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ThePhysiology of Analgesia Stimulation-Produced

Key words AcLtpuDalure, Neu/o-,lrn1or)y, Piin 5lin )uIaIi on-pt odu.-.d.rra/8eila, IFNs. rrajor blood vcsscls surrounrled their peflvas by path$,ay5, cular plerus of ncrVrvasorunr. Thnt sympathclic nerves nray bL.involvcdw,asiirrl denronstrated by Couldef (1921 rvho showed thrt acupuncture ) poifts alongthc sciali. n-of\,e its branches a lntrcduction ancl had Tw,o types ol cLrtaneous stimulirtionhave been lower impcclancc ihan lhe sLrrroLrnding Yoshio sl(in. knor'vn from time immemorial be analgesrc, (1961)has showf lhat Ry.xloraku fts havc o\^, po to ly Th-. iirsl is the instinctiveaction of impedance/high eifective. conductanceMany acupLllcturc "rulhing a pain L)-.ll-.r", ancl the second is pointsare of coufscdcep lo lhe skin:Melzac< a/ ct acupunct!re. Rubbing done rapidlyand lightly, is scr (19//) havc shorvnthat Lhey correlate closclywitlr hi!:hfrequency, pollrls", in physical terms constitutes law T|ave " tri.qtcrl's while Liu eta/. (19/7) have js Acupuncture performed \,"'ith shownthat othcrs irtensit), stimulation. correspond with the motorpoints which is rotated a needle, fairlyslow,ly 2 {about of l ol mLtsclcs. tinresa second),and so constituteslo\,! frequency, What typc oi receplof mtrst be stimulatedlo pfoducc thc acupuf.lure effect?The facl lhal .r h/ghlnten5lfy stimulation. li we accept thatthese two forns of stimulatron arc needle sonre or olher intense sti|rulus ntuslbe used (Bo\'\,sher empiricallyeifective,wc should cnquirc rrhich poifls to a high-thfeshold reccptor I976). p t r r i o l o c , . p , l _ u .r . ' r d m ^ ,l . r ' n r a t , WanS e/ a/. (1985)sholvc'd thal Ab or Crout) lll respons for theiractions. blc fillres in iactlhe groupconccrncd. are Receptorsand primary afferents Spinalcord nechanisms Falr evidently, rubbing activates lar8e lo$,- The fext question bc asked, to ob!iolrsly, whcfc is ), (AB).Sin.e sLrstainedanclhou,do these (RA cutancous aflerenls threshold two typcsoi primaryJiierent pressure nol effective producinB pain reliei, is in the Ap anclA5) s\rnapsc the spinalcord?Foftunatcly in (RA),nncl recent LIlil\ musl be rapidly-adapting eflective advafces anatomy in and l)hvsiology tel can by ly, excited tlansientslimuli. TherapeuLica pain us a Sfeat cleal aboutthcscqueslions. cutaneous Primarv relef by aclivalion of low thresholcl affereftAp fibrcsenterthe spinalcod in is afferents obtainedby LranscutaneoLrs electricirl the medialpaft of the dorsalrool .rndpassrostt'a ly (TENS) (Wall & S\,-.et,1967).lt withoutsyfapscin lhe dorsa (olumnsto cnd n the nervestimLrlation by cana so be obtained the .rpplic.tion vibrator), dorsal column (Sraci an(l clrneate) nuclei. of e stmull (LundeberS,I9Bl), \ ,hich presumably However, cntry 1o lhe spinalcord, thcsc iibres on (AB) rapidly-a.laplinB primaryafferents in give off a segmcrtal ater.rl pencttatcs activate col w,hich the greymatter thc dorsa horf. Some muscle. oi arborsatrons ol presents nrore clifficultproblem, this branchmakc contactin the superfcialparl of AcupunctLrre a it to that the dorsal honr, probablythrough a CABA ergic because was iirst necessary clerfonstrate (To.ldan(l McKenzie,1989), nerves are invoivecl all. Halrpily,experments interncuronc nt w,ilhthe whichshorved that when r nerveis blockedw'ith tenninals small ir](omin8primaryaiicrenlIibres ol is in localanaesthetic, ncLrpunclure inefiective the (se.' Blown ct al., 1978) (Ftlure ir). Such ar1 that the afatomicalarrafBenrent would erplain thc CAIJAterritorysupplied by it, demonstrate effect is conducted along ncrvcs crgic inhibilion observecl Duggan ard Foong acupuncturc by (Chiang a/., 19/3). Also, rt was (nown that ( 1 9 U 5 ) o l l o w i r l S o r s a lc o l L r m n t i m L r l a l i o n e e (s et i d s points thc belo\,v), is on w,hich retrogradely acupunciureonly efectiveat cedain drivcs lhe segnrental body suriace. In iact, comparisof with an .ol alerrlsresponsib for TENS. is oi course e ll rvell atlas shows that many acupurcturc know|l lhat these s),sternsrcspond to high anatomical points with the points at which smali lrcquencies stimulirtion. thus have a circuit oi correspond lvc tre Iascia;Chan (198,1) .apabe oi explirinlng,nt scgme/)tal nervebundlespenctrate /eve/, thc st!dies$/hich showe.lthal 109 analSesic cites two Chinese elfects TENSan.l villration.Note thai of poinlsare situatcd or very closelo 111is on acupuncture cifclrit does not rrvo ve arv opiorderg c 286 nerves, whie a fLrrlhef are on or Verycloselo synapses, therelbrc anal8,"!i.r and thc by l)rocluccd 58

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ACU

Bated on Bo(she,1935 1937

Cinul t lat hi\h incnsit\, b|| ir\tuencr iA.upunctute) rrd h ighJrcqoen.y Io\\' iDteD ry i[N5] !tnru/.rtror. si PAC = Pctiaquachl.tal Btar ,Ri!1 = nu./cus Ripha MrtrlJ = 5ul,starnaOe/.rltnrr. 5C = Itansnission.el! T = Wald.yat c.:ll W CABA = yamino butyti. a.id = s!bnir.. P 5P VIP = Vataactlvcin]rsti'al paptidt: = [nkcphalinetit. intetneuote E by L Fine mrelin,r.i.l Ai f)tinrn,- .t1epnl. are e.tivatcl hi\h intensd|, tptnpti.k) stntuli trithlli the .oft|, they synapsa dirc.tiy trith tnten)e]rcnes tL) \rhi.h, usiDEthe inhibito^ ansnitter enkephnlin (LDk) dD nlhibit trcnsnissioni9!l whi.h rc.eire na.icaptir.: sobsttnti. sel.tino (SC)neDrcnes ntput rni Dnntyeiinile(llC) Vini.]y affercDtt usitlg SubnanccP or rasaa(tne tntestiml Peptl'1elVlP) .1s i tnntmlttct. l5P) (Se entaI a. upun.t trc ). t:m A6 p.nnatr atetents .lso end an l.nliln t .ellt (\\t) \rhi.h ansnit pinpi.k ini.nnutia. dite.tlr to.an..iausne$ ria the spn,ahalani. |..'cl; /la gir,-.soii.ollat-.nk to the petiaqtrc du.titl Ercr mattet (PAG),w/rn:/rJ.riu/es t des.eD./irgt),sterr r:larilg in th.: nrlLltiusl<apheM;lnus t.].tv) ot the ne.lDlla abhil1ata ta ax.it.i rihl)it \r enkephali.etBkintemeureDet lE) thrcuih th.: da^alateral slrniil titni.iilLs theterctopi. 2. LaryL'lavthrcshol.l ptint,nf iltetenls t\lt prs (lne.tly up the dorsal.alumnt on entetin! the.o1l) hut thet rka Eive oii a sclncnkl ollatltai wl1t.h e\.ites 4 nthrl)ibtr GAB^-ery..: interncorcncitut slr\$) whnll ue!,naptir.llr ihhibits lLinttnisskn i.m qimatr aitet.-nt i.-]tive tC) te trnrls(rtNS) na(

(5j0lund TENSi5 nol ]l.:r/o.xore-re!,ersible seEmenta 9 a n dE r i k s s o In , 7 9 ) . AcuplmclLrre analgesia, on lhe olher hand, rs (Mayerel a/., 1977;Sjcjlund na/oxone-reversible & Eriksson, 1979;ChenS Pomeranz, & 1980), and we mustthus look for entifelydillerent explanations of ils elfecls. The Ab primaryailerents which can'ylhe pinpri.k messages lhe spinal cord make lheir !o principal intraspinal coniaclwith thc largc Waldcycr cclls in laminaI of the dorsalhorn (Kumazawa and Perl, l97u). fherc afe inhibitoryenkephalinergic "stalked cells"on the border amifae I and ll (cat: of llennett et a/., I982t mar]: Abclel Ma8uid and Bowsher,1984), with which co latcrals oi A5 primaryaifcrcnts makc synaptjc contact(Coblei a/., 1980). Rudaet al. \l9e'4)haveshownthatthcsc inhibiioryintcrneurons ablc to block thc trans are nrissiorl messages of arri!,inj along C polymodal primary r]ociceplive affcrcnts thc ccllsof originoi to lhe crossedasccnding artcrolatcral iuniculus.ln addilionto this postsynaptic effect,a prcsynaptic inhibitory cffectirom enkephalinergic interneufones of C on the terminals pfjmaryaffcrcnt fibreshasalso (Solodkin a/., 198.1). has been dernor]slrated ei Il also bccn shown that spinal intcrncuroncs not do reacl1ofrequncies stimulation of aboveabout lHz (Bowsher a/., l96U; Harpcrand Lawson,l9ll5). et Thesc mcchanisnrs quitcadcquatc cxplaif thc arc to seElmental mechanisms acupullclure of analgesia. At seSr.eftal level, both high-irec1uenc,v, lorvintensity analgesic stimulation (TENS) and low-irequency, hiBhinlensity annlgesic stimLrlalion (acLrpuncture) adeqLrately are explain-.d the gate by oi controltheory Melzack Wall(1965) and lfl8ule 2). TENSactspres),naptical11, lrt.,or'n* the Satefrom the outside,acLrpuncture pushingit sh!t from by
wrlf n-

Ho\,vever,both mechafisms of srmu arron produced analgesia .rlsoact at higher evelsof the neurrxis. In orrler to stuclythis, we must fifst exilrrrine the relev;nt ascending and descending pathways.

Iha Ba|. inta the .enttal ne^aus tyeenl ts opened Dy a ipatn)fibtes.|t... b." pa.tlr at )rhollv .los.:.1 by pullin:l t. innl outside. bv lrc ..1i\.rion oi l.tge lA[J)b\r threstrottliibres .".u TFN5, )ibtatian; thctr n(hanisns erc nat apiai.ler1i. lhe Eate .an )lsa be pusltetl b hr inhibita.y ner hani tms a(tint a,ithin the .-"ntrel ne^a6 syeen, sD.D .s i.upun.rute. deep bt.tn stinul,ito\ et..;
PEFIPHERY

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laminaV ir] lhc ncck oi the dorsalhorn.n snl.iller Ascending pathways oi anlerolalera iuni(ulu5 ImLow thfeshold mechanoreceptive prinrary conrponcnt thc .rsccnding thc vcntfobasal amus (Bo$rsher tha 1957). aif.'rents, carryinS kinclol iniorr.alionifduccd rcachcs the passup th-.dorsal.olLrrnr)s end rn thc fhis compon..nt 1(l aris.'sprincipallyfrom anrinaeI by TENS, gracileand cuneate nrclei. Fromthe dorsal urrlr and V (\,Villis al.,1979),and givcsoli collarera co ct s (gracle and cuneate) grc,v malier {PA(l) oi lhe nuc ej, nxonsem,.Ilieas lhe to thc pcrlaqucductal medial lemniscus.ind, all-"r dec!s\alirrgin thc midbran in its coursc'(Bo[,shcr ]9571. Manlyh passrostrally enclchieily in ( | 982)shows thisprojcction to that conrcs ffom .rmina I medullaoblongata, Ho\r,evet lhe mcdial of the spinalgrey matter;Pechufa and Li! (1984)) the ventrobasal anrLrstha sLrpplies sLrggested somespinoPACnc'urofes thnt alsorcsiclc lemniscus ofl Sives a rumber oi collatcfal l o n t h e w a t m o s t n o t n b y i n l h e m i d b r a i n . h c i f a m n a V l t i s i n l a m i n a cI a n d V l h a t l i n e i|l conncctionin which we nre rnoslinler-asle.l lhe myclinatcd(A5l prlmary .lffercntiibres lerninate present pleleclalnuclcus ( H e i m e r n d W a l , 1 9 6 8 ; R L ' t h e l y it d / . , 1 9 8 2 ) . a c contextis to the ar)lerior (Bjdr<eland nnd Boivie, 198.1;ltees and Roberts, Thrs there exist path[/ays rvhereby pinprick projecls lhe periaq!cducta ifformaticJn conveved the PAC(Fgul?J). lo is to l989al.This nucleLrs (PACIl (Rees grey m.rtter and llobcits 1989b),and inhibitorysystefr Descendingpathways thence activates the descending It was in 196.1 Tsolr that and Jang d..monstfatcd thal lvhichrsdiscrssecl ou,. be the PAC is the nDst effective spot n the wholc passes thc antero centralnervolrs ior inlofinaliorr up s),stem the abolitjonot pain by 2. High-threshold ,^s ateralfuniculLrs lhe side oppositcto that on rxrlfihe micrcinjection. the P,\C was known on -lhe r ' , , - r' l i r r ' - . : r l - r r l--, . r ' z r . media t \ r ' I l ' - u t N,hichit entersthe .oKl (<ur! 1949). bra nstemretic!larlornralionre.eiveslhe majofity find ng u,as convenientlyignored by \ /cstern 1957).It is nolv resc.archers. However,in 1969 Reyno cls, in the of the ascending axons(llolvsher thnt painless surgery could be canied known that these iibres originalelronr the deep USA,showed e electrical stifrulation thc oi s p i n a l a m i n . r ( V l l . n d V l l l )( K e v e l l e rl a / . ,1 9 8 2 ) , out on the rat dLrring e research possible on having been nctivated lfrolrBh lhe substantia PAC,and this led to intc'nsive in laboratories. gelatinosa by prinrary alierer]t unmvcInated frechanisms Western Invest lations (19/4) showcd thal a principa polynrodal which lenninale ly by Mayer and Liebeskind nociceptofs (SugiLra Perl,1986), descending pathrvay passing inhibitory dor,n frorr in lhe substantra and telatinosa part libfes also arise ironr Ihc caudovcntral of thc PAC to thc spinalcofd lhough some spinoreticLrlar

Ccntnl Dervous machanknsof pain coDtrol Pat, ,cair.1 rhesufsturnaFlatinas. iSC)af thL s]inal (td via unnvelinatcd Uinatr atiercntnociceptatslC). SC ceih .an be postsvnapti.allt inhibitpd hv e]k.phtlineryi. intlneu|'ies ltnk) The la et nrc ..1ivated .jjthL! ria ptinrt.v aita.e)t pinptkk iibtes(A6)a. vi.t lttotoninaryic 15 llf) fibtct d.:s.t:ndinL h thc ,1a6L,latuta1t'uni.Ltlus (DLF) ilonl the nu.lus Rad16 Magnut ltiRv) af the nedDlla abbnNdta fhe /.tr-.r is ,.tierted noni th.- p.ri,\rtc.ltrctal Ercynattcr iPACJin tha midbrain. Ih.: PAC t:an ba activatcd br splnathalani. IST) ilhrc
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MID B F AN

LOWER BRAN STEI\,4 i

SP]NAL CORD

ho.t hishet cent.es, by altete.t tinpti.k" iibta lt\ pirrhw!),r ,..ess/ble h tleep bt.ti, stnulatbn hn, L B., tlic patlrantticulatr4iot,, )nd lran t11,:) ptctaLtaltcabn in thc Th? prcte..t,jl rc$)n is c\.ilerJ by.dbtetrls ttun thL ntldial lannises atienB itl the da6al.:olunn nu.:kli (DCN); Lhctc in thcn [nn ar: a.tivatcd bl latSc ]a||lhteshoid plinary alt'ercrt iihrcs lAp) nevellingin the do\al.alunrB. The AB fibrcs,.n entty lib the iftl. risa e\.te ittltibiLatr CABA eryi. inteneuujes (C) whnlt prcsw)pticallt inhibit input to 5C. Ihus T[Ns and vibtation wa* thrcuih hoh a ,an-aplokk!&1. segn-pntdlne.hannn) lthe CAItA'eryK
'.ttt .,t.. . . 1t l . . . ) . lp' , ,,J" uptu-!-t t rL"t

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activatcdthtrnlgh nt,lial lcnniscal callatctals||hi.h rcach lt)l1ibitory nondtenet\n fibrcs des.eDd frotn the Lo.Dt aberrl-lus /N ) ar(l athet brain stett ..ell gtoups to exett dircd inhibtory attun on tlx: Substentia Cclatinasa ISC); tltk inhibition is nat ]re.liated ria enkpph;linetgic CABA ot

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was responsibje the inhibition of ascending ernotional psychicstateof the individuaJ for or may generated painful dependon the projectionfrom prefrontalcortexvia neurones carryin8messa8es by stimuli.lt is precisely caudoventral the PAC which hypothalamus PAC. to receivesthe ascendingspinothalamic coJlaterals (Mantyh, referred in the previous to section 1982). Dr David Bowsher Furthermore, thereis evidence that thereis a somtaMA MD PhD MRCP Ed FRCPath totopic orSanisation within this part of the PAC Pain Research Institute (Soper and Melzack,1982); this mightexplain why Pain ReliefFoundation heterosegmental acupuncture cannot be obtained RiceLane,Liverpool from any acupuncture point,but mustbe applied to particular points, not necessafilywithin the 1. .Abdel-MaSuid E, and BowsherD (1984)lnrerneurons T and dermatome wherepainreliefis to be obtained. proprioneurons the adult human spinal Srey matrerand in The pathway descendingfrom the PAC relays in g e n e r a ls o m a t i ca f f e r e n tc r a n j a l n e r v e n u c l e i . r , 4 n a r the nucleusRaph6MaSnus(nRM)of the rnedulla 139:9 0 2 oblonsala. probrblv b) '1eu oler.'ne.gi( ribrp. 2 . E e i t z A J ( 1 9 8 2 ) T h s i t e s o f o r i g i n o r D r a r n s r e m (Beitz,1982),at least in the rat. From the nRM, neurotensin serotoninprojections the rodentnuclels and to Raphe Magnus. Neuros.i.2: 829-14 I. mainly serotoninergic fibresdescend the clorso in ) E land , and M lateralfuniculusof the spinal cord to termtnate l . otj o r k eprojections E o i v i eJ ( 1 9 8 , 1A n a n a t o m i c a l the irom the doBal collnrn nuclei directly on enkephalinergic interneurones the at midbrainin the cat. Anrt. Embryol.lTAt 29-43 boundarybetweenlaminaeI and ll of the spinal 4. EloonrF E, Battenberg RossiefJ, E, Ling N, and Cuillemin R '1984) l q I \ F u o n | o , r ' I n 8 0 - - - d o , p L r in ' d dorsalhorn (Clazer and Basbaunr, in the cat. sparately from those containinE There are also serontoninergic nerve terminals lmm!nocytochemical nudies.Proc.Nrt/. Acad ending non synapticaliy the superficial in dorsal 75:1591 5 horn (Maxwellet al., 1983; Leranth a/., 1984; 5. BowsherD (1957)Termination the centra pain pa ef of Hammond a/.,1985). et conscious appreciation pain. Brain.8A: ot 22 l-rrdll),il \^ ll be re' allpd rha lhe PAC .e,piva\ d p-endorphinergic projection (Bloomet 5. Bowsher D (1976) Role of the reuc!lar formation descending r s p o n ste n o r i o u s i m u a I t o n .P a i n . 2 : 3 6 1 A o $ 7 a/-,1978)which turnsout to comefrom the arcuate Eowsher Ma la( A, PetitD, and Albe-Fessnrd (1968) D, D (Mantyh, nucleus the hypothalamus of 1982); and of bulbarrelayto centremedian./. Neurophysio/. 288-3 3I: projections coLrrse hypothalarnic this region receives B r o w nA G , R o s e K , a n d S n o wP G ( 1 9 7 8 ) o F h o k P M and organisation axon collater.ls frcm afferentfibres of from ihe prefrontal cortexin man. lt is because the slowly adaptingtype I units in cat spinal cord. /_ Phtsioi pathwayfrom hypothalamus PAC is endorphin to (Lond.) 277: -27 ergicthat analgesia man produced electrodes 9 . C h a n 5 H 15 ( 1 9 8 4 ) in by H What is beinS implantedin the PAC or in the periventricular acupuncture:evallation of the existence regionanterior it (Richardson, to l982), in which subnrate. Neu,oscl.Ae,5av. Rev.8t25 3f the hypothalamo PAC fibres run, is feversed by 10.ChengR S S, and PomeranzB H (1980)Electroacupunctr analgesia mediatedby stereospecific is opiate feceptors (Hosobuchi al., 1977;Richardson a Daloxone et and is reversed antagonists Type I receptoE.tlrreSci. by oi Akil,1977). 631 3A Withinthe PAC itself, thereare inhibitory interneu l l . C h i a n g C Y , C h a n gC I C h u H L , a n d Y a n eL F ( 1 rones which are inhibited by lonB inhibitory Peripheral afferentpathwayfor acupunct!rc anal8esia. s i n .1 6 . 2 1 0 1 7 afferents, thus accountingfor releaseof activity in n F E e the PAC-nRMpathwayand thus inhibitionin the 1 2 . D u 8 8 a A W a n d F o o n g W ( 1 9 8 5 ) i c u c u l l i na n d p i n h i b i t i o n r o d u c e d y d o r s ac o l u m ns t i m u l a t i oi n i n e b l spinalcordvia the inhibitory pathway. nRM-spinal Pain.22:249 59 Thus the pathwaydescending from the PAC is 1 3 . C l a z eE J , a n d B a s b a u rA | ( 1 9 8 4 ) x o n sw h i c hr a k e r n A responsible a numberof forms of stimulation (rH) serotoninarc presynaplic enkephaijnim for to tive neuronsin cat dorsalhat.. Bnin Res. produced 289:3a9-91 analgesia: 1. Via the dorsal columns and anteriorpretectal l 4 . c o b e l S , F a l l sW M , B e n n e t C J , A b d e l m o u m a n e H a y a s hH , a n d H u m p h r e y { 1 9 8 0 ) A nE . M .a n a l y s io f i E s (seeabove). (i) nucleus Thisexplains why analgesia synaptic connections ho6eEdish peroxidase oi filled stal can be obtained with stimulation the transected celh and islet cel s in the substanria of gelarinosa ihe of dorsalcolumns aboyethe levelof pain;and (ii)why car spinalcord./. .omp. neurcl. 194:7A1aA7 E the literatureis ambivalentabout the effect of 1 5 . C o u l d e n A ( 1 9 2 1 ) T h et r e a t m e not f s c i a t i c a y E a l b acuprnct]rc. qtit. Med.l. 1: 523-24 naloxone on TENS and DCS: the CABAergic 16.Hammond L, Tyle and T Efiecrs of segmental mechanismis rot nalo\one-reversibJe h y d r o x y t rDp t a m i ncnM,n o r aYaksh l L (198s) y ae d drena ininto spinal e becauseit is not opioiderBic, but the long loop . u L , 'r . . ' p d u i n g r 1 r u h r o n o I t h p r d m F o u d . . / . mechanism throughthe anteriorpretectal (Loncl.)359:151 62 nucleus and PAC is opiodergic and therefore naloxone- I 7. HarperA A, and LawsonS N (1985) Electrical prope.ties ral dorsa root ganglion neurones with diflerentpefiphe reversible, neNe condlction ve 359:47 63 2. Na/oxonereversible heterosegmental acupun- 1 8 .H e i m e rL , a n d W a l ocities.I. Physiol.lLand.) t PD (1968) hedorsal oo di T r cture works through the ascending colJateral ' . 1 , . u b r d r r B F I n d n o r Fo l o i projection PAC. spinothalamic to bution in the cat. frp. Neu,o/.6:89-99 , 3. Modulation of pain perceptionthrough the l 9 . H o s o b u c h i YA d a m s l E ,a n d L i n c h z R ( 1 9 7 7 ) P a i n . e l i e f r

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e l e c l r i . a5 l i m u l a l i o o f c e n t r ag r e ym . l l - A irn h ( m . r I a n d n i f h i l ) i l i o n n t r e c a ts p n a l . o . d . - ' . / c n . e . l , l : t . 1 5 t5 j i 2 . + l . S o p e r Y . . d M c l z a c kR 1 1 9 8 2 S t i n r u l a t i o n - l n o ( t u c c d its relcrsa by raloxore. Jc,-.r.e. /9;: l8l 81, W ) 20. Kveller A, H.ber L Fl, Yezie6k R P,Chungj M, A,lartiD C R a r i g c s i a :e ! i d e n . e r o r l o m a t o r o p i .o . g r n i s . r r i o fn L h e 6 , a n d \ 4 / i l l i s D ( 1 9 8 2 ) e ls o f o r i g h o t t h e s p i n o r cc W C l . ' ( b ' r l n . E ' " t r R e r .2 t l : l 0 l l 2 . 1 , 1 l 8 i u ' . Y , l e c C L , a i d P e r lE R ( 1 . 1 8 6 ) C - " n tp . r l i c . t i o n l ! ar rracrin th honkev /. .amp. Neunl 207 61 71 s r ro o 2 1 .K u m a z a w a, . n d P e , ]E R ( l 9 7 8 )E x c i t a t i oo i n ] . ] r s j n .a n d r . l i d e i l i i e d , ! n n v . l i n . l e d ( C l i i r i r c n r I h f e s i n r c ^ , . rn g r g s ! b s t a n t i a e a l i n o s ar e u r o n si n t h c p r n r a t - " p i n . ) l o r d : s c nranrnr.lr.r sl(in.s.icr.. 2l.l: -l-t8 6l r i n d l c ao n so l t h i rp a . - "i n d o 3 i r h o r nf u n . t i o n ao f E a n i s n - . + : . T o d d l , a n d M ( K e n z i . I ( l 9 l l 9 ) C r \ B A m m ! f o f e n . l r v e t l A I o n .I . o t 1 1 p .N c u t o l .I 7 7 : 4 1 7 - 3 1 n c l i o n s i n l h - "d o i i h o , r o i l h . r i l s p i f i r c o r d .N e ! , ? r r ( i . 22.Kur! M (19,19) sersoff paths in the .plnal (aft| and btain l2:799 8t)() , 1 6 l i o ! K , i n d l a n gC 5 ( r 9 6 . 1S l r d i e s n l h e s i t eo i r r i l g c s . rten oinir. Sogisrl,J,Tokyo ) o 2 S . L e r a n t C s , M a x ! v - " 1l J , r n d V c r h o f 5 l aA A I ( l 9 8 . l l h l d a c t i o n i m o ' r t l i n eb v i f t r a . - " r e l ) r n r i c r o i n j e c l i o5 i.- t s n r . o al n U trrstructurcol serolonn- nunoreacti\'bollons r lhe . 1 7 . W a lP D , a n d 5 w e e tW L l 1 1 9 6 r lT e m p o r i . b o l i l o f o i s!bsrantia glal nosa ol lhe rJtt spinal .otd. J. Ph$iol t tLand.) 155: 2DP p a i ni f n r a n s . , e r . e / - t - 51 0 8 , l . : , 1 8W . n 8 l ( M , Y a o5 M , X i i r , Y L , i f d H o u z { r 9 E 5 )A s l u d . v 2 : r . L i ! Y K , V i r e l n M , a n d O $ ! a d R ( 1 9 7 7 )T h e c o f l . s p o i d e n c eb e t \ l e e ns o . r e m o t o r p o i n t sa n . l a . l p u i c t L r r ei o c i . o l L h e r e c e pv c i . l d o i i . ! p o i n t s a n d t h e . e l . t i o n s h i p t Anet I. Chin. Me.l. J: )47-58 betwe-pn ch.rdcteiislcs ol Ie-"d iis sensar .n.l tll]|,ps oi oi ) 25.Lu.debcrg C M (198.1Vib,.torystinrulatiolor the T n altcrentl brer. 5.i sii.26:9()t /-1 Jll,"varlon oi chron c paif. A.t. Ph.rsiol. s.and. 'uppl. 521: 4 9 . [ / i l l s W D . K - " i s h . r l o R , a n d L e o n . ] dR B ( 1 9 7 9 )T h e D t 5l c e ls o l o r g i n o i t h e p ' i n , . r t e p n o t h a . . , i c t i d . / . . o r ) / ) s 2 6 M a n r y hP ! ! ( 1 9 8 2 )r ' - o r c b r ap r o j e c t l o i so t h e p c r i a q l e n t N..0tol.6u:54) 71 l dlctal grcy in rhe monkey,tr,irhobservalions the cat and 5 0 . \ b s h i oN ( 1 9 6 7 ) n t l o d ! . 1 i o rl o R y o d o r a k !/.1 . u p ! r . t D t q . ii l ttrr.I. conp. Neutot.206:116 58 z r . M r n t _ v h W ( 1 9 8 2 )T h - "a s c e n d g r p u t r o t h n r i d b r a i f P n periaqueductal I Ercy ol dre prim.rre. .Lrnp. Ncutol. 21t: 2 8 . M a \ s , eI l ) J , l - e r a n t C s , a n d V e r h o k t i dA A j ( 1 9 8 1 ) i f e h F s t r ! c t u r e l s e h i o n r n o n t an i n Sa x o n s n t h e n r a r S l r a lo n e o c i z o f t h c r a rs p n a l c o r d t h t i n R e s . 2 6 6 : 2 ) ia ' { ) 2 9 M a y c r D J , a n d L l - " b e s k i i J C ( 1 9 7 4 )P a i i r e d l c l i o nb y d ( i o c a e l e c t r i c a li m u l a l i o . o I t h e b r a i n :A n J , r a o f ( . , , i n o b e h . r v i o ! r ra n a l y s i s . r a l nR e s . 6 8 : 7 3 3 l A 9 : l 0 . M a y e fD l , P r i c eD D , a n d R a l i iA ( 1 9 7 7 ) n a t a S os m o i A f a . u p u n . l r r en r a E c s i a n m . n b y l h e n a r c o r i c n t a g o n i ( a n r l . \ o n e l i r . r i n e s ./ 2 / : 1 6 1 r 2 R i l l . M e z a . k l t , . r n dW a l l P D ( 1 9 6 5 J, . i r . , e c h r n i s m s : f e \ ! l A theory.s.ler.-p. 750:971 9 J2 Melzacr R, 5t llw-"1D M, Jnd For E I 11977) ris8er T p o i n t sa n d a c ! p ! n c l ! f e p o i n t st . j p a i n : c o f t ea t r o n s n d a m p l i c a t i o n s .i i , r l : l 2 l P J l . P e c h u f i C M , a n d L i ! R P C ( 1 9 8 6 ) p i n a ln e u r o n s h i c h r S w projecL to the perjrquedlclal Brey and the nred!l .ry r e t i . ! l a rI o r n ) a t i ov i a a x o nc o l l r l e r . l sA d o ! b l c l a b e lf l u o n , r s c e f . e r L d y n t h e r a t ., r a i n R - . s .a l : 3 5 7 6 1 s i J l . 1 . R e e ls , a n d R o b - " f lM H T ( 1 9 8 9 a ) A c t i v a t l o in e l s l n r h f l s . . i n t e r i o p r c t e c t r ln u c l ! sb v d o r s a lc o ! m n s l i . r L r l . r t i orn r the rat I. Physlal. 417: )61-71 1 5 .R e e s , i n d R o b c r t M H | 1 1 9 8 9 b 1 A r ro c i c p r l v--""f f e c t s H s n o i d o r s n . o l u n n i m l l a r i o ni n t h e r r t : i n v o l v m e n tl t h e o l a n t e r i op f e t e . t an u c l c ! s / . P h y s i o 1 t 7 : 3 7 5 A A r . l. 1 6 . R c l h e y i M , L i g n A R , a n d P e r L i l ( 1 9 8 2 )S y n a p t i c conrplcxsform-..]by f!nctionally deliie.l primary aftercnt ! n i t s w i t h f i n c m y e l i n a r idb e s . l . . o n p . N e u r c |2 0 7 : 3 8 1 i . l T . R e t n o l d sD V ( 1 9 6 9 )5 ! e r y i n l h e a r t d u r i n Ee e c l r i . a a n a l s e s lia d u c e d v f o . a l b r a n ( i m u l a t i o n .S c i c n c e1 6 1 : n b . l S . R i c h a r d s o n E , a f d A k i l l l 1 1 . r 7 7 l , . i f r e d u c l i o nb y D ) e e c t r l c ab r r i ns l m u l a l i o nn n r a n / . N e r r o s u r g .7 : I 7 8 9 4 l i 1 J 9 .R i c h . r d s o D [ { ] 9 u 2 tA f a l g s i a r o d u c c d t s r r n r ! t i o i o i n p b a ! a f i o ! s s i t s i n d r c h r m r n 0 - e n d o h i n s y s t m ., 4 p p l p Neurophysial. 116 2) 15: . 1 0 . R u dM A , C o i l i e l d , a n d D ! b n e r l t ( 1 9 8 4 ) D e m o n n r a t i o n a l of poslsynapt oploid nrodulatiool thirlamic role.tion c n p reurons by lhe .onrblncd te.hniq!es of rerro8rade e h o r s c r ar h p e r o x k l a s r n d c n l e p h ai n r n m r n o c y t o c h e m d i s t r y/.. N . u r o r . i ., 1 r l l 7 - 1 2 2 a l . S j o l l n d B H , r n d E r i k $ o n B E ( 1 9 7 9 ) T h f f l l e i c , " o l M i n a l o \ o . eo n a n a l g c s ip r o d ! c e d y p e r i p h c r a o n d i t l o n l n s a b .l s r i m ! l a l i o n6 , a i r R . s . 17 1 : 2 9 5 - l A 2 4 2 . 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T h e B R I T I S HM E D I C A L A C U P U N C T U R E S O C I E T Y a s o c i e t yo f m e d i c a l l yq u a l i f i e d is doctorswho practiseacupuncture have an or i n t e r e s ti n i t . T h e s o c i e t yh a s e x p a n d e dt o include sectionsfor dentists and veterinary surgeonS, The aims of the societyare to promote the s c i e n t i f i c n d e r s t a n d i n g n d p r a c t i c eo f u a acupuncture part of the normal practiceof as medicine. The administrative officer will deal with any enquiries including those regarding membership,coursesof tuition, text books, s c i e n t i f i cm e e t i n g sa n d t h e J o u r n a l o f t h e BritishMedicalAcupuncture Society. P l e a s s e n da l l e n q u i r i ets l e o THEADMINISTRATIVE OFFICER, B.M.A.S. Newton House. Newton Lane, Lower Whltley, Warrington, Cheshire WA4 4JA. Telephone: O925 730727 Faxt 0925 730492

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The physiology of stimulation-produced analgesia

David Bowsher Acupunct Med 1991 9: 58-62

doi: 10.1136/aim.9.2.58

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