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tone
Terence W. Picton, Christopher R. Skinner, Sandra C. Champagne, Adrian J. C. Kellett, and Anita C. Maiste
165 J. Acoust. Soc. Am. 82 (1), July 1987 0001-4966/87/070165-14500.80 1987 AcousticalSociety of America 165
modulationof a continuous
tonewaslargestat modulation The amountof amplitudemodulationwascalculatedas
frequencies
between25 and 50 Hz, and largerfor carrier
(Ama
x -- Amin)/(Amax -{-Amin),
frequencies
at 500 or 1000Hz than for highercarrierfre-
quencies.
The amplitudeof theresponse evokedby a 50-Hz whereAma Xisthemaximumamplitude ofthesignalandAmi n
modulationdecreased
with decreasing
modulationdepth is the minimumamplitude.This formulais the sameasthat
but couldstill be recognized
at a depthof 11%. Reeset al. forthe"modulation index."Theintensityof thisamplitude-
modulated stimulus was calculated on the basis of the mean
(1986) have recentlydescribedhuman steadystate re-
sponsesto amplitudemodulationat frequenciesfrom 0.5 to amplitude.
400 Hz. Hall (1979) and Chamberset al. (1986) have de- The amountof frequencymodulationwascalculatedas
scribed
frequency
followingresponses
to theamplitude
en- (fmax -- fmin)/fcarrier,
velopesof complextones.
wherefmax is the highestfrequency,fmin is the lowestfre-
Pictonet al. (1987a) recordeda steadystateMLR to
quency,andfcarrier is the carrierfrequency.
frequency modulation. Like the AM response reportedby Theseformulasare not equivalentsincethe divisorfor
Kuwadaet al. (1986), this response waslargerfor lower
thefrequency modulation isone-halfof (fmax --fmin). The
carrierfrequenciesandlargerat higherintensities.
At modu-
amountof frequencymodulationcan alsobe measuredby
lation rates of 40 Hz, Picton et al. found that the FM re-
AF whichisthedifference between fmax andfcarrier, or by a
sponsecould be reliably recognizeddown to modulation
modulationindexwhichequalsAF/Fmod,whereFmoaisthe
depthsof 10%.
frequencyof the modulatingsignal.
The presentarticledescribes severalexperiments evalu- Figure 1 presents the acousticwaveformsandthe spec-
ating the humansteadystateresponses to the sinusoidal tra for AM and FM toneswith a carrierfrequencyof 1000
modulationof the amplitudeor frequencyof a tone. The Hz and a modulatingfrequencyof 40 Hz. For-theAM tone,
goalsof thesestudies were:to determine theoptimalrecord- theenergyof thestimulusisconcentrated at thefrequencyof
ingconditions fortheresponse withrespect to thefrequency the carrierandat two sidebands(gcarrie r --gmod), with the
of modulation andthelocationof therecording electrodes; relativeenergiesin thecarrierandthesidebandsdepending
to assess therelationsoftheresponse to changing theintensi- uponthe amountof amplitudemodulation.The spectrumof
ty andfrequency of thetoneor thedepthof themodulation; an FM signalvarieswith the modulationindex.For modula-
to relatetheresponse to the subject's
abilityto discriminate tion indicesbelow0.25, the amplitudespectraof AM and
frequencyand intensity;and to investigate the relations FM tonesareverysimilar.However,with largermodulation
betweenAM andFM responses.
indices(asin ourexperiments), thespectraof thefrequency-
modulatedstimuluscontainenergyin bandsout to F
I. Methods from the carrierfrequencyin bandsoccurringat intervals
A. Stimuli
equalto the modulatingfrequency.
The sinusoidalsignalcontrollingthe amplitudeor fre-
A continuous tonewaspresented to thesubject's right quencymodulationwasgeneratedby the MINC 11/23 com-
earthroughan unshielded TDH-49 earphone. Pilotwork puter and smoothedusinga low-passfilter. The sameclock
usingearplugs hadshown thattheresponse wasnota result controlledboth the modulationsignaland the A-D conver-
of electromagnetic artifact.Thetonewaspresented with sion.Thisclockhada frequencyof 125Hz for thefirstpartof
briefpauses ofbetween 1and2 soccurring every10s.The experiment1 and 500 Hz for all otherrecordings.Because
amplitudeor thefrequency of thetonecouldbemodulated the modulationsignalwasnot a pure tone,therewassome
usinglaboratory-constructed devices. distortionin theAM stimulicausingextrabandsof energy
166 J.Acoust.
Soc.Am.,Vol.82,No.1,July1987 Picton
eta/.'Potentials
evoked
bysinusoidal
AMorFM 166
(near 1500 and 500 Hz in the spectrumfor the 1000-Hz doesnot containthe origin,the recordedresponse can be
carrier) that were about 40 dB below the intensity of the considered reliablydifferentfromzeroat theprobabilityfor
carrier. which the ellipseis calculated.
Althoughthe analysis wasperformed off-linefor most
B. Recording oftheexperiments, anon-lineprotocolwasusedin thestud-
iesof thresholdand in evaluatingthe relationshipsbetween
Electroencephalographic (EEG) signalswererecorded AM andFM. The on-lineprotocolrecordedonlyonechan-
from the scalpuing gold-platedsurfaceelectrodes
applied nel of EEG (Cz-M2). A smallnumber (e.g., 40) of sweeps
with collodion. The active electrodes were located at the ver-
wereaveragedon-lineto form a "subaverage"waveform.
tex (Cz) or at the left and right central regions(C3, C4). The X and I/valueswerecalculatedfor thissubaverage
in the
The referenceelectrodeswere located on the right or left briefpausepriorto beginninganothersubaverage. Several
mastoids (M 1 and M2) or the larynx (La). Two separate (e.g., 10-20) suchsubaverageswerecombined to givethe
channelsof EEG signalswererecordedin mostexperiments. finalresponse.A T 2valuecouldthenbecalculated
fromthe
ThesewereC3-La and C4-La in the first experimentand Cz- setof X and I/measurementsobtainedfrom the subaverages.
M 1andCz-M2 in subsequent experiments. The EEG signals Onevariantof theon-lineprotocolallowedthesimultaneous
were amplifiedand filtered using Beckmanmodel R611 calculationof responses
at twodifferentfrequencies
of stim-
polygraphamplifierswith a bandpassof 0.5-100 Hz (first ulation.
experiment)or 5-100 Hz (subsequent experiments).The The spectrum of theaverageresponsewasexaminedus-
polarityconventionof the recordingwasthat negativityat ing a fast Fouriertransform.This providedinformation
the scalprelativeto thelarynxor mastoidwasrepresented
as aboutthe activityin the response
at frequencies
otherthan
an upwarddeflectionin thewaveformandasa positivevalue the frequencyof stimulation.
in the analysis. An "apparentlatency" (Regan, 1982) was estimated
The analysiswindowlasted1024msfor the firstpart of whenresponses wereobtainedat severalratesof stimulation.
the firstexperimentand256msfor all otherrecordings. The If thephasedelaysarelinearlyrelatedto therateof stimula-
numberof data pointsper analysiswindowwas256. When tion,theapparent latencycanbecalculated fromtheslopeof
two channelswererecorded,this represented128pointsper theregression of phasedelayon rate.The apparentlatency
channel.Rotating buffersallowedcontinuousstimulation equalsthisslopedividedby 360.
and recording--while one sweepwas being recorded,the The resultsof the experimentswere evaluatedwhere
previoussweepwasbeinganalyzedand/or storedon disk. appropriatewith repeatedmeasures analysesof variance
The first two sweepsoccurringafter any pausein the tone (BMDP2V) usingGeisser-Greenhouse adjustments when
were not analyzed. calculatingthe probabilitylevels.Posthoctestingwasper-
formedusingNewman-Keulsprocedures. The significance
C. Analysis level wasp < 0.05.
In the off-line recordingprotocols,a total of 500 (120
for the firstpart of the experiment)sweepswererecordedon D. Subjects
disk in each experimentalcondition.Pairs of sweepswere
then averagedto provide250 setsof data (this stepwasnot Seventeensubjects(8 female) participatedin various
necessaryfor the first part of experiment1). Each point in partsof theseexperiments.Their agesrangedbetween22
the sweepwasthenseparatelymultipliedby thesinefunction and41 (mean28) years.All subjectsdemonstrated hearing
at that latencyfor the frequencyof stimulationand by the thresholdsof lessthan 20 dB HL at the frequenciestested.
cosinefunction. The averagesof each of these products Most subjects
participatedin severalexperiments.During
acrossthe sweepweremultipliedby a scalingfactor to pro- an experiment,the subjectsat in a comfortable
chairin a
vide the values"X" and "." The amplitudeof the response sound-attenuated
room.In mostof the experiments,the sub-
at the frequency of stimulation was then calculated as jectsreadwhiletheirresponses
wererecorded.The EEG was
monitored to ensurethat they did not fall asleep.
x/X2'+ y 2 andthephase
oftheresponse
wastan- (Y/X).
This procedurerepresentsFourier analysislimited to a sin-
glefrequency.A "phasedelay" (Rodriguezet al., 1986) was II. RESULTS
calculatedas 360minus the phaseof the responseof zero
time. This measurementis morehomologousto latencythan A. Experiment 1: Modulation frequency
the phaseat zero time. Phasemeasurementswere expressed This experimentevaluatedthe steadystateresponses
to
in degrees. AM or FM tonesat differentmodulation frequencies.The
The repeatedX- measurementscan be representedas stimuluswasa 1000-Hztonepresented
at anintensityof 76.5
a cloudof pointson a polar plot. Simplecalculationsof the dB SPL (70 dB HL). The amountof modulationwas 100%
meanvaluefor X and yield the coordinatesof the average for both AM and FM. In the first part of the experiment,a
responseover the recordingsession.The reliability of the 1024-ms window was used and modulation occurred at fre-
response
canbe assessed
usingthe Hotelling'sT 2 statistic quenciesof 2.0, 2.9, 4.9, 6.8, 8.8, 10.7,and 12.7 Hz. Re-
(Picton et al., 1987b). Using the variance-covariancema- sponseswereevaluated over120sweeps (about2 min). The
trix of the results,an ellipseis constructedto outline the secondpart of the experimentuseda 256-mswindowand
confidencelimits for the mean measurement.If this ellipse evaluatedresponses at frequencies of 15.6, 19.5,27.3, 35.2,
I'O
0'5 0-5
., C3
PHASE (') o [] C4 PHASE onC4
180
// ,
0 5 I0 15 $0 45 60 0 5 I0 15 30 45 6O
AM FM
Modulation Subjects Group Subjects Group
168 J. Acoust. Soc. Am., Vol. 82, No. 1, July 1987 Pictoneta/.' Potentialsevoked by sinusoidalAM or FM 168
cent responsesthat were reasonablyreliable. The average
6-8 apparent latency over this region was 149 ms (range 53-
350).
-1'OpV 0-3.uv The FFT analysesof the averageresponses over all sub-
o24r,
s 0 61.5 Hz
169 J. Acoust.Soc. Am., Vol. 82, No. 1, July 1987 Pictoneta/.: Potentialsevokedby sinusoidalAM or FM 169
2. Threshold for detecting modulation TABLE II. Thresholdsfor detectingmodulation.[Modulationdepthisex-
pressedasa percentage.The numbersrepresentthe meansandranges(in
The secondpart of thisexperimentestimatedthethresh- parentheses)for eightsubjects.
]
old for recognizingthe evokedpotentialto eitherAM or FM
Thresholds
andcomparedthisto psychoacoustic thresholds.The stimu-
lus was a 70-dB HL, 1000-Hz tone modulated at 39.1 Hz. Type of modulation Behavioral Evokedpotential
On-line recordingprotocolswere used.The responsewas Amplitude 5.1 4.9
judgedpositive
if theT2 valueattained
a significance
levelof (2-7) (3-8)
p <0.05 within 1200sweeps(in 30 blocks). For the higher Frequency 2.8 5.8
(2-3) (2-13)
modulationlevels,the analysiswas often stoppedafter 400
sweeps( 10blocks).The subjectmaintainedalertness by de-
tecting occasional(p- 0.1) blockswherein there was no
modulation.The stimuliwerepresentedto the right ear and
significance(F= 5.1,df= 1,7,0.05<p <0.10). The differ-
the recordingsweretakenfrom Cz-M2. The psychoacoustic
ence between these two thresholdsvaried acrosssubjects
thresholdswereestimatedon separatetrials, usinga forced-
between 0% and d- 10%.
choiceresponseto the equallyprobablepresenceor absence
of modulationin 2-s periods.Nonparametrictechniques
C. Experiment 3: Intensity of the carrier
(Pollack and Norman, 1964) were usedto evaluatethe sub-
1. From 30 to 70 dB HL
ject's detectionperformance.Modulation wasjudgedto be
suprathreshold if A ' wasgreaterthan 0.7. This experiment evaluatedthe effectsof changing the
Samplerecordings areshownin Fig. 6. The amplitudeof intensityof the carrier.The firstpart of the experimentvar-
the responsedecreasedas the depth of modulationde- ied the intensityfrom 30 to 70 dB HL. The carriertoneof
creased.There were no significantchangesin phaseas the 1000Hz waspresented to therightearandrecordings were
responseapproachedthreshold.At 5% modulation,the takenfrom Cz-M 1 andCz-M2. On thebasisof thepreceding
averagephasewas 169for AM and 108for FM (cf. Fig. 5). experiment,
wedecidedto usea modulation
depthof 50%
The thresholds obtainedin thesecond part oftheexperiment for FM and 90% for AM, sincethesegaveresponses
of com-
areshownin TableII. The thresholdfor recognizing theAM parablemagnitude.
Sixsubjects
participated
in thisexperi-
responsewas not significantlydifferentfrom the psycho- ment.The evokedpotentialswereobtainedover500sweeps
acousticthreshold.The differencebetweenthe evokedpo- of 256 ms usinga modulationrate of 39.1 Hz.
tential and the behavioralthresholdvariedacrosssubjects The resultsof thefirstpartof thisexperimentareshown
between -- 3% and d- 2%. The thresholdfor recognizing in Fig.7. Therewasa significantincreasein theamplitudeof
an FM responsewas higherthan the perceptualthreshold theresponse withincreasing intensityofthecarrierforboth
althoughthis differencedid not reachthe level of statistical AM (F = 32.3, df= 4,20, p < 0.001) and FM (F = 9.7,
df= 4,20,p < 0.01). Theamplitude-intensity functions
were
differentbetweenthetypesof modulation. The AM response
increasedmonotonically,whereasthe FM responsein-
AM FM
creasedrapidlyup to an intensityof 40 dB and thenin-
creasedlessrapidly (intensityX modulationtypeinterac-
tion: F= 3.9, df= 4,20, p <0.05). The phasedecreased
significantlywith increasingintensity for both AM
(F=33.3, df=4,20, p<0.001) and FM (F=30.8,
df= 4,20,p <0.001 ). The phaseof the FM response was
significantlyshorterthan the phaseof the AM response
(F= 74.4, df= 1,5,p<0.001).
0-3pV
20, I0, ,5,4, 3,2 .
20, I0, ,5,3,2, I AMPLITUDE (pV) PHASE (*)
180]
dot. On the left are shownthe resultsof a studyto measurethe thresholdfor
recognizingthe evokedpotentialwhenthe modulationdepthfor a 70-dB
o ' 5o 5 7o 3o 5o
HL AM toneis decreased from 20% to 2%. The response becomessmaller
(approachedthe origin) as the modulationdepth decreases. At 3%, the INTENSITY (dB HL)
responsewasno longerrecognizableas differentfrom zero; i.e., the confi- AM FM o
dence-limitellipseincludedthe origin. The EP thresholdwas therefore
measuredas 4% (arrow). The psychoacoustic thresholdfor this subject FIG. 7. Effectsof stimulusintensity.The stimuluswaseithera 90% AM
was 6%. On the right of the figure are shownthe resultsobtainedwhen tone or a 50% FM tone. On the left are shownthe effectsof intensityon the
measuringthe thresholdfor a 70-dB HL FM tone. The EP thresholdwas amplitudeof theresponse andontherightareshowntheeffects
onphase
2% (arrow). The psychoacoustic thresholdwasalso2%. delay.Thisfigureplotsthevector-averaged
datafromsixsubjects.
AMPLITUDE (pV)
PHASE
[.O'
AM
FM IOO
0-5.
o
5o 40 50
ioi/
50 40 5O
O
0 0 'o
IOO]
O!
0 30 'o
CARRIER
[] 500
FIG. 9. Effectsof carrier frequency.Theseef-
fectswerestudiedat modulationdepthsof 10%,
30%, and 50%. The carrierfrequencywas 500,
o IOOO 1000, 2000, or 4000 Hz. The intensity of the
Hz
2000 stimuluswas76.5 dB SPL. This figureplotsthe
I'O
v 4000 vector-averageddata from eight subjects(six
subjectsfor the 30% responses).
FM o.5
IO 30 50
individualsubjectsfrom the slopesof the regressionlinesfor phase delay showed significant effects of frequency
phasedelayversusmodulationfrequency.There wasa ten- (F = 4.8, df-- 3,15,p <0.05) but no effectsof modulation
dencyfor the apparentlatencyto beslightlyshorterfor high- depth.
er carrierfrequencies with the 2000-Hz responsesoccurring The responses at 10% modulationdepthswere very
on average3.2 ms earlierthan the 500-Hz responses. How- small.For the AM responses, only 3 of 8 weresignificantly
ever, statisticalanalysisshowedno significanteffectfor ei- (p < 0.05) differentfrom zeroat 500 Hz and 1 at 4000Hz.
ther carrier frequencyor type of modulation.The average For theFM responses, all 8 weresignificantlydifferentfrom
apparentlatencyover all conditionswas 39.8 ms (range 29- zero at 500 Hz but only 3 at 4000 Hz.
52 over all conditions).
172 J. Acoust.Soc. Am., Vol. 82, No. 1, July 1987 Pictoneta/.' Potentialsevokedby sinusoidalAM or FM 172
ALONE SAME DIFFERENT
2. Selective adaptation
The secondpart of thisexperimentderivesfrom the con-
cept of selectiveadaptation (cf. Regan and Tansley, 1979;
AM Tansleyand Suffield,1983). The prolongedpresentationof
an amplitude-modulatedtonecausesan increasedthreshold
for detectingamplitudemodulationbut doesnot affectthe
thresholdfor detectingfrequencymodulation. The pro-
FM
longedpresentationof a frequency-modulated tone causesa
substantialincreasein the thresholdfor detectingfrequency
:55 51 Hz . 0.SpV
IOO Hz modulationand a slightincreasein the thresholdfor detect-
ing amplitudemodulationbecauseit is impossibleto modu-
FIG. 10. Concurrent stimulation. The stimuli were either AM or FM tones
late the frequencyof a tonewithoutalsomodulatingits am-
presentedat either 35 or 51 Hz either ALONE or in combinationwith an-
othermodulationat the otherfrequencythat waseitherthe SAME or DIF-
plitude at the level of the receptor.The adaptationoccurs
FERENT in type. The basiccomparisons(cf. Table IV) are betweenthe overa periodof 5-15 min. Recoveryoccurswith a periodof
responses
whichhaveamplitudes
thatareindicatedbysimilartriangles(ei- 1-3 min.
ther open or closed). When there is concurrentstimulation with another In this part of the experiment,the techniqueof simulta-
modulation,the responseissmallerthan whenit is alone.However,thereis neous stimulation was used to record concurrent AM and
no significant
difference
betweenthe amplitudeof the responses
whenthe
concurrent modulation is either the same or different. FM responses beforeand after exposureto prolongedAM or
FM tones.The responses wereevokedat frequencies of 39.1
and 43.0 Hz (9 and 11 cyclesper sweep). Responseswere
acrossthe differentratesto give three basicmeasurements analyzedover 500 sweeps(about 2 min). The depthof AM
for eachtypeof modulation:A lmthe amplitudeof the re- was 50% and of FM was 30%. The tone intensitywas 60 dB
sponsewithoutanyconcurrent stimulation; A2mthe ampli- HL. The post-adaptationrecordingcommencedwithin 15 s
tudeof the responsewhentherewasconcurrentstimulation of theendof a 10-minperiodof adaptationwith a continuous
of the sametype;and A3the amplitudeof the response 60-dB HL tone that was amplitude (50%) or frequency
whentherewasconcurrent stimulationwith the othertype (30%) modulatedat a modulationfrequencyof 40 Hz.
of modulation.The primaryhypothesis wasthat therewould There were four sets of before and after measurements
be someindependence of the FM and AM responses, i.e., determinedby the selectionof the frequencyfor AM and
that A2 wouldbesignificantly smallerthanA3. The second- FM, andby the typesof adaption(AM or FM ). The orderof
ary hypothesiswas that there would neverthelessbe some thesefour setsof recordingswasbalancedacrossthe eight
interactionbetweenthetworesponses,
i.e.,thatA3 wouldbe subjects.There was a 5- to 10-minrest periodbetweenthe
significantlysmallerthan A 1. sets.
173 J. Acoust.Soc.Am.,Vol. 82, No. 1, July 1987 Pictonot a/.' Potentialsevokedby sinusoidal
AM or FM 173
periods) betweenthesetwo recordingsshowedchangesof (1986). Kuwadaet al. (their Fig. 2) reportfor onesubject
between -- 58% and -+-55% (mean: -+- 14%) for the AM anamplitude of 0.35/iV fora 60-dBHL, 1-kHztonemodu-
response and between-- 36% and -F 69% (mean: -+-8%) latedat a depthof 91% anda frequency of 50Hz. Reesetal.
for the FM response.(For thesecomparisons, the percent- (theirFig. 8) reportedfor twosubjectsan amplitudeof ap-
ageswere calculatedusingthe larger measurements as the proximately 1/iV for a (?55dB SL) 1-kHztonemodulated
denominatorand usingthe sign to indicatethe direction of at a depthof 100%anda frequency of40Hz. Ourresults(60
change.) dB HL, 1 kHz, 90%, 40 Hz) showan averageamplitudeof
0.85/iV. Kuwadaetal. reported
littlechange
in phase(per-
III. DISCUSSION haps30or 40) with changein modulationdepthandwe
wereunableto demonstrate any significantchange(Fig. 5).
A. Frequency of modulation
Our resultsalsoreplicatethe strikingeffectof carrierfre-
Our resultsindicatethat human steadystateresponses quencyon the response.
The apparentlatencyreportedby
evokedby the modulationof the amplitudeor frequencyof a Kuwada et al. (30.8, 31.8 ms) was somewhatshorterthan
pure tone are mostreliablyrecordedat frequenciesnear 40 our average(37.2 ms) but of the sameorderof magnitude.
Hz. At lower frequencies,
the responsesare moredifficultto Reesetal. reportedan approximately linearincrease
in am-
recordbecausethe EEG noiselevelsare higher at thesefre- plitudewith log intensityand a tendencyfor saturationat
quenciesand becausethe very slowness of the modulating intensities of above 60 dB. Similar results are shown in
signalincreases thedurationof theanalysis. However,there Fig. 7.
are someregionswherethereare clearresponses: between2 The 40- to 50-Hz AM response is probablyverysimilar
and 5 Hz for the AM responses andbetween3 and 7 Hz for to the steadystateMLR elicitedby rapidlypresenteddis-
the FM responses. The FM responses in thefrequencyrange crete tones (Galambos et al., 1981; Stapellset al., 1984;
3-7 Hz appearthemostpromisingfor furtherinvestigation. Rodriguezetal., 1986).The amplitudesof theresponses are
With an apparentlatencyof approximately150 ms, these similar.The phasedelayof the response to discretetonesis
responses may be relatedto the N1-P2 components of the somewhatshorter.Rodriguezet al. reporta phasedelayof
evoked potential to transient changesin frequency(re- 145between40 and 70 dB HL for a 1-kHz tone, whereasthe
viewedby Arlinger et al., 1976). phasedelayfortheAM response wasslightlyover200. This
Previousreports(RickardsandClark, 1984;Reesetal., difference
is probablyrelatedto the morerapidrisetime (4
1986) havesuggested that with increasingmodulationfre- ms) of the toneburstscomparedto the sinusoidalmodula-
quencythe AM response showsa low-passfunctionwith a tion.
small resonanceeffectat frequenciesnear 40 Hz. One diffi- The cerebralsourceof the 40-Hz responseis not yet
culty with this formulationis that it doesnot considerthe known.Galambos(1982) suggested that theresponse might
increasedbackgroundnoisein the recordingat the lower be generatedin the polysensory regionsof the thalamus.
frequencies. Signal-to-noise estimates basedontheT2statis- Spydellet al. (1985) foundthat the phaseof the 40-Hz re-
tic (Picton et al., 1987b) certainlyindicatehighernoiselev- sponse wasaffectedby lesionsof the thalamusor midbrain
elsat low frequencies. Part of themeasuredamplitudeat low butnotbylesions ofthetemporal lobe.However, magnetic
modulationfrequenciesmay thereforereflect the residual responses to stimulimodulatedor presented at frequencies
noisein the recording. near 40 Hz (Romani et al., 1982; Mfikelfi and Hari, 1987)
One way to decreasethe noisein the recordingsis to andepidural recordings(Leeetal., 1984)suggest anactive
averagethe responses acrosssubjectspayingdue regardto source in theregionof theprimaryauditorycortex.A simi-
phase.As seenin Fig. 2, thischanges themodulationtransfer laroriginforthetransient MLR hasbeensuggested byscalp
functionfor the AM responsefrom a simplelow-passfunc- distributionanalyses(Cohen, 1982;Schergand von Cra-
tion to onewith differentresonantfrequenciesin the 2- to 5- mon,1986),andbytheeffects of lesionsin thetemporallobe
Hz and the 20- to 50-Hz regions.However,combiningdata (Krauset al., 1982;SchergandvonCramon,1986).
acrosssubjectsdoesnot distinguishbetweennoiseandinter- Studiesin animalshavesuggested thatcorticalneurons
subjectvariability.Prolongedaveragingof theresponses of a donotrespond tofluctuations
in amplitude atfrequenciesof
singlesubjectwould be necessaryto differentiatetheseef- greater than20Hz (Creutzfeldt etal.,1980)although brain-
fects.
stemneuronscaneasilyfollowveryrapidchanges in ampli-
The presenceof a secondharmonicsuggests that there tude(orfrequency)
(ReesandM411er, 1983). Nevertheless,
are potentialsevokedby both the increaseand the decrease althoughthemajorityofcorticalneurons
donotrespond to
in the amplitudeor frequencyof the tone.Reeset al. (1986) rapidfluctuations,
therearesomecorticalneurons
thatdo
foundsecondharmonicAM responses to belargestat modu- respond to frequencies
of greater
than30Hz (Fastletal.,
lation frequencies
between5 and 20 Hz. We foundthat the 1986;Schreinerand Urbas, 1986).
second harmonic was much smaller and much less consis-
The 40-Hz response is quitevariable.Galamboset al.
tent than the fundamentalresponseat all modulation fre- (1987) havedescribed minute-to-minute fluctuations
in the
quencies. amplitudeof the response. In our experiments,it wasnot
unusualto observethe amplitudechangeby a factorof two
B. AM responses and the 40-Hz potential fromonerecordingsession to another.The natureof these
The 40-Hz responseto AM tonesis similar to the re- fluctuationsis not knownalthoughtheymightbe relatedto
sponsereportedby Kuwadaet al. (1986) andby Reeset al. concomitantchangesin arousal.Certainly,drowsiness and
175 J.Acoust.
Soc.Am.,Vol.82,No.1,July1987 Picton
eta/.:Potentials
evokedbysinusoidal
AMorFM 175
postulatedthat an FM neuroncouldhavethe synapticin- tion frequency
increases
to 40 Hz. Our averagethreshold
putsfrom differentfrequency-specificneuronsarrangedac- wasequivalentto 28 Hz.
cordingto frequencyin an orderly sequenceof distances The steadystateresponsescan be detectedby modu-
from the triggerzonefor the actionpotential.As the stimu- lationdepthscloseto thebehavioralthresholds.
Theymay
lusfrequencychanged,spatiotemporal summationwouldbe therefore
behelpfulin theobjective
evaluation
of a patient's
more efficientif the stimulussweptthe synapticbombard- ability to discriminatesounds.
ment toward,rather than awayfrom, the triggerzone. The responses mayalsobeusefulin objectively evaluat-
inghearingthresholds, Our resultsindicatethat in normal
D. Relations between AM and FM responses subjectsthe thresholdcanbe estimatedto within 5-20 dB.
Similarresultshavebeenreportedby Kuwadaet al. (1986)
The resultsof concurrentstimulationsuggestthat there in patientswith hearinglosses.
Oneparticularadvantage of
is a similargeneratorfor bothFM andAM responses. If the usingAM tonesratherthan discretetonesis that theyare
generatormechanisms wereindependent,concurrentmodu- morefrequency specific.
The amplitudespectrumof an AM
lation of both frequencyandamplitudeshouldhaveresulted tonecontainsenergyonlyat the carrierfrequencyandat two
in larger responses than concurrentmodulationof only one sidebands,separated from the carrierfrequencyby the fre-
parameter.A possible modelfor theseresponsescouldbegin quencyof the modulatingsignal.They are thereforemuch
with an initial frequency/intensity
analysisthroughthe hair more frequencyspecificthan brief tone bursts.The FM
cells and auditory-nervefibers.Certain brain-stemneurons tonesmay alsobe usedalthoughthey are lessfrequencyspe-
whichare specifically responsive to changesin eitherampli- cific than the AM tones.One difficulty in usingthe 40-Hz
tude or intensityare the neurophysiological basisfor sepa- response asanobjective measurement for hearingthresholds
rate AM and FM channels(cf. Tansleyand Regan, 1979). is that it varies with the state of the subject (reviewed by
Once the auditory information has been analyzedin these Linden et al., 1985). Furthermore, it is difficult to record in
separatesystems,both channelsactivatea muchlessspecific infants(Stapellset al., 1987), the patientsfor whomobjec-
mechanismthat generatesthe 40-Hz steadystateresponse. tive audiometryis most important.
One of the characteristics
of a channelis its susceptibil-
ity to selectiveadaptation.Our experimentsshow no evi- ACKNOWLEDGMENTS
dencefor such adaptationin the 40-Hz response.This is
This researchwassupportedby the Medical Research
quite differentfrom the clear adaptationshownfor the de-
Council of Canada and by the Ontario DeafnessResearch
tection of modulation (Tansley and Suffield, 1983). These
Foundation.Marjorie Berry typed the manuscript.Edna
resultssuggestthat the neuralbasisfor adaptationoccursin
Leech assistedin the data analysis.Gilles Hamel provided
a systemseparatefrom that responsible for generatingthe
essentialtechnicalsupport.
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