S P O N S O R E D D O C U M E N T F R O M

CORTEXAJOURNALDEVOTEDTOTHESTUDY
OF THE N ERVOUS S YSTEM AND B EHAVIOR
Cortex.2015Jan62:182202.

PMCID:PMC4298656

doi:10.1016/j.cortex.2014.10.023

Thewhitematterofthehumancerebrum:PartITheoccipitallobebyHeinrich
Sachs
StephanieJ.Forkel,a,b,,1SajedhaMahmood,c,1FrancescoVergani,c,dandMarcoCatanie
a
UniversityCollegeLondon,DepartmentofPsychologyandLanguageSciences,ResearchDivisionofClinical,EducationalandHealth
Psychology,London,UK
b
Natbrainlab,DepartmentofNeuroimaging,InstituteofPsychiatryPsychologyandNeuroscience,King'sCollegeLondon,London,UK
c
DepartmentofNeurosurgery,RoyalVictoriaInfirmary,NewcastleuponTyne,UK
d
InstituteofNeuroscience,NewcastleUniversity,NewcastleuponTyne,UK
e
Natbrainlab,DepartmentofForensicsandNeurodevelopmentalSciences,InstituteofPsychiatryPsychologyandNeuroscience,King's
CollegeLondon,London,UK
StephanieJ.Forkel:stephanie.forkel@kcl.ac.uk:stephanie.forkel@gmx.net

Correspondingauthor.Natbrainlab,DepartmentofNeuroimaging,InstituteofPsychiatryPsychologyandNeuroscience,King'sCollege
London,London,UK.Email:stephanie.forkel@gmx.net,Email:stephanie.forkel@kcl.ac.uk
1
Theseauthorsequallycontributedtothemanuscript.
Received2014Sep6Revised2014Oct31Accepted2014Oct31.
Copyright2014TheAuthors
ThisisanopenaccessarticleundertheCCBYlicense(http://creativecommons.org/licenses/by/3.0/).
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copyediting,andpublishing.TheU.S.NationalLibraryofMedicineisresponsibleforalllinkswithinthedocumentandforincorporatingany
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availableforfree,onScienceDirect,at:http://dx.doi.org/10.1016/j.cortex.2014.10.023

Abstract
ThisisthefirstcompletetranslationofHeinrichSachs'outstandingwhitematteratlasdedicatedtothe
occipitallobe.ThisworkisaccompaniedbyaprologuebyProfCarlWernickewhoformanyyearswas
Sachs'mentorinBreslauandenthusiasticallysupportedhiswork.
Keywords:Whitematter,HeinrichSachs,Wernicke,Postmortemdissection,Occipitallobe
1.Introduction
Bornin1863,HeinrichSachswasaGermanneurologistandneuroanatomistwhoobtainedhis
specialisationinneurologyandpsychiatrywithCarlWernickeinBreslau(Forkel,2014).Sachspublished
onamyotrophiclateralsclerosis(1885),aphasia(18931905),andtraumaticneurosis(1909),butarguably
hismostdistinctivecontributionwasinthefieldofwhitematterneuroanatomy.Whilststilladoctorin
traininghespentmostofhistimelookingatseriesofcrosssectionsobtainedfromhumanbrains.This
painstakingeffortresultedinthepublicationofthefirstatlasoftheoccipitallobeconnectionsinthe
humanbrain(Sachs,1892).Sachs'satlascontainsdetaileddescriptionsofthemethodologicalapproaches
heemployed,whichmakesthetextnotalwaysaneasyreadingbutthefiguresarebeautifullyinformative
andincludemanypreviouslyundescribedtracts.Forexample,Sachsdefinedthreedifferentlayersinthe
deepwhitematteroftheoccipitallobes,whichhecorrectlyidentifiedasfibresofthesplenium(forceps
corporiscallosi,seepage15),opticradiations(stratumsagittalisinternum,seepage18)andthe
associationpathways(stratumsagittalisexternum,seepage20).Thesewerelaterindicatedwiththename
stratumsagittalisofSachsinrecognitionofhiswork.Healsointroducedanewnomenclatureforthevast
numberofUshapedfibresrunningnearthecorticalsurfaceoftheoccipitalcortex.Theknowledgeof
thesetractshaddirectclinicalrelevanceasdifferencesbetweenapperceptiveandassociativevisual
agnosiacouldbeexplainedintermsofprimaryvisualcortexdamageanddamagetoassociativeUshaped
fibres,respectively(Lissauer,1890).Incontemporaryneurosciencewehaveunderstoodthatwithinthe

occipitallobetheseUshapedfibresmediatecrosstalkbetweentheventralvisualstreamdedicatedto
objectsperception(thewhatpathway)andthedorsalvisualstreamdedicatedtoplacelocationand
motionperception(thewherepathway).
Sachs'mentorWernickewasanenthusiasticadvocateofhisanatomicalinsightsandencouragedhis
traineetofurtherpursuethisresearch.Theatlaswasinfactintendedtobeamultivolumeprojectinwhich
subsequentbookswouldhavebeendedicatedtothefunctionandclinicalcorrelatesofeachtract.Thiswas
anambitiousprojectinthefootstepsofthegreatclinicalanatomistsofthetime.Unfortunately,Sachsdid
notcompletewhathehadsetouttoaccomplishandneverreturnedtohismasterplaninthefourdecades
hecontinuedworkingasphysicianattheneurologyandpsychiatryclinicinBreslau.
Despiteitsimportance,Sachs'satlaswentunnoticedfordecades.Thisisinpartduetotheavailabilityof
moredetailedinformationonconnectionalanatomyderivedfromaxonaltracingstudiesperformedin
animals.AlsothelackofanintegraltranslationfromGermantoEnglishdidnotfacilitateits
dissemination.
WebelievethatwiththeadventofnovelMRIbasedmethodstostudyconnectionsinthehumanbrain,the
workofSachscouldbeofgreatrelevancetocontemporaryneuroscience.Thisisparticularlytrueforthose
tractsthatmayunderlieuniquelyhumanabilities.TheverticalfasciculusofWernicke,forexample,
connectsrelevantareasforreading.Sachsdescribesthistractindetailandcreditshisdescriptionto
Wernicke(seepage26).Despitethistractbeingoneofthelargestintraoccipitalconnections,itsfunction
hasremainedunknown.Morerecentstudiesinpatientswithlesionstothiswhitemattertractoritscortical
projectionssuggestthatitmayhavearoleinreading(Yeatman,Rauschecker,&Wandell,2013).Other
tractsdescribedbySachsarestillwaitingtobeascribedaspecificfunctionalcorrelate.
Sachs'soccipitaltractshavebeenrecentlyreplicatedusingpostmortemKlingerdissection(Vergani,
Mahmood,Morris,Mitchell,&Forkel,2014).Detailedtractographystudiesareneededtocharacterisethe
invivoanatomyofthesetractsintermsofinterindividualvariabilityaspreviouslyshownfortractsof
otherlobes(Catanietal.,20072012Forkeletal.,2014LopezBarrosoetal.,2013).
2.Translation
InMemoriamtoDr.HeinrichLissauer,AssistantatthepsychiatricclinicinBreslau.Publicationsfromthe
psychiatricclinicinBreslau.
Thewhitematterofthehumancerebrum.Part1.TheOccipitalLobebyDr.med.HeinrichSachs,
neurologistinBreslauwithaprologuebythemedicalofficerofhealthProf.Dr.C.Wernicke,including3
figuresand8plates.
3.ProloguebyWernicke
Thepresentworkisthefirstcontributiontoaseriesofpublicationsdedicatedtotheinvestigationofthe
brainanditsfunctionsinhealthandpathology.Thisfieldofresearchisstillheavilyunderinvestigatedand
nearlyeverycontributiontoitisastepforwardsimilartoanexpeditionintounknownterritorycomparable
tothedeepestAfrica.Theintegrationofclinicalobservationsandanatomicalaspectshasconstantly
proventobeareliablemethodtomoveforward.Theadvancesinanatomy,whicharenaturallyslow,will
befollowedpromptlybyourclinicalexperience.
Theanatomyofthewhitematterofthecerebrumalwaysintriguedmeasthelinkbetweenalldelicate
clinicalmethodshence,IappreciatewithgreatsatisfactionthatourcolleagueSachsmadesuchan
encouragingstartwiththepresentwork,whichisofthehigheststandardintermsofitscontentand
structure.Mayfuturepublicationsbeequallywellreceivedbycolleagues.Breslau,January1892.
4.Sachs'atlas
Thisworkcanbeconsideredasthefirstpartofamoreextensiveworkonthewhitematterfibretrajectory
inthehealthyadulthumanbrain.Thedissectionspresentedherewereobtainedinthepsychiatricclinicin
Breslau.IshalltakethelibertytoexpressmygratitudetowardsProfessorWernickeforkindlygrantingme

permissiontoundertakethisworkandforhissuggestions.Further,Ithanktheassistant,Dr.Lissauer,for
hisfriendlyandactivesupport.
Theaimoftheworkistoprovideamacroscopicoverviewofthefibreconnectionsoftheoccipitalcortex
aswellasadjacentpartsoftheparietalandtemporallobes.Detailsandsubtletiescanbeaddedtothiswork
inthefuture.
Informationonthewhitematteranatomyofthecerebralhemisphereisrelativelyscarce.Inordertogainan
overviewofthisfieldonehastogobacktothebeginningofthecentury,namelytoBurdach'sgreatwork
(181918261822),asfibretrajectoriesareonlyhintedatinmorerecenttextbooks.TheworkbyMeynert
(1884)isdifficulttounderstandandisnotentirelyevidencebased.Furthermore,theavailablecasereports
arebasedonpathologicalspecimens.Foundationworkdemonstratingthewhitematteranatomyinthe
healthyadultbrainisentirelymissing.However,inordertoassigneachcasereportitsaptplaceinthe
system,thehealthyhumanbrainshouldbethereferenceforallotherstudiesofpathological,foetal,and
animalbrains.
Identifyingthedirectionalityandtrajectoryoffibreswithinthewhitematterusingonlyasinglemethodis
insufficientaseachmethodhasitsinherentlimitations.Inordertoovercometheselimitationstheresults
ofatleasttwomethodsoughttobecompared.Forthecurrentstudythreecomplementarymethodswere
thereforeemployed:
1. TheoldestmethodisZerfaserung[postmortembluntdissection],whichwasusedexclusivelyby
BurdachandhonouredinparticularbyMeynertandhisstudents.Forthecurrentwork,Iusedbrains
thatweretreatedwithalcohol,yetwerenottoohardened.ThemethodintroducedbyStilling(1882)
whichusesHolzessig[woodvinegar]returnsbrilliantresultsforthebrainstembutwas,however,
notsuitableforthewhitematterofthehemispheres.Thedifferenceisthatforthisworkitisnot
importanttosegmentsmallpartsofthebrainintoitsfibrepathwaysbuttorelatetheoverall
directionoffibresandconnectionsbetweenwhitematterbundleswithinalobe.Incontrast,blunt
dissectionsreturnperfectresultsifthemajorityoffibresarerunningalongthesamedirection,whilst
theubiquitouscrossingfibresarenotformingsubstantialbundlesbutarepresentinisolationor
smallnumberswhenpiercingthroughthemainpathways.Insuchcasestheywouldfallapart
smoothlyoronedoesnotnoticethematallunlessalreadyfamiliarwiththem.Additionally,the
presenceoflargefibrecrossingscanbeidentifiedusingthismethod.However,itisnotpossibleto
followwithconfidencethetrajectoryofthefibresbeyondtheirpointofconvergence.Further,if
fibresthatthusfarruninparallelstartadheringtoeachother,asitisthecaseforcallosalfibres
towardsthemidline,thismethodwillalsofail.Inboththesecasestearingthetissuecancreate
arbitraryartefacts.Coarsecrossoversarenotfoundintheoccipitallobeandmatting[occurs]onlyin
thecorpuscallosum.
Themostimportantdrawbackofthemethodisthatitonlygivesustwodimensionalviews.Thedirection
andthewidthofalayercanonlybeidentifiedwithcertaintyifthelayerisentirelydestroyed.Therefore,
bluntdissectionsareonlyfordemonstrationinthiscasetheyareobviouslyinvaluabletoappreciate
specialorganisationandrelationshipsbuttheyareneversufficientasevidenceintheirownright.
2. ThesecondmethodistheinspectionoffreshlypreparedsectionsofspecimenshardenedinMller
solutionandobservedunderdirectlight.Thesesectionsshowthefibresorlayerscuthorizontallyas
purewhitewithonlyahintofgreen.Areaswherefibresarecutobliquelyappearblackgreenandare
darkerintheirshadethanthedarkgreencolourofthegreymatter.Betweenthesetwoextremesall
shadesofcolourscanbefounddependingonthecuttingangleinrelationtothedirectionofthe
fibresandwhetherthemajorityoftheexposedfibreswerecutstraightorobliqueinregionsof
multiplefibreorientations.Additionally,differencesinfibres,suchastheirwidthorthechemical
natureofthemyelinsheath,influencethetoneofcolour,sothatthevariouslayerscanbeclearly
differentiated.Ifthebrainiscutalongdifferentdirections,thesamelayercanappearlightordark
dependingonthedirectionofthecutting.
3. Themostimportantmethod,thatallowsmagnificationandinspectionofindividualfibresisthe

methodintroducedbyStillingandperfectedbyMeynert,whereaseriesofsequentialtranslucentand
stainedsectionsisprepared.TomyknowledgeonlyPal'sHaematoxylinissuitableasstain.In
comparison,whensectionsof1/20mmthicknessarepreparedusingWeigert'sstainthewhitematter
appearsevenlyblue,thusmakingitimpossibletodifferentiatethevariouslayers.Thepreparationof
sectionsasthinas1/10mmforthewholehemispherewasonlypossiblewithgreateffort[asit
requires],timeandskill.Extremelyrareisthesuccessfultransferontoaglassplateafterthe
specimenwasexposedtothestrainsofstainingandbrightening.Incontrast,1/10mmsectionswere
suitableforstainingaccordingtoPal.Pal'sstainhastheadvantageofstainingsomefibresverydark
anddeepblue,whilstotherfibresareonlylightlystainedinbrownglow.Hence,itispossibleto
differentiatethem.Inthebrainstemfivedifferentshadesofcolourcanbedistinguished.These
shadesdependpartlyonthevariablethicknessofthefibresandpartlyonthechemicalpropertiesof
themyelinsheath.Anotheradvantageofthismethodis,thatsectionsstainedwiththismethodcan
bephotographed.Thephotographsenclosedtothiswork[8platesattheendofthispublication]are
anexampleofsuchsectionsandwereprovidedbyDr.Lissauer.
ThePicrocarminestainallowsidentifyingvariouswhitematterlayerswiththenakedeyeandthenuclei
canbeseenunderthemicroscope.StructuresthatareusuallycoloureddarkandbluebyPal'sstainare
stainedyellowbypicrocarmine.WhatappearslightandbrownusingPalisreddishwithpicrocarmine.
Thedrawbackisthatinbraintissue,unlikeperipheralnervesorcord,theaxonalconesarenotdistinctly
stainedinredthereforetheindividualfibrescannotbedifferentiated.
Note:WhenusingPal'sstainforlargespecimens,suchasasectionofthewholehemisphere,amultitude
ofstratagemsarerequiredandnegligenceofeachofthemendangersthefinalresult.Ishalltherefore
carefullydescribethemethodbelow.
Thebrainisremovedfromtheskullassoonaspossibleafterdeath,ideallyinthewinterandthen
preservedinMllersolutionasawholeoronlycutinhalves(toavoidlosingitsshape).Inthefirstfew
days,thesolutionneedsdailychanging.Thespecimenisreadytobecutafterthreetofourmonths.Slices,
cutasthinasthemicrotomeallows,aredriedbysoakingthemindilutedalcoholandpurealcohol,eachfor
aperiodof24h.Slicesarethenimmersedincelloidinsolutionandstucktowoodenplates.Forthe
sectionsIusedthelargestSchanzmicrotomeandanespeciallydesignedheavyknife.Ididnotcutunder
spiritus.Slicesof1/10mmthicknesscanbepickedandtransportedeasilyifnotyetstained.Ifthebrainis
rathercrumbly,thesurfacecanbecoveredwithcollodionorcelloidinbydrippingonathinlayerofthe
solutionpriortoeachcut.Theslicesareplacedwithoutcopperinwaterfor24handsubsequentlyina
1%haematoxylinsolution(Haematoxylin1,alcoh.Abs5,ofwhich5ccmonto100ccmwaterand1ccm
saturatedlithiumcarboniumsolution)forthesamelengthoftime.Onecansimultaneouslystain10or20
slicesinalargeamountofsolution,butthesamesolutioncannotbeusedtwice.Theslicesarethenwashed
withplentyofwateranddestaineditisbesttoletthemsoakinwaterforaperiodof24h.Theycan,
however,beleftinwaterforlongerwithoutanyconcerninwhichcasetheslicesonlydestainfaster.The
individualsliceisthenplacedontoaglassplateorinaglassdishwithfattymarginsandispouredonto
witha.51%manganeserichacidicpotassiumsolutionandgentlyturnedaroundmultipletimes.The
solutionhastobechangedrepeatedlyandisonlyactivelydestainingaslongasitshimmersbluishwhen
heldagainstawhitepaper.Assoonasthebluecolourischangingtowardsviolet,thesolutiondoesnotde
stainanylonger.Onthecontrary,itratherstainspermanentlybrown.Consequently,ifthesectionisnot
stronglydehydratedfollowingthestainandcontainsrestsoffreehaematoxylin,oneswiftlyhastodiscard
thefirstportionofthepotassiumsolution.Incaseswherethemarginofasectionistransparentandfreeof
blackstainswhenitisheldagainstsunlightorabrightflame,thesectioniscarefullywashedwithwater
andpouredontowithanacidicandideallyhotsolution(Ac.Ocalic..5,Nat.sulfuros..5,Aq.200).The
sectionisthengentlyswunginthesolutionuntilthemarginisperfectlywhiteandstainfree.Ifnecessary,
theacidsolutioncanbechanged.Shouldstainsstillpersist,onehaseithertheoptiontobesatisfiedwith
theresultorotherwiserestarttheprocesswithpotassiumsolutionafterwashingthesliceinwater.A
repetitionisalsoadvisableifthestainingwasveryintenseandthelayersarethusnotdistinguishableafter
thefirststaining.Insuchamanner,destainingcanbecarriedtotheextreme.Themoredestainingis
carriedoutthebrightertheentireslicebecomes.Thishoweveralsoappliestothedelicatefibres,especially

corticalfibres,whichcanbedestainedtothepointwheretheywillfade.Ifaslicethatistoobrightand
brownitcanbestaineddarkerandbluewhencoveredinalkalinesolution,anammoniasolutionor
carboniclithium.Theslicefromnowonwardsplacedonanobjectslideisdriedinabsolutealcoholand
thecelloidinisremovedwithetheralcohol.
Iftheslicewascoveredwithcelloidinpriortocutting,itisbesttomakesurethatthesideoftheslicethat
wascoveredwithcelloidinisplacedfacedownonthestage.Itisthenlightenedincarboxylox(ac.Carbol.
2.Xyl.6.).Onedrainsthecarboxylolalittleandpressesatleasteightlayersofblottingpaperquicklyand
stronglyontheslice.Theuppermostpageofblottingpapershouldnotbecomewet,aspartsoftheslide
willsticktoit.ThesliceisthenpouredoverwithwarmorXylolthinnedCanadabalmandcoveredwitha
thinglassplate.Duringmicroscopy,itisbesttolookwithoutapertureusinganAbbmicroscope.
Thecortex,whosewhitematterconnectionsaretobedescribedhere,isdelimitedanteriorlybyafrontal
plane[fr],whichpassestangentialtotheposteriorendofthesplenium(Figs.1and2).Thenatural
boundaryforthewhitematteroftheoccipitallobe,theconfluenceoftheposteriorhorninthecella
lateralisofthelateralventricletheopeningoftheposteriorhornliesjustbehindthisplane.
Ontheconvexityofthemedialsurfacethisplanecutsthemostanteriorpartoftheprecuneus(Fig.2).On
thelateralconvexity(Fig.1)itcutsthegyrusattheendoftheSylvianfissure[supramarginalgyrus],
whosemostposteriorcorticalindentationextentsintothedepths.Onthelateralconvexityofthisthree
sidedpieceofbrain,twosulcicanbeseenrunningdorsoventrally[e,k],andthreesulcirunningposterior
anteriorly[s.o.IIII],whichallimpactontheshapeoftheunderlyingwhitematterduetotheirdepth.The
moreanteriorsulcus[e]ofthetwoverticalsulciistheascendingbranchoftheparallelsulcus[superior
temporalsulcus](Fig.1e).Thisascendingbranchliesentirelywithintheparietallobeandisconsideredas
partoftheangulargyrus.Theadjacentposteriorverticalsulcusistheanterioroccipitalsulcus[posterior
intermediateparietalsulcus](kseeWernicke(1881)).Thissulcusconsideredrepresentingtheborder
betweentheparietalandtheoccipitallobes.Thissulcuscanappearindifferentshapes.Usually,it
continuousventrallyintothecontinuationofthesuperiortemporalsulcus[e]andthusgivesrisetoa
secondascendingbranchofthelatter.Attimes,however,itappearsasaveryshortindentationwithout
connectiontoanyothergyri.Itis,nonetheless,foundineverybrainandisreadilyidentifiable,when
followingtheoccipitoparietalsulcus(o)ontheconvexity(Fig.1)totheinferiortransitionalgyrus(above
k)(Fig.1)betweentheparietalandtheoccipitallobes.Theopeningofthisgyrusistheanterioroccipital
sulcus.
Withintheoccipitallobetherearethreedeepsulcithatarealmosthorizontaltoeachotherbeforethey
separateanteriorly(Ecker,1869).Thesuperior/firstoccipitalsulcus(s.o.I)isanextensionofthe
intraparietalsulcus(i),whichusuallyreachestheoccipitalpole,thoughinterrupted.Themiddle/second
occipitalsulcus(s.o.II)reachesanteriorlytowardsthehorizontalbranchofthesuperiortemporalsulcus
(e).Theinferior/thirdoccipitalsulcus(s.o.III)runstowardsthesecondorthirdtemporalsulcus.The
inferioroccipitalsulcusoftenrunsadjacenttotheinferiorconvexityofthehemisphereandsometimes
evenatthebasalsurface.Themiddleoccipitalsulcuscorrespondsmostlytotheloweroccipitalsulcusof
Wernicke.Whereasbothverticalsulciandthefirsthorizontalsulcusareconsistentandreadilyidentifiable
themiddleandinferioroccipitalsulciareofteninterruptedandbranchoff,andarethereforelessclear.
Theoccipitallobeisdelineatedonthemedialsurfaceofthehemisphere(Fig.2)bytheoccipitoparietal
sulcus[o]separatingthecuneusandprecuneus,andbythecalcarinefissure(f.c.),whichadheresanteriorly
withtheabovementionedsulcus[o].Bothsulciarerarelysimpleincisions.Usually,theirstemformsa
surfacesimilartotheinsulawithsecondarygyri.Nevertheless,thismorphologyisvariable.Theposterior
incisionoftheoccipitoparietalsulcusmayextendmanycentimetresintotheoccipitallobe.Adjacentto
thecalcarinefissureashortgyrusextendingrostrocaudallycanbeseensuperimposedonthetopand
bottomsurfacesfacingeachother.Inthedepthofthefissurethreeverticalshortgyriextenddorso
ventrally.Twoofthesecancontinuetotheconvexityofthesulcusandmergewiththeabovementioned
gyriwhereasthethirdsulcus,thatisthemiddleortheposterior,neverextendstotheconvexity.Sucha
shortgyruscanreachattimestheconvexityandthusinterruptthefissure.Thecalcarinefissure,similarto
theSylvianfissure,hasasuperior(s)andinferior(i)opening(Fig.3)butnoposteriorextension.Rather

thebaseofthefissurebecomesflattenedandcontinuesontothemedialsurfaceofthehemisphere.This
transitionistobeseenintheforkingofthefissureposteriorly.Withinthecuneus,agyrusparalleltothe
calcarinefissureextendsrostrocaudally[cu,Fig.2].
Intheprecuneus,thehorizontal,posteriorlydirectedextensionofthesulcuscallosomarginalis(cm,Fig.2
)[cingulatesulcus]isimportantforwhitematteranatomy.
Onthebasalsurface,themostimportantsulcus,whichshapesthewhitematteristhecollateralsulcus
(coll.,Fig.3),whichisthefifthsulcustoextendcaudorostrallybetweenthecalcarinefissureandthe
inferioroccipitalsulcusandisvariableinitsextensioninbothdirections.Themedialoccipitotemporal
sulcusreachesverycloselytheoccipitalpole.Incaseswherethecalcarinesulcusisasimpleincision,the
medialoccipitotemporalsulcuscanpresentacomplexdivision.
Theoccipitalhornbeginstoformasacanalwithfourwalls,withthindorsalandventralwallsandtwoto
fourfoldwidermedialandlateralwalls.Posteriorly,itrapidlyloosesitsshapeinalldirections.Initially
thelossisprimarilyinheightmorethanwidth,sothatitresemblesalmostasquarebeforeitloosesits
widthandthusbecomesathinsulcuswithitsdorsalandventralwallsturnedintoedges.Duringitscourse
itbendsposteriorlyintwodirections.Initsposteriorpartitbendsgentlyalongaverticalaxisandthusits
posteriorendcomestolieclosertothemedialplanethanitsaperture.Inaddition,itbendsalongasagittal
axisandbecomesaslit,thusbringingthedorsalandventraledgesclosertothemedialplane.Fromits
posteriorendastripofependyma,whichretainsitsform,continuesintotheoccipitalwhitematterfora
shortdistance.Thedoublebendofthehornresemblestheformofthehemisphericconvexityandisdueto
thedeep[occipital]notchclosetothecalcarinefissure.Apartfromthis,onlythemedialoccipitotemporal
sulcushasanimpactontheshapeoftheoccipitalhorn,bybulgingitsinferiorsurfacealittleinthemiddle
partofthehorn.Alltheothersulci,includingthesecondarydeformationsofthecalcarinefissure,areofno
importancetotheshapeoftheoccipitalhorn.Theseinfluencethewidthofthewhitematteronly,andasis
latertobeseen,thethicknessofthefourthandoutermostlayer,whichliesimmediatelyunderneaththe
cortexandisreferredtoasthestratumpropriumcortices.Thedeeperlayersofthewhitematterare
independentofthedepthofthesesulci.
Theoccipitalhornliesclosertothebasalsurfacethantothedorsalconvexityofthehemisphere(Fig.3)
yet,itisequidistantlylocatedbetweenthemedialandlateralsurfaces.Nevertheless,duetothedepthofthe
calcarinefissureitisseparatedfromthecortexatthemedialsurfacebyathinlayerofwhitematter.The
majorityofwhitematter,ontheotherhand,developsatthelateralaspectoftheoccipitalhornbetweenthe
latterandthehemisphericconvexity.
Thefibresoriginatingfromtheoccipitalcortexandcoursingwithintheoccipitalwhitemattercanbe
dividedintotwogroups.Amongstthesegroups,onecanagainsubdividethreegroups:i)fibresthatextend
tosubcorticalcentresandareconsideredasprojectionfibresorcoronaradiata(Stabkranz)(Meynert)ii)
otherfibreshavetheirterminationsincorticalareasandarethereforeassociationfibres.Associationfibres
eitherinterconnectintralobalcorticalareas(shortassociationfibres),orlinktheoccipitalcortexwiththe
cortexofadifferentlobe(longassociationfibres)iii)thethirdgroupcrossestheinterhemispheric
midlineandmightterminatein[contralateral]corticalorsubcorticalareas(callosalorcommissuralfibres).
Thismassoftheoccipitallobefibresisnotatetheredbundle,butisratherorganisedintobundlesand
layersaccordingtocertainrules.Theselayerscanbedistinguishedbasedontheirdirection,groupingand
staining.Thelawoforderisthefollowing(Wernickeascitedabove,p.24):Everyfibrereachesits
destinationviatheshortestpossibleroute,asfarasthisisincorrespondencewithembryological
peculiaritiesofbraindevelopment.Thus,thefollowingtwoconclusionscanbereached:First,shortfibres
arelocatedclosetothecortexwhilstlongerfibresarelocatedclosetotheventricle.Second,fibreswith
roughlythesamedestinationruninparallelorformbundlesforapartoftheircommontrajectory.
Asecond,generallyvalidbiologicallawnottobeignoredinthestudyofbrainstructureisthelawof
variability.Therearenotwobrainsthatareidenticalinalltheirdetails.Variabilityisalsoobservedinthe
arrangementanddevelopmentofwhitematteranatomy.Thecortexandthewhitematteraremutually
dependentoneachother.Ifaparticularareaofcortexisunderdevelopedinabrain,thenthereisalsoa

paucityoffibresoriginatingfromthisarea.
Theoccipitallobefibresformfourlayers,whichenveloptheoccipitalhornlikeanonionskinfromall
sidesexceptitsopening.Theselayers,countedoutwardsfromthemedialtothelateralwallsofthe
ventriclesare(Fig.3):
1. Layerofthecorpuscallosum:Forcepscorporiscallosi(110),a.parsmagnasuperior(1),b.pars
parvainferior(4)
2. Layeroftheprojectionfibres:Stratumsagittaleinternum(1114)
3. Layerofthelongassociationfibres:Stratumsagittaleexternum(15)
4. Layeroftheshortassociationfibres:Stratumpropriumcortices.
a. stratumcalcarinum(16)
b. stratumtransversumcunei(17)
c. stratumpropriumcunei(18)
d. stratumverticaleconvexitatis.
.stratumpropriumfissuraeinterparietalisseealsosulcioccipitalisI.(19)
.stratumpropriumsulcioccipitalisII.(20)
.stratrumpropriumsulcioccipitalisIII.(21)
.stratrumprofundumconvexitatis(23)
e. stratumpropriumsulcicollateralis(22)
f. stratumpropriumpraecunei.
Thislayerisfoundintheregionoftheparietallobe.
Anothertwobundlesarecloselylocatedtotheoccipitallobewithoutjoiningitswhitemattersystem,
namely:
5. BogenbndeloroberesLngsbndel,fasciculusarcuatusseealsolongitudinalissuperior[superior
longitudinalfasciculus]
6. Zwinge,cingulum.
Fibresoriginatingfromtheoccipitalpoleandsurroundingareasbundleupinthemiddleofthewhite
matterandrunanteriorposteriorly.Farbeforetheventricularhorn,thesefibresgrouptogetherintothree
concentriclayers.Theinnersolidlayergivesrisetotheforceps.Theremaininglayers,whichonaxial
sectionslooklikearing,formtheinternalandexternalsagittallayers.Theforcepsappearsclearlyasan
independentlayer,afewmillimetresanteriortotheothertwolayers.Thus,inthesmallstretchthat
includesthestratumpropriumcorticis,onlythreelayersareevident.Thethreelayerswithasagittal
directionthickenanteriorlyasnewcorticalfibresfromalldirectionsjointhem.Atthecaudalendofthe
posteriorhornthesefibrespartlikeafunnel,sothatallthreelayersequallycovertheposteriorhorn.
5.Forcepscorporiscallosi
Iftherewasnoposteriorhorn,theoccipitallobewasasolidstructure,andthecalcaravisdidnotreach
deepintothewhitematterofthelobe,thentheforcepswouldhavetheshapeofaconewithitsheadatthe
occipitalpole,wherecorticalfibreswouldgatherlikeraysequallyfromallsides.However,nowabulgeof
thelateralventricle,whichformstheposteriorhorn,pushesintotheforcepsfromthefront,yetnotalong
theaxisofthecone,butratherclosertothelowersurface.Therefore,theposteriorhornissurroundedfrom
allsidesbylongitudinallyrunningcallosalfibresandtearsapartthelowerpartoftheforceps.Duetothe
positioningofthecorpuscallosumabovetheventricle,amajorpartoftheforcepsrunsanteriorlyoverthe
posteriorhorn(Fig.3.1).Themedialandlateralsurfacesoftheoccipitalhornarecoveredbyathinveilof
longitudinallydirectedcallosalfibres(2,3)withastrongerveilalongtheinferiorsurface(4).
Thelargeupperpartoftheforcepsflexesmediallywheretheposteriorhornopensupatthelevelofthe
quadrigeminalplate,inordertocrosstotheotherhemisphere.Asthispartisattheheightofthesplenium
fromtheverybeginning,itisthenaturalconfluenceforallforcepsfibres.Allforcepsfibresleavethe
cortexinafrontalplane,unlesstheyalreadyhaveenteredthecallosallayer,andthereforecanbetracedin

theirwholelengthtothispointoncoronalsections.Directandunhinderedaccesstotheupperpartofthe
forcepsisonlygiventofibresfromthecuneusandprecuneus,aswellasfibresfromthedorsalandlateral
convexityofthehemispherelocatedabovetheintraparietalsulcus.Thesefibresnotonlyjointheforceps,
butalsodigdeepintoit,beforetheybendfromafrontalplaneintoasagittaldirection.Theythusdivide
themassofsagittalfibresoftheforcepsintoanumberoftractsandlayers.Thelayeringisanexpressionof
theinterweavingofallcallosalfibres,whichcontinuesalmosttothemedialplane.Thus,callosalfibres
fromdifferentpartsoftheoccipitallobelienexttoeachother.Untiltheirinsertion,fibresfromthe
convexityofthehemisphereformatightlypacked,clearlydifferentiablefibremasslayerintheforceps
(5).
Onafrontalplane,theremainingcallosalfibresoriginatingfromthecortexruninferiorsuperiorlyalong
theoccipitalhorn.Whetherthesereachtheirtargetatthelateralormedialsurfaceoftheoccipitalhorn
dependsuponwhetherthecorticalareatheyoriginatefromlieslateralormedialonthesagittalplane
throughthemiddleoftheoccipitalhorn.Thisplaneseparatesthelingualgyrusfromthemedialpartofthe
fusiformgyrusatthebasalsurface.Thefibresystemoriginatingfromthefusiformgyrusoftenatightly
packedlayer,whichisclearlydifferentiablefromtherestofthefibres(6)climbsverticallyandbreaks
throughbothsagittallayersbydividingthemintothreeparts.Theinnermostpart(7)runsatthebasal
surfaceoftheposteriorhornalmosthorizontaltoitandbendsslightlyupwards,toinsertintheyettobe
describedsmallpartoftheforceps.Asmallermiddlepart(8)bendsinsagittaldirectionandstrengthens
theouterhalfoftheforcepsfibresthatrunsagittallyontheinferior[part]oftheposteriorhorn.Thelateral
largestpart(9)runsalongtheoutersurfaceoftheposteriorhorn,adjacentandlateraltothethinlayerof
thehorn.Ishallcallallcallosalfibresattheoutsideoftheoccipitalhornouterforcepslayer.Duringits
coursealongtheoutersurfaceoftheposteriorhorn,thislayeriscontinuouslystrengthenedbyfibres
originatingfromtheconvexityunderneaththeintraparietalsulcus.Thesefibresrundiagonallyfromthe
ventralconvexitytowardsdorsalmedialareas.Amongthemthemostventralfibresareclosetoavertical
direction.Themoredorsalthesefibresreach,themorehorizontaltheyrun,untiltheyjoinfibresthatcross
totheupperpartoftheforcepsdirectlyabovetheintraparietalsulcus.Theyformsmalltracts,visibletothe
nakedeye,thattraversebothsagittallayersinthesamedirectionasbeforeandthusdividethelatterin
evensmallertracts.Theythenbendupwardsinaverticaldirectionandjointheascendingfibres.The
wholelayerthusbecomesthickerasitascendsandbendsfromaverticaltoasagittaldirectionatthelevel
oftheupperpartoftheforceps.Alsothesefibres,likeallcallosalfibres,donotsimplyjoinfrombelowor
outsidethealreadyexistingforcepssystemtheyratherfollowthesamecourseofthecallosalfibres
[originating]fromthedorsalcortex,i.e.,theypenetratetheforcepsfora[certain]distancebeforebending
inasagittaldirection.
Thefibresofthesagittalveilwhicharedirectlyadjacenttothelateralsurfaceoftheposteriorhorn(2)
traversediagonallyalongananteriorsuperior[direction]andmergewiththedorsalbranchofthe
forceps.Inthesameway,thethickenedbundlebendsatthelateralaspectoftheinferioroccipitalhorn(8)
moreanteriorandclosetotheopeningoftheoccipitalhornwhereitrunsupwardsanddiagonallytowards
thefrontandthendirectlyupwardstoreachthesametermination.Thelateralsagittalveilshowsgreater
variability.Attimesitcanbeclearlyseenalongthewholelengthoftheoccipitalhorn,inothercasesit
coversonlytheposteriorpartbecauseitsfibresbentupwardsfarmoreposteriorlyandhencestrengthenthe
layeroftheverticalascendingfibre.Thelatterbordersdirectlytheependyma.
Thesamepositionisnotpossibleforthoseforcepsfibresoriginatingfromtheinnerpartofthefusiform
gyrus,thelingualgyrusandthecalcaravisatthemedialsurfaceoftheoccipitalhorn.Thisisduetothe
prominentcalcaravisthatbulgesintotheoccipitalhornandhindersasoliddevelopmentoffibresthatdo
notbelongtothecalcaravis.Theentireforcepsfibresoriginatingfromthelingualandfusiformgyrithat
shouldascendverticallyatthispointarerunninglongitudinallyalongtheinferiormedialedgeofthe
occipitalhornandtherebystrengthenthemedialhalfofthelongitudinalfibresattheinferioroccipital
horn.Hence,thisformsacordliketract,whichthickenstowardsthefront(4).Justbeforetheanterior
aspectofthecalcaravis,directlybehindtheopeningoftheoccipitalhorn,thistracthasenoughroomto
ascendassmallinnerpartoftheforcepsfromwithintheoccipitalhorn.Onceitreachestheroofofthe
ventricle,thistractbendsinwardstojointhelargerupperpartoftheforcepsandmergewiththecorpus

callosum.Thewhitematterofthefusiformgyrusisadjacenttotheabovementionedfibresthatrun
inferiortotheoccipitalhorn(7),whilstthewhitematterofthelingualgyrusformsadenserlayer(10)
similartotheonefromthedorsalconvexity.Thefibresfromthethinsagittalveilattheinnersurfaceofthe
occipitalhorntheinternalforcepslayer(3),whichareprobablyjoinedbycallosalfibresoriginatingfrom
thecalcaravis,mergeanteriorlyintheascendingpartofthesmallforceps.
Theentireinferiorpartoftheforcepsandthesagittalveilattheinnersurfaceoftheoccipitalhornshow
greatvariability.Bothstructuresaremutuallydependent:Ifthelowerforcepsisstronglydeveloped,than
theveilattheinnersurfacewillbeveryfinetothepointwhereitisdifficulttoappreciateitevenatahigh
magnificationanditmightonlyconsistsoftwoorthreefibrelayers.Inrarecaseshowever,allofthe
inferiorforcepsvanishesandinsteadformsatractmergingwiththeveil,whichdevelopsasarelatively
stronglayerthatuniformlycoverstheinnersurfaceoftheposteriorhorn.Attimes,theinnerforcepsdoes
notascendanteriortothecalcaravisbutascendsmoreposteriorlyinadiagonaldirectionupwardsand
forwards.
Allforcepsfibresarecharacterisedbyastrongfibrediameter.Thelayersoftheforcepsstainratherdark
withhaematoxylin,andstronglyyellowwithpicrocarmin.
6.Stratumsagittaleinternum
Thestratumsagittaleinternumwrapsaroundtheforcepsjustastheforcepsencasestheoccipitalhorn.The
fibresofthislayerdifferfromthefibresoftheforcepsfortheirsmalleraxonaldiameter.Thislayerstains
verylightwithhaematoxylin,whereaspicrocarminstainingcoloursthislayerinredcomparedtothe
surroundinglayers.Fibresofthislayeroriginatefromtheoccipitallobe,seeminglyfromallareasofthe
occipitalcortex,andcontinueanteriorlyintotheposteriorpartofthecoronaradiata.Thesefibresformthe
projectionconnections,namelythecoronaradiataoftheoccipitallobe.Toreachtheirdestination,they
havetogatherattheoutersurfaceoftheventricle.Fibresoriginatingfromtheoccipitalpoleunifyafew
millimetresbehindthebeginningoftheforcepsasasolidtractthatthickensasfurtherfibresjoinandruns
anteriorlyalongalongitudinaldirection.Oncethesefibresreachthetipoftheforcepsthetractfunnelsout
andfromhereonwardsencasestheforcepsfromallsidesintheshapeofananteriorlywideningbelt.On
sections,fibresofthestratumsagittaleinternumwerenottraceablewithoutinterruptionsalongtheirentire
trajectoryfromthecortexthroughthewhitematter.Theycanonlybedifferentiatedwithclarityfromother
fibres,oncetheyformaseparatelayer.Fibresattheinnersurfaceoftheforcepsthatrunlongitudinally
towardsthefront(12)aswellasfibresoriginatingmoreanteriorlyfromthecuneus,precuneus,andlingual
gyruscoursetowardsthelateralsurfaceoftheforcepsstillinthefrontalplanedescribinganarcaround
partsoftheforcepsthatcoursedorsalandventraltotheoccipitalhorn.Oncethesefibresreachtheoutside
oftheoccipitalhorntheybendanteriorlyinalongitudinaldirection.Oncoronalsections,theupperparts
ofthesefibres(13)clingtoforcepsfibresoriginatingfromthecuneusandtheprecuneus.Fibresfromthe
lingualgyrus(14)runinparalleltotheabovedescribedcallosalfibresandcoursefromthelateraltothe
medialsurfaceinoppositedirectionfromthebaseofthehemispheretowardstheinferiorpartofthe
forceps(7).
Asaconsequenceofthisarrangement,thepartofthislayerthatliesoutsidetheoccipitalhorn(11)
becomesthicker,whereasthepartontheinnersidebecomesfinerasthecalcaravisprogressively
penetratestheoccipitalhornanteriorly,suchthatitsoonbecomesonlyamicroscopicallyvisibleveil.
Eventually,theveilwilltearapartjustnearthecallosalbulgetoallowtheforcepstoreachthemedian
surface.
Themostinferiorfibresofthestratumsagittaleinternumrunalmosthorizontalalongtheirentirecourse
towardsthefront.However,themorefibresoriginatedorsoanteriorly,thesharpertheirdiagonalangle
fromadorsalposteriortoananterioinferiordirection.Intheparietallobethecoronaradiataruns
eventuallyverticaloncoronalsectionatthelevelofthetipofthepulvinar.Thusfromhereonwardsthey
canbetracedalongtheirlengthoncoronalsections.
Asalreadymentioned,lateraltotheoccipitalhornthislayerispenetratedbyfibresoftheforcepsthat
divideitinbundlesofequalsize.Thesebundlesarevisibletothenakedeye.Closetotheposteriorarchof

thecaudatenucleusthemiddlepartofthislayerreceivesfurtheradditionsfromtheyettobedescribed
stratumsagittaleexternum.
7.Stratumsagittaleexternum
Thestratumsagittaleexternum(15)enclosesthejustmentionedlayerinthesamewaythestratumsagittale
internumcoverstheforceps.Thislayerconsistsmainlyoffibresoflargeaxonaldiameter.Similartothe
forceps,itstainsverydarkwithhaematoxylin,yellowwithpicrocarmin,andisthusclearlydifferentiated
bothfromthestratumsagittaleinternumandthesurroundingfibres.Whetherthenumerousfinefibresthat
crossthesections,whicharevisibleatthelevelofthislayeroncoronalsections,arepartofitorarejust
traversingitandstrivetowardsthestratumsagittaleinternum,Ihavenotbeenabletoconfirmwithclarity.
Thelatterseemsmoreprobabletome.Fibresofthislayeroriginatefromtheoccipitalcortex,seemingly
fromallitsareas,andcontinuetowardsthetemporalcortexexceptforasmallportion.Theyformthelong
associationtractbetweenthesecortices[inferiorlongitudinalfasciculus].Inordertoreachtheir
destination,whichisthewhitematterofthetemporallobe,theyallhavetogatherattheventralaspectof
theventricle.
Posteriorlythelayerappearsasathinbelt,whichenvelopesthestratumsagittaleinternumequallyfromall
sidesandinitiallydescribesthesamecourse.Thesefibrescouldalsonotbetracedcontinuouslyontheir
wayfromthecortextotheirentranceintothestratum.Itseemsthatthesefibres,similartothoseofthe
stratumsagittaleinternum,donotstrivetotheircollectionpointlikethefibresoftheforcepswhichrun
verticallyfromtheconvexityofthebrainonafrontalplane,inamannersimilartothebranchesofan
appletreetothestem.Rather,theyradiatefromposteriorordiagonallyfromthecortex,anteriorlytowards
theventriclelikethebranchesofapeartreetothestem.Theythereforedonotruninparalleltotheforceps
fibrestowardsthecollectinglayersbutcrossthemlikeclaspedfingers.
Fibresfromtheoccipitalpoleanditsneighbouringareasrunanteriorly,longitudinal,andparalleltothe
ventraledgeoftheventricle.Thefibresunderneaththeoccipitalhornmaintaintheiralmosthorizontal
directionwherebytheycoursetowardsthefrontandslightlydescendinthetemporallobe.Forthejoining
fibresitappliesthatthemorethefibresoriginatefromdorsalanteriorregionsthemoretheirdirection
changesfromadorsalposteriortoananteriorinferiordescendingdirection.Hence,themostanteriorfibres
ofthislayerthatoriginatefromtheconvexitywheretheoccipitoparietalsulcuscutsthrough,meaning
fromthefirsttransitionalgyrus,formanangleofapproximately30withthemostinferiorfibres.Inpost
mortembluntdissectionsthislayerthereforeseemslikeapartiallyopenedfan.Attheinnersurfaceofthe
occipitalhornthislayer,likethetwolayerspreviouslydescribed,thinsouttoaslenderveilduetothedeep
penetrationofthecalcaravis.Thisveilbehavesanteriorlysimilartothethickcoveringonthelateral
surface,inthesensethatalsoherethefibresgraduallytakeaverticaldirection.
Aconsequenceofthisarrangementisthatthestratumsagittaleexternumprogressivelytightenstowards
thebaseofthebrainandformsatrack,whichbecomesbetterdefinedtowardsthetransitiontothe
temporallobe.Thetrackconsistsofasolidfootwithbilaterallyattachedsidepartsinaroundedright
angle.Thisprominentinferioraspectofthestratumsagittaleexternumhasbeentermedinferior
longitudinalfasciculusbyBurdach.Fibresfromthecortexformaridgelikeattachmentinthemiddlepart
oftheoccipitallobeatseveralpointswherecallosalfibrespenetratethelayerasthicktractsthisisthe
casedorsallytowardstheconvexity,attheinferioredgeofthestratumsagittaleexternumandtowardsthe
lingualgyrus.Thisattachmentbecomesveryprominentandelongatedinthelingualgyrusandhasbeen
namedbyBurdachastheinternalbasalbundle[inneresGrundbuendel].
Oncethestratumsagittaleexternumreachesthetemporallobeitquicklythinsoutbysendingfibrestothe
cortexinalldirections.Whenperformingdissections,alargepartofthesefibresfromthelateralaspect
andthefootofthislayercanbefollowedintothefirsttemporalgyrus.Asmallerpartreachesthesecond
temporalgyrus,andtheremainingfibresbecomeinsignificantandreachtowardsthetemporalpolewhere
theyinseparablymergewiththewhitematterofthetemporallobeandcontinueanteriorly.Themost
anteriorfibresofthislayerterminateinthepoleofitslobe.Intheanterioraspectofoccipitallobeandthe
precuneus,sparsefibresdescenddiagonallyfromthemedialsurfaceoftheoccipitalhornandafterjoining

thecingulumtheybendaroundthespleniumandthencontinuewithitandinhealthybraintheyare
inseparablefromittowardsthetemporallobe.
Withintheoccipitallobethestratumsagittaleexternumisdividedintosmall,equalbundlesbypenetrating
forcepsfibresjustlikeforthestratumsagittaleinternum.
Asmallamountoffibresofthestratumsagittaleexternumthatlieslateraltotheoccipitalhorndoesnot
reachthetemporallobeinsteadthemedialpartofthissubdivisionsegregatesintonumeroussmallbundles
thatarevisibletothenakedeye.Thesebundlesareentangledlikeropesandpenetratethestratumsagittale
internum.Theyaredifferentiablewithinthelatterduetothelargeraxonaldiameterandtheirdarkstaining
withhaematoxylin.Bothstructuresjointlyreachthefootofthecoronaradiata.
8.Stratumpropriumcorticis
Themajorityofwhitematterbetweenthestratumsagittaleexternumandthecortexhasalmostthesame
diameterinalldirectionsasthethreeinnerlayerstogether.Thiswhitematterconsistsmainlyofsmall
associationfibres,whichoriginateandterminatewithintheoccipitallobe.Itispenetratedbylongranging
fibresoriginatingfromthecortexandthencemergingwiththethreeinnerlayers.Theshortfibresmainly
runinthefrontal[coronal]plane,andthusinterconnectdorsalandventralormedialandlateralregionsand
onlyrarelydotheyinterconnectdirectlyadjacentcorticalareas.
Threesuchfibroustractsoriginatefromthedorsalaspectofthecorticalregionsabovethecalcaravis.
Themostimportantofthesefibresisthestratumcalcarinum(16),whichconsistsoffibresthat
circumventthecalcaravisinitsfullextension,andthelongestofwhichconnectthecuneustothelingual
gyrus.Inthewhitematterstripsofthethreeabovementionedverticalshortgyri,whichareplacedonthe
insulargroundofthecalcarinefissure,thislayerthickensintothreestrongbundles.Amongthesebundles
themostanteriorisratherprominentandpartiallyreachesthebaseofthehemisphere.Asaresultofthis
fillingofthegyralcomb,therespectivesulcidonotappearatthebedofthecalcaravisontheinner
surfaceoftheoccipitalhorn.Anteriorlythislayerreachesbeyondtheconnectionpointofthecalcarine
fissureandtheoccipitoparietalsulcusintothetemporallobeandenvelopesinasimilarfashionthe
continuationofthecalcarinefissurebyconnectingthecortexoftheuncinategyruswiththelingualgyrus.
Thesecondlayeroriginatesfromthedorsalcorticalregionofthecalcaravis,thestratumcunei
transversum(17).Incontrasttothestratumcalcarinumthislayeronlyexistsintheregionofthecuneus
anddoesnotextendbeyondtheconfluenceofthecalcarinefissureintheoccipitoparietalsulcus.The
fibresofthislayeroriginatetogetherwiththoseofthestratumcalcarinumandinitiallyrunparallelwith
themoverthedorsalcalcaravisfrommedialtolateral.However,ratherthanbendingdownwardsafterthe
calcaravistheycontinueinthesamedirectionabovethedorsalpartofthestratumsagittaleexternumand
benddownwardsontheothersideofthelattertofollowitslateralsurface.Oncoronalsectionscutthrough
theposteriorhalfoftheoccipitalhorn,thislayercanbeseentoreachtheinferiorborderofthestratum
sagittaleexternum:themoreanteriorthelessthesefibresreachinferiorlyandthethinnerthewholelayer
becomesuntiltheyeventuallyvanishintheregionoftheanterioroccipitalsulcus.Thusfarithasnotbeen
possibletotracethesefibresinisolationupontheirexitfromthislayeralongtheirtrajectorythroughthe
stratumpropriumconvexitatistowardsthecortex.Theypotentiallyreachthecortexofthewholeconvex
regionandpartoftheinferioroccipitalcortex,thusforminganassociationpathwaybetweenthecuneus
andtheconvexity.Itseemsthatthesefibreshaveastraightdirectiononlyintheirposteriorandmedialpart
alongamedialtolateralcourse,however,morelaterallyandanteriorly,theybendoutofthefrontalplane
andthancontinuediagonallyalongaposteriormedialtoanteriorlateraldirection.Thusthelongestfibres
ofthislayermightreachthedorsalparietallobeandpossiblytheangulargyrus.
Asubtler,yetattimesquiteprominent,analogouslayeroriginatesfromthelingualgyrusandcontinues
inferiorlyaroundthestratumsagittaleexternum.Inanidealsituationonecanappreciateafifthlayer,
whichenvelopesthelatterstratumfromallsidesintheposteriorfrontalplane,wheretheoccipitalhorn
stillisslitlike,betweenthestratumpropriumcorticesandthestratumsagittaleexternum.
Bothlayers,namelythestratumcalcarinumandstratumcuneitransversum,stainratherdarkwith

haematoxylinyetlessthanthestratumsagittaleexternum.Therefore,theycanbeclearlydifferentiated
fromitandfromtherestofthewhitematter.
Thethirdlayer,namelythestratumpropriumcunei(18),whichoriginatesfromtheupperedgeofthe
calcaravis,ascendsperpendiculartothedorsalhemisphericmarginandencapsulatesthefissureofthe
cuneusthatrunsparalleltothecalcarinefissure.
Thesethreelayersoriginatingfromthecuneus,seeminglyformajointsystemofshortassociationfibres,
whichinterconnectthecortexofthecuneuswiththeentireoccipitalcortex.
Similartotheregionnearthecalcaravis,thespacebetweenthecortexandthestratumsagittaleexternum
lateraltotheoccipitalhornisfilledbyastratumverticaleconvexitatis,whichrunsverticallyinadorso
ventraldirection.Eachofthethreesagittaloccipitalsulciisenvelopedbyasystemofgutterlikefibres[U
shapedfibres],whichconnectthegyriaboveandbelowthesulcistratumpropriumsulcioccipitalisIs.
interparietalis(19),str.Pr.S.o.II(20),str.Pr.S.o.III(21).Afourthsystem,thestratumpropriumsulci
collateralis(22),connectsthelingualgyruswiththefusiformgyrusatthebaseofthebrain.Themore
medialonereachesfromthelateralaspectofthebrainthelongertheverticalfibresbecome.Thefibresthat
followthesuperficialstratapropriaofthesulcihurdleonlyonegyrus.
Thedeepestfibresdirectlyabuttingthestratumsagittaleexternumorstratumtransversumcuneiproject
alongthewholeheightofthelobeandinterconnectasstratumprofundumconvexitatis(23)thedorsaland
ventralmarginsofthehemisphere.
Thisprominentverticalfibresystem,thestratumproprium[verticale]convexitatis,isconsistentacrossthe
wholeposteriorpartofthecerebrum.Anteriorlyitextentsbeyondtheoccipitallobeandgradually
becomesthinnerbeforeitssharpboundaryinthewhitematterstripbetweenthepostcentralandthe
intraparietalsulciwithintheparietallobe.Furtherinferiorthemostanteriorverticalfibresencasethe
insertionpointoftheinterparietalsulcuswiththepostcentralsulcus.Theythanreachthesupramarginal
gyrusfromwheretheycourseanteriorinthedepthtojointheassociationfibresoftheinsulathatascend
fromtheoperculum.Inthetemporallobe,themostanteriorfibresdescendfromtheinferioraspectofthe
angulargyrustowardsthesecond[middle]temporalgyrusandformthefloorofthesuperiortemporal
sulcus,whichatthispointisofteninterruptedbyasmallverticalgyrus.
Thestratumverticaleconvexitatisisalsostronglydevelopedinthemonkeyandhasbeendescribedas
fasciculusoccipitalisperpendicularisbyWernicke(aspreviouslycited,p.23).
Similartothesagittalsulci,bothverticalsulci,namelytheanterioroccipitalsulcusandtheascending
branchofthesuperiortemporalsulcus,areencapsulatedbyaverythingrooveoflongitudinallydirected
shortassociationfibres.
Intheprecuneus,thelayeroffibresadjacenttothecortex,namelythestratumpropriumpraecunei,also
hasaverticaldirectionandencapsulatestheposteriorelongationofsulcuscallosomarginalisindorso
ventraldirection.Moremediallylocatedfibresbendanteriorlyattheirinferiorterminationsandjointhe
dorsalpartofthecingulumwhosedetaileddescriptionisyetoutstanding.Thedeeperthesefibresrun,the
fartheranteriortheypenetratethecortexofthegyrusfornicatus[theupperlimbisthecingulategyrusand
thelowerlimbistheparahippocampalgyrus].Athirdlayerofverticallydirectedfibresisformedbythe
fibrespreviouslydescribedasbelongingtotheanteriormedialpartofthestratumsagittaleexternumand
joiningthedescendingpartoftheventralcingulumreachingthetemporallobe.Thesecondmentioned
layerbelongstotheanteriorpartoftheprecuneus,whereasthethirdbelongstoitsposteriorpart.
Subsequently,fibresofthecoronaradiatafollowthatascendtowardsthehemisphericmargin.
Intheanteriorregionoftheoccipitallobeandatthetransitiontotheparietallobe,wherethestratumcunei
transversumterminates,itremainsawhitemattersystemsurroundedbythestratumpropriumpraecunei
medially,thestratumverticaleconvexitatislaterally,andthestratumsagittaleexternumventrally.This
systemabutsthesuperiorpartofthestratumsagittaleexternumlikearoofridgeandconsistsmainlyof
fibresthatruninalongitudinalcraniocaudaldirection.Thisfibresystemisonlyclearlyvisibleonfresh
coronalsectionsofabrainhardenedintheMllersolution.Itappearsasabrighterarea,whichabutsthe

stratumsagittaleexternumlikeacapandisdistinguishablefromthedeepdarktransversecutofthelatter,
whilstitbecomesgraduallyindistinguishabletowardsthedorsalandlateralwhitematterofthestratum
propriumcorticis.Itseemsthatthiscapdoesnotrepresentanassociationpathwayinitsownrightbutits
curiousappearanceisratherowedtothefibresthatpiercethroughhereandthenadheretoreachintothe
stratumsagittaleexternumandinternum.Itispossiblethatthisareaisalsoinrelationtothemost
anteriorlybendingfibresofthestratumcuneitransversum.Thisisnotnoticeableinstainedsectionsofa
healthybrain.2
Asimilarsmallerfibresystemispresentbetweentheinferiorpartofthestratumsagittaleexternumandthe
stratumpropriumsulcicollateralis.Athirdsystem,attimesincontinuitywiththejustmentionedsystem,
isfoundinthelingualgyrusclosetothecortexofthecalcaravis.
Alltheselayerswithinthestratumpropriumcorticis,exceptthefirstmentionedstratumcalcarinumand
stratumcuneitransversum,stainproportionallyweakwithhaematoxylin.
Withregardstotherelationofsizeandformofallthesewhitematterlayersalookattheattached
photographswillallowabetteroverviewthananythoroughdescription.Here,onlythefollowingwillbe
mentioned,asitseemsimportantwithregardstopathology.
Asmentionedabove,theincisionofthesulciintothewhitematteronlyaffectstheconfigurationofthe
outermostlayer,thestratumpropriumcortices,butonlymarginallytheshapeofthethreeinnerlayers(not
eventhestratumtransversumcunei).Onlythethreelayersofthecalcaravisthinouttoveillikecoverings.
Themedialoccipitotemporalsulcuscausesaconcaveinvaginationofthelowermarginofthestratum
sagittaleexternumwhilstthethicknessofthestratumpropriumcorticisdependsontheproximityofthe
corticalsulcitothestratumsagittaleexternum.Atthemedialsurfaceofthebrainthiseffectisvisiblein
thethickeningofthethreeabovedescribedgyribrevescalcarisavisthatformthestratumcalcarinum.At
theoutersurface,thestratumpropriumispushedtogetherbybothverticalsulcioftheoccipitallobe,less
sobytheanterioroccipitalsulcusbut[even]morebytheascendingbranchofthesuperiortemporalsulcus.
Thestratumverticaleconvexitatisisespeciallythinnedbythecortexofthemostposteriorprotrusionof
theSylvianfissure.Thethinnertheouterlayer,theeasieralesionthatisoriginatingfromthecortexcan
reachtheinnerlayers.Alesionprogressionfromthecortexisthuseasiestattheposteriorendofthe
Sylvianfissureandunderneaththesecondparallelsulcus,hencetheregionoftheinferiorparietallobe.
Consequently,asuperficialsofteningwithinthisregioncan,dependingonitsdepth,isolatethestratum
sagittaleexternumordamageboththestratumsagittaleexternumandthestratuminternum.Thiscancause
transcorticalsyndromessuchasopticaphasia(Freund)orapperceptivesoulblindness[associative
agnosia](Lissauer)duetoaninterruptionoftheconnectionsbetweenvisualandauditorycentres.When
thedisconnectionispresentinassociationwithasubcorticaldisturbancethiscauseshemianopsia.Onthe
otherhand,alesionoriginatingfromtheependymaoftheoccipitalhornmaydisrupts,apartfromasmall
partofascendingforcepsfibresatthelateralsurfaceoftheoccipitalhorn,theentirestratumsagittale
internuminitsfunctionandthuscausepuresubcorticalhemianopsiawithoutcorticalortranscortical
syndromes.
Additionally,twoothertractsremaintobedescribed,whichareinclosespatialrelationshiptotheoccipital
lobe,especiallytothestratumverticaleconvexitatisandstratumcalcarinum.Theydo,however,not
continueaspartofthelobarwhitematterandarenotinrelationwithitscortex.Thesearethearching
fibresandthecingulum(Burdach).Thearchingfibres(respectivelythefasciculusarcuatusordorsal
longitudinalfibresorlongitudinalissuperior)correspondtothestratumverticaleconvexitatisofthe
occipitalbrainintheircoursethroughtheanteriorpartsofthebrain.Itislocatedinthedepthofthedorsal
gyrusoftheSylvianfissure,namelytheoperculumitsfibresextenddorsallyapproximatelyoverhalfthe
heightoftheconvexity.Itconsistsofshortassociationfibres,whichconnectneighbouringgyriwitheach
other.Indeeperlayers,theseassociationfibresbypassonegyrusatmost.Idoubtitcontainslong
associationfibres,whichconnectdistantcorticalareas.Thedeepestfibresofthistractrunninginthebed
ofthedorsalsulcusoftheinsulaseemtohaveaspecialfunction.Thedirectionofthesefibresisalways
perpendiculartothedirectionofthecoronaradiata.Intheregionofthecentralgyrusandthedorsalpartof

themarginalgyrusthesefibresrunhorizontally.Atthetransitionpointfromtheparietallobetothe
temporallobeitbendsdownwardsandjoinsthestratumverticaleconvexitatiswhoseanteriorprojections
areidenticalwiththesefibres.Haematoxylinstainsthearcuatefasciculusrelativelylight.
Alongthemedialaspectofthehemispherethecingulumrunswithatrajectorythatissimilartothearcuate
fasciculus.Itoriginatesunderneaththecallosalrostruminthemostposterioraspectoftheinferiorsurface
ofthefrontallobe[subcallosalgyrus]asathinwidelayerthatisinferiorlyabuttingtothecorpuscallosum.
Initially,thefibrescontinuediagonallyupwardsandthenformabundlethatbendsdorsallyaroundthe
genuandhorizontallyabuttingtothecorpuscallosumdirectlyunderneaththecingulategyrus.The
cingulumrunsalongtheentirelengthofthecorpuscallosumbeforeitbendsaroundthespleniumand
projectstotheparahippocampalgyrusinthetemporallobe.Whendisregardingitsfrontallobetrajectory,
thecingulumcanbesegregatedintoadorsalpart,adescendingpart,andaventralpart.Thecingulum
consistsofnumeroussmallfibresthatonlystainlightlywithhaematoxylinandacompacttractoflong
darkstainingfibres.Thedorsalpartofthecingulumincludestheabovementionedfibresthatconnectthe
cortexoftheprecuneuswiththecingulategyrus.Thedescendingpartseparatesthespleniumfromthe
anteriorfibresofthestratumcalcarinumfromwhichitisdistinguishedbyitsdarkhaematoxylinstaining.
Theinferiorpartisjoinedbyanteriormedialfibresofthestratumsagittaleexternum,whichoriginatefrom
theprecuneus.
Theabovementionedtractsandlayerscanbevisualisedasdetailedbelowbyusingthemethodsdescribed
above.Thestratumverticaleconvexitatiscanbeeasilydemonstratedwithbluntdissectionswherethe
cortexisbrokenoffwithascalpelandthefibresarethenremovedinbundlesusingtweezers.This
procedurepermitstoappreciatethesharpanteriorborderofthelayerandtheanteriortractsofthetemporal
lobethatcometogetherwiththedescendingfibresofthearcuatefasciculus.Ifpreparedcarefullyitis
possibletovisualisefibresthatrunthroughthislayerandprojectmediallyalongthewhitematterstripof
thegyri.OnfreshsectionsthroughabrainhardenedinMllersolution,thislayerappearslightgreenon
coronalorsagittalcutsanddarkgreenonaxialcuts.Thestratumcalcarinumandpropriumsulci
collateraliscanbeequallydissected,although,withmoredifficultyandonlyiftheyarewelldeveloped.
Theyvisualisesimilarlyonfreshsections.Iwasnotabletodemonstratethestratumtransversumcunei
withbluntdissections.Onfreshsectionsofawellhardenedbrain,however,thiscanbedistinguishedfrom
stratumverticaleconvexitatisbyitsdarkercolour.Thestratumpropriumcuneiisespeciallymarkedasa
greenblacktransectiononaxialcutsoffreshsections.Alltheselayersofthestratumpropriumcorticisare
relativelystronglydestainedwhenusingPalstaining.Thestrongestdestainingoccursforthestratum
profundumconvexitatisandalessereffectisseenforthestratapropriaofthesulcithestratumcalcarinum
andstratumcuneitransversumremaindarkblue,thoughstilllighterthanthestratumsagittaleexternum.
Bluntdissectionisparticularlygoodforthelateralsurfaceofthestratumsagittaleexternumwhenthe
fibresofthecortex,whichareperpendiculartothem,arefullyremoved.Thefootofthestratumsagittale
externumisbestdemonstratedbydissectingfrommedialaspectswhereitisthebasalbundleofBurdach
andreachesclosetothecortexofthelingualgyrus.Whenthecalcaravisisreachedthelayerbecomestoo
thinforfurtherdissection.Moreanterior,however,thefibresfromtheprecuneusthatjointheinferiorpart
ofthecingulumarewelldemonstrated.Onfreshsections,thislayerappearsblackonfrontalsectionsand
lighteronparietalsectionsespeciallyinthedorsalpartsduetothemajorityofdescendingfibres.Itis
distinguishedfromthestratumsagittaleinternumaswellasitsabuttingcapbyadifferentshadeofstain.
Onaxialcuts,however,thelayerislightgreenandonlyonmoredorsalcutsinanteriorregionsdoesit
appeardarker.Theinnerborderbetweenstratumsagittaleexternumandinternumcannotbedemonstrated
withbluntdissectionasthefibresofbothlayershavethesamedirectionoverlongdistances.Thestratum
sagittaleexternumisclearlydistinguishableinallitspartsfromsurroundingfibreswhenusingPalstained
sectionsthestrongerthedestainingofthesection,thebetterthedistinction.Thisstainisadequatefor
thislayer.Itstainsstronglydarkblueandcanbefollowedunderthemicroscopeintoitsfinebranchesat
themedialaspectoftheoccipitalhorn.
Asmentionedabove,thestratumsagittaleinternumcannotbeclearlyvisualisedbydissectionsbeginning
fromtheconvexity,however,whenstartingfromthemedialsurfaceitsvisualisationispossiblewhen

removingallcallosalfibres.Onfreshsections,thislayerisdistinguishedfromthestratumsagittale
externumlateraltotheoccipitalhornbyadifferentshadeofcolour.Fibresthatruntransverselyinferior
anddorsaltotheoccipitalhornarewhiteoncoronalcuts.WhenusingPalstainingthislayerstainsonly
lightlyandgainsabrownishshadefromwhichthedarkbluecallosalfibres,thattraversethislayer,canbe
clearlydifferentiated.Picrocarminstainsthislayerreddishcomparedtothesurroundingstructuresand
showsitsnucleiinarowalongthepenetratingcallosalfibres.
Theforcepsisnicelyshowninitsentiretywithbluntdissectionwiththeobviousexceptionofsinglefibres
thatpenetratethesurroundinglayers.Onfreshsectionsthefibresthatrununderneathandlateraltothe
occipitalhorntowardsoccipitalanddorsalregionspenetratethestratasagittalia.Thesefibresappear
whitishonfrontalcutseverythingelseappearsblackgreen.Onaxialsectionstheassociationand
commissuralfibresarewhitishandprojectionfibresareblackgreen.ThePalmethodstainstheselayersof
theforcepsalmostasdarkasthestratumsagittaleexternum.
Itiseasytorevealthearcuatefasciculuswithbluntdissection.Onfreshcoronalcuts,itappearsasadark
slimellipsoidadjacenttothecoronaradiatethatsendsabranchintotheoperculumitcompletely
disappearsbehindtheSylvianfissure.WhenusingPalstaining,thearcuate[fasciculus]isnotdistinctly
visibleanywhere.Theonlychangethatbecomesevidentoncoronalsectionsisthattheregionanteriorto
thecaudalendoftheSylvianfissurewherethearcuateispassingthroughisslightlylighterthanthe
surroundingareaafterstrongdestaining.
Bluntdissectionnicelydemonstratesthecingulumalongitsentirelengthincludingbothitsshortandlong
fibres.Onfreshcoronalcuts,thelongfibresappearasablackgreenfieldthatisabuttingtothecallosum
andpenetratesthecingulategyrus.Behindthespleniumitappearsasawhitegreenthincordwithadorso
ventraldirection.Onfreshaxialcuts,ithasexactlyoppositecolours.Inthetemporallobethecingulum
disappearsasanindependentarea.ThePalmethodstainsitsshortfibreslight,thelongfibresdarkblue,
however,notasdarkasthestratumsagittaleexternum.
Burdach(1822)hasdissectedtheinferiorlongitudinalfasciculuswithabranchtowardsthefrontalpole.
Thisisonlypossible,whencomingfromthestratumsagittaleexternumintheanteriortemporallobetothe
posteriorextensionoftheuncinatefasciculus,whichcoversthelatterandconnectsthetemporalpoletothe
orbitalfrontallobe.Suchatrajectorycanbeartificiallyproducedwithbluntdissection.Thelongestfibres
oftheuncinatefasciculusoriginateattheinferiorlateralmarginofthehemisphere,wheretheshortest
fibresofthestratumsagittaleexternum,comingfromposteriorly,terminate.Thismightthereforebeseen
astheareathatbestdefinestheborderbetweentheoccipitalandthetemporallobe.However,thiscould
evokethefalseimpressionofanuninterruptedtrajectoryoffibresthroughbothbundles.Onhistological
cutsitisimmediatelyevidentthatthisisjustadeception,asbothlayersremainclearlydistinctfromeach
other.Oncoronalsectionsthroughthetemporallobe,thestratumsagittaleexternumbecomesaslim
horizontaldarklineanddisappearsfullytothenakedeyelongbeforeitreachesthetemporalpole.
Meynert(ascited,page41)believesthatitispossibletofollowthefibresoftheanteriorcommissureinto
theoccipitalpoleusingbluntdissection.Iwasnotabletoreplicatethis.Icouldonlyfollowfibrebundles
oftheanteriorcommissureuptotheinferiormarginofthecortexofthetemporallobeandIamconvinced
thatthemajorityofthesefibresendhere(seealsoWenickeascited,page86).Amarginoferrorisgiven
here,asfibresoftheanteriorcommissurecrossthoseofthestratumsagittaleexternumdiagonally,thus
permittingonetoeasilygetfromonefibrelayerintotheotherduringdissection.Anteriorcommissure
fibrescannotbefollowedbeyondthetemporallobeneitheronfreshnoronhistologicalcoronalcuts.
Onufrowicz(1887)andKaufmann(18871888)havestudiedbrainswithcongenitalagenesisofthecorpus
callosuminwhichtheyfoundthatthetapetumofthetemporalandoccipitallobeswaspresent.Both
authorscouldfollowthetapetumanteriorlyasathicklongitudinalfibrebundle,whichtheyreferredtoas
superiorlongitudinalfasciculusorarchingbundle[Bogenbndel]ofBurdachandbelievedittobevisible
duetotheabsenceofthecorpuscallosum.Theythusinferredthatthetapetumisnotpartofthecorpus
callosum,butrathertheposteroinferiorpartofalargefrontooccipitalfasciculus.Thistracthasthence
beenreferredtoasfasciculusfrontooccipitalis[superiorfrontooccipitalfasciculus]intextbooksby

ObersteinerandEdinger.
Itakethelibertytosuggesthere,thatinordertoavoidconfusionalreadyknownstructuresofthebrain
shouldbereferredtousingtheterminologyintroducedbyBurdachuntilafullreviewofanatomicalterms
hasbeenconducted.Burdachonlyreferstothatpartofthecorpuscallosumastapetum,whichruns
towardsthebaseofthebrain,lateraltothelateralventricle,andthencontinuesanteriorlyintothetemporal
lobe.Burdachreferstoallremainingprojectionsfromthecallosumtotheoccipitallobeasforceps.In
morerecentpublications,eventhefibresascendingatthelateralsurfaceoftheoccipitalhornandmerging
withthedorsalforcepsarecalledtapetum.Boththeselayerscorrespondtoeachotherandmergeintoeach
otherattheopeningoftheoccipitalhornyet,theycanbedifferentiatedfromeachother.Theposterior
fibres,whichbendanteriorlyandthusreachthetemporallobe,aretheterminationsofthetapetum.Fibres,
thatfollowafterwards,ofwhichthefirstdescendstraight[while]thelaterruntowardstheoccipitallobe
forashortdistanceinthedorsalforcepsbeforedescending,arepartofforcepsandconstitutetheanterior
partofthislayerthatascendstowardstheforcepsalongthelateralsurfaceoftheoccipitalhorn.Theborder
betweenbothlayersliesjustbehindtheposteriorarchofthecaudatenucleus.
Tomybelieve,bothabovementionedauthorshavemistakenthesuperiorlongitudinalfasciculusor
arcuatefasciculuslocatedclosetothelateralconvexitywiththecingulum,whichislocatedatthemedial
surfaceandseparatedfromthearcuatebythecoronaradiataandthestratumsagittaleexternum.Owingto
theabsenceofthecallosum,thecingulumispositionedmoreinferior.Thearcuatefasciculus3wasnotonly
hintedatbyBurdach,assuggestedbyOnufrowicz,butwasdistinctlydescribedbyhim.Itisindeedeasyto
demonstratethisbundleinthehealthybrainusingbluntdissectionorfreshcrosssections.
Accordingtothedescriptionandthefiguresfrombothpublicationsitcanonlybeinreferedthatthese
fibresbelongtothedorsalpartofthecingulumandposteriorlymergewithascendingfibresoftheforceps.
ThoughIhavelookedwithoutmostcare,Iwasnotabletofollowanyfibresfromthedorsalpartofthe
cingulumtotheoccipitallobe.Thecingulatefibresarelimitedtothecingulategyrus[Randgyrusdes
Balkens].Unlesstheyterminatewithintheanteriorpartoftheprecuneusorthedescendingpartof
cingulategyrus,thesefibresruninanarcharoundthespleniumandreachthetemporallobe.Likewise,on
freshandstainedsectionsitisimpossibletodemonstratethatcingulatefibres,whichareclearlydistinct
everywhere,reachtheoccipitallobe.
OwingtoMr.Kaufmann'scourtesyIwasabletoreexaminehisanatomicalpreparations.Iherebyarrived
attheconclusionthatthisisnotindeedanacallosalbrain.Thefibresofthecorpuscallosumareallpresent
theymerelydonottransversetothecontralateralhemispherebutratherremaininthesamehemisphereand
runanteriorposteriorly.Therebyproducingafrontooccipitalbundleintheacallosalbrainthatis
completelyabsentinthehealthybrain.
Suchheterotopyofthecorpuscallosumisofnoimportanceforthestudyofthenormalbrainanatomy,yet
canbeimportantforthestudyofitsdevelopment.
Morerecently,Mingazzini(1890)indeeddescribedabrainwithcompletecallosalagenesiswherethe
ascendingforcepsfibresandtapetumwerealsoabsent.
WithregardstoHamilton'srepetitionofFoville'sbeliefthatthecorpusisacrossoverofbothinternal
capsules,thefollowingisthecaseintheoccipitallobe:callosalandprojectionfibresareclearly
distinguishablefromeachother.Fibresfromtheposteriorpartofthefootofthecoronaradiatarun
ipsilateraltowardstheoccipitallobewithinthestratumsagittaleinternumand,toasmallerextent,within
thestratumsagittaleexternum.[Also]thereisnoevidencethattheforcepsformsacommissureofboth
occipitallobes.Forthetimebeing,wecannotevenspeculateonthecontinuationoffibresaftertheycome
fromtheforcepsononesideandtraversetotheotherhemisphere.Theymightreachtotallydifferent,
anteriorcorticalregionsorevenreachtheinternalcapsule.Bothmethods,namelybluntdissectionand
histology,failtoanswerthisquestion.Inthefuture,thisquestionmightbeaddressedwithunilaterallesion
studies.
Ibelievethatthewidelyacceptednotionthatthefunctionofthecorpusistoconnecthomotopicalcortical

regions(seeMeynertascitedp.41Wernickeascitedp.23)iswrongoratleastincomplete.Thereisno
evidenceforthisaprioriopinion.Againstthisopinionstandsthefactthatcallosalfibresentanglepriorto
reachingthemidline.Mostlikely,fibresfromcertainareasofonehemispheredisperseindifferent
directionsaftercrossingthemidline.Thereisnoreasontoassumethatthesefibres,insteadofreaching
theirdestinationontheshortestpossiblewaylikeallotherfibres,reachthemidlinetotallyarbitrarilyand
thattheythensoradicallychangetheirpositionthattheycometoliesmoothlyinthesameordernextto
eachotherastheydidatthebeginning.
TheargumentthatHamiltonusesagainstpreviousscientists,especiallyMeynert,namelythatitis
impossibletofollowasinglefibrefromoneareaofthecortextothehomologousareaintheother
hemisphere,alsostandsagainstHamiltonhimself.Itisequallynotpossibletofollowasinglefibrefrom
thecortextotheinternalcapsuleoftheotherhemisphere.
Generally,IagreewithSchnopfhagen's(1891)interpretationofthecorpuscallosumasabedof
associationfibres,whichconnectsstructurallyandfunctionallytotallydifferentregionsofthe
hemispheres.Itisbeyondmyjudgment,ifaminorityofcallosalfibresmightreachtheinternalcapsulein
thefrontallobeaspostulatedbyHamilton.Schnopfhagencontestedthisopinion.Intheposteriorregions
ofthebrainitseemsthatnocallosalfibresenterthefootofthecoronaradiata.
Physiologypostulatesatleasttwotractsintheforceps.Assumingthattheregionsforfocalvisionare
presentedintheoccipitallobes,acommissuraltractwithintheforcepsistobeexpected,whichconnects
thevisualcortices.Additionally,therehastobeadirectconnectionfromtherightoccipitallobetotheleft
temporallobe,whichallowsthenamingofobjectsseenintheleftvisualfieldandwhichhastobe
disruptedincasesofopticaphasiaofFreund.Thistractisprobablyfoundwithintheforcepsontheright
sideandwithinthetapetumontheleft.
IoughttocommentonastatementbySchnopfhagenonthestraightoccipitalbundleofWernicke.
Schnopfhagensays(p.102):Wernickedescribesastraightoccipitaltract,afibrebundlerunningfrom
dorsaltoinferior,whichconnectsthesecondtemporalgyrus(namelythePlicourbe,thedorsalpartthatis
neighbouringontotheprecuneus)withthefusiformgyrus[Spindelwindung].Adrawingofthistract,
basedonanaxialcutthroughamonkeybrain,isavailableinhisbookonbrainpathologies(Fig.19ff).It
seemstomebeyonddoubtthatthisstraightoccipitalbundleisnothingbutaplaitedareaattheconvex
lateralsurfaceoftheoccipitalhorn.
Itseemstomeratherbravetoreachanopinionbeyonddoubtbasedonschematicdrawingsofathird
person,suchasWernicke'sfigure19,fromwhichathirdpartygaineditsassumptions.Thestraight
occipitalbundleisacollectionofassociationfibres,whichareevidentinthemonkeybrainonhorizontal
cutsandespeciallyonsagittalcutswheretheyappearassagittallycutfibres.Atriangularplaitedregionon
axialsections,whichisdistinguishablefromtherestofthefibremassasabaseofagyrusattheconvex
lateralsurfaceofthewalloftheoccipitalhornexistsneitherinthemonkeynorinadulthumanbrain.In
thehumanbrain,theassociationfibresofthestratumprofundumconvexitatisaresoprominentthat
individualfibresfromthecallosum,thecoronaradiataorlongassociationfibresrunningtowardsinner
layersfullydisappearwithinthissystem.
Thefollowingconclusiondoactuallynotbelongherebutareratherdestinedfortheendofthework
dedicatedtotheentirewhitematteranatomyofthecerebrum.Meynert'stheoryaboutthedevelopmentof
psychiatricactivityisbasedupontheanatomicalassumptionthateachpartofthecortexisindirect
anatomicalconnectiontoeachother,suchthatbetweenanytworandomcorticalregionsassociationtracts
canbecarvedout(Meynert,p.138).Myresearchthusfardoesnotsupportsuchanassumptionasa
generalrule.Theoccipitallobehasonlyonelongassociationtract,namelythestratumsagittaleexternum
thatconnectstothetemporallobe[inferiorlongitudinalfasciculus].Possibly,theremightalsobesome
minorconnectionviatheanteriorfibresofstratumtransversumcuneibetweenthecuneusandtheposterior
partoftheparietallobe.Apartfromthesethereisnoevidentconnectionwiththeparietalorfrontallobes
(neithertheirconvexitynortheirmedialsurface),whichwouldbecomparabletothedimensionsofthe
connectionreachingthetemporallobe.Likewise,apartfromthetemporallobe,thereisnosignificantlong

associationtractbetweentwophysiologicallydistantbrainregions.Provableconnectionsarelimitedtothe
vicinityandeventhelongestofthesestaywithinthebordersofeachlobe.Anylong[interlobar]fibres
wouldthereforehavetoberelativelyfewandisolated.
However,thisisratherdifferentforthetemporallobe.Thetemporallobehasastrongconnectionwiththe
occipitallobeviathestratumsagittaleexternum.Animportant,thoughlessprominentconnectiontothe
frontallobeisviatheuncinatefasciculus.Thecingulumconnectsthetemporallobetotheprecuneus,
paracentrallobeandthepartofcingulategyrusthatliesabovethecallosum.Thecingulatefibresmight
evenreachthefrontallobe.Thetemporallobeisinconnectionwiththeparietallobeviatheposteriorpart
ofthearcuatefasciculusortheanteriorfibresofthestratumverticaleconvexitatis.Additionally,itisthe
onlylobetohavecommissuralfibres,meaningthatfortheanteriorcommissureistruewhatisnotthecase
forthecallosum:fibresinbothhemispheresruninthesamewaywithoutcrossingorentangling.
Incomparisontotheseveryprominentassociations,thecoronaradiataofthetemporallobeisrelatively
insignificant.Apartfromthefornix,whichconnectstothemammillarybodiesandpossiblyalsotoa
corticalarea,onlyasmallamountoffibresenterstheinternalcapsule.
Thisarrangementispossiblytheanatomicalexpressionofthepsychologicalfactthatlanguageisofutmost
importanceforhumanthoughtprocess.Wordsandsoundshavedirectanatomicalconnectionswithall
primarycorticalareasforsensoryperception,whereasthoseareasthemselvesareonlyindirectlyconnected
viathespeechcentre.Allseparatepartsofthought,whicheventuallyarecomposedfromthememoryof
varioussensualperceptions,areinessenceconnectedbythemediumword,whichexpressesthethought.
Thustheanatomicalstudyofthebrainmakesoneunderstandtheincrediblepowerthewordhasforhuman
beings,intheireverydaylife,butalsoinhallucinationsofthementallyill,andtheconfabulationsofthe
hypnotised.Thisphysiologicalarrangementofthethinkingorganmightbethereasonforthephenomenon
thatacongenitalblindpersonisabletodevelopallhighercognitivefunctionsdespitethelackofthemost
nobleofsenses,whilstdeafmutepeopleonlyonrareoccasionscanriseabovethelevelofananimal.
9.Annotationstotheenclosedphotographs[18]
TheenclosedphotographsaretakenfromspecimensstainedwiththePalmethodandarerepresented
accordingtotheirnaturalsize.Verticallines16inFigs.1and2representtheapproximatelevelofeach
cut.
1. Thecutislocatedapproximately25mmanteriortotheoccipitalpole.Apartfromatypicalsulci,this
cutshowsthetransectionofthebeginningofthethreeoccipitalsulci(asaboveI,II,III)onthe
lateralconvexityandthecollateralsulcusontheinferiorsurface(coll.).Themedialaspectshowsthe
calcarinefissure(f.c.)withitsprominentdorsalandinferiorgyri.Theseareevidentevenwithstrong
destainingofthespecimenduetothewhitematterstripsrunningwithinthecortexaswellasthe
sagittalrunningsulcusofthecuneus(cu.).Theposteriorhornisnotyetvisibleonthissection.Also
thecallosalfibreshavenotyetunitedtoformadistinctlayer.Themedialaspectofthewhitematter
offersonlytwolayersthatareconcentricandformatrianglewiththedorsaltipandaventralbase.
Thistriangleisroughlyequaltothesizeofthecalcaravis.Thelightlayerinthemiddleisthe
stratumsagittaleinternum(1),thelateraldarkerlayerthestratumsagittaleexternum(2).Thewhite
mattertowardsthecortex,whichisthewhitematteroftheoccipitallobe,thestratumprofundum
convexitas(10)isstainedrelativelylight,thestratapropriaofthecuneus(5)ofthethreeoccipital
sulci(6,7,8)andthesulcuscollateralisarestainedslightlydarker.Evendarker,yetlighterthanthe
stratumsagittaleexternum,isthestratumcalcarinum(4),whichislocatedbetweenthelatterandthe
cortexofthecalcaravis.Thesameshadeisevidentforthestratumtransversumcunei(3),which
jointlyoriginateswiththelatterfibresfromthecuneusandrunsdorsalandlateraltothestratum
sagittaleexternum.This[3]canbefollowedasaslimgreystriponthelateralaspectofthestratum
sagittaleexternumtotheinferiorlateralmargin.Hence,thisplaneshowsatotaloffiveencapsulated
layers,whichdifferintheirstainingintensityandcanthereforebeseparated.
2. Thiscutislocatedapproximately1mmanteriortothepreviousone.Thissectionshowstheforceps
asadarkstipethatisthindorsallyandwidensinferiorly.Itislocatedinthemiddleofthe

homogenousstratumsagittaleinternumasdescribedontheprevioussection.
3. Thiscutislocatedapproximately510mmanteriortothepreviousoneandapproximately30mm
awayfromtheoccipitalpole.Onthemedialmarginthissectiondemonstratesthecalcarinefissure
(f.c.)underneathadeepsulcusofthecuneus.Comparedtotheprevioussection,thefoundationof
thisfissureisdeformedfromawidebasetoathinditch.Onthedorsalcortexofthecalcaravis,as
wellasontheventralcortex,asecondarysulcaldevelopmentisevident.Theinferiormarginofthis
sectionshowsthesectionthroughthewelldevelopedcollateralsulcus(coll.).Onthelateral
convexitytherearefiveevidenttransversesulci.Thefourinferioronesoftheseequalthethree
longitudinaloccipitalsulciwiththedorsaloneforkingintotwobranches(s.o.IIII).Thecalcaravis
isunderdevelopedwhichiscompensatedforbyadominantprotrusionoftheoccipitoparietalsulcus
intothewhitematteroftheoccipitallobe.Thissulcusmergeswiththemainsulcus(o.),which
infringesfromtheconvexityandthenrunsposterior.Onthissectiontheposteriorhorncanalready
beappreciatedasatrianglewithasmallbase,themedialandinferiorbordersconcaveandthelateral
borderconvex(v.).Theindentationsofthemedialandinferiorbordersarenecessitatedbythe
intrusionofthecalcarinefissureandthecollateralsulcus.Amongstthemedialwhitematterlayers
theforceps(1)hasaprominentdorsalpartandalessprominentspoontongueshapedinferiorpart
thatisreachingintothelingualgyrus.Mediallytotheoccipitalhornthesinglelayerscannotbe
differentiatedduetotheoveralldarkstainingonthesection.Thebrightareaattheinferiorhalfof
theforcepsfibreslocatedlateraltotheoccipitalhornisnotagenuineappearancebutoriginatesfrom
anerrorduringthecuttingofthissection.Thereasonforthisisthepresenceofavesselinthe
ependymaloftheoccipitalhorn,whichdamagedsomefibresalongseveralsections.Thestratum
sagittaleinternum(2)canbeappreciatedasabrightareadorsal,lateralandinferiortotheforceps.It
thencontinuesasabeakshapedprotrusionintothelingualgyrus.Medialtotheoccipitalhornthe
layerhasbecometoothinforexaminationwiththenakedeye.Thesameappliestothedarkstained
stratumsagittaleexternum(3).Theinferioraspectofit[3]isthinnedandbendduetothecollateral
sulcus,whilsttheintrudingsulcionlycauseaslightcurvatureonthelateralaspect.Italready
becomesevidentonthissectionthattheinferolateralaspectofthislayerismoredominantthan
otherparts.
Thestratumprofundumoftheconvexity(10)islightlystainedwhilstthelayersofthesulci(6,7,8,9)are
easilydifferentiatedduetotheirdarkerstainaswehavealreadyseenonprevioussections.Exceptionally
darknearlycomparabletothestratumsagittaleexternumarethestratumcalcarinum(5)andthe
stratumtransversumcunei(4)whosecommonorigininthecuneusisclearlyvisible.Atthelateraldorsal
borderofthecortex,thestratumtransversumcuneiformsahelmshapedcap,whichisformedbyits
dorsallyprojectingfibres.Itthencecontinuesasaslimstripeatthelateralsurfaceofthelatterwiththe
resultthatitnearlyreachesitslateroinferiorborder.
4. Thiscutislocatedapproximately30mmanteriortothepreviousoneandapproximately60mm
awayfromtheoccipitalpole.Themajorityofthissectionisnotlocatedintheoccipitallobe
anymore.Thelateralaspectshowsthedorsalparietallobe(I)andunderneathittheinferiorpartof
theangulargyrus(II).Themedialaspectshowstheprecuneus(VIII)andasthemostanteriorpart
oftheoccipitallobetheanteriorterminationofthecuneus,whichisthinningtoremainasslim
stripe(VI).Themainfissuresofthesurfaceanatomythatcanbeappreciatedherearethe
intraparietalsulcus(i)andattheinferiormarginofthehemispherethethirdoccipitalsulcus.The
lattermightalreadybereferredtoasthethirdtemporalsulcushere(s.o.III).Thecollateralfissure
(coll.)isagainvisibleadjacenttotheinferiorpartofthestratumsagittaleexternum.Onthemedial
aspectthefissurecalcarina(f.c.)andtheoccipitoparietalissulcus(o.)areabuttingjustafterthey
havemerged.Thecrosssectionoftheprecuneusshowstheposteriorelongationofthecalloso
marginalsulcus(cm).
Theoccipitalhorninthisparticularspecimenisratherwideinitsanteriorhalf.Incomparisontothe
previoussection,itgainedinwidthandformedaprominentdorsalsurface,whichisprotrudingconvex
intotheventricledorsallyduetotheprotrusionofthedorsalpartoftheforceps.Thedorsalpartofthe

forceps(1)gainedsignificantlyinsizeandcontinuesatthelateralsurfaceoftheoccipitalhorn(2)intothe
verticallyascendingfibres.Thesefibresappearaslongitudinallycutunderthemicroscope(comparefigure
3and9).Theforcepsfibresunderneaththeoccipitalhornarecutlongitudinallywheretheyreachforthe
stratumsagittaleinternumandarecuttransversewheretheyareclosetotheventricle(Fig3.7).The
inferiorpartoftheforceps(4)isstilllocatedattheinferiormarginoftheoccipitalhorn.Theconnection
betweenthisandthedorsalpartisformedbyathinlayeroffibresthatarecuttransverselyandthatrun
alongtheinnersurfaceoftheoccipitalhorn,namelythemedialforcepslayer(3).Thestratumsagittale
internum(5)disappearedwherethecalvaravisispenetratingthewhitematterandisnotvisibleinthis
specimenunderthemicroscope.Thepartofitthatislocatedlateraltotheoccipitalhornisformedby
transverselycutfibres,whilstitsfibresdorsalandinferiortotheforcepsarecutlongitudinallyand
constitutetheadditiontothislayerthatcomesfromthecortexofthemedialoccipitallobe.Thebeaklike
extensionofthestratumintothegyruslingualis,whichwasalreadypresentontheprevioussection,can
stillbevisualisedhere.Thebeakappearsasatransversallycutfibrebundleunderthemicroscope.The
stratumsagittaleexternum(6)issimilartotheinternuminitsshape.Itsinferiorpartisfurtherthinnedand
bendduetothecollateralsulcus.Onthelateralaspectitisalreadyvisibletothenakedeyethatthelayeris
disappearingduetothevariouspenetrationsofthinbundlesoffibresdesignatedtoreachtheforceps.In
theinferiorpartthefibresaretransverselycutwhilstinthedorsalparttheyarecutaslant.Whencomparing
thissectiontothepreviousone,theformationofbundlesfromforcepsfibresisevidentintheregion
betweenthestratumsagittaleinternumandtheexternum.Thestratapriopriaoftheinterparietalsulcus(10)
andthecollateralsulcus(12)areclearlydistinctfromdeeplayersofthecortex(9),asthelatterisstained
lighter.Thedorsalpartofthestratumsagittaleexternumiscoveredbyacapthatappearsdarkercompared
tothesurroundingfibres.Theselighterfibresaretheanteriorremnantofthestratumtransversumcunei,
whichwilldisappearmoreinteriorlytogetherwiththecuneus.
5. (Enlargement9/8)Thiscutislocatedapproximately5mmanteriortotheprevious,approximately
65mmawayfromtheoccipitalpole,andonlyfewmillimetrebeforetheposteriorpartofthecorpus
callosum.Thissectionthereforecoversentirelytheparietallobe.Theremnantofthecuneusthatwas
stillvisibleontheprevioussectionhasnowdisappearedandmaderoomforthedescendingpartof
thecingulategyrus(VIII).Dorsaltothistheprecuneus(IX)iscutalongitslargestdiameter.With
regardstothefissuresontheconvexity,theinterparietalsulcus(i)iscutdiagonallyandthe
ascendingbranchoftheparallelsulcus(e.)iscutlongitudinally.Underneaththelatteronecan
appreciatethetransverselycutsecondandthirdtemporalsulcus.Onthebasalaspectonecanseethe
collateralsulcusagaintheindentsthestratumexternumandonthebordertotheinferiormedial
aspecttheanteriorandsharedpartofthecalcarinefissurewiththeoccipitoparietalsulcus(f.c.).
Thebasalaspectisreducedinsizeinrelationtotheothertwoaswellasinitsabsolutediameterandits
directiongotclosertothemedialsurface,meaningitismorevertical.Theconvexityontheotherhandis
approachingthehemisphericmidlineinferiorjustasitalwayshasdonesuperiorly.Asaconsequenceof
thesedramaticchangesinthearrangementofthegrossanatomythesubcorticalanatomyofthewhite
matterandtheoccipitalhornisrendered.Theoccipitalhorngainedinwidthandheightandhasfourwalls
asitdidontheprevioussection.Amongstthesewallstheinferioroneisverythinandcorrespondstothe
lateralpartoftheinferiorwallfromtheprevioussection.Themedialpartwiththeadjacentcollateral
sulcusisnowthemedialwall.Thedorsalwallis,similartotheprevioussectionbutmoreprominently
indentedduetothedorsalforcepspart.Thissectionshowsthetransitionoftheoccipitalhornintothe
lateralventricles.
Thedorsal(1)andventral(2)forcepspartgainedinvolume.Theventralpartprojectsdorsalalongthe
innersurfaceoftheoccipitalhornandisthereforeonlyseparatedfromthedorsalpartbyathingap.The
fibresoftheinnerforcepslayermergedwithit.Additionallyfibresoriginatingfromtheinferiorconvexity
arejoiningtheforcepsviathemedialwalloftheoccipitalhorn.Likewisethedorsalforcepspartgains
volumefromthenowprominentlayeroffibresthatareascendingverticallyalongthelateralsurfaceofthe
occipitalhorn(3).Bothadditionsarecutlongitudinallyonthissectionwhilstallotherforcepsfibres,
includingthesmallbundleattheinferiorborderoftheoccipitalhorn,arecuttransversely.

Bothsagittallayersmigratedawayfromthemedialsurfaceofthedorsalforcepstoallowthefibres
destinedforthespleniumtopiercethrough.Themajorityofthestratumsagittaleinternum(4)islocated
lateralandtoasmallerextentinferiortotheoccipitalhorn.Onthissectiononecanappreciatemedial
corticalfibresrunningdorsalandventraltotheforcepstowardsthislayer(5).Thestratumsagittale
externum(6)tightenedtowardsitsbaseventraltotheoccipitalhorn.Medialanddorsaltothedorsal
forcepspartnofibresofthislayerareseenonthissection.Thedirectionalityofthefibresisexactlythe
sameasontheprevioussection.Withthecalvaravisthebeaklikeprotrusionsofbothsagittallayers
vanished.Thewhitematterofthecingulategyrus,namelythecingulum,iscutlongitudinally(7)atits
medialaspectwhereitdescendsbehindthecallosum.Thecingulumisstaineddarkhereandtherefore
easilydifferentiated.Thecorticalwhitematterlayersareprominentandslightlydarkerintherestaining.
Theseincludethestratapropriaofthesulcuscollateralis(10),theprecuneus(8),andthefissure
interparietalis(9).
Itshouldbenotedthatthedorsalandlateralareasofthisspecimensaregenerallydarkerstainedcompared
totherest.Thereasonforthisirregularitymightbefoundintheirregularhardeningofthebrainaswellas
theverystrong(andthereforenotnecessarilyeven)destainingnecessitatedbytheintenttophotographthe
sections.
6. Thiscutislocatedapproximately10mmanteriortotheprevious,approximately75mmawayfrom
theoccipitalpole,andanteriortothebrainstructuresofthisexamination.Theintentistoindicate
thesubsequentwhitemattertrajectory.Thissectionshows(i)theposteriorpartofthecentralsulcus
(I)dorsally,(ii)theremnantoftheSylvianfissure(f.s.)laterally,and(iii)thecallosum,fornixand
theposteriorpartofthehippocampusmedially.Withregardstothesulcalanatomy,apartfromthe
Sylvianfissure,theinterparietal(i)andparallelsulcus(e.)aswellasthesecondandthirdparietal
sulci(s.t.IIandIII)areseenonthelateralconvexity.Onthemedialsurfaceonecanappreciatethe
callosomarginalsulcus(cm)dorsallyandthecollateralsulcusventrally.Thecalcarinefissure
alreadyterminatedpriortothissection.
Theoccipitalhorntransitionedintothedescendingpartofthecellalateralisofthelateralventricle,which
isonlyseparatedfromthecorticalsurfaceventrallythroughthefimbriaeofthefornix(12)thatarerunning
intothehippocampus.Thefibresoftheforcepsarefreedfromthewhitematterandthecortex,whichwere
stillseparatingitfromthemidlineontheprevioussection,andarenowlocateddorsomediallytothe
ventricleinthesplenium.Thedorsal(1)andventral(2)partoftheforcepscanstillbeseparated.The
verticallyascendingfibres(3)lateraltotheventriclearenotpartoftheforcepsinthissectionanymorebut
belongtothetapetumofthetemporallobe.Thetapetumhasaposteriorprotrusionandisthinneddueto
thedescendingpartofthecaudatenucleus,whichisnotvisibleonthissection.Thedorsalregionofthe
tapetumisfilledwithcorticalfibresthatpiercethenextlayer(**).Thefibresofthestratumsagittale
internum(4)areallcollectedonthelateralsurfaceoftheventricleandlateraltothetapetum.Thedotted
appearanceinthemiddleofthislayer(4*)isduetomergingwithotherbundlesfromthelateralaspectof
thestratumsagittaleexternumthatarestilldarkerandthereforedifferentiatefromthefibresofthestratum
sagittaleinternum.Underthemicroscopeeachofthesebundlesshowsaropeliketwistarounditsown
axis.Thewholelayerrepresentstheposteriorpartofthebaseofthecoronaradiataandgainsfibres
ventrallyfromthetemporallobeanddorsallyfromtheparietallobe.Thestratumsagittaleexternum(5)is
nowlimitedtotheventralpartoftheventricleintheregionofthetemporallobeandthinsoutasitsends
fibresofftothetemporalcortex.Towardsthehippocampalgyrus,thestratumsendsaprotrusionthatis
long,thin,andastillindentedbythecollateralsulcus.Theterminationofthisprotrusionisjoinedbythe
cingulum.LateraltotheventricleitextentsbarelyuntiltheSylvianfissureasitsdemarcationfadesaway.
Theelongationsofthecorrespondinglayersofthestratumverticalconvexitatisarethestratapropriaofthe
interparietal(9)andparallelsulcus(11)aswellasthewhitematteroftheSylvianfissure(10),whichare
alldarkerstained.Thecortexiscloselyapproachingthecoronaradiataoftheoccipitallobebyafew
millimetresatthedeepestareaintheSylvianfissure.Dorsaltothespleniumatransversecutof
longitudinalfibresshowsthecingulum(7)reachingintothecingulategyrus.Ontheprevioussectionthe
cingulumwascutalongitsdescendinglength.Thelighterareabetweenthelayersoftheinterparietal

sulcusandtheSylvianfissureindicatethelocationofthesuperiorlongitudinalbundleorarchingbundle
(6).Similartotheprevioussection,thedorsalandlateralareasofthisspecimenaredarkerstained
comparedtotherest.
7. Thissectionistakenfromadifferentseriesfromanatrophicfemalebrainofanelderlylady.This
sectionclearlydemonstratesthetriplelayeringoftheoccipitalhorn,includingitsinternalsurfaces,
andtheareabetweenthehornandthecalcaravis(VI.).Thissectionisalsoacoronalcutandistobe
placedbetweentheprevioussections4and5,onlyslightlyanteriortothesection4.The
correspondingphotographydemonstratesthemedialaspectsinaroughlyfourfoldenlargementand
correspondstothesquarethatisindicatedintheschematicdiagramofthesamesection.Thestemof
thecuneus(VII.),whichwasstillvisibleonthesurfaceonsection4,withdrewhereintothedepthof
theoccipitoparietalissulcus(o.)andthecalcarinefissure(f.c.).Thecrosssectionofthewhite
matterisidenticaltosection4,onlythatitispossibletoidentifythemedialforcepslayer(2),the
stratumsagittaleinternum(5),andexternum(6)alongtheentirecrosssectionofthemedialaspect
oftheoccipitalhornandbetweenthehornandthecalvaravis.Thesecanallbeseenwiththenaked
eye.
Thecrosssectionoftheoccipitalhornissquaredwiththedorsalsurfacebeingformedbythedorsal
forceps,themedialsurfacebythecalcarinefissure,andtheinferiorsurfacebythecollateralsulcus.The
dorsalforceps(1)isratherprominent,theventralforcepsratherweak(4),whilstthemedialforcepslayer
(2)isrelativelystrongandequallythickasthelaterallayer(3).Themajorityoffibresofthestratum
sagittaleinternum(5)arecollectedlateraltotheventricle,whilstthefibresoftheexternumarecollected
ventraltoit(6).However,fibresofthelatterarestilltobefoundinthelingualgyrusandtoasmaller
degreeinthestemofthecuneus.Itispossibletotraceaveilfrombothlayersacrossthemedialsurfaceof
theoccipitalhornwiththenakedeye.
8. Thisphotographyshowsacoronalsectionthroughthetemporallobeofabrainthatsuffereda
stroke.Asaconsequenceofthestroketheoccipitalcortexandapartofthetemporalcortex,
especiallythefirsttemporalgyrus,ipsilateraltothelesionaredamaged.Thelevelofthissectionis
comparabletosection5.Thebrainstemwasremovedpriortohardeningthisspecimen.A
ramificationoftheremovalisthatthetemporallobeanatomywasalteredandthecortexshifted
moremedial.Theareaofthecutshowingthecoronaradiataofthetemporallobeisbendmedially
andalmostreachesthehippocampus,whichcausedtheunusualformofthelateralhorn.
TheconvexityofthissectionshowstheSylvianfissurelaterallyandthesulcushippocampi(h.)medially.
Withinthesectionthefollowingstructuresareevident:theparallelsulcus(e.),thesecondandthird
temporalsulci(s.t.II.andIII.),andthecollateralsulcus(coll.),whichindentsthelateralhornfromthe
insidetowardstheeminentiacollateralisMeckelii.Themajorityofthefirsttemporalgyrusandsomeofthe
secondtemporalgyrusareaffectedbythestroke.Fromthesecondtemporalgyrusabrightlayerof
degeneratedfibresrunstowardsthewhitematerditch.
Fromtheoccipitalhornasmallremnantofthetapetumispresent(1)lateraltoitliesthewellmaintained
coronaradiataofthetemporallobe(2)whosepropagationintotheelongationstratumsagittaleinternumis
cutoff.Theveryprominentdarktransversecutthroughthestratumsagittaleexternum,asweknowitfrom
thehealthybrain,isabsent.Instead,wecanappreciateabrightzonewithroughlythesameshape,which
canbefollowedlaterallytotheconvexityandmediallyalsoasasmallbrightstripeuntilthehippocampal
gyrus(3).Underthemicroscope,onlyinsignificantremnantsofwhitemattercanbeseenwithinthiszone.
Thestrokeoftheoccipitallobethereforecausedadegenerationoftheentirestratumsagittaleexternumin
thetemporallobe.Amarkedcontrastisthecinguluminthegyrushippocampi,whichusuallyjoinsthe
stratumsagittaleexternumandisnowstaineddeepblack(Plates14).
Footnotes
2
Inparalyticbrainsthisareaappearsasabrightpatchindegenerativecorticalchanges.Inthesecasesitisfocally
degenerated,neverinitsentirelength(Lissauer).
3

3Notefromtheauthors:TheoriginaltextherementionsBogenbuendel,whichverbatimtranslatestoarchingbundle.
WhenlookingatOnufrowicz'soriginalwork,itbecomesapparentthatheequatedthesuperiorfrontooccipital
fasciculuswithBurdach'sarcuatefasciculus:ThisfibretractmayaptlybecalledFrontooccipitalis
Associationsbuendel[associativefrontooccipitalfasciculus]orcouldbereferredtoasthetruefasciculuslongitudinalis
superior.TheingeniousBurdach[]recognised,orratherguessed,thistractandcalleditthearcuatefasciculusor
superiorlongitudinalfasciculus.Yetneitherhis[Burdach's]norMeynert'sdescriptionsofthistractareobviousanditis
virtuallyimpossibletoproveit[frontooccipitalfasciculus]inthenormalbrain.Wewereabletoappreciatethistract,
locatedamongstcallosalradiationsinthenormalbrainonlyaftercomparisonwithourexperimentofnature,inthecase
ofagenesisofthecorpuscallosum.TranslationinForkeletal.(2014).

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FiguresandTables
Fig.1

Fig.2

Fig.3

Plate1

Plate2

Plate3

Plate4