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AH OFFICE - LONDON, ENGLANDTECHNICAL REPORT

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SOME ASPECTS OF RESEARCH IN PHYSIOLOGY,NEUROPHYSIOLOGY AND NEUROANATO[Y AT THE

UNIVERSITY OF OSLO, NORWAY

by

C.N. PEISS

10 January 1966

This document Is subject to special export controls and each

transmittal to foreign governments or foreign nationals may be'ONDO1 6 made only with prior approval of the Commanding Officer, Office

of Naval Research Branch Office, Box 39, FPO New York, 09510.

Vi

SOME ASPECTS OF RESEARCH IN PHYSIOLOGY, NEUROPHYSIOLOGY ANDNEUROANATOMY AT THE UNIVERSITY OF OSLO, NORWAY

The Institutes of Physiology and Anatomy, and the Neuro-physiology Laboratory, are located in the old University centerin Oslo, a few blocks from the bustling and attractive center ofthe city. Most of the other divisions of the University, whichnow has some 10,000 students, are located at a new, ultramoderncawnus on the outskirts of the city. It is expected that themedical institutes will be located there in about five years.

It is not the intent of this report to summarize super-ficially the research programs of all members o!' each of theseinstitutes. Rather, it is my purpose to report in some detailcertain areas of research by one or more individuals which, inmy opinion, represent significant contributions.

THE INSTITUTE OF PHYSIOLOGY

The Institute of physiology is under the direction ofa new Head, Prof. Bjprne A. Waaler. Waaler io an extremely ener-getic and bright young man, who is capable of establishing aproductive group, provided the necessary support is received. Hehas worked in the past with I. de Burgh Daly in London. He iskenly interested in the teaching of medical physiology, andsome of the problems and ideas with which he and other Europeanphysiologists are concerned will be ,the subject of a later report.Waaler has been Head of the Institute of Physiology for about twoyears, and now has several research projects in operation. Heis still handicapped by a lack of adequate funds, but the situ-ation is improving steadily. The researc'h equipment of the In-stitute is relatively outdated, most recording still being doneby kymograph. -However, a new four-channel recorder is now in use.

The professional staff of the Institute of Physiologyconsists of Waaler (Professor), Leif Erikeon (Associate Professor),Johan Steen, Egil Amundsen and Yngve L~yning (Established ResearchWorkers), two scientific assistants and four research fellows,two of whom are supported by the Norwegian Research Council. Theprimary areas of research include projects on hemoglobin synthe-sis, plasma kinins, gas exchange in fishes, and pulmonary circuls-tiono

WUaler has recently begun work on pulmonary blood flowin an isolated, perfused rabbit lung preparation. He is record-ing pulmonary artery pressure, ventilation and lung weight withfairly crude techniques, but ones adequate to yield the necessarydata. This project is now being converted for rtoording with a

I

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four-channel direct ink-writer, which should provide both more precisemeasurements and the capability of adding other parameters of measure-ment. This work has reached the stage where a first mt ascript isin preparation. The isolated lung is perfused at a constant, non-pulsatile pressure. After a variable period of time, a significantincrease in pulmonary artery pressure occurs spontaneously, Indicat-ing an intense vasoconstriction. During this time no significantchange occurs in ventilation. This vasoconstriction is reversedcompletely by tri-cresol, which appears to inhibit ATP specifically.Vasoconstriction produced by a variety of pulmonary constrictor drugsincluding norepinephrine and several polypeptides is not affected byadmiiistration of tri-cresol. This research, which is in its in-cipient stages, requires considerable expansion. The observed factof this remarkable specificity of tri-cresol in inhibiting a spontan-eously occurring pulmonary vasoconstriction is of great interest andpotential value. However, much remains to be done concerning thenature of the spontaneous constriction in this kind of preparationand concerning the mechanism of action of tri-cresol in reversingthis constriction. To my knowledge there has not been any attemptto assess the histological status of the isolated lung preparationafter varying periods of perfusion. This information should beobtained as well as studies on alveolar gas exchange, etc.

In a previous report it was demonstrated that certailnpolypeptides (bradykinin, kallidin and eledoison) all producepulmonary vasoconstriction in an isolated rabbit lung preparation.Bradykinin was effective in doses as low as 1-5 micrograms, andwas more potent, on a molar basis, than 5-hydroxytryptamine or norepi-nephrine. Kallidin was 30-50% more effective in producing vasocon-striction than was bradykinin, and eledoisin was strikingly morepowerful then either of the other two polypeptides, causing markedpulmonary vasoconstriction in doses smaller than .001 micrograms.Responses to polypeptide injection were not affected by doses ofRegitine which completely blocked vasoconstrictor responses to nor-epinephrine, nor by methysergide which was effective in blockingthe response to 5-hydroxytryptamine. This evidence suggests thatthe mechanism of action of polypeptides in producing pulmonaryvasoconstriction does not operate by way of liberation of catechol-amines or 5-hydroxytryptamine.

A great deal of research effort in the Institute ofPhysiology in concerned with studies on plasma kinins and intestinalobstruction -- areas which I am not competent to assess critically.It might be useful, however, to list some recent publications inthese fields from the Institute of Physiology:

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Amundsen, E. and Gustafsson. B.E. Results-of experimental intest-inal strangulation obstruction in germ-free rats. J. Exp. Med.117, 823-33 (1963).

Amundsen, E. and Midtvedt. T. The toxicity of fluid from exper-imentally strangulated intestinal loops in the rat. J. Surg.Res. k, 306-13 (1964).

Hauge, A., Amundsen, E. and Wasler, B.A. Changes in blood pro-accelerin (Factor V) in rats with intestinal strangulationobstruction. Thromb. et Diath Haemmor. 12, 49-63 (1964).

Amundsen, E.. Waaler. B.A., Dedichen, J.. Laland, P.. Laland, S.,z~n! Thorsdalen, N. Kinase inhibition by a fluorescent sub-stance prepared from liver. Nature 203, 1245-48 (1964).

Amundsen, E. and Nustad, K. Kinin-forming and destroying activ-ities of saliva. Brit. J. Pharm. & Chemotherapy 2, 440-44 (1964).

Amundsen, E. and Rugstad. H.E. Influence of some pathogenicbacteria on kinin formation and destruction. Brit. J. Pharm. &Chemotherapy ?2, 67-73 (1-965).

Amundsen, E. and Nustad. K. Kinin-forming and destroying activ-ities of cell homogenates. J. Physiol. 72, 479-88 (1965).

With the addition of Dr. Johan B. Steen to the Insti-tute, a new area of research has been instituted. Steen hasbeen working in a field best covered by the term "comparativephysiology," including studies on oxygen exchange in smallaquatic animals with a modified Scholander respirometer, thermo-regulation in birds and gas exchange in fish. Together withi. Steen, he published an interesting paper in Acta physiol.scand. §2, 285-91 (1965), entitled, "The Importance of the Legsin the Thermoregulation of Birds." In studies on herons andgulls it was shown that the entire beat production of the birdcould be lost through the legs at high environmental air temp-eratures (5 0C). At 200 C, about one-half the total heat pro-duction could be dissipated from the legs. Heat loss from thelegs to water was found to be about fourfold that from the legsto air. The physiological mechanism responsible for controllingthe rate of heat dissipation from the legs (presumably :-aecularcontrol) responds rapidly to sudden changes in environmentalair temperature. This was studied by measuring the water temp-erature surrounding one leg of a heron while the environmentaltemperature was rapidly lowered from 35°C to 50 C. During thecontrol period at an air temperature of 350C, water temperaturesurrounding the leg increased linearly at a rate of about 0.180 C/min. Thirty seconds after a rapid decrease in environmental

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temperature, heat loss from the leg into the surrounding water haddecreased markedly to the point where the change in water tempera-ture was only O.O3C/mimite. Although the authors made no measure-nent of hypothalamic temperatures, it was felt that the rapidityof this response indicated a reflex initiated by peripheral re-captors. The vascular arrangements of the leg in many birds pro-vide the possibility of countercurrent heat exchange, as well asshunts that can permit a bypass flow of blood around the retia.It would be of great interest if Steen would investigate the nervouscontrol of these blood vessels as an extension of this work.

TIHE NEUROPHYSIOLOGICAL LABORATORY

This laboratory, which is extremely active in research,operates essentially as a fully independent department. It is di-rected by Prof. Birger Kaada, who has made oustanding contributionsin the field of CNS control of autonomic function and various aspectsof behavioral neurophysiology, especially as related to the limbicsystem. Kaada has spent considerable time in the United States,working with Pribram and with'John Fulton at Yale. He has alsoworked at various times with Jasper in Montreal and Granit in Stock-holm. The Neurophysiological Laboratory is presently located indiffuse quarters in the Anatomical Institute. There is a shortageof adequate space, but an obviously maximal use of what is available.Within a few months the Neurophysiological Laboratory will move intolarger quarters in the Institute of Physiology.

The department is well equipped with much of the electronicinstrumentation that is so necessary in modern neurophysiological re-search. However, certain of the research problems under investigationhave reached a stage where a new generation in electronic instrumenta-tion is required. This will essentially involve computers, and willraise problems for the department in obtaining the sizeable funds in-volved. The research funds available to the Neurophysiological Lab-oratory have grown steadily in Kaadata 15-year tenure. In recentyears research support ba been provided by the following sources:

1. The Norwegian Research Council for Science and the Humanities

2. The Norwegian Council on Cardiovascular Diseases

3. The Anders Jahres Foundation

4. The FridtJof Nansen Foundation

5. United States Public Health Service, National Institutesof Health

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6. The Rockefeller Foundation

7. European Office, Air Research and Development

8. The Wellcome Trust, London

The permanent staff consists of Prof. Birger Kadaand Drs. Jan Jansen, Jr., Per Andersen and Holger Ursin.Andersen has had two years in Eccles' laboratory in Australia(1961-63), from which a remarkable series of excellent publica-tions was derived. Ursin was in the United States during 1962-64, spending 18 months 4t the U~iversity of Chicago, two monthsat Michigan and tiaree months at NI.. Inaddition to the perm-anent staff there are 13 research fellows working in the Neuro-physiological Laboratory, as well as several foreign guests.

The research activities of this department can beroughly grouped into six areas:

A. Autonomic nervous system - Endocrinology

B. Control of movement

C. 3leep mechanisms - Reticular formation

D. Behavioral studies in animals

E. Muscle and nerve

F. Physiology of synapses

The current status of research in some of these areas is reviewedbelow.

A. Autonomic nrvous amtes - Endocrinology

The primary interest in recent years he been the influ-oace of the autonomic nervous system on uterine motility Inestrogenined rabbits, and on the responses of the urinary bladderin cats.

I. Uterine Motility

Rapid distention of the myometrium produces aquick contraction followed by inoreoed rhytbhio activity, thefrequency at which is increased with the degree of distention.This pacemaker activity is directly related to the tension inthe myornetrium. The myeenio nature at tOs response to disten-tion is indicated by the lock at effeet of autonomio blooking

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drugs and local or spinal anesthesia, as wel. as the independenceof the two uterine horns in respose to distention. Stimulationof +he hypogastric nerve produces maximal uterine contraction at afroquency of 30-50 cps. Stimulation for only a few seconds resultsin a single contraction, whereas prolonged stimulation at low fre-quency results in a series of rhythmic contractions. Evidence hasbeen gathered which demonstrates the presence of adrenergic fibersin the hypogastric supply to the myometrium. Similar evidence forthe presence of oholinergic fibers is lacking. Splanchnic nervestimulation produces qualitatively similar responses, but of longer.ltency and more gradual onset.

It has been proposed by these workers that the sympatheticnervous system exerts a tonic influence on the uterus. It has beendemonstrated that a variety of sensory inputs can influence uterinemotility, e.g., stimulation of the rectum, and electrical stimulation ofvarious visceral and somatic afferent nerves. In addition, similarresponses in uterine motility can be obtained from asphyxia and car-otid artery occlusion. Responses to all of these stimuli appear tobe the result of sympatho-adrenal activation, the effector actionbeing an initial uterine contraction followed by a period of de-creased rhythmic activity. This effector responee.is eliminatedby sympathetic blocking agents, as is the inhibition of "spontane-ous" rhythmic contractions produced by stimulation of the depressornerve. The latter response presumably is mediated by a sympatho-inhibitory reflex.

Various parts of the brain were explored in order to mapthose areas which influenced uterine motility. In the medull*,excitatory responses were elicited from the lateral part, and in-hibitory responses from the midline region. This distribution issimilar to the general pattern of sypathetic influences of themedulla on other autonomic efteotars such as the blood vessels,Stimulation of the parifornical, dorsal, lateral and posteriorhypothalamus produced excitatory responsese Zxcitation of uterinemotility was also elicited by stimulation of the ventromdial hypo-thalamus, but this was due to oxytocin release. Inhibition wasobserved when a restricted area of the lateral hypothelelaw wasstimulated. Finally, exltcatry responses were also derived fromthe sonsorinotor cortex, anterior 1lmbic region and anygdaleoSeveral oortioil areas produced inhibition of uterine motility,including the senorimoter cortex and orbital surface.

2* Urinary bladder motility

Studies have been carried out to. determine ifiatherthere are antagonistic effects between syupetbetio and peresywps-thetie nerves to t1M urinary bladder. Intravesioular pressure wesrecorded by droot cannuletion of the bladder, leaving the urethral

ONRL-1-66 7

outlet intact. In this preparation, bilateral section of theparasympathetic nerve supply results in abolition or reductionof rhythmic activity. Subsequent denervation of sympatheticfibers causes resumption of rhythmic contractions, althoughthey are of smaller amplitude. Reversal of the sequence ofdenervations produces the expected findings, i.e., hyperactivityafter sympathectomy of the bladder, followed by restoration ofmore normal contractions when the parasympathetic nerves are out.A similar study was performed to determine antagonistic influ-ences of the autonomic innervation on micturition. Section ofparasympathetic fibers abolished the micturition reflex, whereasprimary sympthectomy reduced the threshold for micturition, thusdemonstrating an inhibitory role of sympathetic nerve fibers onthe micturition reflex. Finally, studies were carried out onsupraspinal influences upon the induction or inhibition of mic-turition. In the latter case, micturition was induced by rapidfilling of the bladder. Excitatory areas were found in theamygdala and pyriform cortex (corticomedial area), ventromedialhypothalamus, posterior cingulate gyrus and upper brain stem ret-icular formation. The excitatory response from all regions waseliminated by denervation of parasympathetic nerves to thebladder, but were unaltered by sympthectomy. Inhibitoryresponses from supraipinal areas were abolished by sympatheticdenervation of the bladder.

B. Control of llovoment

A number of experimental approaches are being made toproblems dealing with muscle receptors and fusimotor activityin the cat, much of which cannot be reviewed here due to apacelimitations. However, a brief description can be presented ofa very significant study on the response patterns of nervefibers in the dorsal spinooerebellar tract. Three differenttypes of units in second order neurons of the dorsal spino-cerebellar tract have been identified during the response tomuscle stretch. The response in single units of these secondorder neurons is very similar in general to that of the corre-sponding receptor in the muscle. Thus the cerebellum is pro-vided with threse distinct items of information about the mabon-tcal status of the muscle: (1) a mixed signal of muscle lengthand rate of movement (;.'oup Is unte)l M a snsl primarilyof muscle length (Group 11 units)i nnd 3 a sal related to tbadegree of contraction in the muscle (Group lb units).

C. Slosh Leohaniams - ealticular Formtion

Tt has been known that EM0 synohronization end behavior-.1 sleep can be produced in unansetbetimed animals by low frequency

8 ONRL-1-66

stimulatiorn of the preoptic region, retioular formation, caudoteand medial thalamus. Recent experiments at the NeurophysiologicalLaboratory in Oslo have shown that EEG synchronisation and behavior-al drowsiness can also be evoked by low frequency (6 cpa) stimula-tion of the suballosal cortex. It was not possible to induce deepsleep from this area and the effects were less marked than thosefrom the preoptic region. It has been proposed that this cortical&rep is part of a descending ndeaotiveting" and inhibitory systemwhich sats on auboortical sloep-produoiug mechanism.

In another series of experiments It has been shown thathk frequency (200-300 cps) stimulation in the midbrain in lightlyanesthetized oats may on occasien produce EEG synchronisation, i.e.,immediate anl wide-spread increase in cortical slow wave activity.This response Is not obtainable in unanesthetized animals, and thesynchronisation response shows higher threshold and finally disap-pears as the depth of anesthesia is increased. The response couldbe obtained from points in the entire midbrain reticular formation.An area of low threshol for this response was discovered in thelower midbrain at the decusestion of the brachium conjunctivum.This area of low threshold, however, appears to be independentof the brachium conjunctivum, since the response was still presentin chronic animals following removal of the cerebellum.

). 2havier Studies in Animals

Several years ago eeds and his associates demonstrateddeficit in the perfozmance of rats in mce loarning followug

'bilateral lesions In the hippo mpus. Other workers have failedto observe this effect. Recent experiments by Xands and asesooiateshave oonfirsmed the original findig, and hae also obtained evidencethat the type of mae used to disorilmote lesions is of great im-portonoe. In those experiments it was also found that differentbrain structures are involved in mae learning and in passiveavoidance learning, slum the latter was not affeoted by the hippo-campe lesos.

Another series of experiments has been performed tostudy the effect of torebrain 1.siom on sex incentive. Thecriterion o sex incentive was the nuer of crossings or aneleotrifiod grid toward a rat at olpeite oex. Us mmber orcrossins ws aignifloantly dereesed by bilateral destruotion at(1) upper portion or senwrimoter oartex (2) auidola; and (3) theolfactory bulb. Increased munber o eroossing wre observed terbilateral lesions in (1) septal aree (2) temparel-nsular cortex#end (3) atria mdullaro amd habemuleo The decrease in inoent iveproduced by any at the above lesaion could be counteracted by bi-lateral lesions in any of the above-ummd areas wbioh increasedIncentive. Otbhor aspects at te animals' response and physiology

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remained normal (spontaneous motor activity, sensitivity tothe grid, estrous cycle, etc.).

THE ANATOMICAL LASTITI'E

This Institute ib directed by Prof. Jan Jansen. Ihad hoped to spend considerable time talking with Prof. AlfBrodal about his neuroanatomical studies, but had to settlefor a brief conversation. Brodal at present is serving a three-year term as Dean of the Faculty of Medicine, which has limitedhis time for research in the poet few years.

Brodal has recently carried out experiments on thetermination of primary veatibulocerebellar fibers in the cat.Sterile lesions of the vestibular nerve were made, and animalssacrificed from 5-19 days afterwards. Distribution of degener-ating fibers was studied from brain sections treated by twosilver impregnation methods, those of Nauts and of Glees.Primary degenerating fibers were found in the ipatlaterelcerebellar cortex in the nodulus, floeculus, uvula, and ventralperaflocculus. There was also evidence of termination of somefibers in the dorsal paraflocoulue, lingule and dentate nucleus.A few fibers distributed to the controlateral half of the cere-bellum, reaching corresponding areas. There was no evidence,however, of a contribution of fiers from the vestibular nerveto the medial fastigial nucleus. These findings indicate thatthe functional "vestibulocerebellum"t is not confined to theflooculonodular lobe* Moreover, there are morphological differ-ences between those receiving areas as compared with the restof the oerebellar cortex, which is contrary to the general viewthat this Is uniformly structured throughout,

When his term as Dean of ti Faculty of "edicine ends,Bradal expecte to continue his work on vestibular and cerebellarstructure, He Is currently prepared to undertake a precise studyusiuC oleotron microscopy. The complexities of the nervoussystem and the huge number of struotural elam nts involvedmake one admire the determization of Brodal to carry on hissuperb work at a nw level of resolution. As he pointed outdurirg our conversation, such an undertaking in only possiblewhen ae is thorou~hly familiar In precise detail with thestructure as soon under the light microsoope. lith his took-ground and knowing precisely what he wisheo to examine withelectron microsoopy, Brodal's first published reports in thisfield will be keenly anticipated.

For the peat -1520 years the reticular form tion be*been a subject at intealve li'ostigation by neuroaaetomiets andnurophyiolo~ists. It is an ares to which Brodal has made

10 ONRL-1-66

significant contributions. He has recently written a stimulatingpaper on the role of the retihular formation and other parts ofthe brain as related to consciousness. It is entitled "AnatomicalPoints of View on the Alleged Morphological Basis of Consciousness,"and appeared in Acts Neurochiruraica 12, 166-86 (1964). Perhapsbecause his exposition struck such a responsive chord in my ownfeelings on this subject, I think it worth-while to summarizebriefly the essence of his Wument.

Anatomically the reticular foration can be divided intoseveral cellular or nuclear groups. These groups differ both as totheir connections and cytoarchitecture. The most common element ofthe reticular formation is not the associational cell but, rather,long axons projecting in caudal and/or rostral direction. Brodalpoints out that many of the re-.ptox-a in the periphery give riseto specific fiber systems which do not send off collaterals to thereticular formation. This includes the medial lemniscus. He thenquestions the concept that the reticular formation works as anentity in the maintenance and level of consciousness. Finally,there is a discussion of the connections of the reticular forma-tion with many other areas of the brain such as sensory cranialnuclei, periaqueductal gray, cerebral cortex, limbic system, etc.,all of which may have a role in the process of consciousness.Brodal states the essence of his argument as follows: "We mayindeed ask whether it is not timely to deprive the RF of some ofits supremacy es concerns consciousness, and to consider this,like so many oiher functions, as-:eing more or less dependent onthe whole brain. It would perhaps help to clarify thoughts if weagreed to delete the word 'reticular' from the concept of the'reticular asconding activatir system'."

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"Some Aspects of Research in Physiology, Neurophysiologyand Neuroanatomy at the Univ. of Oslo, Norway"

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13. ABSTRACT

Visits were made by author to the Institutes of Physiologyand Anatomy anc! the Neurophysiology Laboratory, Univ. of Oslo,Norway, and this Tech. Report reports in detail certain re-search areas which are significant, e.g., 'Waaler's work onpulmonary blood flow in isolated, perfused rabbit lung prepara-tion, and production of pulmonary vasoconstriction by certainpolypeptides. Also discussed is Dr. J.B. Steen's work on thermo-regulation in birds, gas exchange in fish. The Neurophysiolo i-cal Lab's work on the autonomic nervous system (endocrinology)is reviewed, also control of movement, sleep mechanisms andreticular formation; behavioral studies in animals, including-effects of forebrain lesions on sex incentive. The work ofA. Brodal at the Anatomical Inst. on termination of primer-vestibulocerebellar fibers in the cat is reviewed.

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pulmonary blood flow, ventilation,9 vaso-consIric tion, norepinephrine, polypeptidefttri-'cres o1, specificity, spontaneous cons-triction, bradykinin. kinese, thermoregu-tation, heat loss, vascular. retia, auto-nlomlic; nervous system, sleep mechanisms,synapses. uterine motility, hypogastricnerve, cholinergic fivers. sympatho-inhibitory, brain, oxytocin, hypottelamus,rtTicular formati on. cerebellum, maze-learninig. vestibulocerebellar, cortex,ne rve~

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