Some Statistical Methods for Random Process Data from Seismology and Neurophysiology Author(s): David R. Brillinger Reviewed work(s): Source: The Annals of Statistics, Vol. 16, No. 1 (Mar., 1988), pp. 1-54 Published by: Institute of Mathematical Statistics Stable URL: http://www.jstor.org/stable/2241421 . Accessed: 23/01/2012 17:53 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Institute of Mathematical Statistics is collaborating with JSTOR to digitize, preserve and extend access to The Annals of Statistics. http://www.jstor.org The Annals of Statistics 1988, Vol. 16, No. 1, 1-54 THE 1983WALD MEMORIAL LECTURES SOME STATISTICAL METHODS FOR RANDOM PROCESS DATA FROM SEISMOLOGY AND NEUROPHYSIOLOGY1 BY DAVID R. BRILLINGER University ofCalifornia, Berkeley To Jeff AustinBrillinger, B.A. Examplesare presentedof statisticaltechniquesforthe analysisof randomprocessdata andoftheirusesin thesubstantive fieldsofseismology and neurophysiology. The problems addressedincludefrequency estimation fordecayingcosinusoids, associationmeasurement, signalestimation, causal connectionassessment, estimationof speed and directionand structural modeling. The techniques includecomplex nonlinear employed demodulation, regression, probitanalysis, maximum value deconvolution, likelihood, singular Fourieranalysisand averaging. decomposition, Table ofcontents I. Introduction II. Seismology 1. The fieldand itsgoals 2. Freeoscillations oftheEarth 3. Estimationoffault-plane parameters 4. Quantification ofearthquakes 5. Arraydata 6. Exploration seismology (reflection seismology) 7. Othertopics 8. Discussion 9. Update III. Neurophysiology 10. The fieldanditsgoals 11. Neuronalsignaling 12. Assessingconnectivities 13. A structural stochastic model 14. Analysisofevokedresponses 15. A confirmed (Fourier)inference 16. Othertopics 2 3 3 4 10 14 18 21 23 24 24 25 25 26 32 34 36 44 45 ReceivedDecember1986;revisedJuly1987. 'Prepared with the partialsupportof NationalScienceFoundationGrantsCEE-79-01642, and whiletheauthorwas a Guggenheim Fellow. MCS-83-16634 AMS 1980subjectclassifications. Primary 62M99,62P99. Key wordsand phrases.Arraydata, averageevokedresponse, binarydata, Fourierinference, complexdemodulation, spatial-temporal processes, neurophysiology, pointprocesses, probitanalysis, timeseries. seismology, systemidentification, D. R. BRILLINGER 2 17. Discussion 18. Update remarks IV. Concluding 47 48 48 I. Introduction Thepurposeofstatistics,...,is todescribecertainrealphenomena. A. Wald(1952) in timeand/orspace. The concernof theselecturesis raw data distributed The basic data are curvesand surfaces.If n denotesthe samplesize and p thentheconcern is withthecase of n muchlessthanp. denotesthedimension, in havedevelopedor are developing thephenomena In the situationsaddressed, so thatsubjectmatterplaysessentialrolesin timeor space. Theyare complex, drawn.Thereneed and conclusions theanalysesmadeand in theinterpretations Indeed,a principal reasoning. ofbothphysicaland statistical to be combinations goal ofthe lecturesis to bringout thekeyrolethatsubjectmatterplaysin the intention is to showthatthefieldsof analysisofrandomprocessdata. A further particuare richin problemsforstatisticians, seismologyand neurophysiology The work larlythose individualswithsome interestin appliedmathematics. presentedinvolvesa mixtureof data analysisand structuralmodeling.The employedare broadlyapplibut thetechniques problemsdiscussedare specific, cable. The data concernedis of high quality,so that detailedanalysesare workand a consistsofpersonal(collaborative) possible.The materialpresented methodsthatserveto tiethedevelopment fewsuccessstoriesofotherparticular An attemptis madeto presentproblems froma unifiedpointofview. together. results. ratherthannovelsubstantive Emphasisis on techniques, with ofstatistics The studyofrandomprocessdata providesa majorinterface in collection of the an there has been scienceand technology. explosion Indeed, of in to much modern measurements part (corresponding spatial-temporal behavingbecomedigital).Some particularissuesand procedures technology with of statistics of the interaction technology. as a result come emphasized Fourier systemsanalysis,inverseproblems, These includesystemidentification, funcaveraging (resolutionversusprecision), bias versusvariability inference, run throughthe study.These strains tions,dynamicsand micro-versus-macro the broad rangeof data examplespresented.Thereis also a desireto display mustnowbe concemed. typeswithwhoseanalysisstatisticians Thereis no claimmade that the analysesare Some provisosare necessary. details.Furtheris an overview, ratherthanspecific definitive. Whatis presented to the is of formalism. The reader referred more,thereis littlepresentation forgreaterdetail. papersreferenced on somestatisticalmethodsin Thereare twolectures.The firstconcentrates It is methodsin neurophysiology. thesecondon somecorresponding seismology, of data in the roles analysis to see thesamemethodsplayingcentral interesting fromtwoquitedisparatefields.Indeed,one oftheprincipalgoalsofthelectures of statisticaltechniques-byexamplesfrom was to bringout the universality thesetwofields. SEISMOLOGY AND NEUROPHYSIOLOGY 3 II. Seismology to scientific methodand statistical detailhas beenone Jeffreys...attention has attaineditspresentlevelof ofthemainforcesthrough whichSeismology precision. BullenandBolt(1985) 1. The field and its goals. The termseismology refersto the scientific of earthquakes and relatedphenomena. It has beendefinedas the investigation whichare recordsof mechanical "science based on data called seismograms, vibrationsofthe Earth"[Akiand Richards(1980)].This latterdefinition allows also studyvibrationscaused by the sea, by the admissionthat seismologists volcanoesor byman.Onefurther definition thathas beengivenis: thescienceof in theEarth. strain-wavepropagation Whateverthedefinition, thebroadgoalsofseismology areto learntheEarth's and a planet'sinteriorcomposition and to predictthe time,size,locationand Workersin the fieldseek to strengthof groundmotionin futureearthquakes. providevalid explanations ofearthquake-related and to understand phenomena thesephenomenaso thatlifemaybe madesafer. locationand quantification Specificproblemsaddressedincludethedetection, of earthquakes,the distinguishing of earthquakesfromnuclearexplosionsand in theEarth'sinterior thedetermination ofwavevelocity as a function ofdepth. in seismology The accumulationof knowledge has displayeda steadybackand-forth thewavesand newinsightconcerning betweennewinsightconcerning the mediathroughwhichthewavespropagate.Amongmajor"discoveries" one can list are the innercore,theliquidcentralcore,the Mohorovicdiscontinuity, the movementof tectonicplatescausingearthquakes themselves and the locatingofnumerousgas and oil fields. The fieldis largelyobservational, withthebasicinstruments theseismogram and clock.There are importantexperiments too, wheretailoredimpulsesare vibrations studied.The fieldexperienced inputto the Earth and the resulting the "digitalrevolution"in the 1950sand now poses problemsexceedingthe capabilitiesofeventoday'ssupercomputers. Statisticalmethodshave playedan important role in seismology formany years-in largepartdue to theefforts ofHaroldJeffreys [seeJeffreys (1977),for example].Vere-Jones and Smith(1981)providea reviewofmanycontemporary instances.Statisticsentersfora varietyof reasons.The data sets are massive. Thereis substantialinherent variability and measurement error.Modelsneedto be refined,fittedand revised.Inverseproblemsneed to be addressed.Experimentsneed to be designed.Sometimestheresearcher mustfallbackon simulations.The basicquantityofconcernis oftena (risk)probability. In particular, it may be pointedout,thatin the construction ofthe Jeffreys and Bullen(1940) traveltime tables,one has an early,perhapsgreatestsuccess,of the use of methods.[B. A. Bolt's (1976) presidential robust/resistant address"Abnormal is wellworthreadingin thisconnection.] seismology" deal withdata of a varietyof types.The important Seismologists formsare fromspatial arraysof seismometers digitalwaveforms of variousdimensions D. R. BRILLINGER 4 in sucha fashionthatan earthquake havebeenarranged (wheretheinstruments listsof entity)and catalogs(containing changing signalmaybe seenas a moving, forgeographic regions an event'stimes,locations, sizesand othercharacteristics) ofinterest. is notwithoutitscontroversies. ones,such Thereare fundamental Seismology as whetheror notplatetectonics is a validatedtheory.Thereare practicalones, such as does the size of the motionof an earthquakeincreasesteadilyas one data approachesthefaultordoesit leveloff?As is so oftenthecase,theexisting and analysismethodsproveinadequateto resolvethesedisputesconclusively. backseismological A generalreference thatprovidesmuchof the pertinent of somespecific groundis Bullen and Bolt (1985).We tum to a presentation problemsand techniques. 2. Free oscillations of the Earth. This subjectis one of the principal in seismology overthe last 25 years.Wheneverthereis a great developments The seismogramthen earthquake,the Earth vibratesfor days afterwards. numberofexponentially ofa sumofan infinite decaying consists,approximately, ofthecosinusoids and cosinusoidsplusnoise;see expression (2). The frequencies ratesof decayrelateto the Earth'scomposition. Measured the corresponding The techniques aboutthatcomposition. values maybe used to makeinferences and regularization may be emof complexdemodulation, nonlinearregression willfollow. Somedetailson thesetechniques ployedin thisconnection. thevibratory motionoftheEarth As is the case withmanynaturalsystems, maybe describedby a systemofequationsoftheform dY(t) dt (1) = AY(t) + X(t), withX(.) a (vector-valued) input.In thecase thattheinputis b8(t), with8(.) to the earthquakeshock)and initial the Dirac delta function (corresponding as conditions Y(O-) = 0, thegeneralsolutionof(1) maybe written Y(t) = exp{At}b = Eiexp{t1t}uj i , t > 0, wheret,uj are the (assumeddistinct)latentsof the matrixA. The spectrum oftheEarthas a body.Focusingon is discretebecauseofthefiniteness occurring ofY(t) and assumingthepresenceofnoise,onehas one ofthe coordinates (2) Y(t) = Eakexp{ k I3kt)CoS(Ykt + Sk) + E(t), with - /k and yktherealand imaginary partsofthepj and ?(*) the noise. This of the seriesY(t) in the model may be checkedby complexdemodulation Providedthe fromtheperiodogram. offrequencies neighborhood Yk,as estimated is not too great,a singlecosinusoidshouldbe bandwidthof the demodulation SEISMOLOGY AND NEUROPHYSIOLOGY 5 1960 Chilean Earthquake 0 L0 0 0 20 40 60 80 timeelapsed (hours) FIG. 1. in the Record of the Chilean great earthquake ofMay 22, 1960, as recordedby the tiltmeter Grotta Gigante at Trieste. The tides have beenpartially removed. included,thelogamplitude shouldfallofflinearly withtimeand thephaseangle at (5). shouldbe approximately constant.Detailsare givenlater,specifically recordedat Triesteofthe 1960Chilean Figure1 is a plotoftheseismogram thetides.Detailsre the data and the greatearthquakeafterpartiallyremoving in be tidal removalprocedure may found Bolt and Marussi(1962).Figure2, a ofthisdata,suggeststhe plotofthelower-frequency portionoftheperiodogram Periodogram- Chilean Data co 0 a) C) co 0 2 4 6 8 frequency(cycles/hour) FIG. 2. The periodogramof the data of Figure1 based on 2548 data values. Onlyordinates correspondingto frequenciesless than 8 cycleslh have beengraphed. The y-axis is loganithmic. D. R. BRILLINGER 6 presenceof a varietyof periodiccomponents. The periodogram of a stretchof time-series valuesY(t), t = 0,..., T - 1,is definedas follows.Set T-1 (3) dT(X) = , Y(t)expt - iXt}, t=o - x < X < oc. X is defined Then theperiodogram at frequency as (4) Iyy(X\)= (2ZrTT)-IdyT(X)12. For data fromthemodel(2), IT (X) maybe expectedto showpeaksforX near the Yk* The basic ideas ofcomplexdemodulation are frequency isolationby narrowband filtering to focuson a singletermin expression (2), followed by frequency translationto slow the oscillationsdown.The specificsteps are: (i) Y(t) -* Y(t)exp{iXt}(modulation), followed by (ii) localsmoothing in t of Y(t)exp{ixt} to obtain Y(t, X), the complexdemodulateat frequency X. In the case that Y( t) = a exp -/3t}cos(yt+ 8), onehas Y(t, A) =1ae8 ePtei(XY-)t, forA neary 0 O, otherwise. Hence loglY(t,X)I = log(a/2) - /Btand arg{Y(t,A)) = 8 + (A - y)t. Plots of these quantitiesversust providecheckson modeladequacyand providepreliminaryestimatesof parameters. Figures3 and 4 presentsuch plots forthe 3.885and 5.6775cycles/h.These frequencies Chileandata at two frequencies, weredetermined thelocationsofpeaksin theperiodogram, bynoticing settingA and thenin somecasesemploying a nearbyAto get equal to them,demodulating in theamplitudeplotcan a morenearlyhorizontal phaseplot.The fluctuations or to splitpeaks be due to noise,to leakagefromotherfrequency components among other things.The rate of decay / is foundto generallyvary with in thepresentseismological situation.ResultsfortheChileandata for frequency a varietyoffrequencies maybe foundin Bolt and Brillinger (1979). fromthecomplexdemodulate for The parameters couldbe estimated pictures, moreeffective lines.It is generally to proceedvia example,by fitting regression This has the further nonlinearregression. advantageof providingestimated one has a model standarderrors.Suppose Y(t) = S(t; 6) + E(t) 0 and e(*) a noiseseries. withS(.) knownup to thefinite-dimensional parameter In the presentcase, S(t) = aexp{-f3t}cos(yt+ 8) and 0 = {a, /3,y,8). For the next step, it is convenientto take Xi = 2'Tj/T and to write Yj = dT(Aj), Ej = dT(Aj) and Sj(6) = dsT(Aj).One willestimateO byminimizing j(6) ()12 E jin J forJ a rangeofsubscripts withXAnear y.The logicofthisis as follows.There forempirical Fouriertransforms are a varietyofcentrallimittheorems [see,for and example,Brillinger (1983)].Supposethatthe noiseseriesE(-) is stationary SEISMOLOGY AND NEUROPHYSIOLOGY 7 at 3.885 cycles/hour Log Amplitude C) LO O S LO N0 0 10 20 30 40 50 60 70 80 60 70 80 time elapsed (hours) Phase Angle LO 0 10 20 30 40 50 timeelapsed (hours) FIG. 3. The result of complexdemodtdtinzgthe data of Figure 1 at a frequencyof 3.885 cycleslh. The upper graph gives the logarithmof the runningamplttude.The lowergraph gives the running phase. The bandwidth ofthefilter eAgloyedis 0.594cycleslh. mixing with power spectrum fee,(A).Then for large T, Ej is approximately Further the variates complex normal with mean O and variance 2vrTf,,,(Xj). It followsthatthe determination of an independent. Ej, Ek are approximately estimate of 0 to minimizeexpression(6) is approxidmately the maximumlikelihood procedure. The statistical propertiesof such estimates were indicated in Bolt and Brillinger(1979) and developedin detail in Hasan (1982). For example, one findsthe asymptoticvariance of yto be proportionalto 4?Tfe -Y) 3c'2 T 8 D. R. BRILLINGER at 5.6775 cycles/hour Log Amplitude 0 co 0 10 20 30 40 50 60 70 80 60 70 80 timeelapsed (hours) Phase Angle cm cn C: CZc' 0 10 20 30 40 50 timeelapsed (hours) as forFigure3, butat thefrequency FIG. 4. Complexdemoduation of 5.6775cycles/h. having considereda limitingprocesswith,B= 4/T as T -x o. The inversecubic It comesfromthe dependenceon samplesize is on firstglancesurprising. ofthepeakswhentheyare present. narrowness is an exploratory Complexdemodulation technique.Hence one has to be of employing it at frequencies of "false"peaks.In consciousof the possibility practice,it is foundthatthe nearnessof the phase plot to a straightlineis a ofthepresenceofa periodiccomponent. highlysensitiveindicator Earlier in the paper,it was noted that progressin seismologyshows a to-and-fro betweennewknowledge ofthestructure ofwavesand newknowledge of the Earth. This occursin the case of freeoscillations. Supposeone has an initialmodelforthe Earthin termsof somephysicalparameters, e.g.,expres- SEISMOLOGY AND NEUROPHYSIOLOGY 9 EarthModel CAL8 .~~~~J.. ~~-S-wavevelocit * P-wavevelocity CJ I I I o t 0 I I 1000 2000 i t I~~~~~~~~~~~~~~~~~ 3000 t 4000 t 5000 6000 7000 depth(km) FIG. 5. The CAL 8 Earth model. The curvesgive theassumed density(grams per cubic centimeter), P-wave velocity(kilometersper second) and S-wave velocity(kilometersper second) as a function of depth assuming a spherical Earth. Given such a model, one can computeimpliedperiods of free oscillation. Of interestis the inverseproblem,givenperiods what is a correspondingEarth model? sionsfordensity, shearwavevelocity and compression wavevelocity as functions r denotingdepth.Figure5, of depth,say p(r), c,(r) and cp(r), respectively, based on the data in Tables 3 and 4 ofBolt (1982),showswhatis meantby an Earth model.Givensucha model,one can computethe impliedfrequencies of freeoscillationYk- How to do thisis describedin Chapter6 of Lapwoodand Usami (1981),forexample.The relationship involvedis nonlinear, butperturbationsmay be expressedlinearlyvia kemels.Specifically, supposeone perturbs theparameters thentheperturbation byamountsAp,Acs and Acp,respectively, ofthe frequency ofthe kthfreeoscillation is givenby AYk jAk(r) Ap(r) dr + jRBk(r) Acs(r) dr + Ck(r) Acp(r) dr, for kemels Ak* Bk and Ck. This expressionis said to lay out the "direct problem":Given Ap,Ac8 and Acp findAyk.Now supposea greatearthquake 10 D. R. BRILLINGER occurs.Then new estimatesof the frequencies Yk are available.One has the "inverseproblem":GiventheobservedAyk,findAp,Acs and Acp. Becausethe new frequenciesare just estimates,one seeks a model only approximately achievingthem.It seemsworthsettingoutthetypeofprobleminvolvedherein a specificnotation.Let Y and () denotenormedspaces.Let X denotea mapfrom e to Y, Y = X0. The valuesY and X are given,a value for0 is desired.Let a denotea scalar.Some,basicallysimilar, methodsforselecting a 0 currently being employedinclude:(a) regularization, choose0 to minimizeIIY - X0112 + allOll2; < a to minimize (b) sieve,choose0 subjectto 11011 IIY- X011; (c) residual,choose 0 subjectto IIY - X0ll< a to minimize11011. A characteristic of the solutions obtainedis that one has to be contentwiththe estimationof some formof averageof the unknown0. Chapter12 of Aki and Richards(1980) containsa discussionofinverseproblems in geophysics. A characteristic thatdistinguishes the presentEarthmodelproblem, fromtheusualinverseproblems, is thatthere are discontinuities presentin the model-corresponding to the Earth'slayers. The aboveperturbation approachofa nonlinear problem to a linearonehas been employedby geophysicists formanyyears;see Jeffreys and Bullen(1940),for example. to thestudyoffreeoscillations Severalotherreferences maybe noted.Hansen of Bolt and Brillinger (1982) extendstheprocedure (1979)to handlethe case of fit.Dahlen (1982) severaleigenfrequencies presentin the nonlinearregression sets down the asymptotic resultsforthe case of tapereddata, that is, when convergence factorshave beenintroduced intothe Fouriertransform computavia bispectral tions.Zadroand Caputo(1968)lookfornonlinearities analysis. 3. Estimation of fault-planeparameters. That thereexistsa see-saw betweenthe studyoftheEarth'sstructure and thestudyofearthquakesources was pointedout earlier.In thissectionit will be indicatedhow a (nonlinear) ofthesource probitanalysismaybe employedto estimatebasiccharacteristics ofan earthquake. An importantquantityread offthe seismictrace of an earthquakeat a is the signof the increment at the arrivalof the first particularobservatory to whether theinitialmotionis a energyfromthe event.This signcorresponds compressionor a dilation.In many cases, followingthe observationof an earthquakeat a numberof stations,if the observedsignsof firstmotionare plottedon a map centeredat the epicenterof the eventa (radiation)pattern results.Figure6, takenfromBrilinger,Udias and Bolt (1980),providessucha plotforone oftheaftershocks (event4) oftheGoodFriday1964Alaskanevent. the due to the locationsof the particularstationsrecording (Unfortunately, event,this figuredoes not providea particularly good exampleof the ideal Werethe stationswell radiationpattem,but the data wereofspecialinterest. one wouldsee mainlysolid circlesin two scattered,in an ideal circumstance oppositequadrantsand mainlyopencirclesin theothertwoquadrants.In this willbe case onlytwo ofthefourquadrantshavebeencovered.The implication thatone oftheplaneswillbe poorlydetermined.) Following Byerly(1926),plots such as this have been employedto leam about the source.Beforedescribing SEISMOLOGY AND NEUROPHYSIOLOGY 11 N 0 0~~~~ 0~~~~ ~~~/ \ 0 0o FIG. 6. The P-wavefirst-motion data fortheearthquakeof theAlaskansequencethattookplace March 30,1964 at 0200. Thesolidcirclesrefertocompressions, i.e., firstmotionupward,theopen circlestodilations,i.e., first motion downward. [Reproduced withpermission fromBriUlinger, Udias and Bolt (1980).] what may be learned,somedetailsof the earthquakeprocesswillbe set down. The usual assumption aredue to (theelasticreboundtheory)is thatearthquakes A crackinitiatesat a pointand (in thecase ofpureslip)spreadsout to faulting. forma faultplane.As the crackpassesa givenpoint,slip takesplace (on the in a stressdropand the radiationof seismicwaves.The faultplane) resulting radiated(P-) wavesmaybe shownto havea quadrantalpatternwithone ofthe axes paralleland the otherperpendicular to the faultplane of the event.It and thisis whatByerly(1926)contributed, follows, thatthedata maybe usedto the estimate fault-plane orientation. Havingan estimateofthe faultplaneand the directionof motionon thatplane is important to geologyand geophysics. Researchersseek to tie togethersurfaceand subsurfacefeatures,to consider regionalstressdirections and to use theresultsto confirm and extendthetheory ofplate tectonics.The resultscan be crucialto seismicriskcomputations. and this has continuedto generallybe the Byerlyproceededgraphically workingapproach.However,the resultsso obtainedare subjective,have no attachedmeasureofuncertainty and maynotbe easilycombined withestimates derivedfromothereventsat thesamesite. 12 D. R. BRILLINGER The problemmaybe approachedin formalstatisticalfashionas follows.The data availableconsistof hypocenter of earthquake, locationsof observatories, directions ofobservedfirstmotions(compressions or dilations)at theobservatoriesand a storeofknowledge the Earth'sstructure concerning [velocitymodels as givenin Jeffreys and Bullen(1940),forexample].It mayfurther be argued noiseis approximately thatseismographic Gaussian[see Haubrich(1965)].Let a faultplanebe describedby threeangles(0T' (kT' Op). Let Aij(OT, 4 T, Op) denote the theoretical forthewaveamplitude on thefocalsphereforeventi expression at stationj. This expression maybe foundin Brillinger, Udias and Bolt (1980). (The focalsphereis a "little"sphereof unitradiusaroundthe hypocenter. In one has to tracethe ray fromthe carryingout the amplitudecomputation, hypocenterto the observatory throughthe focalsphere.)Let Yij denotethe realizedamplitudeof the seismogram at the onsetof the event.Then one can write Yu = aijAij + Eyj,with aij a scale factorand -,j normalmean 0 and vaxiancea?L variate.Here aij reflects the attenuation thesignalexperiences in travelingfromthe sourceto the observing station,whereasEcj represents noise caused by disturbances unrelatedto the earthquakeof concern.Let yj = 1 if = > 0 and It followsthat O otherwise. Yij Prob{yij = 1} = Prob{Yij > 0) = (pijAij), writingpij = aj/vaij,forthis signal-to-noise ratio.The modelmay be further expandedby includinga term-yijto allowforreaderand recorder errors, now writing (7) Prob{yij = 1} = Yij + (I 2yij)4D(pijAij)A to y and a small(hencep large)and imprecise to y near Precisedata correspond 0.5 or p near0. The model is seen to take the formof a nonlinearprobit(witha term-y An exampleofa corresponding to "naturalmortality"). likelihood corresponding is providedby (8) HIj(piAij) ij(1 - 0(piA 1-Yi assumingp to dependon eventalone and y = 0. One can now proceedto likelihood. estimatethe unknown parameters OT' OT' (P, pi bymaximum Figure6 includesthe fittedplanes forthe case of event4 of the Alaska thelikelihood werecomputed estimates sequence.Theseparticular restricting (8) a y termas in (7). ofeventi = 4 and including to the observations Udiasand Bolt(1980), It is criticalto assessthefitofanymodel.In Brillinger, functions. and estimatedprobability thetheoretical thiswas doneby comparing Figure7 is based on a pooledanalysisofsome16 oftheAlaskanevents(labeled It has beenassumedthatthepi are thatseemedto go together. by i previously) thatthe all equal in thefitstudied.The figure providestheempirical probability The ofamplitude. as a function observedfirstmotionagreeswiththetheoretical fittedvalues z = p1Aijhavebeengroupedintocellsofwidth0.1 in theanalysis. SEISMOLOGY AND NEUROPHYSIOLOGY 13 EmpiricalProbability ofCorrectFirstMotion 0 _-. 0 c'J 0~~~~~~~~~~~~ -4 -2 0 2 4 z FIG. 7. A plot of thestatistic(9) ofSection3 and ?(z) forthedata of 16 eventsoftheAlaskan tothevalues ofthemodel(7). Herez refers sequenceof 1964. Theplotis meanttoassessthevalidity pAij. Whatis plottedare ?(z) and (9) (i yj)l sgn Yij = sgnAij, Z - h < Aij<Z + h/{(i, j)z - h < Aij < Z +h, for h = 0.5. Here A refersto A(OT, (PT,0p) and # refersto the countof the numberof elementsin theset.The fitseemsadequate. The resultsof further ofthistypemaybe foundin Brillinger, computations Udias and Bolt (1980)and Buforn(1982).The maximization VA09Aof program the Harwellsubroutine see Hopper(1980),provedeffective in determinlibrary, ingthemaximum values.The estimates likelihood were,however, nonunique and in somecasesofsmalldata sets. poorlydetermined An important ofsuchanalysesis to fonnclusters oflikefault-plane by-product solutionsforeventsin thesameregion,in orderto getat motionsoccurring on the same faultplane; see Udias, Munoz and Buforn(1985),forexample.The maximumlikelihoodstandarderrorsare usefulin thisconnection. The practical oftheworkjust reported implication is thatfirstmotionsforlargecollections of events may be handledroutinelyand that geophysicalconjecturesmay be checkedformally. The finalfault-plane solutionmay be plottedin traditional fashionallowingexamination of the data fordifficulties. What remainsis for morerealisticseismicsourcemodelsthantheone treatedin thepaperslistedto D. R. BRILLINGER 14 An elementary to thesubjectmatterofconcem reference be fittedstatistically. hereis Boore(1977). quesand difficult 4. Quantificationofearthquakes. Oneoftheimportant is how to measurethe "size" of an earthquake.Size is an tionsof seismology makesuse of in attemptsto deal with essentialfeaturethat a seismologist ofconcem.Specifithebasicphenomena earthquakehazardsand to understand is not only interestedin estimatingthe directionof cally, the seismologist that in theoveralldeformation interested movement at thesource,he is further took place and the amountof energythat was released.Amongthe physical fora givenearthquakeare theseismicmoment(a measure quantitiesofinterest of the seismicenergyreleasedfromthe entirefault) and the stressdrop betweentheinitialand finalstress.) (difference haverelatedtheseismic For a varietyofseismicsourcemodels,seismologists IS(X)I,the oftheamplitudespectrum momentand stressdropto characteristics is ofthesignal.Supposethattheseismogram modulusofthe Fouriertransform writtenas Y(t) = S(t; 0) + E(t), where s(.) is the signal, ( is an unknownparameterand E(*) is a noise of s(t; 0), thenwhatis disturbance.If S(X; 0) denotesthe Fouriertransform formof IS(X,0)1. A reasonfor given,fromthe sourcemodel,is the functional is phase information workingin the Fourierdomainhereis that distracting include measurements) eliminated.Commonforms(fordisplacement IS(X; 0)1= a/1/ + (X/Xo)fand a/{1 + (X/Xo) } ofsuchfunctional with0 = {a, /B, X0). The seminalpaperon thedetermination to the"size" oftheearthquake oftheirparameters formsand on therelationship is Brune(1970/1971).Estimatesoftheseismicmomentand stressdropmaybe relate onceestimates ofa and X0areavailable.That theparameters determined in form the discussion will seen for a functional to size and duration be particular graphithat follows.The empirical practicehas beento estimatetheunknowns of the Fourier of the modulus of the empirical amplitude cally froma plot was suggestedin Brillinger formalprocedure transform Idy(X)I.The following and Ihaka (1982). of Idy(X)I may be evaluatedin the case of The asymptoticdistribution in Section2. stationarye usinga centrallimittheoremof the typementioned is foundto dependon IS(X; 0)I and fee(X)alone. The asymptoticdistribution Hence one needsan expression onlyforthemodulusofS, and as statedabove, provides.Next,withthe modelY(t)= this is what the seismologist generally s(t; 0) + e(t) and small noise, idy(X)I= IS(X;0)1+ (dT(X) + dT(-X))/2 + of whendeviations on ISI. However, showingvariationaroundISI notdepending of IdTi froma finalfittedformare plottedversusthefittedvalues,dependence the erroron ISI is apparent.An exampleis providedin Figure8. This is the SEISMOLOGY AND NEUROPHYSIOLOGY 15 TaiwanEvent- TransverseShearWave- 29 January 6.7 1981,Magnitude 8.0 8.5 9.0 9.5 10.0 10.5 11.0 seconds ResidualPlot 5 0 500 1000 1500 2000 fitted values FIG. 8. The uppergraph gives thetransverseS-wave componentof thevibrationsof theJanuary 29, 1981, magnitude 6.7, Taiwan earthquake as recordedby the central accelerometersof the Smart 1 array. The array is approximately30 km northwestof the epicenterof the event. The lowergraph plots the differencesbetweenthe amplitudesof theFourier transformof thedata and corresponding (final) fittedvalues. The data stretchconsistedof 256 points. resultof computations foran earthquakeof magnitude6.7 that occurredin Taiwan on January29, 1981.The data wererecordedby one oftheinstruments oftheSmart1 array;see Bolt,Tsai,Yeh and Hsu (1982).The uppergraphofthe S-waveportionoftherecordedaccelerations. figureprovidesthetransverse The lowergraphprovidesthedeviations plotjust referred to. This plotsuggeststhat the noiseis in part"signalgenerated" in thiscase. Thereare variousphysical phenomenathat can lead to signal-generated noise.These includemultipath reflection and scattering. transmission, The following is an exampleof a model thatincludessignal-generated noise: (10) Y(t) = s(t) + E(YkS(t k - Tk) + akSH(t - Tk)) + e(t), withthe Tk timedelays,withSH the Hilberttransform of s and withYk, 8k 16 D. R. BRILLINGER reflecting the vagariesofthetransmission process.[The inclusionoftheHilbert is discussed, ofphaseshifts. The Hilberttransform transform allowsthepresence forexample,in Brillinger (1975a),page32].Withthe Yk, (k' Tk randomand after thedistribution evaluatingthe largesamplevariance,oneis led to approximate of YJ= dykXj)by a complexnormalwithmean S(Xj; 0) and variancerj = 2iTT(p2/S(X1; 0)12 + a2), wherenow E has been assumedto be whitenoise(of variancea2), and alsoit is assumedthatEyk,E 3k = 0 and thattheprocessTkis of signal-generated Poisson.The ratiop2/a2 measuresthe relativeimportance noise.This varianceis seento dependon the "signal"throughISI and leads to Taiwan Event- Amplitude Spectrum 0 O 0 U) 0 1 0.5 0.1 5 50 10 frequency(cycles/second) FittedPulse 0 0 0 \ O c'J CO 0.0 0.5 1.0 1.5 2.0 2.5 3.0 SeCOndS theamplitudes FIG. 9. The uppergraphprovides transformsoftheTaiwandata of oftheFourietFiure 8 and the correspondinzgfitted expectedvalues as cornputedfor the model of Section 4. Both scales of theplot are logarithmic.The lowergraph provides thefittedpulse s( t). SEISMOLOGY AND NEUROPHYSIOLOGY 17 wedgingof the typepresentin Figure8. One can proceedto estimate0 by derivingthe marginallikelihoodbased on the IYjl. This likelihoodmay be evaluatedand foundto be FH(exp(- ilr2 S IO r;_____ The uppergraphofFigure9 shows Besselfunction. whereIo denotesa modified a fitof the model IS(X)l = all/(l + (X/Xo)4)to the data of Figure8. This functionalformwas settledon afterthe degreeof fitof two moreelementary formswas examined. Details may be foundin Ihaka (1985). We remarkthat this model fit to a timedomainpulses(t) = aXop(X0t), where corresponds p(t) = [sin-i - tsin( + e- t/ for t > 0 and p(t) = 0 otherwise. The expressions(t) = aXop(Xot) indicates how Xo corresponds to the durationof the eventand how a corre(inversely) spondsto size. The lowergraphof Figure9 providesa plotof the fittedpulse. Once estimatesof a, AOare at hand,thesemaybe converted to estimatesofthe seismicmomentand stressdrop via theoreticalrelationships developedby geophysicists. The maximumlikelihoodfitof the modelwas carriedout by a computer programwritten by Ihaka.Thisprogram also generates standarderrorestimates and standardized residuals.Theselatermaybe usedto assessthegoodnessoffit ofthe model.Figure10 providesa plotofthestandardized residualsagainstthe Standardized ResidualPlot CM 0 1000 1500 2000 fitted vaues FIG.10. A standardized residualplot,basedon themodel(10),corresponding tothelowergraphof Figure8. The differences betweentheamplitudes of theFouriertransform valuesand theirfitted expectedvalues have been dividedby theirfittedstandarddeviationsto obtainstandardized residuals. 18 D. R. BRILLINGER fittedvalues of the sameformatas the residualplotof Figure8. The wedging corresponding to signal-generated noisein the laterplot is no longerpresent; however,thereis a definite suggestion that the fitmightbe improvedin the regionwherethe signalhas low amplitude.Luckily,thisis the regionof least It awaitsfutureanalysis.It mightbe handledbyallowingtheseries importance. to E(t) have a nonconstant spectrum. 5. Arraydata. Todayit wouldbe a strangethingindeedforan earthquake In fact,fromthe veryearliestdays, to be recordedon just one seismometer. readingsof the sameeventat geographically scatteredobservatories have been made use of. Since the 1960s,seismometers have beendeliberately arrangedin geometric designsoverdistancesof the orderof milesto hundredsof milesin oftraditional and sometimes orderto allowextraction information ellicitation of newinformation. An importantuse has been the estimationof the directionfromwhicha withwhichit is moving. and thevelocity Onemannerin seismicsignalis arriving ofestimatesoffrequency-wavenumber whichthisis doneis bythecomputation as follows. spectra.The procedure maybe described Supposeonehas arraydata; Y(xj, yj, t), j = 0,..., J and t = 0,..., T - 1. Here (xj, yj) denotes the coordi- natesofthelocationofthejth sensor.The frequency-wavenumber periodogram ofthisdata is givenby (11) 2 |EE j t } , Y(xj,yj,t)expt ii(txj+ vyj+ XAt) < AIv,A.< xo. is thefollowing. A motivation forthisdefinition Supposeone has a planewave = + + + Y(x, y,t) p cos(ax fly yt 8) oftemporalfrequency y and wavenumber willhave a peak near(a, 3,y). (Incidentaly, K = (a, 1). Then the periodogram azimuth withapparent thiswaveis moving velocity y/ja2 + /32 from givenby ofarraydatais givenbyFigure11.Whatis plotted tan0 = /3/a.)Anexample oftheSmart1 arraylocatedin ofnineoftheseismometers are thelocations ofthetracesusedin thecomputations. Taiwan.Alsoplottedaretheportions to thevertical P-wavepart,oftheJanuary Thesetracescorrespond 29,1981 earthquake. (Theinitialnear-flat partis thenoise,savedina buffer, justbefore was30 km ofthisearthquake theonsetofthewave.)Theestimated epicenter ofthefrequency-waveofthearray. southeast Figure12givesa central portion X viaformula number forthisdata,as computed periodogram, (11),at frequency 1.944was pickedon to 1.944cycles/s.(The temporalfrequency corresponding oftheindividual Thereis seen thebasisofa times-series analysis seismograms.) at an azimuth thatturnsoutto to be a largepeakin thesoutheast quadrant, tothatoftheepicenter oftheevent.Theradialdistance correspond corresponds ofP-waves. to thevelocity to employ, withthistypeofdatahaveoften Seismologists working preferred or "Capon"statistic whattheycall,the"high-resolution" [see Capon(1969)] shows statistic insteadof theperiodogram (11).The high-resolution typically Beforedefining moredramatic peaksthantheperiodogram. it, we introduce 19 SEISMOLOGY AND NEUROPHYSIOLOGY Taiwan Arrayand Eventof29 January1981 0 0 C') 0 0 0 E .1 ?A E0 C> o0 0 o) C~)0 o -3000 -2000 I I I -1000 0 1000 2000 3000 distancefromarraycenter(meters) bulletdenotes sensor location as recorded at nineof FIG. 11. The verticalP-waveportionoftheJanuary29 Taiwanearthquake " areplottedat thephysicallocationsofthesensors. thesensorsof theSmart1 array.The " bullets fromtheeventhad beensavedin a buffer. Noise immediately precedingtheamval ofenergy somenotation.Let Y(t) denotethej-vector[Y(xj, yj,t)]. Set T-1 Yk= T- Y(t)exp t=o -I L2k T for k=0,2,.... Furtherlet B=[exp{-i(pxj +vyy)}]. If X=22d/T, 1 an to IBTYII2.Nextdefine thentheperiodogram (11) is proportional integer, M = EykYk, statisticat withthe sum overk with2rk/T near X. Now the high-resolution frequencyX may be definedas 1/B M-'B. If Y(x, y,t) = p cos(ax + fly+ yt+ 8) + noise,thisstatisticmay be expectedto showa peak for(IL,v) near in part,in orderto be (a, fi)and X near y. This statistichas beenintroduced, able to presentthenextexample. D. R. BRILLINGER 20 Periodogram Taiwan Event Frequency-Wavenumber c'LO C\J LO) -0.5 -0.4 -0.3 -0.2 -0.1 0.0 0.1 0.2 0.3 0.4 0.5 wavenumber (cycles/km) FIG. 12. The frequency-wavenumber periodogram of the data of Figure 11. The time series stretchescontain 720 points. The temporalfrequencyemployedis 1.94 cycles/s. Figure13 is reproducedfromScheimerand Landers(1974). It showsthe statisticcomputedfortwo portionsof data recordedby the high-resolution a strip-mining Large ApertureSeismicArray(LASA) in Montanafollowing blast. These computations the validityof the high-resolution confirmed apof proach.The statisticforoneportionshowsa singlelargepeakin thedirection the blast. The statisticforthe following portionshowsenergyarrivingfrom variousdirections.This analysisprovidedempiricalproofof the existenceof of seismicwaves.That thisphenomenon existedhad beentheorized scattering foryears.A frequency-wavenumber theconfirmation. data analysishas provided Spectralanalysesare (too) oftenthoughtof as beingappropriateonlyfor stationarydata. As thepreceding exampleshows,the techniquemaybe highly casesas well.As a secondexamplewemention usefulin nonstationary theresults of Bolt, Tsai, Yeh and Hsu (1982). If, in fact,an earthquakeis caused by thenthe direction of thesourceof seismicenergywillbe changingas faulting, In thepapercited,Bolt, thefaultis ripping, thatis,as thefaulttipis advancing. timestretches Tsai, Yeh and Hsu presenthigh-resolution spectraforsucceeding SEISMOLOGY AND NEUROPHYSIOLOGY A~~~~ t4:1he a , 49 /~~~~~~TM 19 (mrin.: 21 14 20 sec ) wIth tim.Thehighreworkio mayhaven beenthem firsutexedfrimwaendtal measurdemen foflwna seismicdislocationmovingalonga rupturing fault. In each of the preceding two examples,frequency-wavenumber analysishas allowedresearchers to confirm thepresenceofsuspectedscientific phenomena. 6. Exploration seismology (reflection seismology). The problemof learningthe Earth's crustalstructurecan be approachedas one of system identification. The approachto be described takesadvantageofthefactthatthe Earth happens to be made up of layeredstrata.Signals,such as powerful impactsor explosions, can be deliberately inputto theEarthand theconsequent vibrationsrecordedbyan arrayofseismometers or geophones. Suchexperiments may be carriedout in a searchforgas and oil, or in a scientific studyof the generalgeologicalmakeupof a regionof interest.The resultsof theseexperimentsmaybe viewedas oneofthegrandsuccessstoriesforstatistical techniques generally, and ofleastsquaresparticularly. An unusualaspectoftheinferences made is that in manycases one gets to examinetheirvalidity,by the later D. R. BRILLINGER 22 in thissectionofthepaper,in drillingofa well.No workedexampleis presented largepart becausedata sets are hard to comeby. The materialis presented, however,because it providesa case wherea rathercompletesolution(design can be presentedand because the basic experimental throughconfirmation) in theneurophysiological case,whereso muchlessis techniqueis also employed known. propagates ofan initiatedseismicdisturbance theenergy In itssimplestform, an interface it meets When throughthe Earth witha spreadingwavefront. part back and betweengeologicalstrata,part of the energymay be reflected at the interface. in acousticimpedance continueforwarddue to the difference energyechoes.Knowledgeof subreflected The sensorsrecordthe returning ofthe ofthedepthsand anglesofinclination surfacevelocitiesallowsestimation estimaallows ofthelocationsofreflectors whereasknowledge variousreflectors, In tionofvelocities.(One notesagaina see-sawin thecollectionofknowledge.) impactswillbe repeateda numberoftimesat the same practice,the initiating is againused. locationand at pointsofa grid.The powerofaveraging If the inputsignalis takento be X(t) and if Y(t) denotesthe corresponding output,thenthe two maybe modeledas related,assuminglinearityand time invariance,by (12) Y(t) fa(t - s)X(s) ds. sinceiftheDiracdeltafunction The function a(.) is calledtheimpulseresponse, a(-) outputis Y(t) = a(t). The function 8(t) is takenas input,thentheresulting and velocitiesin the earth beneaththe sourceand evidencesthe reflectors as follows. Suppose maybe motivated The modeland itsinterpretation receiver. travels a waveis generated, a pulseis appliedat timeT. Supposein consequence back. a1 is reflected at distanced, and a proportion at velocityv1 to a reflector With X(t) = 8(t - T), then Y(t) = a, 8(t - T - 2d1/v1). (This is actuallythe that the transmitted portion naive modelforradaror sonar.)Supposefurther at distanced2 and a portionof at velocityv2to a reflector continuesdownward to by thefirstreflector back,someofwhichis transmitted its energyis reflected has theformY(t) = a18(t - T - 2dJ1v1)+ Nowtheresponse reachthereceiver. to the system a2 8(t - - 2d1/lv- 2d2/v2). This last is seen to correspond of expression(12) with impulseresponsea(t) = a,8(t - 2dJ1/j)+ a28(t 2d1/v1- 2d2/v2).One can clearlyextendthismodelto situationswithmany and reflection transmission layers,many velocitiesand manycorresponding to reflecPeaks in the functiona(t) maybe seenas corresponding coefficients. is suchan elementary interpretation tors.(It mustbe notedthatunfortunately phenomenasuch as ghost likelyto be complicatedin practiceby interfering havebeendevelopedto handlethese.)The basicsof reflections. Some techniques are discussedin Woodand Treitel(1975),Waters(1978) seismology exploration and Robinson(1983). as one ofsystemidentification; given The problemhas nowbeenformulated an estimateofthe stretchesof corresponding inputX and outputY, determine impulseresponsea(-). In the case that a pulse close to a Dirac 8 may be SEISMOLOGY AND NEUROPHYSIOLOGY 23 generatedand that the functiona(-) drops offto 0 reasonablyquickly,a resultsfromtaking convenient procedure M X(t)= E 8(t- MA) m=1 as input,and the "averageevokedresponse" 1 M M m=1 to applyingpulsesperiodically. as an estimateof a(s). This inputcorresponds to stackingand averaging. The estimatecorresponds Suppose one sets myx(t)= Y*X(t) for some convolutionoperation" ." Then from(12) one has myx(t) = fa(t - s)mxx(s) ds and one has a deconvolution (or inverse)problemto solve.Supposeone decides to seek an X(*) suchthat Ja(t - s)mxx(s) ds z a(t), In termsof Fouriertransforms, to allow elementary the left-hand processing. side heremay be expressedas Jexp{iXt}A(X)Mxx(X) dX, withMXXa Fourier transfornof mxx. Thenwhatis wantedis an X suchthat Mxx(X) - 1 on the supportof A(-). If A(X) is knownto be near0 for0 < X < Xo and forA > A1, is the"chirp"signal thena possiblefunction X(t) = cos([AO + (A - Ao) ]t)- for0 <t < T. In the seismiccase, the values of o, X, have been determined in various in radarworkduringWorldWarII [see The chirpprobeoriginated experiments. Cook and Berenfield (1967)].It maybe seento attachnearequal powerto the frequencies betweenXo and AX.In the seismiccase specialdeviceshave been developedto inputthechirpsignalto theearth.The signalis inputrepeatedly and theresultsaveraged.The responseis thenconvolved withthechirpfunction, that is, myx is formedto estimatea(.). Structurecan appear dramatically described here. duringthe cross-correlation processing In practice,subtlefurther processing is employedto handlewavefront curvaand othernaturalphenomena thatmaybe present. ture,ghostreflections 7. Other topics. Thereare otherproblemsarisingin seismology to which statisticalmethodology can be appliedfruitfully. These includeanalysisof the coda (i.e.,oftheirregular trailing partofthedisturbance), analysisofscattering, riskanalysis,nonlinearphenomena, pointprocessstudies,polarization, cepstral of earthquakesfromexplosions[see, e.g., Tjostheim analysis,discrimination 24 D. R. BRILLINGER laws,earthattenuation study,traveltimetableconstruction, (1981)],seismicity and Vere-Jones quake locationand azimuthaldependenceof characteristics. In somecasesworkhas begun. Smith(1981)discussseveraloftheseproblems. have long been serioususers of statistical 8. Discussion. Seismologists in theentry, statement makingthefollowing methods.One findsHaroldJeffreys "Seismology,statisticalmethods,"in the InternationalDictionaryof Geois as important a partof the resultas the estimate physics:"The uncertainty itself. ... An estimate without a standard error is practically meaningless." traveltimetables Hudson(1981) remarks:"The successofthe Jeffreys-Bullen use of soundstatisticalmethods." consistent was due in largepartto Jeffreys' WhenI asked mycolleagueB. A. Bolt whathe saw as the roleof statisticsin science.... The role he replied:"Seismology is largelyan inferential seismology, observaforturning procedure is to providea rigorous ofstatisticsin seismology ofthe aboutproperties statements tionson seismicwaves,etc.,intoprobabilistic (real) Earth." is characterized One maynotethatworkin seismology by massivedata sets, models, error,defining/fitting/refining inherentvariabilityand measurement needsforrobust/redescription, simulation, probabilistic designofexperiments, of inverseproblemsand combination situations, predictive sistantprocedures, in all theseconnections. Statisticshas muchto offer observations. in 1983,workhas progressed 9. Update. Sincethe lectureswerepresented Smart1 data (1985)has employed on variousofthetopicscovered.Abrahamson of the faultrupturetip. Chiu (1986) studiesthe to bettersee the movement (1986) source.Lindberg ofa movingenergy theparameters problemofestimating ofthefrequencies offree develops"optimal"tapersto employin theestimation data The approachofKitagawaand Gersch(1985)to nonstationary oscillations. The bookbyUdias,Munoz seemslikelyto proveofbroadpracticalapplicability. and Buforn(1985) goes into substantialdetail overthe formalestimationof forthevarianceof Copas (1983)setsdownan expression parameters. fault-plane a statisticlike that of (9). Brillinger (1985) developsa maximumlikelihood ofa planewavegivenarraydata. Donoho, statisticfordetectionand estimation procedureforbetter Chambersand Lamer (1986) developa robust/resistant aligningthe seismictracesof a section.Mendel(1983,1986)presentsmaximum seismollikelihoodstatespace-basedmethodsforhandlingthedata ofreflection to ogy.Shumwayand Der (1985)indicatehowtheEM methodmaybe employed of seismograms deconvolvepulseshiddenin seismictraces.The nongaussianity obtained in reflection seismologyis being taken specificadvantageof; see Giannakisand Mendel(1986).The techniquesof Donoho (1981) and Lii and case. The Rosenblatt(1982) seemboundto proveusefulin the seismological thesis,Ihaka (1985),has been completed.Ogata [e.g.,Ogata (1983) and Ogata analysesof and Katsura (1986)] has carriedout a varietyof likelihood-based on havebegunworking earthquaketimesas a pointprocess.Manystatisticians mention O'Sullivan(1986). We specifically statisticalaspectsofinverseproblems. SEISMOLOGY AND NEUROPHYSIOLOGY 25 will go in the One can speculateon wherethe fieldof statisticalseismology comingyears.It seemsclearthattherewillbe muchconcernwithnon-Gaussian data and analysiswill noise and signals,that vector-valued spatial-temporal becomethe norm,thatlarge-scale conceptualmodelswillbe set downand that strengthin there will be a varietyof techniquesdevelopedfor borrowing situationswithscantydata,e.g.,riskestimation. III. Neurophysiology endeavorto buildstructural modernbiometry is the interdisciplinary stochasticmodelsofbiological phenomena. J.Neyman(1974) is the branchof science 10. The field and its goals. Neurophysiology concernedwith how the elementsof the nervoussystemfunctionand work is seen to involvechemicalmechanisms, electrical together.The functioning The studiesextendfromthemovements mechanisms and physicalarrangement. ofthebrain. to themassbehaviorofthecomponents ofindividualions,through rangeto theheroic:howto explainthingslike The goalsofneurophysiologists behavior.At a less ambitious memory,emotion,learning,sleep,expectation, withhowa singlenervecellrespondsto level,neurophysiologists are concerned oftheenvironment. information and changeswithalterations stimuli,transmits and structural unitofthenervoussystem. The neuronis boththefunctional The elementsof the The brain is a multiprocessor of dramaticcomplexity. fromthosein theseismiccase,in thatthey nervoussystemmaybe said to differ apparentlyhave purposes. withresearchers collecting variedand extenThe fieldis largelyexperimental sive data sets. The data includephotographs made via electronmicroscopes, fluctuating voltagesand currentlevels withinsinglenervecells and finally electroencephalograms (thebrain'selectricalpotentialat pointsnearthe skull.) The studiesare sometimes butoftencomplexexperimental simplyobservational, designsare employed. Importanttechniquesthat are made use of includestainingto identify of microelectrodes within individualneurons,insertion to makemeasurements ofwholesuitesofresponsesto a stimulusof individualcells and the averaging interestin orderto reducewhat can be the dominanteffectsof noise.Many are computer controlled and computer processed. experiments Discoveriesmade by neuroscientists includethe following. Nervecells communicatewitheach otherin botha chemicaland electricalfashion, the voltage pulse that travelsalonga neuron'soutputfiberis of nearconstantshape and thereare a broad varietyof nonlinearphenomenathat occur.A numberof verifiablephysical laws and effectivedeterministic models (such as the Hodgkins-Huxley equations)havebeensetdown.Muchinsighthas beengained, especiallyat thelevelofsmallgroupsofneurons.At thelevelofthebrainitself, Herethebrainis viewedas a blackbox knowledgeis mainlyphenomenological. and studied by systemidentification techniques.Whateverthe approach, discoverieshave been made leadingto lifesavingand life-improving clinical diagnoses. 26 D. R. BRILLINGER Statisticalmethodsenteredwiththe quantification of the field.No single individualscientistseemsto have had a dominating effect, rathertherehave beenmanycontributing workers-researchers concerned withelectroencephalograms(EEGs) and researchers concernedwith small collectionsof neurons. Statisticalmethodsenteredboth because of highnoise levelsand because a varietyofphenomena seemedto be inherently stochastic. Evidenceforthislast is presentedin Burns(1968)and Holden(1976).Pertinent bookson neurophysiologyincludeFreeman(1975),Aidley(1978)and Segundo(1984).Generalreviews ofstatisticalmodelsand methodsin neurophysiology are givenin Moore,Perkel and Segundo(1966)forthecasesofsingleneuronsand ofsmallgroupsofneurons and by Glaserand Ruchkin(1976)forEEGs. Statisticalmethodsforclassificationand pattemrecognition, forhandlingartifacts and fordata summarization are in commonuse. is one ofthemostactivebranchesofscience.The physiological Neurobiology phenomenawithwhichit is concerned arefundamental and in mostcasesbarely understood. 11. Neuronal signaling. One oftheimportant meansby whichnervecells communicate is via spiketrains.The inlaysat the tops of the threegraphsof Figure14 giveexamplesofspiketimesrepresentative ofthreedifferent sortsof neuronalbehavior;pacemaker occursin bursts) (near-periodic), bursting (activity and bursting withacceleration withinbursts). (offiring Suppose that a neuronfiresat timesTn, n = 0, + 1, + 2,.... A convenient ofits temporalbehavioris providedbywriting formalrepresentation Y(t) = n a(t Tn with8(.) theDirac deltafunction. Thisrepresentation leadsto resultsanalogous to ordinary time-series resultsin manycases.In thecase thatthe T, arerandom, one has a stochasticpointprocess{Tn}. A principaldescriptor ofa pointprocess is providedby its ratefunction. This is givenby lim Prob{point in (t, t + h }h, h as h tendsto 0. In the stationary of the case,wherethe stochasticproperties is constantand so processdo not dependon the timeorigin,the ratefunction onlycrudelyusefulthen. is an important The autointensity in thestationary function case. parameter It is definedas lim Prob{point in (t, t + h]lpoint at O}/h, h as h tendsto 0. It is a pointprocessanalogof the autocovariance of function time-series be for analysisin a generalsense.Thisparameter may used, example, to describethe behaviorof spontaneously firingneurons.Figure14 presents examplesforthreecases. In the firstcase, the neuronis firingapproximately The (estimateof)theautointensity is seento oscillate(withperiod periodically. 27 SEISMOLOGY AND NEUROPHYSIOLOGY - L10 as Pacemaker Function Autointensity 0 5 10 15 20 30 40 lag(seconds) co - L10 Burstinrg Function Autointensity 0 10 20 lag (seconds) u) * _ . . , . ~~~~~~~~~~~~~~~~~. . . . . . . . .. . . ... . . ..... . . ............. cM Thecellis FIG. 14. Pointprocessdata (spiketrain)fromthenervecellL10 ofAplysiacalifornica. of fashions.The inlaysat thetopsofthethreegraphsgivebriefstretches behavingin threedifferent Thefunctions thedata (butnotonIthesametimescalesas theautointensities). plottedare estimates of the autointensityfunctionsbased on 1538, 1019, 1631 spikes, respectively. 28 D. R. BRILLINGER equal to the intervalbetweenthe points).In the secondcase, the neuronis evidencing activityin bursts.The probability thatthe neuronfiresagainsoon afterit has firedis high.In the thirdcase,the neuronis also firing in bursts; however,now thereis structure withinthe bursts,the rateof firing is seen to increasetherein.The burstshereare at regularintervals. The autointensity functions have been estimated,forthis figure,by the statistic ITn -Tm - ti < h/2}/Nh, with N the total numberof points,withh a smallbinwidthand witht lag. (Here # refersto the countof the numberof pointsin the set.) The data analyzedare forthe cell L10 of Aplysiacalifornica, the sea hare.They were collectedand previously analyzedby Bryant,Marcosand Segundo(1973).The experimental procedures and detailsof the data preparation may be foundin thatreference. A questionthat arisesin the studyof smallnetworks of neuronsis which neuronsare interacting withwhich?In otherwords,whichspike trainsare associatedwithwhichothers?A usefulparameter to employin thestudyofsuch questionsis providedby the cross-intensity function. Supposingone has spike trainsnamedM and N, thenthecross-intensity function of N givenM at lag t is definedas limProb{N pointin (t, t + h]M pointat 0)/h, h as h tendsto 0. If the M spiketrainconsistsof pointsUrn and the N trainof pointsT thenthiscross-intensity maybe estimatedby # IT,,-,rn - t < h/2)/Mh, with M denotingthe numberof M pointsin the data set, with h a small binwidthand with t lag. Figure 15 presentsthree examplesof estimated functions. The firstgraphrefers cross-intensity to data fromcellsL3 and L10 of Aplysiacalifornica. The behaviorexhibited hereis thatofnegativeassociation; L1O's firingis inhibiting the firing of L3 (forapproximately 0.5 s). If one asks whetherthe values at negativelags differ fromthe level of no-association by morethansamplingfluctuations, one findstheydo not.This resultis consistent withthe cell L10 drivingthecell L3. The middlegraphcorresponds to positive association.It is fora cellin therightvisceropleural connective (RVP) and cell R15. The firstcell tendsto excitethe secondforabout0.25 s. The finalgraph a morecomplicated situation.Thesedata setswerealso represents (polyphasic) detailsmaybe analyzedin Bryant,Marcosand Segundo(1973),wherefurther weredeveloped found.The approximate ofsuchstatistics distributions sampling in Brillinger (1975b).It was found,forexample,thatit couldbe moreconvenient to graphthe squarerootoftheestimatein somecircumstances. The cross-intensity function, beinga pointprocessanalogofcovariance, may be expectedto be an inadequatemeasureof relationship (as usual,correlation statisticaldata,it is usual does notimplycausation).In thecase ofelementary SEISMOLOGY AND NEUROPHYSIOLOGY 29 - L3givenL10 Function Crossintensity 0 -2.5 -2.0 -1.5 -1.0 .0.5 0.0 0.5 1.0 1.5 2.0 2.5 1.5 2.0 2.5 lag (seconds) - R15givenRVP Function Crossintensity -2.5 -2.0 -1.0 -1.5 -0.5 0.0 0.5 1.0 lag (seconds) - L3givenLi0 Function Crossintensity -20 -10 0 10 20 lag (seconds) FIG.15. Estimatesofthecross-intensity functions forthree pairsofAplysianeurons.Theestimates are basedon (1746,302),(1101,288),(1019,993) spikesin thepairs oftrains, respectively. 30 D. R. BRILLINGER to turnto regression In thepointprocesscase it is possible as a bettertechnique. to carryout regression-type analyses.For example,one may fitthe following fonnofmodel: lim Prob{N spikein (t, t + h)IM spike train}/h = i + Ea(t h - am), m to as the as h tendsto 0. The function a(t) appearingin thismodelis referred Set impulseresponse.Thismodelmaybe fitas follows. M dT()= E m=1 exp{-iXam,, Coherence - LI 0 withL3 co o L\. C; *\7- 0 1 _> ,\--- 3 2 4 A 5 frequency (cycles/secnd) ImpulseResponse cm ICM -0 -10 - 54 -5 A 10 0 0 5 10 lag(seconds) and impulseresponseforthedata oftheuppergraphofFigure FIG. 16. The estmted coherence 15. Thehorizontal linegivesan estimate ofthelevelexceededbychanceonly5% ofthehmewhenthe spiketrainsare independent. SEISMOLOGY AND NEUROPHYSIOLOGY 31 with a similardefinition fordT(X). These are point processanalogsof the empiricalFouriertransform (3) oftime-series data.The cross-periodogram ofthe givendata at frequency X is definedas dT(X). INM(X) = (27TT)1dT() If the cross-periodogram is smoothedto obtain fTM(X),then fNMQ(X)is an in thecase that {M, N) is a bivariatestationary estimateofthe cross-spectrum of the impulseresponsea(t), pointprocess.Now A(X), the Fouriertransform may be estimatedby f T (X)fMTm(X)-1. The impulseresponseitselfmay be Coherence - RVP withRl 5 co c C; ~AA 0 1 4 3 2 5 frequency(cycles/second) ImpulseResponse LO LOl -10 -5 0 5 10 lag (seconds) FIG. 17. The estimated coherence and impulseresponseforthedata ofthemiddlegraphofFigure 14. The horizontalline in the uppergraphgives theapproximate upper95% nullpointof the distribution ofthesamplecoherence. 32 D. R. BRILLINGER estimatedby back Fouriertransforming AT. The strength of the relationship proposedin the model may be measured,at frequencyX, by the sample coherencyfunctionR'M(X) = fN(A)/ IMM(X)I NN(X). Its modulussquared is calledthesamplecoherence. The coherence liesbetween0 and 1, beingnearer to 1 the stronger the relationship. Moredetailsof thesecomputations maybe foundin Brllinger(1975b)and Brllinger, Bryantand Segundo(1976).[We here in Wiener(1930).] followtheuse oftheterms"coherency" and "coherence" Figures16 and 17 providetheresultsofsuchan analysisforthefirsttwodata sets of Figure15. In each case the firstgraphis of the samplecoherence. The coherencesare at some distancefromthe value 1.0, but above the 95% null linesin thefigures). significance level(givenbythehorizontal The relationship is inherentlynonlinear,so it could have been anticipatedthat the coherence estimatewould not be close to 1.0. Furtherdiscussionof these and similar analysesmaybe foundin Brillinger, Bryantand Segundo(1976). 12. Assessing connectivities. Questionsthat can arise with small networksofneuronsinclude;is oneneurondriving therestand ifoneapparently is, whichone is it? The nextdata analysisto be presented addressesthisquestion forthreeAplysiacellsL2, L3 and L10. Fromotherexperiments theneurophysiologistsknewthatcellL10 was driving cellsL2 and L3. It wasnotknownifthere wereanydirectconnections betweenL2 and L3. The firstthreegraphsofFigure 18 presentestimatesofthethreecoherences, L10 withL2, L2 withL3 and L10 with L3. As mighthave been anticipated, thesesuggestrelationship existsin each case. ofcellsL2 and It is possibleto addressthequestionofthe directconnection L3, in thepresenceofL10,bypartialcoherence analysis.Supposethat{A, B, C) is a trivariatestationarypoint process.Let RAB(X) denotethe coherency of RACand RBC. Then function of processesA and B, withsimilardefinitions oftheprocessesB and C, havingremovedthe(lineartime the partialcoherency ofprocessA, is givenby invariant)effects (13) RBCIA ~~V(1- RBC- RBARAC IRBAI12) (1 -_IRcAI12) suppressingthe dependenceon X. This definition may be motivatedseveral when betweenthe processesresulting ways. For example,it is the coherency theirbestlinearpredictors basedon A are removed. Or,it is givenby lim corr(d - ddT,dT ; d[T 2 and fBA/fAA' fCA/fAA are Here corrdenotethe(complex)correlation coefficient An estimatemay be determined coefficients. approximateregression by subsideofexpresestimatesforthequantitiesappearingon theright-hand stituting betweentheprocessesB and C beyondtheir sion (13). If thereis no connection SEISMOLOGY AND NEUROPHYSIOLOGY Coherence Li 0 and L2 o Coherence L2 and L3 o CR~~~~~~~~~~~~c o 33 .I , N N 14,0 1 2 3 4 Coherence Li 0 and L3 6 1 2 3 4 Partialcoherence L2 and L3 o 6~~~~~~~~~~~ expectedtoo 0e near zero. 6h fiaorp celsL,Lan 't,0 fFgr L10 2hr 8poie is no sugsto On Onrelecton, relecton, tfrequien tfisquienatoisin (Hz)shngth (Hz) herslso of diec h optto o h 4oncinbigpeet 2 degrizontal whine: linea:95%onullsleve 95heul leveloesn hogrizontalhth a nticshvor Dtatistfor FIG.ra18. anetwr ofpthreenAlysiapneurons esimat hishl highl baedonln baedonln the dpatloerdence FIG.raticDatatis dpatloendence esinmat ofppareentlysiaapturonsthe taeten to be 0nd zerg. the otereample jorinutidepndencrter onsA,sthen the coherence samperxmlso mayub s coherence ofapatial RT coIA partial earoinutiopedncuter onscussion coAmayuearin sitation. fourthern othert patil9ohrececopu eamplSesunof driscsiongand D. R. BRILLINGER 34 so far 13. A struCturalstochasticmodel. The analysesofneuronalfiring, direct with Parameters type. and regression correlation presented,are of the and Segundo In Brillinger have not beenintroduced. biologicalinterpretations and fittedbythemethodofmaximum (1979),a conceptualmodelis constructed elements. likelihood.The modelinvolvesthefollowing flowsto a genesis.Thiscurrent Inputto a nervecellleadsto electrical-current in thecourseofitspassage.Whenthevoltagelevelat zone,beingfiltered trigger the currentzone exceedsa thresholdvalue,the nervecell fires.The neuron Thisprocessmaybe specified backonlyto thetimeofpreviousfiring. remembers potential)at the as follows. Let U(t) denotethevoltage(membrane analytically triggerzone at timet. Let B(t) denotethe timeelapsedsincethe neuronlast fired.Let X(t) denotethe(measured)inputto thecell.Then,assuminglinearity onecan write and timeinvariance, U(t) = f( B (s)X(t - s) ds, (impulseresponse)a(.). The neuronfireswhenthe function forsomesummation level0(t). Dependingon thelevelat whichthe processU(t) crossesa threshold ofthenervecell,theinputwilleither mechanics is set and theinternal threshold and Segundo(1979), In Brillinger accelerate(excite)or slow(inhibit)the firing. as follows.Inputto thecellwas and discretized was completed thismechanism outputwas Yt, t = O,..., T - 1, writtenXt, t = 0,..., T - 1. Corresponding to t and with in the(small)intervalimmediate withYt= 1 iftherewas a firing With Bt denotingthe timeelapsedat t sincethe preceding Yt= 0 otherwise. time that Y = 1, theyset Bt- 1 Ut- E asXt-s. s=O a formoffeedback. ofintroducing The presenceofBt in themodelhad theeffect had theform(t = 0 + Etwith function Finally,theyassumedthatthethreshold distribunormalshavingmean0, variance1 and cumulative the e's independent tion functiond(*). ofthe givendata and themodelthentookthe form The likelihoodfunction T-1 H 4(Ut t=O - )Yt(1 - 1(Ut - 0))l-Yt. withrespect thislikelihood bymaximizing Parameterestimatesweredetermined determined were errors by procedures 0 standard to and the a.. Approximate likelihood. traditionalto maximum Figure19 presentsthe resultsof one such analysis.In thiscase fluctuating level intothecellR2 ofAplysia.The current currentX(t) was injecteddirectly uniform was that (but distribution approximately was takento have marginal The rate was 50 The samples/s. the technique). sampling that is not crucialto the and the noise of a stretch injected of the signal figuregives uppergraph are the fired. Details of the neuron which at experiments times corresponding ifnotimpossible, to see a givenin Bryantand Segundo(1976).It is verydifficult, 35 SEISMOLOGY AND NEUROPHYSIOLOGY NeuronR2 - Noise Driven 0 4 3 2 1 5 tky (seconds) Function Summation C ' 0.0 0.2 0.4 0.6 0.8 1.0 lag (seconds) Probability ofFiring Empirical 0. 40 ci ,r , , -300 -200 I I I/,.| -100 0 X 50 100 200 300 hnearpreditor with FIG. 19. The resultsoffittintheneuron modelofSection13 todata obtainedin an experiment theceUR2 ofAplysia.Theuppergraphis a segment ofthedata. Noise(lowertrace)is injectedinto the cell. The uppertracegies corresponding observed firngtimes.The middlegraphgives the maximum likelihood at 25 lags. Thelowergraph estinateofthesummation function a(-), estimated providesthe statistic(14) of Section13 and the curve4D(U- I) with# the estimatedmean threshold. The verticallineis at U = ?. 36 D. R. BRILLINGER connectionbetweenthesetwo stretchesof data. The middlegraphgivesthe function estimatedsummation a'. The lowergraphis onemeansofassessingthe (9) ofSection3, and givenby fitofthemodel.It is analogousto expression (14) #{Yt=1withU-h<Ut< U+h}l#{twithU-h< forsmall h, plottedversusU. Here U< Ut +h}, Bt- 1 Ut E s=O as t-s The smoothcurveis the corresponding is the fittedlinearpredictor. D(U - 0). werecarriedout by a The fitmaybe describedas adequate.The computations data of variantof the programdevelopedforhandlingthe seismicfirst-motion and Section 3. Furtherexamplesand discussionmay be foundin Brillinger Segundo(1979). Othertypesof inputare employedand alternateestimating procedures comparedthere. The large-sample ofsuchestimatesmaybe studiedas in Sagalovproperties approach,ofthissection,is sky(1982).A greatadvantageofthemodel-building A and estimated havebiologicalinterpretations. introduced thattheparameters furtheradvantageof the maximumlikelihoodapproach,overthat of partial is thatthespiketrainsinvolvedcan be highlynonstationary. coherency, 14. Analysis of evoked responses. A traditionalmeansof studyingthe the sensory stimulito a subjectand examining nervoussysteminvolvesapplying foran evokedresponse.The stimulusmay be ongoingelectroencephalogram somatosensory olfactory, pattern), visual(e.g.,lightflash,checkerboard auditory, thestimulusis applied or a task.Generally, (e.g.,an electricalshock),gustatory fora timeintervalthatis briefin comparison to the durationof the response. play an essentialrole in quantitativebiology. experiments Evoked-response is able to choosewhichstimulito applyand whento Because the experimenter noted,to formalinferences can pass beyondassociations applythem,conclusions thesameas the are formally Theseexperiments causal mechanisms. concerning in Section6. described reflection experiments seismological One is Some dramaticsuccessstoriesof the techniquemay be mentioned. Fra,Gandiglioand Mutani(1967).Siamese Bergamasco, presentedin Bergamini, twinswerejoined in such a way that it was not possibleto determineby traditionalmeans if the peripheralnervouspathwayswere interconnected. of the twins. it was crucialto determine theinterconnections Beforeoperating, OngoingEEGs wererecordedforeach. A seriesof trialswerecarriedout in in turnby electricalshocks.What whicheach ofthe twins'legswas stimulated was a of one twin stimulated, responsewas noted was foundwas thatwhen leg the twins were the of this basis information, only in her EEG. On use of the evoked of the separated-successfully.A secondnotableexample infants for newbom exams is (including responsetechnique providedby hearing Theseare examinedforresponsesafterloud infantsasleep.)EEGs are recorded. clicksare made near the infants'ears. Rapin and Graziani(1967) presentan SEISMOLOGY AND NEUROPHYSIOLOGY 37 bothwearingand notwearinga exampleforan infantwithhearingdifficulties, measurableeffect. hearingaid. The hearingaid is foundto have an objectively data recordedat a 4 x 4 Figure20 presentsan exampleof evoked-response in a rabbit.In thiscase thestimuluswas an odorand arrayofsensorsimplanted in orderto studytherabbit'solfactory system.These thesensorswereimplanted A secondexampleis givenin Figure21. It responseswererecordedconcurrently. gives the 20 successiveresponsesevokedby applyinga currentpulse to the froma sensorimplantedin the tractofa rabbitand recording lateralolfactory in Figure20. In cortex. The is of the fairly pronounced signal pre-piriform depth the is was and the weak of stimulus signal notapparent. Figure21 the strength in the laboratoryof W. J. Freeman, Both of these data sets were collected Somedetailsofhisexperiments ofCalifornia, maybe found Berkeley. University in Freemanand Schneider(1982). is the factthatit is generally Crucialto manyevokedresponseexperiments to apply a stimulusonce. Ratherit must be applied repeatedly insufficient averaged.(Thisis also truein the (perhapsthousandsoftimes)and theresponses case of reflection seismologyas mentionedearlier.)In the twinsand infant if M equaled250 and 100,respectively. Formally, examplesdiscussedpreviously, Y(t) denotes the measuredEEG and the stimulusis applied at times am, m = 1,..., M, thenit is usual to takeas thebasicstatistic,the averageevoked response 1M m M m=1 thedata of The left-hand columnin Figure22 presentstheresultsofaveraging averaginga signalis Figure21 with M = 3, 5, 10, 20 and 38. Withincreasing slowlyappearingfromthe noise.Some alternateevidenceforthe presenceof a column.Theseare averagesof signalis providedbytheresultsoftheright-hand 38 responses,wherethe stimulushas beenappliedat a successionofincreasing strengths. arisein thecourse A varietyofquestions, whichhavestatisticalformulations, (2) ofworkwithevokedresponses. (1) Does an appliedstimuluselicita response? stimulielicitthe same response?(3) Is the same response Do two different (5) If the elicitedat twodifferent sensorlocations?(4) Is theresponsestationary? responsesaltered? orderof stimuliapplicationis altered,are the corresponding stimuliadditive?(7) How does the response (6) Are the effectsof different dependon exogenous (8) Howdo theresponses intensity? dependon thestimulus seekquickefficient researchers variables?To go withanswersto thesequestions, that Difficulties ofvariability. data collection, preciseestimatesand indications commonlyarise includesmall response,large noise,variabilityin response, artifactspresentand superposed effects. Nextin thissection,twoformalset-ups willbe presentedthatmaybe employed to addressthesituation. Suppose, to begin,that thereis a singlestimulusand that it is applied If experiment. at times am. Let a(-) denotethe responsein a single-shock the systemis time invariantand the effectsof the variousshocksadditive 38 D. R. BRILLINGER RabbitOlfactory System- Responses at 4 by4 Array 2. 0.0 2. 0.0 .2 00 .6 00 2 . 0.02 0.04 0.06 0.08 .2 00 0.02 o .6 00 0.04 0.06 0.0 02 . 0.08 0.0 .2 00 .0 2 2. 0.02 0.04 0.06 0.08 0.0 .2 00 0.02 0.04 2.0 0.06 0.08 .2 00 0.0 .6 00 0.0 00 .4 00 .8 0 s 0.02 0.04 0.06 0.08 .2 00 0.02 0.04 FI.2 0.02 0 2wure in 0ec0nd0 0.04 0.0 .6 00 0.06 . 0.08 0.06 00 00 4 00 Th2usso .8 2n 2abt .6 00 0.02 0.04 0.06 0.08 .2 00 0.0 0.02 00 00 .6 00 0.04 0.06 .4 00 0.08 0.0 0 I 0.08 ,2 00 0, inseconds. 0.0 . .6 00 o0 0t;X o . 2. .2 00 0.0 0.02 0.04 lcrecparrpi onigtesito 0.06 0.08 0.0 0.02 0.04 ciiy eodda fterbitb 2 0.06 ot 0.08 h 6snoso noo.Teuiso 0.0 n 0.02 h 0.04 0.06 -n 0.08 ra r SEISMOLOGY AND NEUROPHYSIOLOGY 39 Individual Responses - RabbitPre-piriform Cortex response 1 response 2 response 3 response 4 0 o o3 0.0 0.04 0; 0.08 0.0 response 5 o.0 0.04 0.08 0.o 0.08 0.04 9 0.0 response 13 0.0 0.04 0.08 0.04 0.08 0.08 0.o 0.04 0.08 0.0 0.04 0.08 V 0.0 0.04 0.08 0.0 0.04 0.04 0.08 = 0.04 0.0 0.04 ? E; 0.08 0.04 0.08 0.08 response 12 0.0 :: 0.04 0.08 response 16 0.0 response 19 0.o 0.04 0.o 0 0.08 0.08 response 8 response 15 response 18 0.o 0.08 response 11 0 tA> 0.04 response 7 response 14 response 17 o.o 0.0 response 10 0 0.04 0.08 response 6 response 9 0.0 0.04 0.04 0.08 response 20 0.o 0.04 0.08 FIG. 21. Twentysuccessiveresponsesevokedin thepre-piriform cortexby (electricaey) stimulating a rabbit. The x-axis unitsare in seconds. 40 D. R. BRILLINGER Average Evoked Responses - Several M's, Several Intensities - averageof 3 100% threshold 1 89% threshold - averageof38 I- 0.0 0.02 0.04 0.06 0.08 0.10 0.0 0.02 0.04 0.06 0.08 0.10 0.0 - averageof 10 100% threshold 0.0 0.02 0.04 0.06 0.08 0.02 0.04 0.06 0.08 0.10 0.0 0.02 0.04 0.06 0.08 0.08 0.10 0.02 0.04 0.06 0.08 0,10 0 02 0.04 0.06 0.08 0.10 - averageof38 133% threshold 0.10 0.0 - averageof 38 100% threshold 0.0 0.06 122% threshold - averageof38 - averageof 20 100% threshold 0.0 0.04 - averageof38 111% threshold - averageof 5 100% threshold 0.0 0.02 0.02 0.04 0.06 0.08 0.10 - averageof38 144% threshold 010 0.0 0.02 0.04 0.06 0.08 0.10 FIG. 22. The variousgraphs hereare meantto show theeffects of changingthenumberof responses averaged (left column) and thestrengthofstimulusapplied (rightcolumn) fordata such as thatof Figure 21. SEISMOLOGY AND NEUROPHYSIOLOGY 41 thena modelforconsideration is (superposable), (15) Y(t) = ,u+ 2a(t- am)+ E(t), m noise.In thecase ofthe withY(.) denotingtheongoingEEG and E(.) denoting ratherthanbeingan EEG thismodelseemsto have to be empirically verified, case it cameout ofa concepimplicationof basic biology.(In theseismological For example,the assumptionof superposability tual framework.) may be examinedas followsfortheanimalstudied.To begin,carryout somesingle-shock experiments, i.e.,applytheshocksat timesfarenoughapartthattheirindividual effectsseem likelyto have died off.Let a&(s) denotethe averageof the responsesevoked,withs lag sincestimulusapplication.Now carryout some two-shockexperiments, i.e., apply shockssay A timeunitsapart. Let b(s, A) denote the averageof the responsesevoked.To examinethe assumptionof superposability compared(s) + a(s - A) withb(s, A). The resultsof carrying out such a check,in an experimental situation,are givenin Biedenbachand Freeman(1965). They forn averagesof M = 150 responsesand do not note departurefromsuperposability. We nowturnto one formalanalysisofthemodel(15). If one writes X(t) = E 8(t m rnM), then(15) takestheform Y(t) = ,u+ la(t - s)X(s) ds + e(t)q i.e., it is seen to be the modelof cross-spectral analysis.TakingFouriertransone has forms, dT(X) = A(X) dT(X) + d[(X), forX > 0, withA(X) denoting theFouriertransform of a(-). Considera number of frequenciesXk = 27rk/Tnear X. Then,assumingA(-) smooth,one has the linearmodel approximate Yk A(X)Xk + Ek, with T-1 Yk= 2,,kt\ E Y(t)exp - i T } t=0 and similardefinitions of Xk, Ek. Next,via a centrallimittheorem forempirical Fouriertransforms, thenoisevariatesEk maybe approximated by independent All the inference (complex)normalshavingmean 0 and variance2fTfTee(X). forthelinearmodelbecomeavailable.Forexample,as an estimateof procedures the transfer function A(X), one has A(X)= D. R. BRILLINGER 42 as complexnormalwithmean distributed and thisvariatewillbe approximately has a varietyofconveniA(X) and variance27TTfee(X)/E4lXkI2.Thisformulation It directly extendsto the cases ofmultiplestimuliand multiple ent properties. responses.It handlesstimuliof varyingintensity.It allows the individual suchas procedures, responsesoftheseparateshocksto overlap.Formalinference maybe designedand analyzed-comtests,are available.Complexexperiments meastructure, factorialtreatment rotation, plexitieshandledsuchas blocking, are available.Finally,one can suredcovariates.Formalchecksforinteraction turnto the questionofoptimaldesign. viewpointforthe problem. to adopt a different It is sometimesconvenient Suppose that the shocks are applied at times such that ,m,, -,,, a(s) = 0 fors > V and s < 0. Write = Y.(S) Y(s + > V with am.) Then Yin(S) = ,x+ a(s) + Em(S) for0 < s ? V. The averageevokedresponseis now conveniently denotedY(s). As an exampleof the use of thisformulation, stimuliand thateachare appliedJ times,thenone supposethereare I different is led to set downthemodel Y,j(s) = Alij+ a(s) + b,(s) + eij(s), suchas Othermethodologies, replicates. withi indexingstimuliand j indexing analysis,are seento becomeavailablewiththis growncurvesand discriminant formulation. byartifacts. data maybe contaminated thatevoked-response It wasmentioned available. are directly estimates It is perhapsworthnotingthatrobust/resistant such as of a measure one has distance, Suppose [Y(s) - a(s)] IIY- al2 - ds estimatesis and an estimateof scale p. Then a familyof robust/resistant providedby a(s) = ,WmYm(S) F2Wm, m m with WM= W(MIY - ad/,p) and W(.) a univariateset of multipliersfor An eleThe estimatewillneed to be computedrecursively. robust/resistance. mean" mentaryexampleis providedby the"trimmed a(s) - E'Ym(S)/13M, was proposedin - all. This classofestimates withE' overthe fBMsmallestIYYm in Folledo(1983).The uppergraphof (1979,1981a)and investigated Brillinger (/3 = 0.5),in Figure23 providesan exampleofthisestimatewith50%trimming 122%of the case of data likethatofFigure21 (butwitha stimulusofstrength the thresholdstimulus).The solidcurvedenotesthe averageevokedresponse, the dashed one the trimmedstatistic.The two curvesare nearlyidentical, arefoundto differ theindividual noticeably. responses althoughwhenexamined, a highly becausethisis usuallyconsidered was employed, trimming Fifty-percent SEISMOLOGY AND NEUROPHYSIOLOGY 43 Average Evoked Response and Robust/Resistant Variant If) 0 cmJ 0.0 0.02 0.04 0.06 0.08 0.10 time(seconds) 122% threshold- 50% trimming RunningTrimmedMean and Robust/Resistant Variant 0 LO 0.0 0.02 0.04 0.06 0.08 0.10 time(seconds) 122% threshold- 50% trimming FIG.23. The uppergraphcomparestheaverageevokedresponsewiththe50% trined meanfor thedata takenat 122%ofa threshold stimulation value.Thelowergraphcontrasts the50%trimned meanstatisticwitha valuecomputed recursively. resistantlevel in the case of elementary statistics.The factthat the trimming had suchlittleeffect on thefinalanswersuggeststhattherewereno substantial outlyingcurvesin thedata set.Had a curvebeenfarremoved fromtherest,then it wouldhave beenrejectedfromthe average.It is to be remarked thatin this case ofpresentconcem,wholecurvesare beingeliminated fromtheaverage,not just outlyingpointsthatsomecurvesmighthave. It is to be notedthata "real-time" versionofsucha trimmed meanmaybe computed;see Brillinger (1981a). This statisticis givenrecursively for m = 44 D. R. BRILLINGER 1,2,... by Pm m = a L 1 - - Pm' and if IIYm? -amII < Pm+? = Pm+ L/m and by otherwise, am?i(S) if IIYm+? - Pm, + am(S) (Ym?i(S) Ain(S)), and A ?i(S) = am(S) otherwise. a workedexample,it was foundmoreconvenient in the (In preparing The lowergraphofFigure23 givesthe choiceofL to replacep byitslogarithm.) resultforthe same data as that of the uppergraph.The algorithm was run settingad(s) = Yl(s) and L = 0.15.The real-time estimate,givenby the solid as wellas thedead-time estimatein thiscase. One line,has performed virtually can remarkagain that had therebeen somehighlydissimilarcurvespresent, fromthesampleaverage.Followingthe thenthisestimatewouldhave differed advice sometimesgivenin connectionwith resistantregression estimates,it wouldseemsensibleto computeboththeordinary and theresistant If the forms. If thetwodiffer twoare similar,thenthereis no difficulty. thenthe noticeably, situationshouldbe examinedin somedetail. Brillinger (1979)proposedthe preceding techniquesand variousothers.Brillinger(1981a,b) werebased on that lectureand coversome otherstatistical methods.Tukey (1978) also addresses problemsarisingfromevoked-response statisticalissuesand proposessomeprocedures. 15. A confirmed(Fourier) inference. Muscle cells are electrochemical is appliedat theneuromuscular devices.If the chemicalacetylcholine junction, releasecauses result.Specifically, measurablevoltagefluctuations acetylcholine Katz membrane channelsto openleadingto voltagefluctuations. postsynaptic associatedwith this and Miledi (1971, 1972) measuredvoltagefluctuations and foundthatthepowerspectrum couldbe approximated phenomenon by the to suchdata forma/(/32+ A2).[Anexampleofthefitofthisfunction functional of a fitting and a description procedure maybe foundin Bevan,Kullbergand Rice (1979).]Theyproposedthemodel Y(t) = >a(t -am), m withthe o(Jpointsof a Poissonprocessand witha(t) = exp{-,t}. This a(-) to theeffectiveness of an openchanneldecayingexponenfunction corresponds tiallyand leads to a powerspectrumof the indicatedform.Katz and Miledi but ofrandomduration, thatthepulsesmightactuallybe rectangular mentioned form.Stevens(1972)proposedthe to deal withthe exponential theypreferred SEISMOLOGY AND NEUROPHYSIOLOGY 45 specificmodel Y(t) = Yam(t am), m also with (am) Poisson, but now with am(t) = 1 for0 < t < Tmand am(t) = 0 The Tmare independent otherwise. ofmean1/fBand correspond exponentials to the lengthsof timethatthe channelsare open.Stevensnotedthat thismodel also led to a powerspectrumof the forma/(,2' + X2). The models were withthedata collected. indistinguishable The problemwas laterresolvedby improvedexperimental technique.Neher and Sakman(1976)developeda techniquethatallowedtheopeningand closings of individualchannelsto be seen. They foundthat the channelsremained and openfortimeperiodsofvarying equallyeffective The twoproposed lengths. modelscouldbe distinguished. Examples of single-channel data and the corresponding estimatedpower spectrummaybe foundin Lecar(1981).Jacksonand Lecar(1979)presentresults the exponential confirming durationoftheopenings. 16. Other topics. Spatial-temporal data are commonly collectedby neuroscientists.One formis theelectroencephalogram recordedby an arrayofsensors on the scalp. Figure19 presented an exampleofdata collectedforthe olfactory systemoftherabbit.The stimulus wasreleaseoftheodorethylacetate. An8 x 8 arrayof electrodeswas imbeddedin theanimal.The data, alreadypresented in Figure19,givetheresponsesforthesensorsat thepositionswithx-coordinates 2, 4, 6 and 8 and y-coordinates 1, 3, 5 and 7 of Figure23. One procedure that Freemanhas foundhelpfulforunderstanding thistypeofdata is thecomputing of empiricalorthogonalfunctions;see Freeman(1980). Figure 24 gives an example.These resultsare derivedby stackingthe responsesinto a matrixX withrowscorresponding to sensorand columnsto time,and thencomputing the singularvalue decomposition X = UDVT,ofthatmatrix.The U fora particular component, are thenplottedversussensorlocationas in theupper say thefirst, graphofFigure23. The V valuesaresimilarly plottedversustimeand appearin thelowergraph.The resultsofFigure23 arebasedon 64 series,notjust the16 of Figure19. The contourplot suggeststhe presenceof a focusof activity.The time-series elicitedmaybe seenlurking component in theindividualresponses of Figure 19. (It may be mentionedthat meteorologists have long computed empiricalorthogonalfunctionsforspatial-temporaldata and used themin see Lorentz(1956),forexample.A numberof otherreferences forecasting; are givenin Jolliffe (1986).] Childershas also made use of arraydata in studyingthe neuralsystem.In Childers(1977),he estimates thefrequency-wavenumber spectrum forresponses evokedby visual stimuli(lightflashes)in the humanEEG. He was concerned withestimating thespeedand direction ofpropagating waves.In thepapercited he firstnotes an apparenthigh-velocity wave. Afterthis wave has been "removed,"he notesthepresenceofa pairofwavesmovingin oppositedirections. His researchis directedat developing a diagnosticprocedure forvariousvisual disordersand obtaining insightconcerning howthevisualsystemfunctions. D. R. BRILLINGER 46 LatentSpatial Component N- (0~~~~~~~~~~ LCI) cl-) co 1 0.0 0 2 0.02 3 5 4 0.04 6 7 8 0.06 0.08 time(seconds) FIG. 24. The results of a singular value decomtposition of the full set of the data fromwhich the burstsofFigure20 weretaken.The valuesgraphedare for thefirst components. upper graph give spatial location. The axes in the The decayingcosinemodelof Section2 has also founda use in nseurophysiology.In his workwiththeolfactory system,Freeman(1972,1975,1979)found that the averageevokedresponsecould be weR fittedby the sunnof a few He developeda modelinvolving decayingcosinetertns. spike-to-wave conversion, collections of constantcoefficient second-order differential involving equations, feedforward and feedbackand involving conversion. He involving wave-to-spike in thetimedomainto estimatetheunknowns. In employednonlinear regression one case, involving two cosines, he was led to view the strongerwave as representingintracorticalnegative feedback and the weaker as representinga SEISMOLOGY AND NEUROPHYSIOLOGY 47 secondfeedbackloop. Of interestin thistypeof workis whathappensto the conditions are altered.A and thedecayrateswhentheexperimental frequencies to decayingcosinesis Childersand Pao (1972).They consider secondreference the model t > 0, Y(t) = EakteXp{ 3kt}coS(Ykt + Sk) + e(t), k forvisual evokedresponsesmonitored overthe occipitalregion.In particular, theystudythe data by complexdemodulation. Briefreference will be made to severalothertopics.Dumermuth, Huber, Kleinerand Gasser(1971)estimatethe bispectrum of humanEEGs. de Weerd and Kap (1981) discuss the computationof some time-varying quantities. Marmarelisand Naka (1974)considerthecase ofbiologicalsystemswithseveral inputs.An extremecase of thisoccurswhenthe inputis varyingin bothtime and space.This circumstance is considered in Yasui,Davis and Naka (1979).The bookby Marmarelis and Marmarelis (1978)goesintogreatdetailconceming the identification of systemsthat are polynomial and timeinvariantin the input. Theyemphasizetheadvantagesresulting fromemploying a Gaussianwhite-noise input.The dedicationofthebookis worthmentioning-"To an ambitiousnew breed:SYSTEMS PHYSIOLOGISTS". Anotherarea of researchactivityhas been that of control.The worksby Poggioand Reichardt(1981)and Wehrhahn, Poggioand Bulthoff (1982)maybe noted.They are concemedwithdata thatare three-dimensional trajectories. 17. Discussion. As theexamplespresented indicate,a broadrangeofdata data are collectedat both the typesarise in the neurosciences. Furthermore, microand macrolevel.The procedures to developedoftenhavetheopportunity moveon to directclinicaluse. It is particularly to notetheevolution oftheanalysisin thecase of interesting the neuronalsignalinganalysisas presentedin Sections11 and 13. One can recognizethe stagesof (1) (feature)description; (2) correlation/association; (3) (ad hoc) regression;(4) conceptualmodel. These stages are usual in many situations. elementary The fieldof neurophysiology has the satisfying aspect that in manycases controlledlaboratoryexperiments are possibleand repeatable.Furthermore, thereare opportunities forthedesignofexperiments. In thefield,statisticshas beenseento providetechniques formodelformation and validation, formeasurin conclusions and for of inguncertainty addressing questions causality.Statistical techniqueshave led to insightconcerning In this theunderlying physiology. connectionit seemsimportant to notethe following collaborator provisoofmy J. P. Segundo, however,".... The maximof all of the above is that the powerof available mathematics(and of the instrumentation that implementsthem) should be used exhaustively, gded by an unflagging biologicalrealism,misand keepingin mindthattheultimategoalis understandtrustful and stubborn, ingin strictly biologicalterms."[See Segundo(1984),page 294.] It seems likelythat in the neurosciences, more oftenthan not, notable advanceswillcomefromthecarrying outofnovelexperiments, ratherthanfrom 48 D. R. BRILLINGER thingsat new novelanalyticmethods. Experiments willbe carriedoutmeasuring ordersof smallness.Morecomplexstimuliwillbe invoked.Nonlinearsystems will be the norn. Neuralnetworkswill be a major concern.Luckily,forus havebecomecommonin thelaboratory and this statisticians, digitalcomputers seemsto be bringing a movetowardquantizationof otheraspectsof the work ofthedata. beyondthe simplerecording data, briefly referred to in 18. Update. The analysisof singleion-channel Section 15, has becomea wholeindustry. Modelswithseveralstatesare now routinelyfitted.References includeColoquhounand Hawkes(1983),Labarca, Rice,Fredkinand Montal(1986)and Milne,Edesonand Madsen(1986).Extending the workof Section13,Brillinger (1986)presentsa numberof examplesof corresponding spike themaximumlikelihood fitting ofa neuralmodelemploying traininputand outputdata. Smithand Chen(1986)studya morecomplicated as beingof substantialimporneuralmodel.The chirpsignalwas propounded in physiologitancein seismicexploration. Someuse ofit has beenmaderecently cal studies.In Norciaand Tyler(1985),a 10-sspatialfrequency sweepstimulus is visualevokedpotentialmeasured.Th. Gasser employedand the corresponding and collaborators have nowcarriedout a substantialnumberof statisticaland We mentionin particularthe papers substantiveanalysesof evokedresponses. by Mocks,Tuan and Gasser(1984),Gasser,Mocks,Kohlerand de Weerd(1986) and Gasser,Mocksand Kohler(1986).Finally,we notethat Grajski,Breiman, di Priscoand Freeman(1986) applymodernclassification to study procedures bulb EEGs of rabbits the effectsof applyingdifferent odorson the olfactory and that Gevins,Morgan,Bressler,Cutillo,White,Illes, Greer,Doyle and accuracyto brainelectricalpatterns Zeitlin(1987) relatehumanperformance just beforea task. IV. Concluding remarks. In thisarticlewe have presenteda numberof showingthe use of the examples,drawnmainlyfromour personalexperience, same statisticaltechniquein the ratherseparatesciencesof seismologyand have the to ask what,if anything, It nowseemsappropriate neurophysiology. fromeach other threesciences-statistics, neurophysiology-gained seismology, even thoughtheyare indirect?Havingin minda as a resultof connections broaderclassofexamplesthanthosediscussedin thispaper,onecan saythat:(i) statisticsis richerforhavingbeen led to developand studyvariousnovel or neurophysiology; methodsto handlespecificproblemsarisingin seismology are the richerforthe other'sfield (ii) both seismologyand neurophysiology thatthe to abstractsufficiently havinggenerateda problemforthestatistician or result'sapplicabilityto theirfieldbecameapparent;(iii) eitherseismology becausevariousof their benefitfroma statisticalfornulation neurophysiology to need to be statedin termsof probabilities (e.g., problemsseem necessarily and becausethese nor earthquakesseem deterministic) neitherneuronfirings to validateresultsand to fitconceptualmodels.That the fieldsneedprocedures in submethodsof statisticscan lead to important insightand understanding stantiveproblemsseemsagreed. SEISMOLOGY AND NEUROPHYSIOLOGY 49 It may be remarkedthatthe applicability of statisticalprocedures to these two substantivefieldshas further oftheirmoveto grownin directconsequence greaterquantification and digitaldata collection. The data setsanalyzedwereof highquality.The factthattheanalyseswereinformative to an extentherebodes in fieldswithdata oflesserquality.I needto wellfortheuse ofsuchtechniques remarkhow crucial,in working withthedata setsdiscussed,I have foundit to be to plot the data in its originalform.Something specialseemedto be learned in each case fromdoingso. Thisis whyforthevariousanalyses,I havesoughtto providedata plotsas partsofthepresentation. The readerwillhavenotedthatsomeoftheanalysesweretime-side and some werefrequency-side. Each domainhas its advantages.It seemsworthpointing out specifically thatstationarity was notrequiredforsomeofthefrequency-side procedures.It wouldseemthat mosttimeseriesand pointprocesssituations would benefitfromcarryingout simultaneoustime-sideand frequency-side analyses. On reviewit maybe seenthatthe techniquesemployedfortime-series data and forpointprocessdata in manycasesare notthatdifferent. Brillinger (1978) presentssome comparative discussionof the techniquesforthe two cases. Our presentation is somewhat remissin theseismological case in notpresenting some workedexamplesof auto-and cross-intensity estimation. Examplescouldhave been provided. It shouldbe apparentthatmajordata management and computational efforts wererequiredin thederivation ofall resultspresented. I havebeenimpressed by the way that the neuroscientists couldturnto theirlab book keptduringthe and pulloutcrucialdetails,sometimes experiments manyyearsaftertheexperimentshad been completed.My analysesalso extendovermanyyearsnow. I have foundit veryusefulrecently to maintaina "Readme" filein the various computerdirectories forthedata sets,whereinI listwhatthevariousprograms do, futurewishesconcerning the programs and all of the thingsthat I thinkI willneverforget. Turningto thoughtsconcerning to come,it seemsthat the developments futurewillsee manyofthetraditional statistical techniques extendedto applyto datumofmorecomplicated forms-specifically, to curves,movingsurfaces, point cloudsand thelike.It seemsthattechniques developedin onefieldwillcontinue to be transferred (by statisticians?) to otherfields.For example,I expectto see theresultsdevelopedbyneuroscientists forarrayson a curvedsurface(theskull) to be takenup by the seismologists as theyneed to take specificnote of the Earth's curvature.If I have foundanythinglackingin our currenttoolkitof statisticalmethodsand devices,it is a collection oftechniques thatsuggestwhat to do nextwhena modelfailsa validationcheck.Perhapsthefuturewillsee such techniquesdevelopedin an organizedfashion. Acknowledgments. One wondersif it is possibleto workmeaningfully on problemsfromsubstantive areasunlessoneis in closecontactwithresearchers of those areas. I doubtit. I can say that theselectureswouldneverhave come about,but forthehelpand encouragement thatmycollaborators B. A. Bolt and 50 D. R. BRILLINGER J. P. Segundohave providedthroughthe years.I cannotthankthemenough. The influence ofmydoctoralsupervisor, J.W. Tukey,shouldbe readilyapparent I remember hissaying,whenI was a student, as well.In particular, "One cannot unlessone becomesa chemist."This has certainly consultwitha chemist, been withrespectto thepartsofneurophysiology and seismology myownexperience on whichI have worked.The reversehas also provedto be the case, namely, fromthosefieldshave had to learnpartsofstatisticsas wellas I collaborators knewthem.I also mustthankmystudentswhohaveworkedwithmeon various I mentionM. Folledoand R. Ihaka, of the problemsdescribed.In particular, whosetheseshave beendetailed. Finally,I would like to thankJ. Rice and R. Zuckerforprovidingsome I wouldlike to thankN. Abrahamsonand R. neurophysiological references. 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