physiological	 interpretations	 of	 the	 EEG	 SSRs	reported	 by	 Lenc	 et	 al.	 (1).	 The	 ‘Entrainment’	 account	 postulates	 sustained	oscillatory	 activity	 outlasting	 the	 stimulus	 offset	 (blue	 waveform)	 at	 the	frequency	of	 stimulus	presentation	and	 the	perceived	beat.	The	 ‘ERPs’	 account	postulates	 event-related	 potentials	 (red	 waveform;	 in	 this	 example	 the	 N-P	vertex	wave)	 evoked	by	 each	 stimulus,	with	 greater	 amplitude	 recorded	when	stimuli	reflect	salient	periodicities	like	the	beat.	Both	accounts	predict	a	similar	enhancement	 of	 EEG	 power	 (right)	 at	 the	 frequencies	 corresponding	 to	 both	stimulus	presentation	 (Top,	 purple	 dashed	 lines)	 and	 the	perceived	beat	 (Top,	green	dashed	lines).	ISI,	inter-stimulus	interval.				References			1.		 Lenc	T,	Keller	PE,	Varlet	M,	Nozaradan	S	(2018)	Neural	tracking	of	the	musical	beat	is	enhanced	by	low-frequency	sounds.	Proc	Natl	Acad	Sci	115(32):8221–8226.	2.		 Hove	MJ,	Marie	C,	Bruce	IC,	Trainor	LJ	(2014)	Superior	time	perception	for	lower	musical	pitch	explains	why	bass-ranged	instruments	lay	down	musical	rhythms.	Proc	Natl	Acad	Sci	111(28):10383–10388.	3.		 Nozaradan	S,	Peretz	I,	Missal	M,	Mouraux	A	(2011)	Tagging	the	neuronal	entrainment	to	beat	and	meter.	J	Neurosci	31(28):10234–40.	4.		 Mancini	F,	et	al.	(2018)	Characterizing	the	Short-Term	Habituation	of	Event-Related	Evoked	Potentials.	eneuro	5(5):ENEURO.0014-18.2018.	5.		 Legrain	V,	Iannetti	GD,	Plaghki	L,	Mouraux	A	(2011)	The	pain	matrix	reloaded:	A	salience	detection	system	for	the	body.	Prog	Neurobiol	93(1):111–124.	6.		 Nobre	AC,	Van	Ede	F	(2018)	Anticipated	moments:	Temporal	structure	in	attention.	Nat	Rev	Neurosci	19(1):34–48.	7.		 Breska	A,	Deouell	LY	(2017)	Neural	mechanisms	of	rhythm-based	temporal	prediction:	Delta	phase-locking	reflects	temporal	predictability	but	not	rhythmic	entrainment.	PLoS	Biol	15(2):1–30.	8.		 Wunderlich	JL,	Cone-Wesson	BK	(2001)	Effects	of	stimulus	frequency	and	complexity	on	the	mismatch	negativity	and	other	components	of	the	cortical	auditory-evoked	potential.	J	Acoust	Soc	Am	109(4):1526–1537.	9.		 Hubbard	TL	(2010)	Auditory	imagery:	empirical	findings.	Psychol	Bull	136(2):302–29.	10.		 Barczak	A,	et	al.	(2018)	Top-down,	contextual	entrainment	of	neuronal	oscillations	in	the	auditory	thalamocortical	circuit.	Proc	Natl	Acad	Sci	115(32):E7605–E7614.		

Tagging the musical beat: Neural entrainment or event-related potentials

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Tagging the musical beat: Neural entrainment or event-related potentials

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