Experiments on the Section of the Glossopharyngeal and Hypoglossal Nerves of the Frog, and

Observations of the Alterations Produced Thereby in the Structure of Their Primitive Fibres
Author(s): Augustus Waller
Source: Philosophical Transactions of the Royal Society of London, Vol. 140 (1850), pp. 423-429
Published by: The Royal Society
Stable URL: http://www.jstor.org/stable/108444
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[ 423 ]

XX. Experimentson the Section of the Glossopharyngealand Hypoglossal Nerves of

the Frog, and observationsof the alterationsproduced therebyin the Structure
of their Primitive Fibres. By AUGUSTUS WALLER, M.D.

Communicatedby

ProfessorOWEN,F.R.S.
ReceivedNovember22, 1849,-Read February21, 1850.

THE objectof thepresentobservationsis to describecertainalterationswhichtake

place in the elementaryfibresof the nerve afterthey have been removed fromtheir

connectionwiththe brainor spinalmarrow.

ofthleopinionsand researchesof modernphysioThe followingis a briefsummary
logistson alterationsof thenerve-tubes.
BURDACH* placed a ligatureon the sciatic nerveof a frog,and afterthelapse of
fibreseitherabove or belowtheligature.
a weekfoundno alterationoftheprimitive
STEINRUCKt did the same,and statesthatin threecases thewholenervewas more

slender than on the healthyside, and ascribed it to the atrophyof the neurolema.
fibresin the
VALENTINtdenies likewise that thereis anyalterationoftheprimitive
lower portionof the nerve.
GUNTHERand SCHON?,whose researches are most interesting,state thatthe primitive fibresbeing examined towards the end of a week, after division of the nerve
it was perceived that they had no longer the full
when it had lost its irritability,

roundappearanceofthesoundones. Here and theretheircontentsappearedas if

days aftersectionthesestructuralchangesbecame
curdled; fromeightto fourteen
still moreevident,and continuedto increaseuntilthe fibresappearedflat,broken

up, entirelylosing theirtransparency,theircontentsappealing as if disjointed.

flstates,that fivemonthsaftersectionof the sciatic nerveof a

frog,the
tubesbelow thesectionwere brokenup into granulesand small clumps; that all
were strongly
the nerve-tubes
granulated,in some the small granulesbeing united
intooval bodies,which appearedto be surroundedby a pale cylindricalmembrane,
whichin somewas wanting,owingprobablyto its disorganization.
to the Royal Societydescribedthenervesof
Having in a formercomtnunication
NASSE

the papille and of the muscular fibrein the frog'stongue in their normal condition,
* Beitragzur Mikroskopisclier
Anatomieder Nerven,E. BURDACH. Konigsberg,1837, p. 42.
De
Nervorum
Regeneratione.Berolini,1838, p. 72.
t
NervorCerebral,&c. Lib. iv. 1839, p. 127.
t De Functionibus
iiberRegenerationder Nerven,p. 276.
MULLER'S
Archiv,
1840, Versucheund Bemrerkungen
?
Ueber
die
der
nach ihrerDurchschneidung,
MULLER, Arch. 1839, p. 409.
11
Verdnderungen Nerven-fasern

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424

DR. A. WALJLER ON THE ALTERATION

OF THE

it is myintention
at presentto describevariousalterations,
as seen underthemicroscope,which take place in the structureof the same nervesaftertheircontinuity
withthebrainhas beeninterrupted
ofthe frogstongue
by section. The innervation
is, as I have alreadyshown,derivedfromtwo pairsofnerves,one arisingfromthe
brain,and traversing
a foramenin theposteriorpartof thecranium,accompaniedby
thepneumogastric
nerve. This paircorresponds
to theglossopharyngeal
in Man. In
its courseit descendsuntilit reachesthe hyoglossusmuscle,whenit is accompanied
by the lingualvesselspassingover partof the hyoidbone, and enteringthetongue
withoutgivingoffany branchto thethroat. The otherpair arisesfromtheanterior
partof thespinalmarrow,traverses
thefirstcervicalforarnen,
and constitutes
thefirst
cervicalpair of nerves. Followingthe exampleof BURDACH, I regardthis pair as
to the hypoglossal
in Man, and shallapplythattermtoit. It takesits
corresponding
coursetowardsthetonguetina similardirectionto thatoftheformer
pair,givingoff
severalbranchesto the musclesoftheneck and throat,and whenit reachesthe.hyoglossusmuscleit is considerably
smallerthantheglossopharyngeal.Afterattaining
thismuscleit runsparallelto the formernerve,passingbelowthe hyoidbone in its
transitto thetongue. For a moreminutedescriptionI mustreferto the paper of
E. BURDACH, of whicha translationhas appearedin theAnnalesdes SciencesNaturelies.
Division of the GlossopharyngealNerve.

That divisionof thesenervesproducessome seriouslesion is provedby the death

of the animal,whichgenerallytakesplace a fewdaysaftertheoperation. Considering the well-known
tenacityof lifepossessedby theseanimalsthiswas quitean unexpectedresult,forwhichI am unable at presentto affordany satisfactory
reason.
We can onlysurmise,thatbesidestheirgustatorypowers,theyhaveothersconnected
withrespiration,
in regulating
theactionofthetonguein closingthenares,forforcing
the air intothe lungs. Whatevermaybe the trueexplanation,it is impossiblenot
to regardthis resultwith surprise,whenwe considerthe seriouslesionswhichthis
animalis capableofundergoing
at otherpointsofthe framewithoutloss oflife, The
usual timewhich it survivesis variable,and dependsgreatlyon the seasonof the
year. If the examinationtakes place in summer,death frequently
ensues on the
fifth
day; if in winter,not beforethe twentieth.For the purposeof avoidingthis
loss oflife,I adopttheplan ofdividingtheglossopharyngeal
on one side onlyofthe
tongue,and I findthat it has thedesiredeffect
of preserving
thelifeof the animal,
whilewe can observethe same alterationson thecorresponding
side,as well as when
bothnervesare divided. Anotheradvantagefoundin the divisionof a singlenerve
is, thaton theuninjuredside we have constantly
at hand a meansof comparisonby
whichwe can judge withcertainty
respectingany alterationsthatmaybe produced
in thedividednerve. In cases of any doubt,it will alwaysbe foundof thegreatest
serviceto examineat thesame timea minutefragment
fromeach side ofthetongue.
The firsteffects
ofsectionofa glossopharyngeal
nerveat the throat,are decreased

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PRIMITIVE

NERVE FIBRES

AFTER SECTION.

425

power of movingthe tongue,diminishedsensibility,

generallyveryslight on the
dividedside,and symptoms
indicativeofsomedisturbanceofthenutritive
functions.
The diminution
of motorpoweris veryslight,as is evidentby the almostmolecular
tremorwhichstillexistsin any partirritated,
and bythe capabilityofretracting
the
tongue. The loss ofsensation,which.is also veryslight,arisesfromthesectionofa
fewsensitivefilaments
whichare containedin the glossopharyngeal
nerve,and are
distributed
extremities.The lesionsofnutrition
principally
about thetubercular
and
circulationon theside of the divisionare veryvariableand uncertain. Sometimes
that half is cedematous,particularly
towardsthe tubercle. Sometimesthe papilla
are muchinjectedand congested,while in othercases this side is morepale than
can be detectedbetweenthetwo sides,
theother. In manyinstancesno difference
untiltheorganhas been slightlyirritated,
whenon thedividedside thevessels,and
of
those
the
become
especially
fungiform
papillv,
congestedand ofa deeperred than
causes independent
of
on the other. Some of these differences
probablyarise from.
the nervouslesion,as the vessels of thetonguewhichaccompanythese nervesare
doubtlessinjuredin someof theexperiments.
During thefirsttwo or threedays aftersection,no alterationin the textureand
of the tubes of the papillarynerves can be detected. Generally,at
transparency
the end of the thirdand fourthday, we detect the firstalterationby a slightly
turbidor coagulatedappearanceof the medulla,whichno longerappearscompletely
to fillthetubularmembrane,
whichdoes notappearto be affected.These alterations
to whicha littledistilledwaterhnasbeen
of themedullaare best seen in a fragment
hoursafterdeath,
added to renderit moretransparent.When examinedtwenty-four
the
betweentheseand the nerveson
the difference
healthyside is still more evident. Commencingdecompositionon the healthyside causes the nerve-tubesto
so as to attainnearlydoubletheirordinarysize. On thedivided
swell considerably,
side the disorganizednerveretainsnearlythe same size and appearanceas when
fresh. Causticpotash,whichdissolvesall the tissuesexceptthe nerves,rendersthe
the morphological
and consequently
alterednervesmoretransparent,
changesare less
made simultaneously,
we may
apparent. Nevertheless,
by comparativeexperiments
betweenthenervesofthetwosides. In somecases, in about
stilldetecta difference
threeor fourdaysaftersection,I have tracedtheturbidstateof thenervefromthe
fungiformpapillaeintobranchescontainingfortyor fifty
tubes,whereit did not apbut wheretheopacityof thenervepreventedmyobservingit any
pear to terminate,
in the
further.Aboutfiveor six daysaftersection,the alterationofthenerve-tubes
or
more
of
much
kind
become
a
papillaehas
distinct,by
coagulation curdlingof the
medullaintoseparateparticlesof varioussizes. Sometimesthecoagulatedparticles
have an unevenspongyappearance,as if the componentparts of the medulla,i. e.
thewhitesubstanceand the axis cylinder,were mixedtogether. Oftentheyappear
effused
fromthe
merelylike separatedparticlesofthemedulla,suchas are frequently
ends ofa dividednerve,and presentthe doublecontourand thecentralnucleuschaMDCCCL.

3 I

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426

DR. A. WALLER

ON THE

ALTERATION

OF THE

racteristicof the nervousmedulla. In somecases the coagulatedparticlesare very

uniform
in size and appearance,averaging--looth
ofan inch. In others,thelimits
are fargreater,
betweenthe maximumand minimum
dimensions
namely,from2-5-0
0th
to ,2>0thofan inch. The diameterof the alteredtubes,examinedin theordinary
mannerin water,is about a fourthsmallerthanthatof thesoundones,and in many
instancesthetubularcylindersappearwanting,and the medullaryparticlesto have
escapedfromthecylinder,and to be merelyheld togetherby theneurilemawhich
surroundsthe whole nerve. Afterthe applicationof potash,the diameterof the
alteredand unalterednervesis as nearlyas possiblethesame. This equalizationof
the two is producedalmost entirelyby a decreasein the size of the sound tubes,
contractbythe applicationofthe
in water,and afterwards
whichswellconsiderably
of size at thisstage betweenthe
alkali. It is therefore
probablethat the difference
the
from
arises
former
not absorbingso muchwithin
alteredand unalterednerves
themas thelatter. Whetherthisarisesfroma rupturedstate of themembraneor
froma chemicalchangeofthe medulla,is not evident. Afterthesurrounding
tissues
offers
no signsof rupture,
have been removedby potash,thetubularmtnebrane
and
the medullaappears less disorganizedthan beforethe denudation. The disjointed
A portionofeachnerveconditionofthemedullais greatesttowardstheextremities.
so disorganizedas to be carriedaway amongthetissuesdissolved
tubeis frequently
ofthetubes
presentedbytheextremity
by thealkali. The circularrimso frequently
around
often
observe
the
We
nervous
and thepapillary
is absent.
healthy
branches,
a commonsheathor neurilemna
nervein particular,
fitting
closelyto thenerve. After
disorganizationhas attainedthis degree,it appears to forma kind of loose pouch
around the nerveand separatedfromit by an intervalof- 0th ofan inch. This
ofthecurdledmedullaryparticles,which,
pouchappearsto formthe sole investment
as we have stated,previousto theactionofthealkali,appearvoid ofanytubularinvestment. As we ascend towardsthebrainthedisorganization
appearsto decrease,
the coagulatedmedullais moreapt to assume the oval form,and at some places it
unaffected.The effectof decomposition
in
presentsits double contourapparently
In
in
the
former
to
that
is
similar
nerve
altered
and
unaltered
the
stage. consequence
nerveis moreopake thantheunalteredone. In
of theabovechangesthedisorganized
of the tubes belongingto commonsensation
the tuberclessomeoftheramifications
afterthe sectionofthecorresponding
These
becomedisorganized
glossopharyngeals.
alterationstake place in the same periodand in thesame manneras in the papillary
nerves.Withthisexception,all the othernervesofthe tongue,whichcomprisethose
of commonsensationand ofmuscularaction,remainunaltered.The muscularfibres
of the papille are slightlyaltered at this period; theirtransversestriv are notso
distinctas on thehealthyside,whiletheirlongitudinalonesare moreso. The fibre
itselfis usuallypaler,narrowerand morewavyin its course.
The capillariesare eithermuch congestedwithdarkblood,or theyare completely
emptyand scarcelyto be detected.

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PRIMITIVE NERVE FIBRES AFTER SECTION.

427

and theciliaryfilaments
are unaltered.
The epithelium
On theseventh,
eighthand ninthdaysthedisorganization
ofthenervousstructure
continuesto progress. In thepapillaethe curdledparticlesof medullabecomestill
moredisconnected,
and in partsare removedbyabsorption.The tubularsheathalso
is rupturedand disorganizedneartheextremities
of the tubes. In theotherramificationsoftheglossopharyngeal,
medulla
the
becomesmoreand moredisjointedand
collectedintooval or circularcoagulatedmasses.
On thetenthday and upwardswe perceiveanothermorphological
stateof themedulla. The coagulatedparticleslose theiramorphousstructureand assumea granulated texture. The granules,retainedtogetherby slightcohesion,are dark by
transmitted
light,but of a lightwhitecolourby reflexion,
and average 100th
of an
inch.
Aboutthe twentieth
day themedullaryparticlesare completelyreducedto a granularstate. The conditionofthepapillary
nerveis represented
in PlateXXXI. fig.2,
wherewe findthe presenceof the nervouselementmerelyindicatedby numerous
black granules,generallyarrangedin a row like thebeads of a necklace. In their
it is easyto detectthewavydirection
ofthenerves. They
characteristic
arrangement
are still containedin the tubularmembrane,
whichis but veryfaintlydistinguished,
probablyfromthe loss ofthe medullaand fromatrophyofitstissue. The resistance
of thesegranularbodies to chemicalagentsis mostremarkable,fortheyremainunaffected
by acids, alkaliesand the ethers,whichhave so greatan influenceover the
nervousmedulla. Thesegranulesmay be detectedwithinthe papillarynervesfora
considerableperiodoftime. I have seen themapparently
unalteredin thepapillae
upwardsoffivemonthsafterdivisionof thenerve,reunionnothavingtakenplace.
Divisionof theHypoglossalNerve.
When the hypoglossalnervesare dividedat theirexit fromthespine,all movementsofthethroatand tongueare abolished,and theprocessof respiration
entirely
at an end. The tonguemay be drawnfromthe mouthremainingcompletely
inert,
pinchingor cuttingcausingno appearanceof pain. Hence we mayconcludethat
thisnerveis ofa mixednature,containingsensorialas well as motorfilaments.Anotherexperiment,
whichshowsthismoreplainly,consistsin dividingone hypoglossal
nervenear the spineon one side, and on the otherthe glossopharyngeal
at its exit
fromthecranium. Pinchingand othermodesofirritation
cause no painon the side
wherethe hypoglossalis divided,whileon the otherundoubtedpain is caused.
Afterdivisionof the hypoglossalsat the throat,the motorand sensorialpowersof
thetongueare not entirely
lost. Whendrawnout of themouth,the extremity
onlyremainsfixedbetweenthejaws, flaccidand powerless. At theinnerhalfthefibresare
stillcontractile,
on accountof theirbelongingto thehypoglossus,
whichat its lower
halfreceivesa branchfromthehypoglossalnerveabove thepointofsection.By this
3 I2

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428

DR. A. WALLER ON THE ALTERATION

OF THE

forthe same
means,aided also by the mylohyoidmuscle,whichis leftunaffected
powers.
reason,the innerhalfofthetonguestillenjoyscontractile
Respirationis hurriedand laboured,and deathis the invariableresultofdivision
theanimals
ofthesenerves,whether-madeat the spineor at the throat. In summer,
died at theend oftwoor threedays.
half of
Divisionofa singlehypoglossalonlycauses paralysisof the corresponding
whenat the
the tongue,completewhenthesectionis near the spine,and imperfect
throat. The animalsgenerallysurviveafterthesectionofone nerve.
of the hypoglossalnervesare mosteasilyfoundat the
extremities
The peripheric
surfaceof the tongue. By removinga minutefragment
at thisregion,we
inferior
ofnerves,whichare grapreparation,
ramifications
can observe,withoutany further
duallyreducedto a networkof singletubuleson the surface,among the capillary
either
network.At thesame time,amongthemuscularfibresare otherramifications,
directionor runningparallel.Like theformer,
theyare
crossingthemin a transverse
all
directionswithoutforming
anyfreeends. It
reducedto singlefibres,runningin
is at theundersuifacethatthealterationsof thehypoglossalmustbe studied.
Duringthefourfirstdays,aftersectionof thehypoglossalnerve,no changeis observedin its structure. On thefifth
day the tubesappear morevaricosethan usual,
and themedullamoreirregular.Aboutthetenthdaythe.medullaformsdisorganized,
masses at intervals,and wherethe whitesubstanceofSCHWANN cannotbe
fusiform
whichare mostevidentin the singletubules,maybe
detected. These alterations,
days manyof thesingletubules
foundalso in the branches. Aftertwelveor fifteen
have ceased to be visible,theirgranularmedullahavingbeenremovedby absorption.
The branchescontainmassesofamorphousmedulla.
We are naturallyled to inquire,whetherextraneouscircumstanceshave any inwhenthe
fluenceoverthe'removalofthetissue. We findthatin thesummer-time,
be
must
inoreactive,in consequenceof the inconsiderably
renewalofthe tissues
of
the
animals
and activity
upon,that the alteraexperimented
creasedrespiration
tionis morerapidthanin winterwhentheyare in a stateoftorporand hybernation.
our observationsto thealterationswhichtake place
At presentwe have restricted
in the ramifications
originatingfromtwotrunks,but we cannotsupposethatthisis
and thatothernervesdo not participatein similaralterations,
a local phenomnenon,
mustbe excluded.
the
brain
that
and
itself,composedin greatpartof tubularfibres,
thatsuch is not thecase,
enable me to affirm
lxperimentson othernervesaldready
and thattheyare to be.foundon othernerves,suchas thesciatic &c.,and, moreover
thattheyare as extensiveas thenervoussystemitself. It is impossiblenot to antiresultsfromtheapplicationofthisinquiryto thedifferent
nervesof
cipateimportant
to nervousdiseasesthatit
withreference
theanimalsystem. But it is particularly
to
extend
one
convictionimpressesitself
theseresearches. If
will be mostdesirable
on the mindthananother,it is thatwhatwe termfunctional
diseasesof
morefirmly

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PRIMITIVE

NERVE FIBRES

AFTER SECTION.

429

the nervesare in realityowing to certain organic and physicalchanges in the

.tubularfibre,whichit willbe theprovinceof themicroscopeto ascertain. If a fewv
days' inactivity
ofa nervoustrunk,such as is producedby ligatureor section,is sufficientto cause such disorganizationof themedulla,howcan we refuseto admit of
its beingalteredin cases ofprolongedparalysis?
October27, 1849.
Kensington,

DESCRIPTION

OF THE PLATE.

PLATE XXXI.
Fig. 1. Papillarynerveoffrog,six daysafterligature.
Fig. 2. Papillarynerve,threeweeksaftersection,withmuscularfibresin theinterior
ofthecapillarycoil at thesummitofthefungiform
papilla.
Fig. 3. DisorganizedMuscularnerve,fromthe inferior
surfaceof the tongue,five
daysaftersection. The muscularfibrehas been omittedin thisdrawing.

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