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Walter Rudolf Hess Leben und Werk

Anlässlich des 70 Jahre Jubiläums der Verleihung des Nobelpreises

von Anton Valavanis und Alexander Borbély

Klinisches Neurozentrum USZ 2019 Impressum:

Herausgeber Klinisches Neurozentrum, Universitätsspital Zürich

Copyright Copyright © 2019 Klinisches Neurozentrum, Universitätsspital Zürich, 8091 Zürich, Schweiz

Gestaltung Susanna Sigg, Klinisches Neurozentrum, Universitätsspital Zürich

Text Anton Valavanis, Alexander Borbély

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Auflage 200

Adresse Klinisches Neurozentrum Zentrumsadministration Frauenklinikstrasse 10, 8091 Zürich Telefon +41 44 255 56 20 [email protected], [email protected]

Website www.neurozentrum.usz.ch

Inhaltsverzeichnis

1. Vorwort 5

2. Leben und Werk von Walter Rudolf Hess 7 a) Biographische Daten von Walter Rudolf Hess 9 b) Autobiographie von Walter Rudolf Hess 11 c) Hess CW: Walter Rudolf Hess (17.3.1881-12.8.1973) 35 d) Akert K: Walter Rudolf Hess (1881-1973) and his Contribution to 43 e) Waser PG: Walter Rudolf Hess: aus seinem Leben und seiner Tätigkeit an der Medizinischen Fakultät der Universität Zürich (100-Jahr-Feier seines Geburtstages: 14. März 1981) 59 f) Jung R: Walter R. Hess (1881-1973) 67

3. Der Nobelpreis für Walter Rudolf Hess 89 a) Nominationen für die Nobelpreisverleihung an Walter Rudolf Hess 91 b) Ankündigung der Nobelpreisverleihung in der Zeitschrift NATURE 93 c) Olivecrona H: Award Ceremony Speech 95 d) Hess WR: Nobel Lecture: The Central Control of the Activity of Internal Organs 99 e) Hess WR: Funktion und nervöse Organisation der inneren Organe 105 f) Wyder M: Walter Rudolf Hess: Nobelpreis für Medizin oder Physiologie 1949 121

4. Walter Rudolf Hess und die Tradition der Zürcher Neurowissenschaft 125 a) Valavanis A: Zur Tradition der Zürcher Neurowissenschaft: vom «Monakowschen Kränzchen» zur von Walter Rudolf Hess gegründeten «Zürcher Arbeitsgemeinschaft für Hirnforschung» und darüber hinaus 127 b) Valavanis A: Zur Interaktion zwischen Walter Rudolf Hess und M. Gazi Yasargil 133 c) Borbély A: Beiträge von Walter Rudolf Hess zur Schlafforschung 135 d) Huber A: W. R. Hess, the ophthalmologist 137 e) Puelles L: Hess’s experiments on the diencephalon and hypothalamus in the light of 141 modern neuromeric genoarchitectonics

5. Von der Neurowissenschaft zur Neurophilosophie 145 a) Hess WR, Fischer H: Brain and Consciousness: A Discussion About the Function of the Brain 147 b) Hess WR: Causality, Consciousness, and Cerebral Organization 157 c) Hess WR: Biological Order and Human Society 161

6. Das Vermächtnis des Hirnforschers Walter Rudolf Hess 175 a) Valavanis A: Das Vermächtnis von Walter Rudolf Hess und kritische Gedanken zur gegenwärtigen Situation der Neurowissenschaft 177

7. Die Bibliographie von Walter Rudolf Hess 187

8. Nachrufe auf Walter Rudolf Hess 203 4 1. Vorwort

Professor Walter Rudolf Hess (1881-1973), herausragenden Pioniere der tiefen Hirn- zunächst freipraktizierender Augenarzt, stimulation, gehalten. dann Direktor des Physiologischen Insti- tutes der Universität Zürich (1914-1951), Mit dem vorliegenden Band würdigt das hat sich der Hirnforschung verschrieben Klinische Neurozentrum Walter Rudolf und enge Verbindungen zum damaligen Hess als eine aussergewöhnliche Persön- Kantonsspital Zürich gepflegt. Als Prota- lichkeit, die selbstkritisch, bescheiden und gonist der universitär verankerten Zürcher unbeirrt von den jeweiligen Konjunktur- Neurowissenschaften erhielt er 1949 den strömungen während einer 40-jährigen Nobelpreis für Medizin für seine experi- Forschungstätigkeit beharrlich ihr Ziel mentellen Untersuchungen über die funk- verfolgte. Sein umfassendes neurowis- tionelle Organisation des Zwischenhirns. senschaftliches Wissen, seine intellektuel- Damit jährt sich in diesem Jahr zum 70. le Kreativität und methodische Präzision Mal die Verleihung des Nobelpreises an wurden mit der Verleihung des Nobelprei- Walter Rudolf Hess. ses gekrönt.

Aus Anlass des 70-Jahre Jubiläums dieses Der Band enthält eine Zusammenstellung besonderen Ereignisses würdigt das Klini- ausgewählter Veröffentlichungen von und sche Neurozentrum des Universitätsspitals über Walter Rudolf Hess, die sein Leben, Zürich das Leben und Werk des Nobel- seine Persönlichkeit, seine neurowissen- preisträgers Hess mit einer Sondersitzung schaftlichen Leistungen und neurophilo- im Rahmen seines diesjährigen Jahressym- sophischen Gedanken beleuchten. Er dient posiums, welches am 14. November 2019 auch dazu, die Erinnerung an diesen Pionier im Monakow Hörsaal des Klinischen Neu- und Förderer der theoretischen und klini- rozentrums zum Thema ‘Tiefe Hirnstimu- schen Neurowissenschaft wach zu halten. lation/Deep Brain Stimulation‘ stattfindet. Die Ursprünge der klinisch angewandten Die Herausgeber danken dem Enkel von tiefen Hirnstimulation finden sich in den Walter Rudolf Hess, Christian W. Hess, tierexperimentellen Untersuchungen von Professor emeritus der Universität Bern Walter Rudolf Hess und stellen ein frühes und ehemaligen Direktor der Neurologi- Paradigma der translationalen Forschung schen Klinik des Inselspitals Bern, für sei- in der Neurowissenschaft dar. Die tradi- ne wertvollen Hinweise sowie der Sekre- tionelle «Yasargil Lecture», die jeweils im tärin des Klinischen Neurozentrums des Rahmen des Jahressymposiums stattfin- USZ, Frau Susanne Sigg, für die hilfreiche det, wird von Professor Alim Louis Bena- Unterstützung bei der Erstellung und Ge- bid, Professor emeritus der Universität staltung dieser Festschrift. Grenoble und Membre de l’Institut de France (Académie des Sciences), einem der Anton Valavanis Alexander Borbély

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2. Leben und Werk von Walter Rudolf Hess

7

Biographische Daten von Walter Rudolf Hess

1881 Born March 17 in , 1900-1905 Studied medicine in Lausanne, Bern, , , and 1906 Graduated as M.D. in Zurich 1906 Assistant in Surgery, Kantonsspital Münsterlingen (Conrad Brunner) 1907-1908 Assistant in Ophthalmology, (Otto Haab) 1908 Married Louise Sandmeier; children: Gertrud (1910) and Rudolf Max (1913) 1908-1912 Practiced Ophthalmology in Rapperswil 1912-1913 Assistant in , University of Zurich (Justus Gaule) 1913-1917 Privatdozent, University of Zurich 1915 Assistant in Physiology with Max Verworn in Bonn 1917-1951 Professor of Physiology and Director of the Physiology Department in Zurich 1931-1937 Cofounder (1931) and Chairman of the Board of the International High Altitude Research Laboratory at Jungfraujoch 1932 Prix Marcel Benoist (Switzerland) 1934 Doctor honoris causa, University of Berne 1938 President of the Sixteenth International Physiological Congress held in Zurich 1938 Carl Ludwig Medal of the German Society of Circulation Research 1944 Doctor honoris causa, 1949 1953 Doctor honoris causa, McGill University, Montreal 1959 Doctor honoris causa, im Breisgau 1967 Moved to Ascona, Switzerland 1973 Died , in Ascona, Switzerland

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From Medical Practice to Theoretical Medicine: An Autobiographic Sketch FROMMEDICAL PRACTICE TO THEORETICAL MEDICINE Walter Rudolf Hess AN AUTOBIOGRAPHIC SKETCH Nachdruck der vom Deutschen ins Englische übersetzten Autobiographie, erschienen 1963W. inR. derHESS* Zeitschrift Perspectives in Biology and Medicine, 6 (4): 400-423, 1963

Ancestry FROMMEDICALI should likePRACTICEto begin thisTOshortTHEORETICALsketch of myMEDICINElife with a few comments ANconcerningAUTOBIOGRAPHICmy forebears.SKETCHThis will come as no surprise to readers inter- ested in biology, who are well aware of the decisive influences one's in- heritanceW. R. HESS*has in the shaping of one's life. In my own case it is rather inter- esting, as two different folk-groups are represented in my pedigree. My father'sAncestry ancestors belonged to the Alemanni, a Germanic tribe ofthe upper RhineI shouldvalleylike to beginformerlythis shortundersketchRomanof my lifecontrol.with a fewIn severalcommentsexpansive move- concerning my forebears. This will come as no surprise to readers inter- estedmentsin biology,these peopleswho are drovewell awarethe ofRomansthe decisivesouthwardinfluencesandone'soverin- the Alps; they heritancethen occupiedhas in the shapingthe nowof one'sGerman-speakinglife. In my own caseregionit is ratherof northeasterninter- Switzer- esting,landasandtwoaredifferentconsideredfolk-groupsthe areprincipalrepresentedancestorsin my pedigree.of its presentMy population. father'sMy mother'sancestors belongedpeopletocamethe Alemanni,from atheGermanicregiontribeofofthethe formerupper kingdom of RhineSaxonyvalleywhereformerlyEastunderEuropeanRoman control.typesIn severalwere expansiveto be foundmove-along with the ments these peoples drove the Romans southward and over the Alps; they thenNordic.occupiedThisthe nowdivergenceGerman-speakingbetweenregiontheofmaternalnortheasternandSwitzer-paternal inheritance landgenerallyand are consideredis of a certainthe principalgeneticancestorsinterest.of its present population. My mother'sA ratherpeoplebroadcamerangefromofthetradesregion andof theprofessionsformer kingdomis to ofbe found in my Saxonyfamilywheretree.EastIt includesEuropeanfarmers,types werecraftsmen,to be foundandalongmerchantswith theas well as more Nordic. This divergence between the maternal and paternal inheritance generallydistantlyis ofrelateda certainclergymengenetic interest.and politicians who were rather gifted orators. A Myratherfather,broad rangea collegeof tradesteacherand professionsin physics,is combinedto be found ainremarkablemy sense of familyordertree.withIt includesa teacher'sfarmers,talent.craftsmen,Althoughand merchantsfundamentallyas well as morea very serious man, distantlyhe alwaysrelatedretainedclergymena andgoodpoliticianssense ofwhohumor.were ratherMygiftedmotherorators.had an altogether happyMy father,anda collegepleasantteacherdisposition,in physics, combinedwhich stooda remarkablein strikingsense ofcontrast to the orderratherwithsobera teacher'smoodtalent.andAlthoughreservedfundamentallyattitude soacharacteristicvery serious man,of the inhabitants he always retained a good sense ofhumor. My mother had an altogether happyof easternand pleasantSwitzerlanddisposition,at thatwhichtime.stoodItinisstrikingno wondercontrastthatto thewe had little con- rathertactsoberwithmoodour neighbors,and reserved attitudewho lookedso characteristicupon usofalmostthe inhabitantsas strangers with dif- of eastern Switzerland at that time. It is no wonder that we had little con- * Goldauerstrasse 25, Zürich, Switzerland. This autobiographic sketch was translated from the tactoriginalwith Germanour neighbors,manuscriptwhobylookedDr. E. uponCastagnoli,us almostan Americanas strangersphysician,with dif-presently in Zürich. * Goldauerstrasse 25, Zürich, Switzerland. This autobiographic sketch was translated from the original German manuscript by Dr. E. Castagnoli, an American physician, presently in Zürich.

W. R. Hess · From Medical Practice to Tlteoretical Medicine 4OOW. R.PerspectivesHess · FrominMedicalBiologyPracticeandtoMedicineTlteoretical·MedicineSummer 1963 4OO Perspectives in Biology and Medicine · Summer 1963 11 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

ferent customs and social standards. My mother also had an unusually lively disposition. She bubbled over with energy—so much so, in fact, that she often had a hard time falling asleep. I, too, was to suffer from this difficulty throughout my life. I have always had to struggle to shut out the impressions and experiences of the day. Early Environment The editors who invited me to write a short autobiography encouraged me to include a psychological background to my life story. This is in keep- ing with modern autobiographic writing and means that the role played by the external environment in developing, shaping, and limiting an individual's inherited capabilities must not be overlooked. My own childhood was provided with ample opportunities to become aware of nature at firsthand. My early curiosity about the world around me was stimulated and supported by my father. Even at the age of five, I used to explore the fields and meadows to collect plants, and every new specimen meant an exciting experience, for it was brought home and care- fully classified with Father's help. Soon I had my herbarium and a collec- tion of butterflies. As time went on, I became aware of the significance of the ecological setting, that is to say, of the specific interrelationship be- tween flora and fauna. I could see, for example, that a particular species of caterpillar was always to be found on a certain plant. Such observations were unforgettable. Even much later I remembered the exact place where they had occurred. More and more it became clear that functional mani- festations, such as the germination of a seed or the rapid sprouting of a shoot from a willow, were more apt to capture my mind than purely morphological features. Perhaps most fascinating to me was the observa- tion of the metamorphosis from caterpillar to chrysalis to butterfly. I just couldn't get enough of nature. How beautiful were those solitary walks through fields and forests and along streams and rivers. By the time I en- tered elementary school I had thoroughly discovered the countryside sur- rounding my village. Student Years The phase in my life that did much to form my personality began with my entrance into the Gymnasium at the age of twelve. My father soon al- lowed me to visit his physics laboratory and to help out in setting up the

401 12 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

experiments for his classes. This early contact with classical physics was of great value to me in later life. It was during this time that I saw how the seemingly stationary processes in so-called static systems were in reality a system of antagonistic forces resulting in a dynamic equilibrium. During my free time I used to make toys such as bows and arrows, sail boats, and airplanes from improvised materials to be found in and around experimentsthe house.forThishis classes.did muchThis earlyto developcontact withnotclassicalonly manualphysics wasskillsof but also a cer- greattainvaluepracticalto me insenselater andlife. Itinventiveness.was during this timeOn thatthe Iothersaw howhand,the I had to over- seeminglycome manystationarydifficultiesprocesses within so-calledmy studies.static systemsIt waswerehardin forrealityme to remember a systemisolatedof dataantagonisticwhichforcescouldresultingnot bein aassociateddynamic equilibrium.with significant events. I also During my free time I used to make toys such as bows and arrows, sail boats,hadandtroubleairplanesmemorizingfrom improvisedthe materialsvocabulariesto be foundin Greek,in and Latin,around and French. In theshort,house.inThisthedidlinguisticmuch to developsubjectsnotI wasonly manualnot a goodskills butstudent;also a cer-I was poor also in taincomposition.practical senseThroughoutand inventiveness.myOnlifetheI haveother hand,had difficultiesI had to over-in expressing my comethoughtsmany difficultiesin writing.withOnmythestudies.otherIt washand,hardI forhavemealwaysto rememberbeen able to express isolated data which could not be associated with significant events. I also hadmyselftroubleclearlymemorizingand convincinglythe vocabularies in Greek,speaking.Latin,Whenand French.involvedIn in an inter- short,estingin thedebatelinguisticor subjectsdiscussion,I was nottheawordsgood student;flow Ifreely.was poorButalsowhenin I try to put composition.these sameThroughoutideas on mypaper,life IIhaverunhadintodifficultiestrouble,in expressingthe sentencesmy usually being thoughtstoo longin writing.and complicated.On the other hand,I haveI havesoalwaysoftenbeenenviedable to expressthe writing skills of myselfothers.clearly and convincingly in speaking. When involved in an inter- esting debate or discussion, the words flow freely. But when I try to put theseThesamequestionideas on paper,naturallyI run intoarisestrouble,howtheit camesentencesaboutusuallythatbeingsomeone like my- tooself,longwhoseand complicated.true interestsI havelie soin naturaloften enviedscience,the writingshouldskillscomeof to study medi- others.cine. I think that this can be attributed to the influence ofour family doctor The question naturally arises how it came about that someone like my- self,whowhosetreatedtrue interestsme forlie inpleurisy.natural science,This shouldbout comewithtotuberculosisstudy medi- cast the only cine.shadowI thinkonthatmythis otherwisecan be attributedhappyto theyouth.influenceBeingofourveryfamilylivelydoctorby temperament, whoit wastreatedallmetheforharderpleurisy.forThismeboutto withheedtuberculosisthe warningscast theagainstonly overexerting shadowmyself.on my otherwise happy youth. Being very lively by temperament, it was all the harder for me to heed the warnings against overexerting myself.I decided during the last semesters in the Gymnasium on the path I wouldI decidedtakeduringfollowingthe last semestersgraduation.in theTheGymnasiummedicalonprofessionthe path Iseemed to offer woulda coursetake followingof traininggraduation.which Thewouldmedicalnotprofessionbe too severeseemedintoitsofferdemands. In addi- a coursetion, bothof trainingseeingwhichthewouldapplicationnot be tooofseveresciencein itsfordemands.the benefitIn addi-ofpeople and the tion, both seeing the application ofscience for the benefit ofpeople and the doctor'sdoctor'srolerolein theincommunitythe communityappealed toappealedme. to me. UniversityUniversitylife openedlife openedup new updimensionsnew dimensionsfor an essentiallyfor ansmall-townessentially small-town boy.boy.FirstFirstof all,ofI all,had Itohaddecidetoatdecidewhich universityat whichI universitywould study.I Mywould study. My firstfirstchoicechoicewas thewasUniversitythe UniversityofLausanne,ofLausanne,located in thelocatedFrench-speakingin the French-speaking areaareaof Switzerland.of Switzerland.There ITherewould Ialsowouldhave thealsoopportunityhave theofopportunitygetting to ofgetting to knowknowthe theessentiallyessentiallydiffèrentdiffèrenttraditionstraditionsand customsandofmycustomsfellow citizensofmy fellow citizens

W. R. Hess ' From Medical Practice to Theoretical Medicine 402 PerspectivesW. R.inHessBiology' andFromMedicineMedical· SummerPractice1963to Theoretical Medicine 402 Perspectives in Biology and Medicine13· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

who were ofRoman descent. Following the first year, I took advantage of an opportunity to study for one semester in the capital city ofBern, where I was exposed to the well-functioning democracy of my country. I then went to Zürich, where I completed the preclinical courses in anatomy, histology, biochemistry, and physiology. The urge to travel—inherent in all youth—overcame me and I went to Berlin, at that time still under the Kaiser's rule. In addition to its famous clinics, Berlin had many interesting things to offer, such as museums, a zoological garden, an oceanarium, and an astronomical observatory. The countryside of Berlin, with its wide plains and open spaces, was especially interesting and novel for a Swiss ac- customed to the hills and mountains ofhis country. Everything in the fields ofart and theater was also at one's disposal. Despite all these varied possi- bilities, I remained faithful to my love for nature. In 1903 I had an excellent opportunity to experience the sea in the Ger- man city ofKiel, whose harbor also served as a base for the German navy. How longingly my eyes followed the ships as they disappeared over the horizon into distant lands. For me, the most unforgettable personality on the medical faculty of Kiel was the internist Quincke. He had the talent ofpresenting his cases in a very interesting way; I missed hardly any of his lectures. I also learned a great deal of surgery under Helferich, who was an excellent teacher, al- though to be sure an air ofcynicism wafted through his lectures or demon- strations. The lectures in dermatology and venereology were excellent; the clinical case presentations were especially impressive, as more than ade- quate material was available from the port district. Following the completion ofmy studies in Berlin, and after having de- voted much time to personal scientific interests during the summer se- mester spent in Kiel, I returned to Switzerland. At the turn ofthe century, the length of study for doctors was ten semesters—that is, for an indus- trious student. Because I had devoted much time to my side interests, I felt that my prospects of passing the final examinations were rather poor. Nevertheless, I made up the classes I had missed in Berlin and Kiel by adhering to a rigid work schedule during the last two semesters in Zürich. I took no notes whatsoever, but gave my full attention to the lectures and demonstrations. For reading material, I consulted textbooks and atlases. This study procedure was supplemented with periodic discussions with my fellow students who were also preparing for final exams and who were

14 403 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

informed about the "specialties" of the examiners. I successfully passed the examinations in 1905 and received the diploma bestowing the right to practice medicine in Switzerland. Inklings ofResearch Looking back on my days in medical school, I can still remember one informedincidentaboutwhichthe "specialties"is worthyofofthemention.examiners. IDuringsuccessfullya dissectingpassed the session in the examinationsAnatomicalin 1905Instituteand receivedin Zürichthe diplomathe instructorbestowingcalledthe rightourto attention to an practiceanomalymedicinein thein Switzerland.arterial blood supply of the foot. The arteria dorsalis pedis was not formed as a direct continuation of the arteria tibialis anterior as it Inklings ofResearch usuallyLooking is.backInstead,on my daysa well-developedin medical school, anastomosisI can still rememberwith onethe arteria tibialis incidentposteriorwhichwasis worthypresent.ofThismention.fact,Duringpointeda dissectingout withoutsessionfurtherin the elaboration by Anatomicalthe anatomist,Instituteinterestedin Zürich themeinstructorand I askedcalledforouranattentionexplanation.to an The teacher's anomaly in the arterial blood supply of the foot. The arteria dorsalis pedis wasanswernot formedthatasthea directanomalycontinuationwas notofattheallarteriainfrequenttibialis anteriordid notas itsatisfy my scien- usuallytific curiosity.is. Instead, aI well-developedfinally gatheredanastomosisup enoughwithcouragethe arteriaandtibialispresented my own posteriorinterpretation.was present.ItThisseemedfact, pointedclear tooutmewithoutthatfurtherthe differentialelaborationresistanceby between thetheanatomist,arteriainterestedtibialis anteriorme and I askedand forposterioran explanation.must Thebe ofteacher'sconsiderable signifi- answercance.thatThethe anomalypath withwas nottheatleastall infrequentresistivediddropnot satisfyin pressuremy scien-carries the blood tific curiosity. I finally gathered up enough courage and presented my own interpretation.more effectivelyIt seemedtoclearthetoperipheralme that the differentialsegmentsresistancesince itbetweenmaintains the greater thepressurearteria tibialisreserves.anteriorIt isandthisposteriorpressuremustreservebe of considerablewhich procuressignifi- blood for the cance.vascularThe pathsystemwith theofleasttheresistivefoot viadrop thein pressureanastomosiscarries thebetweenblood two arterial morebranches.effectivelyThus,to thetheperipheralabovesegmentsmentionedsince anomalyit maintainsisthea directgreater example of the pressure reserves. It is this pressure reserve which procures blood for the vascularrelationshipsystem ofbetweenthe foot hemodynamicsvia the anastomosisandbetweenmorphogenesistwo arterial in the arterial branches.system.Thus, the above mentioned anomaly is a direct example of the relationshipI can nobetweenlongerhemodynamicsremember theand contentmorphogenesisof theindiscussionthe arterialwhich followed system.in the dissecting room. In any event, the question remained open and arose againI can noinlongerBerlinrememberin a diffèrentthe contentcontext.of the discussionThere I metwhichW.followedRoux, the anatomist in the dissecting room. In any event, the question remained open and arose againat thein Berlinnearbyin a Universitydiffèrent context.of Halle,There I metwhoW.wasRoux,thethefounderanatomistof the science of atdevelopmentalthe nearby Universitymechanics.of Halle, whoBecausewas theI wasfounderinterestedof the sciencein theof question, I was developmentalkeen to learnmechanics.the opinionBecauseofI wassuchinteresteda distinguishedin the question,scientist,I was who had also keenworkedto learnonthetheopinionproblemof suchofa distinguishedthe relationshipscientist,betweenwho hadhemodynamicsalso and worked on the problem of the relationship between hemodynamics and thethemorphologicalmorphologicalformationformationof the arterialof thetreearterialwith specialtreereferencewith specialto reference to thetheangleangleof itsofbranches.its branches.I thereforeI thereforepresented mypresentedideas to himmyinideasthe formto him in the form ofofa lettera letterand wasandverywashappyverytohappyreceivetoa replyreceiveinvitinga replyme toinvitingHalle formea to Halle for a personalpersonaldiscussion.discussion.As a result,As aRouxresult,suggestedRouxthatsuggestedthe theorythatbe pub-the theory be pub- lishedlishedin thein Archivthe Archivfür Entwicklungsmechanik.für Entwicklungsmechanik.It is understandableIt is understandablethat my that my interestinterestin scientificin scientificwork receivedwork receiveda strong stimulus.a strongIt ledstimulus.to a student'sIt led to a student's

W. R. Hess · From Medical Practice to Theoretical Medicine 404 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 404 Perspectives in Biology and Medicine15· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

paper entitled "Eine mechanisch bedingte Gesetzmässigkeit im Bau des Blutgefässsystemes" ("A Mechanically Induced Conformity in the Struc- ture of the Vascular System"). My concern with this problem led to further consequences, as the reader will discover later. Choice of the place for my first internship was dictated by financial cir- cumstances, which did not permit me to pursue a course ofstudy in theo- retical medicine, as I would have liked, following the completion of my training in 1905. From then on I had to earn my own living. My first as- signment was a residency in the state hospital of my home canton rather than an unpaid position in a university clinic. The department of surgery at the hospital was under the direction of Dr. Konrad Brunner, whose reputation extended beyond the borders of Switzerland. Retrospectively, I can see how much I profited from having worked in the well-disciplined team under Brunner. In addition to the routine daily chores, I came to understand and appreciate the skilled tech- nique and high sense ofresponsibility ofthis great surgeon. Furthermore, I had the opportunity to see parts of the vascular system in vivo, which in- tensified my early interests in hemodynamics. The relationship between the morphological organization of the vascu- lar system and the flow characteristics of blood factors became more ap- parent in determining circulatory phenomena. I could also see that in prac- tical medicine more attention was paid to the dynamic action of the heart pump than to peripheral factors such as the resistance and blood viscosity. The role of the latter was almost completely ignored, and therefore I de- cided to construct a convenient apparatus to measure blood viscosity for clinical usage. The problem was intriguing because the existing viscosi- metric apparatus (e.g., Ostwald) applicable to so-called idealized fluids could not be used for measuring blood viscosity. Due to the coagulability of blood, the measurements would have to be completed within 2 to 3 minutes, and, for practical purposes, I to 2 drops of blood—as used in de- termining hemoglobin values—would have to suffice. In addition, it was desirable to develop a procedure which would allow the results to be read offdirectly, thereby avoiding calculations. How to construct an apparatus which could fulfil all these requirements occupied almost all my free time. I would concentrate on the problem especially during walks. After I had probed many varied constructions in my mind, a possible solution all at once occurred to me. It had the character of a sudden inspiration whereby

16 405 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

the latent psychological tension becomes discharged. Unforeseen difficulties had to be overcome in order to actualize the mental concept into a practi- cal, usable instrument. Much patience was required and modifications had to be made often to arrive at the goal. At last the plan was fulfilled and measurements were made. The results and conclusions were published in a small treatise entitled "Viscosität des theBluteslatent psychologicalund Herzarbeit"tension becomes("Viscositydischarged.of theUnforeseenBlooddifficultiesand the Work of the hadHeart").to be overcomeDr. Konradin orderBrunner,to actualizemythe mentalchief,conceptwho freelyinto a practi-admitted that he had cal,anusableinadequateinstrument.knowledgeMuch patienceofwastherequiredproblem,and modificationsleft the futurehad of the work to be made often to arrive at the goal. entirelyAt last theinplanmywashands.fulfilledI thereforeand measurementspersonallyweresubmittedmade. Thetheresultspaper for publica- andtionconclusionsin Pßüger'swereArchivpublishedfürinPhysiologie.a small treatiseI hadentitledto undergo"Viscositätadesrather painful ex- Blutesperienceund Herzarbeit"because of("Viscositydie authoritarianof the Bloodattitudeand thethenWorkprevalentof the toward youth- Heart").ful scientists.Dr. KonradToday,Brunner,ofcourse,my chief,thewhonecessityfreely admittedand dutythat ofhe promotinghad and en- ancouraginginadequate youngknowledgepeopleof theinproblem,scientificleftresearchthe futureareofwellthe workestablished. Pflüger entirelywantedin mytohands.knowI thereforeunderpersonallywhose directionsubmitted thethepaperworkfor publica-had been carried out. tion in Pßüger's Archivfür Physiologie. I had to undergo a rather painful ex- perienceUponbecauseexplainingofdie authoritarianto him thatattitudeit hadthenbeenprevalentdonetowardentirelyyouth-on my own, he fulrejectedscientists.theToday,paperofcourse,withoutthe necessitycomment.and dutyFortunatelyofpromotingforandme,en-a member ofthe couraging young people in scientific research are well established. Pflüger wantedZürichto knowmedicalunderfacultywhosewhodirectionheardtheofworkthe hadsituationbeen carriedinterestedout. himselfon my Uponbehalf.explainingIt wastoProfessorhim that itZanggerhad been whodone entirelytook overon mytheown,printinghe of the work rejectedin Vierteljahresschriftthe paper without comment.der ZürcherFortunatelyNaturforschendenfor me, a memberGesellschaft.ofthe ZürichThemedicalmainfacultyfindingwhoinheardtheofpaperthe situationwas theinteresteddemonstrationhimselfon mythat the viscosity behalf.of theIt wasbloodProfessorvariesZanggerwith thewhonumbertook overofthecirculatingprinting of theerythrocytes,work which is in Vierteljahresschrift der Zürcher Naturforschenden Gesellschaft. oneThe mainof thefindingfactorsin thethatpapermaywasinfluencethe demonstrationperipheralthat theresistance.viscosity It further sug- ofgestedthe bloodthatvariesdecreasedwith theviscositynumber ofofcirculatingthe blooderythrocytes,can lead whichto turbulenceis and may onebeofresponsiblethe factors thatformaytheinfluenceso-calledperipheralanemicresistance.heart murmur.It further sug- gestedInthatthedecreasedfollowingviscosityyearsofthethebloodsciencecan leadof viscosimetryto turbulence andofmayblood plasma and beserumresponsibleexpandedfor the so-calledconsiderablyanemic and,heart murmur.in combination with refractometry, it In the following years the science of viscosimetry of blood plasma and serumbecameexpandedpossibleconsiderablyto differentiateand, in combinationthe albuminwith refractometry,and globulinit fractions in becameplasmapossibleor serum.to differentiateAn increasingthe albuminclinicaland interestglobulin fractionsin bloodinviscosity mani- plasmafestedoritselfserum.betweenAn increasingthe clinicalyears interestof 1908in andblood1920.viscosityThereaftermani- interest de- festedclined,itselfprobablybetween thebecauseyears of more1908 andconvenient1920. Thereafterand interestpracticalde- methods (e.g., clined, probably because more convenient and practical methods (e.g., measurementmeasurementof theofsedimentationthe sedimentationrate) had beenrate)developed.had beenNeverthe-developed. Neverthe- less,less,forforspecialspecialcases, cases,interestinterestin the viscosimetryin the viscosimetryof blood and otherof bloodbody and other body fluidsfluidshashasremainedremaineduntil theuntilpresentthetime.present time. DuringDuringmy firstmyyearfirstofyearinternshipof internshipI was naturallyI wasconcernednaturallyaboutconcernedmy about my future, so I set a firm goal for myself. I wanted to work in some branch of medicinefuture,whichso I setwoulda firmgivegoalme anforadequatemyself.incomeI wantedwhiletoatworkthe samein some branch of medicine which would give me an adequate income while at the same

W. R. Hess · From Medical Practice to Theoretical Medicine 406 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 406 Perspectives in Biology and Medicine17· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

time allowing me sufficient time to devote myself to problems of basic research. A practice in ophthalmology seemed ideal to meet these two re- quirements. I therefore seized the first opportunity to enter the department of ophthalmology at the University of Zürich as a resident early in 1906. My new chief, Professor O. Haab, was an outstanding clinician and skilled surgeon. The somewhat one-sided morphological orientation prevalent in the clinic did not quite suit my views; however, I had ample opportunities to make diagnostic observations of a more dynamic character—for ex- ample, in analyzing oculomotor disturbances. According to the technique employed at that time, the examiner relied on the patient's report as to the site of the double image when a moving light with changing directions was flashed before his eyes. With such a method the examiner was, of course, too dependent on the patient's intel- ligence, and quite often an exact diagnosis had to be foregone. I decided, therefore, to develop a method which would give more objective results concerning the co-ordination ofeye muscles. I shall not describe the proce- dure in detail. Suffice it to recall that by employing an apparatus called "co-ordimeter," it was possible to determine the motor defect quantita- tively. Furthermore, it was possible to evaluate with satisfactory accuracy the effects of treatment through repeated controls. The apparatus has un- dergone improvements and modifications since its inception, but the novel principle has held its own till now. More important to me personally was the clear insight I gained into the problem ofmotor co-ordination which— many years later—added greatly to my understanding of the analysis of motor responses during stimulation in the region of the thalamus and subthalamus. The same was true of a thorough investigation of stereoscopic vision, which presents an instructive example of the integrative function of the central nervous system. In conjunction with the above interests, I tried to develop a procedure for making stereoscopic photographs, with the hope that I might capitalize on it and obtain thereby a certain degree offinancial freedom. In fact, I succeeded in making three-dimensional transparencies which gave a real view into the depths of the object. The pictures, how- ever, lacked certain desirable qualities which I was trying to incorporate when World War I halted my attempts. At the end ofthe war my position was more secure and other problems were occupying my attention. Before the war I had completed my residency in ophthalmology and

18 407 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

had furthered my education through a briefperiod of study in Paris. Soon thereafter I took over the practice of an ophthalmologist in an attractive city and important railroad intersection in the vicinity of Zürich. By the spring of 1908 I was living rather comfortably as a practicing eye specialist and was, therefore, in a position to marry my fiancée, Louise Sandmeier, who had worked in the outpatient department of the Zürich Eye Clinic. hadAsfurthereda formermy doctor'seducation throughaide, shea briefwasperiodwellofpreparedstudy in Paris.to offerSoonefficient help in thereafterher husband'sI took overoffice,the practiceand thusof anweophthalmologistwere fortunatein anenoughattractiveto overcome our cityfinancialand importantdifficultiesrailroadforintersectionthe timein being.the vicinity of Zürich. By the spring of 1908 I was living rather comfortably as a practicing eye specialist and was,Leaptherefore,into theinUnknowna position —to Newmarry Yearsmy fiancée,of TrainingLouise Sandmeier, who had worked in the outpatient department of the Zürich Eye Clinic. As a formerThe securitydoctor's aide,and shecompensationwas well preparedderivedto offerfromefficientan extensivehelp in practice had hernegativehusband's aspects.office, andIthushadwesetwereup fortunatea laboratoryenoughintowhichovercomeI spentour my free time financialworkingdifficultieson problemsfor the timeofbeing.hemodynamics, particularly in relation to blood viscosimetry,Leap into the Unknownand on—NewproblemsYears of ofTrainingmotor co-ordination. The ever increas- ingThe demandssecurity andofcompensationthe practice,derivedand especiallyfrom an extensivethe manypracticeprofessionalhad consulta- negativetions andaspects.opinions,I had set uplefta melaboratorywith lessin whichandIlessspenttimemy freeto pursuetime my scientific workinginterests.on problemsI wouldof hemodynamics,come homeparticularlyfrom stimulatingin relation toscientificblood meetings in viscosimetry, and on problems of motor co-ordination. The ever increas- ingZürichdemandsandofthefeelpractice,downrightand especiallyunhappythe manyandprofessionaldissatisfiedconsulta-because my hands tionswereandtied.opinions,A deepleft meconflictwith lessresultedand less thentime tobetweenpursue mymyscientificsense ofduty to pro- interests.vide forI wouldmy familycome homeand frommy longingsstimulatingforscientificpure meetingsscience. inFinally, a critical Zürichdecisionand feelhaddownrightto be made,unhappyas a positionand dissatisfiedbecamebecauseavailablemy handsfor me at the Insti- were tied. A deep conflict resulted then between my sense ofduty to pro- videtuteforofmyPhysiologyfamily and myin Zürich.longings forI canpurerememberscience. Finally,so wella criticalthe exact spot in my decisiongardenhadwhereto be made,I decidedas a positionin favorbecameof science,available fordespiteme at the Insti-fact that I now also tutehadofPhysiologya child toinsupport,Zürich. Iandcan rememberthat the sodecisionwell the entailedexact spotainconsiderablemy reduc- gardentionwherein income.I decidedOfin favorcourseof science,I discusseddespite thefactproblemthat I nowbeforehandalso with my had a child to support, and that the decision entailed a considerable reduc- tionwife,in income.who agreedOf coursetoIthediscussedstep whichthe problemwasbeforehandso significantwith formy my later life. wife, Thewho agreedmovetowasthe madestep whichin 1912.was so Itsignificantwroughtfor mentalmy later tensionslife. which were notThe alwaysmove waseasymadeto inbear.1912.MyIt wroughtnew teachingmental tensionsassignmentwhichmadewere it necessary for notmealwaysto revieweasy to bear.scientificMy newmaterialteaching assignment—old andmadenewit necessary—and muchfor was actually melearnedto reviewwhilescientificteaching.materialBecause—old andofnewthe—andpoormuchhealthwas ofactuallymy chief, Professor learned while teaching. Because of the poor health of my chief, Professor Gaule,Gaule,a pupila pupilof CarlofLudwig,Carl Ludwig,personal contactpersonalwithcontacthim for purposeswith himof for purposes of scientificscientificdiscussiondiscussionwas ratherwaslimited,ratherandlimited,I was moreandorI wasless onmoremy own.or less on my own. I foundI foundno lackno oflackpressingof pressingproblems problemsto tackle. Predominantto tackle. amongPredominant among thesethesewaswasthe economythe economyof energyofinenergythe mechanicsin theofmechanicscirculation, ofwhichcirculation, which presents a dynamic parallel, for example, to the architectural patterns of spongeouspresentsbone,a dynamicknown toparallel,be builtforso example,as to insuretooptimalthe architecturalbearing patterns of spongeous bone, known to be built so as to insure optimal bearing

W. R. Hess · From Medical Practice to Theoretical Medicine 408 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 408 Perspectives in Biology and Medicine19· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

strength. Take the head of the femur as an example. While the conditions obtaining in hemodynamics are not so obvious as in static systems, they are, nevertheless, offundamental significance. In order to understand prop- erly the hemodynamic situation, we must realize that the energy output of the heart is greatly dependent upon the configuration of the vascular sys- tem, especially of the arteries. Significant criteria are (a) the relation be- tween the cross-sectional area of a vessel and the sum of the areas of its branches; (b) the angle between the branching vessel and the axis of its trunk; and (c) the distance to be covered between the heart and the capil- lary bed of an organ. Given the above parameters, it was possible to set up an idealized model for an optimal vascular energy system for purposes ofcomparison with the performance of conditions prevailing in a living circulatory system. The formulation of such an ideal model may be regarded as an example of theoretical biology. The second procedure, testing the theory in vivo, rep- resents, on the other hand, unequivocal experimental biology. I presented the above work for my Habilitationsschrift, which was accepted by the medical faculty in 1913. My presentation was not as convincing as the evi- dence found in osteology in regard to the adaptive structuralization of spongeous bone, partly because the former requires mathematical treat- ment before it becomes obvious and the latter presents itselfdirectly to the eyes of a scientifically naïve observer. As I glance back after fifty years of experience, I would now tackle the problem somewhat differently; never- theless, my basic attitude of laying a theoretical foundation before setting up an experimental procedure would remain unaltered. In my position as Privatdozent I was authorized to give laboratory courses, which I organized according to my own inclinations—i.e., bio- physicaily, but not without clinical perspectives. This ideal situation was, unfortunately, terminated by the outbreak of World War I. In the general mobilization of the Swiss army which followed, I, along with everyone else fit for military service, was called to active duty. My work at the physiology institute was completely interrupted. As the war went on, and the increasing superiority of the Allies became apparent, it was sometimes possible to return to work for a few months. I even contemplated con- tinuing my scientific training abroad. Above all, I wanted contact with general physiology. I found it in the laboratory of Max Verworn, Pflüger's successor in

20 409 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

Bonn. Verworn had extensive experience with animal physiology, ac- quired as a pupil of Haeckel. He was especially familiar with so-called ele- mentary organisms and the basic requirements for life. A widely read book by Verworn entitled Allgemeine Physiologie (1922) contains more informa- tion concerning his work. Along with the formal lectures and laboratory work, the almost daily private contact at tea time with Verworn, when we Bonn.discussedVerwornteachinghad extensivemethodsexperiencein general,with animalthe workphysiology,of ourac- colleagues, and quiredevenastopicsa pupilinof Haeckel.art, literature,He was especiallyand philosophy,familiar withdidso-calledmuch ele-to broaden me and mentaryprepareorganismsme forandthethefuture.basic requirementsI will be foreverfor life. Athankfulwidely readforbookthe kindness shown bytoVerwornme byentitledthis man.AllgemeineHe neverPhysiologielet (1922)me becontainsawaremoreof theinforma-vast distance which tion concerning his work. Along with the formal lectures and laboratory work,separatedthe almosthimdaily—aprivatefamous,contactexperienced,at tea time withandVerworn,wise whenmanwe—from myself, a discussedyoungteachingand inquisitivemethods inscientist.general, theI wasworkallowedof our completecolleagues, andfreedom in experi- evenmentaltopics workin art, literature,and wasandable,philosophy,therefore,did muchto concentrateto broaden mefurtherand on problems prepareof hemodynamics.me for the future. I will be forever thankful for the kindness shown to me by this man. He never let me be aware of the vast distance which separatedAs ahimready—a supplyfamous, ofexperienced,dogs wasandavailablewise manfor—experimentalfrom myself, a purposes, I was youngableandto inquisitiveinvestigatescientist.in livingI wasanimalsallowed completethe regulationfreedom inofexperi-regional blood flow, mentali.e., itsworkadaptationand was able,to thetherefore,changingto concentrateneeds offurtherspecificon organs.problemsAt that time the ofconcepthemodynamics.of the "peripheral heart" was a topic of much discussion, espe- As a ready supply ofdogs was available for experimental purposes, I was ableciallyto investigateamong inclinicians living animalswhotheunderstoodregulation oftherebyregional bloodthe totalflow, muscular com- i.e.,ponentits adaptationof thetoarterialthe changingsystem.needsTheofspecificpulseorgans.was ascribedAt that timepartiallythe to an active conceptforce,ofi.e.,the the"peripheralcentrifugalheart"wavewas aoftopicarterialof muchcontractiondiscussion, releasedespe- by the con- ciallytractionamongofcliniciansthe heartwhoandunderstoodresultingtherebyin thetheforwardtotal muscularpropulsioncom- of the blood. ponent of the arterial system. The pulse was ascribed partially to an active force,Thei.e., thenonbranchingcentrifugal wavesectorof arterialof thecontractioncommonreleasedcarotidby thearterycon- seemed to me tractionto beofwellthe heartsuitedandforresultingexperimentallyin the forwardtestingpropulsiontheofabovethe blood.concept. The tech- niqueThe nonbranchingused was sectoras follows.of the commonThroughcarotida smallarterylongitudinalseemed to me incision in the toproximalbe well suitedpartforofexperimentallythe commontestingcarotidthe abovearteryconcept.a glassThestoppertech- was so inserted niqueas tousedstopwastheas follows.flow ofThroughblood. Toa smallpreventlongitudinalany backflowincision inintothe the area under proximal part ofthe common carotid artery a glass stopper was so inserted as investigation,to stop the flow oftheblood.systemTo preventwas alsoany closedbackflowofFcraniallyinto the area underwith a stopper pro- investigation,vided withtheasystemsystemwasforalsoopticallyclosed ofFcraniallyregisteringwithpressurea stopper changespro- in the thus videdisolatedwith aarterialsystem forsection.optically Theregisteringsystemicpressurebloodchangespressurein the thuswas occasionally isolatedmonitoredarterial insection.the femoralThe systemicartery,bloodparticularlypressure wasin occasionallyconnection with artificial monitored in the femoral artery, particularly in connection with artificial pressurepressureincreasesincreasesin the carotidin the artery.carotidAllartery.attemptsAllto alterattemptsthe bloodto alterpres- the blood pres- sure,sure,eithereitherin theinisolatedthe isolatedsection ofsectionthe carotidoftheor incarotidthe peripheralor in arterialthe peripheral arterial system,system,endedendedin failure.in failure.At the endAtofthemyendyear'sofleavemy ofyear'sabsenceleaveI couldof absence I could onlyonlyreportreportnegativenegativeresults. results. Soon thereafter I learned by reading the papers of Hering, Jr., and Hey- mansSoonthat I thereafterhad just missedI learnedan opportunityby readingto makethe papersan importantof Hering,dis- Jr., and Hey- mans that I had just missed an opportunity to make an important dis-

W. R. Hess · From Medical Practice to Theoretical Medicine 410 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 410 Perspectives in Biology and Medicine21· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

co very. My own procedure involved the common carotid up to the point where it bifurcates, thus excluding the carotid sinus. Had I included the sinus, I could not have failed to observe a massive fall in pressure due to the now well-known reflex control of blood pressure. This oversight really irked me, but I consoled myselfwith the thought that similar failures had certainly happened to many other investigators. In any case, I didn' t allow myself to be detracted from the study of hemodynamics. Upon returning to Zürich, it was not possible for me to continue the experimental work aimed at clarifying the adaptation of blood supply to the changing requirements of individual organs. I was called upon for further military service in addition to my teaching schedule, which took up all my time. Further research, then, had to wait until the end ofWorld War I. An Unexpected Turn ofEvents As mentioned previously, my chief, Professor Justus Gaule, head of the department of physiology in Zurich, had suffered from poor health for many years, which hindered him from being very active in teaching and prevented him completely from doing research. Conditions finally became so critical that Gaule announced his resignation just before the winter se- mester of 1916. 1 was asked to take charge of the lectures and laboratories temporarily until the appointment of the permanent holder of the chair. I would have personally preferred to spend several more years as a Pri- vatdozent, with adequate time for research, rather than to be acting chair- man with all the administrative and teaching duties. Nevertheless, with healthy optimism and steadfast determination I began the work ahead of me. In relying on the experience gained during my years with the handling of mechanical and optical equipment, I planned to use such extensively in the laboratory experiments. I was also counting on drawing from my in- ternship experience, and particularly on the operative skills I had acquired as an assistant surgeon and as an ophthalmologist. Finally, conceptual refer- ences between normal and pathological physiology as well as the insights gained into general physiology under Verworn turned out to be most helpful in the new situation. Zürich Faculty It should be noted that in the German-speaking region of Switzerland appointments to professorial chairs were at that time rather complicated

22 411 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

and entailed far greater attention to the so-called "school" of the prospec- tive candidate than, for example, in the United States, where much more liberal principles for selection operate. Additional difficulties arose in gen- eral from the fact that the relatively young University of Zürich lacked a corps ofgraduates trained in science. Therefore, German candidates usually had an advantage from the beginning, and many positions were occupied andbyentailedGermans,far greatersomeattentionof whomto thewereso-calledreally"school"outstanding.of the prospec-This method for tivegatheringcandidateacademicthan, for example,personnelin thewasUnitedin vogueStates, whereduringmuchthemoretime of the expan- liberalsive tendenciesprinciples forofselectionthe Wilhelmianoperate. Additionalepoch.difficultiesIn anyaroseevent,in gen-the German pro- eral from the fact that the relatively young University of Zürich lacked a corpsfessorsofgraduatestook ittrainedfor grantedin science.thatTherefore,a SwissGermanpersoncandidatescouldusuallynot be considered hadseriouslyan advantagefor anfromacademicthe beginning,positionand manyin apositionstheoreticalwere occupiedmedical discipline; a bySwissGermans,scientistsomewasof whommorewerelikelyreallyto beoutstanding.intrustedThiswithmethoda practicalfor assignment. gatheringUnderacademicthese personnelcircumstances,was in voguethen,duringa proposalthe timewasof thepreparedexpan- at the end of sive tendencies of the Wilhelmian epoch. In any event, the German pro- fessorsthe wintertook itsemesterfor grantedwhichthat aforesawSwiss personcallingcoulda notGermanbe consideredprofessor to the chair seriouslyin physiology.for an academicWith positiondue considerationin a theoreticalofmedical"his senioritydiscipline;anda much larger Swisstraining,"scientisthewashadmoreproducedlikely to befarintrustedmore withscientifica practicalpapersassignment.than I and had unde- niablyUnder goodthese circumstances,qualities, especiallythen, a proposalin thewasfieldpreparedof physiologicalat the end of chemistry. In thespitewinterof semesterthis, hewhichhad notforesawyetcallingbeen acalledGermantoprofessora leadingto thepositionchair in Germany inandphysiology.was stillWithin linedue considerationfor one. of "his seniority and much larger training," he had produced far more scientific papers than I and had unde- niablySince,good however,qualities, especiallythe administrationin the field of physiologicaland managementchemistry. Inof the institute spiteduringof this,thehe winterhad not yetsemesterbeen calledhadto gonea leadingalongpositionsmoothlyin Germanyunder my acting andchairmanship,was still in lineandfor one.because the lectures and laboratories had been well at- Since, however, the administration and management of the institute duringtended,thethewinteruniversitysemesterauthoritieshad gone alongdecidedsmoothlyto re-examineunder my actingthe entire situation chairmanship,more thoroughly.and becauseAntheinvitedlecturesopinionand laboratorieswrittenhadbybeenVerwornwell at-which had been tended,withheldthe universityby the facultyauthoritiestilldecidedthen towasre-examinealso broughtthe entiretosituationmy support. Finally, morethe summerthoroughly.semesterAn invitedturnedopinionoutwrittento beby anotherVerworn whichsuccess,hadtobeenwhich two very withheld by the faculty till then was also brought to my support. Finally, thecapablesummerassistantssemester turnedgenerouslyout to becontributed,another success,E.toRothlinwhich twoandveryA. Fleisch—sub- capablesequentlyassistantsprofessors,generouslyrespectively,contributed, E.ofRothlinpharmacologyand A. Fleischat—Baslesub- and of physi- sequentlyology atprofessors,Lausanne.respectively,The decisionof pharmacologyofthe nominatingat Basle andcommitteeof physi- came out in ologymy favor,at Lausanne.andThein decisionthe fallofofthe1917nominatingthe chaircommitteefor physiologycame out in was officially myintrustedfavor, andto inme.the fall of 1917 the chair for physiology was officially intrusted to me.

DevelopmentDevelopmentof a Programof a Programfor Instructionfor Instructionand Researchand Research NowNowthatthatI wasI wasresponsibleresponsiblefor the developmentfor the developmentof physiology,ofitphysiology,was it was mandatory for me to become familiar with the management and setup in othermandatoryuniversities.forI hadmetheto greatbecomefortunefamiliarofbeingwithable tothegetmanagementto know and and setup in other universities. I had the great fortune ofbeing able to get to know and

W. R. Hess · From Medical Practice to Theoretical Medicine 412 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 412 Perspectives in Biology and Medicine23· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

to observe the English physiologists of the classical period, as they were Betriebstill working in their laboratories. Starling gave me complete freedom to gain an insight into his programing and methodology for lecture sessions Untertiteland lab work. Through Langley I came to understand how the institute in Auch inCambridge betriebswirtschaftlicherwas run. I sawHinsichtSherrington hat at work only briefly, as he was das Klinischevery busy Neurozentrumand implied dasthat Berichtsjahrhe did not want to be bothered. With the help erfolgreichand kindnessabgeschlossen.of one Mitof his insgesamtassistants, however, I was allowed to find out 21’951 stationären und ambulanten Patien- tenbehandlungenabout everything wurde ofeininterest Zuwachsto me.der Many years later I met this assistant— Fallzahlenwho gegenübernow holds dema leading Vorjahrposition um 4.3%in the United States—at an international erzielt, congresswas sich in(I einerbelieve Zunahmein Montreal). des Ertra- Naturally the topic of conversation ges umturned 2.4% widerspiegelt.to Professor Sherrington.Ebenfalls wurBecause- of my own later experience, I de eine deutliche Zunahme der ambulanten Tätigkeithad in longden Ambulatoriensince come to undunderstand Sprech- Sherrington's reserve and aloofness. stunden derIn addition Klinikento desthis Neurozentrumsvisit to England and later travels, I also participated in verzeichnet,various diecongresses sich in einerto which SteigerungI owe a great deal for the broadening of my der erwirtschaftetenknowledge. I canTaxpunktestill vividly um remember8.7% the demonstration of dogs suffer- gegenübering fromdem Vorjahrrickets presentedniederschlägt.by E. DieMellanby, later Lord Mellanby, with the markante Zunahme der stationären Tätig- keit umhelp 18.5%of hattehis wife, einelater AufwandszunahLady Mellanby,- at the International Physiological me vonCongress 5.6% gegenüberin Paris indem1920. VorjahrAt that zurtime Mellanby demonstrated the effects Folge. ofmalnutrition—the concept ofvitamins was still controversial. One could hear the differences of opinion being argued more critically in the cor- ridors than in the demonstration hall in the presence of the speakers them- selves. A great scientific event also occurred at the congress in Edinburgh in 1923, where, under the leadership of J. J. R. MacLeod, a series oflectures included those by Banting and Best in which the chain ofevents which led to the discovery of insulin was unfolded. I returned from still other sci- entific meetings enriched in experiences and knowledge which could be incorporated into my lectures by way of illustrating them with experi- mental material. The meetings of the Deutsche Gesellschaftfür Naturforscher und Ärzte were particularly stimulating and important for me. At these sessions I attended primarily the general lectures. Otherwise, I sat in on those given by the internists. As a teacher, I was much concerned with the advances in internal medicine, as my students, upon entering the clinical semesters, should be able to correlate clinical material with their knowl- edge of physiology. A special feature ofthese meetings was the chance to become personally acquainted with other scientists in the more relaxed atmosphere outside the lecture halls. Apart from shop talk, these personal contacts provided im-

413 24 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

portant insights for evaluating, to some extent, the publications of the Betriebvarious institutions. Such first-hand impressions saved me a lot of time in selecting literature for study. Research Projects For the time being hemodynamics remained the chief subject of re- portantsearchinsightsin myforinstitute.evaluating,It includedto some extent,researchthe workpublicationsconcerningof the the autonomic various institutions. Such first-hand impressions saved me a lot of time in selectingpropertiesliteratureandforthestudy.reactivity of isolated arteries. It was clear to me that information obtained on an isolated arterial section could not by itself be decisiveResearch Projectsfor the intact living arterial apparatus. On the other hand, I was convincedFor the timethatbeingthehemodynamicsinherent functionalremained propertiesthe chief subjectof theofarterialre- tube, even searchthoughin mynormallyinstitute. It includedsubjectresearchto nervouswork concerninginfluences,the autonomicwould be of great sig- propertiesnificanceandinthethemselves.reactivity ofInisolatedline witharteries.theseIt wasthoughts,clear to mearterialthat rings were re- information obtained on an isolated arterial section could not by itself be decisivemovedforfromthe intactvariouslivingpartsarterialofapparatus.the arterialOn thesystemother andhand,testedI was for their spon- convincedtaneousthatbehaviorthe inherentwhilefunctionalsubmersedpropertiesin physiologicalof the arterial tube,saltevensolution. In addi- thoughtion, normallythe ringssubjectweretostimulatednervous influences,mechanicallywould andbe ofelectricallygreat sig- with varying nificanceparametersin themselves.as wellIn lineas withchemicallythese thoughts,witharterialbiologicallyrings wereactivere- substances— movedamongfromothers,various adrenalinparts of the arterialand acetylcholine.system and testedThesefor theirexperimentsspon- were ex- taneous behavior while submersed in physiological salt solution. In addi- tion,tendedthe ringsoverwerea periodstimulatedofmechanicallyyears and yieldedand electricallythe materialwith varyingwhich, with inclu- parameterssion of asthewellreflexas chemicallymechanisms,with biologicallyprovided activea generalsubstancesinsight— subsequently amongelaboratedothers, adrenalinin 1930 andin theacetylcholine.monographTheseRegulationexperimentsofwerethe Circulatoryex- System. tendedFromoverthea closeperiodrelationshipofyears and yieldedand synergicthe materialfunctioningwhich, withofinclu-the circulatory and sion of the reflex mechanisms, provided a general insight subsequently elaboratedrespiratoryin 1930apparatus,in the monographit followedRegulationautomaticallyof the CirculatorythatSystem.The Regulation of FromRespirationthe close relationshipbe handledandinsynergican analogousfunctioningmannerofthe circulatory(193 1). and respiratoryFromapparatus,discussionsit followedwith a fewautomaticallycolleagues,that ITheoccasionallyRegulation ofheard that refer- Respirationence to bethehandledconceptin anofanalogousa meaningfulmannerorganization(193 1). of physiological systems From discussions with a few colleagues, I occasionally heard that refer- encewasto herethe conceptand thereof a meaningfulregardedorganizationas an expressionof physiologicalof teleologica!systems tendencies. In wasreplyinghere and totheresuchregardedcomments,as an expressionI haveofalwaysteleologica!maintainedtendencies.thatIn living matter replyingcan onlyto suchexistcomments,in the formI haveofalwaysorganizationmaintainedorientedthat livingtowardmattersuccess; indeed, canthatonlytheexistinvestigationin the form of organizationof such organizationoriented towardposessuccess;theindeed,central problem of that the investigation of such organization poses the central problem of biology.biology.TheTheknowledgeknowledgeand insightandgainedinsightby experimentalgained by experimentalmethods and methods and thethecollectedcollectedpool poolof factsofallowingfacts allowinga synthesisabetweensynthesisorganizationbetweenandorganization and performanceperformancetrulytrulyconstituteconstitutethe grandtheaimgrandof an integralaim ofphysiology,an integralwhichphysiology, which cancanneverneverbe fulfilledbe fulfilledwith thewithmerethecollectingmere ofcollectingfacts. The ofdeleteriousfacts. The deleterious effects of disturbances, as in sickness, confirm the above viewpoint. The developmenteffects ofofdisturbances,this theme was asworkedin sickness,out and laterconfirmformallytheexpressedaboveinviewpoint. The development ofthis theme was worked out and later formally expressed in

W. R. Hess · From Medical Practice to Theoretical Medicine 414 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 414 Perspectives in Biology and Medicine25· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

the monograph (1948) Diefunktionelle Organisation des vegetativen Nerven- systems (Functional Organization ofthe Vegetative Nervous System). The shift to a seemingly new topic—namely, examination ofactive reg- ulatory dispositives at the higher level ofthe diencephalon—will readily be understood as a logical consequence of a long-range project. From an earlier programmatic work entitled "Correlations between Psychic and Vegetative Functions" (1924), in conjunction with and from knowledge of certain clinical experiences and from exploratory experimental investiga- tions ofKarplus and Kreidl, certain conclusions could be drawn. It became apparent that before further progress could be expected, a new experi- mental technique which could adequately fulfil the more differentiated demands of the modern investigation would first have to be developed. For it was my goal to test the behavior ofexperimental animals (cats) sub- jected to electrical stimulation and diathermic lesions without impairing the freedom of movement of the animal by immobilization or narcosis. For this purpose, the very finest electrodes were needed which could be inserted into the hypothalamus without flexing and bending. In addition, to use the induction coil as a stimulating device was out ofthe question, for the relatively slowly arising autonomic effects in the face of the rapidly occurring potential deflections produced by induction could be all too easily overshadowed by reactions from somatic sensorimotor systems and could, under certain conditions, not even have time to be formed. I was more hopeful for a form of stimulation with pulsating direct current, pro- vided that the cathodic electrode points could be depolarized between the single impulses, thus maintaining a consistent stimulus intensity also during prolonged series of stimulation. The decisive advantage therein proved to lie in the possibility of adapting duration and shape of the stimulating pulses to the time properties of the nervous elements under examination. Apart from these precautionary measures concerning stimulation tech- nique, the problem ofanatomical identification of the stimulated structure had to be solved so as to allow the closest possible correlation between symptom and reacting substrate. Since required accuracy can never be achieved on the basis ofelectrode placements alone, microscopic control of serial histological sections of each experimental brain is necessary. Three atlases composed of photomicrographic reproduction of serial sections cut perpendicularly to each other gave invaluable assistance in the topographi- cal analysis of stimulation sites. In addition, these atlases served to register

26 415 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

in each individual case the histological finding as well as to prepare cumula- tive map files of all points stimulated during the many years of investiga- tive work. Cross-references among these three standard atlases allowed localization ofeach site ofstimulation in all three planes, thereby providing three-dimensional registration, which in many cases was indispensable to demonstrate the relation between topographical and functional organiza- intion.each individualCinematographycase the histologicalwas usedfindingextensivelyas well as tofromprepare1927cumula-on as a further aid tiveinmaprecordingfiles of allobservations.points stimulatedIt enabledduring theusmanyto makeyears ofdetailedinvestiga-comparisons be- tivetweenwork.symptomsCross-referencesof experimentsamong these threeseparatedstandardfromatlasesoneallowedanother by years. localization ofeach site ofstimulation in all three planes, thereby providing three-dimensionalThis would haveregistration, been whichimpossiblein manybycasesreliancewas indispensableon verbaltonotes alone. Fi- demonstratenally, a filingthe relationsystembetweenwastopographicaldeveloped andwhichfunctionalcontainedorganiza-references to the tion.symptomsCinematographyproducedwas usedby stimulation;extensively fromthis1927enabledon as a furtherone inaidthe train of data inintegrationrecording observations.to obtainIt enabledfull particularsus to make ofdetaileda desiredcomparisonscase be-together with the tweenverbalsymptomsprotocol,of experimentsphotomicrographicseparated fromlocalization,one another byandyears.cinematographic Thisdocuments.would have been impossible by reliance on verbal notes alone. Fi- nally, a filing system was developed which contained references to the symptomsThe technicalproduced byaspectsstimulation;ofthisthisentireenabledprogramone in thetotraininvestigateof data the functional integrationorganizationto obtainof thefull particularsdiencephalonof a desiredwere reportedcase togetherin withdetailthein a monograph verbal protocol, photomicrographic localization, and cinematographic documents.(1932) entitled Die Methode der lokalisierten Reizung und Ausschaltung sub- kortikalerThe technical Hinabschnitteaspects ofthis entire("TheprogramMethodologyto investigateoftheLocalizedfunctionalStimulation and organizationDestructionof theofdiencephalonSubcorticalwereBrainreportedAreas").in detailThisin aasmonographwell as the second part, (1932)publishedentitled1938,Die Methodeconcerningder lokalisiertenbloodReizungpressureundandAusschaltungrespiratorysub- effects elicited kortikalerby localized Hinabschnittediencephalic("The Methodologystimulationof Localizeddid not Stimulationbecome tooandwell known be- Destruction of Subcortical Brain Areas"). This as well as the second part, publishedcause of1938,WorldconcerningWar bloodII. Thepressuregroupandofrespiratorycompetenteffectsworkerselicitedin this field was bysmalllocalizedat thatdiencephalictime, andstimulationall thedidtypenotofbecomethis editiontoo well wasknowndestroyedbe- in Leipzig causeduringofWorldWorldWarWarII. TheII.group ofcompetent workers in this field was smallAtat thatthetime,veryandbeginningall the type ofofthistheeditionexploratorywas destroyedexperimentsin Leipzig (1925) I had duringreckonedWorld withWar II.from two to three years for gathering data sufficient to At the very beginning of the exploratory experiments (1925) I had reckonedobtain witha certainfrom twoinsight.to threeButyearsthe formeregatheringmanualdataworksufficientinvolvedto in the tech- obtainniquea certaindescribedinsight.aboveBut theusedmere upmanualthisworkmuchinvolvedand ineventhe tech-more of my time. niqueAlso,describeda heavyaboveteachingused upschedulethis muchdidandnotevenmakemorethingsof mygotime.any faster. Never- Also,theless,a heavyinteaching1927, schedulethe firstdidresultsnot makewerethingsmadego anyknownfaster. Never-at a meeting of the theless, in 1927, the first results were made known at a meeting of the DeutscheDeutschePhysiologischePhysiologischeGesellschaftGesellschaftin Frankfurt.in OfFrankfurt.course, followingOf course, following thetheconclusionconclusionof theoffirsttheseriesfirstofseriesabout fiftyofaboutcases, nofiftyinsightcases,intonogeneralinsight into general lawslawsgoverninggoverningthe functionalthe functionalorganizationorganizationwas obtained;wasthe obtained;symptoms the symptoms wereweretootooinconsistent.inconsistent.Therefore,Therefore,a second experimentala second experimentalseries was carriedseries was carried out—which did not, however, do much to clarify the situation. outOnly—littlewhichby littledidandnot,everhowever,so slowly diddothemuchveil lifttoa bitclarifyhere andthetheresituation. Only little by little and ever so slowly did the veil lift a bit here and there

W. R. Hess · From Medical Practice to Theoretical Medicine 416 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 416 Perspectives in Biology and Medicine27· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

so that twenty-five years had passed before I could finally think ofputting together all the many single research publications that had appeared over the years which had been concerned with individual symptoms. The vast number of experiments turned out to be decisive; for generalization con- cerning symptoms, syndromes, and localizations could be supported only by such a large body ofdata. An integrative review ofall these efforts was published in two monographs (1947, 1948). They were well received, and in 1949, "for your discovery of the functional organization of the dien- cephalon and its role in the coordination of the functions of the inner or- gans," the Nobel Prize was bestowed upon me. The reward was shared with Egaz Moniz, who had developed the prefrontal as a thera- peutic method in the management of certain psychoses. The long way which culminated in this high distinction has been pre- sented here in a rather detailed manner because during the war Switzerland was totally closed offand precious little information concerning my inves- tigative methods and results had reached readers abroad. Even with the war's end the fact that all my publications were written in German, the accepted written language ofcentral and eastern Switzerland, proved to be for many years an almost insurmountable barrier in re-establishing a close scientific contact with the Anglo-Saxon countries. Apart from behavioral and autonomic symptoms, interesting somatic motor effects came to light during the course ofthe experiments. Through these the relationships of certain formations of the thalamus as well as of the mesencephalon with the so-called extrapyramidal motor system were established. Last but not least, our attention was drawn to psychomotor phenomena on which I reported in a most recently published treatise (1962) entitled Psychologie in biologischer Sicht ("Psychology from the Standpoint of a Biologist"). Incidents, Personalities, and Anecdotes During my active years as professor of physiology on the medical fac- ulty at Zürich, the above described work was often interspersed with note- worthy incidents and problems which have remained vividly in my mem- ory. One of these could be cited as an example of "fortune in misfor- tune." When I first came to the physiological institute of our university, there was a noticeable shortage of both equipment and space, without which a successful work program was rather difficult—or, more plainly,

28 417 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

impossible. Significant improvement was out of the question for the time being. The basement and first floor ofthe four-storied building which had been constructed during the time ofthe famous physiologist Hermann, the predecessor of Gaule, were occupied by the physics department, which was also cramped for space. As a result of these unsatisfactory conditions, the department oftheoretical physics and a succession ofits famous staffmem- impossible.bers, suchSignificantas Einstein,improvementvon Laue,was outandof theSchrödinger,question for thehadtimeto work in the being.smallestThe basementspace ofandall.firstA floor"higherofthepower"four-storiedintervenedbuilding whichin thishadprecarious situa- beention.constructedOn theduringeveningthe timebeforeoftheNewfamousYear'sphysiologistEve inHermann,1923, justthe before retiring, predecessor of Gaule, were occupied by the physics department, which was alsoI receivedcramped fora telephonespace. As a callresultinformingof these unsatisfactoryme that ourconditions,institutethewas going up in departmentflames. AsoftheoreticalI arrivedphysicson theandspot,a successionthe entireofits topfamousfloorstaffwasmem-already burning. bers,ThesuchfireasdepartmentEinstein, von wentLaue, andto workSchrödinger,immediately.had to workThein captainthe of the fire smallestsquadspacewasofassuredall. A "higherthat itpower"wouldintervenednot be necessaryin this precariousto evacuatesitua- material from tion.theOnlowerthe eveningfloors.beforeAs thoughNew Year'sto spiteEve inthis1923,comfortingjust before retiring,prognosis, however, I received a telephone call informing me that our institute was going up in flames.the fireAs Isuddenlyarrived on brokethe spot,throughthe entirethetop floorceilingwasandalreadythreatenedburning. to destroy the Themainfire portionsdepartmentofwentourtoinstitute.work immediately.It was highThe captaintime, then,of the tofireget books—and squadanythingwas assuredelsethatof itvaluewould—notoutbeofnecessarythe buildingto evacuateas materialquicklyfromas possible and to thetakelowerthemfloors.throughAs thoughtheto deepspite thissnowcomfortingto neighboringprognosis, however,institutes. During this the fire suddenly broke through the ceiling and threatened to destroy the main"operationportions ofevacuation"our institute. Itourwasbasichigh time,sensethen,of democracy,to get books—andvalid for all social anythinglevels, elsewasofdramaticallyvalue—out ofillustrated.the building For,as quicklyin additionas possibletoandthetostaffmembers of takethethemphysicsthroughdepartmentthe deep snowandto otherneighboringinstitutes,institutes.membersDuring thisof the executive "operationbranch ofevacuation"our stateourgovernmentbasic sense ofweredemocracy,also onvalidhandfordoingall socialtheir very best to levels, was dramatically illustrated. For, in addition to the staffmembers of thehelpphysicsout.departmentSince onlyandmyotherchiefinstitutes,assistantmembersand myselfof thewereexecutiveinformed about the branchvalueofofourthestateinventory,governmentwewerebecamealso on handresponsibledoing theirforverygivingbest to directions as to helptheout.prioritySince onlyformyobjectschiefassistantto be saved.and myselfThuswereitinformedarose thataboutthetheshoe came to be valueon theof theotherinventory,foot,weinbecamethat ourresponsibleemployers,for givingthe directionspoliticalasauthorities,to had to thetakepriorityordersfor fromobjectstheirto be saved.employees,Thus it arosei.e., fromthat thetheshoeprofessorcame to beand his assistants. on the other foot, in that our employers, the political authorities, had to takeOnordersthe fromfollowingtheir employees,day, afteri.e., thefromfirethe professorwas finallyand hisextinguished,assistants. we had to Oncollectthe followingour possessionsday, after thefromfire thewas neighboringfinally extinguished,buildings.we hadOfto course, every- collectthingourstoredpossessionsin thefromtopthefloorneighboringwas lost.buildings.In addition,Of course,theevery-building itself had thingbeenstoredseriouslyin the topdamaged.floor wasThislost. Instateaddition,of affairsthe buildinggave itselfthe cuehad to the second, been seriously damaged. This state of affairs gave the cue to the second, happy,happy,partpartof myofstory;my story;for, duringfor,theduringrepair work,the repairan additionalwork, flooran additional floor waswasaddedaddedand andthus ourthusneedourforneedspaceforalleviated.space Wealleviated.moved intoWethemoved into the newlynewlyrepairedrepairedand expandedand expandedinstitute withinstitutethe sincerewithhopethe ofsincerebeing ablehope of being able totodevotedevoteourselvesourselvesto ourtoworkourwithoutworkdisturbance.without disturbance. This expectation, however, all too soon proved to be illusory. In our directThisdemocracyexpectation,a constitutionalhowever,law existsall toowherebysoonactionprovedcan beto initi-be illusory. In our direct democracy a constitutional law exists whereby action can be initi-

W. R. Hess · From Medical Practice to Theoretical Medicine 418 Perspectives in Biology and Medicine · Summer 1963- W. R. Hess · From Medical Practice to Theoretical Medicine 418 Perspectives in Biology and Medicine29· Summer 1963- From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

ated by the people provided that a sufficient number of signatures are sub- mitted. A movement of the community touching at the very heart of medicine and biochemistry arose banning all animal experiments in the canton of Zürich. Members of the physiology institute in Zürich were called upon by the medical faculty to combat this attack, which was aimed at destroying a technique indispensible to scientific progress. We had to prepare ourselves for the battle, which, unfortunately, would take up a lot of time. In place of scientific work, it now became necessary to write popular newspaper articles and to speak at public gatherings in order to present the true interest of the people. The problem of financing the cam- paign, in the face ofthe opposite community drive on behalfofthe protec- tion ofanimals, was not a simple matter. When the polls opened, our side was understandably on pins and needles. Members of our campaign head- quarters and many interested colleagues had assembled in a hotel room to keep informed on the incoming results from the city and the rural areas. Happily, our efforts had not been in vain. The popular vote resulted in a two-thirds majority in favor of freedom of research with reasonable re- strictions. It is impossible to imagine what the fate of our faculty might have been had animal experimentation been declared a criminal offense. Another equally laborious but more constructive undertaking into which I was drawn by chance turned out to be more than I had bargained for. The Naturforschende Gesellschaft (Swiss Society for Scientific Re- search) had taken upon itself, due to the promptings ofone ofits members, a competent meteorologist, to set up a committee for the purpose of con- structing a research station at the highest point in Switzerland accessible by railroad—namely, on the Jungfraujoch, over 11,000 feet above sea level. The scientific promoter of the plan died before he could carry it out. Therefore, a new presidium had to be selected for the commission, and it came about that the job fell to me. I had suspected that acceptance would entail no small sacrifice, but in various respects this was a great underesti- mate. Originally a Swiss undertaking was envisioned. I myself tried to have the project internationalized. One ofthe main reasons was the thought that thereby new contacts between representatives of countries which had been enemies during World War I could arise, which could help promote un- derstanding among peoples ofdifferent nations. A second point in my pro-

30 419 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

gram was to include all natural sciences, and particularly biology. One source of resistance was laid down in the initial formulation of objectives of the research station, according to which it was to be used primarily for meteorological studies and for work on geophysical problems. Fortunately the branch ofthe federal government charged with the main authority was farsighted enough to interpret the formulation ofthe objectives ofthe plan gramso thatwas allto includedisciplinesall naturalof scientificsciences, andresearchparticularlyhavingbiology.an establishedOne interest in sourceworkofinresistancethe highwas Alpinelaid downregionin the initialshouldformulationreceive equalof objectivesconsideration. of theWithresearchagreementsstation, accordingon internationalizationto which it was to beandusednonrestrictionprimarily for as to field of meteorologicalresearch secured,studies andit wasfor worktimeontogeophysicallook aroundproblems.forFortunatelypartners who were pre- the branch ofthe federal government charged with the main authority was farsightedpared toenoughmaketoainterpretfinancialthecontributionformulation offorthe objectivesthe project.oftheWeplanhad the satisfac- sotionthat allofdisciplinesreceivingofpositivescientific researchresponseshavingfroman establishedmany countries,interest in among which workmayinbethementionedhigh Alpine regionthe thenshouldKaiserreceiveWilhelmequal consideration.Society in Berlin (presently Max-Planck-Gesellschaft),With agreements on internationalizationUniversityand nonrestrictionof Paris, Royalas to fieldSocietyof of London, research secured, it was time to look around for partners who were pre- paredAcademyto make aoffinancialSciencescontributionin Vienna,for theNationalproject. WeFoundationhad the satisfac-for Scientific Re- tionsearchof receivingin Brussels,positiveand,responsesfinally,fromthemanyRockefellercountries, amongFoundationwhich in New York, maywhich,be mentionedhowever,the thendeclinedKaisertoWilhelmparticipateSocietyonin Berlinthe international(presently committee. Max-Planck-Gesellschaft),Naturally, it was ofprimeUniversityimportanceof Paris, Royalthat theSocietyJungfrauof London,Railway Company Academy of Sciences in Vienna, National Foundation for Scientific Re- searchreaffirmedin Brussels,its and,activefinally,andthefinancialRockefellerco-operation,Foundation in NewwhichYork,it had promised which,beforehowever,the grantdeclinedfor constructionto participate onofthetheinternationalrailway hadcommittee.been made, as well as Naturally,its readinessit wastoofprimehelp outimportancein anythatothertheJungfrauproblemsRailwayconcerningCompanyconstruction and reaffirmedoperation.its active and financial co-operation, which it had promised before the grant for construction of the railway had been made, as well as its readinessNo effortto helpcouldout inbeanysparedother problemsin overcomingconcerningtheconstructionconditionsand which were to operation.be expected in working on the rugged, fissured precipices. One had to reckonNo effortwithcouldunexpectedbe spared in overcomingstone slidesthe conditionsand waterwhichbreakthroughswere to as well as beunseasonalexpected in workingheavy snowfall.on the rugged,Anotherfissuredproblemprecipices. wasOne hadthe toextreme diurnal reckonvariationwith unexpectedin temperature.stone slidesTheandstonewatermaterialbreakthroughsto beas usedwell asfor the building unseasonal heavy snowfall. Another problem was the extreme diurnal variationwas carefullyin temperature.selectedThetostonewithstandmaterial tothebeintensiveused for theheatingbuildingby high-altitude wassolarcarefullyradiationselectedduringto withstandthe daythe andintensiveexposureheatingtobyfreezinghigh-altitudetemperatures dur- solaringradiationthe night.duringA completelythe day and exposureunexpectedto freezingeventtemperaturesmade a jokedur- of all our care- ingfulthecalculationsnight. A completely—for unexpectedin an areaeventwhichmadehada jokebeenof allstableour care-and inactive for ful calculations—for in an area which had been stable and inactive for seventyseventyyears,years,an avalanchean avalanchesuddenlysuddenlybroke lose,brokeinterruptinglose, theinterruptingflow of the flow of suppliessuppliesfor valuablefor valuableweeks.weeks. ButButat longat longlast, inlast,the insummerthe summerof 193 1, theofresearch193 1, thestationresearchwas dedi-station was dedi- catedcatedin thein thepresencepresenceof prominentof prominentrepresentativesrepresentativesof the participatingof the participating countriescountriesand andinstitutionsinstitutionsas well as Swisswell governmentas Swiss governmentauthorities of theauthorities of the highesthighestrank.rank.A specialA specialvolumevolumepreparedprepared for the memorable for theoccasionmemorablein- occasion in-

W. R. Hess · From Medical Practice to Theoretical Medicine 420 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 420 Perspectives in Biology and Medicine31· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

eluded contributions from W. Nernst and v. Ficker (Berlin), Ch. Fabry and L. Lapicque (Paris), Sir F. G. Hopkins and G. M. B. Dobson (Oxford), A. Durig and W. Schmidt (Vienna), E. de Wildeman, Gand and J. Jau- motte (Liège), as well as from many others. Every donor country was rep- resented by two delegates. With the reading of the declaration statement "For the Scientific Brotherhood" which had been prepared by the physi- ologist Lapicque, we fraternally joined hands. After being responsible for directing the commission through five years of development and construction in addition to a further five years of service as director ofthe newborn institute, I felt relieved ofa great burden when I finally stepped down. A. v. Murait, from Bern, proved himself more than adequate and an excellent choice for the succession. I should mention another event which brought additional honors and duties to our institute—the Sixteenth International Congress ofPhysiology in Zürich (1938). It should not be forgotten that we stood at that time at the brink of World War II. The annexation of Austria by Hitler was then an accomplished fact. At the last minute, partly out ofpolitical, partly out of racist motives, we were exposed to a considerable change in attitude from foreign colleagues and officials. Also, personal tensions between par- ticipants of the congress could not be ignored. It was not even certain whether the congress would be held at all. Nevertheless, it must be credited to those attending this memorable meeting that proper discipline was maintained and unpleasant incidents were avoided by suppressing any al- lusions of a political nature in the lectures and in the other events of the congress. In connection with the deterioration of the warlike situation, the Swiss physiologists were put into an unusual position. Till then they had been associated partly with the German Physiological Society and partly with its French-speaking counterpart. A new way had to be found in loosening our ties from both of the warring factions. With the support of K. Spiro, the biochemist on the Basle medical faculty, I made a motion for the founding of a Swiss Physiological Society. Inevitably, the next step was the founding in 1943 of a periodical, Helvetica Physiologica et Pharmacologica Acta. It is understandable that at first the circulation of the new periodical was rather limited. Everyone's inter- est was completely directed toward the war. In addition, the passage of

32 421 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

printed matter out of Switzerland into, for instance, the United States was barred. Even after the war, the demand for the new journal was small. Yet we had to be satisfied with what had been achieved. Today we note that the journal is enjoying an increase in circulation. To end this sketch, I have the satisfaction of announcing a gratifying printedundertakingmatter outwhichof Switzerlandis intimatelyinto, forconnectedinstance, the Unitedto myStateslife'swaswork. As early as barred.1940EvenI hadafterassumedthe war,thetheinitiativedemand forforthefoundingnew journala wasstudysmall.group devoted to Yetbrainwe hadresearch.to be satisfiedPeriodically,with what hadphysiologists,been achieved.psychiatrists,Today we notebrain surgeons, thatinternists,the journalpharmacologists,is enjoying an increaseneuro-anatomists,in circulation. psychoanalysts, and psycho- therapistsTo end thiswouldsketch, Icomehave thetogethersatisfactionat theof announcingphysiologicala gratifyinginstitute. At first, we undertakingwanted towhichget tois intimatelyknow oneconnectedanother'sto mypointslife's ofwork.viewAs andearlytoas find a common 1940languageI had assumedfor thethevariousinitiativetermsfor foundingand concepts.a study group devoted to brainThanksresearch.toPeriodically,the financialphysiologists,support psychiatrists,of the Rockefellerbrain surgeons,Foundation and later internists, pharmacologists, neuro-anatomists, psychoanalysts, and psycho- therapiststo thatwouldofthecomeSwisstogetherAcademyat theofphysiologicalMedical Sciences,institute. Atthroughfirst, weits Commission wantedfor Brainto getResearch,to know onethisanother'sstudypointsgroupofviewachievedand to itsfindgoala commonwith ever expanding languageparticipationfor the variousuntil 1950.terms andTodayconcepts.the original plan has culminated in an insti- tuteThanksfortobrainthe financialresearch,supportto ofbetheopenedRockefellerin 1963.Foundation and later to that ofthe Swiss Academy ofMedical Sciences, through its Commission for BrainIf oneResearch,considersthis studymygroupownachievedinterestitsingoalthewithfunctionalever expandingorganization of the participationbrain alonguntilwith1950. Todayits psychicthe originalmanifestationplan has culminated—an ininterestan insti-which was ex- tutepressedfor brainas farresearch,backtoasbe1924openedandinwhich1963. was never lost sight of—one will well understandIf one considersthatmytheowndecisioninterest inofthethefunctionalfacultyorganizationand the responsibleof the authorities brain along with its psychic manifestation—an interest which was ex- pressedto createas far backa chairas 1924forandbrainwhichresearchwas neverhaslostfilledsight ofmy—oneyoungwill wellold heart with the understandvery deepestthat thesatisfaction.decision of theEquallyfaculty gratifyingand the responsibleis the factauthoritiesthat I can regard the tofuturecreate a ofchairtheforinstitutebrain researchwithhasconfidence,filled my youngas oneold heartof mywithearlierthe co-workers, veryK.deepestAkert,satisfaction.presentlyEquallyon thegratifyingmedicalis thefaculty,fact thathasI canbeenregardnamedthe its director. future of the institute with confidence, as one of my earlier co-workers, K. Akert,A lastpresentlypoint onremainsthe medicalto befaculty,mentionedhas been—namedresearchits director.costs money and I am gladA last topointacknowledgeremains to be mentionedthe help —I researchreceived.costsWhilemoney andthe Ismallam communities gladwhichto acknowledgesupport ourthe helpSwissI received.universitiesWhilewerethe smallnotcommunitiesable to provide adequate whichmeans,supportfinancialour Swissassistanceuniversitiesbywerethe notRockefellerable to provideFoundationadequate and by a per- means, financial assistance by the Rockefeller Foundation and by a per- sonalsonalbenefactor,benefactor,BenjaminBenjaminStern, wasStern,of decisivewas ofvalue.decisiveIn the lastvalue.few In the last few yearsyearsthetheSwissSwissNationalNationalFoundationFoundationfor ScientificforResearchScientificfurtheredResearchour furthered our endeavors.endeavors.

In Inanticipationanticipationof a possibleof a possiblequestion fromquestionmy readers,from especiallymy readers,thoseespecially those in America, I must answer whether or not during my active period I found in America, I must answer whether or not during my active period I found

W. R. Hess · From Medical Practice to Theoretical Medicine 422 Perspectives in Biology and Medicine · Summer 1963 W. R. Hess · From Medical Practice to Theoretical Medicine 422 Perspectives in Biology and Medicine33· Summer 1963 From Medical Practice to Theoretical Medicine: An Autobiographic Sketch

time for a hobby. Through fortunate circumstances, in view of my in- clination to financially unpromising basic research, I was awarded a cash prize from the Swiss Marcel Benoist Foundation during the relatively early years of my work. This money was invested in a simple summer house south of the Alps in the region of the so-called "sun terrace of Switzerland." There I tended my vines and cared for my garden whenever I found it necessary or possible to relax a bit. My reward was not only a harvest of grapes and the pleasure given by the southern plants, but also acquisition of a variety of bits of information as well as the manual skills practiced by the farmers there since, presumably, as far back as Roman times. And the Italian attitude and language contributed much to my re- laxation and enjoyment. I would quickly recover from overwork and in the presence of lizards, scorpions, Aesculapius snakes, and other crawling beasts so alien to eastern Switzerland, feel myselfa new man. In this rustic setting, my mind would often turn to contemplating questions of pre- history, and I would become mentally refreshed and rested. And now, after sixty years of service on behalf of science, I contentedly put down my pen with the hope that perhaps my work has contributed to scientific progress in my own field, especially toward the understanding of regulatory functions at the integrative level of the organism.

34 423 SNS jubilee

Walter WalterR. Hess R.(17.3.1881–12.8.1973) Hess (17.3.1881–12.8.1973) SNS jubilee n C. W. Hess Christian W. Hess NeurologischeWalter Universitätsklinik, R. Hess Inselspital(17.3.1881–12.8.1973) Bern n C. W.Persönliche Hess Darstellung des Lebenslaufes und der wissenschaftlichen Beiträge von NeurologischeWalter Universitätsklinik, R. Hess, verfasst Inselspital von seinemBern Enkel und Neurologen Prof. Dr. Christian W. Hess. Nachdruck aus Schweizer Archiv für Neurologie und Psychiatrie, 159 (4):255-261, 2008

This article is based on Hess’s autobiographical who initially encouraged him to go into scientific sketches [1], on obituaries [2–4], articles about his research [1]. The family doctor Dr. Elias Haffter life work [5–10], his work and last but not least, (1851–1909), who treated young Walter for tuber- on writtenThis andarticle oral is traditions based on of Hess’s his family.Because autobiographicalculouswho pleurisy,initially encouraged also had a him major to go effect into scientificon him. of limitationssketches of[1], space, on obituaries only a [2–4],few aspects articles ofabout his hisHeresearch decided [1].to study The family medicine, doctor beginning Dr. Elias his Haffter stud- scientificlife workoeuvre [5–10], will hisbe worktouched and upon,last but and not anleast,ies (1851–1909),in Lausanne who in 1899treated and young later Walter pursuing for tuber-them extensiveon written discussion and oral of his traditions scientific of hisachievements family.Becausefurtherculous in pleurisy, Berlin, also Kiel had and a major Zurich. effect During on him. a of limitations of space, only a few aspects of his He decided to study medicine, beginning his stud- will not be attempted. For further information on summer term in Kiel, he was greatly impressed scientific oeuvre will be touched upon, and an ies in Lausanne in 1899 and later pursuing them these subjects, the reader is referred to the exten- by the riveting lectures of the legendary internist extensive discussion of his scientific achievements further in Berlin, Kiel and Zurich. During a Heinrich Quincke sive writingswill not beof attempted.Richard Jung For further [5] and information Konrad onProf.summer term in Kiel, he(1842–1922). was greatly Inimpressed Zurich Akert these[8], both subjects, of whom the reader collaborated is referred with to the Hess. exten-he bywas the fascinated riveting lectures by a vascular of the legendary anomaly internistthat he Hess’ssive more writings important of Richard original Jung articles [5] and can Konrad be observed Prof. Heinrich in the dissection Quincke (1842–1922).room, which Inhe Zurichinter- found Akertin an English[8], both translation of whom collaborated in the monograph with Hess.pretedhe was as fascinatedthe product by ofa vascular haemodynamic anomaly thatforces. he editedHess’s by Akert more [11]. important A complete original bibliography articles can of beHeobserved approached in the the dissection anatomist room, Prof. whichWilhelm he Rouxinter- his 300found articles in an appeared English translationin an obituary in the article monograph of (1850–1924)preted as ofthe Halle, product the offounder haemodynamic of developmen- forces. the Vienneseedited by Academy Akert [11]. [12] A and complete can also bibliography be found oftal Hemechanics, approached who the encouraged anatomist Prof.him toWilhelm publish Roux his in Akert’shis 300 monograph articles appeared [11]. in an obituary article ofobservation(1850–1924) [13]. of Halle, the founder of developmen- the Viennese Academy [12] and can also be found talAfter mechanics, passing whothe medicalencouraged qualifying him to publish examina- his in Akert’s monograph [11]. tionobservation in Zurich [13].in 1906, his first post was in Mün- Life sterlingenAfter as passing an assistant the medical in surgery qualifying to Dr. examina-Conrad Brunnertion in(1859–1927) Zurich in 1906, whose his firstefficient post wasand in tightly Mün- WalterLife R. Hess (fig. 1) was born on March 17, 1881, runsterlingen clinic served as an as assistant an example in surgery to him to Dr.thereafter.Conrad Brunner (1859–1927) whose efficient and tightly in Frauenfeld in the Canton of Thurgau in eastern He proposed introducing walking casts to encour- Walter R. Hess (fig. 1) was born on March 17, 1881, run clinic served as an example to him thereafter. Switzerland, the second of three children of age tissue growth through gradually increased in Frauenfeld in the Canton of Thurgau in eastern He proposed introducing walking casts to encour- Clemens Hess,originally from Zug,and of Gertrud weight-bearing, but, as he was still a neophyte, the Switzerland, the second of three children of age tissue growth through gradually increased Hess, néeClemens Fischer, Hess,originally originally from Zug,andMitweida of nearGertrudideaweight-bearing, was not accepted. but, as Nonetheless, he was still a surgeryneophyte, gave the ChemnitzHess, in née Thuringia. Fischer, originallyAlready asfrom a small Mitweida child, nearhimidea ample was opportunitynot accepted. to Nonetheless, pursue his surgeryreigning gave in- he mustChemnitz have possessed in Thuringia. a healthy Already degree as a smallof self- child,teresthim at ample that time opportunity and observe to pursue the vascular his reigning system in- confidence:he must having have learntpossessed the Germana healthy language degree of of self-in theterest living at that organism. time and Forobserve his clinicalthe vascular studies system he Germanyconfidence: from his having mother, learnt and the still German speaking language with ofdevelopedin the living a device organism. to measure For his clinical blood studiesviscosity he a SaxonGermany accent,he from criticised his mother, the teacher and still on speaking the first with(nameddeveloped “viscosimeter”), a device to whichmeasure later blood was usedviscosity ex- day of aschool Saxon for accent,he his faulty criticised pronunciation! the teacher Later on the he firsttensively(named in “viscosimeter”), clinical medicine which until later the introductionwas used ex- derivedday an of enthusiasm school for his for faulty nature pronunciation! from his father, Later heof blood-sedimentationtensively in clinical medicine measurement.To until the introduction his great a physicsderived teacher an enthusiasm in grammar for school nature whofrom didhis father,re- disappointment,of blood-sedimentation however, measurement.To his paper on the his subject great searcha in physics meteorology teacher andin grammar ran a weather school whostation. did re-wasdisappointment, rejected by Pflügers however, Archiv, his paperthe onleading the subject phy- As a pupil,search hein meteorologytook part in and experiments ran a weather in station.his siologywas rejectedjournal byof thePflügers day, Archiv, becausethe he leading was an phy- un- father’sAs physics a pupil, laboratory he took part and inhelped experiments him elec- in hisknownsiology beginner journal withoutof the day, a scientific because he mentor. was an Theun- trify thefather’s family’s physics apartment. laboratory It was and also helped his fatherhim elec-publication,known beginner which hewithout submitted a scientific as a dissertationmentor. The trify the family’s apartment. It was also his father publication, which he submitted as a dissertation in 1906, was finally published in the journal of the in 1906, was finally published in the journal of the scientific research society of Zurich [14]. scientific research society of Zurich [14]. Correspondence: He could not devote himself fully to his scien- Correspondence: He could not devote himself fully to his scien- Prof. Dr. med. Christian W. Hess Prof. Dr. med. Christian W. Hess tific pursuits because, in that era, assistants in Neurologische Universitätsklinik tific pursuits because, in that era, assistants in Neurologische Universitätsklinik theoretical university institutes were not paid a Inselspital theoretical university institutes were not paid a Inselspital liveable wage. Therefore, in 1907, he went to Prof. CH-3010CH-3010 Bern Bern liveable wage. Therefore, in 1907, he went to Prof. e-mail: [email protected]: [email protected] OttoOtto Haab Haab(1850–1931)(1850–1931) of of the the Zurich Zurich UniversityUniversity

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35 Walter R. Hess (17.3.1881–12.8.1973)

Figure 1

Figure 1

W. R. Hess as a medical student (left), as chairman of the Institute (centre) andW. afterR. Hess retirement as a medical with hisstudent parrot (left), “Joko” as (right).chairman of the Institute (centre) and after retirement with his parrot “Joko” (right). Ophthalmologic Department for two years to be offered him the position of an assistant in the trained as an ophthalmologist. Haab taught him Physiological Institute at the University of Zurich. both theOphthalmologic art of diagnosis Department of eye disease for two andyears the to beHeoffered decided him to theaccept position the offer of an after assistant careful in de-the surgicaltrained skills asthat an servedophthalmologist. him well notHaab justtaught in his himliberationPhysiological and Institutediscussion at thewith University his wife, of Zurich. whose practiceboth as anthe ophthalmologist, art of diagnosis butof eye also disease later in and his theagreementHe decided understandably to accept the came offer withafter mixed careful feel- de- animalsurgical experiments.Having skills that served no himone towell guide not himjust in hisingsliberation because ofand the discussion loss of material with his wealth. wife, whose his scientificpractice interests,he as an ophthalmologist, independently but alsodeveloped later in his agreementAt the Physiological understandably Institute came he with obtained mixed feel- the a systemanimal to identify experiments.Having the paretic noeye one muscle(s) to guide himin inrankings of because “Privatdozent” of the loss (scientific of material associate) wealth. one diplopiahis andscientific to quantify interests,he the independently deviation [15]. developed The year laterAt the (1913) Physiological with a thesis Institute on haemodynamics he obtained the a system to identify the paretic eye muscle(s) in rank of “Privatdozent” (scientific associate) one apparatus, which he called a “coordimeter”, is still [17], although the work displeased the then chair- diplopia and to quantify the deviation [15]. The year later (1913) with a thesis on haemodynamics in use among ophthalmologists today and known man of internal medicine,Prof.Hermann Eichhorst apparatus, which he called a “coordimeter”, is still [17], although the work displeased the then chair- as “Hess screen”. In the same period he invented (1849–1921), who considered the mathematical in use among ophthalmologists today and known man of internal medicine,Prof.Hermann Eichhorst a methodas “Hess for screen”.making Instereoscopic the same period photographs he inventedformulae (1849–1921), found in who it toconsidered be “foreign the to mathematical physiology” with aa finelymethod corrugated for making prism stereoscopic glass [16], photographs which [6].formulae After the found outbreak in it to of be the “foreign First Worldto physiology” War in has recentlywith a come finely into corrugated use again.He prism later glass intended [16], which1914, [6]. his After scientific the outbreak work was of thefrequently First World interrupt- War in to perfecthas recently this apparatus, come into establish use again.He a company later intended to ed 1914,by calls his toscientific active dutywork as was a captainfrequently in theinterrupt- Swiss produceto perfectit and takethis apparatus,out a patent establish on it, a but company these toArmyed byMedical calls to Corps. active duty The oneas a advantagecaptain in the the Swiss war plans cameproduce to nothingit and take because out a ofpatent the outbreakon it, but ofthesegaveArmy him Medicalwas that Corps. it enabled The one him advantage to take athe scien- war the Firstplans World came War.to nothing A few because specimens of the ofoutbreak these oftificgave leave him in was Germany, that it enabled as it was him relatively to take aeasy scien- to “stereothe pictures” First World decorated War. Athe few dining specimens room ofof his thesefindtific a good leave place in Germany, to do research as it was in viewrelatively of the easy many to apartment“stereo on pictures”the Zurichberg. decorated the dining room of hispositionsfind a good left placevacant to doby researchthe fighting. in view He of wasthe many thus In apartmentthe spring onof the1908, Zurichberg. after a brief sojourn in ablepositions to spend left the vacant year 1915/16by the fighting. at the Physiological He was thus Paris forIn further the spring training of 1908, in aftervenereology a brief sojourn and inInstituteable to ofspend the theUniversity year 1915/16 of Bonnat the Physiologicalunder Prof. neurology,Paris Hessfor furthertook over training an ophthalmologist’s in venereology andMaxInstitute Verworn of (1863–1923), the University at thatof Bonn time underthe leading Prof. practiceneurology, in RapperswilHess took (Canton over ofan Sanktophthalmologist’s Gallen) neurophysiologistMax Verworn (1863–1923), in Germany. at thatVerworn’s time theconcep- leading practice in Rapperswil (Canton of Sankt Gallen) neurophysiologist in Germany. Verworn’s concep- and thereby attained the necessary financial tions of physiology, his wide-ranging knowledge and thereby attained the necessary financial tions of physiology, his wide-ranging knowledge security to marry his fiancée Louise Sandmeier and his synthetic mode of thinking left lasting security to marry his fiancée Louise Sandmeier and his synthetic mode of thinking left lasting and start a family. Lisy, as she was called, was the impressions on Hess. Having been largely a scien- and start a family. Lisy, as she was called, was the impressions on Hess. Having been largely a scien- daughterdaughter of a Frauenfeld of a Frauenfeld lawyer; lawyer; she had she hadworked workedtifictific autodidact autodidact up up to to that that time, time,HessHess hadhad nownow as a doctor’sas a doctor’s assistant assistant in the in Ambulatorythe Ambulatory Eye Eye found, found, for for a shorta short time, time, a ascientific scientific mentor.mentor. The The Clinic Clinicat the at Zurich the Zurich Cantonal Cantonal Hospital Hospital and andnow nowhappyhappy situation situation did did not not last last because because Verworn Verworn suf-suf- becamebecame his devoted his devoted assistant assistant in his in outpatienthis outpatientferedfered the the first first of ofseveral several cerebral cerebral strokes strokes duringduring practicepractice and inand the in operatingthe operating room. room. His Hissister sisterHess’sHess’svisit.visit. HannaHannakept housekept house for them for them and lookedand looked after after their their BackBack in inZurich, Zurich, his his chief chief Gaule Gaulewaswas illill asas wellwell daughterdaughter Gertrud, Gertrud,who whowas bornwas born in 1910. in 1910. Their Their and and went went into into premature premature retirement retirement in in the the autumn autumn son Rudolfson Rudolf Max wasMax bornwas born3 years 3 years later. later. The Theoph- oph-of 1916.of 1916.HessHessbecamebecame acting acting chief chief of of the the InstituteInstitute thalmologythalmology practice practice flourished, flourished, with with a satellite a satelliteandand faced faced a good a good deal deal of of extra extra work work with with teaching teaching office officein the in Cantonal the Cantonal Hospital Hospital of Glarus,of Glarus, and andandand organisational organisational matters. matters. In In the the spring spring ofof 1917,1917, allowedallowed Hess Hessto buildto build up his up financialhis financial reserves. reserves.thethe faculty faculty proposed proposed taking taking on on asas newnew chiefchief a He keptHe on kept visiting on visiting scientific scientific meetings meetings and andalso also did didGermanGerman physiologist physiologist who who was was olderolder andand moremore experimentsexperiments on arterial on arterial blood blood pressure pressure and andwith withexperiencedexperienced than than Hess Hess. At. At that that time, time, in in Zurich,Zurich, artificial lenses on rabbits in the washhouse of the most of the “important” chairs were held by Ger- artificial lenses on rabbits in the washhouse of the most of the “important” chairs were held by Ger- garden. In 1912 Prof. Justus Gaule (1849–1939) mans, and a Swiss could hardly even be considered garden. In 1912 Prof. Justus Gaule (1849–1939) mans, and a Swiss could hardly even be considered

256 SCHWEIZER ARCHIV FÜR NEUROLOGIE UND PSYCHIATRIE www.sanp.ch 159 n 4/2008 256 SCHWEIZER ARCHIV FÜR NEUROLOGIE UND PSYCHIATRIE www.sanp.ch 159 n 4/2008

36 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

for an ordinary professorship and the direction Hess also made a valuable contribution to re- and head of an Institute in a theoretical discipline search in meteorological physiology. The Interna- like physiology. Hess was offered an extraordinary tional Foundation for the High Alpine Research for an ordinary professorship and the direction Hess also made a valuable contribution to re- professorship (i.e. non-chair full professor without Station Jungfraujoch was founded in 1930 under and head of an Institute in a theoretical discipline search in meteorological physiology. The Interna- being likethe physiology.head of theHess Institute),was offered but andeclined. extraordinary As his tionaldirection.The Foundation research for the station High wasAlpine inaugurated Research Hess hadprofessorship led the Institute (i.e. non-chair successfully full professor during without the onlyStation one yearJungfraujoch later andwas led founded by him in until1930 under1937. interregnumbeing the and head was wellof the liked Institute), by the students but declined. and AsThis,his too, direction.The was the period research of hisstation active was engagement inaugurated trainees,Hess thehad faculty’s led the decision Institute raisedsuccessfully a good during deal theagainstonly onethe yearso-called later “anti-vivisectionists”and led by him until, 1937. who of discontent.interregnum The andresponsible was well likeduniversity by the adminis-students andwishedThis, to too, forbid was the all period experimentation of his active onengagement animals. trativetrainees, body discovered the faculty’s that decision the faculty raised had a good con- dealHess againstfought the on so-called the front “anti-vivisectionists” lines of this struggle, who in sulted ofMax discontent. Verworn Thefor responsibleadvice about university the selection adminis-thewished name ofto theforbid faculty all experimentation and often bore on the animals. brunt of a newtrative head body of discoveredphysiology, that that the Verworn faculty hadhad con-of Hesshostilityfought [18]. on Thethe frontattacks lines culminated of this struggle in the in recommendedsulted Max Hess Verwornas thefor top advice candidate about theand selection that naivelythe name absurd of thedemand faculty of and one often embittered bore the oppo- brunt this expertiseof a new had head been of suppressed.The physiology, that head Verworn of thehadnentof that,hostility if animal [18]. Theexperimentation attacks culminated could notin thebe Cantonalrecommended Department Hess ofas Education, the top candidate Counsellor and thatcompletelynaively absurd dispensed demand with, of itone should embittered at least oppo- be Heinrichthis Mousson expertise (1866–1944,Educationalhad been suppressed.The head Direc- of theperformednent that, on if creatures animal experimentation “inimical to mankind,such could not be Cantonal Department of Education, Counsellor completely dispensed with, it should at least be tor from 1914 to 1929), actually went in person to as tigers and lions”! Heinrich Mousson (1866–1944,Educational Direc- performed on creatures “inimical to mankind,such discuss the situation with the students and then He faced the major challenge of organising the tor from 1914 to 1929), actually went in person to as tigers and lions”! attended one of Hess’s lectures during the summer 16th International Congress of Physiology in Zurich discuss the situation with the students and then He faced the major challenge of organising the term,attended obtaining one a offavourable Hess’s lectures impression during the [6]. summer In in 1938,16th International which only Congress took place of Physiology at all because in Zurich of the autumnterm, of obtaining 1917,Hess a favourablewas chosen impressionOrdinary Pro- [6]. Inhis inuncompromising 1938, which only leadership took place and at steadfastness.all because of fessor theand autumn Chairman of 1917, of theHess Physiologicalwas chosen Ordinary Institute Pro-Onhis the uncompromising eve of the Second leadership World War,and and steadfastness. right after by unanimousfessor and vote Chairman of the of Cantonalthe Physiological Governing InstitutetheOn “Anschluss” the eve of theof Austria Second toWorld Germany,attempts War,and right after by Councilby (Regierungsrat). unanimous vote of the Cantonal Governinga fewthe colleagues “Anschluss” toof exert Austria pressure to Germany,attempts of a political and by Hess’sCouncilfirst (Regierungsrat). step in office was to modernise anti-Semitica few colleagues nature to endangered exert pressure the of entire a political event and at the teachingHess’s methods.first step He in introduced office was a tocourse modernise in the anti-Semitic last minute nature [1]. Another endangered very the unpleasant entire event con- at experimentalthe teaching methods methods. and, He as introduced one of the a coursefirst, insequencethe last of minute the war [1]. was Another that all very of the unpleasant original page con- movingexperimental pictures as anmethods instructional and, as medium one of the(and, first,proofssequence and ofillustration the war was blocks that all forof the Hess’s originalmono- page later, asmoving a research pictures tool as anas instructionalwell). As the medium Institute (and,graphproofs on theand diencephalic illustration regulationblocks for ofHess’s circulationmono- staff waslater, very as asmall research until tool 1946 as (twowell). academic As the Institute as- andgraph respiration on the diencephalic (ed. 1938) were regulation destroyed of circulation by fire sistants,staff one was mechanic very small [Max until Jenny] 1946 and(two one academic secre- as-on andthe premisesrespiration of (ed. the Thieme 1938) were publishing destroyed house by fire in tary [Minasistants, Eugster] one mechanic), his wife [Max served Jenny] as hisand private one secre-Leipzig.on the Yet premises another of theresult Thieme of the publishing war was housethat the in tary [Mina Eugster]), his wife served as his private Leipzig. Yet another result of the war was that the secretary. At that time, his two first assistants Swiss physiologists, who had previously been asso- secretary. At that time, his two first assistants Swiss physiologists, who had previously been asso- and disciples Dr. Alfred Fleisch (1892–1973) and ciated with the German and French physiological and disciples Dr. Alfred Fleisch (1892–1973) and ciated with the German and French physiological Dr. Ernst Rothlin (1888–1972) were reliable and societies, founded their own national society on Dr. Ernst Rothlin (1888–1972) were reliable and societies, founded their own national society on diligentdiligent support support for him. for Hehim. also He hiredalso hired and andtrained trainedHess’sHess’sinitiative.initiative. untaughtuntaught workers workers who providedwho provided essential essential support support AnotherAnother scientifically scientifically productive productive phase phase ensued ensued for hisfor research: his research: histologist histologist Verena Verena Bucher Bucher, data, datain thein the post-war post-war years years with with the the support support of of the the Can-Can- managermanager Anna AnnaJaussi. Jaussi. tonaltonal authorities authorities (more (more staff), staff), the the Swiss Swiss NationalNational In his Inresearch his research he at hefirst at concentratedfirst concentrated on cir-on cir-ResearchResearch Foundation Foundation and and the the Rockefeller Rockefeller Foun-Foun- culatoryculatory and respiratory and respiratory regulation.He regulation.He was wasaward- award-dationdation of Newof New York.After York.After Hess Hessretiredretired to to the the rankrank ed the edMarcel the Marcel Benoist Benoist Prize Prizein 1932in 1932 for work for work in this in thisof Professorof Professor Emeritus Emeritus in in 1951, 1951, the the anatomist anatomist Prof.Prof. area. Hearea. used He the used prize the prizemoney money to buy to abuy plot a plot of land of landGianGian Töndury Töndury(1906–1985)(1906–1985) put put an an office office in in his his ownown in Asconain Ascona (in the (in Canton the Canton of Ticino), of Ticino), on which on which he heInstituteInstitute at Hess’sat Hess’sdisposal,disposal, where where he he continued continued to built abuilt small a smallsummer summer house house in 1934 in 1934 that thatbecame became work work on onthe the evaluation evaluation of of his his experimental experimental findings findings his hobby.his hobby. In 1934 In the1934 University the University of Berne of Berne made madefor fora numbera number of ofyears. years. His His disciple, disciple, Prof. Prof. OskarOskar him anhim honorary an honorary Doctor Doctor of Natural of Natural Philosophy; Philosophy;A.A. M. M. Wyss Wyss(1903–1992),(1903–1992), succeeded succeeded himhim asas headhead Hess furtherfurther honorary honorary degrees degrees (Geneva, (Geneva, McGill McGill [Mont- [Mont-of ofthe the Institute Institute of of Physiology. Physiology.Hess waswas highlyhighly real], Freiburg [Germany]) and prizes followed. satisfied to see his vision of a modern brain re- real], Freiburg [Germany]) and prizes followed. satisfied to see his vision of a modern brain re- It was also in the 1930s that he began his epoch- search institute become reality in 1962, under It was also in the 1930s that he began his epoch- search institute become reality in 1962, under making experiments with diencephalic stimulation the leadership of his last and most active disci- making experiments with diencephalic stimulation the leadership of his last and most active disci- on freely moving cats, for which he won the Nobel ple Prof. Konrad Akert (*1919). After moving to on freelyPrize moving(shared cats, with for Egas which Moniz) he won in the1949 Nobel “for thepleAscona Prof. Konrad in 1967, Akert he continued(*1919). to After carry movingon a wide- to Prize (shareddiscovery with of theEgas functional Moniz) inorganisation 1949 “for theof theAsconaranging in correspondence1967, he continued from to there carry and on to a receive wide- discoverydiencephalon of the functional as a coordinating organisation centre of visceralthe rangingvisitors correspondence from all over the from world, there including and to frequent receive diencephalonfunction”. as a coordinating centre of visceral visitorsvisits from from all his over children the world, and includinggrandchildren. frequent He function”. visits from his children and grandchildren. He

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37 Walter R. Hess (17.3.1881–12.8.1973)

died of heart failure in August 1973, at the age of ings in a larger biological context, with order and 92. His widow outlived him by 14 years. economy of conception being important guiding considerations. died of heart failure in August 1973, at the age of ings in a larger biological context, with order and Until the Second World War German physio- 92. His widow outlived him by 14 years. economy of conception being important guiding W. R. Hess as a person: logy was his intellectual home; he regularly visited considerations. his view of the world and mode of thinking the GermanUntil the physiological Second World congresses, War German at physio-which W. R. Hess as a person: he presentedlogy was his his intellectual results [5]. home; Nonetheless, he regularly he visitedmade Hess washis certainlyview of the a strict world chief and modewho placed of thinking heavy severalthe Germanacademic physiological trips to Britain congresses, and America at which to demands on himself, on the researchers working visithe thepresented Anglo-American his results [5]. physiologists Nonetheless, hewhose made under Hesshim wasand certainly on his astudents, strict chief some who ofplaced whom heavyworkseveral he attentively academic tripsfollowed. to Britain The andfact Americathat he atto occasionallydemands came on tohimself, fear him. on the Once researchers he gave a working stu- firstvisit published the Anglo-American only in German physiologists had the conse-whose dent theunder worst him possible and on grade his students, in the first some examina- of whomquencework thathe attentively his physiological followed. concepts The fact remainedthat he at tion ofoccasionally the university came course to fear when him. he Once had he detected gave a stu-unknownfirst published in the English-speaking only in German world had thefor aconse- long academicdent dishonesty the worst possible in the performance grade in the first of an examina- ex- time.quence that his physiological concepts remained periment.tion Heof the was university obviously course already when a he personage had detected unknownAs a speaker in the, heEnglish-speaking had a clear manner world for of a longoral demandingacademic respect dishonesty quite early in the on performance in his career, of yetan ex-expression;time. his extemporaneous speaking was pre- he regularlyperiment. held He discussions was obviously on important already a matters personagecise andAs direct.He a speaker found, he had writing a clear difficult,however, manner of oral with alldemanding of those working respect quite under early him. on in his career, yetas heexpression; often remarked. his extemporaneous His sentences speaking were was some- pre- he regularly held discussions on important matters cise and direct.He found writing difficult,however, Like his father before him, his worldview was times convoluted, and he generally rewrote his with all of those working under him. as he often remarked. His sentences were some- that of a freethinker, informed by natural science, manuscripts several times. Like his father before him, his worldview was times convoluted, and he generally rewrote his and hethat was of nota freethinker, religious. informed He repeatedly by natural empha- science, manuscriptsIn private, several he loved times. the Mediterranean and sised, however,and he was that not a religious.scientist Heshould repeatedly always empha- ac- often spentIn private, holidays he lovedwith his the family Mediterranean in Bordighera and knowledgesised, the however, limits of that scientific a scientist discovery,and should always he ac-on oftenthe Ligurian spent holidays coast. with In hishis familyyounger in Bordighera years, he thereforeknowledge explicitly the limitsrefused of scientificto rule outdiscovery,and the pos- hewason accompanied the Ligurian almost coast. everywhere,including In his younger years, the he sibilitytherefore of unknown explicitly powers refused and effects. to rule Uncondi-out the pos-Institute,was accompanied by his faithful almost dog. everywhere,including In my own school-age the tional sibilityrespect of for unknown religion powers and its and symbols effects. was Uncondi- as years,Institute, during by my his weeklyfaithful dog.visits, In Imy experienced own school-age my importanttional to respect him as for reverence religion andfor allits livingsymbols crea- was asthenyears, retired during grandfather my weekly as visits, a patriarchal I experienced family my tures,important including tohis him experimental as reverence animals. for all living It was crea-headthen who retired dominated grandfather conversations as a patriarchal around family the very importanttures, including to him his that experimental they should animals. be treated It wasfamilyhead dining who dominatedtable. This hadconversations become all around the more the respectfullyvery important and humanely. to him that they should be treatedinevitablefamily dining because table. of Thishis severe had become hardness all theof hear-more Hessrespectfullyheld great and respect humanely. for the Professor of ing.inevitable He was stillbecause driven of hisby severescientific hardness curiosity of hear- and NeurologyHess in Zurich,held greatConstantin respect for von the Monakow Professor ofbreding. snakes He was and still dissected driven by crabs scientific in his curiosity apartment and (1853–1930),Neurology and in attended Zurich, Constantinhis legendary von weeklyMonakowon thebred Zurichberg,while,in snakes and dissected the crabs luxuriant in his apartmentgarden of (1853–1930), and attended his legendary weekly on the Zurichberg,while,in the luxuriant garden of colloquia [8]. In accordance with von Monakow’s his summer house, he experimented with exotic colloquia [8]. In accordance with von Monakow’s his summer house, he experimented with exotic ideas, Hess did not believe in the existence of plants and cared for various species of grapevine, ideas, Hess did not believe in the existence of plants and cared for various species of grapevine, anatomically circumscribed nuclear “centres” in pears, apples and figs. His pet at that time, the talk- anatomically circumscribed nuclear “centres” in pears, apples and figs. His pet at that time, the talk- the brain.the brain. For Hess For Hessthe “centres”the “centres” were, were, rather, rather,inging parrot parrot “Joko”, “Joko”, whom whom he he had had brought brought upup fromfrom relativelyrelatively diffuse diffuse and sometimes and sometimes interpenetrating interpenetratingthethe beginning, beginning, was was always always by by his his side. side. functionalfunctional networks. networks.Hess’sHess’s scientific scientific mode mode of think- of think- ing wasing morewas moresynthetic synthetic than thananalytic, analytic, system- system- oriented,oriented, teleological, teleological, and also and alsointuitive. intuitive. His Hisform formScientificScientific achievements achievements of teleologyof teleology was basedwas based on biological on biological considera- considera- tions: ittions: was itconcerned was concerned with thewith goal-directed the goal-directed inte- inte-HisHis more more scientific scientific achievements achievements cancan bebe sum-sum- grativegrative performance performance of the of entire the entire organism, organism, rather rathermarisedmarised as asfollows follows (periods (periods of of activity): activity): than involvingthan involving any formany formof transcendental of transcendental ins- ins- pirationpiration [5, 8, [5, 19]. 8, Starting 19]. Starting from from this thispoint, point, he heA.A. The The organisation organisation and and regulation regulation of of the the circula-circula- developeddeveloped conceptions conceptions and formulated and formulated hypotheses hypotheses tiontion and and respiration respiration as as autonomic autonomic functionsfunctions that hethat tested he tested by experiment.Clarity by experiment.Clarity of conception of conception (1913–1931).(1913–1931). and stubbornand stubborn diligence, diligence, spurred spurred on by on goal-direct- by goal-direct-B.B. The The adrenergic-sympathetic adrenergic-sympathetic (“ergotropic”) (“ergotropic”) and ed formulation of hypotheses and reined in by cholinergic-parasympathetic (“trophotropic”) ed formulation of hypotheses and reined in by cholinergic-parasympathetic (“trophotropic”) methodological limitations, were the characteristic components of a dual, antagonistic autonomic methodological limitations, were the characteristic components of a dual, antagonistic autonomic features of his mode of working and the basis of (vegetative) system in the hypothalamus (1924– features of his mode of working and the basis of (vegetative) system in the hypothalamus (1924– his successes. Though he could become interested 1949). his successes.in new ideas Though and hefindings could in become conversation interested and dis- C.1949). The central representation of motivational and in newcussion, ideas and he restrictedfindings in his conversation practical research and dis- to at-C. Theinstinctive central representation behaviours such of motivationalas hunger, thirst, and cussion,tainable he restricted goals [5]. his He practical always tried research to put to his at- find- instinctivefear and ragebehaviours (1941–1943). such as hunger, thirst, tainable goals [5]. He always tried to put his find- fear and rage (1941–1943).

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38 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

D. The mechanisms of sleep as an active process [20] and the degenerated nerve fibres emanating and its induction by weak medial thalamic from the microcoagulated stimulation points were stimulation near the massa intermedia (1929– traced [20, 21]. D. The mechanisms of sleep as an active process [20] and the degenerated nerve fibres emanating 1944). The representation of the autonomic functions and its induction by weak medial thalamic from the microcoagulated stimulation points were E. The oculomotor system as a pre-cybernetic in the hypothalamus that Hess discovered placed stimulation near the massa intermedia (1929– traced [20, 21]. model1944). (1944–1946). these Thefunctions representation in two of anatomical the autonomic zones: functions the F. TheE. diencephalic The oculomotor postural system system as a of pre-cybernetic the body “trophotropic”in the hypothalamus (parasympathetic)that Hess discovered components placed and themodel reciprocal (1944–1946). relations between postural werethese found functions to be located in two in anatomical the anterior zones: (lateral) the andF. goal-directed The diencephalic motion postural (1941–1965). system of the bodyhypothalamus“trophotropic” as far (parasympathetic) as the septal nuclei,components while G. The biologicaland the reciprocal correlates relations of mental between functions posturalthewere “ergotropic” found to be (sympathetic) located in the components anterior (lateral) were and consciousnessand goal-directed (1943–1973). motion (1941–1965). foundhypothalamus to be located as infar the as posteriorthe septal ventromedial nuclei, while G. The biological correlates of mental functionshypothalamusthe “ergotropic” and perifornical (sympathetic) region.Stimulation components were Hess’s stimulationand consciousness technique: (1943–1973).The decisive break- in thefound posterior to be located hypothalamus in the posterior led, e.g. ventromedial to extreme through became possible with his development, in excitement,hypothalamus sometimes and perifornical with defence-like region.Stimulation behav- the lateHess’s 1920s, stimulation of a method technique: of stimulatingThe decisive the brain break-iour,in the ranging posterior all thehypothalamus way to flight led, e.g.or to extremea well- in non-anaesthetised,through became possible freely moving with his animals development, with indirectedexcitement, attack. sometimes Bilateral with lesions defence-like in this region,behav- electrodesthe late located 1920s, at of precisely a method defined of stimulating anatomical the brainon iour,the other ranging hand, all ledthe to way apathy to flight and anor adynamicto a well- sites, enablingin non-anaesthetised, him to explore freely systematically moving animals the withstatedirected or sleep-like attack. behaviour. Bilateral lesions Meanwhile, in this stimula- region, electrodes located at precisely defined anatomical on the other hand, led to apathy and an adynamic “vegetative” neuronal networks of the thalamus, tion in the rostral “trophotropic” zone led to a sites, enabling him to explore systematically the state or sleep-like behaviour. Meanwhile, stimula- hypothalamus and adjacent regions of the midbrain fall of blood pressure,slowing of respiration,pupil- “vegetative” neuronal networks of the thalamus, tion in the rostral “trophotropic” zone led to a and telencephalon. Hess’s hypothalamusOne and needsadjacent to regionsunderstand of the midbrainlaryfall constriction, of blood pressure,slowing and sometimes of respiration,pupil- even cardio-in- specialand stimulating telencephalon. techniqueOne needs[20], to which understand differed Hess’shibition.lary constriction, Hunger, thirst, and sometimes defecation even and cardio-in- micturi- fundamentallyspecial stimulating from the thentechnique usual[20], Faradic which stimu- differedtionhibition. could also Hunger, be provoked thirst, defecation by stimulation and micturi- in this lation,fundamentally in order to interpret from the the then stimulation usual Faradic effects stimu-region.tion could also be provoked by stimulation in this that helation, found in and order compare to interpret them the with stimulation the findings effects region.For Hess it was obvious that the stimulations al- of otherthat researchers. he found andHess compareaimed them at with specifically the findingsso evokedFor Hess concordantit was obvious emotions, that the and stimulations the induced al- targetingof otherthe smallresearchers. (poorlyHess myelinatedaimed at andspecifically un- behaviourso evoked was concordant appropriately emotions, referring and theto theinduced sur- myelinated)targeting fibres the smallof the (poorly autonomic myelinated system and (in un-roundings;behaviour e.g., was the appropriately cat which was referring put into to the an sur-ag- particularmyelinated) of the periventricular fibres of the grey),andautonomic at system avoid- (ingressiveroundings; mood e.g., attacked the cat the which nearest was putexperimenter into an ag- ing orparticular minimising of the stimulating periventricular effects grey),and on thickly at avoid-withgressive due precision, mood attacked when itthe was nearest allowed experimenter to do so. myelinateding or fibres minimising that might stimulating obscure effects the effects on thickly on Likewise,with due a catprecision, which was when stimulated it was allowed near theto do “fear so. the autonomicmyelinated system.To fibres that this might end, obscureHess developed the effects onzone”Likewise, watched a cat out which for awas suitable stimulated hiding near place, the “fear and a specialthe technique autonomic which system.To he labelled this end, “interruptedHess developedin onezone” case watched discovered out for a slightlya suitable open hiding roof place, window and a special technique which he labelled “interrupted in one case discovered a slightly open roof window direct-current (DC) stimulation”. Rather than the through which it promptly escaped. direct-current (DC) stimulation”. Rather than the through which it promptly escaped. brief (<0.5 ms) square-wave impulses of Faradic The localisation of autonomic and instinctual brief (<0.5 ms) square-wave impulses of Faradic The localisation of autonomic and instinctual stimulation, Hess used stimuli of long duration, functions in the hypothalamus has meanwhile been stimulation, Hess used stimuli of long duration, functions in the hypothalamus has meanwhile been typicallytypically 12.5 or 12.5 25 orms, 25 with ms, withramp-like, ramp-like, attenuated attenuatedconfirmedconfirmed in man,within man,with roughly roughly analogous analogous sites sites cor-cor- upwardupward and downward and downward slopes. slopes. To avoid To avoid polarising polarisingrespondingresponding to toeach each type type of of function: function: for for example,example, effectseffects that might that might damage damage the tissuethe tissue adjacent adjacent to tobilateral bilateral posterior posterior hypothalamic hypothalamic lesions lesions can can lead lead to the electrodes,the electrodes, he often he often used useda very a very weak weak “coun- “coun-apathyapathy and and hypothermia,while hypothermia,while bilateral bilateral rostral rostral andand tercurrent”tercurrent” between between the stimuli the stimuli so as so to as neutralise to neutraliselaterallateral hypothalamic hypothalamic lesions lesions can can lead lead to to anorexiaanorexia the cumulativethe cumulative net electrical net electrical charge. charge. The Thetrains trains of ofandand sometimes sometimes adipsia adipsia resulting resulting in in dehydrationdehydration stimulationstimulation were typicallywere typically lasting lasting 30 seconds 30 seconds or one or one[22,[22, 23]. 23]. Stereotactic Stereotactic lesions lesions in in the the posteromedial posteromedial minute.minute. Bipolar Bipolar and monopolarand monopolar stimulation stimulation was washypothalamushypothalamus have have even even been been used used to to free free patients patients used. Furthermore,used. Furthermore, the stimuli the stimuli were were weak weak (around (aroundfromfrom compulsive compulsive aggressive aggressive behaviour behaviour [24]. [24]. 0.5–1.50.5–1.5 V) and V) of and low of frequency low frequency (2–12 (2–12 Hz, Hz, usually usually Hess’sHess’sfindingsfindings with with regard regard to to sleep sleep[25,[25, 26], 26], on on 8 Hz).8 Great Hz). Greatcare was care taken was taken to limit to limit the spreadthe spread of ofthethe other other hand, hand, were were controversial controversial from from thethe veryvery the stimulatingthe stimulating currents, currents, which which were were estimated estimated to tobeginningbeginning and and remained remained so so for for many many years,years, even even be on thebe on order the orderof 0.1–0.15 of 0.1–0.15 mA. mA. He also He alsoused used much muchthoughthough his his son son Rudolf Rudolf M. M. Hess Hess, an, an electroencepha- electroencepha- finer electrodes than were customary at that time, lographer,reproduced these experiments using the finer electrodes than were customary at that time, lographer,reproduced these experiments using the with a diameter of 0.25 mm. Afterwards he went same stimuli in 1950 in collaboration with Konrad with a diameter of 0.25 mm. Afterwards he went same stimuli in 1950 in collaboration with Konrad to great effort to localise the anatomical site of Akert and Werner Koella and was able to record to great effort to localise the anatomical site of Akert and Werner Koella and was able to record stimulation precisely. He used electrocoagulation typical sleep-EEG patterns [27].Other researchers stimulationby the precisely.stimulating He electrodes used electrocoagulation to localise the site andtypicalwere sleep-EEG unable to patternsreplicate [27].Otherthe induction researchers of sleep by the atstimulating the same time electrodes to produce to localise small lesions the siteallowing and wereby unablemedial thalamicto replicate stimulation. the induction However, of unlikesleep at the sameobservation time to of produce the induced small behavioural lesions allowing changesby Hess,medial they thalamic used conventional stimulation. Faradic However, stimuli unlike or observation of the induced behavioural changes Hess, they used conventional Faradic stimuli or

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39 Walter R. Hess (17.3.1881–12.8.1973)

middle-frequency alternating currents which are amply confirmed [36]. In view of the unsettled not appropriate to excite unmyelinated nerve concepts on the mechanism of cataplectic loss of fibres [28–30]. Furthermore, because of the laten- muscle tone in patients with narcolepsy and the middle-frequency alternating currents which are amply confirmed [36]. In view of the unsettled cy between the salvo of stimulation and the ani- role of REM atonia, it is interesting to note that not appropriate to excite unmyelinated nerve concepts on the mechanism of cataplectic loss of mal’s actually falling asleep, it was suspected Hess induced muscle atonia when stimulating in a fibres [28–30]. Furthermore, because of the laten- muscle tone in patients with narcolepsy and the that Hess’s cats merely fell asleep because they felt circumscribed area of the anterior ventrolateral cy between the salvo of stimulation and the ani- role of REM atonia, it is interesting to note that comfortable.mal’s actually For Hess, fallinghowever, asleep, it was it was precisely suspectedhypothalamus Hess induced just muscle above atonia the whenchiasm stimulating [37]. Atonia in a the factthat that Hess’s the catscats curledmerely upfell comfortably asleep because before they feltstartedcircumscribed shortly after area onset of the of anteriorstimulation ventrolateral and out- fallingcomfortable. asleep that Forconfirmed Hess, however, that the it wasobserved preciselylastedhypothalamus stimulation just up aboveto 10 minutes.the chiasm [37]. Atonia sleep wasthe factphysiological, that the cats in curled addition up comfortablyto the fact that before startedIn the shortlyview of after Hess onsetit willof stimulation be of paramount and out- the catsfalling could asleep be immediately that confirmed reawakened. that the observedHess importancelasted stimulation to incorporate up to 10 the minutes. biological aspects consideredsleep wasthe physiological,slow response in additionto be typical to the factof a thatof psychicIn the functions view ofinto Hess theit domainwill be ofof physiologyparamount “trophotropic”the cats could function be immediately like sleep. reawakened. He explainedHessin orderimportance to understand to incorporate human the behaviour.biological aspects Con- the cats’considered not falling the asleep slow responseduring the to stimulationbe typical of asciousof psychic experience functions seemedinto theto himdomain to represent of physiology the train,“trophotropic” which lasted 30 function to 60 like seconds, sleep. He by explainedtheir supremein order level to understandof behavioural human integration. behaviour. How- Con- possiblythe being cats’ notkept falling awake asleep by concomitant during the stimulation stimu- ever,scious he experiencebelieved that seemed the tonature him to ofrepresent subjective the lation train,of other which systems lasted besides 30 to 60the seconds, “vegetative” by theirexperience supremewould level ofnot behavioural be causally integration.intelligible [8, How- 38, hypnogenicpossibly system being [26].kept awake In fact, by mild concomitant myoclonus stimu- 39].ever, In his he view, believed “the processthat the of nature becoming of subjective [subjec- in synchronylation of with other the systems stimulation besides was the sometimes “vegetative”tively]experience consciouswould cannot not be in causally principle intelligible be explained [8, 38, hypnogenic system [26]. In fact, mild myoclonus 39]. In his view, “the process of becoming [subjec- observed during stimulation trains. It is interesting in terms of itself”, due to fundamental limits of our in synchrony with the stimulation was sometimes tively] conscious cannot in principle be explained to note,in this context,that Hess regularly achieved mental potencies [38]. observed during stimulation trains. It is interesting in terms of itself”, due to fundamental limits of our the opposite effect (arousal) when he raised the to note,in this context,that Hess regularly achieved mental potencies [38]. stimulusthe intensity opposite aboveeffect (arousal)1.5 V [26]. when This hemay raised indi- the cate thatstimulus different intensity networks above with1.5 V opposite [26]. This effectsmay indi-References on vigilance and sleep are co-localised in the medial cate that different networks with opposite effects References thalamuson vigilance(midline and nuclei sleep / areintralaminar co-localised system in the medial of 1 Hess WR. From medical practice to theoretical medicine: an autobiographic sketch. the centralthalamus grey).(midline Perhaps nuclei this / isintralaminar the main reason system of 1 Hess WR. From medical practice to theoretical medicine: Perspect Biol Med. 1963;6:400–23. for thethe discrepant central grey). results. Perhaps In fact, this recent is the observa-main reason an autobiographic sketch. tions infor patients the discrepant have once results. again In fact,raised recent the ques-observa-2 CaspersPerspect H, Hess Biol Med. R, Kugler 1963;6:400–23. J, Petsche O, Struppler A. tion oftions the inrole patients of the have medial once thalamusagain raised in thesleep ques- 2Nachruf Caspers auf H,W. Hess R. Hess. R, Kugler Z EEG J, EMG.Petsche 1973;5:139–41. O, Struppler A. Nachruf auf W. R. Hess. Z EEG EMG. 1973;5:139–41. [31, 32],tion indicating of the role that of thenetworks medial withthalamus opposite in sleep3 Wyss OAM. Walter Rudolf Hess 17.3.1881–12.8.1973; [31, 32], indicating that networks with opposite 3Die WyssSelbständigkeit OAM. Walter der Rudolf Physiologie. Hess 17.3.1881–12.8.1973; effects are indeed probably to be found there. Die Selbständigkeit der Physiologie. effects are indeed probably to be found there. Schweiz Med Wochenschr. 1973;103:1758–9. According to this hypothesis, lesions in the medial Schweiz Med Wochenschr. 1973;103:1758–9. According to this hypothesis, lesions in the medial4 Akert K. Obituary: Walter Rudolf Hess (1881–1973). thalamus, depending on their precise site, extent 4 Akert K. Obituary: Walter Rudolf Hess (1881–1973). thalamus, depending on their precise site, extent Brain Res. 1974;68:V–VIII. and affected type of neurones, can cause either an Brain Res. 1974;68:V–VIII. and affected type of neurones, can cause either an5 Jung R. Walter R. Hess (1881–1973). arousal deficit (decreased vigilance, hypersomnia) 5 Jung R. Walter R. Hess (1881–1973). arousal deficit (decreased vigilance, hypersomnia) Rev Physiol Biochem Pharmacol. 1981;88:1–21. or a sleep deficit (insomnia), or a mixture of these Rev Physiol Biochem Pharmacol. 1981;88:1–21. or a sleep deficit (insomnia), or a mixture of these6 Waser PG. Walter Rudolf Hess: Aus seinem Leben 6 Waser PG. Walter Rudolf Hess: Aus seinem Leben two states, i.e. de-arousal without physiological, und seiner Tätigkeit an der Medizinischen Fakultät der two states, i.e. de-arousal without physiological, und seiner Tätigkeit an der Medizinischen Fakultät der deep non-REMdeep non-REM sleep. sleep. UniversitätUniversität Zürich Zürich (100-Jahr-Feier (100-Jahr-Feier seines seines Geburtstages: Geburtstages: 14. März 1981). Gesnerus. 1982;39:279–86. Hess’sHess’sconceptionsconceptions of the of organisation the organisation of motor of motor 14. März 1981). Gesnerus. 1982;39:279–86. functionfunctionhave havealso receivedalso received more more attention attention in in77 Huber Huber A. W.A. W.R. Hess,R. Hess, the the ophthalmologist. ophthalmologist. recentrecent years. years. In the In diencephalon the diencephalon he foundhe found the the Experientia.Experientia. 1982;38:1397–400. 1982;38:1397–400. representationrepresentation of spatio-temporal of spatio-temporal coordination coordination 8 8 Akert Akert K. WalterK. Walter Rudolf Rudolf Hess Hess (1881–1973) (1881–1973) and and his his of headof and head body and bodymovements movements in the in three the three cardinal cardinal contributioncontribution to neuroscience.to neuroscience. J HistJ Hist Neurosci. Neurosci. 1999;8:248–63. 1999;8:248–63. planesplanes of space of space[33]. [33].HessHessconsideredconsidered the themotor motor effectseffects elicited elicited from fromthe diencephalon the diencephalon to be to frag-be frag-99 Davison Davison J, Wilson J, Wilson G. G.Walter Walter Rudolf Rudolf Hess. Hess. ClinClin Experiment Experiment Ophthalmol. Ophthalmol. 2005;33:517. 2005;33:517. ments mentsof an ofintegrated an integrated motor motor system system [8] and[8] andem- em- 10 Koella WP. A tribute to W. R. Hess (1881–1973). phasisedphasised the reciprocal the reciprocal relations relations between between postural postural10 Koella WP. A tribute to W. R. Hess (1881–1973). Experientia.Experientia. 1982;38:1383. 1982;38:1383. (extrapyramidal)(extrapyramidal) and goal-directed and goal-directed motor motor systems. systems. 11 Akert K, editor. Biological Order and Brain Organization. Posture, in his view,not only stabilises and supports11 Akert K, editor. Biological Order and Brain Organization. Posture, in his view,not only stabilises and supports Selected Works of W. R. Hess. goal-directed movements, but also supplies the Selected Works of W. R. Hess. goal-directed movements, but also supplies the Berlin, Heidelberg, New York: Springer; 1981. necessary initial conditions (“motorische Bereit- Berlin, Heidelberg, New York: Springer; 1981. necessary initial conditions (“motorische Bereit- 12 Auerswald W. Nachruf auf Walter Rudolf Hess. schaft”) for voluntary action by providing a pro-12 Auerswald W. Nachruf auf Walter Rudolf Hess. schaft”) for voluntary action by providing a pro- Almanach Österr Akad Wissenschaft. 1975;124:414–33. active, anticipatory control [5, 11, 34, 35]. This con- Almanach Österr Akad Wissenschaft. 1975;124:414–33. active, anticipatory control [5, 11, 34, 35]. This con- 13 Hess WR. Eine mechanisch bedingte Gesetzmäßigkeit cept clearly went beyond Sherrington’s classical13 Hessim WR. Bau Einedes Blutgefäßsystems.mechanisch bedingte Gesetzmäßigkeit cept clearlyviews ofwent postural beyond reflexes Sherrington’s and has recently classical been im BauArch des Entwicklungsmech Blutgefäßsystems. Org. 1903;16:632–41. views of postural reflexes and has recently been Arch Entwicklungsmech Org. 1903;16:632–41.

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14 Hess WR. Zum Thema Viskosität des Blutes und 28 Ranson SW, Magoun HW. The hypothalamus. Herzarbeit. Dissertation. Vierteljahresschr Naturforsch Ergeb Physiol. 1939;41:56–163. Ges Zürich. 1906;51:236–51. 29 Hunsperger RW, Leisinger-Trigona M-C. Further studies 15 Hess14 WR. Hess Eine WR. neue Zum Untersuchungsmethode Thema Viskosität des Blutes bei Doppel-und 28on the Ranson alleged SW, hypnogenicMagoun HW. zone The hypothalamus.in the thalamus. bildern.Herzarbeit. Arch Augenheilkunde. Dissertation. 1908;62:233–8. Vierteljahresschr Naturforsch PerspectivesErgeb Physiol. in Brain 1939;41:56–163. Sciences. 1972;1:100–8. Ges Zürich. 1906;51:236–51. 16 Hess WR. Direkt wirkende Stereoskopbilder. 3029 Wyss Hunsperger OAM. Präzisierung RW, Leisinger-Trigona der Reizwirkung M-C. Furthermittelfrequenter studies Z Wiss15 Photogr Hess WR. Photophys Eine neue Photochem.Untersuchungsmethode 1914;14:33–8. bei Doppel- Wechselströme.on the alleged hypnogenic zone in the thalamus. bildern. Arch Augenheilkunde. 1908;62:233–8. PflügersPerspectives Arch Ges in Physiol.Brain Sciences. 1967;295:136–46. 1972;1:100–8. 17 Hess WR. Das Prinzip des kleinsten Kraftverbrauches im Dienste16 Hess hämodynamischer WR. Direkt wirkende Forschung Stereoskopbilder. [Habilitation]. 3130 Tinuper Wyss P, OAM. Montagna Präzisierung P, Medori der ReizwirkungR, Cortelli P,mittelfrequenter Zucconi M, Leipzig:Z Veit Wiss Arch Photogr Anat PhotophysPhysiol; 1914. Photochem. S. 1–62. 1914;14:33–8. BaruzziWechselströme. A, et al. The thalamus participates in the regula- Pflügers Arch Ges Physiol. 1967;295:136–46. 17 Hess WR. Das Prinzip des kleinsten Kraftverbrauches tion of the sleep-waking cycle. A clinico-pathological 18 Jäggi M. In primo loco. Geschichte der Medizinischen im Dienste hämodynamischer Forschung [Habilitation]. 31study Tinuper in fatal P, Montagnafamilial thalamic P, Medori degeneration. R, Cortelli P, Zucconi M, Fakultät Zürich 1833–2003. Zürich: Rüffer+Rub; 2004. Leipzig: Veit Arch Anat Physiol; 1914. S. 1–62. ElectroencephalogrBaruzzi A, et al. The Clin thalamus Neurophysiol. participates 1989;73:117–23. in the regula- tion of the sleep-waking cycle. A clinico-pathological 19 Akert18 K, Jäggi Hess M. WR. In primo Symposion loco. Geschichte über das Zwischenhirn.der Medizinischen 32 Bassettistudy C,in fatalMathis familial J, Gugger thalamic M, Lövbladdegeneration. K-O, Hess CW. Helv PhysFakultät Pharm Zürich Acta. 1833–2003. 1950;Suppl Zürich: VI:9–80. Rüffer+Rub; 2004. HypersomniaElectroencephalogr following Clin paramedian Neurophysiol. thalamic 1989;73:117–23. stroke: 20 Hess19 WR. Akert Beiträge K, Hess zur WR. Physiologie Symposion des über Hirnstammes. das Zwischenhirn. I. a report of 12 patients. Ann Neurol. 1996;39:471–80. Die Methodik der lokalisierten Reizung und Ausschaltung 32 Bassetti C, Mathis J, Gugger M, Lövblad K-O, Hess CW. Helv Phys Pharm Acta. 1950;Suppl VI:9–80. 33 Hess WR. Charakter der im Zwischenhirn ausgelösten subkortikaler Hirnabschnitte. Hypersomnia following paramedian thalamic stroke: 20 Hess WR. Beiträge zur Physiologie des Hirnstammes. I. Bewegungseffekte.a report of 12 patients. Ann Neurol. 1996;39:471–80. Leipzig: Thieme; 1932. S. 1–122. Die Methodik der lokalisierten Reizung und Ausschaltung Pflügers Arch Ges Physiol. 1941;244:767–86. 33 Hess WR. Charakter der im Zwischenhirn ausgelösten 21 Weisschedelsubkortikaler E, Jung Hirnabschnitte.R. Die anatomische Auswertung (EnglishBewegungseffekte. in [11], p. 223–42). und dasLeipzig: Studium Thieme; der sekundären 1932. S. 1–122. Faserdegeneration 34 HessPflügers WR. Teleokinetisches Arch Ges Physiol. und 1941;244:767–86. ereismatisches nach lokalisierter subcorticaler Ausschaltung durch 21 Weisschedel E, Jung R. Die anatomische Auswertung Kräftesystem(English in in[11], der p. Biomotorik. 223–42). Helv Physiol Acta. Elektrokoagulation.und das Studium der sekundären Faserdegeneration 341943;1:C62–3. Hess WR. Teleokinetisches (English in [11], und p. ereismatisches 265–8). Z Anat Entwicklungsgesch.nach lokalisierter subcorticaler 1939;109:374–95. Ausschaltung durch Kräftesystem in der Biomotorik. Helv Physiol Acta. Elektrokoagulation. 35 Hess WR. Cerebrale Organisation somatomotorischer 22 Plum F, Van Uitert R. Nonendocrine diseases and 1943;1:C62–3. (English in [11], p. 265–8). Z Anat Entwicklungsgesch. 1939;109:374–95. Leistungen. I. Physikalische Vorbemerkungen und disorders of the hypothalamus. 35Analyse Hess konkreterWR. Cerebrale Beispiele. Organisation somatomotorischer Res22 Publ Plum Assoc F, Van Res Uitert Nerv R. Ment Nonendocrine Dis. 1978;56:415–73. diseases and Leistungen. I. Physikalische Vorbemerkungen und disorders of the hypothalamus. Arch Psychiatr Nervenkr. 1965;207:33–44. 23 Carmel PW. Vegetative dysfunctions of the hypothalamus. Analyse konkreter Beispiele. Res Publ Assoc Res Nerv Ment Dis. 1978;56:415–73. 36 Stuart DG. Integration of posture and movement: Acta Neurochir (Wien). 1985;75:113–21. Arch Psychiatr Nervenkr. 1965;207:33–44. 23 Carmel PW. Vegetative dysfunctions of the hypothalamus. contributions of Sherrington, Hess, and Bernstein. 36 Stuart DG. Integration of posture and movement: 24 Sano J, ActaMayanagi Neurochir Y. Posteromedial (Wien). 1985;75:113–21. hypothalamotomy Hum Mov Sci. 2005;24:621–43. in the treatment of violent, aggressive behaviour. contributions of Sherrington, Hess, and Bernstein. 24 Sano J, Mayanagi Y. Posteromedial hypothalamotomy 37 HessHum WR, Mov editor. Sci. 2005;24:621–43.Hypothalamus und Thalamus. Acta Neurochir Suppl (Wien). 1988;44:145–51. in the treatment of violent, aggressive behaviour. Experimental-Dokumente. 37 Hess WR, editor. Hypothalamus und Thalamus. 25 Hess WR.Acta The Neurochir mechanism Suppl of (Wien). sleep. 1988;44:145–51. Stuttgart: Thieme; 1968. S. 63. Experimental-Dokumente. Am J Physiol. 1929;90:386–7. 25 Hess WR. The mechanism of sleep. 38 HessStuttgart: WR. Interdisciplinary Thieme; 1968. discussion S. 63. of selected Am J Physiol. 1929;90:386–7. 26 Hess WR. Das Schlafsyndrom als Folge dienzephaler 38problems Hess WR. with Interdisciplinary reference to “The discussion biology of of selected mind”. Reizung.26 Hess Helv WR. Physiol Das SchlafsyndromActa. 1944;2:305–44. als Folge dienzephaler Perspectproblems Biol with Med. reference 1970;13:267–93. to “The biology of mind”. (English in [11], p. 131–70). Reizung. Helv Physiol Acta. 1944;2:305–44. 39 HessPerspect WR. Causality, Biol Med. consciousness 1970;13:267–93. and cerebral (English in [11], p. 131–70). 27 Hess RM Jr, Akert K, Koella W. Les potentiels bio- 39organization. Hess WR. Causality,Science. 1967;158:1279–83.consciousness and cerebral électriques27 Hess du RM cortex Jr, Akert et duK, thalamusKoella W. Leset leur potentiels altération bio- organization. Science. 1967;158:1279–83. par stimulationélectriques du ducentre cortex hypnotique et du thalamus chez leet chat.leur altération Rev Neurol.par stimulation 1950;83:537–44. du centre hypnotique chez le chat. Rev Neurol. 1950;83:537–44.

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41

Walter R. Hess (1881–1973) and his Journal of the History of the 0964-704X/99/0803-248$15.00 1999,Contribution Vol. 8, No. 3, pp. 248–263 to Neuroscience © Swets & Zeitlinger

KonradWalter Akert Rudolf Hess (1881-1973) and His Contribution to Journal of the History of the Neurosciences 0964-704X/99/0803-248$15.00 1999, Vol. 8, No. 3, pp. 248–263 Neuroscience* © Swets & Zeitlinger Beitrag von Prof. Konrad Akert, Gründer und Direktor des Instituts für Hirnforschung und ehemaliger Rektor der UniversitätKonrad Zürich Akert zu den neurowissenschaftlichen Beiträ- genWalter von W. R. Rudolf Hess. NachdruckBrain Hess Research (1881-1973)aus Institute Journal of theof Universitythe and History Hisof Zürich of Contributionthe Neurosciences, to 8(3): 248-263, 1999 Neuroscience*

Konrad Akert Brain Research InstituteABSTRACT of the University of Zürich

The present account includes a brief life history of Walter Rudolf Hess (1881-1973) and a review of his major neurophysiological contributions. Hess belonged to the nearly extinct generation of great universal- ists among students of human physiology. He began his scientific work in hemodynamics and ophthalmol- ogy, then studied the functional organizationABSTRACT and regulation of circulation and respiration and developed a number of highly sophisticated instruments which fostered his international reputation. By electrically Thestimulating present the account brainincludes of the unanesthetized, a brief life history freely of moving Walter animal Rudolf he Hess explored (1881-1973) the functional and a review organization of his majorand localization neurophysiological of the cat contributions. diencephalon Hess in terms belonged of autonomic, to the nearly extrapyramidal extinct generation motor of functions, great universal- and in- istsstinctive among behavior, students e.g. of human hunger, physiology thirst, fear,. He and began rage. his His scientific thoughts work on biological in hemodynamics order led and him ophthalmol- to consider ogy,the problems then studied of psychic the functional forces. organization He was convinced and regulation of the close of circulation correlation and of respiration behavioral and research developed and aneurophysiology number of highly and sophisticated believed that instruments neuronal patterns which determinefostered his the international content of consciousness reputation. By without electrically pro- stimulatingviding clues the concerning brain of the the unanesthetized, transformation offreely such moving patterns animal into subjective he explored experience. the functional organization and localization of the cat diencephalon in terms of autonomic, extrapyramidal motor functions, and in- Keywords:stinctive behavior,Neurophysiology, e.g. hunger, Thalamus, thirst, fear, Hypothalamus, and rage. His Autonomic thoughts on nervous biological system, order Extrapyramidalled him to consider motor system,the problems Motivational of psychic behavior, forces. Psychophysiology, He was convinced Consciousness, of the close Hess, correlation W. of behavioral research and neurophysiology and believed that neuronal patterns determine the content of consciousness without pro- viding clues concerning the transformation of such patterns into subjective experience. BRIEF LIFE HISTORY role of the internal resistance of blood as an en- Keywords: Neurophysiology, Thalamus, Hypothalamus,ergetic Autonomic factor nervous which system, could in Extrapyramidalfluence the effi motorcacy system, Motivational behavior, Psychophysiology, Consciousness, Hess, W. Walter Rudolf Hess (Fig. 1) was born March 17, of cardiac work. To facilitate research along this 1881 in Frauenfeld, a small town in German line he designed a new method for the determi- BRIEFspeaking LIFE Eastern HISTORY Switzerland. He studied medi- rolenation of ofthe blood internal viscosity, resistance an ofinstrument blood as anwhich en- cine in Lausanne, Bern, Berlin and Kiel and ergeticconquered factor the which medical could laboratories influence the under efficacy the Waltergraduated Rudolf from Hess the (Fig. University 1) was born of MarchZürich 17, in ofname cardiac of ‘Hess work. viscosimeter To facilitate’ research(Hess, 1907). along this He 1905.1881 in Already Frauenfeld, as a a medical small town student in German he per- linethen he decided designed to take a new a residency method for in ophthalmol-the determi- formedspeaking his Easternfirst piece Switzerland. of research He in studied the anatomi- medi- nationogy and of soon blood came viscosity, out with an a newinstrument technique which for calcine dissecting in Lausanne, room: Bern, an anomaly Berlin ofand the Kiel arterial and conqueredthe exact planimetric the medical identi laboratoriesfication of under eye mus- the treegraduated which from in his the opinion University was due of to Z hemody-ürich in clename deviations of ‘Hess in viscosimeter diplopic patients’ (Hess, (Hess, 1907). 1908). He 1905.namic Alreadyforces. He as approached a medical the student then he famous per- Thetheninstrument decided to takeis still a residency in use in in modi ophthalmol-fied form pioneer in developmental mechanics, Wilhelm under the name of ‘Hess screen’ (Fig. 2). A sim- formed1 his first piece of research in the anatomi- ogy and soon came out with a new technique for calRoux dissecting, who kindly room: invited an anomaly him to a of personal the arterial dis- theilarly exact ingenious planimetric invention identi concernedfication of the eye manu- mus- treecussion which and in encouraged his opinion publication was due to of hemody- his re- clefacture deviations of photographic in diplopicfi patientslm coated (Hess, with 1908). a spe- namicmarkable forces. observation He approached (Hess, 1903). the then In his famous medi- Thecial emulsion instrument containing is still in a use contiguous in modi setfied of form tiny pioneercal thesis in of developmental 1906 he discussed mechanics, the important Wilhelm underlenses the (much name like of an‘Hess insect screen eye)’ which(Fig. 2). gave A sim- rise Roux1, who kindly invited him to a personal dis- toilarly direct ingenious stereoscopic invention images. concerned This early the interest manu- in the oculomotor system and binocular vision 1cussionWilhelm and Roux encouraged (1850-1924), publication Professor of of Anatomy, his re- facture of photographic film coated with a spe- Universitymarkable observationof Halle. (Hess, 1903). In his medi- cialshould emulsion later come containing to play a a contiguous significant roleset of in tiny the cal thesis of 1906 he discussed the important lenses (much like an insect eye) which gave rise * Address correspondance to: Konrad Akert, Hirslanderstr.to 33, direct CH-8032 stereoscopic Zürich, Switzerland. images. This Tel.: early 0041-1-382 interest 15 83. in the oculomotor system and binocular vision 1 Wilhelm Roux (1850-1924), Professor of Anatomy, University of Halle. should later come to play a significant role in the

* Address correspondance to: Konrad Akert, Hirslanderstr. 33, CH-8032 Zürich, Switzerland. Tel.: 0041-1-382 15 83.

43 Walter R. Hess (1881–1973) and his Contribution to Neuroscience

NEUROSCIENCE CONTRIBUTIONS OF W.R.HESS 249

formation of general concepts of sensorimotor tions by Max Verworn7 which he obtained in coordination. From 1908-1912 he practiced oph- Bonn during a postdoctoral leave of absence in thalmology in the nearby lakeshore town of 1915. His university teaching assignment con- Rapperswil. Yet, his keen sense of scientific sisted of physiology in the broadest sense, even research and his profound interest in fundamen- including physiological chemistry. Similarly tal mechanisms in biology and medicine broad was the spectrum of research topics: aside prompted him to abandon prosperity in the life from brain research it ranged from circulation, of a country doctor and to return to medical respiration and temperature regulation to sport – school in Zürich as an assistant in physiology. high altitude (Hess, 1929) – and work physiol- His elegant viscosity studies were extended into ogy. The comprehensive treatment of the non- quantitative investigations on the geometry and neurophysiological topics of a man who worked cross-sectional relations of the vascular system. seven days per week all his life, and even spent This led to the ‘Habilitationsschrift’ on the prin- his holidays excessively writing, would exceed ciple of minimal energy dissipation in hemody- the space allotted for this essay. namics (Hess, 1914) and four years later to the The main topics in neurophysiology were: (1) chair of physiology; a remarkable achievement the diencephalon and its relations to the auto- in view of the sequence of brilliant German pre- nomic (vegetative) nervous system8, (2) sleep decessors such as Ludwig2, Moleschott3, Fick4, mechanisms, (3) the problem of biological order Hermann5 and von Frey6. exemplified in the diencephalic postural system He held this position until his retirement in of the body, (4) the oculomotor system as a 1951. This was of course not meant to be the end precybernetic model, (5) the mutual relation- of the scientific work. The main concern during ships of autonomic (vegetative) and psychic the following years was an outlook on psycholo- functions, and (6) the biological correlates of gy taken from the biological point of view causality and consciousness. (Hess, 1962). At this time he changed his retreat Attendance at scientific meetings in Germany from Zürich to Ascona where he profited from a and his close contact to a group of foreign col- milder climate and enjoyed his exuberant gar- leagues in the neighboring countries, i.e., Fré- den. Death came to him in 1973. déric Brémer (Brussels); Paul Hoffmann Hess belonged to the generation of universa- lists of the previous century among the represen- 7 Max Verworn (1863-1923), Professor of Physiology tatives of human physiology. His theories and in Bonn. experiments concentrated on the systemic level. 8 Throughout his scientific life he remained an Hess’s terminology ‘Vegetatives system’ deserves a short comment since it is not usually used in the cur- autodidact, although in his autobiographic rent world literature. Clearly, it means the internal sketch (Hess, 1963) he acknowledged inspira- organs (viscera) as opposed to the ‘Animals’ system’ meaning the sensory organs and the motor system: a dichotomy dating back to the concept of the great French physiologist Bichat (1800: ‘La vie organique’ 2 Carl (K.F.W.) Ludwig (1816-1895), Professor of vs ‘La vie animale’) and its translations by his Ger- Anatomy and Physiology in Zürich 1849-56. man colleague Reil (1807: ‘Das vegetative vs das animale Nervensystem’). Bichat’s term: ‘Le système 3 Jakob Moleschott (1822-1893), Professor of Physiol- des ganglions’ became converted into the ‘autonomic ogy in Zürich 1856-61. nervous system’ by the British pharmacologist Langley (1898). Hess acknowledged the automatism 4 Adolf Fick (1829-1901), Professor of Physiology in but not the autonomy of the internal organs, and in- Zürich 1861-68. stead emphasized their obvious anatomic and func- tional dependence on the central nervous system – the 5 Ludimar Hermann (1868-1914), Professor of Physi- latter being quite manifest in normal emotional life ology in Zürich 1868-84. and particularly in psychosomatic diseases. Neverthe- less, this author shall use the term ‘autonomic’ which 6 Max von Frey (1852-1932), Professor of Physiology is overwhelmingly upheld today – not only in the in Zürich 1898-99. Anglosaxon literature.

44 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

250 KONRAD AKERT

Fig. 1. Walter Rudolf Hess in his seventies.

(Freiburg i.B.); Richard Jung (Freiburg i.B.); (Utrecht); Hugo Spatz (Munich) and others, pro- Charles Kaiser (Strasbourg); Ernst Kretschmer vided important opportunities for the exchange (Tübingen); Gösta Liljestrand (Stockholm); of ideas. Yet, the two world wars severely inter- (Seewiesen); Rudolf Magnus rupted the communication with colleagues in

Fig. 2. The ‘Hess screen’. Graphic coordimeter demonstrating the paralysis of rectus externus muscle. Dashed square shows limited motility of paralysed muscle. (W.R. Hess, 1950, Fig.2).

45 Walter R. Hess (1881–1973) and his Contribution to Neuroscience

NEUROSCIENCE CONTRIBUTIONS OF W.R.HESS 251

foreign countries. An additional handicap was were summarized in monographs (Hess, 1930, the fact that Hess’s work appeared almost exclu- 1931, 1948). sively in the German language. Hess’s name In all this endeavor Hess was searching for therefore remained virtually unknown in the the driving forces, as well as for the organizing Anglosaxon literature for a long time. Even after principles at the systemic level, thereby coming the wars one rarely finds Hess’s original papers close to Walter B. Cannon’s concept of homeo- and monographs quoted. stasis (Cannon, 1915). Hess went further in an- other direction by asking for aim and meaning behind bodily activities (although the concept of FROM CIRCULATION AND RESPIRATION teleology was at that time extremely controver- TO THE ORGANIZATION OF THE AUTO- sial by leading naturalists). To demonstrate na- NOMIC NERVOUS SYSTEM ture’s effort of optimizing functional perfor- mance in the economic context was one of the During the initial phase of his research career characteristics in Hess’s thinking. The dual an- Hess was engaged mainly in the field of hemo- tagonistic components, the adrenergic-sympa- dynamics and respiratory functions, as already thetic and the cholinergic-parasympathetic in- indicated. He began with experimental studies of nervation of the internal organs, appeared to him the dynamics of peripheral blood flow, then as an example of synergism allowing periods of turned step by step to the segmental level, i.e., maximal tension under stressful conditionsalter- the reflex control of arteries and heart, and fi- nating with periods of rest, sleep, and recovery; nally to the cerebral representation of circula- the former guided by the ergotropic system in tion. which the adrenergic component was leading, A similar approach from vagal reflex action the cholinergic subdued (Fig. 3); in the opposite to central regulation characterizes Hess’s exper- case, when in the context of economy and pro- imental work on respiration. These early efforts tective exigencies, the trophotropic system pre-

Fig. 3. Regulation of the responsiveness (‘Leistungsbereitschaft’) of the ‘Animals’ System’ (A) (= sensorimo- tor system) by the two antagonistic divisions of the ‘Vegetative System’.E=Ergotropic division, T = Trophotropic division of the vegetative system. Dashed lines indicate inhibition. Further explanation see text. (W.R. Hess, 1933, Fig.2).

46 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

252 KONRAD AKERT vailed (Hess, 1933). So far, however, this con- the smallest possible stainless-steel electrodes to cept has not been widely accepted; neither has it minimize the size of the brain lesions. Second, gained access to the textbooks, even though systematic experiments were undertaken to esta- Hess’s analysis of biological energetics and eco- blish the current form (low frequency, direct nomics at the systemic level deserves growing current pulses with special wave form) that attention especially in present times of molecu- proved to be effective at minimal threshold and lar dominance. minimal current spread and therefore most suit- This new concept led to the question of the able for the thinly myelinated periventricular site of the central operational elements which grey (Hess, 1932). His new approach led to sen- was to be sought somewhere in the forebrain sational results which made him widely known. where somatomotor and autonomic inputs are Much attention was given to the precise guiding processed and correlated with psychologic func- of the electrodes and the localization of stimu- tions (Hess, 1924/25). Encouraged by the pio- lated ‘‘points’’. For this purpose he prepared his neering studies of Karplus & Kreidl (1910) and own histological atlas on the basis of complete Bard (1929) on the connections between the serial histological sections in the three cardinal sympathetic nervous system and the hypothala- planes of the cat diencephalon, thus facilitating mus, Hess chose the diencephalon for his monu- the positioning of the stimulating electrodes. mental work in brain research. Indeed, the pres- (Fig. 4). The Hess atlas was equally important ence of important receptors (e.g., thermo- and for the point-to-point mapping of the representa- osmoreceptors), the convergence of visceral, tion of the virtually thousands of stimulation gustatory, and olfactory inputs as well as the effects recorded cinematographically between potent outflow to the pituitary, the powerful au- 1927 and 1949. tonomic and extrapyramidal motor output sys- The basic and still controversial problem of tems seemed to predispose diencephalic regions brain localization accompanied Hess throughout for regulatory mechanisms. It should be noted these initial years of experimentation. He did not that similar thoughts had been brought forward believe in circumscript centers as anticipated by by his contemporary, S.W. Ranson and his phrenologists and their followers, but much like school in Chicago, whose electrical stimulation von Monakow9 he preferred widespread fibre and ablation studies of hypothalamic regions networks whose connections he hoped to dis- paralleled those of Hess for some time and led close with combined physiologic and anatomic partly to similar conclusions (Ranson & Ma- goun, 1939). Only subsequent experiments by 9 Fulton (1939), Hunsperger & Bucher (1967) and Constantin von Monakow (1853-1930), founder and director of the Neurological Policlinic and of the Insti- others identified the limbic allocortex and the tute of Brain Anatomy of the University of Zürich orbitofrontal neocortex as significant autonomic (1894-1928), had reached the climax of fame when representation areas at the highest level. Hess began his scientific career. Monakow had been the highly respected leader of a group of students and admirers who met periodically in his home under the name of ‘Monakow’s Kränzchen’ where problems of BRAIN RESEARCH: A NEW TECHNICAL brain localization and the brain-mind enigma were APPROACH vividly discussed from neurological, psychiatric, and philosophical points of view. Hess frequented the in- Instead of using conventional procedures dating formal gatherings of this group and held Monakow in back to the time of electrical brain stimulation high esteem. Relations with Monakow’s successsor, Mieczyslaw Minkowski (1884-1972) were relatively by Fritsch & Hitzig (1870), Hess’ basic inten- cool. An attempt at collaboration with Minkowski on tion was to use a near natural stimulus to excite the anatomical analysis of Hess’ experimental brain the living unanesthetized brain of freely moving material concerning extrapyramidal motor symptoms animals. Thus, the first step consisted in the de- (Minkowski, 1943) ended when the physiologist and velopment of new hardware for the exploration the anatomist could not agree on matters of interpreta- tion, whereupon Hess enlisted Verena Bucher (1901- of the depths of the cat brain. First, he prepared 1986) as an expert on the anatomy of the cat brain.

47 Walter R. Hess (1881–1973) and his Contribution to Neuroscience

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Fig. 4. Stimulation electrodes mounted to the skull. Wires attached. The animal can move around in ‘‘trolleybus fashion’’ during stimulation. (W.R. Hess, 1942, Fig.2).

methods. Together with Ewald Weissschedel & Aside from the comfortably relaxed posture of Richard Jung (1939) and assisted by Verena the sleeping cat and the shielding of the organ- Bucher he began to trace by means of the ism from the outside world, like closure of the Marchi method the degenerated nerve fibers em- eyes and the selective elevation of sensory anatingfrommicrocoagulatedstimulationpoints thresholds, he put considerable weight on (Hess, 1932). A huge systematic collection of changes of the autonomic (parasympathetic) functional and morphological data involving nervous system, e.g., decrease of pulse rate and thalamus, hypothalamus and adjacent regions of blood pressure, contraction of the sphincter telencephalon and midbrain was analyzed and muscles (pupil, anus, bladder) as well as the interpreted (Hess, 1949, 1956). This relentless dark adaptation of the retina. All these signs experimental and synthetical effort, aided by a seemed to mark increased (and not decreased) small team of only 2 assistants, 1 mechanic, 1 activity. Thus, for Hess the bodily reactions dur- secretary/ technician, and 1 graduate student ing sleep were manifestations of an active state (Fig. 5), led to nearly 300 original articles and a and in contrast to anesthesia not a passive state number of monographic reviews and eventually with clearly diminished ergotropic functions. to the Nobel prize in physiology or medicine in The electrical exploration of the diencephalon 1949 ‘‘for his discovery of the functional orga- revealed a relatively widespread region in which nization of the interbrain as a coordinator of the positive sleep behavior with all the typical activities of the internal organs’’. symptoms could be induced at low threshold. He worked with strictly standardized conditions with respect to the experimental surroundings SLEEP INDUCED BY ELECTRICAL STIM- and time of stimulation: before applying electri- ULATION cal impulses the cats were kept in a normal ex- citatory state. The sleep-positive and negative Hess conceived sleep as a typical form of regions of the diencephalon and its surroundings trophotropically regulated behavior. He derived were marked in his atlas on the basis of histolog- this notion primarily from natural observations: ical verification. The sleep region extended from

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Fig. 5. The experimental team in place (R. Jung, 1981, Fig.3b).

the medial thalamus well towards the caudate identified. He suspected ultrafine unmyelinated nucleus. Short reports on electrically induced fibers which would escape the available micro- sleep appeared in 1927; they were presented be- scopic techniques and belonged to cerebral ana- fore the International Congress of Physiological logues of the fibers of the autonomic nervous Sciences in Boston (Hess, 1929). His early find- system which regulate the readiness (‘Leis- ings (1927) were confirmed and extended in tungsbereitschaft’) of cortical neurons in a syn- subsequent experiments (Hess, 1933). His son, ergistic fashion. Much of his work was ex- Rudolf M. Hess who introduced electroencepha- panded after the Second World War by Moruzzi lography in Switzerland, combined electrical & Magoun (1949), Jasper (1949), Magoun stimulation in the freely moving animal with (1954), Jouvet (1961), Moruzzi (1963) and oth- EEG recordings and found the induced sleep ers by means of electroencephalographic and stages of the cats to be accompanied by typical histochemical methods. It turned out that the normal EEG sleep patterns (Hess jr. et al., cortex indeed received an ascending input from 1950). lower brainstem nuclei carrying not only adren- Critical arguments have been raised by subse- ergic and cholinergic, but a host of other, modu- quent investigators (see Ranson & Magoun, lating transmitters influencing the impulse traf- 1939; Hunsperger & Leisinger-Trigona, 1972) fic within cortical grey matter. Today, we real- who claimed that Hess’s cats fell asleep sponta- ize that sleep mechanisms are far more complex neously. Consensus at that time could not be than anticipated, but the basic ideas of Hess are reached. How then to interprete Hess’s findings still compatible with present day findings in retrospect in the light of present knowledge? (McCormick & Bal, 1997; Borbély & Tononi, He himself postulated sleep-mediating nervous 1998). impulses arising from the depth of the brain. In those days such a projection system could not be

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THE CENTRAL REPRESENTATION OF turned off. Other types of complex behavioral MOTIVATIONAL AND INSTINCTIVE BE- responses elicited from different hypothalamic HAVIOR and septal regions concerned the species-speci- fic expression of hunger- and thirst-like pat- The most striking stimulation effect was at low- terns, locomotor and cleaning drives, as well as est threshold elicited from the perifornical re- characteristic urination and defecation behavior. gion of the cat hypothalamus: the typical reac- Hess discussed the question in detail as to tion normally presented to the appearance of a whether these experimental syndromes were dog. The animal’s extreme excitement was simply due to an electrically-induced release of shown by crouching, tail lashing, retraction of an unconscious executive automatism, or the ears, widening the pupils, piloerection at whether relationships to conscious motivational back and tail, hissing, spitting and growling; not behavior might be assumed. In his opinion, the infrequently there was urination; eventually the highly integrated behavioral responses display- defense-like behavior ended up with flight or ing a perfect coordination of somatomotor and well directed attack. The behavior was ex- autonomic changes and the animal’s manifest pressed so dramatically that the cat jumped vi- integration of environmental perception favored ciously toward the observers within reach and thesecondinterpretation(Hess,1949,1962).The clawed the towel held for protection (Fig. 6). corresponding stimulation points were carefully Thus, the behavior seemed typically goal-di- mapped but no correlation between the then rected and emotionally affected as if in a state of identified hypothalamic nuclei and the function- anxiety, rage or fear (Hess & Brügger, 1943). ally identified regions was found. Hess was not Very impressive was the crescendo of the ‘af- unhappy for – as already mentioned – he did not fective defense reaction’ as designated by Hess believe in ‘nuclear centers’. In the meantime, and the sudden break when the stimulator was highly promising anatomical as well as func-

Fig. 6. Affective defense reaction elicited by electrical low threshold stimulation of the perifornical region of the hypothalamus. Hissing, ear retraction. The cat is about to attack the experimenter. (W.R. Hess & M. Brügger, 1943, Fig. 1-3).

50 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

256 KONRAD AKERT tional imaging methods (e.g. nuclear magnetic with weak if any reactions to sensory stimuli resonance) had been elaborated for the detailed and lack of spontaneous movements. Basal met- topographical localization of brain functions. abolic rate and body temperature were found to In this context it is to be expected that Hess’s be diminished. This syndrome representing the hypothalamic regions of motivational and in- exact reverse to the extreme excitement of the stinctive behavior may soon be revisited. From affective defense reaction was called ‘adynamia’ the surroundings of the perifornical focal zone, by Hess (Hess, 1944). clear signs of general excitement, such as motor The ‘‘trophotropic zone’’ was defined more unrest, pupillodilation, and rapid respiration, anteriorly in the hypothalamus extending into and sometimes abortive defense movements, the septal region where electrical stimulation were observed. When stimulating the animal elicited effects that are normally attributed to the under deep anesthesia, Hess observed a prompt representation of the parasympathetic nervous rise in blood pressure and respiration in this re- system: fall of blood pressure as if it were gion (Hess, 1938). Subsequent investigators caused by the carotid sinus reflex, slowing of found a classical behavioral and EEG cortical respiration, pupilloconstriction, sometimes even arousal. It seemed therefore appropriate to des- cardioinhibition (Fig. 8). In the waking animal ignate the relevant region of the ventral poste- analogous changes in the visceral realm were rior hypothalamus as the ‘‘ergotropic zone’’ coupled with the dampening of general behavior (Fig. 7). The argument is strenghtened by the including acute muscular relaxation turning fact that bilateral, nearly symmetrical lesions eventually into a sleep-like condition. Lesions in within this zone gave rise to a marked loss of this zone as reported by other authors (see attention and tonus of the musculature combined Akert, 1959) gave rise to vital disabilities, e.g.,

Fig. 7. Sagittal section of the hypothalamus from Hess’s Atlas of the diencephalon (unpublished). Localisation of the ergotropic zone with pupillodilatation (circles) and locomotor unrest (up arrows). The tropho- tropic zone partly overlapping the thalamic sleep region represented by pupilloconstriction (squeezed circles) and decreased locomotion (down arrows). (R. Jung, 1983, Fig.4).

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Fig. 8. Blood pressure responses elicited from circumscribed hypothalamic regions. Above: Marked increase by stimulation of the ergotropic zone. Below: Decrease from the trophotropic zone. R = Stimulation. (W.R. Hess, 1950, Fig.7).

pulmonary edema, hyperthermia, rapidly fol- head-trunk movements of a totally different lowed by death. character (Fig. 9). Hess’ interest in the organization of the motor system dated back to his time as ophthalmolo- ORGANIZING PRINCIPLES OF THE MO- gist and his clinical studies on eye movements TOR SYSTEM (see Fig. 2). The motor system of the eye with three pairs of antagonistic muscles oriented in Hess never aimed at specific stimulation re- the three cardinal planes of space, linked with sponses but collected the data as they came in the analogous vectorial organization of the ves- by patiently and systematically positioning his tibular apparatus seemed to him an ideal para- electrodes in millimeter steps through the en- digm of the general biomotor control system. tire diencephalon. At the hypothalamic level he Likewise, in the diencephalon he found the rep- predominantly encountered autonomic effects resentation of spatio-temporal coordination of (see above) partly of instinctive character, head and body movements in the three cardinal whereas extrapyramidal motor responses pre- planes of space (Hess, 1941). Hess considered vailed in thalamus, subthalamus, and rostral the motor effects elicited from the diencephalon midbrain (Hess, 1941). And again, in contrast as fragments of an integrated motor system con- to cortical stimulation there were no synchro- trolling complex postural functions. In addition, nous (clonic) twitchings of isolated muscles or he emphasized the reciprocal relations between muscle groups but smooth, integrated eye- postural (‘ereismatic’ = supporting) and goal

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Fig. 9. Responses of the extrapyramidal motor system: eye, head, and body movements along horizontal and vertical axes elicited by electrical stimulation of circumscribed diencephalic regions representing the cardinal planes of space. (W.R. Hess, 1942, Figs.5 & 6).

directed (‘teleokinetic’) motor systems. Posture BIOLOGICAL ORDER AND PSYCHO- in his view, however, consisted of dynamic PHYSIOLOGY equilibria of forces not only stabilizing and sup- porting goal-directed movements but also pro- It seems of considerable interest that not only viding adequate start conditions (‘Bereitschaft’) Hess but also other eminent for teleokinesis (Hess, 1943; Jung 1983). In among his contemporaries were deeply con- other words, his ereismatic system included a cerned with the brain-mind problem (Eccles, proactive, anticipatory control as well. This con- 1966). Hess’s approach in his monograph of cept clearly surpassed classical views (Sherring- 1962 and especially in its second edition of 1968 ton) of postural reflexes and has been amply evolved directly from his experimental research: confirmed by recent advances (see Georgopou- 1) early work on the mutual relationships be- los, 1994). tween the activity of the autonomic nervous sys- To develop and understand the regulating tem and psychic functions in health and disease mechanisms in which spatial perception, vestib- (Hess, 1924/25); 2) motivational behavior elic- ular, proprioceptive, and intracerebral feedbacks ited by electrical impulses delivered to circum- intervene, he designed three-dimensional mod- scribed hypothalamic regions, like aggres- els in which the experimental and theoretical sive/defensive behavior (Hess, 1943); 3) his vectors of muscular forces could be demon- large-scale search for the diencephalic control of strated (Fig. 10). With the aid of these geometri- motor behavior (Hess, 1942). cal models (Hess, 1942) it was possible to base Psychic functions were to him an essential analysis and interpretation of experimental data aspect of the living organism, especially in on a more solid and useful foundation and de- higher forms, whereby conscious experience velop a comprehensive precybernetic concept of seemed to represent the supreme and indispens- biomotor systems. able level of behavioral integration. Insofar as willful psychic activities such as conscious thinking and willful impulses were concerned, Hess envisaged a special, i.e., unique force of unknown nature, equally unknown being its transformation into a biological form of force. The only common denominator so to speak

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Fig. 10. Model of the central organisation of the biomotor system derived from clinical (see Fig. 2) and experi- mental studies of eye movements. Peripheral and central feedback systems according to Hess’ precybernetic thinking. B = Blickfeld (field of gaze), M = Muskelkerne (oculomotor nuclei), Starlets = site of convergence of multimodal afferents, + = representation of eye position, x = stimulus position in the visual brain area (W.R. Hess, 1941, Fig.7).

seemed to him the order that prevails on both However, as already indicated, the nature of sides. subjective experience according to Hess is not Order itself according to Hess is not a force, known, not causally intelligible. How nature has but is manifest in biological and psychic forces. managed to transform objective organic events, It may be compared with information, which is coded by neural information into subjective psy- not bound on mass and energy like the various chic experiences, such as feelings, thoughts, and carriers of information, e.g., mechanical or elec- willful impulses, may possibly never be under- tromagnetic forces. Psychic forces create psy- stood. chic patterns. According to Hess, the order of One of his last papers (Hess, 1967) deals with psychic patterns corresponds directly with the the problem of causal thinking as a drivelike order of nervous patterns which form the or- compulsion and its relation to the inborn ‘‘An- ganic background of behavior. lage’’ of the . In view of its para- Take as example the formulation of a word – mount importance for the understanding of hu- imagined and expressed: verbal communication man behavior, Hess emphasized the necessity of between individuals relies on the correspon- incorporating the biological aspects of psychic dence between orders of psychic and organic functions into the domain of the physiological patterns. sciences.

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HESS AS ACADEMIC PERSON AND MEN- ments of editor and reviewers should be acknowled- TOR ged.

Of course, he belonged to that dignified genera- tion of pre-worldwar ‘Herr Professors’, yet dis- REFERENCES tinctly more accessible to students than the 19th century German ‘Geheimrats’. I remember his Selected scientific papers and monographs of W.R. Hess lectures with mixed feelings. There were unfor- gettable lectures (‘Sternstunden’) though rather Hess WR (1903): Eine mechanisch bedingte Ge- too often alternating with painful hours where he setzmässigkeit im Bau des Blutgefässystems. Wil- seemed virtually absent minded and obviously helm Roux’Arch Entwickl Mech Org 16: 632-641. longing to return as fast as possible to the Hess WR (1907): Ein neuer Apparat zur Bestimmung der Viskosität des Blutes. Münch Med Wochenschr laboratory. Unforgettable were his daily rounds 54: 1590-1591. through the Physiology Institute. The length of Hess WR (1908): Eine neue Untersuchungsmethode time spent with each of us was an indicator of bei Doppelbildern. Arch Augenheilkd 62: 233-238. one’s standing with the chief. He was most Hess WR (1914): Das Prinzip des kleinsten Kraftver- feared for his sharp and sometimes merciless brauches im Dienste haemodynamischer For- schung. Arch Anat Physiol Physiol Abt (Lpz.) criticisms (especially toward female students), 1914: 1-62. but again some of the intimate discussions with Hess WR (1924/1925): Über die Wechselbeziehungen him turned out to be most fascinating and mem- zwischen psychischen und vegetativen Funktionen. orable. He would then open his inquiring mind Schweiz Arch Neurol Psychiat 15: 260-277, 16: 36- for fundamental problems to the junior listener. 55, 285-306. Hess WR (1927): Stammganglien-Reizversuche. Ber Hess as an experimenter was extremely con- ges Physiol exp Pharmakol 42: 554-555. cerned with technical details. He never failed to Hess WR (1929): The Alpine Research Station of be present during each brain stimulation session Switzerland. J.State Med. 37: 11-14. and called for utmost discipline during the pro- Hess WR (1929): The mechanism of sleep. Amer J cedure. One of his frequent complaints was his Physiol 90: 386-387. Hess WR (1930): Die Regulierung des Blutkreislaufs, ‘endless war with his pen’ as he used to say; and gleichzeitig ein Beitrag zur Physiologie des vege- indeed his writings – even after numerous and tativen Nervensystems. Leipzig, Thieme, pp. 1- painful editings – were often unusually heavy 162. and involved: quite cumbersome for the reader. Hess WR (1931): Die Regulierung der Atmung, After retirement he became a relaxed and gleichzeitig ein Beitrag zur Phyiologie des vege- tativen Nervensystems. Leipzig, Thieme, pp. 1- charming old gentleman full of new and old 137. ideas about nature and fundamental forces Hess WR (1932): Beiträge zur Physiologie des Hirn- within the universe and with a lively interest in stammes. I. Die Methodik der lokalisierten Reizung the progress of brain research, which he dis- und Ausschaltung subkortikaler Hirnabschnitte. cussed intensively with his frequent visitors. Leipzig, Thieme, pp. 1-122. Hess WR (1933): Der Schlaf. Klin Wochenschr 12: 129-134. Hess WR (1938): Beiträge zur Physiologie des ACKNOWLEDGMENTS Hirnstammes. II. Das Zwischenhirn und die Regu- lation von Kreislauf und Atmung. Leipzig, Thieme, pp. 1-127. Special thanks are due Prof. Hans-Rudolf Lüscher, Hess WR (1941): Charakter der im Zwischenhirn Physiology Department, University of Berne, Prof. ausgelösten Bewegungseffekte. Pflügers Arch Ges Alexander Borbély, Department of Pharmacology and Physiol 244: 767-786. Dr. Caroline Jagella, Department of the History of Hess WR (1941): Die Motorik als Organisations- Medicine, University of Zürich for helpful comments. problem. Biol Zbl 61: 545-572. I am particularly grateful for the hospitality and the Hess WR (1942) Biomotorik als Organisations- facilities offered by my colleagues of the Brain Re- problem. Naturwissenschaften 30: 441-448, 537- search Institute and to Roland Schoeb for preparing 541. the illustrations. Last but not least, the helpful com-

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Hess WR (1943): Teleokinetisches und ereismatisches Fritsch G, Hitzig E (1870): Über die elektrische Kräftesystem in der Biometrik. Helv Physiol Phar- Erregbarkeit des Grosshirns. Arch Anat Physiol makol Acta 1: C 62-63. Wiss Med (Lpz) 37: 300-332. Hess WR, Brügger M (1943): Das subkortikale Zen- Fulton JF (1939) Levels of autonomic functions with trum der affektiven Abwehr. Helv Physiol particular reference to the cerebral cortex. Res Pharmakol Acta 1: 33-52. Publ Nerv Ment Dis 19: 219-236. Hess WR (1944): Hypothalamische Adynamie. Helv Hess RM jr, Akert K, Koella W (1950): Les potentiels Physiol Acta 2: 137-147. bioéléctriques du cortex et du thalamus et leur Hess WR (1948): Die funktionelle Organisation des altération par stimulation du centre hypnique chez vegetativen Nervensystems. Basel, Schwabe, pp. 1- le chat. Rev Neurol 83: 537-544. 226. Hunsperger RW, Bucher VM (1967): Affective Hess WR (1949): Das Zwischenhirn. Syndrome, behaviour produced by electric stimulation in the Lokalisationen, Funktionen. Basel, Schwabe, pp. forebrain and brain stem of the cat. Progr Brain 1-187. 2. erw. Aufl. 1954,p.p. 1-218. Res 27: 103-127. Hess WR (1950): Funktion und nervöse Regulation Hunsperger RW, Leisinger-Trigona M.-C. (1972): der innern Organe. Vierteljahrschr Naturforsch Further studies on the alleged hypnogenic zone in Ges Zürich 95: 249-264. the thalamus. Perspectives in Brain Sciences 1: Hess WR (1954): Diencephalon. Autonomic and ex- 100-108. trapyramidal functions. New York, Grune & Strat- Jasper HH (1949): Diffuse projection systems. EEG ton, pp. 1-79. Clin Neurophysiol 1: 405-420. Hess WR (1956): Hypothalamus and Thalamus. Doc- Jouvet M (1961): Telencephalic and rhombencephalic umentary pictures. – Hypothalamus und Thalamus. sleep in the cat. In: Wolstenholme GEW, Experimental-Dokumente. Atlas with German and O’Connor M, eds., CIBA Foundation Symposium English legends. Stuttgart, Thieme, pp. 1-70. 2nd on the nature of sleep. Boston, Little Brown, pp. ed. 1969, pp. 1-77. 188-208. Hess WR (1962): Psychologie in biologischer Sicht. Karplus JP, Kreidl A (1910): Gehirn und Sympa- Stuttgart, Thieme, p.p. 1-120. 2. erw. Aufl. 1968, thicus. II. Ein Sympathicuszentrum im Zwischen- pp. 1-132. hirn. Pflügers Arch ges Physiol 135: 401-416. Hess WR (1963): From medical practice to theoretical Langley JN (1893): Preliminary account of the ar- medicine. An autobiographic sketch. Perspect Biol rangement of the autonomic nervous system based Med 6: 400-423. chiefly on observations upon pilomotor nerves. Hess WR (1964): The Biology of Mind. Chicago, Uni- Proc Roy Soc 52: 547-556. versity of Chicago Press, pp. 1-203. Magoun HW (1952): The ascending reticular activat- Hess WR (1967): Causality, Consciousness and Cere- ing system. Ass Research Neurol Ment Dis 30: bral Organization. Science 158: 1297-1283. 480-492. McCormick DA, Bal T (1997): Sleep and Arousal: References to authors quoted in this essay. Thalamocortical mechanisms. Ann Rev. Neurosci 20: 185-215. Akert K (1959): Physiology and Pathophysiology. In: Minkowski M (1943): Zur Kenntnis des Verlaufs, der Schaltenbrand G, Bailey P eds., Introduction to Verbindungen und der extrapyramidalen Bahnen, Stereotaxis with an Atlas of the Human Brain, besonders in ihren Beziehungen zum Vestibula- Vol.I, Stuttgart, Thieme, pp. 152-229. rapparat, im Zusammenhang mit physiologischen Bard P (1928): A diencephalic mechanism for the ex- Versuchen von W.R. Hess. Schweiz Arch Neurol pression of rage with special reference to the sym- Psychiat 51: 99-138. pathetic nervous system. Amer J Physiol 84: 490- Moruzzi G, Magoun HH (1949): Brain stem reticular 513. formation and activation of the EEG. Electro- Bichat MFX (1800): Recherches physiologiques sur la enceph Clin Neurophysiol 1: 455-473. vie et la mort. Paris, Brossson & Gabon, p.p.1-321. Moruzzi G (1963): Active processes in the brain stem Borbély AA, Tononi G (1998): The quest for the es- during sleep. Harvey Lectures 58: 233-297. sence of sleep. Daedalus 127: 161-196. Ranson SW, Magoun HW (1939): The Hypothalamus. Cannon WB (1915): Bodily changes in pain, hunger, Ergeb. Physiol. 41: 56-163. fear and rage. New York, Appleton, 2nd ed (1929) Reil JC (1807): Über die Eigenschaften des Ganglien- pp. 1-404. Systems und sein Verhältnis zum Cerebralen-Sys- Eccles JC (1966): Brain and Conscious Experience. tem. Reil’s Arch f Physiol 7: 189-254. Study week September 28 to October 4, 1964, of Weissschedel E, Jung R (1939): Die anatomische the Pontificia Academia Scientiarum. Eccles JC, Auswertung und das Studium der sekundären ed., Berlin-Heidelberg-New Work, Springer, pp. 1- Faserdegeneration nach lokalisierter subcorticaler 591. Ausschaltung durch Elektrokoagulation. Z Anat Entwickl gesch 109: 374-395.

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Biographical writings on W.R. Hess and his work Caspers H, Hess R, Kugler J, Petsche O, Struppler A (1974): Nachruf auf W.R. Hess. EEG EMG 5: 139- Akert K (1974): Walter Rudolf Hess (1881-1973). 141. Obituary. Brain Research 68: 5-8. Jung R (1981): Walter R. Hess (1881-1973). Rev Akert K (1981): Biological Order and Brain Organi- Physiol Biochem Pharmacol 88: 1-21. zation. English translations of selected works. Ber- Jung R (1983): W.R. Hess und das Ordnungsprinzip lin, Heidelberg, New York, Springer, pp. 1-147. in der Physiologie. Schweiz Arch Neurol Psychiat Auerswald W (1975): Walter Rudolf Hess. Nachruf 132: 277-308. mit Schriftenverzeichnis. Almanach Oesterr Akad Wyss von W (1948): 50 Jahre Psychophysiologie in Wiss 124: 414-433. Zürich. A. Forel. E. Bleuler. C. von Monakow. W.R. Hess. 111. Neujahrsblatt zum Besten des Waisenhauses in Zürich. Beer, Zürich, pp. 1-69.

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Walter Rudolf Hess: aus seinem Leben und seiner WalterTätigkeitRudolf an derHess Medizinischen Fakultät der Universität Zürich (100-Jahr-Feier seines Geburtstages: 14. März 1981) Has seinem Le&en und seiner Tätigkeit an der Medizinischen Tafeu/tät derWalterPeterUniversität G. WaserRudolfZürichHess f 200-J'^ahr-h'eier seines Geburtstages: id. März 1981 J HasWürdigungseinem derLe&en akademischenund seiner TätigkeitTätigkeit von W.an R.der HessMedizinischen durch den ehemaligenTafeu/tät Direktor derdes PharmakologischenUniversität Zürich Institutes der Universität Zürich, veröffentlicht in der Zeitschrift Von Peter G. Waser fGesnerus200-J'^ahr-h'eier 39 (1982), seinespp. 279-286.Geburtstages: id. März 1981 J

EsVonistPeterimmerG. Waserschwierig, über einen bedeutenden Mann zu berichten, auch wenn man einige Jahrzehnte mit ihm zusammen gelebt und gelegentlich auch gearbeitet hat. Wenn man wie ich zuerst sein Student war, später AssistentEs ist immerin einemschwierig,nahenüberForschungsinstituteinen bedeutendenundManndannzu berichten,Kollege inauchder Fakultät,wenn manisteinigees besondersJahrzehnteschwierig,mit ihmkritischzusammenzu bleibengelebtmitundallergelegentlichBewunde- rungauch fürgearbeitetdie außerordentlichehat. Wenn manwissenschaftlichewie ich zuerstLeistung,sein Studentfür diewar,menschli-später cheAssistentGröße undin einemdie kollegialenahen ForschungsinstitutVerbundenheit, die erundeinemdannengenKollegeKreisinseinerder MitarbeiterFakultät, istundes besondersjungen schwierig,Freunde entgegenbrachte.kritisch zu bleibenWasmit allerdavonBewunde-in der Erinnerungrung für die zurückbleibt,außerordentlicheist wenigerwissenschaftlichedas Sachliche,Leistung,die genauenfür dieDatenmenschli-des Lebens,che Größesondernund diederkollegialeganz persönlicheVerbundenheit,menschlichedie erAspekteinem engeneiner Begegnung,Kreis seiner dieMitarbeiterSpuren hinterläßt,und jungenEindrückeFreundemachtentgegenbrachte.und AnregungenWasvermittelt.davon inSoderist meinErinnerungBerichtzurückbleibt,sicher lückenhaft,ist wenigereinseitigdas Sachliche,und persönlichdie genauen— aber indemDaten wirdes einesLebens,unssondernprägendender ganzKollegenpersönlicheund Mitmenschenmenschlichegedenken,Aspekt einergeschiehtBegegnung,es aus einerdie Spurensubjektiven,hinterläßt,unvollständigenEindrücke machtSicht,unddieAnregungenzum Gesamtbildvermittelt.einesSoFor-ist schersmein Berichtund Menschensicher lückenhaft,gehört. einseitig und persönlich — aber indem wir einesZuerstuns prägendenkurz einigeKollegenbiograp/iisc/ieund MitmenschenDaten: W.R.Hessgedenken,wurdegeschiehtam 17.Märzes aus 1881einer insubjektiven,Frauenfeldunvollständigen(Kt.Thurgau) Sicht,als Sohndie eineszum GesamtbildMittelschullehrerseines For-für Physikschers undgeboren.MenschenEr verlebtegehört. dort eine glückliche Jugend in Naturverbun- denheit,Zuerstkamkurzbereitseinigedurchbiograp/iisc/ieseinen VaterDaten:in KontaktW.R.Hessmit wurdenaturwissenschaft-am 17.März liehen1881 inProblemen,Frauenfeldbesonders(Kt.Thurgau)in deralsPhysikSohn undeinesMeteorologie,Mittelschullehrersund mitfür einigenPhysik geboren.technischenEr verlebteBetriebendortdereineerwachendenglückliche JugendIndustrie.in Naturverbun-Nach dem Gymnasiumdenheit, kambegannbereits erdurchsein seinenMedizinstudiumVater in Kontaktin Lausannemit naturwissenschaft-in einem kulturell undliehensprachlichProblemen,französischbesondersorientiertenin der PhysikLandesteilundderMeteorologie,Schweiz. Dochundwarenmit aucheinigenanderetechnischenMedizinschulenBetriebenwie Bern,der erwachendenZürich und vorIndustrie.allem BerlinNachund Kieldem wichtigeGymnasiumAusbildungsstätten,begann er sein Medizinstudiumdie nicht nurinfachlich,Lausannesondernin einemvorkulturellallem kulturellund sprachlichauf denfranzösischjungen orientiertenMedizinstudentenLandesteilgroßender Schweiz.EindruckDochmachten.waren Nebenauch andereKonzert-Medizinschulenund Theatervorstellungenwie Bern, Zürichbesuchteund vorerallemauch dieBerlinMuseen.und KielVor allemwichtigedieAusbildungsstätten,Meeresaquarien unddieeinnichtastronomischesnur fachlich,Institutsondernmitvor einemallem kulturell auf den jungen Medizinstudenten großen Eindruck machten. GesnerusNeben Konzert-2 (1982) und Theatervorstellungen besuchte er auch die Museen. Vor279 allem die Meeresaquarien und ein astronomisches Institut mit einem

Gesnerus 2 (1982) 279

59 Walter Rudolf Hess: aus seinem Leben und seiner Tätigkeit an der Medizinischen Fakultät der Universität Zürich

Riesenfernrohr hinterließen einen tiefen Eindruck. Hess schreibt selbst: «Es beschäftigt mich die Frage der Unendlichkeit, das, was sich hinter den sichtbaren Sternen befindet!» Also beschäftigte ihn schon von Anfang an die Frage, was hinter den Objekten steht, nicht nur das, was in der Natur einfach beobachtet wird, sondern die Zusammenhänge dahinter oder darüber hinaus, die Kraft und die Organisation, welche die äußere Welt zusammen- halten und bewegen. Dazu ein Beispiel: Im Präpariersaal des Zürcher Anatomischen Institutes wurde eine arterielle Anomalie demonstriert, die Hess als Student dazu bewegte sich zu fragen, wie die Natur eine Optimie- rung der Gewebedurchblutung erreiche. Wenn sich eine Arterie in zwei Äste aufspaltet, muß bei konstantem Gesamtvolumen der Hauptarterie und ihrer beiden Äste der Gesamtwiderstand ein Minimum werden. Das einfache mathematische Problem wurde zusammen mit dem Zürcher Mathematiker Prof. Rudolf Fueter gelöst und experimentell bestätigt. Wir mußten später als Studenten der Naturwissenschaften dieses Problem in der Vorlesung über höhere Mathematik bearbeiten, und es ist mir deshalb durch seine Klarheit und Logik in Erinnerung geblieben. Der ehemalige geniale Gerichtsmediziner unserer Fakultät Prof. Hein- rieh Zangger beurteilte die kurze Abhandlung: «Eine mechanisch bedingte Gesetzmäßigkeit im Bau des Blutgefäßsystems» sehr positiv — und förderte wohl damit die wissenschaftlichen Interessen des jungen Studenten. Nach einigen klinischen Auslandsemestern, wie sie damals noch üblich — und möglich! — waren, legte Hess ein Staatsexamen für Medizin in Zürich 1905 ohne Schwierigkeiten ab. Leider konnte Hess aus finanziellen Gründen nicht sofort als Assistent am Physiologischen Institut tätig sein, wie er sich das wünschte. Damals reichte eine Assistentenbesoldung für Wissenschafter auch bei einfachster Lebensführung nicht aus, und eigene Mittel standen nach dem Studium nicht zur Verfügung. So begann er seine Ausbildung im Allgemeinen Krankenhaus in Münsterlingen am Bodensee. Doch war der chirurgisch orientierte Chef (Konrad Brunner) sehr streng, die Freizeit knapp. Sie reichte daher nur zur Ausarbeitung einer selbstgewählten Dissertation über: «Viscosität des Blutes und Herzarbeit» (1906). Das damals entwickelte Viscosimeter (eine relative Bestimmungsmethode zum Vergleich der Visko- sitäten von Blut und Wasser) benutzten wir alle als Medizinstudenten im Physiologie-Kurs während Jahrzehnten! Die eigentliche Ausbildung er- folgte nun in der Zürcher Augenklinik (Prof. Otto Haab) zum Facharzt für Ophthalmologie, wobei sich Hess wieder für ein physiologisches Problem,

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Walter Rudolf Hess 1881-1973 die Koordination der Bewegung der Augen und die Diagnostik der Augen- muskellähmungen, interessierte und ein entsprechendes Gerät zur genauen Messung dieser Funktion nach einem neuen Prinzip konstruierte. Nach einer weiteren praktischen Ausbildung in Paris in Neurologie, Venerologie und innerer Medizin eröffnete Hess 1908 eine Praxis für

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61 Walter Rudolf Hess: aus seinem Leben und seiner Tätigkeit an der Medizinischen Fakultät der Universität Zürich

Augenheilkunde in Rapperswil. Das gute Einkommen ermöglichte ihm endlich die Heirat. Aber alle praktischen Tätigkeiten, auch die nebenamtli- che Betreuung einer klinischen Abteilung für Augenkranke im Kantonsspi- tal Glarus, befriedigten ihn nicht, so daß er im Herbst 1912 nach Zürich zurückkehrte und Assistent beim Physiologen Prof. Justus Gaule, einem Schüler Carl Ludwigs, wurde. Der sich zunehmend verschlimmernde psychi- sehe Zustand seines Chefs brachte rasch eine steigende Belastung im Unterricht und im Kurs. Dazu reichte das Assistentengehalt für die junge Familie, inzwischen mit einer Tochter und einem Sohn, nur für eine äußerst sparsame Lebensführung. Eine Erfindung zur stereoskopischen Betrach- tung photographischer Bilder in räumlicher Dimension sollte finanzielle Hilfe bringen. Doch dann kam 1914 der Weltkrieg und verlagerte die medizinische Tätigkeit des Sanitäts-Hauptmannes Hess in die Armee, ohne die Möglich- keit, neben den gelegentlichen Urlauben für den Medizinerunterricht noch Forschung zu betreiben. Ein ganzjähriger Auslandsurlaub (1915/16) in Bonn beim Physiologen Prof. Max Yerworn erlaubte trotzdem eine wesentli- che Weiterbildung und Forschungsarbeit im Kreislaufgebiet über die Regulierung des Blutdruckes und Probleme der Haemodynamik, die später in Zürich fortgesetzt wurde. Die Habilitation (1913) hatte schon als Thema die physikalisch-biologische Basis der Zusammenhänge verschiedener Fak- toren für die Kreislaufregulierung enthalten! Der damalige Internist war anscheinend bei der Beurteilung der Habilitationsarbeit über die mathema- tischen Formulierungen «als der Physiologie ferne», wie er sagte, Unglück- lieh, wahrscheinlich weil die mathematische Interpretation biologischer Vorgänge für den praktischen Mediziner neu war. Die Entscheidung über die Zukunft von Hess fiel im Wintersemester 1916/17 durch den unerwarteten Rücktritt von Prof. Gaule. Zuerst las Hess die ganze Physiologie-Vorlesung vertretungsweise, als Lückenbüßer, wie er selbst sagte, lehnte aber ein Extraordinariat unter einem neuen Chef, der aus Deutschland berufen worden wäre, ab. Im September 1917 wurde er dann durch die Fakultät zögernd, aber durch den Erziehungs- und Regierungsrat einstimmig, zum Nachfolger und Ordinarius gewählt. Bestimmend war damals wohl der Erziehungsdirektor, Regierungsrat Mousson, der nicht nur die Studenten über die Vorlesung befragte, sondern einmal selbst die sehr lebendige Vorlesung über Experimentalphysiologie des Privatdozenten Hess besuchte und davon sehr beeindruckt war. Die Antrittsrede vom 15. Juni 1918 befaßte sich mit der «Zweckmäßigkeit im Blutkreislauf».

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62 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

Die Le/irféitig/ceû möchte ich aus eigener Erfahrung beurteilen. Meine Generation begann das Medizinstudium vor dem Zweiten Weltkrieg und kam während der Aktivdienstjahre in die klinische Ausbildung. Wir waren gerade während dieser Übergangszeit drei Semester lang mit Physiologie und Anatomie beschäftigt. Die Vorlesung war für uns anregend, lehrreich, systematisch. Sie wurde begleitet von zahlreichen Experimenten und Modellen, die uns das funktionelle Äquivalent der detaillierten Vorlesung über die Anatomie des menschlichen Körpers gaben. In dieser Hinsicht waren die Stunden im alten Hörsaal des Physiologischen Institutes für mich ein weiter Ausblick in die eigentlichen Lebensvorgänge, die ich später in der Klinik in einer ganz anderen Perspektive, mit Störungen oder Abnormitä- ten, kennenlernen sollte. Neu, und für viele auch ein vergnügliches Schau- spiel, waren die Filme ohne Ende, bei denen der gleiche funktionelle Ablauf einer Magen-Darm-Bewegung oder der Hcrz-Kreislauf-Tätigkcit sich im- mer wiederholte. Da das Endstück des 9-mm-Schmalfilms an den Anfang der Sequenz angeklebt war — eine Schlaufe ohne Ende —, wiederholte sich der gleiche Vorgang stereotyp, natürlich von Hess mit lebhaftem Kommentar begleitet, bis wir die Sache ganz kapiert hatten. Hess war ein didaktisch begabter Lehrer, der es verstand, uns auch schwierige Dinge in anschaulicher Weise verständlich zu machen und uns zu eigenem selbständigen Denken und Experimentieren anleitete. Wenn dann aber die Filme der Katzenversuche gezeigt wurden, bei denen eine elektri- sehe Hirnreizung immer wieder die gleiche Wirkung, z.B. Erbrechen oder Defäkation oder Schlaf, dann wieder Erregung hervorrief, wurde es still im Auditorium. Denn mancher überlegte sich, was nun wirklich Entscheidung und freier Wille sei, wenn ein Druck auf eine Taste über eine Hirnreizung ein Tier zu einem bestimmten, immer wieder gleichartigen Verhalten bringt, ohne daß das Tier diese Tätigkeit eigentlich ausführen will. Diese Experi- mente waren überzeugend, und wir jungen Medizinstudenten ahnten, welche großartigen Probleme unser Professor in Bearbeitung hatte, um die funktionellen Zentren des Gehirns zu verstehen. Auch der Experimentierkurs war für uns eine wertvolle Instruktion. Er war gut organisiert, von Assistenten überwacht und hatte zahlreiche interessante Arbeitsplätze aus den verschiedenen Gebieten der Physiologie. Es wurden dabei nicht immer leichte Aufgaben gestellt. «Papa Hess», wie wir ihn unter uns nannten, mit Baskenmütze auf dem weißen Haupt von Platz zu Platz schreitend, befragte uns, demonstrierte, diskutierte oft über Themen, die weit entfernt vom Fachgebiet lagen, aber von großem menschli-

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63 Walter Rudolf Hess: aus seinem Leben und seiner Tätigkeit an der Medizinischen Fakultät der Universität Zürich

c.hem Interesse waren: über Ethik, Leben, Krankheit und Tod, oder über die Notwendigkeit von Tierversuchen in der Medizin und in den biologischen Wissenschaften und über viel anderes mehr. Das Thema über Tierversuche wurde oft angesprochen, weil Hess fortwährend von den Yivisektionsgeg- nern verfolgt wurde und Mühe hatte, ihre gehässigen Angriffe abzuwehren. Diese Wirrköpfe hätten seine Forschungsarbeit verunmöglicht, wenn sie dazu den richtigen demagogischen Einfluß auf die Politiker gefunden hätten. Nicht zu vergessen ist, daß Hess schon 1922—24 Dekan der Medizinisc/ien Fakultät der Universität Zürich war und später auch wesentlichen Anteil an deren Entwicklung zur heutigen Leistungsfähigkeit auf hohem Niveau hatte. Während der ersten 15 Jahre war in seiner Lehrverpflichtung auch die Physiologische Chemie eingechlossen. Erst 1932 wurde ein separater Lehr- stuhl für Pysiologische Chemie errichtet. Zuerst war er mit einer ungeeigne- ten Persönlichkeit besetzt, aber später wurde er unter Prof. Franz Leuthart als Biochemie zum neuen Grundpfeiler der Medizin und Biologie ausgebaut. Die Gründung des hochalpinen Forschungsinstitutes Jungfraujoch wurde durch Hess als Präsident einer Kommission der Schweizerischen Naturforschenden Gesellschaft gefördert. Auch war Hess Mitbegründer der Schweizerischen Physiologischen Gesellschaft und deren wissenschaftlichen Zeitschrift. Er organisierte zudem 1938 als Tagungspräsident den Interna- tionalen Kongreß für Physiologie, Physiologische Chemie und Pharmakolo- gie in Zürich. Hess pflegte einen regen Kontakt mit vielen Wissenschaftern. Er war auch oft in Zürich mit Paul Karrer und Leopold Ruscizka zusammen. Die Forsckang hat Hess während seiner ganzen akademischen Tä tigkeit voll in Anspruch genommen. Neben der anstrengenden Lehrtä tigkeit war sie nur möglich durch gute Planung in einem speziellen, begrenzten Wissensge- biet und mit dem vollen Einsatz aller Kräfte. Bei dem begrenzten Personal- und Finanzetat der damaligen Zeit konnte dies nur durch Bildung eines über lange Zeit zusammenarbeitenden Teams, aus meistens nicht studierten Technikern, ermöglicht werden, die über Jahrzehnte beisammen blieben und die neue Forschungstechnik weiter ausbauten (Frl. Verena Bucher, Herr Max Jenny, Frl. Anna Jaussi und Frau Mina Eugster). Dazu kamen während längerer Zeitperioden jüngere Forscher wie Ernst Rothlin und Alfred Fleisch, später dann Oskar Wyss, Marcel Monnier, Werner Koella, Konrad Akert und Robert Hunsperger als Oberassistenten. Alle wurden Professoren für Physiologie in Zürich, Basel oder an anderen Universitäten, zwei von ihnen Nachfolger auf dem Lehrstuhl und Leiter des Physiologi-

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64 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich sehen Institutes in Zürich. Eine große Zahl von Assistenten und Doktoran- den nahm nur während kürzerer Zeit an einem wissenschaftlichen Problem teil. Ihre Beiträge waren wichtige Steine im großen Gebäude des For- schungsplanes, der während Jahrzehnten konsequent und inlensiv verfolgt wurde. So war auch eine maximale Effizienz der eingesetzten Arbeitskräfte und der damals, vor der Gründung des Schweizerischen Nationalfonds, noch beschränkten staatlichen Forschungsmittel möglich. Neben kleineren Stif- tungen und einer Unterstützung durch die Industrie, war es vor allem die Rockefeller Foundation, die wiederholt finanziell mithalf. Zahlreiche For- schungsurlaube ermöglichten es Hess, seine Resultate in Monographien zusammenzufassen. Das Forschungsinteresse war vor 1930 auf den Blutdruck, die Haemody- namik und vor allem auf die Regulation von Kreislauf und Atmung konzentriert. Es wechselte dann, entsprechend der Verfolgung der zentralen Steuermechanismen, und konzentrierte sich nach Beschreibung der funktio- nellen Organisation des vegetativen Nervensystems auf das Zwischenhirn. Dieses zentrale, subcorticale Gebiet zur Regulation der Tätigkeit der inneren Organe wurde mit der hauseigenen Technik während über 20 Jahren unermüdlich erforscht. Durch Reizung und Ausschaltung diskreter, kleiner Hirngebiete, ihre genaue histologische Fokalisation und Eintragung auf Karten der Hirnquerschnitte wurde der Zusammenhang mit den beobachte- ten peripheren Wirkungen am Tier festgestellt und damit die funktionelle Organisation dieses Hirnteiles, des Diencephalon, aufgeklärt. Diese müh- same Analyse eines dichten, neuronalen Hirngebietes und die Methode waren so erfolgreich, daß später das ganze Gehirn von anderen Forschungs- gruppen bei verschiedenen Tierspezies untersucht und in verschiedenen Ebenen kartographiert wurde mit allen Angaben der Reizpunkte und der eintretenden Wirkungen. Auf dieser Grundlage beschäftigen sich heute die Neurochemiker mit den vielfältigen Formen chemischer Impulsübertra- gung durch Neurotransmittoren und Neurohormone, die Neurophysiologen mit den komplizierten neuronalen Verbindungen, dem Schaltplan der Neurone, und die Molekularbiologen mit den Vorgängen, welche das Lernen und die Gedächtnisbildung ermöglichen. Das Verständnis für das Wesen der Denkvorgänge auf molekularer Ebene ist dadurch in Sichtweite gekommen, zum Teil auch auf der Grundlage der Hessschen Forschungsresultate. Hess hat für seine Leistungen auch zahlreiche Ehrungen erhalten: Ehrendokto- rate, Mitgliedschaft in Akademien, nationale und internationale Preise und insbesondere 1949 den Nobelpreis für Physiologie und Medizin.

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65 Walter Rudolf Hess: aus seinem Leben und seiner Tätigkeit an der Medizinischen Fakultät der Universität Zürich

Nach der Emeritierung (1951) bearbeitete Hess weiter sein immenses Forschungsmaterial und publizierte die Zusammenfassung der Resultate in verschiedenen Monographien. Sein Interesse wendete sich nun noch höheren Leistungen des Gehirns zu: der Psyche! So verfaßte er ein bemerkenswertes Buch über «Psychologie in biologischer Sicht» (1962), das bald in verschie- dene Weltsprachen übersetzt wurde. Wir durften als junge Forscher bei der zweiten, erweiterten Auflage (1968) in einem interdisziplinären Gespräch über ausgewählte Themen mitarbeiten. Es war für uns natürlich äußerst interessant, anregend, ehrenhaft, aber auch amüsant, mit unserem verehr- ten und geliebten Lehrer nochmals diesen letzten Gedankenaustausch zu haben. So wurden viele von uns ausgebildet und geprägt durch die großar- tige Forschung, aber auch erzogen und gefördert durch den persönlichen Kontakt mit diesem selbstkritischen, exakt denkenden und logisch handeln- den Mann. Den tiefsten Eindruck machen uns heute wohl kaum die zahlreichen Ehrungen, die ihm zuteil wurden, sondern die Tatsache, daß LIess als «Seifmade man» sich ohne Lehrer oder Schule zum Forscher entwickelte und seinen Weg zum Erfolg ohne äußere Hilfe mit einer unglaublichen Energie und Selbstdisziplin allein ging. Dadurch bleibt er für uns ein großes Vorbild.

Ein vollständiges Verzeichnis der wissenschaftlichen Arbeiten von W. R. Hess ist im Buch «Biological Order and Brain Organization, Selected Works of W.R.Hess», edited by K. Akert, Springer Verlag, 1981, enthalten.

Summary

The life and scientific career of Walter Rudolf Hess is described from a personal viewpoint of a former student and later colleague of the medical faculty of the University of Ziirich. His selfeducation, his unlimited energy and discipline in hard research work led him to pioneering new techniques and knowledge of brain functions and behaviour. He was always held in the highest esteem throughout his life by his faithful and successful pupils in different fields of biological and medical research and officially honoured by the endowment of the Nobelprize in Physiology and Medicine in 1949.

Prof. Dr. med. et phil. II Peter G. Waser Pharmakologisches Institut der Universität Zürich Gloriastraße 32 CH-8006 Ziirich

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66 Walter R. Hess (1881–1973)

Richard Jung

Würdigung der Leistungen und Beiträge von W. R. Hess durch den prominenten Neuro- logen Professor Richard Jung, Universität Freiburg i. Br., veröffentlicht in Rev. Physiol. Biochem. Pharmacol., Vot. 88 (1981)

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3. Der Nobelpreis für Walter Rudolf Hess

89

Nominationen von Walter Rudolf Hess für den Nobelpreis

Walter Rudolf Hess wurde seit 1933 bis zur Nominationen (chronologisch 1933-1949): Verleihung des Nobelpreises an ihn im Jahr • Physiology or Medicine 1933, by F Hartmann 1949 insgesamt 31 Mal von Kollegen aus • Physiology or Medicine 1935, by L Bouman dem In- und Ausland dem Nobelpreis-Ko- • Physiology or Medicine 1939 mitee empfohlen bzw. nominiert. An der by Alexandre Besredka Nomination beteiligten sich auch 9 seiner • Physiology or Medicine 1939, by H Zangger Kollegen aus der Zürcher Medizinischen • Physiology or Medicine 1939, by A Vandevelde Fakultät, nämlich der Gerichtsmediziner • Physiology or Medicine 1939, by E Anderes Heinrich Zangger (1939, 1944), der Internist • Physiology or Medicine 1939, by W Löffler Wilhelm Löffler (1939, 1944), der Neurochi- • Physiology or Medicine 1939, by A Vogt rurg Hugo Krayenbühl (1949), der Psychia- • Physiology or Medicine 1942 ter Manfred Bleuler (1949), der Neurologe by Mieczyslaw Minkowski (1949), der Physio- • Physiology or Medicine 1943, by Emil Bürgi loge Oskar Wyss (1947), der Ophthalmolo- • Physiology or Medicine 1943 ge Alfred Vogt (1939), der Zahnmediziner by Corneille Heymans Alfred Gysi (1949) und der Physiologe Emil • Physiology or Medicine 1944, by W Löffler Feer (1949), was auf die hohe Anerken- • Physiology or Medicine 1944, by H Zangger nung, die er von seinen Fakultätskollegen • Physiology or Medicine 1945, by J Nicod genoss, hinweist. • Physiology or Medicine 1945, by A Fleisch • Physiology or Medicine 1945, by L Michaud Es fällt auch auf, dass kein einziger Deut- • Physiology or Medicine 1945, by A Vannotti scher je Walter Rudolf Hess für den Nobel- • Physiology or Medicine 1945 preis nominiert hat, obwohl die deutsche by Corneille Heymans Physiologie seine wissenschaftliche Heimat • Physiology or Medicine 1946, by E Luescher und – neben der Schweiz – sein primärer • Physiology or Medicine 1947, by F Naville Wirkungsort war. Das ist wohl der belaste- • Physiology or Medicine 1947 ten Beziehung während der Nazi-Zeit ge- by Ferdinand Morel schuldet. Anlässlich des von Hess geleite- • Physiology or Medicine 1947, by E Frommel ten 16. Physiologenkongress 1938 in Zürich • Physiology or Medicine 1947, by Oscar Wyss wurden die Angelsachsen und Franzosen • Physiology or Medicine 1947 bevorzugt (Kommentar Christian W. Hess). by Georges de Morsier • Physiology or Medicine 1948, by I Abelin • Physiology or Medicine 1949, by E Feer • Physiology or Medicine 1949, by W Löffler • Physiology or Medicine 1949, by H Krayenbühl • Physiology or Medicine 1949, by M Bleuler • Physiology or Medicine 1949, by Alfred Gysi • Physiology or Medicine 1949, by M Minkowski

91

Ankündigung der Nobelpreisverleihung in der Zeitschrift NATURE

Bericht über die Verleihung des Nobelpreises für Physiologie und Medizin an Prof. W.R. Hess und Prof. A. E. Moniz in der Zeitschrift NATURE 4179: 947, 1949

NEWS and VIEWS (Quelle: No 4179, December 3, 1949 NATURE Publishing Group)

Nobel Prize for Physiology and stimulating current, so that the animals can Medicine for 1949 move freely. Motor effects of the stimula- THE Nobel Prize for Physiology and Medi- tion are recorded cinematographically and cine for 1949 has been awarded jointly to analysed with the help of the film record. Prof. Walter Rudolf Hess, of Zurich, and Prof. The brain tissue between the electrodes is Antonio Egas Moniz, of Lisabon. then destroyed by electrocoagulation. The resulting disturbances, together with the Prof. W. R. Hess effects of stimulation, allow conclusions to Prof. Hess is professor of physiology in the be drawn on the functions of the part of University of Zurich. Thanks to his exceptio- the brain concerned. The latter is identified nal ability, Hess, originally a school teacher, with certainty by making histological pre- obtained medical qualifications and adop- parations, plotted as an atlas of brain func- ted an academic career. His great energy tion. Last year Hess published a monograph and far-sightedness were responsible for which clearly outlines his conclusions on the the establishment of the High Altitude Re- relation of the diencephalon to motor acti- search Station of the Jungfraujoch. This pro- vity. By these systematic and important rese- ject met a long-felt need of scientific wor- arches, covering almost twenty-five years of kers and was realized in 1931 with the help tireless work, Prof. Hess has richly deserved of an international foundation of which the the present award. Royal Society is a member. The earliest of Hess’s physiological researches to attract wi- despread notice was his clear statement of Prof. A. E. Moniz the principles of autonomic nerve function. Prof. Moniz is a most distinguished neu- The medical world of the Continent owes to ro-surgeon whose international fame rests him the recognition of the “ergotropic”, po- on two contributions to medical practice wer-increasing action of sympathetic nerve especially. Between 1927 and 1937 he intro- impulses, and the energy-saving or “histo- duced and developed cerebral angiography tropic” action of parasympathetic stimula- as a method of diagnosis of intracranial di- tion promoting cell growth and repair. So sease. All the essentials of the method at early as 1925 he developed the technique present in occasional use in neuro-surgical which he has continued to use for investi- clinics are due to him. He published in 1936 gating the function of individual regions of a monograph on the operative treatment the brain. The method is to introduce need- of certain psychoses, which described the le electrodes into the brain of a cat and to operation of prefrontal leucotomy and the stimulate very localized areas through these impressive results of its application to twen- electrodes. The animals survive very well; ty cases of psychosis. Imagination, boldness special means are used for leading in the and skill were necessary to demonstrate,

93 Ankündigung der Nobelpreisverleihung in der Zeitschrift NATURE as he did, the feasibility of the injection of Although leucotomy is applicable in fewer radio-opaque substances into the internal cases and is more drastic than either insulin carotid artery, and of the operative destruc- comas or electroplexy, the two other inno- tion of the white matter of the prefrontal vations which have recently revolutionized lobes. He has described in his monograph the treatment of mental disorders, the work how he came to devise leucotomy. He had of Prof. Moniz has exerted at least as im- noted how little the extensive removals of portant an influence, because he directed brain tissue by Brickner and Clovis Vincent the attention of neuro-surgeons to psychia- had impaired social behaviour, and had con- tric problems. The procedure which he devi- sidered the possibility of lesser removals of sed may eventually be superseded; but the brain substance as a means of reversing the lesson which he has taught will not quickly rigid functioning of groups of brain cells, be forgotten, for, with the neuro-surgeons which he thought to be responsible for the who have followed him, he has played a persistence in psychosis of painful ideas. The large part in reviving in psychiatry a traditi- impetus to the first attempt came in London on of courageous and energetic treatment, in August 1935, when he heard Jacobsen’s and has demonstrated that skilful interven- description of his work with Fulton on the tion may yield a degree of success even in effects of frontal ablations on the so-called the most serious and advanced cases of psy- neurotic behaviour of chimpanzees. Assis- chosis. ted by Dr. Almeida Lima, he then perfected the technique of leucotomy within a few months.

94 Award Ceremony Speech

Herbert Olivecrona

Laudatio von Prof. H. Olivecrona, Neurochirurg am Karolinska Institut anlässlich der Zeremonie zur Ver- leihung des Nobelpreises an Prof. W.R. Hess und Prof. A.E. Moniz, From Nobel Lectures, Physiology or Medicine 1942-1962, Elsevier Publishing Company, Amsterdam, 1964

Presentation Speech by Professor H. Olive- Although we have long been aware of the crona, member of the Staff of Professors of function of the midbrain as a higher center the Royal Caroline Institute. of the autonomous nervous system, we have only now, through the research of Walter Royal Highnesses, Ladies and Gentlemen. Rudolf Hess, had the localization of these The Caroline Institute, through the presen- functions mapped out in detail. Through tation of this year‘s Nobel Prize, desires to the use of a refined and accurate technique reward two important discoveries in the he has succeeded in applying stimulation fields of neurophysiology and its clinical to or causing the destruction of very small application, neurology. Both of these disco- areas, and thus he has been able to study veries touch upon the connection between the effect of the stimulus, as well as of the function and localization in the brain. disappearance of a function. For his experi- ments he has used cats into which, under The primary centers of vital functions such anesthesia, a thin metal wire has been intro- as respiration, control of blood circulation, duced. The wire has been electrically insula- the secretory and motor activity of the di- ted along its entire length except at the very gestive organs, and the like, as was already end, which has been placed in the center of well known, are all located in the medulla the brain area which is being studied. When oblongata. It has also been understood for the animal has regained consciousness and some time that the functions of these and has recovered entirely from the effect of several other primary reflex centers are in- narcosis, it is possible to stimulate the area tegrated in the mesencephalon, that is, of the midbrain where the end of the need- the grey matter, grouped in several nuclei, le is situated by conducting a weak electric which is located below the hemispheres of current through the metal wire. There were the cerebrum, in immediate proximity to various effects, depending upon the loca- the hypophysis, which exercises a regulating tion of the needle-end, each effect always influence on a great many vegetative func- connected to one definite center of stimu- tions. It has been found, inter alia, that a lation. By electrical stimulation of certain surgical operation for the purpose of remo- strictly limited parts of the midbrain it was ving tumours situated in or near this region possible to reproduce spontaneous autono- is accompanied by very great dangers due to mous functions. By stimulating one definite the fact that even slight operative traumas part, the animal was caused to curl up and in this sensitive area may cause disturbances go into a sleep from which it could be awa- of vital functions such as temperature regu- kened just as easily as from natural sleep. Sti- lation, blood pressure, and the like. mulation of other areas gave rise to defen- sive reactions which would normally occur if the cat were threatened by a dog: the hairs on the back stood up on end, the tail was

95 Award Ceremony Speech ruffled, the cat spat and, if the stimulus was The American physiologist, Fulton, and his not discontinued, it attacked. Other locali- collaborators have proved by experiments zations of the stimulus caused evacuation of on anthropoid apes that neuroses caused the intestines or the bladder, accompanied experimentally disappeared if the frontal lo- by the characteristic body postures. From bes were removed and that it was impossib- other areas, blood circulation and respira- le to cause experimental neuroses in animals tion could be influenced. It was common to deprived of their frontal lobes. all these experiments with stimuli that they released complex functions, not limited to It occurred to Moniz that psychic morbid the reactions characteristic of the autono- states accompanied by affective tension mic nervous system, such as evacuation of might be relieved by destroying the frontal the intestines, secretion of saliva, changes lobes or their connections to other parts of of the pupils, and the like, but were accom- the brain. On the basis of this idea Moniz panied by reactions of the skeletal muscles gradually worked out an operative method necessary and appropriate to the autonomic whose purpose was to interrupt the lines of function, such as characteristic body pos- communication of the frontal lobes to the tures, attempts at escape or attack, and so rest of the brain. Since these lines of com- forth. It was apparent from these facts that munication run through the white matter, in the midbrain we have higher centers of this operation was called frontal or prefron- autonomic functions which coordinate the- tal leucotomy. It was soon found that mor- se with reactions of the skeletal musculatu- bid conditions in which emotional tension re adapted to the individual functions. An was a dominating part of the pathological ingenious method made possible the exact picture reacted very favorably to such ope- determination of the anatomic localization rations. To this group of diseases belong, after the discontinuance of the experiment, primarily, states of depression accompanied and thus the anatomic substrata of complex by fear and anxiety, obsessive neuroses, autonomic functions could be mapped out. certain forms of persecution mania, and Through his research Hess has brilliantly ans- a considerable part of the most important wered a number of difficult questions re- and common of all mental diseases, schizo- garding the localization of body functions phrenia: those cases, namely, in which the in the brain. schizophrenic pattern of behaviour and the emotional condition is affectively charged The lines of thought along which Antonio to a high degree, as for instance in states of Egas Moniz has advanced to the discovery of anguish or anxiety, refusal to take food, ag- the prefrontal leucotomy refer primarily to gressiveness, and the like. Great subjective the localization of certain psychic functions suffering and invalidism are characteristic in the brain. It has long been known that of this group of diseases. Many of the di- the frontal lobes are of great importance seased, especially within the schizophrenic for higher cerebral activity, especially in re- group, are very difficult patients and are of- gard to the emotions, and that the destruc- ten dangerous to the people around them. tion of the frontal lobes, by bullet wounds When it is remembered that other methods or brain tumours, lead to certain typical of treatment have failed or have been follo- changes of the personality, primarily on the wed by recurrence of the disease, it is easy affective plane, but sometimes also affec- to understand the immense importance of ting the intellect, especially highly integra- Moniz‘ discovery for the problems of psy- ted intellectual functions such as power of chiatric treatment. As was expected, the judgement, social adaptability, and the like. results are best for the non-schizophrenic

96 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich groups, that is to say, among those suffering use a great number of suffering people and from depression, obsessive neurosis, and the total invalids have recovered and have been like, where the great majority of patients socially rehabilitated. operated upon have recovered and become capable of working. Within the schizophre- Professor Hess. On behalf of the Caroline nic group, where the disintegration of the Institute I extend to you our warm congra- personality has often advanced very far, the tulations and invite you to receive the me- prospects are less favourable, but even in dal and diploma from the hands of His Royal this group quite a few cases can be released Highness the Crown Prince. from the mental hospitals, some of them after having fully regained the capacity for The Caroline Institute much regrets that Pro- work. In other less favourable cases, the nur- fessor Moniz has not had the opportunity of sing problem will be much simplified by the being present on this occasion to receive in fact that the patient, after operation, can be person his Nobel Prize. The prize will now kept in a «quiet» ward. instead be delivered to the charge d‘affaires of the Legation of Portugal. The interesting observation has also been made that serious, bodily conditioned pain Mr. Patricio. Permit me to request you to can be successfully treated through frontal receive on behalf of Professor Moniz the leucotomy. As the operation does not touch Nobel Prize in Physiology or Medicine from any pain-communicating tracts, and the ca- the hands of His Royal Highness the Crown pacity of the patient to feel pain is unim- Prince. paired, the effect must be due to a change in the psychic experience of pain. Anguish From Nobel Lectures, Physiology or Medici- and anxiety due to pain and the affective ne 1942-1962, Elsevier Publishing Company, tension which accompanies pain disappe- Amsterdam, 1964 ar. When asked, the patient admits that he feels pain, but he does not care about the Copyright © The Nobel Foundation 1949 pain: he has become indifferent to it. These observations of psychically normal persons, on whom leucotomy has been performed in order to remove pain, have contributed in a high degree to the clarification of the influence of leucotomy on the normal men- tal functions. Without doubt there are, after double-sided leucotomy, changes of perso- nality of the same type as observed after the destruction of the frontal lobes through ot- her causes. When it becomes a question of persons who are complete invalids because of sickness, this may be of small importan- ce, while in other cases a very strict inter- pretation of indications is necessary. Frontal leucotomy, despite certain limitations of the operative method, must be considered one of the most important discoveries ever made in psychiatric therapy, because through its

97

Nobel Lecture: The Central Control of the Activity of Internal Organs

Walter Rudolf Hess

Nachdruck der Nobelpreisrede von Prof. Walter Rudolf Hess gehalten am 12. Dezember 1949 (www.nobelprize.org)

A recognized fact which goes back to the ear- After this short introduction, we now come on liest times is that every living organism is not to the concrete complex of questions, to which the sum of a multitude of unitary processes, has been devoted the work whose results have but is, by virtue of interrelationships and of earned me this great distinction and have thus higher and lower levels of control, an unbro- brought me into this high circle. The initial si- ken unity. When research, in the efforts of tuation is defined by the findings, which have bringing understanding, as a rule examines now been turned to the common good, con- isolated processes and studies them, these cerning the morphological and physical struc- must of necessity be removed from their con- ture of the so-called vegetative - „autonomic“ text. In general, viewed biologically, this ex- in English usage - nervous system. This was - I perimental separation involves a sacrifice. In believe - a good start, and was brought about fact, quantitative findings of any material and by the achievements of the great masters W.H. energy changes preserve their full context Gaskell and J.N. Langley, and had been given only through their being seen and unders- shape in an easily comprehensible and stimula- tood as parts of a natural order. This implies tingly graphic exposition by the distinguished that the laws governing organic cohesion, pharmacologists H.H. Meyer and R. Gottlieb. the organization leading from the part to the Of particular interest here in this conception, whole, represent a biological uncertainty, in- for which there is good experimental confirma- deed an uncertainty of the first order. It beco- tion, is the paired antagonistic innervation of mes all the more acute, the more rapidly the the internal organs, and their grouping accor- advances of specialization develop and threa- ding to the separate regions in which the peri- ten the ability to grasp, or even to appreciate pheral organization is linked to the central ner- it. While this state of affairs has just been re- vous system. (The keywords sympathetic and ferred to, our subject is defined by its general parasympathetic characterize the relationships content. In particular it deals with the neural under consideration.) In contrast to the explo- mechanisms by which the activity of the inter- ration of the vegetative nervous system, which nal organs is adapted to constantly changing is very far-reaching (even if it is not still without conditions, and by which they are adjusted to certain inner contradictions) stands a relatively one another, in the sense of interrelated sys- limited understanding of the central organiza- tems of functions. It only remains to be added tion of the whole mechanism of control. This is that broadening of our knowledge in these not easily understood, because the informative respects is of benefit not only with regard to experiments must explore a segment in which the human compulsion to understand, but elements are also assembled and integrated, also to the practical healing art. For man also, which subserve special sensory functions and in health and sickness, is not just the sum of the movement of the body. Things that are dis- his organs, but is indeed a human organism. tributed over a wide area in the body itself, lie close together in the central nervous system.

99 Nobel Lecture: The Central Control of the Activity of Internal Organs

Correspondingly an unequivocal differentia- yond doubt (especially Ranson and Magoun), tion is disproportionately more difficult. The have followed another path, beset with vari- direct contiguity of functionally multivalent ous avoidable experimental errors. But, here pathways and nuclei confuses the experimen- as there, fundamental to the investigations lay tal elucidation of related symptoms. One thing the same principle which enables to obtain in- had nevertheless become clear, namely that formation, if one wishes to explore the layers the parts of the brain communicating direct- and connections which lie below a surface: one ly with the spinal cord at the upper end - the applies „probes“. In our case, artificial foci of medulla oblongata, and the segment lying di- excitation were produced with electrical im- rectly beneath the cerebrum, the so-called di- pulses, and their effects noted. In addition, the encephalon - exert a decisive influence on the technique of localized exclusion was also ap- vegetative controlling mechanisms. As regards plied; as with disease foci, the reactions of this the diencephalon further orientation had been on the behaviour were interpreted in terms of achieved to the extent that it was realized that disappearance of symptoms and were linked the parts of it lying nearest the base of the indirectly to the functional significance of the skull, i.e. the hypothalamus, were particularly excluded substrata. important to the nature of the whole enquiry. Observations at the sickbed in conjunction with If I had thought, by observing the effects of what was ascertained in the deceased at autop- artificial electrical stimuli in small doses on sy, and in addition experiments which provided some dozen or so experimental animals with some rough pointers, had led to this insight altogether some hundred points of stimulation (Karplus and Kreidl et al.). Something which distributed over the diencephalon, to achieve still, however, lay in obscurity, when my own in due course the looked-for elucidation, then investigations were started, was the allocation the first result was a thorough disappointment. of definite functions to particular morphologi- The only positive finding which could be drawn cal substrata, was, in other words, the organic from the first series, was the conclusion that structure of the diencephalic vegetative control the relationships obviously had a more com- system. To throw as much light as possible on plicated lay-out than had been thought, for this was the task which I set myself. the effects were so varied that no obedience to any law could be discovered. To meet this si- At the beginning of all experimental work tuation, the experiments had to be carried out stands the choice of the appropriate techni- on a considerably expanded basis. This was a que of investigation. In many cases it has first simple enough conclusion; its realization was a to be created; it was so here as well. Although different matter. It must be born in mind that the method used to produce the results was of one does not see directly - as is the case in the decisive importance, it can only be gone into exploration of the surface of the brain - where in outline here. In the first place, satisfaction the electrodes are attacking. Exact information of two requirements was decisive, these two about the functional significance of the deep being a condition of the special circumstances sections of the brain is only obtained by wor- in the central nervous system, which have been king through the brain histologically in serial described: one concerns the technical devices, section. To avoid far too great delays, the expe- which were taken practically to the limits of re- riments must be fitted in together as it were in finement; the other is related to securing the time, and it is only possible to keep the material minimum obstruction to the experimental an- collected under control by using a carefully or- imal‘s modes of expression. We would like to ganized system of registration. The difficulty of emphasize these two, because in these particu- finding one‘s way around in the abundance of lars American investigators, whose merit is be- individual observations was overcome by a gra-

100 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich phic method. The extension of the experiments variety of types of phenomenon stands the on the widest basis means time. To this is added unity of the organism. On stimulation within the expenditure - for Swiss circumstances - of a circumscribed area of the ergotropic (dyna- considerable funds. I make these last references mogenic) zone, there regularly occurs namely on the one hand to make the slow growth of a manifest change in mood. Even a former- knowledge comprehensible, and on the other ly good-natured cat turns bad-tempered; it to be able to thank the various Swiss Founda- starts to spit and, when approached, laun- tions and above all the Rockefeller Foundation ches a well-aimed attack. As the pupils simul- of New York for their financial support. taneously dilate widely and the hair bristles, a picture develops such as is shown by the cat This short outline of the method of working if a dog attacks it while it cannot escape. The brings us to the question of the results. As we go dilation of the pupils and the bristling hairs into these, certain motor effects, although they are easily comprehensible as a sympathetic merit great interest, must be left on one side. effect; but the same cannot be made to hold good for the alteration in psychological be- Now, concerning the influence of the dience- haviour. For this, only connections between phalon on the activity of the internal organs, hypothalamus, thalamus and cerebral cortex the following facts could be disclosed: first of come into consideration. Functionally, the to- all, it has turned out that the functions which tal behaviour of the animal illustrates the fact are mediated by the sympathetic section of that, in the part of the diencephalon indica- the vegetative nervous system, are related to ted, a meaningful association of physiological the posterior and middle parts of the deepest processes takes place, which is related on the section of the hindbrain, i.e. of the hypotha- one hand to the regulation of the internal or- lamus. So the latter is to be considered, as it gans, and on the other involves the functions were, as the central area of origin of the sym- directed outwards towards the environment. pathetic system. In order to give this discovery In other words: we know the key position in its full physiological import, some more eluci- the diencephalon which has one aspect di- dation is required. The goal of physiological rected inwards and one aspect directed out- research is functional nature. So in the course wards. The sympathetic system is thereby, of this preoccupation with the vegetative ner- within the framework of a far-reaching orga- vous system, among other things the question nization, the mediating agent which interve- has arisen of whether a circumscribed role is nes particularly in the activity of the internal associated with the classical sympathetic sys- (vegetative) organs. With regard to the mani- tem, which is defined primarily in terms of its fest influence of the psychomotor system and area of origin, which is restricted to the tho- the psychological processes of association, a racic spinal cord. An investigation undertaken bridge is thrown over a gap, still wide open from the viewpoint of the effect of its activity today, which lies between the purely somati- has yielded the finding that this is the case to cally oriented physiology and psycho-physio- a considerable extent. Where the sympathetic logy. It completes and broadens the insight intervenes, it assists the body‘s efficiency and into psychosomatic relationships, in the way it aids the organism to greater success in its they had been demonstrated by the great conflicts with its environment. It is functional, Russian physiologist Pavlov, who approached in so far as it behaves like an ergotropic or dy- them from another side. To him also fell the namogenic system. In addition to this item of great honour of speaking from this position. knowledge there are still more findings which will interest the psychiatrist in particular, but In spite of the necessary restrictions on our also everyone who realizes that behind the exposition, observations of a different kind

101 Nobel Lecture: The Central Control of the Activity of Internal Organs will induce us briefly to touch on the theme Often a profuse flow of saliva occurs; further of somato-motor phenomena once again. Be- symptoms are choking and vomiting, mictu- fore this, another striking finding must be re- rition, defaecation. In other cases panting is ported. The individual, vegetatively innerva- caused, i.e. the mechanism most often seen ted, organ gets its differentiated innervation in the dog under natural conditions, when it in the known peripheral organization of the is hot. While the tongue, with its rich blood sympathetic ergotropic system; correspon- supply, moistened by a copious flow of saliva, dingly it can also be brought into action in iso- hangs out of the wide-open mouth with the lation. This fact and the relationships, as they flow of air due to rapid respiration streaming are met with, for example in the projection over it, the discharge of excess heat takes place. of the peripheral organs reacting to nervous This function, which is also common to the cat, stimulation in the motor area of the cerebral serves the regulation of temperature and is in cortex, could give grounds for supposing that this sense equivalent to sweating (e.g. in man) the individual internal organs also have a di- when heat accumulates. Another effect of sti- screte representation in the diencephalon. In mulation not mentioned so far is constriction of such an order of things the ergotropic zone the pupils, followed by a drawing-across of the would also be organized as it were by organs. nictitating membrane.

The experimental findings offer proof that To summarize, we are dealing with symptoms in reality the relationships are disposed dif- which are characteristic of a decrease in the in- ferently. The fact is that even the most nar- fluence due to the sympathetic system, and of rowly circumscribed forms of excitation and an increase of parasympathetically transmitted the most delicate stimulus dose never bring excitation. Regarding the physiological effect, to light an isolated symptom related to one such reactions bring functional relaxation to organ. In every case a group symptomatolo- the individual organs, or protection against gy makes its appearance. It is always groups overloading, but indeed protection above all. of organs that are called into action, and in- Where the digestive processes are concerned, deed in such a way that the individual effects the complex serving restoration is mediated by are combined, namely in accordance with the separate mechanisms. Since we give these re- principle of synergistic coordination. Controls lated effects a common denominator in their issue from the diencephalon which harness functional aspect, the term „trophotropic sys- the functional capacities of individual organs tem“ is appropriate. Moreover the experimen- in viable responses. This order of things holds tal findings show that a circumscribed region good quite markedly in the ergotropic zone. of the diencephalon corresponds to it, namely But under the influence of circumscribed sti- the anterior part of the hypothalamus, the area muli applied to the hypothalamus, and partly praeoptica, and the septum pellucidum as well. also to the layers of the thalamus lying clo- With this, a central, fairly clearly demonstrable se above it, symptoms have also appeared division of the two partners of the vegetative which do not permit of classification in the nervous system becomes manifest. This is all the sympathetic-ergotropic system of functions, more noteworthy, as they are most intimately and indeed rather act in opposition to this. interwoven in their peripheral terminal territo- The blood pressure, for example, does not ries. Important, too, is the establishment that respond by a rise, but by a fall; the heart there is no evidence in the trophotropic zone rate does not increase, but rather decreases. of a central organization corresponding to par- At the same time respiration slows down, as ticular organs. It emerges conclusively from the opposed to the speeding-up which is obtai- effects obtained from the most varied sites of ned from the ergotropic-dynamogenic zone. stimulation falling in the area named that here

102 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich too no grouping is found in contrast, for exam- applied to the most rostrally situated regions, ple in the formation of nuclei; each particular the cat adopts the normal posture for the phy- syndrome shows a fairly large scatter. This does siological deposition of faeces; therefore the not, however, conceal the fact that some ef- stimulus activates the skeletal musculature, fects are preferentially released from certain which is innervated by the cerebrospinal axis, larger areas. It is different in this respect from and which is also responsible for the abdominal the ergotropiczone, where the group organi- muscular pressure. It was possible to make the zation is more consistent. same observation during micturition. The syn- ergistic coordination between the function re- We take a step forward, when we turn our at- leased through the vegetative nervous system tention to the observations from which it emer- and the somatomotor complement is manifest. ges that reciprocal mutual connections operate An improvement in the result is obtained, part- between the sympathetic-ergotropic and the ly in the form of a speeding-up of the process, parasympathetic-trophotropic areas, indeed in and partly in the way soiling is avoided. the sense that at each moment they produce a dynamic equilibrium adapted to the situati- Particularly impressive is the synergistic coor- on at any given moment of the organism as a dination of mechanisms controlled by the ve- whole. In this equilibrium the unity of the cen- getative nervous system with cerebrospinal in- tral regulation of the whole vegetative system nervated activities in the defence mechanism is expressed. In the broad view, competition is a which is accompanied by emotion. This had constructive principle. already been the subject of discussion, but wit- hout any particular light being thrown on the With this statement, we could conclude our ex- structure of the action as a whole in relation position, that is, if we wanted to confine it to to the systematic involvement of somatomotor a more narrowly conceived theme. But among phenomena. Dilation of the pupils, bristling of the fundamental results of the experimental the hair are the vegetative components, and exploration of the diencephalons alone was snuffling and spitting as somatomotor proces- the finding that the effects produced from ses complete the picture intended to scare off this part of the brain are not restricted to the the opponent. The aimed blow with the paw vegetative system. Indeed the rule is that they conclusively presupposes a visual orientation, are associated with somatomotor symptoms. which, while compulsive in affect, nevertheless When I refer to this, I do not have in mind the fits in adaptively because of the intervention functions which testify to a higher order regu- of the cortex. Thus one sees how the various lation of bodily posture within the framework levels make their contribution to the full suc- of the extra-pyramidalmotor system. What we cess of a complete activity, and one unders- are discussing here concerns the motor sympto- tands how individual functions are associated matology which stands in a specific relationship in stages in an activity carried out by the orga- to the vegetative function. This is where our nism as a whole. interest lies, if we attend briefly to this state of affairs. So let us look at a new stage in the According to this view of how the diencepha- integration of the organ functions in the total lon plays a decisive role in activity which pro- performance. In this way, for example, we can gresses from the part to the whole, it will come understand the experimental finding, that in as no surprise if still other observations could many cases where the stimulus applied to the be made which lead in another direction. Thus diencephalon causes defaecation, this is not it has been seen that under defined experi- brought about simply by peristalsis of the colon mental conditions a constriction of the pupil and rectum. In particular, where a stimulus is and a drawing-across of the nictitating mem-

103 Nobel Lecture: The Central Control of the Activity of Internal Organs brane may be caused from a certain region of undisturbed recovery. The latter - judging by the diencephalon. A slowly developing nar- the physiological sleep - particularly concerns rowing of the palpebral fissure accompanies the higher centres. For the rest, as regards the these events, which reflect a decrease in the where and the why, electroencephalography, sympathetic innervation, which - particularly in e.g. in the line of research by Jasper, and bio- the pupil - occurs with an increase in parasym- chemistry have the say in the matter now. So pathetic influence. These effects also are not the investigations, which have been reported infrequently associated with certain symptoms here, have, as is the rule, in addition to a clear under the nervous control of the central ner- understanding also brought into focus the for- vous system: the drooping of the upper eyelid mulation of new questions. develops into an active closing of the lid. Simul- taneously the head droops, whereupon, as the Now that we have come full cycle with the in- syndrome develops further, the whole animal vestigation in which the integrating activity lies down. It is necessary to observe this pro- of the diencephalon was experimentally dif- cess accurately in its development and in the ferentiated, I would very much like to show final stage; it will be noticed, as you will realize, some more pictures, with which the spoken that the cat does not simply collapse, as is the word will be clarified. The time available, of case with generalized loss of tone. Under the course, permits only a limited selection of sli- influence of gradual relaxation the animal cho- des and a film. But I think they are enough to ses its place and curls up. Altogether a pictu- provide an objective representation of what re results like that known under physiological has been said. conditions only in the sleeping cat. In a certain sense one is presented with the mirror-image 1. The following were shown, as lantern sli- of the emotionally aroused cat with increased des: rise in blood pressure, fall in blood excitability; for the readiness to react to senso- pressure, increase of respiration, slowing of ry stimuli is markedly decreased, whereby - as respiration; in the film: methodology -dila- in the normal sleeping cat - the tickle reflex of tion of pupils, retraction of the nictitating the ear stays „awake“. Behaviour towards ol- membrane, defaecation, retching, panting factory stimuli is also relatively little inhibited, - licking movements, chewing movements, which are effective as a more potent arousal sniffing movements - affective reactions: stimulus, thereby proving the reversibility of spitting, bristling of hair, leaping to attack, the whole process. In other respects, as has urge to eat, urge to flee - atonia as an effect been mentioned, the preparedness for ener- of stimulation - adynamia as an effect of ex- getic activity is reduced to a minimum. Clearly, clusion - sleep as the effect of stimulation. in the competition between ergotropic and trophotropic systems the former forfeits some 2. The following localizing findings were pre- of its influence on the organism as a whole in sented: medium section of the cat‘s brain favour of an excess of the latter. But it must and further sagittal sections with symbols always be borne in mind that we are dealing marked in (lowering blood pressure, raising with the result of artificial stimulation, and in- blood pressure, pupil dilation, pupil cons- deed in a limited area. To explain the gradual triction, affective defence, hunger drive). development of this inhibition of activity as the result of destruction in the hypothalamus is misleading. On the other hand it is correct that one is dealing with a protective function con- trolled from the diencephalon, which avoids exhaustion and produces the conditions for an

104 Funktion und nervöse Organisation der inneren Organe

Walter Rudolf Hess

105 Funktion und nervöse Organisation der inneren Organe

106 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

107 Funktion und nervöse Organisation der inneren Organe

108 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

109 Funktion und nervöse Organisation der inneren Organe

110 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

111 Funktion und nervöse Organisation der inneren Organe

112 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

113 Funktion und nervöse Organisation der inneren Organe

114 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

115 Funktion und nervöse Organisation der inneren Organe

116 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

117 Funktion und nervöse Organisation der inneren Organe

118 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

119 Funktion und nervöse Organisation der inneren Organe

120 Walter Rudolf Hess: Nobelpreis für Medizin oder Physiologie 1949

Margrit Wyder

Erinnerung an die Verleihung des Nobelpreises an Prof. Walter Rudolf Hess, von Margrit Wyder: Einstein und Co.-Nobelpreisträger in Zürich, Verlag NZZ libro, Zürich 2015

Walter Rudolf Hess Nobelpreis für Physiologie oder Medizin 1949

brachte er ihm etwa bei, zur passenden Universität Zürich hielt. Einen zufälli- Gelegenheit «Guten Appetit» zu sagen. gen Fund bei der Sektion, nämlich eine Das Interesse am Lebendigen zieht Anomalie in den Arterien des Fusses, sich wie ein roter Faden durch das Le- konnte der Student auf eine mechani- ben von Walter Rudolf Hess. Doch er sche Gesetzmässigkeit zurückführen. war kein sentimentaler Tierliebhaber. Für seine Dissertation an der Uni- Walter RudolfNeben der Hess Unterhaltung dienten ihm versität Zürich erforschte Hess 1901 Nobelpreis für PhysiologieTiere als Studien- und oder Forschungsob Medizin- die Viskosität, 1949 also die Zähflüssigkeit, jekte. Mensch und Tier waren für Hess des Blutes, und erfand dafür gleich physiologisch gut vergleichbar. Er be- ein neues Instrument, das «Hess’sche brachte er ihm etwa bei, zur passenden Universität Zürich hielt. Einen zufälli- wunderte als Forscher die sinnreiche Viskosimeter», das über Jahrzehnte in Gelegenheit «Guten Appetit» zu sagen. gen Fund bei der Sektion, nämlich eine Organisation des Körpers, sah darin der medizinischen Praxis verwendet Das Interesse am Lebendigen zieht Anomalie in den Arterien des Fusses, eine sich selbst erhaltende, durch die wurde. Als Augenarzt in Rapperswil sich wie ein roter Faden durch das Le- konnte der Student auf eine mechani- Nobelpreis für Physiologie oder Medizin 1949 Evolution perfektionierte Maschine. In – eine Gedenktafel am Haus «Zum Pe- ben von Walter Rudolf Hess. Doch er sche Gesetzmässigkeit zurückführen. «für seine Entdeckung der funktionalen den 1930er-Jahren fand er sein frucht- likan» weist dort auf sein Wirken hin war kein sentimentaler Tierliebhaber. Für seine Dissertation an der Uni- Organisation des Zwischenhirns barstes Forschungsfeld in der Funkti- – führte er 1909 eine neue Methode zur Neben der Unterhaltung dienten ihm versität Zürich erforschte Hess 1901 als Koordinator der Aktivitäten der onsweise des Gehirns. Tiere als Studien- und Forschungsob- die Viskosität, also die Zähflüssigkeit, inneren Organe» Begonnen hatte alles mit Naturbe- jekte. Mensch und Tier waren für HessWalter des Rudolf Blutes, Hess und sah erfand im dafür gleich obachtungen im ländlichen Thurgau * 17. 3. 1881 in Frauenfeld physiologisch gut vergleichbar. ErKörper be- ein eine neues durch Instrument, Evolution das «Hess’sche und Laborexperimenten, die der 1881 † 12. 8. 1973 in Ascona wunderte als Forscher die sinnreicheperfektionierte Viskosimeter», Maschine. das über Jahrzehnte in geborene Walter Rudolf mit seinem Organisation des Körpers, sah darin der medizinischen Praxis verwendet 1913 – 1916 Privatdozent, 1917 – 1951 Professor Vater, Physikprofessor am Frauenfel- eine sich selbst erhaltende, durch die wurde. Als Augenarzt in Rapperswil für Physiologie an der Universität Zürich der Gymnasium, durchführen konnte. einfachen Erkennung von Bewegungs- Nobelpreis für Physiologie oder Medizin 1949 Evolution perfektionierte Maschine. In – eine Gedenktafel am Haus «Zum Pe- Dieses doppelte Interesse, in Kombina- störungen der Augen ein, und 1912 ent- «für seine Entdeckung der funktionalen den 1930er-Jahren fand er sein frucht- likan» weist dort auf sein Wirken hin tion mit «ursprünglichster Originali- wickelte er eine Technik der Stereofoto- Organisation des Zwischenhirns barstes Forschungsfeld in der Funkti- – führte er 1909 eine neue Methode zur tät», wie es in der Festschrift zu seinem grafie, die nur wegen des bald darauf als Koordinator der Aktivitäten der onsweise des Gehirns. 60. Geburtstag heisst, machte ihn zum ausbrechenden Weltkriegs nicht zur inneren Organe» Begonnen hatte alles mit Naturbe- genialen Erfinder und zu einem der praktischenWalter Anwendung Rudolf Hess kam. sah im obachtungen im ländlichen Thurgau Das* 17.Rätsel 3. 1881 in Frauenfeld des wichtigsten Vertreter der experimen- Um seineKörper Verlobte eine durch Louise Evolution Sand- und Laborexperimenten, die der 1881 † 12. 8. 1973 in Ascona tellen Biologie. meier – perfektionierte sie stammte ebenfalls Maschine. aus Schlafs geborene Walter Rudolf mit seinem Das Medizinstudium führte Walter Frauenfeld – heiraten zu können, war 1913 – 1916 Privatdozent, 1917 – 1951 Professor Vater, Physikprofessor am Frauenfel- Der alte Professor hatte einen Vogel Rudolf Hess an mehrere Universitäten Hess in die Praxis gegangen. Doch er für Physiologie an der Universität Zürich der Gymnasium, durchführen konnte. einfachen Erkennung von Bewegungs- – buchstäblich: Papagei Joko sass mit im In- und Ausland: Lausanne, Bern, war als Arzt nicht glücklich, wollte Dieses doppelte Interesse, in Kombina- störungen der Augen ein, und 1912 ent- Vorliebe auf der Schulter von Walter Berlin, Kiel und Zürich. In Erinnerung zurück in die universitäre Forschung. tion mit «ursprünglichster Originali- wickelte er eine Technik der Stereofoto- Rudolf Hess und zerbiss immer wieder an «den ersten Kontakt mit Interessen Trotz Bedenken seiner Frau wegen der tät», wie es in der Festschrift zu seinem grafie, die nur wegen des bald darauf die Kabel seines Hörgeräts. Der Vogel von wissenschaftlichem Charakter … finanziellen Folgen nahm er 1912 eine 60. Geburtstag heisst, machte ihn zum ausbrechenden Weltkriegs nicht zur durfte sich alles erlauben, zum gros- sehe ich mich auf dem Präpariersaal der Stelle als Assistent am Physiologischen genialen Erfinder und zu einem der praktischen Anwendung kam. sen Ärger von Frau Hess. Den Profes- Zürcher Anatomie», sagte Hess in der Institut bei Professor Justus Gaule an. Das Rätsel des wichtigsten Vertreter der experimen- Um seine Verlobte Louise Sand- sor interessierten die Hirnleistungen Rede, die er nach Bekanntwerden der So zog die Familie mit der zweijährigen tellen Biologie. meier – sie stammte ebenfalls aus des Papageis,Schlafs die er eifrig trainierte. So Nobelpreis-Verleihung in der Aula der Tochter Gertrud an die Winterthurer- Das Medizinstudium führte Walter Frauenfeld – heiraten zu können, war Der alte Professor hatte einen Vogel Rudolf Hess an mehrere Universitäten Hess in die Praxis gegangen. Doch er – buchstäblich: Papagei Joko sass mit im In- und Ausland: Lausanne, Bern, war als Arzt nicht glücklich, wollte www.nobelpreis.uzh.ch/hess.html Walter Rudolf Hess – Nobelpreis für Physiologie oder Medizin 1949 1/4 Vorliebe auf der Schulter von Walter Berlin, Kiel und Zürich. In Erinnerung zurück in die universitäre Forschung. Rudolf Hess und zerbiss immer wieder an «den ersten Kontakt mit Interessen Trotz Bedenken seiner Frau wegen der die Kabel seines Hörgeräts. Der Vogel von wissenschaftlichem121 Charakter … finanziellen Folgen nahm er 1912 eine durfte sich alles erlauben, zum gros- sehe ich mich auf dem Präpariersaal der Stelle als Assistent am Physiologischen sen Ärger von Frau Hess. Den Profes- Zürcher Anatomie», sagte Hess in der Institut bei Professor Justus Gaule an. sor interessierten die Hirnleistungen Rede, die er nach Bekanntwerden der So zog die Familie mit der zweijährigen des Papageis, die er eifrig trainierte. So Nobelpreis-Verleihung in der Aula der Tochter Gertrud an die Winterthurer-

www.nobelpreis.uzh.ch/hess.html Walter Rudolf Hess – Nobelpreis für Physiologie oder Medizin 1949 1/4 Walter Rudolf Hess: Nobelpreis für Medizin oder Physiologie 1949

strasse 27 in Zürich. 1913 kam hier der Medium ein. So filmte er Sportler in Kinder ein Aquarium und ein Terra- Sohn Rudolf Max zur Welt. «Lisy» Hess Bewegung, um das perfekte Zusam- rium ein, damit sie die Natur studieren unterstützte ihren Mann zeitlebens bei menspiel der aktiven, haltenden und lernten. Zur Familie gehörte auch ein der Arbeit. Sie wirkte als Privatsekretä- stützenden Muskeln zu dokumentie- kleiner Wolfshund namens Jürg. rin und tippte seine wissenschaftlichen ren. Zur Verdeutlichung seiner phy- Als Leiter des Instituts konnte Hess Aufsätze auf der Schreibmaschine. siologischen Erkenntnisse liess Hess seine Forschungsfelder frei wählen. Die HabilitationsschriftWalter Rudolfvon Walter Trickfilme Hess und ausgeklügelte Modelle Nachdem er zunächst Kreislaufthe- Rudolf NobelpreisHess behandelte die physika für- Physiologieherstellen. Seine Physiologie-Vorlesun oder Medizin- men behandelt 1949 hatte, wollte er ab 1929 lisch-biologische Basis der Kreislauf- gen waren stets akribisch vorbereitet. mehr über die Natur und den Sinn des regulierung. Dabei zeigte sich erneut Ging eine Vorführung schief, bekam Schlafes herausfinden. Dafür brauchte brachte er ihm etwa bei, zur passenden Universität Zürich hielt. Einen zufälli- seine aussergewöhnliche Fähigkeit, der Labordiener etwas zu hören – Hess es Tierversuche, und Hess wählte Kat- Gelegenheit «Guten Appetit» zu sagen. gen Fund bei der Sektion, nämlich eine Gesetzmässigkeiten logisch abzulei- hatte das lebhafte Temperament seiner zen, weil sich diese Tiere im Körper- Das Interesse am Lebendigen zieht Anomalie in den Arterien des Fusses, ten. 1915 nutzte er die Möglichkeit, für sächsischen Mutter geerbt. bau wenig voneinander unterschei- sich wie ein roter Faden durch das Le- konnte der Student auf eine mechani- ein Jahr in Bonn bei dem berühmten Der langjährige Arbeitsort von den und auch gern am Tag schlafen. ben von Walter Rudolf Hess. Doch er sche Gesetzmässigkeit zurückführen. Physiologen Max Verworn zu arbeiten. Hess war das Physiologische Institut Er entwickelte eine Methode, um mit war kein sentimentaler Tierliebhaber. Für seine Dissertation an der Uni- Bei seiner Rückkehr nach Zürich hatte im Obergeschoss des universitären elektrischen Reizen einzelne Areale Neben der Unterhaltung dienten ihm versität Zürich erforschte Hess 1901 sich die berufliche Situation drama- des Zwischenhirns zu stimulieren. Die Tiere als Studien- und Forschungsob- die Viskosität, also die Zähflüssigkeit, tisch verändert: Institutsleiter Gaule aufwendigen und diffizilen Versuche Für Literatur,jekte. Mensch Kunst und oder Tier waren für Hess des Blutes, und erfand dafür gleich war ernsthaft erkrankt und trat vorzei- führte Hess mit wenigen Hilfskräften Musikphysiologisch war in der Familie gut vergleichbar. Hess Er be- ein neues Instrument, das «Hess’sche tig zurück. Hess übernahm seine Ver- über Jahre hinweg mit grösster Sorgfalt kaumwunderte Platz – das als MikroskopForscher die sinnreiche Viskosimeter», das über Jahrzehnte in tretung. Doch konnte er sich als junger durch. Eine Kinderschwester, die er war Kultur.Organisation des Körpers, sah darin der medizinischen Praxis verwendet Privatdozent Hoffnung auf die Nach- zuhause nicht mehr benötigte, wurde eine sich selbst erhaltende, durch die wurde. Als Augenarzt in Rapperswil folge Gaules machen? Die Berufung zur treuen Mitarbeiterin: Anna Jaussi Nobelpreis für Physiologie oder Medizin 1949 Evolution perfektionierte Maschine. In – eine Gedenktafel am Haus «Zum Pe- entwickelte sich zum Politikum, denn Physikgebäudes an der Rämistrasse protokollierte die Versuche und führte «für seine Entdeckung der funktionalen den 1930er-Jahren fand er sein frucht- likan» weist dort auf sein Wirken hin zunächst wurde ihm von der Fakultät 69. Am Jahresende 1923 war es akut über 20 Jahre die statistische Doku- Organisation des Zwischenhirns barstes Forschungsfeld in der Funkti- – führte er 1909 eine neue Methode zur ein erfahrener deutscher Physiologe gefährdet: Wegen einer Nachlässigkeit mentation der Hirnreizexperimente als Koordinator der Aktivitäten der onsweise des Gehirns. vorgezogen. Dies sorgte für Unmut am des Hauswarts kam es zu einem nächt- durch. So gewann man schrittweise inneren Organe» Begonnen hatte alles mit Naturbe- Institut, und es zeigte sich, dass eine lichen Dachstockbrand. Als positive grundlegendeWalter Erkenntnisse Rudolf Hess über sah dieim obachtungen im ländlichen Thurgau sehr positive* 17. 3. 1881 Empfehlung in Frauenfeld Verworns Folge der Verwüstungen wurde das Gehirnfunktionen.Körper eine durch Evolution und Laborexperimenten, die der 1881 für Hess† 12. unterdrückt 8. 1973 in Ascona worden war. Der Gebäude gründlich umgebaut; durch Mit 68perfektionierte Jahren, noch mitten Maschine. in seiner geborene Walter Rudolf mit seinem damalige Zürcher Erziehungsdirektor eine Aufstockung erhielt auch die Tätigkeit, erhielt Walter Rudolf Hess 1913 – 1916 Privatdozent, 1917 – 1951 Professor Vater, Physikprofessor am Frauenfel- Heinrich Mousson griff nun persönlich Physiologie mehr Platz. Mit der ge- für seine Forschungen am Zwischen- für Physiologie an der Universität Zürich der Gymnasium, durchführen konnte. einfachen Erkennung von Bewegungs- in die Berufungsfrage ein, und Hess gen viele Widerstände durchgesetzten hirn den Nobelpreis. Die Preisverlei- Dieses doppelte Interesse, in Kombina- störungen der Augen ein, und 1912 ent- wurde im September 1917 zum Ordent- Gründung der internationalen Hochal- hung in Stockholm im Dezember 1949 tion mit «ursprünglichster Originali- wickelte er eine Technik der Stereofoto- lichen Professor für Physiologie und pinen Forschungsstation auf dem Jung- empfand er als einen Höhepunkt sei- tät», wie es in der Festschrift zu seinem grafie, die nur wegen des bald darauf Institutsleiter gewählt. Er dankte dies fraujoch konnte Hess einen weiteren, nes Lebens. Er liebte aber auch lokale 60. Geburtstag heisst, machte ihn zum ausbrechenden Weltkriegs nicht zur der Universität, indem er sein ganzes einmaligen Ort für die Wissenschaft Feiern und gesellige Anlässe. Für Li- genialen Erfinder und zu einem der praktischen Anwendung kam. Forscherleben hier verbrachte. gewinnen. teratur, Kunst oder Musik war in der Das Rätsel des wichtigsten Vertreter der experimen- Um seine Verlobte Louise Sand- Ein besonderes Augenmerk richtete Privat bewohnte der Institutsdirek- Familie hingegen kaum Platz. «Das tellen Biologie. meier – sie stammte ebenfalls aus Walter RudolfSchlafs Hess von Beginn an auf tor mit seiner Familie nun eine standes- Mikroskop ist Kultur», bemerkte Hess, Das Medizinstudium führte Walter Frauenfeld – heiraten zu können, war die Lehre. Unter den Studierenden war gemässe Wohnung an der Susenberg- wenn seine Tochter sich darüber be- Der alte Professor hatte einen Vogel Rudolf Hess an mehrere Universitäten Hess in die Praxis gegangen. Doch er er als begeisternder, aber strenger und strasse 198, nahe beim Kurhaus Rigi- klagte. Während der Sohn Rudolf Max – buchstäblich: Papagei Joko sass mit im In- und Ausland: Lausanne, Bern, war als Arzt nicht glücklich, wollte manchmal ungeduldiger Dozent be- blick. Der Haushalt blieb aber betont als Neurologe die Hirnstudien des Va- Vorliebe auf der Schulter von Walter Berlin, Kiel und Zürich. In Erinnerung zurück in die universitäre Forschung. kannt. Als einer der Ersten setzte er einfach, privaten Luxus gab es nicht. ters durch Elektroenzephalogramm- Rudolf Hess und zerbiss immer wieder an «den ersten Kontakt mit Interessen Trotz Bedenken seiner Frau wegen der in Lehre und Forschung den Film als Auf der Veranda richtete Hess für die Untersuchungen ergänzte, studierte die Kabel seines Hörgeräts. Der Vogel von wissenschaftlichem Charakter … finanziellen Folgen nahm er 1912 eine durfte sich alles erlauben, zum gros- sehe ich mich auf dem Präpariersaal der Stelle als Assistent am Physiologischen sen Ärger von Frau Hess. Den Profes- Zürcher Anatomie», sagte Hess in der Institut bei Professor Justus Gaule an. www.nobelpreis.uzh.ch/hess.html Walter Rudolf Hess – Nobelpreis für Physiologie oder Medizin 1949 2/4 sor interessierten die Hirnleistungen Rede, die er nach Bekanntwerden der So zog die Familie mit der zweijährigen des Papageis, die er eifrig trainierte. So Nobelpreis-Verleihung in der Aula der Tochter Gertrud an die Winterthurer-

www.nobelpreis.uzh.ch/hess.html Walter Rudolf Hess – Nobelpreis für Physiologie oder Medizin 1949 1/4

122 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

TrudiTrudi BiologieBiologie undund fandfand schliesslichschliesslich InIn Zürich Zürich aufgeklärt: aufgeklärt: sichtensichten auf auf das das Versuchstier, Versuchstier, zu zu denen denen der der ihren eigenen Weg: Sie liess sich 1948 Forscher durch das hohe Ziel seiner Arbeit ihren eigenen Weg: Sie liess sich 1948 DieDie Organisation Organisation des des Forscher durch das hohe Ziel seiner Arbeit am neu gegründeten Jung-Institut zur verpflichtet ist, wissentlich verletzt sind, da am neu gegründeten Jung-Institut zur ZwischenhirnsZwischenhirns verpflichtet ist, wissentlich verletzt sind, da AnalytikerinAnalytikerin ausbilden.ausbilden. stellenstellen wir wir uns uns rückhaltlos rückhaltlos auf auf die die Seite Seite der der AlsAls Walter Walter Rudolf Rudolf Hess Hess 1951 1951 von von DasDas Zwischenhirn Zwischenhirn liegt, liegt, wie wie sein sein Name Name schon schon Tierschützer.»Tierschützer.» seinemseinemWalter AmtAmt alsals InstitutsdirektorInstitutsdirektor Rudolf zuzu-- sagt,sagt, zwischen zwischenHess dem dem auf auf dem dem Rückenmark Rückenmark DassDass seine seine Grundlagenforschung Grundlagenforschung zu zu rücktrat,rücktrat, hattehatte erer ZürichZürich zu zu einemeinem welt -welt- aufsitzendenaufsitzenden Hirnstamm Hirnstamm und und dem dem Gross Gross- - therapeutischentherapeutischen Anwendungen Anwendungen führen führen weitweit beachtetenbeachtetenNobelpreis ZentrumZentrum derder fürHirnforHirnfor- -Physiologiehirn,hirn, das das beim beim Menschen Menschen oder besonders besonders Medizin stark stark könnte,könnte, war war für1949 für Hess Hess kein kein Thema. Thema. Eingriffe Eingriffe schungschung gemacht,gemacht, undund erer konntekonnte mit mit entwickeltentwickelt ist. ist. Vom Vom Zwischenhirn Zwischenhirn werden werden le le- - insins Gehirn Gehirn verliefen verliefen in in der der ersten ersten Hälfte Hälfte des des Genugtuung feststellen, dass seine benswichtige, so genannt vegetative Funk- 20. Jahrhunderts noch brachial. So erhielt Genugtuung feststellen, dass seine benswichtige,brachte soer ihmgenannt etwa vegetative bei, zur Funkpassenden- 20. JahrhundertsUniversität noch Zürich brachial. hielt. So erhieltEinen zufälli- Schüler die Forschung weiterführten. tionen und Gefühlsverhalten gesteuert: die der spanische Neurologe Egas Moniz den Schüler die Forschung weiterführten. tionenGelegenheit und Gefühlsverhalten «Guten Appetit» gesteuert: zu die sagen. der spanischegen Fund Neurologe bei der Egas Sektion, Moniz nämlich den eine Der Emeritus behielt im Anatomischen Körpertemperatur, der Wasserhaushalt des Nobelpreis 1949 gemeinsam mit Hess, und Der Emeritus behielt im Anatomischen Körpertemperatur,Das Interesse der Wasserhaushalt am Lebendigen des ziehtNobelpreis Anomalie 1949 gemeinsam in den Arterien mit Hess, unddes Fusses, Institut beim Universitätsspital ein Körpers, Hunger- und Sättigungsgefühl, de- zwar «für die Entdeckung des therapeuti- Institut beim Universitätsspital ein Körpers,sich Hunger- wie ein und roter Sättigungsgefühl, Faden durch de- daszwar Le- «fürkonnte die Entdeckung der Student des auftherapeuti eine mechani- - Büro, wo er weiterhin wissenschaft- fensive Abwehr, Angriff und auch der Schlaf. schen Wertes der präfrontalen Leukotomie Büro, wo er weiterhin wissenschaft- fensiveben Abwehr, von Walter Angriff undRudolf auch Hess.der Schlaf. Doch schen er Wertessche derGesetzmässigkeit präfrontalen Leukotomie zurückführen. liche Aufsätze schrieb. Oft wurde er Der biologische Sinn des Schlafs war für die bei gewissen Psychosen». Bei dieser Ope- liche Aufsätze schrieb. Oft wurde er Der biologischewar kein sentimentalerSinn des Schlafs war Tierliebhaber. für die bei gewissenFür Psychosen». seine Dissertation Bei dieser Opean -der Uni- dahin von seinem Hund «Schnäuzli» Wissenschaft lange ein Rätsel. Walter Rudolf ration werden die Nervenbahnen in den dahin von seinem Hund «Schnäuzli» WissenschaftNeben der lange Unterhaltung ein Rätsel. Walter dienten Rudolf ihmration versität werden die Zürich Nervenbahnen erforschte in den Hess 1901 begleitet. Von der Goldauerstrasse 25 Hess verstand ihn als lebensnotwendige vorderen Hirnregionen durchtrennt. Durch begleitet. Von der Goldauerstrasse 25 HessTiere verstand als Studien- ihn als lebensnotwendige und Forschungsob vorderen- die Hirnregionen Viskosität, durchtrennt. also die Zähflüssigkeit, Durch Einrichtung, um dem Körper Erholung von das Verfahren sollten Wahnvorstellungen Einrichtung,jekte. Mensch um dem Körperund Tier Erholung waren von für Hessdas Verfahrendes Blutes, sollten und Wahnvorstellungen erfand dafür gleich der täglichen Aktivität zu verschaffen. Bei geheilt werden, es führte aber bei den Hess hat Zürich zu einem der täglichenphysiologisch Aktivität gutzu verschaffen.vergleichbar. Bei Ergeheilt be- ein werden, neues es Instrument, führte aber beidas den«Hess’sche Hess hat Zürich zu einem äusserer Ruhe können so «Instandstellungs- Patienten zu irreparablen Schäden und weltweit beachteten Zentrum äussererwunderte Ruhe können als Forscher so «Instandstellungs die sinnreiche- Patienten Viskosimeter», zu irreparablen das Schäden über Jahrzehnte und in weltweit beachteten Zentrum arbeiten» verrichtet werden. Persönlichkeitsveränderungen. der Hirnforschung gemacht. arbeiten»Organisation verrichtet werden.des Körpers, sah darinPersönlichkeitsveränderungen. der medizinischen Praxis verwendet der Hirnforschung gemacht. Die Erforschung des Zwischenhirns Die Hess’sche Forschungsmethode der Dieeine Erforschung sich selbst des erhaltende, Zwischenhirns durch dieDie wurde.Hess’sche Als Forschungsmethode Augenarzt in Rapperswil der war sehr aufwendig. Die Versuchstiere, in tiefen elektrischen Hirnstimulation ist Nobelpreis für Physiologie oder Medizin 1949 war Evolutionsehr aufwendig. perfektionierte Die Versuchstiere, Maschine. in tiefen In – elektrischen eine Gedenktafel Hirnstimulation am Haus ist «Zum Pe- – in den 1940er-Jahren war man etwas diesem Fall Katzen, wurden zunächst durch jedoch in den letzten Jahren erfolgreich – in den«für 1940er-Jahren seine Entdeckung war der man funktionalen etwas diesemden Fall 1930er-Jahren Katzen, wurden fand zunächst er sein durch frucht jedoch- likan» in den weist letzten dort Jahren auf erfolgreich sein Wirken hin näher zur Stadt gezogen – fuhr Hess Füttern an den Versuchstisch gewöhnt. angewendet worden bei der Behandlung näher Organisationzur Stadt gezogen des Zwischenhirns – fuhr Hess Fütternbarstes an denForschungsfeld Versuchstisch gewöhnt.in der Funkti angewendet- – führte worden er 1909 bei eine der Behandlungneue Methode zur meist mit der Rigiblick-Seilbahn und In Narkose schraubte man ihnen einen von Symptomen der Parkinson-Krankheit meist mitals Koordinator der Rigiblick-Seilbahn der Aktivitäten der und In Narkoseonsweise schraubte des Gehirns. man ihnen einen von Symptomen der Parkinson-Krankheit dem Tram zur Arbeit, und der Hund Elektrodensockel auf den Schädel, von und anderen Bewegungsstörungen. Elekt- dem Traminneren zur Organe» Arbeit, und der Hund ElektrodensockelBegonnen auf hatte den alles Schädel, mit vonNaturbe und- anderen Bewegungsstörungen. Elekt- rannte aus-serhalb des öffentlichen Ge- dem die feinen Stahlfäden ins schmerzun- roden werdenWalter dabei Rudolf chirurgisch Hess ins Gehirn sah im rannte aus-serhalb des öffentlichen Ge- demobachtungen die feinen Stahlfäden im ländlichen ins schmerzun -Thurgauroden werden dabei chirurgisch ins Gehirn fährts mit* 17. 3.– 1881ein Umstand,in Frauenfeld der bei den empfindliche Gehirn gesenkt wurden. Bei eingepflanztKörper und übereine dünne durch Kabel Evolution mit fährts mit – ein Umstand, der bei den empfindlicheund Laborexperimenten, Gehirn gesenkt wurden. die Beider 1881eingepflanzt und über dünne Kabel mit Passanten† 12. zu 8. 1973Diskussionen in Ascona Anlass gab. leichter Reizung mit Strom zeigte das Tier einem Impulsgeberperfektionierte im Brustbereich Maschine. ver- Passanten zu Diskussionen Anlass gab. leichtergeborene Reizung Waltermit Strom Rudolf zeigte dasmit Tier seinem einem Impulsgeber im Brustbereich ver- Enkel Christian W. Hess, später selbst dann je nach der Lage der Elektrodenspitze bunden. Durch die Impulse können bei den Enkel Christian1913 – 1916 Privatdozent, W. Hess, später 1917 – 1951 selbst Professor dannVater, je nach Physikprofessor der Lage der Elektrodenspitze am Frauenfel bunden.- Durch die Impulse können bei den Professor für Neurologie, erinnert sich zum Beispiel Schläfrigkeit oder aggressi- Patienten zum Beispiel das störende Zittern Professorfür Physiologie für Neurologie, an der Universität erinnert sichZürich zumder Beispiel Gymnasium, Schläfrigkeit durchführen oder aggressi -konnte.Patienten einfachen zum Beispiel Erkennung das störende von Zittern Bewegungs - auch an lehrreiche Spaziergänge des ves Verhalten. Am getöteten Versuchstier verhindert oder die Bewegungshemmung auch an lehrreiche Spaziergänge des ves Verhalten.Dieses doppelte Am getöteten Interesse, Versuchstier in Kombina verhindert- störungen oder die der Bewegungshemmung Augen ein, und 1912 ent- Grossvaters mit den drei Enkelkin- liessen sich die Spuren der Elektroden im überwunden werden. Wie dieser «Hirn- Grossvaters mit den drei Enkelkin- liessention sich mit die «ursprünglichster Spuren der Elektroden Originaliim überwunden- wickelte werden. er eine Wie Technik dieser «Hirn der Stereofoto- - dern, die oft mit einer Ovomaltine im Zwischenhirn beobachten. Millimeter für schrittmacher» im Detail funktioniert, ist dern, die oft mit einer Ovomaltine im Zwischenhirntät», wie esbeobachten. in der Festschrift Millimeter fürzu seinemschrittmacher» grafie, imdie Detail nur funktioniert,wegen des baldist darauf Kurhaus Rigiblick endeten. Millimeter wurde so das Hirngewebe erkun- noch ungeklärt. (MW) Kurhaus Rigiblick endeten. Millimeter60. Geburtstag wurde so das heisst, Hirngewebe machte erkun ihn- zumnoch ungeklärt.ausbrechenden (MW) Weltkriegs nicht zur 1967 verliess das Ehepaar Hess Zü- det. Daraus entstanden Karten, auf denen 1967 verliess das Ehepaar Hess Zü- det. genialenDaraus entstanden Erfinder Karten, und auf zu denen einem der praktischen Anwendung kam. rich und zog samt Papagei ins Ferien- die lokalisierten Areale im Hirn einzelnen rich undDas zog samtRätsel Papagei insdes Ferien - die lokalisiertenwichtigsten Areale Vertreter im Hirn dereinzelnen experimen - Um seine Verlobte Louise Sand- haus im Tessin. Mit 92 Jahren starb Funktionen zugeordnet werden konnten. haus im Tessin. Mit 92 Jahren starb Funktionentellen Biologie.zugeordnet werden konnten. meier – sie stammte ebenfalls aus Walter SchlafsRudolf Hess dort an Herzver- Dabei legte Hess immer Wert dar- Walter Rudolf Hess dort an Herzver- DabeiDas legte Medizinstudium Hess immer Wert führte dar Walter- Frauenfeld – heiraten zu können, war sagen. Margrit Wyder auf, dass man möglichst schonend mit sagen. DerMargrit alte WyderProfessor hatte einen Vogelauf, Rudolf dass man Hess möglichst an mehrere schonend Universitäten mit Hess in die Praxis gegangen. Doch er den Katzen umging. Der «Tierschutz im – buchstäblich: Papagei Joko sass mitden im Katzen In- und umging. Ausland: Der «Tierschutz Lausanne, im Bern, war als Arzt nicht glücklich, wollte Tierversuch» war ihm ein Anliegen, wor- Quelle: MargritVorliebe Wyder: auf Einsteinder Schulter und Co. – von WalterTierversuch»Berlin, Kiel war undihm Zürich.ein Anliegen, In Erinnerungwor- zurück in die universitäre Forschung. Quelle: Margrit Wyder: Einstein und Co. – über er in der Zeitschrift «Der Tierfreund» NobelpreisträgerRudolf Hess in Zürich; und Verlag zerbiss NZZ immer libro, wieder an «den ersten Kontakt mit Interessen Trotz Bedenken seiner Frau wegen der Nobelpreisträger in Zürich; Verlag NZZ libro, über er in der Zeitschrift «Der Tierfreund» Zürich 2015 Illustration: Aline Telek schrieb: «Wo und wann immer die Rück- Zürich 2015die KabelIllustration: seines Aline Hörgeräts. Telek Der Vogelschrieb:von «Wowissenschaftlichem und wann immer die Rück Charakter- … finanziellen Folgen nahm er 1912 eine durfte sich alles erlauben, zum gros- sehe ich mich auf dem Präpariersaal der Stelle als Assistent am Physiologischen sen Ärger von Frau Hess. Den Profes- Zürcher Anatomie», sagte Hess in der Institut bei Professor Justus Gaule an. www.nobelpreis.uzh.ch/hess.html Walter Rudolf Hess – Nobelpreis für Physiologie oder Medizin 1949 3/4 www.nobelpreis.uzh.ch/hess.htmlsor interessierten die Hirnleistungen Rede, die er nach WalterBekanntwerden Rudolf Hess – derNobelpreis So zog für die Physiologie Familie oder mit Medizinder zweijährigen 1949 3/4 des Papageis, die er eifrig trainierte. So Nobelpreis-Verleihung in der Aula der Tochter Gertrud an die Winterthurer-

www.nobelpreis.uzh.ch/hess.html Walter Rudolf Hess – Nobelpreis für Physiologie oder Medizin 1949 1/4

123 Walter Rudolf Hess: Nobelpreis für Medizin oder Physiologie 1949

Walter Rudolf Hess Nobelpreis für Physiologie oder Medizin 1949

brachte er ihm etwa bei, zur passenden Universität Zürich hielt. Einen zufälli- Gelegenheit «Guten Appetit» zu sagen. gen Fund bei der Sektion, nämlich eine Das Interesse am Lebendigen zieht Anomalie in den Arterien des Fusses, sich wie ein roter Faden durch das Le- konnte der Student auf eine mechani- Begeisternd, aber streng: Walter Hirnleistungen interessiertenben von Walter Walter Rudolf Rudolf Hess:Hess. Seinem Doch Papagei er sche Joko Gesetzmässigkeit zurückführen. Rudolf Hess im Hörsaal. brachte er bei, beiwar passenden kein sentimentaler Gelegenheiten Tierliebhaber.«Guten Appetit» zu sagen.Für seine Dissertation an der Uni- Bild: Privatarchiv Christian W. Hess Neben der Unterhaltung dienten ihm versität Zürich erforschte Hess 1901 Tiere als Studien- und Forschungsob- die Viskosität, also die Zähflüssigkeit, jekte. Mensch und Tier waren für Hess des Blutes, und erfand dafür gleich physiologisch gut vergleichbar. Er be- ein neues Instrument, das «Hess’sche wunderte als Forscher die sinnreiche Viskosimeter», das über Jahrzehnte in Organisation des Körpers, sah darin der medizinischen Praxis verwendet eine sich selbst erhaltende, durch die wurde. Als Augenarzt in Rapperswil Nobelpreis für Physiologie oder Medizin 1949 Evolution perfektionierte Maschine. In – eine Gedenktafel am Haus «Zum Pe- «für seine Entdeckung der funktionalen den 1930er-Jahren fand er sein frucht- likan» weist dort auf sein Wirken hin Organisation des Zwischenhirns barstes Forschungsfeld in der Funkti- – führte er 1909 eine neue Methode zur als Koordinator der Aktivitäten der onsweise des Gehirns. inneren Organe» Begonnen hatte alles mit Naturbe- Walter Rudolf Hess sah im obachtungen im ländlichen Thurgau * 17. 3. 1881 in Frauenfeld Körper eine durch Evolution und Laborexperimenten, die der 1881 † 12. 8. 1973 in Ascona perfektionierte Maschine. geborene Walter Rudolf mit seinem 1913 – 1916 Privatdozent, 1917 – 1951 Professor Vater, Physikprofessor am Frauenfel- für Physiologie an der Universität Zürich der Gymnasium, durchführen konnte. einfachen Erkennung von Bewegungs- Dieses doppelte Interesse, in Kombina- störungen der Augen ein, und 1912 ent- tion mit «ursprünglichster Originali- wickelte er eine Technik der Stereofoto- tät», wie es in der Festschrift zu seinem grafie, die nur wegen des bald darauf 60. Geburtstag heisst, machte ihn zum ausbrechenden Weltkriegs nicht zur genialen Erfinder und zu einem der praktischen Anwendung kam. Das Rätsel des wichtigsten Vertreter der experimen- Um seine Verlobte Louise Sand- Schlafs tellen Biologie. meier – sie stammte ebenfalls aus Das Medizinstudium führte Walter Frauenfeld – heiraten zu können, war Der alte Professor hatte einen Vogel Rudolf Hess an mehrere Universitäten Hess in die Praxis gegangen. Doch er – buchstäblich: Papagei Joko sass mit im In- und Ausland: Lausanne, Bern, war als Arzt nicht glücklich, wollte Ein KatzenexperimentVorliebe auf in derWalter Schulter Rudolf Hess’ von Labor, Walte protokolliertr Berlin, von Kiel Anna und Jaussi. Zürich. Bild: Privatarchiv In Erinnerung Christian zurückW. Hess in die universitäre Forschung. Rudolf Hess und zerbiss immer wieder an «den ersten Kontakt mit Interessen Trotz Bedenken seiner Frau wegen der die Kabel seines Hörgeräts. Der Vogel von wissenschaftlichem Charakter … finanziellen Folgen nahm er 1912 eine durfte sich alles erlauben, zum gros- sehe ich mich auf dem Präpariersaal der Stelle als Assistent am Physiologischen sen Ärger von Frau Hess. Den Profes- Zürcher Anatomie», sagte Hess in der Institut bei Professor Justus Gaule an.

www.nobelpreis.uzh.ch/hess.htmlsor interessierten die Hirnleistungen Rede, die er nachWalter Bekanntwerden Rudolf Hess – Nobelpreis der So fürzog Physiologie die Familie oder mit Medizin der zweijährigen 1949 4/4 des Papageis, die er eifrig trainierte. So Nobelpreis-Verleihung in der Aula der Tochter Gertrud an die Winterthurer-

www.nobelpreis.uzh.ch/hess.html Walter Rudolf Hess – Nobelpreis für Physiologie oder Medizin 1949 1/4

124 4. Walter Rudolf Hess und die Tradition der Zürcher Neurowissenschaft

125

Zur Tradition der Zürcher Neurowissenschaft: vom «Monakowschen Kränzchen» zur von Walter Rudolf Hess gegründeten «Zürcher Arbeitsgemeinschaft für Hirnforschung» und darüber hinaus Anton Valavanis

Ausgangspunkt der Entwicklung der Zürcher lichen «Kränzchen» den «Psychiatrisch-Neuro- Neurowissenschaften ist der an der Universität logischen Verein in Zürich». Unter der Leitung Zürich ausgebildete Arzt und Hirnforscher rus- von Monakows beteiligten sich daran mit Vor- sischer Abstammung Constantin von Monakow trägen und Diskussionen zu Fragen der Hirn- (1853-1930). Er war Neurologe, Gründer der forschung seine Mitarbeiter und Schüler, die schweizerischen, Mitgründer der deutschen Psychiater des Burghölzlis und andere neuro- neurologischen Gesellschaft und erster Pro- wissenschaftlich interessierte Neurologen und fessor und Direktor des von ihm gegen er- Psychiater, Vertreter benachbarter Gebiete hebliche Widerstände seitens der Zürcher sowie zahlreiche Gäste aus dem Ausland. Ein Medizinischen Fakultät gegründeten Hirnana- regelmässiger Teilnehmer an den periodischen tomischen Institutes in Zürich, welches er unter Versammlungen des Vereins war auch Walter schwierigsten Arbeitsbedingungen zu Weltruf Rudolf Hess, der seit 1917 Direktor des, wenige geführt hat (3-6, 8-11). Schritte vom Haus Belmont entfernten, Physio- logischen Institutes der Universität Zürich war. Bereits vier Jahre nach der Gründung seines In einem Brief an den damals 64-jährigen, welt- Hirnanatomischen Institutes (1894), rief von bekannten Hirnforscher von Monakow, datiert Monakow 1898 ein gemeinsames Diskus- vom 20. September 1917, bekundete der im sionsforum ins Leben mit der Absicht einen Herbstsemester 1917 frisch zum Professor für fortbestehenden Austausch zwischen Neuro- Physiologie und Direktor des physiologischen logen, Neuroanatomen, Neurophysiologen Institutes berufene, 36-jährige Walter Rudolf und Psychiatern zu etablieren (12). Es ging in Hess seine Bereitschaft und sein Interesse an die lokale Medizingeschichte als das berühmte den angekündigten Versammlungen des neu- «Monakowsche Kränzchen» ein (6). Forschung gegründeten Vereins teilzunehmen: «Ich kann und Klinik waren aber weiterhin örtlich ge- mir keine idealere Gelegenheit denken, mich trennt, was den Alltag sehr erschwerte. Von auf sicherer Grundlage und unter fachkundi- Monakow empfand dies als eine Schwäche ger Führung in die Physiologie des Zentralner- und ein Hindernis für die Weiterentwicklung vensystems – des Nervensystems überhaupt der Neurowissenschaften und bemühte sich – einzuleben, als wenn ich ein möglichst regel- beharrlich, Hirnforschung und klinisch-neuro- mässiger Teilnehmer ihres psychiatrisch-neuro- logische Tätigkeit gemeinsam an einem Ort logischen Vereins werde. Ich werde dort, so zusammenzuführen. Dies gelang ihm erst viel ist mir klar, in erster Linie als Nehmender 1913 als das Hirnanatomische Institut und die und nicht als Gebender erscheinen können. Ich Neurologische Poliklinik unter einem Dach, im darf dies wohl sagen, ohne mir eine Blösse zu Haus Belmont an der Rämistrasse 67, in unmit- geben. Denn bei der Einseitigkeit mit welcher telbarer Nähe zur Universität und zum Kan- die Physiologie des Zentralnervensystems von tonsspital untergebracht wurden (3, 11, 12). meinen Fachgenossen behandelt wird, und Nach Bezug der Räume im Haus «Belmont» bei der riesigen Ausdehnung unseres Gebietes gründete von Monakow aus seinem ursprüng- überhaupt, kann ich mir nichts anderes vor-

127 Zur Tradition der Zürcher Neurowissenschaft: vom «Monakowschen Kränzchen» zur von Walter Rudolf Hess gegründeten «Zürcher Arbeitsgemeinschaft für Hirnforschung» und darüber hinaus stellen, als dass ich bei der Gelegenheit unserer nisation des Zwischenhirns erhalten sollte, im Besprechungen viel Neues hören werde.» (12). Geiste von Monakows die Zürcher Arbeitsge- Neben seiner Beteiligung an den dort durchge- meinschaft für Hirnforschung gründete. Die- führten Diskussionen, hielt er mit der Zeit auch ses Konzept wurde zum Vorbild für die 1961 selbst Vorträge zu seinen Forschungsergeb- erfolgte Gründung des Institutes für Hirnfor- nissen, wovon manche von ihnen im von Mo- schung als Nachfolgeinstitution des von Mona- nakowschen Schweizer Archiv für Neurologie kowschen Hirnanatomischen Institutes. Hess und Psychiatrie veröffentlicht wurden, so zum beschreibt dieses Ereignis in seiner Autobio- Beispiel sein Vortrag «Über die Wechselbezie- graphie wie folgt (englische Übersetzung, 7) hungen zwischen psychischen und vegetativen Funktionen», welcher am 31. Mai 1924 gehal- «I have the satisfaction of announcing a grat- ten und im Band XV, Heft 2, 260-177, 1924 des ifying undertaking, which is intimately con- Schweizer Archivs erschienen ist. Mit der Zeit nected to my life‘s work. As early as 1940, wurde Hess mit seiner aktiven Partizipation an I had assumed the initiative for founding a den Versammlungen des Vereins von einem study group devoted to brain research. Pe- ‘Nehmenden’ auch zu einem ‘Gebenden’. So riodically, physiologists, psychiatrists, brain wurde der psychiatrisch-neurologische Verein surgeons, internists, pharmacologists, neu- die Plattform für eine Interaktion und einen ro-anatomists, psychoanalysts, and psycho- Austausch von Ideen und Ansichten über Fra- therapists would come together at the phys- gen der Hirnforschung zwischen von Mona- iological institute. At first, we wanted to get kow und Hess. Sein Schüler Konrad Akert, in to know one another‘s points of view and seiner Würdigung der Beiträge von Hess zur to find a common language for the various Neurowissenschaft geht mit folgender Bemer- terms and concepts. kung auf diese Interaktion ein: “Constantin von Monakow (1853-1930), founder and di- Thanks to the financial support of the Rock- rector of the Neurological Policlinic and of the efeller Foundation and later to that of the Institute of Brain Anatomy of the University Swiss Academy of Medical Sciences, through of Zürich (1894-1928), had reached the climax its Commission for Brain Research, this study of fame when Hess began his scientific career. group achieved its goal with ever expanding Monakow had been the highly respected lea- participation until 1950. Today the original der of a group of students and admirers who plan has culminated in an institute for brain met periodically in his home under the name research, to be opened in 1963. of ‘Monakow’s Kränzchen’ where problems of brain localization and the brain-mind enigma If one considers my own interest in the were vividly discussed from neurological, psy- functional organization of the brain along chiatric, and philosophical points of view. Hess with its psychic manifestation—an interest frequented the informal gatherings of this which was expressed as far back as 1924 and group and held Monakow in high esteem.” which was never lost sight of—one will well (13). Diese Interaktion erwies sich später als understand that the decision of the faculty besonders wichtig für die Weiterentwicklung and the responsible authorities to create a und vielschichtige Entfaltung der Zürcher chair for brain research has filled my young Neurowissenschaften. So ist auch zu verste- old heart with the very deepest satisfaction. hen, dass im Jahr 1940, ein Jahrzehnt nach der Equally gratifying is the fact that I can regard Emeritierung von von Monakow, Hess, der in- the future of the institute with confidence, zwischen in der Hirnforschung tätig war und as one of my earlier co-workers, K. Akert, 1949 den Nobelpreis für seine experimentellen presently on the medical faculty, has been Untersuchungen über die funktionelle Orga- named its director”. (From: Hess W.R.: From

128 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

Medical Practice to Theoretical Medicine: und der Neurologischen Poliklinik ernannt. An- An Autobiographic Sketch. Perspectives in gesichts der Tatsache, dass Lüthy schwerpunkt- Biology and Medicine. 6 (4): 400-423, 1963). mässig der Neuropathologie und klinischer Neurologie und weniger der neurobiologi- Nach der Emeritierung von Monakows im schen Grundlagenforschung zugewandt war, Jahr 1927 wurde sein langjähriger und enge- sah sich die Fakultät vor das Dilemma gestellt rer Mitarbeiter Mieczyslaw Minkowski 1928 entweder das Hirnanatomische Institut auf- zu seinem Nachfolger als Extraordinarius ad zuheben oder einen neuen, eigenständigen personam für Neurologie und Direktor des Lehrstuhl für Hirnforschung zu schaffen. Es ist Hirnanatomischen Institutes und der Neuro- dem energischen Einsatz des Neurochirurgen logischen Poliklinik ernannt. Erst im Jahr 1944 Hugo Krayenbühl und von Walter Rudolf Hess wurde sein persönliches in ein étatmässiges Ex- (s. oben) zu verdanken, dass nach 6-jährigen traordinariat umgewandelt. Nebst Weiterfüh- kontroversen Verhandlungen schlussendlich rung seiner Forschungstätigkeit konzentrierte der Zürcher Regierungsrat sich entschied ein er seine Kräfte hauptsächlich auf die neuro- neues Institut für Hirnforschung zu schaffen. logisch-ärztliche, also klinische Tätigkeit. 1952 Als dessen Direktor und Lehrstuhlinhaber (Or- folgte die Übersiedlung des Hirnanatomischen dinariat) wurde 1960 der Schüler von Walter Institutes mit der Nervenpoliklinik in das neu Rudolf Hess, Konrad Akert, der zu jener Zeit als gebaute Kantonsspital. Dabei wurde eine ers- Professor für Neuroanatomie und Neurophy- te, kleine Bettenstation eingerichtet, was An- siologie an der Universität von Wisconsin (USA) lass zur Gründung einer eigentlichen Neurolo- tätig war, gewählt. Der neu gewählte Akert gischen Klinik, der ersten in der Schweiz, war. würdigte die Leistungen seines Lehrers Hess, Auch während der Amtszeit Minkowskis wur- anlässlich seiner Antrittsvorlesung mit dem Ti- den die periodischen, interdisziplinären Ver- tel «Ziele und Ergebnisse der Hirnforschung», sammlungen des von von Monakow gegrün- gehalten am 27. Oktober 1962 an der Universi- deten «Psychiatrisch-Neurologischen Vereins tät Zürich, mit folgenden Worten: «Der große in Zürich» unter aktiver Partizipation von Wal- Wurf, 1949 mit dem Nobelpreis ausgezeichnet, ter Rudolf Hess fortgeführt (11), wobei Akert, gelang dem Zürcher Physiologen W. R. Hess. das Verhältnis zwischen Hess und Minkowski Eine überragende Grundkonzeption und eine eher als kühl beschreibt: «Relations with Mo- bis ins feinste Detail ausgearbeitete Technik nakow’s successsor, Mieczyslaw Minkowski des experimentellen Vorgehens bildeten die (1884-1972) were relatively cool. An attempt Voraussetzungen. In einer beispiellosen Sys- at collaboration with Minkowski on the ana- tematik der punktförmigen Abtastung bisher tomical analysis of Hess’ experimental brain unerreichbarer und unerschlossener Gehirnab- material concerning extrapyramidal motor schnitte in der Tiefe des Zwischenhirns wurde symptoms (Minkowski, 1943) ended when the Element um Element aus dem gesamten Ver- physiologist and the anatomist could not ag- haltensrepertoire der Katze herausgeholt, was ree on matters of interpretation, whereupon normalerweise durch natürliche Umgebungs- Hess enlisted Verena Bucher (1901-1986) as an reize ausgelöst wird. Vor allem die angebore- expert on the anatomy of the cat brain.” (13). nen, eingebauten Schablonen der Selbst- und Im Jahr 1954 wurde Minkowski emeritiert und Arterhaltungstriebe kamen zum Vorschein, der frei praktizierende Neurologe, neurolo- und deren funktionelle Organisation und die gische Berater der Medizinischen Klinik und Lokalisierung der verantwortlichen Nervenele- Leiter der neurohistologischen Station des Pa- mente konnten weitgehend aufgeklärt wer- thologischen Institutes Fritz Lüthy wurde als den.» (1). Nachfolger Minkowskis zum Extraordinarius und Direktor des Hirnanatomischen Institutes

129 Zur Tradition der Zürcher Neurowissenschaft: vom «Monakowschen Kränzchen» zur von Walter Rudolf Hess gegründeten «Zürcher Arbeitsgemeinschaft für Hirnforschung» und darüber hinaus

Unter Akert erfuhr die Zürcher Hirnforschung Entwicklung des Interdisziplinaritätskon- einen regelrechten Schub, was für die Weiter- zeptes der Zürcher Neurowissenschaft entwicklung der Zürcher Neurowissenschaft förderlich war. Treu zu der von Monakow und 1898 v. Monakow gründet den psychiatrisch- Hess initiierten Tradition, pflegte auch Akert neurologischen Referierabend die Kollaboration mit der klinischen Neuro- (Monakowsches Kränzchen) logie. Dieses Konzept wurde, mit der in der 1904 v. Monakow gründet den Psychiatrisch- Amtszeit Akerts, 1972 von Alexander Borbély, Neurologischen Verein in Zürich Walter Lichtensteiger und Hans Zeier ins Le- ben gerufenen «Arbeitsgemeinschaft Neuro- 1940 Walter Rudolf Hess gründet die Interdis- biologie Zürich» fortgeführt. Akert führte im ziplinäre Arbeitsgemeinschaft für Rahmen der akademischen Lehrtätigkeit die Hirnforschung wöchentliche, gemeinsame Lehrveranstaltung 1972 Alexander Borbély, Walter Lichtenstei- «Probleme der Hirnlokalisation mit klinischen ger und Hans Zeier gründen die Arbeits- Demonstrationen» unter Mitwirkung von gemeinschaft Neurobiologie Zürich Akert selbst, G. Baumgartner, Hugo Krayen- bühl, R.B. Livingston, R. Hess, M.G. Yasargil, M. 1998 Martin Schwab gründet das Zentrum Wiesendanger, G. Dumermuth, W. Isler und J. Neurowissenschaften Zürich (ZNZ) der Siegfried (s. XI. Jahresbericht 1971/1972, Hirn- UZH und ETHZ forschungsinstitut der Universität Zürich, p.16). 2014 Anton Valavanis, Michael Weller und Luca Regli gründen das Klinische Neu- Die so erzielte enge Verknüpfung zwischen rozentrum des USZ klinischer Neurologie und Hirnforschung er- wies sich als ein charakteristisches Merkmal der Weiterentwicklung der Zürcher Neurowis- Es ist der uneigennützigen Weitsicht von Wal- senschaften, die fast ein Jahrhundert später ter Rudolf Hess, der sich 1960-1961 für die Grün- mit der Errichtung zunächst der Neurochirur- dung des Institutes für Hirnforschung und der gischen Klinik durch Hugo Krayenbühl (1937), Übertragung dessen Leitung an seinen Schüler dann des Institutes für Hirnforschung durch Konrad Akert dezidiert eingesetzt hat, zu ver- Walter Rudolf Hess und Konrad Akert (1961) danken, dass die von Monakowsche Tradition und schliesslich der Klinik für Neuroradiologie an der Universität Zürich und am Universitäts- durch Anton Valavanis (1994) in die Gründung spital Zürich erfolgreich weitergeführt werden des Zentrums für Neurowissenschaften der konnte. Auch Akert, so wie sein Lehrer Hess, Universität und ETH Zürich durch den Hirn- hat die Nachwuchsförderung in den Neuro- forscher Martin Schwab (ZNZ, 1998) sowie des wissenschaften zu einer seiner Hauptaufgaben Klinischen Neurozentrums des USZ durch den gemacht, wie aus den letzten Sätzen seiner An- Neuroradiologen Anton Valavanis, den Neuro- trittsvorlesung hervorgeht: «Noch hat die mo- logen Michael Weller und den Neurochirurgen derne Hirnwissenschaft erst eine kurze Strecke Luca Regli (KNZ, 2014) mündete. zurückgelegt, und sie hat einen weiten Weg bis zum Ziel vor sich. Aber es scheinen genug An- haltspunkte da zu sein, welche die Richtung der naturwissenschaftlichen Arbeit klar vorzeich- nen. Um die großen Aufgaben der Zukunft zu lösen, braucht es aber nicht nur teure Appara- te und zeitraubende Methoden, sondern jene Kräfte und Energien, welche begeisterungsfä-

130 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich Die Protagonisten der Zürcher neurowissenschaftlichen Tradition Die ProtagonistenDie ProtagonistenDie Protagonisten der Zürcherder Zürcher der neurowissenschaftlichenZürcher neurowissenschaftlichen neurowissenschaftlichen Tradition Tradition Tradition Die Protagonisten der Zürcher neurowissenschaftlichen Tradition

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Zur Tradition der Zürcher Neurowissenschaft: vom «Monakowschen Kränzchen» zur von Walter Rudolf Hess gegründeten «Zürcher Arbeitsgemeinschaft für Hirnforschung» und darüber hinaus higen jungen Gehirnen entstammen.» (1). 8. Jagella C., Isler H., Hess K.: 100 Jahre Neurologie an der Literatur Universität Zürich, 1894–1994: Constantin von Mona- 1. Akert K.: Ziele und Ergebnisse der Hirnforschung. Neue kow (1853–1930), Hirnforscher – Neurologe – Psych- Zürcher Zeitung, 16. Dezember 1962, Blatt 5, Nr. 5045 iater – Denker, Schweiz. Arch. Neurol. Psychiatr. (Swiss (134) Archives of Neurology and Psychiatry) 145 1–60, 1994 2. Akert K. Das Hirnforschungsinstitut. Die Universität Zü- 9. Kesselring J.: Developments in neurology from the rich 1933-1983. Festschrift zur 150-Jahr-Feier der Uni- 19th to the 20th century with special reference to vari- versität Zürich. Hrsg. Rektorat der Universität Zürich, ous contributions from Switzerland. Schweiz Rundsch 1983 Med Prax 83:491–6 1994 3. Akert K.: Das Institut für Hirnforschung der Universität 10. Minkowski M.: Constantin von Monakow 1853–1930. Zürich. Physiologie: Forschung, Lehre, Öffentlichkeit 5 Schweiz Arch Neurol Psychiatr 27: 1–58 1931 6-12 1995 11. Minkowski M.: Die Poliklinik für Nervenkranke und das 4. Bassetti CL and Valko PO: History of the Swiss Neuro- Hirnanatomische Institut. In: Zürcher Spitalgeschichte, logical Society. Schweiz Arch Neurol Psychiatr 160(2): Band II. Hrsg. Regierungsrat des Kantons Zürich, 427- 52–60 2009 474 1951 5. Gubser AW. Aus den Papieren Constantin von Mona- 12. von Monakow C.: Vita mea – Mein Leben, Huber Ver- kows (1853-1930). Gesnerus. 24 (3-4); 1967 lag, Bern, Switzerland, 1970 6. Hess K.: Geschichte der Neurologischen Klinik und Poli- 13. Akert K.: Walter Rudolf Hess (1881-1973) and His Con- klinik Zürich. Schweiz Arch Neurol und Psychiatr. 159 tribution to Neuroscience. Journal of the History of the (4); 191-197 2008 Neurosciences. 8 (3): 248-263; 1999 7. Hess WR. From Medical Practice to Theoretical Medici- ne: An Autobiographic Sketch. Perspectives in Biology Anschrift des Verfassers and Medicine 6, (4), 1963, pp. 400-423 Prof. em. Dr. med., Dr. h. c., Anton Valavanis Klinisches Neurozentrum USZ Frauenklinikstrasse 10 CH-8091 Zürich E-Mail: [email protected]

132 Zur Interaktion zwischen Walter Rudolf Hess und M. Gazi Yasargil

Anton Valavanis

Nach vierzigjähriger (1952-1992), ununterbro- und seine Assistenz im Operationssaal bei der chener Tätigkeit während welcher Professor Durchführung stereotaktischer Eingriffe im Gazi M. Yasargil die neurochirurgische Klinik Hypothalamus, die neuroanatomische Region des USZ zur weltweit unbestrittenen führen- welche er erforscht und wofür er den Nobel- den Institution herangeführt hatte, entschied preis erhalten hatte. Dies ist ein frühes Beispiel er sich nach seiner im Januar 1993 erfolgten der fruchtbaren Interaktion zwischen Grund- Emeritierung als Direktor der Neurochirurgi- lagenforschung und klinischer Anwendung in schen Klinik des USZ und Professor für Neuro- der Neurowissenschaft, realisiert Jahrzehnte chirurgie an der Universität Zürich, aus einer bevor der Begriff ‘translationale Forschung’, Fülle von Angeboten mehrerer Universitäten der heute in aller Munde ist und von Woolf Europas und der USA, den Ruf auf eine or- wie folgt umschrieben wurde: «Translational dentliche Professur der University of Arkan- research means different things to different sas Medical School (UAMS) in Little Rock an- people, but it seems important to almost every- zunehmen, wo er am 1. Oktober 1994 seine one6», eingeführt wurde. neue Tätigkeit begonnen hat. Bereits am 2. November 1994 führte er dort seine erste Textauszug aus Yasargil M. G.: A legacy mikroneurochirurgische Operation durch1. of microneurosurgery: Memoirs, lessons Während seiner Tätigkeit in Little Rock fällt and axioms. Neurosurgery 45 (5): 1999, auch seine ehrenvolle Auszeichnung als ‘Neu- pp. 1030-1031 rosurgery’s Man of the Century2,3,4. In ihrer «On January 4, 1953, I began my training in Ausgabe vom November 1999 kündigte die neurosurgery with Professor Krayenbühl in renommierte Zeitschrift Neurosurgery, als Zurich and lived at the University Hospital for das offizielle Organ des Congress of Neurolo- 40 very intense working years. I am still ac- gical Surgeons, die Ernennung von Professor tive there in my daytime reveries and in my Harvey Cushing(†) als den Neurosurgery’s nighttime dreams. Both the university and Man of the Century 1900-1949 und Professor the Kantonsspital (later University Hospital) M. Gazi Yasargil als den Neurosurgery’s Man in Zurich were founded in 1830, during a of the Century 1950-1999 an. Anlässlich dieser very dynamic socioeconomic era. Zurich was Ehrung und auf Einladung des Herausgebers the home of the great Swiss reformer Ulrich der Zeitschrift Neurosurgery veröffentlichte Zwingli (1484-1531). The political turmoil in Yasargil seine Autobiographie mit dem Titel: Europe during the 19th century exercised a «A Legacy of Microneurosurgery: Memoirs, favorable influence on the development of Lessons and Axioms5». Darin widmet er einen the university, the hospital, and the cultural Abschnitt der Würdigung der Persönlichkeit, evolution in Zurich. Numerous emigrants, der Leistungen und der Bedeutung von Hess among them scientists, philosophers, writers, für die Neurowissenschaft. Im letzten Ab- composers, and artists, were welcomed by schnitt seiner Ausführungen erwähnt Yasargil the liberal democratic government in Zurich. die vom theoretischen Hirnforscher Hess ge- The small but dynamic city soon became a währte Unterstützung bei der Weiterentwick- world center for business, finance, medicine, lung der damaligen neurochirurgischen Klinik surgery, and the neurosciences. A century of

133 Zur Interaktion zwischen Walter Rudolf Hess und M. Gazi Yasargil sustained, assiduous work by genius scientists Professor Hess essentially supported the de- culminated in Nobel Prizes in physics and che- velopment of the neurosciences (at present, mistry, and in 1949, in medicine, in the per- there are seven centers in Zurich) and the son of Professor W.R. Hess. neurosurgical department. He impressed me immensely as being a sincere scientist and During the course of 40 years, Professor Hess most gentle individual. In 1959, he assisted performed painstaking minute microsurgical me at a stereotactic procedure to define the work in the laboratory of the physiology insti- hypothalamic areas in a patient and assisted tute in Zurich, introducing perfectly targeted in the stereotactic application of bilateral hy- microelectrodes into the cat hypothalamus, pothalamic microlesions». which allowed him and his team to perform physiological and pharmacological experi- Literatur ments. Using a special staining technology, 1. Al-Mefty O.: M. Gazi Yasargil: The Time in Little Rock. they mapped the investigated region of the Neurosurgery 45 (5): 1019-1024, 1999 hypothalamus. This research was financially 2. Apuzzo L.M.J.: Summa Cum Laude. Neurosurgery 45 supported by the Rockefeller Foundation. As (5): 975-976, 1999 a consequence of his paradigmatic questio- 3. Tew Jr. J.M.: M.Gazi Yasargil: Neurosurgery’s Man of ning, Professor Hess opened a new dimen- the Century. Neurosurgery 45 (5): 1010-1014, 1999 sion in biology and neurobiology. He could 4. Flamm ES.: Professor M. Gazi Yasargil: An Appreciation by a Former Apprentice. Neurosurgery 45 (5): 1015- explore the function of the hypothalamus, as 1018, 1999 well as the principles of servomechanism and 5. Yasargil M.G.: A legacy of microneurosurgery: Memoirs, autoregulation, within this investigated re- lessons and axioms. Neurosurgery 45 (5): 1025-1091, 1999 gion and examine its relationship with other 6. Woolf S.H.: The Meaning of Translational Research and functional systems. His hitherto not widespread Why It Matters. JAMA. 2008;299(2):211-2 scientific concepts and research will surely find more resonance and appreciation in the Anschrift des Verfassers coming century. Prof. em. Dr. med. Anton Valavanis Klinisches Neurozentrum UniversitätsSpital Zürich CH-8091 Zürich e-mail: [email protected]

134 Beiträge von Walter Rudolf Hess zur Schlafforschung

Alexander Borbély

In seiner Laudatio zur Verleihung des Nobelprei- verborgen und nicht vollständig geklärt. Dass ses an W.R. Hess erwähnt Herbert Olivecrona sie vorhanden sind könne zwar lediglich aus auch seine bahnbrechenden Experimente zur ihrer Wirkung abgeleitet werden. Dennoch Schlafforschung. Mittels feinster ins Zwischen- seien sie der Kern des Schlafproblems. Der hirn von Katzen implantierter Elektroden habe Ruhezustand der Sinnesorgane, der Muskeln Hess gezeigt, dass nach elektrischer Stimulation und der psychischen Vorgänge seien lediglich bestimmter Regionen die Tiere sich einrollen Begleitfaktoren, welche den Erholungsvor- um anschliessend einzuschlafen. Aus diesem in- gang in den Geweben erleichterten. Dass dem duzierten Schlaf konnten sie ebenso leicht ge- Schlaf eine für den Ablauf metabolischer Pro- weckt werden wie aus dem natürlichen Schlaf. zesse unterstützende Funktion zukommt wird auch in einer neuen Übersichtsarbeit über die Hess scheute sich nicht eine kontroverse Schlaf- Schlafregulation hervorgehoben.3 So ist die theorie zu vertreten. So beschreibt er in seiner Theorie von Hess auch heute aktuell. 1932 in der Zeitschrift Lancet erschienen Ar- beit den Schlaf als einen vegetativen Prozess, Es ist bemerkenswert, dass Hess den Schlafvor- in welchem das autonome Nervensystem die gang bereits systemphysiologisch betrachtete Aktivität höherer Bereiche des ZNS reguliere.1 und sich damit der vorherrschenden Meinung entgegenstellte. So schreibt er in der Arbeit Hess unterschied im Tagesablauf zwei funktio- von 1932, die Physiologie des ZNS sei bis an- nelle Phasen des autonomen Nervensystems: hin von einer hierarchischen Betrachtungswei- Die auf Interaktionen mit der Umwelt aus- se geprägt gewesen mit dem Cortex als dem gerichtete ergotrope Phase mit Prädominanz Hohepriester. Diese Auffassung sei so verbrei- des Sympathicus („fight and flight“) und die tet, dass sie infrage zu stellen bereits als an- energiesparende, Restitution und Erholung stössig gelte. Es sei nahezu unvorstellbar, dass begünstigende, trophotrope Phase mit Prä- die geistigen Fähigkeiten unter dem Einfluss dominanz des Parasympathicus. Die moderne des autonomen Nervensystems stünden. Hess Rhythmusforschung hat das Konzept von Hess plädiert dafür, sich nicht durch Emotionen lei- wiederentdeckt indem sie den circadianen ten zu lassen und die Verhältnisse unvoreinge- Rhythmus in zwei Phasen des Metabolismus nommen zu betrachten. unterteilt: In der Phase des Wachseins und der Nahrungsaufnahme herrschen katabole Die Untersuchung des durch elektrische Stimu- Stoffwechselprozesse vor, in welchen unter lation induzierten Schlafs bei der Katze wurde anderem Glykogen und Cholesterin syntheti- von Rudolf Hess, dem Sohn von Walter Rudolf siert wird. In der Phase des Schlafs und Fastens Hess, gemeinsam mit Werner Koella und Kon- dominieren anabole Prozesse wie Gluconeo- rad Akert weitergeführt.4 Dieser wendete sich genese und Glykogenolyse.2 Heute wissen wir, später dem Schlaf-EEG des Menschen zu5 und dass diese Abfolge ein Ausdruck des circadia- wurde zu einem Pionier des klinischen EEGs. nen Rhythmus ist, der von den suprachiasmati- Der Schlaf zog auch Christian W. Hess, den En- schen Kernen im Zwischenhirn reguliert wird. kel von Walter R. Hess, in seinen Bann. Er ver- wendete als einer der ersten die transkranielle Hess schreibt, die Mechanismen der Erholungs- Magnetstimulation, um die Erregbarkeit des funktion des Schlafs seien in den Geweben Cortex in den verschiedenen Schlafstadien zu

135 Beiträge von Walter Rudolf Hess zur Schlafforschung verfolgen.6 Zudem war er 1991 Mitbegründer 3. Borbély et al. The two-process model of sleep regula- der Schweizerischen Gesellschaft für Schlaffor- tion: a reappraisal J Sleep Res 25: 131-143, 2016. schung, Schlafmedizin und Chronobiologie. 4. Hess R, Koella WP, Akert K Cortical and subcortical re- cordings in natural and artificial sleep in cats. Electroen- ceph clin Neurophysiol 5: 75-90, 1953 So war Walter Rudolf Hess mit seinen ein- zigartigen experimentellen Arbeiten und 5. Hess R The electroencephalogram in sleep. Electroen- ceph clin Neurophysiol 16: 44-55, 1964 wegweisenden Theorien ein Pionier der 6. Hess CW et al. Excitability of the human motor cortex Schlafforschung und Wegbereiter einer For- is enhanced during REM sleep. Neurosci Letts 82: 47-52, schungsdisziplin, die durch seine Nachkom- 1987 men und Schüler erfolgreich weiterentwickelt wurde. Prof. em. Dr. med. Alexander Borbély Universität Zürich Literatur Neuwies 28 1. Hess WR The autonomic nervous system. The Lancet 8700 Küsnacht 1932; Dec 3: 1199-201; Dec 10: 1259-1261 e-mail: [email protected] 2. Bass J, Takahashi JS Circadian integration of metabolism and energetics. Science 330: 1349-1354, 2010.

136 W. R. Hess, the ophthalmologist

Alfred Huber

Würdigung der wissenschaftlichen Beiträge von W. R. Hess aus seiner frühen beruflichen Tätigkeit als Ophthalmologe veröffentlicht durch den Zürcher Ophthalmologen Professor Alfred Huber (†) in der Zeitschrift Experientia 38 (1982), pp. 1397-1400.

137 W. R. Hess, the ophthalmologist

138 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

139 W. R. Hess, the ophthalmologist

140 Hess’s experiments on the diencephalon and hypothalamus in the light of modern neuromeric genoarchitectonics

Luis Puelles

Vortrag des Neuroanatomen und Neurobiologen Prof. Luis Puelles, Universität Murcia, Spanien, gehal- ten am 6. Jahressymposium des Klinischen Neurozentrums des USZ zur Würdigung der neurowissen- schaftlichen Leistungen von Walter Rudolf Hess

Seen from the viewpoint of a neuroana- (under ventral thalamus, dorsal thalamus and tomist, the extensive mappings by W.R. Hess epithalamus). Hess divided the hypothalamus of places in the cat forebrain where charac- into functionally distinct ‘anterior’ parasym- teristic physiologic visceral and somatic reac- pathetic and ‘posterior’ sympathetic parts, tions are obtained by stimulation show inte- which correspond to the preopto-supraoptic resting features (the publications Hess, 1932, and tubero-mamillary regions, respectively. 1948, 1954, 1957, 1968 recapitulate his physio- logical work on the brain). He obviously did Probably induced by practical considerations not intend to revolutionize neuroanatomy, as regards the best route of approach for his though he did advise his colleagues in phy- electrodes, which penetrated deeply into the siology that more attention to neuroanatomy brain, Hess systematically analyzed his stimu- was essential. Following the suggestion of his lation sites in sections oblique to the colum- colleagues R.Jung and E. Weisschedel some nar diencephalon model (Hess, 1932). These 70 of his experiments finished with localized were oriented roughly parallel to the plane electrocoagulation and were subjected to in- that connects the anterior and posterior com- dependent anatomic analysis with the Marchi missures, as identified in paramedian sagittal degeneration technique by his collaborators sections. This plane is also roughly parallel to V.M.Bucher and S.M.Bürgi (e.g., Bucher and the optic tract coursing from the chiasma into Bürgi, 1945, 1951, 1955a,b; Bürgi and Bucher, the superior colliculus. 1960 [with prologue of Hess and full list of reports]; Verena Bucher was the laboratory These oblique sections turn out to be approxi- technician of Hess from 1929 until his retire- mately horizontal with regard to the modern ment in 1951, but she was still active at the alternative conception of the subtelencepha- Zürich department of physiology in 1960; she lic forebrain which is embodied within the also collaborated briefly with W.J.H.Nauta so-called prosomeric model. The latter was in the meantime [2 papers]; S.M. Bürgi wor- proposed by Rubenstein and myself in 1993- ked with Hess in several papers, some on his 1994, and has been developed further since own] –collaborated with V.Bucher between then (latest update in Puelles et al., 2012 and 1945-1960, and became Professor of Neuro- Puelles and Rubenstein, 2015). This model logy at the Univ. of Bern in 1958). As regards defines on the basis of gene expression and localization, Hess interpreted his data strictly patterning studies a forebrain axis bent at the within the then standard paradigm on the di- cephalic flexure (under the midbrain), which encephalon (Herrick’s, 1910 columnar model), then runs successively through pretectum, in which the forebrain axis ends in the telen- thalamus, and prethalamus into the hypotha- cephalon and the hypothalamus figures as a lamus, where it ends roughly behind the op- ventral longitudinal diencephalic component tic chiasma (the axis does not extend into the

141 Hess’s experiments on the diencephalon and hypothalamus in the light of modern neuromeric genoarchitectonics telencephalon). This conceptual change in the natomic usage of his contemporaries, whose axial dimension of the forebrain compared to conventional coronal sections cut obliquely the classic columnar model is fundamented in the same tracts, rendering them hardly use- causal analysis (accrued developmental pat- ful as landmarks). Accordingly, his schematic terning data suggest that the axial dimension sagittal section maps of the stimulation sites is generated strictly by the notochord, and where given reactions were obtained can be does not involve the prechordal plate). This easily reinterpreted within the modern pro- leads first to conceiving the hypothalamus someric model, thanks to the recorded land- jointly with telencephalon as the rostralmost mark tracts. A better model typically reinter- forebrain compartment, which accordingly prets older results in an advantageous way, is placed rostral to the diencephalon proper and it is remarkable that Hess’s data from the rather than under it (the telencephalon is a thirties and forties acquire new depth and dorsal vesicle that bulges out of the hypo- pregnancy when studied from this recent thalamus). The prosomeric model corrobora- viewpoint. tes molecularly, and reintroduces, the basic division of the forebrain neural wall into alar In the first place, his reference to anterior and and basal plates, which was first proposed by posterior hypothalamus (e.g., Hess, 1957) im- W.His (1893, 1895), but was later disdained plies distinguishing alar from basal hypotha- by Herrick (1910) and his followers. Further, lamus in the new model (Puelles et al., 2012; it forces reinterpreting the classic ‘longitudi- Puelles and Rubenstein, 2015). A number of nal’ diencephalic subdivisions as transversal other differential mapping data within the di- anteroposterior subdivisions, namely as neu- encephalon proper (caudal to hypothalamus) romeres (as had already been suggested by also refer clearly to alar-basal differences in some classic authors). The new era of descrip- localization. Secondly, a number of localized tive and experimental molecular brain data ‘functions’ correlate with specific forebrain (brain genoarchitectonics), which started in neuromeres (i.e., some reaction types are lar- the eighties, has brought in forty years very gely contained within specific forebrain neu- weighty evidence in support of the prosome- romeres). Thirdly, some mapped distributions ric model, which seems widely assumed now of functional properties make use of the ra- in the field of developmental neurobiology as dial dimension within a neuromeric field (i.e., well as in comparative neuroanatomy. Clinical differences map onto the radial dimension colleagues are slowly starting to adapt their extending from the periventricular stratum thinking to this fundamental change of para- into the superficial stratum across interme- digm, which enhances significantly our mor- diate elements). phologic understanding of the brain. These new topologic interpretations of the The prosomeric model typically visualizes function-structure relationships discovered some well-known forebrain tracts, such as the by Hess in the light of the molecularly and posterior commissure, the retroflex tract, the causally based prosomeric model of the fo- mamillothalamic tract and the fornix tract as rebrain add renewed impact to Hess’ experi- transversal landmarks running parallel and mental contributions. This goes well beyond adjacent to interneuromeric boundaries. It so the salience emerged under the non-causal happens that Hess’s oblique sections intersect neuroanatomic dogma of the columnar mo- optimally in the horizontal plane these inter- del, which already won him the Nobel prize in neuromeric transverse tracts, which he clearly 1949. It is to be expected that modern func- used as his main topographic references (in tional and anatomic studies of autonomic this he departed from conventional neuroa- and reflex/instinctive regulatory mechanisms

142 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich in the brain will again examine in detail and Hess, W.R. (1954) Das Zwischenhirn (2. Aufl.) Basel: B. try to reproduce Hess’s data with the help of Schwalbe Co., pp 1-218. modern molecular-based methods. One is re- Hess, W.R. (1957) The Functional Organization of the Di- minded of the use made by Kepler of the un- encephalon. (English transl. of ‘Das Zwischenhirn’ 1949 [1. usually precise earlier data of Tycho Brahe for Aufl.], ed. by J.R.Hughes), New York/London: Grune and Stratton. pp 1-180. his revolutionary analysis of the solar system. Hess, W.R. (1968) Hypothalamus und Thalamus. Experi- mental Dokumente. (2. erweit. Aufl.). Suttgart: G. Thieme, pp 1-77. References His W (1893) Vorschläge zur Eintheilung des Gehirns. Arch Bucher, V.M. and Bürgi, S.M. (1945) Untersuchungen über Anat Entwickl-Gesch Jahrg 1893:173-179. die Faserverbindungen im Zwischenhirn und Mittelhirn der His W (1895) Die Anatomische Nomenclatur, Nomina Ana- Katze. Confinia Neurol. 6:317-340 tomica. Neurol Suppl Bd, Arch Anat Entwickl-Gesch Jahrg Bucher, V.M. and Bürgi S.M. (1950) I. Fiber connections of 1895:155-177. the tectum opticum. JCN 93:139-172 Puelles, L. and Rubenstein, J.L.R. (1993) Expression patterns Bucher, V.M. and Bürgi S.M. (1952) II. Fiber connections of of homeobox and other putative regulatory genes in the the pretectal region and the posterior commissure. JCN embryonic mouse forebrain suggest a neuromeric organi- 96:139-178 zation. Trends Neurosci. 16: 472-479. Bucher, V.M. and Bürgi S.M. (1953) III. The supraoptic de- Puelles L, Martinez-de-la-Torre M, Bardet S, Rubenstein, cussations. JCN 98:355-380 JLR. (2012) Hypothalamus. Chapter 8 in “The Mouse Ner- Bucher, V.M. and Bürgi S.M. (1953) IV. The ansa lenticula- vous System”. Watson C, Paxinos G, Puelles L (eds). Acade- ris, pars ascendens mesencephalica, with observations on mic Press/Elsevier 2012; pp 221-312. other systems ascending from and descending to the mes- Puelles L., and Rubenstein J.L.R. (2015). A new scenario of encephalon. JCN 99:415-436 hypothalamic organization: rationale of new hypotheses Bürgi, S.M. and Bucher, V.M. (1960) Markhaltige Faserver- introduced in the updated prosomeric model. Front.Neu- bindungen im Hirnstamm der Katze. Monograph. aus dem roanat. doi: 10.3389/fnana.2015.00027. In: Alvarez-Bolado, Gesamtgeb. der Neurol. u. Psychiat. Heft 87, Berlin:Sprin- G., Grinevich, V., Puelles, L., (eds) Development of the Hy- ger Verlag., pp. 1-127 pothalamus. Lausanne: Frontiers Media. doi: 10.3389/978- 2-88919-634-0. Frey, E and Bucher, V.M. (1947) Degenerationstudien über extrapyramidale Bahnen und Zentren nach Ausschaltungs- Rubenstein, J.L.R.; Martínez, S., Shimamura, K., Puelles, L. versuchen. Schweiz. Arch. Neurol. u. Psychiat. 60:80 (1994) The embryonic vertebrate forebrain: the prosomeric model. Science 266: 578-580. Herrick CJ (1910) The morphology of the forebrain in am- phibia and reptilia. J Comp Neurol 20:413-547. Hess, W.R. (1932) Die Methodik der lokalisierten Reizung Anschrift des Verfassers und Ausschaltung subcorticaler Hirnabschnitte. Leipzig: G. Prof. Dr. med. Luis Puelles Thieme, pp 1-122. Director Department of Human Anatomy Hess, W.R. (1948) Zwischenhirn und Motorik. Helv. Physiol. University of Murcia Pharmacol. Suppl. V, pp 1-89. Spain e-mail : [email protected]

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5. Von der Neurowissenschaft zur Neurophilosophie

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Brain and Consciousness: BRAINA DiscussionAND CONSCIOUSNESS: About the FunctionA DISCUSSION of the Brain ABOUT THE FUNCTION OF THE BRAIN* Walter Rudolf Hess, Hanns Fischer

NachdruckW. R. HESS einerand DiskussionH. FISCHERf zwischen Prof. W.R. Hess und Prof. H. Fischer, ehemaligen Direktor des Pharmakologischen Instituts der Universität Zürich zum Thema Hirn und Bewusstsein, erschienen in Perspectives in Biology and Medicine 17(1): 109-118, 1973

Dr. Hess to Dr. Fischer I should like to add a few thoughts to our discussion. At the time when we both were members of the medical faculty, your opinion struck me as concurring in many ways with mine. It soon became evident, however, that in questions of philosophy of life we differ considerably. This refers to the so-called ultimate questions which every scientist is bound to face sooner or later and which may even cause him conflicts of conscience. I shall try to point out where our respective opinions diverge. You are relying on ancient traditions, whereas I have been brought up in the belief of, and am working with, the laws of natural science. In other words, my opinions are based on observation and experience. In particu- lar, I have always respected the limits set by the organization of the brain, and I hold that it is basically impossible to come to valid conceptions about matters which are inaccessible to our means of perception. Here is an example. For me, the contents of visual perception are represented by the excitation of some retinal elements and their propagation through fiber tracts to the occipital neuron populations. The fiber paths can be seen under the microscope, the propagation of the excitation can be fol- lowed, and their arrival at the occipital area determined by electrical re- cording. The transition from neuronal activity to the subjective experi- ence is not, however, accessible to rational understanding. It is, neverthe- less, a reality. You, for example, suggest that language is an objective replica of a feeling or of a train of thought. To my mind, the represen- tation is merely formal and is comparable to a photograph of a landscape with mountains, waterfalls, lakes, flowers, etc., in short, of those features * Editor's note,—Drs. W. R. Hess and H. Fischer have carried on a friendly debate on the functions of the brain for some years. They have not come to agree on all points. Each desires to publish his present position on brain function.—D. J. I. t Professor Hess died August 12, 1973. His address was 6612 Ascona, via Gabbio 6, Zurich, Switzerland. Professor Hans Fischer, Witellikerstrasse 60, 8702 Zollikon, Zurich, Switzerland.

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147 Brain and Consciousness: A Discussion About the Function of the Brain

of an object displayed in space which are apt to be represented in the photograph. I refer to the short monograph Psychologie in biologischer Sicht [1, 2] and to an atlas [3, pp. 66-67] which show how I arrived at the experi- mental basis of our subject. Illustrations 201 and 203 [3] should be studied. I consider the evidence adequate to show that the electrical stimulation applied in a defined area of the hypothalamus is able to produce behavior which closely resembles that which under natural cir- cumstances accompanies rage and aggressiveness. The question arises here as to whether this behavior is the concomitant of subjective experience. If one considers the further information that a cat in this situation turns toward the experimenter and purposively hits his hand, or even prepares to jump at him, one cannot but assume that the appropriate subjective experience is integrated into the whole purposeful behavior. It is likely to be substantially the same as in the case of a dog approaching a cat, or of a man engaging in a fight with a counterpart. When cat or man fights back and experiences the corresponding feelings, our interpretation reaches the critical point. The excitation of defined cerebral areas is a fact; the neurophysiological organization of behavior is accessible to rational un- derstanding. How the electrical excitation of nervous matter is converted into subjective experience, on the other hand, is an open question. You hold the opinion that the answer must come from philosophical thought. In my own view the gap in our knowledge must be admitted and ex- plained by the fact that our brain is not organized to analyze its own mechanisms, although it is able to assess certain results of its activity. In other words, its organization is aimed at only a limited number of perfor- mances out of all possible performances. Let us take the other example of electrical stimulation which induces the cat to purr, as if lying cozily in her mistress's lap. It is equally inter- esting to note how vigilant behavior can be suppressed as in natural sleep [3, p. 64], and how the drive to eat or eagerly lap up some liquid can be produced by stimulation. These are behavior patterns which are natu- rally induced by drives, accompanied by feelings of hunger and thirst, and yet are clearly assignable to defined areas in the stimulation experiment. Let me repeat that I abstain from discussing possible different inter- pretations of situations or processes which are beyond rational compre- hension. I am, however, firmly convinced that the existing functional organization of the brain should be given preference over other possible explanations.

Dr. Fischer to Dr. Hess I fully agree when you say that "the transition from neuronal activity to the subjective experience is not, however, accessible to rational under-

110 W . R. Hess and H . Fischer · Brain and Consciousness

148 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich standing [but is] nevertheless a reality." The visual process described by you corresponds exactly with this concept. I would like to go one step further and say that not only is the process which enters our consciousness an experience, but the entire neuronal process from the periphery of the sensory nervous system (the receptor) to the central nervous system is accompanied by an active system (inaccessible to our consciousness) which mediates between the stimulus of the receptor and the adequate content of the consciousness. This is of course a hypothesis, and I see no way of verifying that the process is transcendent. By transcendent I mean processes which are inaccessible to our causal-analytical methods of cog- nition. There are other "solutions" which are accessible to us, such as compar- ing the nervous process to an electrical one. The electrical process, how- ever, does not lead to such knowledge as the objectively colored experi- ences in our consciousness, transmitted by the sense organs. As an example I choose the synaptic function. The neurobiochemical synaptic process is rather well known, but we shall probably never understand how the synapse transmits the specific content of the impulse which becomes con- scious in the brain. The content of the sensory impulse which reaches the sense organ and thus alters its physiologic state cannot be replaced by an electrical explanation. We find ourselves in the same position as the molecular biologist who for similar processes in biochemical genetics uses the term "information," which suggests the participation of a "thinking being" in the genetic process. We could presume the same in the realm of the senses, possibly even in every neuronal process, and thus lose the grounds for exact understanding. This indicates that in every living organism one meets with processes which manifest themselves as real but remain hidden to our limited cognition. We can, however, assume that the entire, extremely compli- cated nervous process, apart from the functions explainable by causal- analytical methods, also takes part in those other functions which are inaccessible to man and are therefore transcendental or metaphysical. The consequence of this conception is that in the living organism we have to reckon with the fact that its biochemical and physical nature is linked with qualities which, although they have the status of reality, are at the same time transcendental. Molecular geneticists have followed (more or less unconsciously) the same direction, with the conviction, however, that they are dealing with physically accessible processes. But in their interpretation they use words which indicate with all desirable clarity that mere biochemical and bio- physical terms are insufficient for the explanation of processes transcend- ing biochemistry. They therefore use expressions like "information,"

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"messenger RNS," "code," etc., for processes inseparably connected with the biochemistry of the living cell but biochemically unexplainable. We are facing something incomprehensible, the reality of which we cannot deny and which is part of the innermost events of life. But it is precisely this incomprehension which may produce a creative force in us, possibly only a longing, leading beyond that which can strictly be experienced. The creative force often lies in the direction of philosophy, art, and religion. I sympathize most with your psychology and philosophy when you go beyond what is now explainable and name it, and therefore acknowl- edge its existence. Wre do not know for instance what an engram is and how ecphorization occurs; yet both expressions have a real background in cerebral physiology although we cannot imagine their nature. You hold that feelings and often powerful emotions which are of tran- scendental character are a privilege of the human being. As they are in- accessible to sense perception and examination, they are also beyond judgment from a biological view. From this standpoint we are bound in space and time. I would like to answer this. Metaphysics plays a much more important role in the natural sciences than is generally accepted. The most im- pressive example is the partial transcendence of genetic processes which is very impressive on all levels of life. In the future we shall possibly have to operate with even more expressions which lie on the borderline between natural and spiritual sciences. The "eternity" of germ plasma is one of these: we do not know what eternity is. The crossing point of objective understanding and subjective experi- ence is our consciousness, which we consider a function not to be analyzed scientifically. The reality of our experiences proves that our brain is capable of apprehending and digesting more than the perceptions gained by scientific methods. It is by all means in agreement with the reality of experiences when you say that the living brain has command over still another dimension which cannot be defined by itself. Here you hit the crucial point. There is a dimension which is inaccessible to our cognition, unlike the cerebral processes which are understandable by causality and analysis. This in- accessible dimension has a real value which cannot be controlled by us. The metaphysical dimension is just as familiar to our thinking as the physical one. As a rule we consider it as belonging to the field of spiritual sciences. Molecular-biology geneticists as well as your brain experiments have shown that the natural sciences need transcendental expressions as well in order to describe transcendental facts. There are enigmas which we sometimes describe but cannot understand. Now to your observations on language as an organ of cognition. According to you, language cannot

112 W. R. Hess and H. Fischer · Brain and Consciousness

150 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich state an objective cognition: it is a symbol for communication without reality, with symbolic value only. From my point of view, thinking is objectified in language. If language has only symbolic value we cannot recognize anything but can produce a likeness of reality expressed in letters and numbers. I agree with you that no identity does or can exist between the expression of a thing and the thing (or organism) itself. We cannot recognize the objective reality of an organism (as in Platonism we cannot recognize the "True Being" of a thing). In this respect I have to admit that language represents a world of symbols, a kind of explanation by signs, which man over a long period of time has adopted and by which the anthropomorphic character of the language is secured. But as scientists we behave as if, with the interpretation of factual findings, we would have experienced and characterized the real world. The natural scientist in particular ignores the world of symbols and be- lieves in the reality of his characterizing word. Language is the only way of dealing with "reality"; it is the only way of thinking by which we can approach reality without ever attaining an identity. This also applies to the language of mathematics. If we want to assign truth to philosophy we find ourselves in the same position as Plato with his parable of the cave: we do not discern the "True Being" but only its image in its shadow. If we were able to discern the "True Being," we would possess divine powers. If language is pure symbolism, our cognition also is pure symbolism and loses its content of truth to us. A symbolic language could be some- thing immensely beautiful, especially in poetry and in art, but not in the natural sciences. If the language of the natural sciences dealt with symbols only, we could only build symbolic houses, machines, tools, etc. We therefore have to say that thinking is objectified by language. Only language enables us to think as millenniums of human culture testify. The signs of language alter with their meaning. The important thing is that language is the basis for mutual understanding. In this sense language signs are conventions that are unable to reproduce the true content of a thing in its true being. The truth remains humanly limited. Only our yearning for the "thing-in-itself" reaches beyond it. We are children of an ideality which we will never attain, and perhaps in this lies our happiness. Used as symbol, language loses its credibility, especially concerning logic and causality. I do not, however, want to reject the expression "symbol": language represents the mediator between thinking and self- assertion. The self-understanding of thinking occurs in linguistic symbols. But language contains its own value of cognition. It also copies reality in the natural sciences; here I agree with you. It is a way that leads to the

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truth that is possible for (or permitted) us. This truth will always remain relative, but it obtains a certain general validity by the actual consensus omnium, and thereby it may approach real objectivity.

Your brain experiments have opened a psychophysiologic world that explains many things which were inaccessible before. The brain and its functions are thus much clearer to us. In the previously described experi- ments, you have asked the crucial question of whether the electric impulse in the hypothalamus is produced without the subjective experience (as, for instance, the fit of rage of the cat), and you add that how the stimu- lation of a nervous element becomes a subjective experience within us remains an open question. This means that in your opinion, as in mine, the subjective experience also represents a nervous process, but you forgo explaining how the subjective experience comes about and do not comment on other interpretations of inconceivable situations or processes, although you prefer with conviction the significance of a definite func- tional organization of the brain. And to this you add that philosophical consideration may yield an answer. I do not hesitate to consider the cat's fit of rage caused by hypothalamic stimulation as a subjective experience for the cat; the directed aggressive movements especially indicate this. The situation to me seems similar to an external excitation of rage (the dog). The process of defense toward aggression is too complicated to be merely automatic; it is more likely a willful experience. We certainly agree on one point: there is no thinking and no experi- ence whatsoever without a living brain, and all the experiences of the world and the individual are connected with neural processes of the brain. But the problem of our main topic, the brain and its functions, is much deeper. Every process of the brain includes something unexplainable. Generally speaking, the discernible cerebral process that can be measured somehow represents only the covering of a process inaccessible to us. We do not know, for instance, how a stimulus of the senses is transmitted from the nerve periphery to the central nervous system, how it enters our consciousness, or how a mental process passes over the threshold of consciousness. It may be presumptuous to say that all our experienced impressions, mental acts, emotions, and everything that pertains to what we call the soul contain, according to our comprehension, transcendent elements inaccessible to our analytic understanding. From my point of view, this is also the way to interpret your animal experiments leading to experiences. Not much may be gained thereby for our understanding, but the en-

114 W.R.Hess and H.Fischer · Brain and Consciousness

152 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich tire problem has been raised to another level. The "experience" which resists a causal explanation has been given the status of a process that philosophical thinking can conceive or at least describe. Thinking of synaptic transmission, you said that in the question of quasi-synaptic transmission one cannot at the same time coordinate a pat- tern and analyze the mechanisms of transmission [4, pp. 933-965]. The situation would therefore be similar to the observation of an atomic com- plementarity process in physics. This is correct insofar as we can indeed study the synaptic process, including corresponding biochemical altera- tions, but not the "pattern," meaning, I suppose, the excitatory pattern produced in the nerve which enters consciousness in the brain as an impulse. We do not know in what condition this pattern, having been present throughout the whole path of the nervous system, finally becomes conscious in the brain. And yet it is obvious that this pattern (experience, emotional pattern), somehow already on the way to the brain, must possess a reality even if it is inaccessible to us and therefore transcendent. Without becoming unscientific, we have to learn to accept the fact that in all nervous and mental processes features occur which are transcendent to our knowledge. They precede what then becomes available in a greater or lesser variety of choice to our consciousness. Consciousness and memory represent at the same time possibilities for the understanding of the connections of the meaning of a process. No animal or human being can live without these connections of the meaning of physical and metaphysi- cal processes which are only indirectly perceivable to us, nor could animals or human beings live without these transcendental processes. On principle, you prefer not to comment on incomprehensible situations and processes even if the brain should be involved. And yet you confess to the classical scientific view, which you have gained by your impressive experimental brain research. Nevertheless, based on your enor- mous experience through brain research, you say that the living brain controls still another dimension which cannot be defined by itself. Even more positively, you say that in the transmission of neuronal organization another force of a different kind might take part in the content of con- sciousness. We are, however, incapable of ascertaining which cerebral process leads to the result which enters our self-consciousness as reality, or as our "truth." First of all, our limits of perception lie within the limits of our understanding of the cerebral process, as far as we are dealing with the recognition of the way of transmission of psychic contents passed on by nerves which "appear" in our consciousness. Second, we do not know the process or the "transition" leading from the nervous process to the consciousness. Third, the content of consciousness can only be expressed by words, that is, in a language of signs whose identity with the process of

Perspectives in Biology and Medicine · Autumn 1973 | 115 153 Brain and Consciousness: A Discussion About the Function of the Brain

consciousness is not ascertained; thus, the signs (according to you) prob- ably have a symbolic value only and can only transmit one truth, which is the human truth. You say that the living brain controls another dimen- sion not to be defined by itself, so this could be an idea which aims at a hypothetical explanation for the inexplicable (the transcendent) of the living brain and which, as an idea, probably is of great future importance.

You say that you are strongly opposed to the notion of an "electric brain," for the living brain controls by its capacity of experience a special dimension. There I fully agree with you. The notion of an electric brain comes from a group of scientists ready to subject biologic functions to a mechanistic system, thereby entirely misunderstanding or disregarding specific biological dimensions. Among these mechanists (Lam- metrie also was a mechanist in his "homme machine") we find a consider- able number of molecular biologists. This can be understood insofar as they acknowledge only the physicochemical molecular process advanced by themselves. Thus, they are unaware of the totality of the animal organiza- tion. I confirm your opinion on Darwinism regarding evolution: Darwinism is not the last word indeed, and the thoughts on evolution by natural selection are still in proof. I fully agree with you that there are indeed possibilities to take in certain experiences but that the organization of our brain is responsible for our not being able to comprehend their cause. We have for instance musical experiences of a spiritual nature. They are just as "true" as our understanding of the production of the sound, of the pitch of the tone, that is, waves of a specific quality, which we can measure quantitatively in various ways. Their causal reality is undoubted and can be repeated experimentally at will. It has, however, nothing in common with the musical experience as such, for this occurs at an entirely different level of (perceptible) understanding than the physical event. There is no doubt that we daily meet with psychic and emotional experiences, as for instance when we read a work of poetry with a fully understandable content whereby, however, other forces become effective and only they lead to the poetical experience (e.g., Shakespeare's Hamlet). If this were not so, all poetry would be senseless. We are confronted with the experience of an inner reality of a spiritual nature. You also comment on the question of teleology when you say that under normal circumstances an instinctive tension underlies purposeful behav- ior. I wonder whether ideological behavior can be called an instinct. Teleology is a basic quality of all living things. We can observe and

116 I W. R. Hess and H. Fischer · Brain and Consciousness

154 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich describe its effect in fertilization, in multiplication, in growth, etc., with- out knowing the underlying forces of the event. Its reality is beyond any doubt. It would be senseless to speak of teleology connected with the inorganic world, crystalline growth, for instance. The sphere of activity of teleology includes an enigma which accompanies the living, psychoso- matic process at every single moment. Therefore, you justly say that teleology is as real in the realm of the liv- ing as gravity is in the inorganic world. This thought is followed by another with which I also fully agree: that there are still so many things existing in the world that are beyond understanding. When furthermore you state that in fact we do not reach a convincing argument—tradition on one side and pure natural science on the other—this formulation of our viewpoints may be somehow misleading. If, according to my conception, the natural sciences are drawing near the spiritual sciences, it does not mean tradition but future, because I think that even pure natural science is not pure natural science, but also contains spiritually explainable or to a certain degree recognizable elements of a metaphysical character tran- scending our real perception. At the same time, it represents real processes (e.g., consciousness, teleology, certain elements of the genetic processes and growth, and maybe also evolution) [5]. We are now leaving the grounds of exact natural science and are trying, at least if only tentatively, to understand the uniqueness of the being and behavior of the living. We are trying to understand areas where natu- ral science gives us no real help. This selfness, which undoubtedly exists in all living things, is of future importance for the comprehension of the living organism, or altogether with regard to cognition of life, which we want to serve by natural and spiritual sciences. We have seen that natural science, in its distinct recognition of the mental capacities of man, operates with classical methods which have resulted in a century of tremendous success in biological research. You have indicated that the scope of possible perception might be enlarged, by saying that the living brain controls also another dimension undefinable by itself which, as I conceive it, can be experienced with the same intensity as the purely scientific part of life but which has an entirely different structural truth. This part is not less "real" than the world of natural science in its classical forms and is somehow inseparably connected with it. An "unperceivable reality" is in the sense of natural science a paradox but not unrealistic. I am convinced that no living organism can exist with- out the partly inaccessible realities which might be described but not fully understood. Future scientific research might be able to gain a certain insight into the teleological nature of the living being if it will not shrink from paying attention to events attributed to the more spiritual experience.

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Dr. Hess to Dr. Fischer I am strictly opposed to the conception of the brain as an "electronic instrument"; the living brain encompasses, with its capability for sub- jective experience, a particular dimension. Such a confession might be considered heretic. I hold, however, that every man's honest conviction must be respected but that one also should unabashedly proclaim one's own. When I look for reasons for the different convictions of the two of us, I find them—apart from different personality structures—in exogenous influ- ences, in particular those that took effect in earliest youth when we were lacking the capability of critical assessment. A permanent flow of influences from the environment was thereafter superimposed. Darwinism, which you are referring to, is of restricted validity in my own view also; it does not, in all events, give an explanation for the phenomenon of sub- jective experience. Years ago, when discussing these problems with Profes- sor A. Fleisch, I ventured the thought that a particular, hitherto unknown force might be involved. Fleisch rejected this idea firmly, and I left the question open, unprovable as the hypothesis is. It is, however, not an unthinkable one, since transformation of one form of energy into another is quite common (one only need recall the transformation of water pressure into electricity). To sum up, I hold that we have to content ourselves with the recog- nition that much exists and evolves in this world which is not accessible to our comprehension, since our cerebral organization is primarily devised so that it secures survival of the individual in its natural surroundings. Over and above this, modest silence is the appropriate attitude, I believe, in connection with the problems under discussion.

REFERENCES 1. W. R. Hess. Biology of mind. Chicago: Univ. Chicago Press, 1964. 2. ------. In : Psychologie in biologischer Sicht. 2d ed. Stuttgart: Georg Thieme Verlag, 1968. 3. ------. Hypothalamus und Thalamus: Experimental-Dokumente. 2d ed. Stutt- gart: Georg Thieme Verlag, 1968. 4.H. Fischer. Vergleichende Pharmakologie der Überträgersubstanzen in tiersys- tematischer Darstellung. Berlin: Springer-Verlag, 1971. 5. ------.Naturwissenschaften, 59:425, 1972.

118 I W.R.Hess and H. Fischer · Brain and Consciousness

156 Causality, Consciousness, and Cerebral Organization

Walter Rudolf Hess

Eine der letzten Publikationen von W.R. Hess über seine Ansichten zur Cerebralen Organisa- tion erschienen in Science, 158, 3806: 1279-1283, 1967

157 Causality, Consciousness, and Cerebral Organization

158 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

159 Causality, Consciousness, and Cerebral Organization

160 Biological Order and Human Society

Walter Rudolf Hess

Anlässlich der 100-Jahr-Feier des Geburtstages von Walter Rudolf Hess, gab sein Schüler und Direktor des Hirnforschungsinstitutes der Universität Zürich, Konrad Akert, eine Festschrift mit dem Titel «Biological Order and Brain Organization: Selected Works of W. R. Hess», Springer-Verlag, 1981, heraus. Darin wird eine Auswahl von 17 her- ausragenden Werken aus dem gesamten Spektrum der wissenschaftlichen Tätigkeit von Hess (304 Publikationen), einschliesslich seiner Autobiographie und Bibliographie, wiedergegeben. Hier wird das erste Kapitel «Biological Order and Human Society», welches grundlegende neurowissenschaftliche Konzepte von Hess verdeutlicht, wie- dergegeben.

Aus dem Vorwort von Konrad Akert: The thread that winds through his seemingly heterogeneous topics is the concept of „Biological Order“ on the one hand, and that of the Leistungs- und Erfolgsbezogene Physiologie, on the other. Hess became aware of the role of biological order in his early work on hemodynamics, and he later recognized the same principle as the governing factor in motor coordinati- on, psychic functions, and even in the behavior of human society.

161 Biological Order and Human Society

[email protected] Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

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[email protected] 6. Das Vermächtnis des Hirnforschers Walter Rudolf Hess

175

Das Vermächtnis von Walter Rudolf Hess und kritische Gedanken zur gegenwärtigen Situation der Neurowissenschaft

Anton Valavanis, Klinisches Neurozentrum USZ

«Ich glaube wir müssten uns darin finden, deter Kritik, geäussert seitens prominenter dass in unserer Welt vieles sich entwickle und Neurowissenschaftler, wurde das Projekt mit existiere, was unserem Verständnis nicht zu- gewissen methodologischen Anpassungen gänglich sei; denn unser Hirn sei in erster Linie fortgeführt2,5,6,8,13,17,18. darauf angelegt, das Überleben des Individu- ums in seiner natürlichen Umwelt zu sichern. Begonnen hatte diese von Optimismus be- Es sei nicht so organisiert, dass es seine eigene gleitete Entwicklung mit dem Ausrufen des Arbeitsweise völlig analysieren könne. Jen- ‘Jahrzehnts des Gehirns’ durch den damaligen seits der Effekte seiner Tätigkeit, die sich er- Präsidenten der Vereinigten Staaten George forschen lassen, sei bescheidenes Schweigen H. W. Bush im Juli 1990. Ziele waren die um- die angemessene Haltung». fassende Erforschung des Gehirns und seiner Erkrankungen, was zu deren Heilung oder Mit diesen Worten beendete Walter Rudolf mindestens zu deren Linderung führen soll- Hess eine längere und kritische Diskussion te30. Eine Beurteilung der erzielten Ergebnis- über die Funktion des Gehirns mit seinem Kol- se am Ende des Jahrzehntes veröffentlicht in legen Professor Hans Fischer, dem damaligen der Zeitschrift Science fällt eher bescheiden Direktor des Pharmakologischen Institutes der aus: «Even when a cure or effective treat- Universität Zürich, welche im Jahr 1973 statt- ment does not yet exist, as in Alzheimer‘s fand und in der Zeitschrift Perspectives in Bio- disease, the attention attracted by recent logy and Medicine veröffentlicht wurde12. breakthroughs such as those in stem cell re- search and the genetics of Parkinson‘s disease Vier Jahrzehnte später, im Oktober 2013, kün- has induced a clear expectation that treat- digte die Europäische Kommission die Lancie- ments are not too far away. Identification of rung des 1 Milliarde Euro schweren ‘Human the pathophysiology underlying chronic brain Brain Project’ an11,14,20. Dieses ambitiöse Projekt and mind disorders has also helped to reduce hat sich als Ziel die Entschlüsselung der struk- the stigma attached to these conditions» 31. turell-funktionellen Organisation des mensch- Dennoch hat diese Initiative die Proklamation lichen Gehirns und seiner Erkrankungen auf ähnlicher Projekte in verschiedenen Ländern der Grundlage einer zu schaffenden extensi- wie Japan, China, Indien, Korea, Deutschland, ven informations-technologisch gewichteten u.a. ausgelöst und im Jahr 2013 zur Ausrufung Forschungsinfrastruktur gesetzt. Das Projekt des oben erwähnten Europäischen Human wurde von der ETH Lausanne (EPFL) initiiert Brain Project geführt. Wenige Wochen später und beteiligt mehr als 100 Institutionen in 24 folgte die Proklamation des mit rund 3 Milliar- Ländern1,20. Angesichts seines Ausmasses, sei- den Dollar zu finanzierenden Brain Activity ner Zielsetzung und der ausgelösten Euphorie Map Project oder BRAIN Initiative durch den in breiten Teilen der neurowissenschaftlichen damaligen Präsidenten der USA Obama mit Gemeinschaft wurde es, in Anlehnung an den dem erklärten Ziel unser Verständnis über von Galison 1992 eingeführten Begriff ‘Big das menschliche Gehirn zu revolutionieren Science’, mit dem neugeschaffenen Begriff (!)32. Diese Initiativen wurden sogar als „mega ‘Big Neuroscience’ gekoppelt3,7,22. Trotz begrün- brain projects“ definiert. In einer umfassen-

177 Das Vermächtnis von Walter Rudolf Hess und kritische Gedanken zur gegenwärtigen Situation der Neurowissenschaft den Analyse der Konzeption dieser Projekte hingewiesen hat. In der Beurteilung der Dis- kommt Poo (2014) zu folgender kritischen sertationsarbeit von Monakows 1881 hielt er Schlussfolgerung: „The mega brain projects fest: «…und gratuliere ich Ihnen herzlich für announced by the US President and European die schöne, umsichtige und von scharfer und Commission in 2013 offer new initiatives for wahrer Kritik zeugende Arbeit. Derartige gibt developing innovative neurotechnology, but es eben in dem immer mehr anschwellenden the prospect of effective therapies for brain Meere des Papierdruckes nicht gar viele, und disease remains uncertain33». Ungeachtet der sie werden leider zu sehr in diesem Ocean des geäusserten Kritik wurden beide Initiativen schwulstigen ‘Schwafel’ ersoffen. Umso mehr als die treibenden Kräfte eines New century of freut man sich, schöne Arbeiten zu treffen.» the brain bezeichnet22. Bei Persistenz dieser in Dieses Phänomen wurde später vom Begrün- Teilen der neurowissenschaftlichen Gemeinde der der Zürcher Neurowissenschaften Cons- proliferierenden euphorischen Haltung kann tantin von Monakow ebenfalls identifiziert extrapoliert werden, dass nach dem Jahrzehnt und als Gefahr für die Weiterentwicklung des des Gehirns der 1990er Jahre (The decade of Gebietes erkannt . In seinem berühmten Refe- the brain) und dem anschliessenden Jahrhun- rat, vorgetragen an der 5. Versammlung der dert des Gehirns der 2010er Jahre (The new Schweizerischen Neurologischen Gesellschaft century of the brain) bald das Jahrtausend des am 29. April 1911 in Aarau, welches anschlies- Gehirns (The new millenium of the brain) auf- send in der von ihm gegründeten Zeitschrift brechen wird. «Arbeiten aus dem Hirnanatomischen Institut in Zürich» (Heft VI, 1912, pp. 1-27) veröffent- Obwohl die moderne Neurowissenschaft licht wurde, hält er diesbezüglich fest: «Wie dank ihrer wissenschaftlichen Durchbrüche, auf allen Gebieten der Medizin, so häuft sich neurotechnologischen Innovationen und er- heute auch in der Hirnlehre rastlos Stoff auf heblichen klinischen Fortschritten in der Pa- Stoff an. Wer auf diesem Gebiete selbsttätig tientenbehandlung, sich vielschichtig entfal- mitarbeitet und nicht zurückbleiben will, muss tet und unbestrittenerweise den Status einer dem Studium der Literatur übermässig viel Zeit medizinischen Leitdisziplin erreicht hat, ist opfern. Die Fülle des neu Dargebotenen, zu- sie mit gegenwärtig weit über 100.000 Pub- mal in Gestalt von Einzelbeobachtungen, lässt likationen pro Jahr unterschiedlicher Qualität dem Forscher kaum Zeit, sich mit letzteren ge- und Relevanz nicht nur unübersichtlich son- nügend vertraut zu machen, geschweige denn dern, bedingt durch die modisch geworde- sie auf ihren wirklichen Wert zu prüfen. Unter nen ‘open access’-Publikationen10 in gewissen der fortgesetzten und hastigen Orientierung ihrer Teilbereiche anfechtbar geworden. So über die -ihrem Inhalt nach- so verschiedenen kommt es, dass gleichermassen wichtige und Leistungen anderer Autoren, leidet nicht sel- irrelevante Forschungsergebnisse in der gan- ten die Übersicht über das Ganze, vor allem zen Weltliteratur zerstreut und nicht selten aber die eigene ausgereifte Produktivität, die mangelhaft besprochen sind. So entstehen ja bekanntlich ihren Ursprung vorwiegend aus in der Neurowissenschaft vorübergehende eigenen Beobachtungen und Erfahrungen ‘Konjukturströmungen’, die gelegentich so- nimmt und durch fremde Gedankengänge gar als die Forschung fehlleitende Dogmen eher gestört als gefördert wird». In ähnlicher weiter bestehen. Die Unübersichtlichkeit der Weise drückte sich Hess 37 Jahre später aus, Neurowissenschaft ist allerdings kein neu- als er, auf Anregung und Einladung des da- es, sondern ein akzentuiertes Phänomen, maligen Rektors, am 1. November 1949 einen auf welches bereits im letzten Viertel des 19. Vortrag in der Aula der Universität Zürich, an- Jahrhunderts der damalige Direktor der psy- lässlich des Festaktes zur Feier der Verleihung chiatrischen Klinik Burghölzli, August Forel, des Nobelpreises für Medizin und Physiologie,

178 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich hielt. Er schloss den Vortrag mit einer persön- ten Neurowissenschaftlern und die jahrmarkt- lichen und bescheidenen Begründung, wie ähnliche Gestaltung neurowissenschaftlicher seine Forschungstätigkeit zur für ihn überra- Kongresse. Treffend haben Hagner und Borck schenden Verleihung des Nobelpreises führ- (1999/2006)43 diese Entwicklung wie folgt be- te: «Der gezeichnete Entwicklungsgang ver- schrieben: «Die grossen Fragestellungen ver- lief scheinbar so programmgemäss, dass man stellen – bei aller qualitativer Bedeutung – den an die Realisierung eines von Anfang an dis- Blick dafür, dass die Neurowissenschaften ponierten Planes denken möchte. Es ist aber eine heterogene Wissenschaftslandschaft dar- nicht so. Führend war von Schritt zu Schritt stellen, die keinen allgemeinen Überblick er- einzig der Vorrang des Interesses, welchem, laubt. Wer die Jahrestagungen der American wie es scheint, eine logische Induktion eigen- Association for Neuroscience mit ihren mehr tümlich ist. Soweit bewusster Wille im Spiele als 30’000 Teilnehmern besucht und sich den war, betrifft dies die Ablehnung, mich nach zahllosen Themengebieten zuzuwenden ver- Konjunkturströmungen zu richten, und im sucht, wird wohl noch bemerken, dass die von Kampfe mit einer widerstrebenden Feder den Vertretern der verschiedenen Disziplinen nicht nachzugeben.»27 Die evidente Paralleli- – Anatomie, Physiologie, Neurologie, Psychia- tät der Ansichten Forels, von Monakows und trie, Molekularbiologie, Biochemie, Physik, Hess’ zeigt wie sich über die Jahre ein für die Statistik, Computerwissenschaften, Linguistik, Zürcher Neurowissenschaft charakteristischer Psychologie usw. – vorgestellten Forschungen Denkstil entwickelt und durch deren Nach- zwar alle in irgendeiner Weise um das Gehirn folger bis in unsere Zeit weitergeleitet hat. oder ein anderes Nervensystem situiert sind; Wesentlich dazu beigetragen haben die Me- doch man kann mit gutem Grund daran zwei- dizinische Fakultät und die Erziehungsdirek- feln, dass sie alle es mit ein und demselben oder tion des Kantons Zürich, welche mit ihren wei- sogar einem einheitlichen Forschungsgegen- sen Entscheidungen in der Ausbauphase der stand zu tun haben. Auch solche Mammutta- Zürcher Neurowissenschften herausragende gungen vermögen die Neurowissenschaft nur Schüler und engste Mitarbeiter als Nachfolger noch als Jahrmarkt der Unübersichtlichkeiten der Gründergeneration zu berufen wussten. zusammenzuhalten»43. Hagner weist in diesem Dies betrifft zum Beispiel die Berufung des Zuammenhang auch dauaf hin, «dass die ver- Neurologen Mieczyslaw Minkowski (1933) als schiedenen Forschungszweige innerhalb der Nachfolger von Constanin von Monakow, des Neurowissenschaften trotz solcher gemeisamer Hirnforschers Konrad Akert (1961) als Nach- Veranstaltungen bis zur gegenseitigen Ver- folger von Water Rudolf Hess, des Neurochi- ständnislosigkeit auseinanderdriften»43. Cha- rurgen M. Gazi Yasargil (1973) als Nachfolger rakteristisch für diese Entwicklung ist auch der von Hugo Krayenbühl und des Neuroradiolo- stattgehabte Wandel, vom Singular zum Plural, gen Anton Valavanis (1984), der zwar keinen in der Bezeichnung der Disziplin: aus der Leit- Vorgänger hatte aber wissenschaftlich und disziplin ‘Neurowissenschaft’ und ihrer inter- klinisch im Zürcher neurowissenschaftlichen agierenden Subdisziplinen ist ein loser, nur be- Umfeld aufgewachsen war38,39,40. schränkt interdisziplinär zusammenarbeitender Verbund der ‘Neurowissenschaften’ geworden. Die oben erwähnte Unübersichtlichkeit be- schränkt sich mittlerweile keinesfalls nur auf Mit seiner eingangs zitierten Aussage, de facto eine überbordende, nicht mehr überschau- sein Vermächtnis, hat der bescheidene Hirnfor- bare Publikationserzeugung, sondern umfasst scher Hess eine Skepsis in die und eine Haltung auch die arbiträr wachsende ‘subdisziplinäre’ gegenüber der Neurowissenschaft hineingetra- Zusammensetzung der Neurowissenschaft, gen, die der Mehrheit der prominenten Neuro- die rasant wachsende Zahl von selbsternann- wissenschaftlern der Gegenwart im unentwirr-

179 Das Vermächtnis von Walter Rudolf Hess und kritische Gedanken zur gegenwärtigen Situation der Neurowissenschaft baren Dickicht der Unübersichtlichkeit der sog. Die von Steno eingeleitete kritisch-skeptische ‘Big Neuroscience’ abzugehen scheint oder be- Haltung fand ihren nächsten Kulminationspunkt reits abgegangen ist. Es scheint als ob die Ver- mit Santiago Ramon y Cajal, dem eigentlichen fechter der ‘Big Neuroscience’ der Illusion ver- Begründer der modernen Neurowissenschaft fallen sind zu einem allumfassenden, perfekten und gemeinsam mit Nobelpreis- Bild einer schönen neuen Neurowelt hinsteuern empfänger für Medizin oder Physiologie (1906), zu können, in welchem der Eindruck erweckt der gegen Ende seiner langen Forschungstätig- wird, als wollte man mit diesem Ansatz alle Be- keit festgehalten hat: lange des menschlichen Gehirns in das Korsett von Molekülen, Genen, Stammzellen, Transmit- «The complexity of the nervous system is so great, tern, Synapsen, Impulsen und neuronalen Ver- its various association systems and cell masses schaltungen stopfen9. so numerous, complex, and challenging, that understanding will forever lie beyond our most Solche Änderungen der Grundhaltung und des committed efforts»16 und «As long as our brain Denkstils sind in der 2.500-jährigen Entwick- is a mystery, the universe, the reflection of the lungsgeschichte der Neurowissenschaft wieder- structure of the brain, will also be a mystery»23. holt vorgekommen9. So vertrat beispielsweise der Philosoph René Descartes Mitte des 17. Jahr- Zum Vermächtnis von Hess gehört, neben der hunderts in seinem 12 Jahre nach seinem Tod Haltung der skeptischen Bescheidenheit, auch erschienenem Werk ‘De homine’ (1650) die An- der Hinweis auf die Priorität der Erforschung der sicht, dass alle Funktionen des Gehirns mit Hilfe Struktur vor derjenigen der Funktion als essen- seiner Maschinentheorie vollständig verstanden zielle methodologische Voraussetzung zum Ver- werden können4. Descartes’s optimistische Hal- ständnis der von der Hirnarchitektonik abhängi- tung gegenüber der Erforschung des Gehirns gen funktionellen Organisation des Gehirns. Die wurde einige Jahre später vom dänischen Ana- strikte Anwendung dieses Fundamentalprinzips tomen und Naturforscher Nicolaus Steno in der Hirnforschung hat es ihm ermöglicht, eine seiner Abhandlung zur Anatomie des Gehirns präzise morphologische Unterteilung und damit (1669) mit folgender Stellungnahme entgegnet: eine Kartographie des Zwischenhirns zu erstel- len, die die Zuordnung der durch die Elektro- «Es wäre ein grosses Glück für die Menschheit, denstimulationen evozierten unterschiedlichen wenn dieses Organ, das, von allen am feinsten funktionellen Reaktionen zu den spezifischen gebaut, sehr häufig gefährlichen Krankheiten anatomischen Untereinheiten des Zwischen- ausgesetzt ist, ebenso gut erkannt würde, wie hirns und so zur Entschlüsselung der komple- viele Philosophen und Anatomen es sich vor- xen, multifunktionellen Organisation des Zwi- stellen. Nur wenige von ihnen drücken sich mit schenhirns und zur Verleihung des Nobelpreises derselben Ehrlichkeit aus wie Sylvius, der über führte42. In seiner Autobiographie beschreibt er das Gehirn nie ohne Vorbehalt spricht, obwohl diese Forschungsmethodologie wie folgt: er sich mit ihm mehr befasst hat als irgend je- mand den ich kenne. Die Zahl derer, denen «Since required accuracy can never be achieved nichts schwierig vorkommt, ist zweifellos viel on the basis of electrode placements alone, grösser. Die Leute, die so prompt Behauptungen microscopic control of serial histologic sections aufstellen, werden Ihnen eine Beschreibung des of each experimental brain is necessary. Three Gehirns und der Lage der einzelnen Teile mit atlases composed of photomicrographic repro- derselben Sicherheit geben, als wären sie beim duction of serial sections cut perpendicularly Bau dieser wunderbaren Maschine zugegen ge- to each other gave invaluable assistance in the wesen und in alle Pläne des grossen Baumeisters topographical analysis of stimulation sites. In eingeweiht worden»19. addition, these atlases served to register in each

180 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich individual case the histological finding as well Auch der Begründer der Zürcher neurowissen- as to prepare cumulative map files of all points schaftlichen Schule und Mitbegründer der Neu- stimulated during the many years of investiga- rologie überhaupt, Constantin von Monakow, tive work. Cross-references among these three hielt 1912 diesbezüglich fest: standard atlases allowed localization of each site of stimulation in all three planes, thereby pro- «Sie alle wissen, dass für das Studium von krank- viding three-dimensional registration, which in haften Hirnerscheinungen (einschliesslich der many cases was indispensable to demonstrate höchsten) einzig die Anatomie die ganz feste the relation between topographical and func- Basis liefert und dass unsere Kenntnisse über nor- tional organization»26. male und pathologische Seelenzustände im näm- lichen Umfang an Festigkeit und Klarheit gewin- Auf diesem Fundamentalprinzip neurowissen- nen, in welchem die physiologisch-anatomische schaftlicher Forschungsmethodologie hat als Hirnforschung eine feinere Ausgestaltung und erster der oben erwähnte Nicolaus Steno in sei- Vertiefung erfährt.»37 ner berühmten kritischen Rede über die Ana- tomie des Gehirns, gehalten 1669 in Paris, hin- Auch die Hirnforscher und Neuroanatomen der gewiesen. Im Abschnitt IV dieser Rede entwirft klassischen Epoche der Neurowissenschaft wie er ein «Neues Programm für die Hirnanatomie» Ramon y Cajal, Korbinian Brodmann, Constan- und hält mit Nachdruck fest: tin von Economo, Georg Koskinas, sowie Cecile und Oskar Vogt befolgten dieses Grundprinzip «Zuerst sichere Anatomie, dann funktionell- in ihrer Forschungsarbeit. So hielt beispielswei- physiologische Deutung». se Koskinas, der mit von Economo das klassische Werk “Die Cytoarchitektonik der Hirnrinde des Zu Beginn des 19. Jahrhunderts kam der Dichter erwachsenen Menschen, 1925” entwickelt hat- und Naturforscher Johann Wolfgang von Goe- te, diesbezüglich 1931 folgendes fest: the (1749-1832) im Rahmen seiner ausgiebigen naturwissenschaftlichen Studien zur Evolutions- “As a general principle, each physiological func- theorie zur Erkenntnis, dass: tion presupposes a corresponding anatomical basis. From the precise knowledge of the struc- «Also bestimmt die Gestalt die Lebensweise des ture of the cerebral cortex we may expect to shed Thieres, light on issues of the utmost importance, such as Und die Weise zu leben, sie wirkt auf alle Gestal- the relationship between mental attributes and ten mächtig zurück34,35» brain structure.” 44, 45

In der zweiten Hälfte des 19. Jahrhunderts, der Und zu Beginn der modernen Ära der Entwick- Blütezeit der klassischen Neurowissenschaft, be- lung der Neurowissenschaft hat der australische kräftigte Bernhard von Gudden, der erste Direk- Neurophysiologe, Philosoph und Nobelpreisträ- tor der 1870 eröffneten psychiatrischen Klinik ger des Jahres 1963 Sir John Eccles die Notwen- Burghölzli und Lehrstuhlinhaber für Psychiatrie digkeit der Befolgung dieses Forschungsprinzips an der Universität Zürich, in seiner im Jahr 1886 mit folgenden Worten bekräftigt: veröffentlichten klassischen Arbeit «Ueber die Frage der Localisation der Functionen der Gross- «In the nervous system, we physiologists are hirnrinde» diese Maxime mit dem Satz: more dependent upon what the anatomists tell us than we are anywhere else»25. «Zuerst also Anatomie und dann Physiologie, wenn aber zuerst Physiologie, dann nicht ohne Der zeitgenössische und einer der führenden Ex- Anatomie»24. ponenten der molekularbiologischen und geno-

181 Das Vermächtnis von Walter Rudolf Hess und kritische Gedanken zur gegenwärtigen Situation der Neurowissenschaft architektonischen Erforschung der Evolution des allerdings mit einer bislang nicht genau be- Gehirns, Luis Puelles, weist mit Nachdruck auf stimmbaren zeitlichen Verschiebung und ohne die Notwendigkeit der primären Erforschung präzise Korrelation mit der betreffenden ana- der Struktur des Gehirns als Voraussetzung für tomischen Hirnstruktur. Die Faszinationskraft, die nachfolgende Zuordnung der Funktion, die die von dieser und ähnlichen Visualisierungsme- ein Derivat und somit ein Epiphänomen der thoden ausgeht, verdrängt die Tatsache, dass sie Struktur ist: lediglich den Eindruck vermitteln, neue Fenster zur Funktionsweise des Gehirns und damit auch «…function is irrelevant because it is context-de- zum geistigen Leben, ja sogar zum sozialen Ver- pendent and epiphenomenal relative to brain halten des Menschen zu öffnen. Damit entsteht structure36». eine verwässernde Popularisierung der Neuro- wissenschaft, die in historischer Perspektive an Als ein herausragendes Paradigma der Befol- die Gallsche Phrenologie des frühen 19. Jahr- gung dieses Prinzips und dessen Anwendung in hunderts erinnert. Die in den letzten Jahren er- die klinische Neurowissenschaft ist die Entwick- folgte Verschiebung des Akzentes der Hirnfor- lung der Mikroneurochirurgie durch M. Gazi Ya- schung mit Prioritätssetzung in die Hirnfunktion sargil, was das Gebiet der Neurochirurgie revolu- lässt sich am objektivsten in der Literatur erken- tioniert und die Zürcher Klinik für Neurochirurgie nen. Eine Literaturrecherche des Autors im Pub- zur Weltgeltung geführt hat. Die, mit Blickwin- med ergab für die Periode 1990-2019 219‘700 kel auf den zu wählenden chirurgischen Zugang Publikationen zur fMRT des Gehirns gegenüber zur atraumatischen Erreichung des von einer be- 19’760 Publikationen zur strukturellen MRT des stimmten Erkrankung befallenen Gehirns und Gehirns. Hier sei nochmals und wie oben darge- deren vollständige Entfernung, detaillierte Er- legt daran erinnert, dass Hess, für die Hirnfor- forschung der komplexen und variantenreichen, schung paradigmatisch, zuerst und abgekoppelt phylogenetisch abgeleiteten Hirnanatomie, ge- von der Funktion, die detaillierte anatomische koppelt mit der Überzeugung, dass jede Stelle Unterteilung des Zwischenhirns erforscht und des Gehirns eine funktionelle Bedeutung hat identifiziert hat um auf dieser Grundlage tier- und damit zu schonen ist, bildet die Grundlage experimentell die Funktionen des Zwischenhirns der Yasargilschen Mikroneurochirurgie. Ihre An- präzis zuzuordnen. Der umgekehrte Forschungs- wendung am Patienten erfordert Behutsamkeit weg, zuerst Funktion und dann anatomische und Präzision des chirurgischen Tuns und stän- Zuordnung, hätte zu approximativen oder gar digen Respekt vor dem menschlichen Gehirn41. falschen Ergebnissen geführt.

Während der letzten zwei Jahrzehnte ist aller- Die Fortschritte, welche in den letzten Jahr- dings in Teilen der neurowissenschaftlichen Ge- zehnten auf den Gebieten der Hirnforschung meinschaft eine Abweichung von diesem fun- und den klinisch-neurologischen Wissenschaf- damentalen Forschungsprinzip festzustellen. ten erzielt wurden sowie die grosse öffentliche Diese Tendenz lässt sich beispielsweise an der Aufmerksamkeit, welche diese andauernd und Anwendung der sog. funktionellen Magnet- berechtigterweise finden, begründen einerseits resonanztomographie (fMRT) beobachten. Mit die Neurowissenschaft als eine neue Leitdisziplin dieser Methode wird die minimale Erhöhung an der Schnittstelle zwischen Medizin und Na- der Durchblutung und damit der Sauerstoff- turwissenschaft, lenken andererseits etwas von zufuhr in einer ‘Hirnregion’ gemessen. Die so den grossen Herausforderungen ab, mit denen gemessene und dann mit komplizierten ma- die Neurowissenschaft konfrontiert ist. Diese thematischen Operationen in allen möglichen betreffen sowohl die nach wie vor vielen unge- Farbvarianten visualisierte Aktivität lässt sich lösten Fragen in der Forschungsbemühung die mit erhöhter neuronaler Aktivität korrelieren, Struktur und die davon abhängige Arbeitswei-

182 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich se des komplexesten lebenden Systems, näm- aber faszinierenden Gebiet der klinischen und lich des Zentralnervensystems, zu verstehen, als theoretischen Neurowissenschaft zu fördern. Im auch die Ursachen und Mechanismen gewisser Schlusswort seines Vortrags über die «Funktion akuter, mancher bösartiger und vieler chroni- und nervöse Regulation der inneren Organe», scher Hirnerkrankungen zu eruieren und folg- gehalten am 1. November 1949 in der Aula der lich erfolgreich zu behandeln. Angesichts dieser, Universität Zürich, anlässlich des Festaktes zur den Neurowissenschaften inhärenten Hürden Feier der Verleihung des Nobelpreises für Medi- und Limitationen muss der klinische Neurowis- zin und Physiologie hält er folgendes fest: senschaftler, der häufig leicht und schnell ent- fachten Euphorie über positive präliminäre For- «Es soll speziell an die angehenden Jünger der schungsergebnisse oder Behandlungserfolge, Wissenschaft gerichtet sein, die berufen sind, zu eine im Hess’schen Sinne kritische und zugleich ihrer Zeit, in ihrem Gebiet, unter Einsatz ihrer bescheidene Haltung gegenüberstellen kön- Veranlagung und unter Ausnützung der ihnen nen. Wie in keinem anderen Gebiet der Medizin gebotenen äusseren Möglichkeiten die Erkennt- sind Selbstkritik und Bescheidenheit als heraus- nis um einen Schritt voranzubringen»27. ragende Persönlichkeitsmerkmale des klinischen Neurowissenschaftlers gefragt. Dazu gehört Entsprechend ist es eine wesentliche Aufgabe des gemäss Hess aber auch die Erkenntnis, dass die Klinischen Neurozentrums als einer universitär Durchführung eines Forschungsprojektes nicht angesiedelten Institution, die Bedingungen zu immer als Realisierung eines von Anfang an dis- schaffen und nachhaltig zu gewährleisten, welche ponierten Planes verläuft sowie die Bereitschaft eine ungehinderte Forschungsarbeit der daran In- und der Wille sich nicht nach Konjukturströmun- teressierten und dafür Qualifizierten ermöglichen. gen zu richten27. Das Vermächtnis von Walter Ru- dolf Hess nimmt einen festen Platz in der langen Im Klinischen Neurozentrum wird der vom Tradition der Zürcher Neurowissenschaft ein, die Hess’schen Vermächtnis abgeleiteten Grund- mit Constantin von Monakow begonnen hat haltung in der klinischen Neuronosologie, in der und über Hess, Krayenbühl, Akert, Baumgart- neurowissenschaftlichen Forschung und in der ner, Yasargil und Schwab, um nur einige der be- Lehre konsequent nachgelebt21. Dies obwohl teiligten Pioniere zu nennen, zu dem heutigen derzeit festzustellen ist, dass der fröhliche Op- Zentrum Neurowissenschaften der Universität timismus, welcher von einer grösseren Gruppe und ETH Zürich (ZNZ, gegründet 1998) sowie prominenter und erfolgreicher Exponenten der zum Klinischen Neurozentrum des Universitäts- neurowissenschaftlichen Forschergemeinschaft spitals Zürich (KNZ, gegründet 2014) geführt propagiert wird, die Hess’schen Tugenden der hat28. Die enge Verknüpfung der beiden Insti- selbstkritischen Haltung und der skeptischen Be- tutionen, der theoretischen (ZNZ) und der klini- scheidenheit im Klammergriff hat. Aber wie die schen (KNZ), die historisch auf die von Hess 1940 Geschichte der Neurowissenschaft mit Bezug auf initiierten Gründung der Zürcher Gemeinschaft die intrinsischen, sich periodisch wiederholen- für Hirnforschung zurückreicht, erwies sich den Oszillationen in der Haltung der Forscher ge- für die Interaktion (in heutiger Terminologie: genüber der Erforschung des Gehirns lehrt, wird Translation) als besonders fruchtbar. Sie bildet es irgendwann einmal wieder umgekehrt sein. ein historisch gewachsenes, charakteristisches Merkmal der Zürcher Neurowissenschaften. Ziel und Kriterium für jede wissenschaftliche Arbeit und damit auch für die Hirnforschung Schliesslich sei hier auf die von Hess hervorge- ist in erster Linie die Suche nach der Wahrheit. hobene Verpflichtung der akademischen Lehrer Sie kann nur gedeihen, wenn die Hirnforscher, der Neurowissenschaft hingewiesen, den dafür sich im Sinne der von Hess propagierten und ge- geeigneten Nachwuchs in diesem komplexen lebten Bescheidenheit bewusst sind, dass, wie

183 Das Vermächtnis von Walter Rudolf Hess und kritische Gedanken zur gegenwärtigen Situation der Neurowissenschaft schon der vorsokratische Philosoph Xenopha- wenn sie aber aus innerer Verpflichtung trotz- nes29 in seinen sogenannten skeptischen Frag- dem mit aller zu Gebote stehenden intellektuel- menten vor über 2000 Jahren postuliert hat, len Redlichkeit danach suchen. niemand die reine Wahrheit wissen wird und

Sichere Wahrheit erkannte kein Mensch und wird keiner erkennen über die Götter und alle die Dinge, von denen ich spreche. Selbst wenn es einem einst glückt, die vollkommenste Wahrheit zu künden, wissen kann er sie nie. Es ist alles durchwebt von Vermutung. (Xenophanes ca. 520 vChr., übersetzt von Karl Popper in: Logik der Forschung, 1934)

Literatur 1. Amunts K., Ebell C., Muller J., Telefont M., Knoll 12. Hess W. R., Fischer H.: Brain and Consciousness: A A., Lippert T.: The Human Brain Project: Creating Discussion About the Function of the Brain. Per- a European Research Infrastructure to Decode the spectives in Biology and Medicine, 17 (1): 109-118; Human Brain. Neuron, 92 (3): 574-581; 2016 1973 2. Amunts K, Knoll AC, Lippert T, Pennartz CMA, Ryv- 13. Lim D.: Brain simulation and personhood: A concern lin P, Destexhe A, et al. (2019) The Human Brain Pro- with the Human Brain Project. Ethics and Informa- ject—Synergy between neuroscience, computing, tion Technology 16: 77-89; 2014 informatics, and brain-inspired technologies. PLoS 14. Markram H., Frackowiak R., and Meier K.: Big di- Biol 17(7): e3000344. https://doi.org/10.1371/jour- gital science-A roadmap for the brain. 2014. Avai- nal.pbio.3000344 lable at: http://ec.europa.eu/archives/commissi- 3. Christen M., Biller-Adorno N., Bringedal B., Grimes on_2010-2014 K., Savulescu J., & Walter H.: Ethical challenges of 15. Marquardt W.: Human Brain Project mediation re- simulation-driven Big Neuroscience. AJOB Neuro- port. 2015. Available at: http://www.fz-juelich.de/ science 7 (1): 5-17; 2016 SharedDocs/Pressemitteilungen/ 4. Des Cartes Renatus.: De homine. Ex officina Hackia- 16. Ramon y Cajal S.: Reglas y Consejos sobre Investiga- na, 1664 cion Biologia (los tonicos de la voluntad), 4th edi- 5. Editorial. The Human Brain Project: mutiny on the tion (Fortanet : Madrid). See English translation by flagship. The Lancet Neurology. 13 (9): 855; 2014 N. Swanson and L.W.Swanson, Advice for a young Investigator, MIT Press: Cambridge, 1999, p.63) 6. Frégnac Y., and Laurent G.: Where is the brain in the Human Brain Project? Nature 513: 27-29; 2014 17. Requarth T.: The big problem with “big science” ventures -Like the Human Brain Project. 2015. Avai- 7. Galison P.: The many facets of Big Science. In: Big lable at: http://nautil.us/blog/ science: The growth of large-scale research, ed. P. Galison and B. W. Hevly, 1-17; 1992. Stanford CA: 18. Rose N.: The Human Brain Project: Social and ethical Stanford University Press challenges. Neuron 82 (6): 1212-1215; 2014 8. Grillner S.: Megascience efforts and the brain. Neu- 19. Steno N.: Discourse de Monsieur Stenon sur l’ Ana- ron 82 (6): 1209-1211; 2014 tomie du Cerveau. 1669. Paris: Ninville 9. Hagner M.: Der Geist bei der Arbeit. Historische 20. Technical Review Report of the HBP. 2015. Available Untersuchungen zur Hirnforschung. 2. Aufl, 2007, at: https://www.humanbrainproject.eu/-/hbp-tech- Wallstein Verlag, Göttingen 2006 nical-revew-report-now-available 10. Hagner M.: Open access, data capitalism and acade- 21. Valavanis A.: Jahresbericht 2014 des Klinischen Neu- mic publishing. Swiss Med Wkly 2018; 148w14600 rozentrums USZ. 2015, p.1. UniversitätsSpital Zürich 11. HBP Report 2012. The Human Brain Project. A re- 22. Yuste R., Church G. M.: The new century of the port to the European Commission. Available at: brain. Sci Am 310 (3): 38-45; 2014 https://www.humanbrainproject.eu/documents/

184 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

23. Ramón y Cajal, S. : Charlas de café. Pensamientos, 37. von Monakow C.: Über Hirnforschungsinstitute und anécdotas y confidencias / Santiago Ramón y Cajal; Hirnmuseen. Arbeiten aus dem Hirnanatomischen ed., introd. y notas de Francisco Fuster. – Madrid Institut in Zürich. VI: 1-27, 1912 :FCE, 2016 38. Die Universität Zürich 1933 – 1983. Festschrift zur 24. von Gudden B.: Über die Frage der Localisation der 150-JahrFeier der Universität Zürich. Herausgege- Functionen der Hirnrinde. Allgemeine Zeitschrift ben vom Rektorat der Universität Zürich, 1983 für Psychiatrie 42: 478-499, 1986 39. Zürcher Spitalgeschichte, Band 3. Herausgegeben 25. Eccles SJ: in CIBA Foundation Symposium on the vom Regierungsrat des Kantons Zürich 2000, Zürich Neurological Basis of Behavior. Little, Brown: Bos- 40. Jäggi M.: in primo loco. Geschichte der Medizini- ton, 1958, p. 24 schen Fakultät Zürich 1833-2003. Rüffer und Rub,

Sachbuchverlag, Zürich, 2004 26. Hess W.R.: From medical practice to theoretical medicine: an autobiographic sketch. Perspect Biol 41. Yasargil M.G.: A Legacy of Microneurosurgery: Me- Med. 6: 400-423, 1963 moirs, Lessons, and Axioms. Neurosurgery 45 (5): 1025-1091, 1999 27. Hess W.R.: Funktion und nervöse Regulation der in- neren Organe. Vierteljahrsschr. der Naturforsch. 42. Hess, W.R.: Photogrammatlanten von Stammgan- Gesellsch. Zürich. 4: 250-264, 1950 glien und Zwischenhirn der Katze, Standardserie. Physiologisches Institut Zürich 1926 28. Valavanis A.: Meilensteine der Entwicklung der Zür- cher Neurowissenschaften. In: Jahresbericht 2017 43. Hagner M. und Borck C.: «Brave Neuro Worlds. des Klinischen Neurozentrums USZ. pp. 5-12 Historische Untersuchungen zur Hirnforschung». In: Hagner M. Der Geist bei der Arbeit. Historische 29. Xenophanes.: In: M. Laura Gemelli Marciano: Die Untersuchungen zur Hirnforschung. Göttingen, S. Vorsokratiker, Band 1, pp. 222-283, Artemis und 17-37, 1999/2006 Winkler Verlag, Düsseldorf, 2007 44. Koskinas, G. N. Scientific Works Published in Ger- 30. Anderson A.: Brain research. Heading for the nine- man: their Analyses and Principal Assessments by ties. Nature. 346 (6282): 304 Eminent Scientists. Athens: Pyrsus Publishers. 1931 31. Jones E.G., Mendell L.M.: Assessing the Decade of 45. Triarhou, L. C.: The Comparative Neurology of Neo- the Brain. Science. 284 (5415):739, 1999 cortical Gyration and the Quest for Functional Spe- 32. https://braininitiative.nih.gov/ cialization. Front Syst Neurosci. 2017 Dec 18;11:96. doi: 10.3389/fnsys.2017.00096. eCollection 2017 33. Poo M.: Whereto the mega brain projects. National Science Review. 1: 12-14, 2014 Anschrift des Verfassers 34. Goethe J.W.von.: Bildung und Umbildung organi- Prof. em. Dr. med. Anton Valavanis scher Naturen, 1807 Klinisches Neurozentrum UniversitätsSpital Zürich 35. Russell E.S.: Form and Function. London: Murray; 1916. CH-8091 Zürich 36. Puelles L., Alonso A., Garcia-Calero E., Martinez- e-mail: [email protected] de-la-Torre, M.: Concentric ring topology of mam- malian cortical sectors and relevance for patterning studies. J Comp Neurol. 1-22, 2019

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7. Bibliographie von Walter Rudolf Hess Das Publikationsverzeichnis (1903-1973) von Walter Rudolf Hess

1. Hess, W.R.: Eine mechanisch bedingte Gesetzmässigkeit im Bau des Blutgefäßsystems. Arch. Entwicklungsmech. Org. 16, 632–641 1903 2. Hess, W.R.: Zum Thema: Viskosität des Blutes und Herzarbeit. M.D. Dissertation Vierteljahrsschr. Naturforsch. Ges. Zürich 51, 236–251 1906 3. Hess, W.R.: Ein neuer Apparat zur Bestimmung der Viskosität des Blutes. Münch. Med. Wochenschr. 54, 1590–1591 1907

4. Hess, W.R.: Die Bestimmung der Viskosität des Blutes. Münch. Med. Wochenschr. 54, 2225– 2229 1907

5. Hess, W.R.: Ein neuer Apparat zur Bestimmung der Viskosität des Blutes. Corresp.Bl. Schweiz. Ärzte 37, 73 – 75 1907

6. Hess, W. R.: Un nuovo apparecchio per la determinazione della viscosità del sangue. Policlinico 15, 4–6 1908

7. Hess, W. R.: Eine neue Untersuchungsmethode bei Doppelbildern. Arch. Augenheilkd. 62, 233–238 [p. 55] 1908

8. Hess, W.R.: Die Viskosität des Blutes bei Gesunden. Dtsch. Arch. Klin. Med. 94, 404–408 1908

9. Hess, W.R.: Der Einfluß warmer Bäder auf die Viskosität des Blutes. Wien. Klin. Rundschau 32, 595 - 597 1908

10. Hess, W.R.: Die Viskosimetrie des Blutes (Beitrag zur Apparatenfrage). Med. Klin. 5, 1397– 1400 1909

11. Hess, W.R.: Reibungswiderstand des Blutes und Poiseuillesches Gesetz. Z. Klin. Med. 71, 421–427 1910

12. Hess, W.R.: Blutviskosität and Blutkörperchen. Pflügers Arch. Ges. Physiol. Menschen Tiere 140, 354–362 1911

13. Hess, W.R.: Zur Kritik der Viskosimetrie. Dtsch. Med. Wochenschr. 37, 1854–1855 1911

14. Hess, W.R.: Die graphische Darstellung von Bewegungsstörungen der Augen mit Beispieltafeln zur Diagnose von Augenmuskellähmungen (+ 14 Erläuterungstafeln für den Text). Arch. Augenheilkd. 70, 10–16 1911

15. Hess, W.R.: Über Relationen zwischen Blutbeschaffenheit und Hämodynamik. In: Festschrift zur Eröffnung des gerichtlich-medizinischen Institutes der Universität Zürich, pp. 91–108. Berlin: Hirschwald 1912. Vierteljahrsschr. Gerichtl. Med. 3. Fol. 44, 93–108 1912

16. Hess, W.R.: Reibungswiderstand des Blutes und Poiseuillesches Gesetz. Z. Klin. Med. 74, 428–432 1912

17. Hess, W.R.: Aufzeichnungs-Formulare für die graphische Darstellung von Bewegungsstörungen der Augen nach Dr. W. R. Hess. Fünfzig Blatt. Wiesbaden. Bergmann 1912

18. Hess, W.R.: Herstellung plastischer Photographien (mit Demonstrationen). Vierteljahrsschr. Naturforsch. Ges. Zürich 57, XXXIV–XXXVII 1912

19. Hess, W.R.: Der Strömungswiderstand des Blutes gegenüber kleinen Druckwerten. Arch. Anat. Physiol. (Abt. Physiol.) 197–214 1912

20. Hess, W.R.: Der Einfluß des Druckes auf den Koeffizienten der Blutviscosität. Berl. Klin.

187 Bibliographie von Walter Rudolf Hess

Wochenschr. 50, 197 1913

21. Hess, W.R.: Das Prinzip des kleinsten Kraftverbrauches im Dienste hämodynamischer Forschung. Master’s thesis. Leipzig: Veit. Arch. Anat. Physiol. (Abt. Physiol.) 1914, 1–62 1913

22. Hess, W.R.: Direkt wirkende Stereoskopbilder. Z. Wiss. Photogr. Photophys. Photochem. 14, 33–38 [p. 61] 1914

23. Hess, W.R.: Unmittelbar wirkende Stereoskopbilder. Vierteljahrsschr. Naturforsch. Ges. Zürich 1914 24. Hess, W.R.: Über das stereoskopische Bild. Wissen und Leben 424–430 1914

25. Hess, W.R.: Über die funktionelle Bedeutung der Arterienmuskulatur. Corresp.-Bl.Schweiz. Ärzte 44, 993–1000 1914

26. Hess, W.R: Die Wärme im Energiehaushalt des Organismus. Corresp.-Bl. Schweiz. Ärzte 44, 1281–1292 1914

27. Hess, W.R.: Gehorcht das Blut dem allgemeinen Strömungsgesetz der Flüssigkeiten? Pflügers Arch. Ges. Physiol. Menschen Tiere 162, 187–224 1915

28. Hess, W.R.: Die Arterienmuskulatur als „peripheres Herz“? Pflügers Arch. Ges. Physiol. Menschen Tiere 163, 555–593 1916

29. Hess, W.R.: Ein einfaches messendes Verfahren zur Motilitätsprüfung der Augen. Z. Augenheilk. 35, 201–219 1916

30. Hess, W.R.: Über die periphere Regulierung der Blutzirkulation. Pflügers Arch. Ges. Physiol. Menschen Tiere 168, 439–490 1917

31. Hess, W.R.: Die Kinematographie in Unterricht und Forschung. Verh. Schweiz. Naturforsch. Ges. Zürich 99. Vers., 307–308 1917

32. Hess, W.R.: Die Zweckmäßigkeit im Blutkreislauf. Akademische Antrittsvorlesung, gehalten am 15. Juni 1918. Basel: Schwabe [p. 35] 1918

33. Hess, W.R.: Der erste Herzton. Verh. Schweiz. Naturforsch. Ges. Lugano 100. Vers., 143–144 1919

34. Hess, W.R.: Untersuchungen über den Antrieb des Blutstromes durch aktive Gefäßpulsationen. Pflügers Arch. Ges. Physiol. Menschen Tiere 173, 243–264 1919

35. Hess, W.R.: Viscosimeter mit Temperaturregulierung. Pflügers Arch. Ges. Physiol. Menschen Tiere 180, 61–67 1920

36. Hess, W.R.: Beitrag zur Theorie der Viskosität heterogener Systeme. Kolloid Z. 27, 1–11 1920

37. Hess, W.R.: Die innere Reibung gelatinierender Lösungen. Kolloid Z. 27, 154–163 1920

38. Hess, W.R.: Die graphische Aufzeichnung der Herztöne nach neuer Methode. Pflügers Arch. Ges. Physiol. Menschen Tiere 180, 35–60 1920

39. Hess, W.R.: Die Entstehung des ersten Herztones. Dtsch. Arch. Klin. Med. 132, 69–95 1920

40. Hess, W.R., Gundlach, R.: Der Einfluß des Adrenalins auf die Sekretion des Magensaftes. Pflügers Arch. Ges. Physiol. Menschen Tiere 185, 122–136 1920

41. Hess, W.R., Gundlach, R.: Der Einfluß von Hypophysenextrakt auf die Magensaftsekretion. Pflügers Arch. Ges. Physiol. Menschen Tiere 185, 137–140 1920

42. Hess, W.R.: Die physiologischen Grundlagen für die Entstehung der reaktiven Hyperämie und des Kollateralkreislaufes. Beitr. Klin. Chir. 122, 1–19 1921

188 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

43. Hess, W.R.: Die Rolle der Vitamine im Zellchemismus. Hoppe Seylers Z. Physiol. Chem. 117, 284–308 1921

44. Hess, W.R.: Die Sensibilität der Kreislaufregulierung. Verh. Schweiz. Naturforsch. Ges. Schaffhausen 102. Vers., 160–161 1921

45. Hess, W.R., Takahashi, N.: Nachweis eines stofflichen Defizites im Gewebe an Avit-aminose erkrankter Tiere. Biochem. Z. 722, 193–203 1921

46. Hess, W.R.: Photographische Konzentrationsbestimmung einer Farbstofflösung. Hoppe Seylers Z. Physiol. Chem. 779, 172–175 1922

47. Hess, W.R.: Die Rolle der Vitamine im Zellchemismus. Bemerkungen zu der „Ergänzung“von Emil Abderhalden. Hoppe Seylers Z. Physiol. Chem. 120, 277–280 1922

48. Hess, W.R., Messerle, N.: Untersuchungen über die Gewebeatmung bei Avitaminose. Hoppe Seylers Z. Physiol. Chem. 119, 176–189 1922

49. Hess, W.R., Wyss, W.H. v.: Beitrag zur Kenntnis der Eingeweidesensibilitäten. Pflügers Arch. Ges. Physiol. Menschen Tiere 194, 195–205 1922

50. Hess, W.R.: Viscosimetrische Untersuchungen an lyophilen Kolloiden. Ree. Trav. Chim. Pays Bas 42, 1097–1103 1923

51. Hess, W.R.: Eine Station für Höhenforschung an der Jungfrau. Q. J. Exp. Physiol. Suppl. Vol.: Proc. Xlth Int. Physiol. Congress, Edinburgh 143–144 1923

52. Hess, W.R.: Über die Wirkung von Acetylcholin auf den Skelettmuskel. Q. J. Exp. Physiol. Suppl. Vol.: Proc. Xlth Int. Physiol. Congress, Edinburgh 144–146 [p. 67] 1923

53. Hess, W.R.: Die Regulierung des peripheren Blutkreislaufes. Ergeb. Inn. Med. Kinderheilkd. 23, 1–65 1923

54. Hess, W.R.: Die physiologischen Grundlagen der pathologischen Blutdrucksteigerung. Schweiz. Med. Wochenschr. 53, 1077–1082 1923

55. Hess, W.R.: Die Blausäurevergiftung als Methode der Avitaminoseforschung. Pflügers Arch. Ges. Physiol. Menschen Tiere 198, 483–489 1923

56. Hess, W.R.: „Die Rolle der Vitamine im Zellchemismus“. Erwiderung auf die Antwort Emil Abderhaldens. Hoppe Seylers Z. Physiol. Chem. 727, 196–198 1923

57. Hess, W.R.: Unbewußte Sinnesleistungen. Vierteljahr sehr. Naturforsch. Ges. Zürich 68, III–IV 1923

58. Hess, W.R.: Metodo grafico per lo studio dei disturbi délia motilità oculare. Boll. Oculistica 2, 614–616 1923

59. Hess, W.R., Rohr, K.: Über den Einfluß thermischer Vorbehandlung von Trockenhefe auf ihre Reduktionsleistung und ihre Vitaminwertigkeit, mit einem Beitrag zur Kenntnis der Vogelberiberi. Hoppe-Seylers Z. Physiol. Chem. 129, 268–283 1923

60. Hess, W.R., Ryffel, W.: Atmungsversuche an überlebender Fischmuskulatur. Ein Beitrag zur Biologie des Muskels. Pflügers Arch. Ges. Physiol. Menschen Tiere 200, 124–131 1923

61. Hess, W.R., Zimmermann, A.: La viscosité du suc pancréatique. Arch. Intern. Physiol. 27, 191–203 1923

62. Hess, W.R.: Über die Wirkung der Vitamine. Dtsch. Med. Wochenschr. 50, 163–164 1924

63. Hess, W.R.: Toxikologische Untersuchungen im Dienste der Avitaminoseforschung. Arch. Exp. Pathol. Pharmakol. 103, 366–379 1924

189 Bibliographie von Walter Rudolf Hess

64. Hess, W.R.: Zur Physiologie der Vasomotoren. Schweiz. Arch. Neurol. Psychiatr. 14, 20–29 1924

65. Hess, W.R.: Die reflektorische Ruhigstellung schmerzender Körperteile. Pflügers Arch. Ges. Physiol. Menschen Tiere 203, 539–541 1924

66. Hess, W.R.: Die Prüfung der motorischen Koordination der Augen im graphischen Verfahren. Laupen (bei Bern ): Polygraphische Gesellschaft 1924

67. Hess, W.R.: Méthode graphique pour l’examen de la motilité oculaire. Rev. Gen. Ophtalmol. 38, 5–20 1924

68. Hess, W.R.: Die Physiologie der Arbeit. Naturwissenschaften 12, 1031–1039 1924

69. Hess, W.R.: Diskussion zu Michaud, L.: Le rôle du système nerveux autonome dans la pathogénèse des maladies internes. Schweiz. Arch. Neurol. Psychiatr. 14, 308–309 1924

70. Hess, W.R., Neergaard, K. v.: Die Beziehungen der Acetylcholinverkürzung des Skelettmuskels zur Einzelzuckung und zum Tetanus. Pflügers Arch. Ges. Physiol. Menschen Tiere 205, 506–517

71. Hess, W.R.: Uber die Wechselbeziehungen zwischen psychischen und vegetativen Funktionen. Schweiz. Arch. Neurol. Psychiatr. 15, 260–277 (1924); 16, 36–55, 285–306 [p. 71] 1925

72. Hess, W.R.: Über die Bedeutung der Fusion für die motorische Koordination der Augen. Klin. Monatsbl. Augenheilkd. 75, 289–294 1925

73. Hess, W.R.: Die physiologische Bedeutung des Sportes für die Jugenderziehung. Schweiz. Z. Gesundheitspflege 5, 1–6 1925

74. Hess, W.R.: Die Wirkung von Ergotamin auf das Auge. Klin. Monatsbl. Augenheilkd. 75, 295– 300 1925

75. Hess, W.R., Messerle, N.: Untersuchungen über die motorische Koordination der Augen. Pflügers Arch. Ges. Physiol. Menschen Tiere 210, 708–720 1925

76. Hess, W.R.: Klima und Schlaf. In: Verhandlungen der klimatologischen Tagung in Davos 1925, pp. 1–8. Basel: Schwabe 1926

77. Hess, W.R.: Untersuchungen über den Einfluß der Peripherie auf die Regulierung der Herztätigkeit. Schweiz. Med. Wochenschr. 56, 690–692 1926

78. Hess, W.R.: Untersuchungen über den Mechanismus der Kreislaufregulierung. Pflügers Arch. Ges. Physiol. Menschen Tiere 213, 163–170 1926

79. Hess, W.R.: Funktionsgesetze des vegetativen Nervensystems. Klin. Wochenschr. 5, 1353– 1354 1926

80. Hess, W.R.: Die Temperaturregulierung im Bienenvolk. Z. Vergl. Physiol. 4, 465–487 1926

81. Hess, W.R.: Experimentelles zur Dynamik der Augenmuskeln. Arch. Augenheilkd. 97, 460– 466 1926

82. Hess, W.R.: Vegetatives Nervensystem und Auge. Klin. Monatsbl. Augenheilkd. 77, 708 1926

83. Hess, W.R.: Praktische Erfahrungen in der Diagnostik von Augenmuskelstörungen. Klin. Monatsbl. Augenheilkd. 77, 708–709 1926

84. Hess, W.R.: Photogrammatlanten von Stammganglien und Zwischenhirn der Katze, Standardserie. Physiologisches Institut Zürich 1926

190 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

85. Hess, W.R., Lehmann, F.E.: Der Einfluß vegetativer Reizstoffe auf den Adaptierungszustand der Retina. Pflügers Arch. Ges. Physiol. Menschen Tiere 211, 603–611 1926

86. Hess, W.R., Rehsteiner, R.: Die Wirkung von Acetylcholin auf den Zuckungsablauf des Froschmuskels. Pflügers Arch. Ges. Physiol. Menschen Tiere 214, 463–470 1926

87. Hess, W.R.: Die innere Reibung des Blutes. In: Handbuch der physikalischen und technischen Mechanik, Vol. V. Auerbach, F., Hort, W. (eds.), pp. 471–472. Leipzig: Barth 1927

88. Hess, W.R.: Das Löschphänomen der Acetylcholin-Contraktur. Pflügers Arch. Ges. Physiol. Menschen Tiere 217, 511–515 1927

89. Hess, W.R.: Methodisches zur Untersuchung von Störungen der Augenmotilität. Klin. Monatsbl. Augenheilkd. 79, 289–292 1927

90. Hess, W.R.: Die Gesetze der Hydrostatik und Hydrodynamik. In: Handbuch der normalen und pathologischen Physiologie, Vol.VII/2: Blutzirkulation II. Bethe, A. et al. (eds.), pp. 889 – 903. Berlin: Springer 1927

91. Hess, W.R.: Die Verteilung von Querschnitt, Widerstand, Druckgefälle und Strömungsgeschwindigkeit im Blutkreislauf. In: Handbuch der normalen und pathologischen Physiologie, Vol. VII/2: Blutzirkulation II. Bethe, A. et al. (eds.), pp. 904–933. Berlin: Springer 1927

92. Hess, W.R.: Stammganglien-Reizversuche. Ber. Ges. Physiol. Exp. Pharmakol. 42, 554–555 1927

93. Hess, W.R.: Bewegungsanalytische Filme. Ber. Ges. Physiol. Exp. Pharmakol. 42, 574–575 1927

94. Hess, W.R.: Das Schlafzentrum. Verh. Schweiz. Naturforsch. Ges. Basel 108. Vers., 247–248 1927

95. Hess, W.R., Büsch, J.: Der Einfluß von Atropin auf die Reaktion des Skelettmuskels bei direkter Reizung. Pflügers Arch. Ges. Physiol. Menschen Tiere 216, 644–650 1927

96. Hess, W.R.: Hirnreizversuche über den Mechanismus des Schlafes. Arch. Psychiatr. Nervenkr. 86, 287–292 1928

97. Hess, W.R.: Praktische Ernährungslehre auf wissenschaftlicher Grundlage. Führer durch die Wanderausstellung für Ernährungslehre. Zürich: Schweizer. Zentralstelle für Gesundheitspflege 1928 98. Hess, W.R.: Über den Schlaf. Vierteljahrsschr. Naturforsch. Ges. Zürich 74, XIX–XX 1929

99. Hess, W.R.: Versteckte Feinde - Wissen gibt Macht - Von den Vitaminen - Gewissenskonflikt - Tierschutz im Tierversuch. In: Aussprache zwischen Vertretern der experimentellen Medizin und Vivisektionsgegnern über die Zweckdienlichkeit und Zulässigkeit des wissenschaftlichen Tierversuches. Tierfreund 55, 32–36 1929

100. Hess, W.R.: The Alpine research station of Switzerland. J. State Med. 37, 11–14 1929

101. Hess, W.R.: The mechanism of sleep. Am. J. Physiol. 90, 386–387 1929

102. Hess, W.R.: Lokalisatorische Ergebnisse der Hirnreizversuche mit Schlafeffekt. Arch. Psychiatr. Nervenkr. 88, 813–816 [p. 131] 1929

103. Hess, W.R.: Die Funktionen des vegetativen Nervensystems. Klin. Wochenschr. 9, 1009– 1012 1930

104. Hess, W.R.: Kritik der Hering-Breuer’schen Lehre von der Selbststeuerung der Atmung. Pflügers Arch. Ges. Physiol. Menschen Tiere 226, 198–211 [p. 211] 1930

191 Bibliographie von Walter Rudolf Hess

105. Hess, W.R.: Die Regulierung des Blutkreislaufes, gleichzeitig ein Beitrag zur Physiologie des 105. Hess,vegetativen W.R.: Nervensystems,Die Regulierung pp.des 1 Blutkreislaufes,–162. Leipzig: Thieme gleichzeitig 1930 ein Beitrag zur Physiologie des vegetativen Nervensystems, pp. 1–162. Leipzig: Thieme 1930 106. Hess, W.R., Arthus, M.: Traité pratique d’alimentation rationnelle à base scientifique. Guide de 106. Hess,l’exposition W.R., itinérante Arthus, M.: d’alimentation Traité pratique rationnelle. d’alimentation Zürich: rationnelle Centrale Suisseà base pourscientifique. la Propagande Guide de l’expositiond’Hygiène 1930 itinérante d’alimentation rationnelle. Zürich: Centrale Suisse pour la Propagande d’Hygiène 1930 107. Hess, W.R.: Die Verwendung des Schmalfilms für biologische Zwecke. In: Handbuch der 107. Hess,biologischen W.R.: DieArbeitsmethoden, Verwendung des Abt. Schmalfilms II: Physikalische für biologische Methoden, Zwecke. Vol. 2/II. In: Abderhalden, Handbuch der E. (ed.),biologischen pp. 2317 Arbeitsmethoden,–2336. Berlin: Urban Abt. &II: SchwarzenbergPhysikalische Methoden, 1930 Vol. 2/II. Abderhalden, E. (ed.), pp. 2317–2336. Berlin: Urban & Schwarzenberg 1930 108. Hess, W.R.: Biologie und Universitas. Vierteljahrsschr. Naturforsch. Ges. Zürich 76, 7–15 108. Hess,1931 W.R.: Biologie und Universitas. Vierteljahrsschr. Naturforsch. Ges. Zürich 76, 7–15 1931 109. Hess, W.R.: Le sommeil. C.R. Soc. Biol. (Paris) 107, 1333–1364 1931 109. Hess, W.R.: Le sommeil. C.R. Soc. Biol. (Paris) 107, 1333–1364 1931 110. Hess, W.R.: Der Schlaf. Schweiz. Med. Wochenschr. 61, 849 1931 110. Hess, W.R.: Der Schlaf. Schweiz. Med. Wochenschr. 61, 849 1931 111. Hess, W.R.: Die Regulierung der Atmung, gleichzeitig ein Beitrag zur Physiologie des 111. Hess,vegetativen W.R.: Nervensystems,Die Regulierung pp.der 1Atmung,–137. Leipzig: gleichzeitig Thieme ein 1931 Beitrag zur Physiologie des vegetativen Nervensystems, pp. 1–137. Leipzig: Thieme 1931 112. Hess, W.R.: Entstehungsgeschichte und Ziele des Forschungsinstitutes. In: Hochalpine 112. ForschungsstationHess, W.R.: Entstehungsgeschichte Jungfraujoch. Internationale und Ziele des Stiftung Forschungsinstitutes. Bern, pp. 11–16. In:Zürich: Hochalpine Orell Füssli Forschungsstation1931 Jungfraujoch. Internationale Stiftung Bern, pp. 11–16. Zürich: Orell Füssli 1931 113. Hess, W.R.: Die Grundlagen der Arbeitsphysiologie. Tech. Rundschau (Bern) 23/45, 1 – 2 113. Hess,1931 W.R.: Die Grundlagen der Arbeitsphysiologie. Tech. Rundschau (Bern) 23/45, 1 – 2 1931 114. Hess, W.R.: The autonomic nervous system. Lancet 223, 1199–1201 1931 114. Hess, W.R.: The autonomic nervous system. Lancet 223, 1199–1201 1931 115. Hess, W.R.: Hirnstamm-Reizversuche. Dtsch. Z. Nervenheilkd. 124, 132–136 1932 115. Hess, W.R.: Hirnstamm-Reizversuche. Dtsch. Z. Nervenheilkd. 124, 132–136 1932 116. Hess, W.R.: Beiträge zur Physiologie des Hirnstammes. I. Die Methodik der lokalisierten 116. ReizungHess, W.R.: und BeiträgeAusschaltung zur Physiologie subkortikaler des Hirnabschnitte, Hirnstammes. I.pp. Die 1– Methodik122. Leipzig: der loThiemekalisierten 1932 Reizung und Ausschaltung subkortikaler Hirnabschnitte, pp. 1–122. Leipzig: Thieme 1932 117. Hess, W.R.: Demonstration: Der Hering-Breuer’sche „Schaltreflex“in neuer Deutung. Schweiz. 117. Hess,Med. Wochenschr. W.R.: Demonstration: 62, 263 1932 Der Hering-Breuer’sche „Schaltreflex“in neuer Deutung. Schweiz. Med. Wochenschr. 62, 263 1932 118. Hess, W.R.: Demonstration der Methodik zur lokalisierten Reizung und Ausschaltung 118. subkortikalerHess, W.R.: Demonstration Hirnabschnitte. der Schweiz. Methodik Med. zur Wochenschr. lokalisierten Reizung62, 1042 und 1932 Ausschaltung subkortikaler Hirnabschnitte. Schweiz. Med. Wochenschr. 62, 1042 1932 119. Hess, W.R.: Motorische Zentren im Hypothalamus. Schweiz. Med. Wochenschr. 63, 457 1933 119. Hess, W.R.: Motorische Zentren im Hypothalamus. Schweiz. Med. Wochenschr. 63, 457 1933 120. Hess, W.R.: Über den Mechanismus der Bewegungsinnervation. Schweiz. Med. Wochenschr. 120. Hess,63, 1030 W.R.: 1933 Über den Mechanismus der Bewegungsinnervation. Schweiz. Med. Wochenschr. 63, 1030 1933 121. Hess, W.R.: Der Schlaf. Klin. Wochenschr. 28, 129–134 [p. 115] 1933 122.121. Hess, W.R.: TeilDer Schlaf.und Ganzes Klin. Wochenschr.im Organismus. 28, Schweiz. 129–134 Lehrerztg. [p. 115] 1933 [Suppl. to No. 44] 78, 21– 122. Hess,25 1933 W.R.: Teil und Ganzes im Organismus. Schweiz. Lehrerztg. [Suppl. to No. 44] 78, 21– 25 1933 123. Hess, W.R.: Modell-Demonstrationen zum Zentrenproblem. Schweiz. Med. Wochenschr. 64, 123. 583Hess, 1934 W.R.: Modell-Demonstrationen zum Zentrenproblem. Schweiz. Med. Wochenschr. 64, 583 1934 124. Hess, W.R.: Plastizitätslehre und Lokalisationsfrage. Verh. Dtsch. Ges. Inn. Med. 46. Kongr. 124. Hess,212–218 W.R.: 1934 Plastizitätslehre und Lokalisationsfrage. Verh. Dtsch. Ges. Inn. Med. 46. Kongr. 125. Hess,212–218 W.R.: 1934 Diskussionsvotum zum Vortrag Muralt, A. v.: Über die Blutregulation im 125. Höhenklima.Hess, W.R.: DiskussionsvotumSchweiz. Med. Wochenschr. zum Vortrag 64, Muralt, 1174 1934A. v.: Über die Blutregulation im Höhenklima. Schweiz. Med. Wochenschr. 64, 1174 1934 126. Hess, W.R.: Die Struktur der starren und der gesteuerten Motorik. Schweiz. Med. 126. Wochenschr.Hess, W.R.: Die 64, Struktur 1176 1934 der starren und der gesteuerten Motorik. Schweiz. Med. Wochenschr. 64, 1176 1934 127. Hess, W.R.: Medizinisch-biologische Fragen und Allgemeines über den Betrieb des 127. Hess, W.R.: Medizinisch-biologische Fragen und Allgemeines über den Betrieb des Internationalen Forschungsinstitutes. Vierteljahrsschr. Naturforsch. Ges. Zürich 79, XIX–XX 1934

128. Hess, W.R.: Die hochalpine Forschungsstation192 Jungfraujoch. Vierteljahrsschr. Naturf. Ges. 79, 1–26 1934

129. Hess, W.R.: Beispiele funktionellen Denkens. In: Festschrift Heinrich Zangger, pp. 769–775. Zürich: Rascher 1935

130. Hess, W.R.: Das physiologische Zusammenspiel von Kreislauf und Atmung. Verh. Dtsch. Ges. Kreislaufforsch. 8. Tagung, 9–31 1935

131. Hess, W.R.: Diskussionsvotum zum Vortrag Sauerbruch, F.: Physiologie und Klinik des Zwerchfells. Schweiz. Med. Wochenschr. 65, 567 1935

132. Hess, W.R.: Physiologische Varia. Schweiz. Med. Wochenschr. 65, 648 1935

133. Hess, W.R.: Aktuelle Probleme der Physiologie. Schweiz. Med. Wochenschr. 65, 852–854 1935

134. Hess, W.R.: Filmdemonstrationen zur Physiologie des Zwischenhirns. Verh. Schweiz. Physiol. (Juni) pp. 18–19. Basel: Schwabe 1935

135. Hess, W.R.: Zentrale Vertretung von Hilfsfunktionen des vegetativen Systems. Schweiz. Med. Wochenschr. 65, 729 1935

136. Hess, W.R.: Diskussionsvotum zu Leuthardt, F.: Allgemeines über die Regulierung der Reaktion und ihre physiologische Schwankungsbreite. Schweiz. Med. Wochenschr. 65, 1035 1935

137. Hess, W.R.: Kritisches zum Zentrenbegriff. In: Problèmes de biologie et de médecine. Volume jubilaire dédié au Prof. Lina Stern, pp. 43–49. Leningrad, Moscow: Governmental Publications in Science and Medicine [p. 205] 1935

138. Hess, W.R.: Hypothalamus und die Zentren des autonomen Nervensystems: Physiologie. Arch. Psychiatr. Nervenkr. 104, 548–557 1936

139. Hess, W.R.: Die Rolle des Vagus in der Selbststeuerung der Atmung. Pflügers Arch. Ges. Physiol. Menschen Tiere 237, 24–39 1936

140. Hess, W.R.: Ziel- und Fertigkeitsbewegungen in der Zeitlupe. Verh. Ver. Schweiz. Physiol. (June), pp. 6–7. Basel: Schwabe 1936

141. Hess, W.R.: Diskussion zu Ruzicka, L.: Über die Sexualhormone, unter besonderer Berücksichtigung der männlichen. Schweiz. Med. Wochenschr. 66, 1020 1936

142. Hess, W.R.: Zentrale Regulierung von Kreislauf und Atmung. Schweiz. Med. Wochenschr. 66, 1078 1936

143. Hess, W.R., Wyss, O.A.M.: Die Analyse der physikalischen Atmungsregulierung an Hand der Aktionsstrombilder des Phrenicus. Pflügers Arch. Ges. Physiol. Menschen Tiere 237, 761–770 1936

144. Hess, W.R.: Die Lokomotion von Tier und Mensch in der Zeitlupe. Verh. Ver. Schweiz. Physiol. (January), p. 6. Basel: Schwabe 1937

145. Hess, W.R.: I. Reflexbeziehungen zwischen afferente Vagus und Innervation der respiratorischen Thoraxmuskulatur. II. Beziehungen zwischen der Innervation synergistischer Muskeln. Verh. Ver. Schweiz. Physiol. (June), pp. 5–6. Basel: Schwabe 1937

146. Hess, W.R.: O pokusném vysetrovâni vegetativniho nervového systému. Biol. Listy 22, 77–82 1937 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich Internationalen Forschungsinstitutes. Vierteljahrsschr. Naturforsch. Ges. Zürich 79, XIX–XX 1934

128. Hess, W.R.: Die hochalpine Forschungsstation Jungfraujoch. Vierteljahrsschr. Naturf. Ges. 79, 1–26 1934

129. Hess, W.R.: Beispiele funktionellen Denkens. In: Festschrift Heinrich Zangger, pp. 769–775. Zürich: Rascher 1935

130. Hess, W.R.: Das physiologische Zusammenspiel von Kreislauf und Atmung. Verh. Dtsch. Ges. Kreislaufforsch. 8. Tagung, 9–31 1935

131. Hess, W.R.: Diskussionsvotum zum Vortrag Sauerbruch, F.: Physiologie und Klinik des Zwerchfells. Schweiz. Med. Wochenschr. 65, 567 1935

132. Hess, W.R.: Physiologische Varia. Schweiz. Med. Wochenschr. 65, 648 1935

133. Hess, W.R.: Aktuelle Probleme der Physiologie. Schweiz. Med. Wochenschr. 65, 852–854 1935

134. Hess, W.R.: Filmdemonstrationen zur Physiologie des Zwischenhirns. Verh. Schweiz. Physiol. (Juni) pp. 18–19. Basel: Schwabe 1935

135. Hess, W.R.: Zentrale Vertretung von Hilfsfunktionen des vegetativen Systems. Schweiz. Med. Wochenschr. 65, 729 1935

136. Hess, W.R.: Diskussionsvotum zu Leuthardt, F.: Allgemeines über die Regulierung der Reaktion und ihre physiologische Schwankungsbreite. Schweiz. Med. Wochenschr. 65, 1035 1935

137. Hess, W.R.: Kritisches zum Zentrenbegriff. In: Problèmes de biologie et de médecine. Volume jubilaire dédié au Prof. Lina Stern, pp. 43–49. Leningrad, Moscow: Governmental Publications in Science and Medicine [p. 205] 1935

138. Hess, W.R.: Hypothalamus und die Zentren des autonomen Nervensystems: Physiologie. Arch. Psychiatr. Nervenkr. 104, 548–557 1936

139. Hess, W.R.: Die Rolle des Vagus in der Selbststeuerung der Atmung. Pflügers Arch. Ges. Physiol. Menschen Tiere 237, 24–39 1936

140. Hess, W.R.: Ziel- und Fertigkeitsbewegungen in der Zeitlupe. Verh. Ver. Schweiz. Physiol. (June), pp. 6–7. Basel: Schwabe 1936

141. Hess, W.R.: Diskussion zu Ruzicka, L.: Über die Sexualhormone, unter besonderer Berücksichtigung der männlichen. Schweiz. Med. Wochenschr. 66, 1020 1936

142. Hess, W.R.: Zentrale Regulierung von Kreislauf und Atmung. Schweiz. Med. Wochenschr. 66, 1078 1936

143. Hess, W.R., Wyss, O.A.M.: Die Analyse der physikalischen Atmungsregulierung an Hand der Aktionsstrombilder des Phrenicus. Pflügers Arch. Ges. Physiol. Menschen Tiere 237, 761–770 1936

144. Hess, W.R.: Die Lokomotion von Tier und Mensch in der Zeitlupe. Verh. Ver. Schweiz. Physiol. (January), p. 6. Basel: Schwabe 1937

145. Hess, W.R.: I. Reflexbeziehungen zwischen afferente Vagus und Innervation der respiratorischen Thoraxmuskulatur. II. Beziehungen zwischen der Innervation synergistischer Muskeln. Verh. Ver. Schweiz. Physiol. (June), pp. 5–6. Basel: Schwabe 1937

146. Hess, W.R.: O pokusném vysetrovâni vegetativniho nervového systému. Biol. Listy 22, 77–82 1937

193 Bibliographie von Walter Rudolf Hess

147. Hess, W.R.: Über experimentelle Untersuchungen des vegetativen Nervensystems. Med. Klin. 34, 323–326 1938

148. Hess, W.R.: Inverser Vaguseffekt auf das Zwerchfell. Verh. Ver. Schweiz. Physiol. (January) p. 7. Basel: B. Schwabe 1938

149. Hess, W.R.: Diskussion zu Wagner, H.: Der heutige Stand der operativen Therapie der Netzhautablösung. Schweiz. Med. Wochenschr. 68, 703 1938

150. Hess, W.R.: Wo steht heute die Physiologie und welches sind ihre aktuellen Probleme? Schweiz. Med. Wochenschr. 68, 945–947 1938

151. Hess, W.R.: Eröffnungs-Ansprache des Kongreß-Präsidenten. 16. Int. Physiol. Kongr. Zürich 3, 96–103 1938

152. Hess, W.R.: Rückblicke auf den XVI. Internationalen Physiologen-Kongreß. Schweiz. Med. Wochenschr. 68, 1331 1938

153. Hess, W.R.: Beitrag zur Physiologie des Hirnstammes. II. Das Zwischenhirn und die Regulation von Kreislauf und Atmung, pp. 1–127. Leipzig: Thieme 1938

154. Hess, W.R.: Beziehungen zwischen Winterschlaf und Außentemperatur beim Siebenschläfer. Z. Vgl. Physiol. 26, 529–536 1939

155. Hess, W.R.: Die funktionelle Organisation des Nervensystems. Vierteljahrsschr. Naturforsch. Ges.Zürich 84, 197–210 1939

156. Hess, W.R.: Diskussion zu Rossier, P.: Die Bedeutung des Carotis-Sinus für die Klinik. Schweiz. Med. Wochenschr. 69, 329 1939

157. Hess, W.R.: Kollaboration der Hirnforschung durch Standardisierung der Hilfsgeräte. Schweiz. Arch. Neurol. Psychiatr. 43, 221–224 1939

158. Hess, W.R.: La coordination des mécanismes régulateurs du système respiratoire. J. Physiol. Pathol. Gén. 37, 1233–1238, 1939

159. Hess, W.R.: Eine Stellungnahme der Schweizerischen Medizinisch-biologischen Gesellschaft zum Tierversuch. Schweiz. Med. Wochenschr. 69, 892–894 1939

160. Hess, W.R.: Pupille und Zwischenhirn. Klin. Monatsbl. Augenheilkd. 103, 407–413 1939

161. Hess, W.R.: Zentrogene Einflüsse auf Iris und Augenmuskeln. Schweiz. Med. Wochenschr. 69, 843 1939

162. Hess, W.R.: Prof. Justus Gaulet † 1849–1939. Verh. Ver. Schweiz. Physiol. (July), pp. 3–4. Basel: Schwabe 1939

163. Hess, W.R.: Beziehungen des Hypothalamus zu Trieb- und Stimmungsäußerungen. Verh. Ver. Schweiz. Physiol. (July), pp. 5–9. Basel: Schwabe 1939

164. Hess, W.R.: Georg Justus Gaule (1849–1939); Mitglied der Gesellschaft seit 1887. Vierteljahrsschr. Naturforsch. Ges. Zürich 84, 375–377 1939

165. Hess, W.R.: Ergebnisse von Reizversuchen in Zwischenhirn und Nachbargebieten. Pflügers Arch. Ges. Physiol. Menschen Tiere 243, 409–430 1940

166. Hess, W.R.: Beitrag zur Technik des zentralen Reizversuches. Pflügers Arch. Ges. Physiol. Menschen Tiere 243, 431–438 1940

167. Hess, W.R.: Die Ernährung der Zivilbevölkerung. Schweiz. Z. Gemeinnützigkeit 79, 190–199 1940

194 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

168. Hess, W.R.: Statistische Analyse zentraler Reizeffekte. Verh. Ver. Schweiz. Physiol. (June), pp. 15–16. Basel: Schwabe 1940

169. Hess, W.R.: Zwischenhirn und Motorik. Pflügers Arch. Ges. Physiol. Menschen Tiere 243, 634 – 650 1940

170. Hess, W.R.: Motorische Symptome im Gesicht bei Reizung im Zwischenhirn. Pflügers Arch. Ges. Physiol. Menschen Tiere 243, 678–685 1940

171. Hess, W.R.: Diencephale Reizsymptome am Körperstamm; Beziehungen derselben zu Bewegungseffekten im Gesicht und an den Extremitäten. Pflügers Arch. Ges. Physiol. Menschen Tiere 243, 741–747 1940

172. Hess, W.R.: Aufbau und Aufbauordnung physiologischer Funktionen. Umschau 47, 131–139 1940

173. Hess, W.R.: Der Reflextonus des Zwerchfells und der Einfluß der Lungenentfaltung auf den Atmungstypus. Veröff. der Reichsstelle für den Unterrichtsfilm zu dem Archivfilm Nr. B 438 /1940

174. Hess, W.R.: Zwischenhirn und Gleichgewichtsregulierung. Verh. Schweiz. Naturforsch. Ges. Locarno, 120. Vers., 212 1940

175. Hess, W.R.: Weitere Beobachtungen über den tonischen Vaguseinfluß bei verschiedenem konstantem Lungenvolumen. Pflügers Arch. Ges. Physiol. Menschen Tiere 244, 360–364 1941

176. Hess, W.R.: Zwischenhirn und Gleichgewichtsregulierung. Schweiz. Med. Wochenschr. 71, 893–895 1941

177. Hess, W.R.: Beziehung von Zwischenhirn zur Organisation der Bewegung. Verh. Ver. Schweiz. Physiol. (July), p. 4. Basel: Schwabe 1941

178. Hess, W.R.: Experimenteller Beitrag zur Frage der extrapyramidalen Motorik. Z. Ges. Neurol. Psychiatr. 172, 639–642 1941

179. Hess, W.R.: Die Motorik als Organisationsproblem. Biol. Zentralbl. 61, 545–572 1941

180. Hess, W.R.: Charakter der im Zwischenhirn ausgelösten Bewegungseffekte. Pflügers Arch. Ges. Physiol. Menschen Tiere 244, 767–786 [p. 223] 1941

181. Löffler, W., Hess, W.R.: Über Isthmusstenose der Aorta als pathologisch-physiologisches Problem. Schweiz. Med. Wochenschr. 71, 636–647 1941

182. Hess, W.R.: Das vegetative Funktionssystem. In: Schweizer Medizinisches Jahrbuch, pp. 51– 67. Basel: Schwabe,1942

183. Hess, W.R.: Biomotorik als Organisationsproblem. Naturwissenschaften 30, 441–448, 537– 541 [p. 243], 1942

184. Hess, W.R.: Der naturwissenschaftliche Film für Forschung und Unterricht. Vierteljahrsschr. Naturforsch. Ges. Zürich 86, XXVI-XXVIII,1942

185. Hess, W.R.: Die S.E.D.-Methode in der Hirnforschung. Verh. Ver. Schweiz. Physiol. (June), pp. 22–23. Basel: Schwabe,1942

186. Hess, W.R.: Vegetatives Nervensystem. Fragen der Organisation der Begriffe und Bezeichnungen. Schweiz. Arch. Neurol. Psychiatr. 50, 1–5, 1942

187. Hess, W.R.: Physiologische Aspekte der extrapyramidalen Motorik. Nervenarzt 15, 457–466, 1942

195 Bibliographie von Walter Rudolf Hess

188. Hess, W.R., Bartorelli, C., Bucher, V.: Diencephale Motorik und Marchi-Degeneration. Z. Ges. Neurol. Psychiatr. 174, 340–352, 1942

189. Hess, W.R.: Die Regulation der Laktogenese als Beispiel allgemein-physiologischer Gesetzlichkeit. Schweiz. Med. Wochenschr. 73, 53–60, 1943

190. Hess, W.R.: Induzierte Störungen der optischen Wahrnehmung. Nervenarzt 16, 57–66, 1943

191. Hess, W.R.: Alte und neue Ziele der Physiologie. Vierteljahrsschr. Naturforsch. Ges. Zürich 88, 67–76, 1943

192. Hess, W.R.: Von den Gefühlen und Stimmungen eines Hundes. Ein Beitrag zur vergleichenden Psychophysiologie. Z. Vgl. Physiol. 30, 123–138, 1943

193. Hess, W.R.: Symptomatik des durch elektrischen Reiz ausgelösten Schlafes und die Topographie des Schlafzentrums (Film) Helv. Physiol. Acta 1, C61 - C62, 1943

194. Hess, W.R.: Teleokinetisches und ereismatisches Kräftesystem in der Biomotorik. Helv. Physiol. Acta 1, C62-C63 [p. 265], 1943

195. Hess, W.R.: Das Zwischenhirn als Koordinationsorgan. Helv. Physiol. Acta 1, 549–565, 1943

196. Hess, W.R., Brügger, M.: Das subkortikale Zentrum der affektiven Abwehrreaktion. Helv. Physiol. Acta 1, 33–52 [p. 183], 1943

197. Hess, W.R., Brügger, M.: Der Miktions- und der Defäkationsakt als Erfolg zentraler Reizung. Helv. Physiol. Acta 1, 511–532, 1943

198. Hess, W.R., Magnus, W.O.C.: Leck- und Kau-Automatismen bei elektrischer Reizung. Helv. Physiol. Acta 1, 533–547, 1943

199. Hess, W.R.: Diskussion zu Sulzer, W.: Arbeitsschutz und Arbeitsmedizin. Schweiz. Med. Wochenschr. 74, 270, 1944, 1944

200. Hess, W.R.: Wandernde Tonusstörungen nach zentraler Reizung. Helv. Physiol. Acta 2, 131– 135, 1944

201. Hess, W.R.: Hypothalamische Adynamie. Helv. Physiol. Acta 2, 137–147 [p. 171], 1944

202. Hess, W.R.: Das Schlafsyndrom als Folge dienzephaler Reizung. Helv. Physiol. Acta 2, 305– 344 [p. 131], 1944

203. Hess, W.R.: Alfred Vogt †. Von seinem Leben und Schaffen. Vierteljahrsschr. Naturforsch. Ges. Zürich 89, 136–140, 1944

204. Hess, W.R.: Kollektive Ordnung in biologischem Aspekt. In: Festschrift Max Huber - Vom Krieg und vom Frieden, pp. 151–172. Zürich: Schulthess, [p. 3], 1944

205. Hess, W.R., Stoll, W.A.: Experimenteller Beitrag betreffend die Regulierung der Körpertemperatur. Helv. Physiol. Acta 2, 461–480, 1944

206. Hess, W.R.: Der extrapyramidal-motorische Innervationsapparat. Helv. Physiol. Acta 3, C 3-C 4, 1945

207. Hess, W.R.: Von den höheren Zentren des vegetativen Funktionssystems. Bull. Schweiz. Akad. Med. Wiss. 1, 138–166, 1945

208. Hess, W.R.: Experimentelle Darstellung des mesencephalen Blickzentrums, seiner Struktur und Funktionsweise. Helv. Physiol. Acta 4, C 3-C 4, 1946

209. Hess, W.R.: Demonstrationen und Erläuterungen zur Funktion des Tectum opticum. Bull. Schweiz. Akad. Med. Wiss. 2, 51–54, 1946

196 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

210. Hess, W.R.: Statistik betreffend die Ernährungsverhältnisse während der Kriegszeit. Bull. Eidgen. Gesundheitsamtes Nr. 45, 1–8, 1946

211. Hess, W.R., Müller, H.R.: Schnupperbewegungen als zentrale Reizeffekte. Helv. Physiol. Acta 4, 339–345, 1946

212. Hess, W.R., Müller, H.R.: Einflüsse des Mittel- und Zwischenhirns auf die Atmung. Helv. Physiol. Acta 4, 347–358, 1946

213. Hess, W.R., Bürgi, S., Bucher, V.: Motorische Funktion des Tektal- und Tegmentalgebietes. Monatsschr. Psychiatr. Neurol. 112, 1–52 [p.269], 1946

214. Hess, W.R., Brügger, M., Bucher, V.: Zur Physiologie von Hypothalamus, Area praeoptica und Septum, sowie angrenzender Balken- und Stirnhirnbereiche. Monatsschr. Psychiatr. Neurol. 111, 17–59, 1946

215. Hess, W.R.: Die Relativität biologischer Reaktionen. Schweiz. Med. Wochenschr. 77, 3–9, 1947

216. Hess, W.R.: Stufen im Aufbau des vegetativen Nervensystems. Schweiz. Med. Wochenschr. 77, 659–665, 1947

217. Hess, W.R.: Vegetative Funktionen und Zwischenhirn. Helv. Physiol. Acta [Suppl.] IV, 1–65, 1947

218. Hess, W.R.: Das endophylaktisch-trophotropische System und seine Beziehung zur Symptomatik der Überempfindlichkeit. Schweiz. Med. Wochenschr. 77, 1151–1154, 1947

219. Hess, W.R.: Die funktionelle Organisation des vegetativen Nervensystems, pp. 1–226. Basel: Schwabe, 1948

220. Hess, W.R.: Zwischenhirn und Motorik. Helv. Physiol. Acta [Suppl.] V, 1–55, 1948

221. Hess, W.R.: Korrespondierende Symptome aus Stirnhirn, innerer Kapsel und vorderem Thalamus. Helv. Physiol. Acta 6, 731–738, 1948

222. Hess, W.R.: Kleine Beiträge zur Biologie der Wirkstoffe. Helv. Physiol. Acta 7, 178–188, 1949

223. Hess, W.R.: Das Zwischenhirn. Syndrome, Lokalisationen, Funktionen, pp. 1–187. Basel: Schwabe, 1949

224. Hess, W.R.: Die Bedeutung des Hypothalamus für die Regulierung vegetativer Funktionen. Dtsch. Arch. Klin. Med. 195, 55–57, 1949

225. Hess, W.R.: Das Zwischenhirn als Organisator vegetativer Funktionen. Nervenarzt 20, 533– 536, 1949

226. Hess, W.R.: Le sommeil comme fonction physiologique. J. Physiol. (Paris) 41, 61-A-67-A, 1949

227. Hess, W.R.: Motorik und Zwischenhirn. Arch. Psychiatr. Nervenkr. 184, 267–270, 1949

228. Hess, W.R.: Funktionen des Zwischenhirns im Rahmen der extrapyramidalen Motorik. Bull. Schweiz. Akad. Med. Wiss. 5, 221–226, 1949

229. Hess, W.R., Weisschedel, E.: Die höheren Reflexzentren der regulierten Körperhaltung. Helv. Physiol. Acta 7, 451–469, 1949

230. Hess, W.R., Koella, W., Szabo, Th.: Experimentelle Studien über die antagonistische Innervation. Z. Ges. Exp. Med. 115, 481–490, 1949

231. Hess, W.R.: Physiologie du thalamus. Rapport présenté au IVe Congrès Neurologique International, Paris 1949. Paris: Masson, 1950

232. Hess, W.R.: Symposion über das Zwischenhirn mit einleitenden Referaten und Demonstrationen. Helv. Physiol. Acta [Suppl.] VI, 5–80, 1950 197 233. Hess, W.R.: Die zentrale Regulation der Tätigkeit innerer Organe. In: Le Prix Nobel en 1949, pp. 115–123. Stockholm: Norstedt, 1950

234. Hess, W.R.: Die Kontrolle von Wirkstoffen am Irispräparat im isometrischen Verfahren. Helv. Physiol. Acta 8, C49–C50, 1950

235. Hess, W.R.: Überblick über die Dokumente von 25 Jahren Hirnforschung. Helv. Physiol. Acta 8, C50–C51, 1950

236. Hess, W.R.: Funktion und nervöse Regulation der inneren Organe. Vierteljahrsschr. Naturforsch. Ges. Zürich 95, 249–264 [p. 17], 1950

237. Hess, W.R., Akert, K.: Die Koordination morphologischer und physiologischer Untersuchungsmethoden in der Hirnforschung. Folia Psychiatr. Neurol. Neerl. 53, 268–274, 1950

238. Hess, W.R., Koella, W.: Experimentelle Studien über die antagonistische Innervation II. Z. Ges. Exp. Med. 116, 431–443, 1950

239. Hess, W.R., McDonald, D.: Film “Basilar artery”. Helv. Physiol. Acta 8, C51, 1950

240. Akert, K., Hess, W.R., McDonald, D.A.: Autonomy and somatic activity evoked by Stimulation of frontal cortex and thalamus in cats. (Film). J. Physiol. (London) 113, 19 P–20 P, 1951

241. Hess, W.R.: Die automatische Regulierung der Körperhaltung. Experientia 7, 51–58, 1951

242. Hess, W.R.: Prinzipien organischer Ordnung am Beispiel des vegetativen Nervensystems. Klin. Wochenschr. 29, 105–111, 1951

243. Hess, W.R.: Beitrag zur experimentellen Analyse des Stirnhirnes. Bull. Schweiz. Akad. Med. Wiss. 7, 295–306, 1951

244. Hess, W.R.: Methodische Erfahrungen über die experimentelle Erforschung des Zentralnervensystems. Helv. Physiol. Acta 9, 367–377, 1951

245. Hess, W.R., Akert, K.: Die corticale Repräsentation von Gesicht, Oral- und Pharyngealsphäre bei der Katze. Helv. Physiol. Acta 9, 269–289, 1951

246. Hess, W.R., Weisschedel, E.: Aus dem Zwischenhirn ausgelöste motorische Symptome an den Extremitäten und im Gesicht. Nervenarzt 22, 14–22, 1951

247. Hess, W.R., Akert, K., McDonald, D.A.: Beziehungen des Stirnhirnes zum vegetativen System. Helv. Physiol. Acta 9, 101–124, 1951

248. Hess, W.R., McDonald, D.A., Livingston, R.B.: Head-turning and eye deviation evoked by cortical excitation in the freely moving cat. Fed. Proc. 10, 63, 1951

249. Hess, W.R.: Experimentalphysiologie und Psychologie. Helv. Physiol. Acta 10, 85–92, 1952

250. Hess, W.R.: Fundamentos de la regulación vegetativa. Folia Clin. Int. 2, 1–6, 1952

251. Hess, W.R.: Aktuelle Aufgaben der Physiologie. Wien. Klin. Wochenschr. 64, 621–623, 1952

252. Hess, W.R.: Vom Lichtreiz zur bildhaften Wahrnehmung. Helv. Physiol. Acta 10, 395–402, 1952

253. Hess, W.R.: Physiologische Grundlagen der Ästhetik. Helv. Physiol. Acta 10, 462–468, 1952 Bibliographie von Walter Rudolf Hess

International, Paris 1949. Paris: Masson, 1950

232. Hess, W.R.: Symposion über das Zwischenhirn mit einleitenden Referaten und Demonstrationen. Helv. Physiol. Acta [Suppl.] VI, 5–80, 1950

233. Hess, W.R.: Die zentrale Regulation der Tätigkeit innerer Organe. In: Le Prix Nobel en 1949, pp. 115–123. Stockholm: Norstedt, 1950

234. Hess, W.R.: Die Kontrolle von Wirkstoffen am Irispräparat im isometrischen Verfahren. Helv. Physiol. Acta 8, C49–C50, 1950

235. Hess, W.R.: Überblick über die Dokumente von 25 Jahren Hirnforschung. Helv. Physiol. Acta 8, C50–C51, 1950

236. Hess, W.R.: Funktion und nervöse Regulation der inneren Organe. Vierteljahrsschr. Naturforsch. Ges. Zürich 95, 249–264 [p. 17], 1950

237. Hess, W.R., Akert, K.: Die Koordination morphologischer und physiologischer Untersuchungsmethoden in der Hirnforschung. Folia Psychiatr. Neurol. Neerl. 53, 268–274, 1950

238. Hess, W.R., Koella, W.: Experimentelle Studien über die antagonistische Innervation II. Z. Ges. Exp. Med. 116, 431–443, 1950

239. Hess, W.R., McDonald, D.: Film “Basilar artery”. Helv. Physiol. Acta 8, C51, 1950

240. Akert, K., Hess, W.R., McDonald, D.A.: Autonomy and somatic activity evoked by Stimulation of frontal cortex and thalamus in cats. (Film). J. Physiol. (London) 113, 19 P–20 P, 1951

241. Hess, W.R.: Die automatische Regulierung der Körperhaltung. Experientia 7, 51–58, 1951

242. Hess, W.R.: Prinzipien organischer Ordnung am Beispiel des vegetativen Nervensystems. Klin. Wochenschr. 29, 105–111, 1951

243. Hess, W.R.: Beitrag zur experimentellen Analyse des Stirnhirnes. Bull. Schweiz. Akad. Med. Wiss. 7, 295–306, 1951

244. Hess, W.R.: Methodische Erfahrungen über die experimentelle Erforschung des Zentralnervensystems. Helv. Physiol. Acta 9, 367–377, 1951

245. Hess, W.R., Akert, K.: Die corticale Repräsentation von Gesicht, Oral- und Pharyngealsphäre bei der Katze. Helv. Physiol. Acta 9, 269–289, 1951

246. Hess, W.R., Weisschedel, E.: Aus dem Zwischenhirn ausgelöste motorische Symptome an den Extremitäten und im Gesicht. Nervenarzt 22, 14–22, 1951

247. Hess, W.R., Akert, K., McDonald, D.A.: Beziehungen des Stirnhirnes zum vegetativen System. Helv. Physiol. Acta 9, 101–124, 1951

248. Hess, W.R., McDonald, D.A., Livingston, R.B.: Head-turning and eye deviation evoked by cortical excitation in the freely moving cat. Fed. Proc. 10, 63, 1951

249. Hess, W.R.: Experimentalphysiologie und Psychologie. Helv. Physiol. Acta 10, 85–92, 1952

250. Hess, W.R.: Fundamentos de la regulación vegetativa. Folia Clin. Int. 2, 1–6, 1952

251. Hess, W.R.: Aktuelle Aufgaben der Physiologie. Wien. Klin. Wochenschr. 64, 621–623, 1952

252. Hess, W.R.: Vom Lichtreiz zur bildhaften Wahrnehmung. Helv. Physiol. Acta 10, 395–402, 1952

253. Hess, W.R.: Physiologische Grundlagen der Ästhetik. Helv. Physiol. Acta 10, 462–468, 1952

198 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich

254. Hess, W.R., Akert, K., McDonald, D.A.: Functions of the orbital Gyri of cats. Brain 75, 244– 258, 1952

255. Hess, W.R.: Vergleichende Physiologie und pharmakologische Untersuchungen der Iris- Muskulatur. Arch. Exp. Pathol. Pharmakol. 219, 284–290, 1953

256. Hess, W.R.: Die physiologische Interpretation pathologischer Symptome aus dem Hirnstamm. Schweiz. Arch. Neurol. Psychiatr. 72, 405–406, 1953

257. Rüegg, J.C., Hess, W.R.: Die Wirkung von Adrenalin, Nor-Adrenalin und Acetyl-cholin auf die isolierten Irismuskeln. Helv. Physiol. Acta 11, 216–230, 1953

258. Hassler, R., Hess, W.R.: Experimentelle und anatomische Befunde über die Drehbewegungen und ihre nervösen Apparate. Arch. Psychiatr. Nervenkr. 192, 488–526, 1954

259. Hess, W.R.: Das Zwischenhirn, Syndrome, Lokalisationen, Funktionen, 2nd ed. pp. 1–218. Basel: Schwabe, 1954

260. Hess, W.R.: Das vegetative Funktionssystem im Lichte der experimentellen Forschung. Neuralmed. 2, 240–251, 1954

261. Hess, W.R.: Diencephalon. Autonomy and extrapyramidal functions, pp. 1–79. New York: Grüne & Stratton, 1954

262. Hess, W.R.: Übergeordnete corticale Zentren vegetativer Regulationen? Ber. Ges. Physiol. Exp. Pharmakol. 172, 116, 1955

263. Hess, W.R., Akert, K.: Experimental data on role of hypothalamus in mechanism of emotional behavior. Arch. Neurol. Psychiatr. (Chicago) 73, 127–129, 1955

264. Hess, W.R.: Hypothalamus and Thalamus. Documentary pictures - Hypothalamus und Thalamus-Experimental-Dokumente. Atlas with German and English legends, pp. 1–70. Stuttgart: Thieme, 1956

265. Hess, W.R.: Beziehungen zwischen psychischen Vorgängen und Organisation des Gehirnes I. Stud. Gen. 9, 467–479, 1956

266. Hess, W.R., Meyer, A.E.: Triebhafte Fellreinigung der Katze als Symptom diencephaler Reizung. Helv. Physiol. Acta 14, 397–410, 1956

267. Hess, W.R.: Beziehungen zwischen psychischen Vorgängen und Organisation des Gehirnes II. Stud. Gen. 10, 327–339, 1957

268. Hess, W.R.: Experimentelle Medizin der Gegenwart. Schweiz. Med. Wochenschr. 87, 1177– 1188, 1957

269. Hess, W.R.: Die Formatio reticularis des Hirnstammes im verhaltensphysiologischen Aspekt. Arch. Psychiatr. Z. Neurol. Nervenkr. 196, 329–336, 1957

270. Hess, W.R.: Physiologische Aspekte extrapyramidaler Motorik. Schweiz. Arch. Neurol. Psychiatr. 80, 299–305, 1957

271. Hess, W.R.: Die vegetative Innervation des Kortex. Acta Neuroveg. 16, 5–8, 1957

272. Meyer, A.E., Hess, W.R.: Diencephal ausgelöstes Sexualverhalten und Schmeicheln bei der Katze. Helv. Physiol. Acta 15, 401–407, 1957

273. Hess, W.R.: Von den integrierenden Funktionen des Diencephalon. In: Pathophysiologia Diencephalica. Internationales Symposium Mailand, May 1956. Curri, S.B., Martini, L. (eds.), pp. 210–216. Wien: Springer, 1958

274. Hess, W.R.: Objektive und subjektive Aspekte des Lärms. Z. Präventivmed. 4, 1–2, 1959

199 Bibliographie von Walter Rudolf Hess

275. Hess, W.R.: Kritische Bemerkungen zum Hirnreiz versuch. Pflügers Arch. Ges. Physiol. Menschen Tiere 270, 9–11, 1959

276. Hess, W.R.: Zentrale Einwirkungen auf die Muskeltätigkeit. Bull. Schweiz. Akad. Med. Wiss. 16, 398–401; 421–422, 1960

277. Hess, W.R.: Schlaf, sein Mechanismus und seine physiologische Funktion. Natur- wiss. Rundschau 14, 1–7, 1961

278. Hess, W.R.: Die physiologische Grundlage der Psychosomatik. Dtsch. Med. Wochenschr. 86, 3–8, 1961

279. Hess, W.R.: Psychophysiology and Psychopharmacology. In: Neuropsychopharmacology, Vol. 2. Rothlin, E. (ed.), pp. 3–8. Amsterdam: Elsevier, 1961

280. Akert, K., Hess, W.R.: Über die neurobiologischen Grundlagen akuter affektiver Erregungszustände. Schweiz. Med. Wochenschr. 92, 1524–1530, 1962

281. Hess, W.R.: Psychologie in biologischer Sicht, pp. 1–120. Stuttgart: Thieme, 1962

282. Hess, W.R.: „Kausalität“als psychophysiologischer Begriff. Schweiz. Arch. Neurol. Neurochir. Psychiatr. 91, 59–64, 1962

283. Hess, W.R.: Etude des corrélations entre l’activité psychique et le cerveau. J. Psychol. Norm. Pathol. 59, 369–378, 1962

284. Hess, W.R.: From medical practice to theoretical medicine: An autobiographic sketch. Perspect. Biol. Med. 6, 400–423 [p. 309], 1963

285. Hess, W.R.: Prinzipien cerebraler Organisationen. Experientia 19, 511–513, 1963

286. Hess, W.R.: Psicologia y biologia. La psicologia desde el punto de vista biologico. Morata: Filosofla, Psicologia, Pedagogica. Madrid (Registro No. 3112/63), 1963

287. Hess, W.R.: Problemas planteados en los limites de la psicofisiologia. Folia Clin. Int. 14, 3–6, 1964

288. Hess, W.R.: Die Motivierung bewußten Verhaltens. Schweiz. Arch. Neurol. Neurochir. Psychiatr. 93, 338–341, 1964

289. Hess, W.R.: The biology of mind, pp. 1–203. Chicago: University of Chicago Press, 1964

290. Hess, W.R.: Psychische Manifestation cerebraler Organisation. Arch. Psychiatr. Nervenkr. 205, 333–348, 1964

291. Hess, W.R.: Der Schlaf als Phänomen des integralen Organismus. Praxis 53, 1355–1357, 1964

292. Hess, W.R.: Gehirn und psychische Begabung. Bull. Schweiz. Akad. Med. Wiss. 21, 43–50, 1965

293. Hess, W.R.: Von Beziehungen zwischen Psychologie und Psychiatrie. Schweiz. Med. Wochenschr. 95, 317–319, 1965

294. Hess, W.R.: Sleep as a phenomenon of the integral organism. In: Sleep mechanisms. Progr. Brain Res. 18, 3–8. Akert, K., Bally, C., Schadé, J.P. (Eds.). Amsterdam: Elsevier, 1965

295. Hess, W.R.: Cerebrale Organisation somatomotorischer Leistungen. I. Physikalische Vorbemerkungen und Analyse konkreter Beispiele. Arch. Psychiatr. Nervenkr. 207, 33–44, 1965

296. Hess, W.R.: Cerebrale Organisation somatomotorischer Leistungen. II. Experimentelle Untersuchungen über die zentrale Koordination der Motorik. Arch. Psychiatr. Nervenkr. 208, 209–233, 1966

297. Hess, W.R.: Causality, consciousness, and cerebral organization. Science 158, 1279–1283, 1967 200

298. Hess, W.R.: Hypothalamus und Thalamus. Experimental-Dokumente. 2nd ed. pp. 7–77. Stuttgart: G. Thieme, 1968

299. Hess, W.R.: Psychologie in biologischer Sicht. Mit kritischen Beiträgen von K. Akert, G. Baumgartner, R. Hess, jr., R. Hunsperger, R. Jung, P.G. Wasser, G. Weber. 2nd ed. pp. 1– 132. Stuttgart: Thieme, 1968

300. Hess, W.R.: Hypothalamus and thalamus. Experimental documentation. 2nd ed. pp. 1–77, enlarged edition. Stuttgart: G. Thieme, 1969

301. Hess, W.R.: Polarität regulierender Funktionen und zerebrale Organisation. Dtsch. Med. Wochenschr. 95, 1672–1675, 1970

302. Hess, W.R.: Interdisciplinary discussion of selected problems with reference to “The biology of mind”. Perspect. Biol. Med. 13, 267–293, 1970

303. Hess, W.R.: Ein Beitrag zur Physiologie des Kleinhirns. Zentrale Organisation somatomotorischer Leistungen. Biol. Zbl. 89, 457–466, 1970

304. Hess, W.R., Fischer, H.: Brain and consciousness. A discussion about the function of the brain. Perspect. Biol. Med. 17, 109–118, 1973 Klinisches Neurozentrum | Prof. em. Dr. med. A. Valavanis | UniversitätsSpital Zürich Untersuchungen über die zentrale Koordination der Motorik. Arch. Psychiatr. Nervenkr. 208, 209–233, 1966

297. Hess, W.R.: Causality, consciousness, and cerebral organization. Science 158, 1279–1283, 1967

298. Hess, W.R.: Hypothalamus und Thalamus. Experimental-Dokumente. 2nd ed. pp. 7–77. Stuttgart: G. Thieme, 1968

299. Hess, W.R.: Psychologie in biologischer Sicht. Mit kritischen Beiträgen von K. Akert, G. Baumgartner, R. Hess, jr., R. Hunsperger, R. Jung, P.G. Wasser, G. Weber. 2nd ed. pp. 1– 132. Stuttgart: Thieme, 1968

300. Hess, W.R.: Hypothalamus and thalamus. Experimental documentation. 2nd ed. pp. 1–77, enlarged edition. Stuttgart: G. Thieme, 1969

301. Hess, W.R.: Polarität regulierender Funktionen und zerebrale Organisation. Dtsch. Med. Wochenschr. 95, 1672–1675, 1970

302. Hess, W.R.: Interdisciplinary discussion of selected problems with reference to “The biology of mind”. Perspect. Biol. Med. 13, 267–293, 1970

303. Hess, W.R.: Ein Beitrag zur Physiologie des Kleinhirns. Zentrale Organisation somatomotorischer Leistungen. Biol. Zbl. 89, 457–466, 1970

304. Hess, W.R., Fischer, H.: Brain and consciousness. A discussion about the function of the brain. Perspect. Biol. Med. 17, 109–118, 1973

201

8. Nachrufe auf Walter Rudolf Hess Nachrufe auf Professor Walter Rudolf Hess

Akert K. Obituary: Walter Rudolf Hess (1881–1973). Brain Res. 1974;68: V–VIII.

Akert K. Walter Rudolf Hess (1881–1973) and his contribution to neuroscience. J Hist Neurosci. 1999; 8:248–63.

Akert K, editor. Biological Order and Brain Organization. Selected Works of W. R. Hess. Berlin, Heidelberg, New York: Springer; 1981 . Auerswald W. Nachruf auf Walter Rudolf Hess. Almanach Österr Akad Wissenschaft. 1975; 124:414– 33 . Caspers H, Hess R, Kugler J, Petsche O, Struppler A. Nachruf auf W. R. Hess. Z EEG EMG. 1973; 5:139–41.

Davison J, Wilson G. Walter Rudolf Hess. Clin Experiment Ophthalmol. 2005; 33:517.

Huber A. W. R. Hess, the ophthalmologist. Experientia. 1982; 38:1397–400.

Jung R. Walter R. Hess (1881–1973). Rev Physiol Biochem Pharmacol. 1981; 88:1–21.

Koelbing H.M.F.: Walter Rudolf Hess. Historisches Lexikon der Schweiz 33, 2007

Koella WP. A tribute to W. R. Hess (1881–1973). Experientia. 1982; 38:1383.

Masneuf S., Baumann C.R., Hess C.W., Valko P.O.: Walter Rudolf Hess (1881-1973) J Neurol 262:2198-2199, 2015

Shampo M.A., Kyle R.A., and Steensma D.P.: Walter Hess-Nobel Prize for Work on the Brain. Mayo Clin Proc. 86 (10): e49, 2011

Waser PG. Walter Rudolf Hess: Aus seinem Leben und seiner Tätigkeit an der Medizinischen Fakultät der Universität Zürich (100-Jahr-Feier seines Geburtstages: 14. März 1981). Gesnerus. 1982; 39:279– 86.

Wyss O.A.M.: Walter Rudolf Hess 17.3.1881–12.8.1973; Die Selbständigkeit der Physiologie. Schweiz Med Wochenschr. 1973; 103:1758–9.

Wyss O.A.M.: Zum Hinschied von Walter Rudolf Hess. Neue Zürcher Zeitung 382:20. August 1973

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