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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 vcssclssurrounrledby their peflvas
path$,ay5, cular plerus of ncrVrvasorunr.Thnt sympathclic
nervesnray bL.involvcdw,asiirrl denronstrated
by
Couldef (1921) rvho showed thrt acupuncture
poifts alongthc sciali. n-of\,e
lntrcduction
anclits branches
had a
Tw,o types ol cLrtaneous
stimulirtionhave been lower impcclancc
ihan lhe sLrrroLrnding
sl(in.Yoshio
knor'vnfrom time immemorialto be analgesrc,
ly (1961)has showf lhat Ry.xlorakupo fts havc o\^,
Th-.
iirsl
is
instinctive
eifective.
the
action of impedance/high
conductance-Many acupLllcturc
"rulhing a pain L)-.ll-.r", ancl the second is pointsare of coufscdcep lo lhe skin:Melzac<ct a/
acupunct!re.
Rubbingis done rapidlyand lightly,scr (19//) havc shorvnthat Lheycorrelateclosclywitlr
physical
hi!:hfrequency,law
pollrls",while Liu eta/. (19/7) have
in
terms constitutes
T|avel's " tri.qtcrjs
performed
Acupuncture
\,"'ith shownthat othcrscorrespond
irtensit),
stimulation.
with the motorpoints
a needle,which is rotatedfairlyslow,ly{about2 of l ol mLtsclcs.
tinresa second),and so constituteslo\,! frequency, What typc oi receplof mtrst be stimulatedlo
pfoducc thc acupuf.lure effect?The facl lhal .r
h/ghlnten5lfy
stimulation.
li we acceptthatthesetwo forns of stimulatron
arc needleor sonreolher intensesti|rulusntuslbe used
empiricallyeifective,wc should cnquirc rrhich poifls to a high-thfeshold
reccptor(Bo\'\,sher
I976).
p t r r i o l o c , . p , l _ u .r .
' r d m ^ ,l . r ' n r a t , WanS e/ a/. (1985)sholvc'dthal Ab or Crout) lll
respons
blc for theiractions.
fillresarein iactlhe groupconccrncd.
Receptorsand primary afferents
Spinalcord nechanisms
Falr), evidently, rubbing activates lar8e lo$,- The fext questionto bc asked,ob!iolrsly,is whcfc
cutancousaflerenls(AB).Sin.e sLrstainedanclhou,do thesetwo typcsoi primaryJiierent(RA
threshold
pressure
is nol effectivein producinBpain reliei,the Ap anclA5) s\rnapsc
in the spinalcord?Foftunatcly
(RA),nncl recentadvafcesin anatomyand l)hvsiology
eflectiveLIlil\ musl be rapidly-adapting
can tel
ly, pain us a Sfeatclealaboutthcscqueslions.
excitedby tlansientslimuli. TherapeuLica
relef by aclivalion of low thresholclcutaneous PrimarvaffereftAp fibrcsenterthe spinalcod in
is obtainedby LranscutaneoLrs
afferents
electricirl the medialpaft of the dorsalrool .rndpassrostt'a
ly
,-.et,
(TENS)
(Wall
1967).lt withoutsyfapscin lhe dorsa (olumnsto cnd n the
nervestimLrlation
& S\
cana so be obtainedby the .rpplic.tionof vibrator), dorsal column (Sracie an(l clrneate) nuclei.
stmull (LundeberS,I9Bl), \ ,hich presumably However,on cntry 1o lhe spinalcord, thcsc iibres
(AB) primaryafferentsin give off a segmcrtalcol ater.rlw,hichpencttatcs
activaterapidly-a.laplinB
the
greymatteroi thc dorsa horf. Somearborsatrons
muscle.
ol
presentsa nrore clifficultproblem, this branchmakc contactin the superfcialparl of
AcupunctLrre
it was iirst necessary
to clerfonstrate
that the dorsal honr, probablythrough a CABA ergic
because
(To.ldan(l McKenzie,1989),w,ilhthe
nervesare invoiveclnt all. Halrpily,experments interncuronc
whichshorvedthat when r nerveis blockedw'ith tenninalsol small ir](omin8primaryaiicrenlIibres
is inefiectivein the (se.' Blown ct al., 1978) (Ftlure ir). Such ar1
localanaesthetic,
ncLrpunclure
would erplain thc CAIJAterritorysupplied by it, demonstratethat the afatomicalarrafBenrent
acupuncturceffect is conducted along ncrvcs crgic inhibilion observeclby Duggan ard Foong
(Chianget a/., 19/3). Also, rt was (nown that ( 1 9 U 5 )i o l l o w i r l Sd o r s a lc o l L r m ns t i m L r l a l i o( n
see
points
is only efectiveat cedain
on thc belo\,v),w,hich retrogradelydrivcs lhe segnrental
acupunciure
body suriace. In iact, comparisof with an .ol alerrlsresponsib
e for TENS.ll is oi courservell
anatomicalatlas shows that many acupurcturc know|l lhat these s),sternsrcspond to high
pointscorrespond
with the points at which smali lrcquencies
oi stimulirtion.
lvc thus have a circuit
nervebundlespenctratetre Iascia;Chan (198,1) .apabe oi explirinlng,nt scgme/)tal/eve/, thc
citestwo Chinesest!dies$/hich showe.lthal 109 analSesic
elfectsof TENSan.l villration.Note thai
poinlsare situatcdon or very closelo 111iscifclrit does not rrvo ve arv opiorderg
acupuncture
c
286 are on or Verycloselo synapses,
nerves,
whie a fLrrlhef
and therelbrcthc anal8,"!i.r
by
l)rocluccd
58
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(5j0lund
seEmentaTENSi5 nol ]l.:r/o.xore-re!,ersible
a n dE r i k s s o In9, 7 9 ) .
AcuplmclLrreanalgesia,on lhe olher hand, rs
(Mayerel a/., 1977;Sjcjlund&
na/oxone-reversible
Eriksson,
1979;ChenS& Pomeranz,
1980),and we
mustthus look for entifelydillerentexplanations
of
ils elfecls.The Ab primaryailerents
which can'ylhe
pinpri.k messages
!o lhe spinal cord make lheir
principalintraspinal
coniaclwith thc largcWaldcycr
cclls in laminaI of the dorsalhorn (Kumazawa
and
Perl, l97u). fherc afe inhibitoryenkephalinergic
"stalkedcells"on the borderof amifae I and ll (cat:
llennett et a/., I982t mar]: AbclelMa8uid and
Bowsher,1984), with which co latcrals oi A5
primaryaifcrcntsmakc synaptjccontact(Cob€lei
a/., 1980).Rudaet al. \l9e'4)haveshownthatthcsc
inhibiioryintcrneurons
are ablc to block thc trans
nrissiorlof messages
arri!,in€jalong C polymodal
primaryaffcrcnts
r]ociceplive
to thc ccllsof originoi
lhe crossedasccndingartcrolatcraliuniculus.ln
addilionto this postsynaptic
effect,a prcsynaptic
inhibitorycffectirom enkephalinergic
interneufones
of pfjmaryaffcrcntC fibreshasalso
on the terminals
(Solodkinei a/., 198.1).Il has
been dernor]slrated
also bccn shown that spinal intcrncuroncs
do not
reacl1ofrequ€ncies
of stimulation
aboveabout lHz
(Bowsheret a/., l96U; Harpcrand Lawson,l9ll5).
Thescmcchanisnrs
arcquitcadcquatcto cxplaif thc
seElmental
mechanisms
of acupullclure
analgesia.
At seSr.eftal level, both high-irec1uenc,v,
lorvintensity analgesic stimulation (TENS) and
low-irequency,
hiBhinlensityannlgesicstimLrlalion
(acLrpuncture)
are adeqLrately
explain-.dby the gate
controltheoryoi MelzackandWall(1965)lfl8ule 2).
TENSactspres),naptical11,
lrt.,or'n* the Satefrom
the outside,acLrpuncture
by pushingit sh!t from
b
9
I
d
ACU
Bated on Bo(she,1935 1937
Cinul t lat hi\h incnsit\, b|| ir\tuencr iA.upunctute) rrd
h ighJrcqoen.y Io\\' iDteDsiry i[N5] !tnru/.rtror.
PAC = Pctiaquachl.talBtar
,Ri!1 = nu./cus Ripha MrtrlJ
= 5ul,starnaOe/.rltnrr.
5C
=
T
Itansnission.el!
= Wald.yat c.:ll
W
CABA = yamino butyti. a.id
= s!bnir.. P
5P
VIP = Vataactlvcin]rsti'al paptidt:
= [nkcphalinetit. intetneuote
E
wrlf n-
L Fine mrelin,r.i.l Ai f)tinrn,- .t1epnl. are e.tivat€clby 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
sobsttnti. sel.tino (SC)neDrcneswhi.h rc.eire na.icaptir.:
ntput rni Dnntyeiinile(llC) Vini.]y affercDttusitlg SubnanccP
l5P) or rasaa(tne tntestiml Peptl'1elVlP) .1s i tnntmlttct.
(Set:mentaI a. upun.t trc ).
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.
Ho\,vever,both mechafisms of srmu arron
producedanalgesia
.rlsoact at higher evelsof the
neurrxis. In orrler to stuclythis, we must fifst
exilrrrinethe relev;nt ascendingand descending
pathways.
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.-ntna( i.-]tive tC) te trnrls(rtNS)
59
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
Ascendingpathways
anlerolaleraiuni(ulu5
ImLow thfeshold mechanoreceptive prinrary conrponcntoi thc .rsccnding
tha amus (Bo$rsher
1957).
aif.'rents,
carryinSthe kinclol iniorr.alionifduccd rcachcsthc vcntfobasal
1(lend rn thc fhis compon..ntaris.'sprincipallyfrom anrinaeI
by TENS,passup th-.dorsal.olLrrnr)s
gracileand cuneatenrclei. Fromthe dorsalco urrlr and V (\,Villis
ct al.,1979),and givcsoli collarera
s
(gracle and cuneate)nuc ej, nxonsem,.Ilieas lhe to thc pcrlaqucductal
grc,v malier {PA(l) oi lhe
medial lemniscus.ind, all-"r dec!s\alirrgin thc midbran in its coursc'(Bo[,shcr ]9571. Manlyh
conrcsffom .rminaI
medullaoblongata,passrostrallyto enclchieily in ( | 982)showsthatthisprojcction
the ventrobasaltha anrLrs-Ho\r,evet lhe mcdial of the spinalgrey matter;Pechufaand Li! (1984))
thntsomespinoPACnc'urofesalsorcsiclc
lemniscusSivesofl a rumber oi collatcfalsLrpplies sLrggested
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 .l 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 lhe myclinatcd(A5l prlmary .lffercntiibres lerninate
conncctionin which we nre rnoslinler-asle.l
presentcontextis to the ar)leriorpleleclalnuclcus ( H e i m e ra n d W a l , 1 9 6 8 ; R L ' t h e l ycit d / . , 1 9 8 2 ) .
(Bjdr<elandnnd Boivie, 198.1;ltees and Roberts, Thrs there exist path[/ays rvhereby pinprick
projeclslo lhe periaq!cducta ifformaticJn
is convevedto the PAC(Fgul?J).
l989al.This nucleLrs
(PACIl
(Rees
grey m.rtter
and llobcits 1989b),and
thence activatesthe descendinginhibitorysystefr Descendingpathways
It was in 196.1thatTsolrand Jangd..monstfatcd
thal
lvhichrsdiscrssecl
be ou,.
the PAC is the nDst effectivespot n the wholc
passes
ior the abolitjonot pain by
inlofinaliorr
up thc antero centralnervolrss),stem
2. High-threshold
,^s the P,\C was known
to that on rxrlfihe micrcinjection.
ateralfuniculLrs
on lhe side oppositc-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
However,in 1969 Reynocls, in the
of the ascendingaxons(llolvsher1957).It is nolv resc.archers.
surgerycould be canied
known that these iibres originalelronr the deep USA,showedthnt painless
s p i n a l a m i n . r (eV l l . n d V l l l )( K e v e l l eerl a / . ,1 9 8 2 ) , out on the rat dLrringelectricalstifrulationoi thc
research
on possible
having been nctivated lfrolrBh lhe substantia PAC,and this led to intc'nsive
in Westernlaboratories.
gelatinosa by prinrary alierer]t unmvcInated frechanisms
Invest€lations
polynrodalnociceptofs
which lenninaleprincipaly by Mayer and Liebeskind(19/4) showcd thal a
inhibitorypathrvaypassingdor,n frorr
in lhe substantra
and Perl,1986), descending
telatinosa(SugiLra
partof thc PAC to thc spinalcofd
libfes also arise ironr Ihc caudovcntral
lhough some spinoreticLrlar
Ccntnl Dervousmachanknsof 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.l€us 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
MID
B F AN
,.t
LOWER
BRAN
STEI\,4
i
SP]NAL
CORD
a.nt.
.tt...a t,n
r.."t..1.
-.
.a.t 1/
t..t..,..
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
}Y
-
nk'j
_
i
\ ,r-,j ,'-/
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'ercrtiihrcs 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
activatcdthtrnlgh nt,lial lcnniscal callatctals||hi.h rcach
lt)l1ibitorynondtenet\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;linetgicot CABA
Downloaded from http://aim.bmj.com/ on October 2, 2016 - Published by group.bmj.com
was responsibjefor the inhibition of ascending ernotionalor psychicstateof the individuaJmay
generatedby painful dependon the projectionfrom prefrontalcortexvia
neuronescarryin8messa8es
stimuli.lt is preciselythe caudoventral
PAC which hypothalamus
to PAC.
receivesthe ascendingspinothalamiccoJlaterals
referredto in the previoussection(Mantyh,1982).
Dr David Bowsher
Furthermore,
thereis evidencethat thereis a somtaMA MD PhD MRCP Ed FRCPath
totopic orSanisation
within this part of the PAC
Pain ResearchInstitute
(Soperand Melzack,1982);this mightexplainwhy
Pain ReliefFoundation
heterosegmental
acupuncturecannot be obtained
RiceLane,Liverpool
from any acupuncture
point,but mustbe appliedto
particular points, not necessafilywithin the
1. .Abdel-MaSuid
T E, and BowsherD (1984)lnrerneurons
and
dermatome
wherepainreliefis to be obtained.
proprioneuronsin the adult human spinal Srey matrerand
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
1 3 9 : 92 0
oblonsala. probrblv b) '1euoler.'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,
neurotensinand serotoninprojectionsto the rodentnuclels
RapheMagnus.I. Neuros.i.2: 829-14
mainly serotoninergic
fibresdescendin the clorso
l. Ejorkeland
M , a n d E o i v i eJ ( 1 9 8 , 1A) n a n a t o m i c a l
lateralfuniculusof the spinal cord to termtnate
ot the projectionsirom the doBal collnrn nuclei
directly on enkephalinergic
interneurones
at the
midbrainin the cat. Anrt. Embryol.lTAt 29-43
boundarybetweenlaminaeI and ll of the spinal 4. EloonrF E, BattenbergE, RossiefJ,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 (Clazerand Basbaunr,
in the cat.
s€parately from
those
containinE
There are also serontoninergic
nerve terminals
lmm!nocytochemicalnudies.Proc.Nrt/. Acad
ending non synapticaliyin the superficialdorsal
7 5 : 1 5 9 15
horn (Maxwellet al., 1983; Leranthef a/., 1984; 5. BowsherD (1957)Terminationof the centra pain pa
Hammondet a/.,1985).
consciousappreciationot pain. Brain.8A:
22
l-rrdll),il \^ ll be re' allpd rha lhe PAC .e,piva\ d
projection(Bloomet 5. Bowsher D (1976) Role of the reuc!lar formation
descendingp-endorphinergic
r € s p o n steo n o r i o u s$ i m u a I t o n .P a i n . 2 : 3 6 17 A
a/-,1978)which turnsout to comefrom the arcuate
EowsherD, Ma la( A, PetitD, and Albe-Fessnrd
D (1968)
(Mantyh,1982);and of
nucleusof the hypothalamus
bulbarrelayto centremedian./. Neurophysio/.3I: 288-3
projections
coLrrse
thishypothalarnic
regionreceives
B r o w nA G , R o s eP K , a n d S n o wP G ( 1 9 7 8 )M o F h o k
and organisationof axon collater.ls frcm afferentfibres
from ihe prefrontalcortexin man. lt is because
the
slowly adaptingtype I units in cat spinal cord. /_ Phtsioi
pathwayfrom hypothalamus
to PAC is endorphin
(Lond.)
15-27
ergicthat analgesia
in man producedby electrodes 9 . C h a n 5277:
H H (1984) What is beinS
implantedin the PAC or in the periventricular acupuncture:evallation of the existence
regionanteriorto it (Richardson,
l982), in which
subnrate.Neu,oscl.Ae,5av.
Rev.8t25 3f
the hypothalamo
PAC fibres run, is feversedby 10.ChengR S S, and PomeranzB H (1980)Electroacupunctr
analgesiais mediatedby stereospecific
opiate feceptors
a
Daloxone(Hosobuchiet al., 1977;Richardson
and
is reversedby antagonistsoi Type I receptoE.tlrreSci.
Akil,1977).
631 3A
Withinthe PAC itself,thereare inhibitoryinterneu 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
Peripheralafferentpathwayfor acupunct!rc anal8esia.
s i n .1 6 . 2 1 0 1 7
afferents,thus accountingfor releaseof activity in
n W a n d F o o n gF W ( 1 9 8 5 )E i c u c u l l i naen d
the PAC-nRMpathwayand thus inhibitionin the 1 2 . D u 8 8 a A
i n h i b i t i o np r o d u c e d
b y d o r s acl o l u m ns t i m u l a t i oi n i n e
spinalcordvia the inhibitorynRM-spinal
pathway.
Pain.22:249 59
Thus the pathwaydescendingfrom the PAC is 1 3 . C l a z eEr J , a n d B a s b a u rAn | ( 1 9 8 4 )A x o n sw h i c hr a k e
responsible
(rH) serotoninarc presynaplicto enkephaijnim
for a numberof forms of stimulation
tive neuronsin cat dorsalhat.. Bnin Res.289:3a9-91
producedanalgesia:
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
i , a n d H u m p h r e yE { 1 9 8 0 ) A nE . M .a n a l y s iosf
nucleus(seeabove).Thisexplains(i) why analgesia
synapticconnectionsoi ho6eEdish peroxidase
filled
can be obtainedwith stimulationof the transected celh and islet cel s in the substanriagelarinosaof ihestal
dorsalcolumnsaboyethe levelof pain;and (ii)why
car spinalcord./. .omp. neurcl. 194:7A1aA7
the literatureis ambivalentabout the effect of 1 5 . C o u l d e nE A ( 1 9 2 1 ) T h et r e a t m e not f s c i a t i c ab y E a l
acuprnct]rc. qtit. Med.l. 1: 523-24
naloxone on TENS and DCS: the CABAergic
16.
Hammond
L, Tylec M, andYaksh
T L (198s)Efiecrs
of
segmentalmechanismis rot nalo\one-reversibJe h y d r o x y t rD
y p t a m i naen d n o r a d r e n a l i n ien t o s p i n a l
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 . . / .
mechanismthroughthe anteriorpretectalnucleus
(Loncl.)359:151 62
and PAC is opiodergic and therefore naloxone- I 7. HarperA A, and LawsonS N (1985) Electricalprope.ties
ral dorsa root ganglion neuroneswith diflerentpefiphe
reversible,
neNe condlction ve ocities.I. Physiol.lLand.)359:47 63
2. Na/oxonereversibleheterosegmental
acupun- 1 8 .H e i m e rL , a n d W a l P D ( 1 9 6 8 )T h e d o r s a lr o o t
di
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
projectionto PAC.
spinothalamic
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 hr z R ( 1 9 7 7 ) P a i n . e l i e f
6l
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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 .
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a c t i o no i m o ' r t l i n eb v i f t r a . - " r e l ) r n
a rl i c r o i n j e c l i o5ni.- t s n r .
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s!bsrantia g€lal nosa ol lhe rJtt spinal .otd. J. Ph$iol
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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 pvt 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
Anet I. Chin. Me.l. J: )47-58
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ttrr.I. conp. Neutot.206:116 58
z r . M r n t _ v hP W ( 1 9 8 2 )T h - "a s c e n dn g r p u t r o t h € n r i d b r a i f
periaqueductalErcy ol dre prim.rre.I .Lrnp. Ncutol. 21t:
2 8 . M a \ s , eI l ) J , l - e r a n t C
h s , a n d V e r h o k t i dA A j ( 1 9 8 1 )F i f e
s t r ! c t u r eo l s e h i o n r nc o n t an i n Sa x o n si n t h e n r a r S l r azlo n e
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 ' { )
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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 ral n a l y s i sA. r a l nR e s . 6 8 : 7 39 3
: 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 )A n a t a S osf m o i
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l l . M e z a . k l t , . r n dW a l l P D ( 1 9 6 5 Jl , . i r . , e c h r n i s m sA: f e \ !
theory.s.ler.-p. 750:971 9
J 2 M e l z a c r R , 5 t l l w - " 1 D M , J n d F o r E I 1 1 9 7 7 )T r i s 8 e r
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 sa n d
m p l i c a t i o n sP.i i , r l : l 2 l
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a n a l s e s lian d u c e db 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 :
l S . R i c h a r d s o nD 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
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Neurophysial.15: 116 2)
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62
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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
SOCIETY
i s 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
doctorswho practiseacupunctureor have an
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 cu n d e r s t a n d i n ga n d p r a c t i c eo f
acupunctureas part of the normal practiceof
medicine.
The administrativeofficer 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
BritishMedicalAcupunctureSociety.
P l e a s see n da l l e n q u i r i etso l
THEADMINISTRATIVEOFFICER,
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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