thorax in history 2. hellenistic experiment and human · thorax, 1978, 33, 153-166...

Thorax, 1978, 33, 153-166

The thorax in history 2. Hellenisticexperiment and human dissectionR.K. FRENCH

From the Wellcome Unit for the History of Medicine, University of Cambridge

The first anatomical revolution occurred in Alex-andria in the third century before Christ. To thatperiod we can trace the continuous historical lineof our own ideas on the structure and function ofthe body, important among which are those on thethorax. For the first time, the human body wassystematically explored in an attempt to under-stand its structure, and, with some notable errorsfrom animal anatomy, the only source for 'human'anatomy in the preceding period, these anatomicalideas are recognisably similar to those of to-day.The physiological ideas of the Alexandrians, onthe other hand, are strikingly different from ours,and their subsequent transformation will be exam-ined in later articles.

Human dissection

The most important single reason behind this firstanatomical revolution was the introduction ofsystematic dissection of the human body. It wassuggested at the beginning of the first article inthis series (Thorax, 33, 10-18, 1978) that humandissection has been historically very rare and thatthere were normally a number of taboos operatingagainst it. We must briefly consider then why itwas that the human body could be dissected inAlexandria.

Alexandria was the city of Alexander the Greatand shared fully in the cultural and commercialexchanges of political empire. A cosmopolitanpopulation, the mixture of Hellenistic andEgyptian culture, and the patronage of the artsand sciences by the Ptolemies all contributed to acultural ferment. The changes that made Alex-andria the centre of Hellenistic culture after thedeath of Alexander meant that no single system ofphilosophy or belief held a dominant position,nor were its taboos universally accepted.Apart from such general considerations, there

are a number of particular reasons why humandissection should have flourished. The opening of

the dead body was essential to the ancient practiceof embalming, an Egyptian technique well knownto the Greeks. The Greek philosophers, Plato andAristotle, emphasised the distinction between souland body:' it was the immaterial soul that sur-vived the death of the corporeal body, and thesoul, sharing no characteristics with the body,could not be affected by the postmortem mutila-tion of the body; there was no quasi-material after-life in human form, as so many cultures believed.In a word, the old religious taboos no longerapplied.Of fundamental importance in the development

of human dissection in Alexandria was the exist-ence of a medical school (or schools). Even in theabsence of specific cultural taboos, the dissectionof the body of a person recently dead naturallyexcites a feeling of revulsion; only in a medicalschool, where anatomy is considered a basic medi-cal science and becomes an established discipline,is there sufficient mutual support within a groupto preserve an unpleasant practice, a sort of pro-fessional rejection of the normal taboos. Humandissection was a teaching device as well as a re-search tool.

Also associated with the development of medicalschools were medical and anatomical texts. Galen2tells us that the ancient method of teachinganatomy-he was thinking of the Hippocraticperiod-was oral, when the father (or master)taught his sons (or apprentices) within the familygroup over a period long enough for the assimila-tion of the great detail. Only when this systembegan to break down and adult men were admittedinto the circle to learn anatomy in a shorter periodwas it necessary to write down the anatomy.Perhaps this increasing formalisation of teachingwas becoming evident by Aristotle's time, for thefirst textbook of anatomy is said to have beenwritten by Diocles of Carystos, a contemporary ofAristotle.3 Many of the anatomical texts up tothe time of Galen (second century AD) were

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school textbooks, or at least the texts of pro-

fessional anatomy teachers. A written tradition,of course, is open to detailed and precise modifica-tion, and Galen's own treatment of the work ofhis predecessors is the beginning of textual andanatomical scholarship, upon which later develop-ments depended.A natural result of teaching within a school, as

opposed to a father-son line of oral instruction, is,in addition to texts, a comparatively large numberof students with similar ideas. Future teachers,chosen from the students, would perpetuate theseideas alongside the texts, so that the physicalschool becomes also a 'school of thought'.Aristotle's Lyceum became a school of peripateticphilosophy, opposed to others, like the atomist or

stoic. In the same way the medical schools de-veloped traditional ideas: from theoreticalpositions already available in the Hippocraticcorpus, and in parallel with the medical schoolsthat had already been in existence for two or threecenturies on the periphery of the Greek world,there developed in Alexandria, during the periodwe are discussing, a number of medical 'sects'with profoundly different views on the usefulnessof anatomy in medicine. The Dogmatists or

Rationalists, inclined to natural philosophy, heldthat it was possible in principle to understand theprocesses going on inside the body, and that there-fore the physician was justified in intervening inthese processes to put right what had gone wrong

in illness. As function was closely related to struc-ture, it followed that anatomy was a fundamentalmedical science. It is possible that this attitudewas reinforced by the observational, empirical,and inductive methodology that Aristotle hadannounced and put into practice in his zoologicalworks.

It appears that the Rationalists of Alexandriatook this methodology one step further andpractised not only dissection, to discover humananatomy, but vivisection,4 to discover the functionof the human body. In other words, they intro-duced experiment into the descriptive and observa-tional method. Both dissection and vivisectionwere opposed by the Empiricists,5 who fromHippocratic times had believed that the processesof the body were ultimately incapable of explana-tion. The business of the physician was not, there-fore, to intervene but to observe; empiricalmedicine emphasised prognosis based on case

history and treatment based on regimen. The non-

investigative empirical physician, rejecting theneed of a knowledge of function, had no need ofknowledge of structure, and he held not only that

anatomy was unnecessary but that the means of

acquiring anatomical knowledge-dissection-wasdisgusting and unnatural. It was, moreover, mis-leading, since the organs of the body changed indeath. It was undoubtedly to see the organs of theliving body that the Rationalists employed vivi-section, a technique that the Empiricists con-sidered inadmissible on the grounds of cruelty.The Empiricists argued that the only form of ana-tomical knowledge necessary and proper to a physi-cian was that gleaned 'accidentally' from wounds.

The influence of Aristotle

The question of the extent of Aristotle's influenceon the Alexandrian experimenters is not agreedamong historians, but a brief discussion of thetopic will serve to highlight some important issuesand to introduce the achievements of theAlexandrians themselves.Wellman and Jaegar argue that there was a

considerable line of influence from Aristotlethrough Diocles,6 whom we have already met asthe author of the first anatomical textbook. If it isindeed true that Diocles thought of himself as afollower of Aristotle, then his book on humananatomy might well have been designed to fill themost obvious gap in Aristotle's biological works.The fragments of Diocles' work that survive showthat his anatomy was in fact derived from animals,and it may be that, like Aristotle, he was unable todissect the human body and relied on amethodology that consciously extrapolated fromcomparative anatomy to human. All this, how-ever, is supposition, and all we can say with confi-dence of Diocles is that his treatment in generalis like that of Aristotle, and like Aristotle heplaced the soul in the heart. On the other hand,some important physiological and anatomicalideas of the Alexandrians and some of their widerbeliefs are quite opposed to Aristotle, andSteckerl7 argues that his influence has been over-emphasised.Whether or not a direct transmission of

Aristotelian ideas can be identified, there is nodoubt of the background importance of hisphilosophy. His argument that even terrestrialobjects were worthy of the attention of a philo-sopher was new to Greek science and a turningpoint within it.8 His programme of analysing thewhole natural world encouraged his successor,Theophrastus, to work on plants, at the descrip-tion of which Aristotle had stopped short, and ledhis student, Clearchus of Soloi, to write a book on

the anatomy of the human bones and muscles,another gap in Aristotle's world picture. Clearchuswas contemporary with the first of the Alexandrian

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anatomists and may have had access to humanmaterial in Alexandria. Certainly Aristotle's em-phasis on the primacy of the senses in scientificinvestigation must have greatly encouraged thedevelopment of human dissection in Alexandria.9(We have already seen how Aristotle's philosophydisplaced some old taboos.) Although Aristotle'smethod was in a sense empirical (as opposed to thedeductive system of Plato) it was part of a highlyrational attempt to understand the natural world.It was rational in seeking to establish the causesof things, and this was an attitude that appealedto the medical Rationalists of Alexandria.

This rationality of Aristotle and others lay inuncovering the plan according to which nature hadbeen put together. Perhaps the first appearance ofthis rationality appears in the 'Hippocratic' workon the heart, Peri Kardies, the author of whichadmires the skill with which the valves of theheart have been constructed. Peri Kardies, betterknown under the Latin title De Corde, is prob-ably closer in time to Aristotle than to Hippo-crates. By then it was accepted that the reasonemployed by Nature the craftsman, or thedemiurge who, Plato said, created the body, wasof the same kind as human reason, and mancould, therefore, in principle understand how thenatural world had been constructed. Anatomy nowhad a new function: it was no longer simply amedical affair, useful in understanding the struc-ture and function of the body, and useful insurgery; it was also a philosophical exercise inlearning how the body was constructed.

The results of human dissection

The major achievement of the Alexandrians was

the discovery of the central nervous system. Ofcourse, as we have already seen, the brain, spinalcord, and nerves had already been recognisedanatomically, and some dim awareness of themotor and sensory powers of the brain had beenthe cause of a traditional placing of at least part ofthe soul in the head. But the Alexandrians knewthat the central nervous system was the organisingcentre of the body; they distinguished betweenmotor and sensory nerves, and we have every

reason to suppose that they experimentallysectioned nerves in the living animal-perhaps,indeed, in living man-and learned from theresult the function and distribution of the nerves.

Discoveries thus made by experimental vivisectionwere taught by the descriptive anatomy of dis-section.10 The central nervous system is not ofdirect consequence to the history of the thorax,except that Galen, in developing the work of the

Alexandrians, made some dramatic and sophisti-cated experiments sectioning the nerves controllingthe motions of respiration. Its importance for ourstory is that the discovery of the central nervoussystem at once threw into relief the traditionaldebate about the most important organ of thebody, the seat of the soul.

Quite apart from the other influences of theAristotelian philosophy that we have discussed, amajor influence was that Aristotle had said thatthe heart was the seat of the soul, the organisingcentre of the body. This position became increas-ingly difficult to maintain as the function of thebrain and nerves became clear. The nerves werenow distinguished from the other fibres of thebody; neura became a technical term meaning'nerves' only, and it was recognised that onlyneura had a sensitive and motor function. Yet sogreat was the influence of Aristotle that someanatomists attempted to reconcile Aristotle's non-nervous neura of the heart (regarded as theorganising centre) with the new anatomy andphysiology. Thus Praxagoras, who is placed bysome historians in a direct line of influence fromAristotle and Diocles to the great Alexandrians,Herophilus and Erasistratus, claimed that thearteries, arising from the heart, finally became toonarrow to admit blood and became nerves. Sincethe work of Herophilus and Erasistratus exists onlyin fragments" we cannot tell what directed theirattention to the central nervous system; mostprobably it was a continuation of older Greekwork on the sense organs, and particularly theeye. Some of this interest may have been originallygenerated as a result of philosophical problems ofperception. We have already noticed a previouslyunrecognised addition to our knowledge of Hero-philus's work on the optic nerves in a passage ofChalcidius,'2 and it is possible that he generalised,from this and other examples of the connectionsbetween the sense organs and the brain, to thestatement that the brain is responsible for allsense and motion.Whatever led Herophilus to these ideas, it was

clear enough that the heart, the traditional seat ofthe soul and the prime organ of the thorax, couldnever be the same again. Two other discoveriesjust before or at the beginning of the Alexandrianperiod further transformed ideas about it and thepattern of blood flow through it. The first of thesewas the final clarification of the distinction ofarteries and veins. While earlier authors had someglimmering of this, a striking difference betweenthe two sets of structures was made by Praxagoras,who derived the veins from the liver (which wasto be Galen's view) and said that the arteries

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contain only air or spirit, pneuma, an opinionshared by Erasistratus and bitterly attacked byGalen. So the difference between arteries and veinswas functional as well as structural. Herophilussaid that the arterial coat was six times as thickas the venous, and Erasistratus, quite free fromany ambiguities between nerves and other fibres,and between arteries and veins, elevated all threetypes of structure to the status of three greatsystems serving the body. The veins suppliednourishment, the arteries vital spirit, and thenerves psychic spirit; each organ of the body wasactually composed of the fine terminations of thethree kinds of ramifying vessel.The second important discovery of the pre- or

early Alexandrian period was that of the cardiacvalves. We have seen that these structures aredescribed in the work De Corde, which was onceattributed to Hippocrates but which was probablywritten after Aristotle, perhaps in Alexandria. Itis in the nature of a valve that it allows only aunidirectional flow, and this at once imposedsevere constraints upon traditional ideas of themovement of blood and spirit through the heart.We have seen that these ideas are somewhat vaguein the existing texts, but what is clear enough tous is that some symmetrical relationship was heldto exist between the lungs and the heart, bloodand air being exchanged on both sides. But thevalves changed all this, those of the vena cava onthe right 'entering', and those of the aorta on theleft 'leaving' the heart.These two discoveries involved the demise of

the traditional fundamental pair of blood vessels,anciently assumed to run throughout the length ofthe body with symmetrically disposed branches tothe organs. The discovery that one of these vesselswas a vein (the vena cava) and one an artery (theaorta), the assumed origin of the vena cava fromthe liver and the aorta from the heart, and theassumed function of the arteries of carrying spirit,not blood, all combined to destroy the ancientsymmetry. The picture was particularly compli-cated in relation to the vessels between the heartand the lungs: the vessel that arose from the hearton the same side as the veins looked like an arterybut was considered to have a venous function aspart of the great nutritive system of liver-venacava-heart-veins. Correspondingly, the vesselbetween the heart and the lungs on the left, thearterial side of the heart, looked like a vein buthad to have the arterial function of the great res-piratory system of lungs-left ventricle-aorta-arteries. The names 'venous artery' and 'arterialvein' thus came into existence during the Alex-andrian period.

Herophilus and Erasistratus

Against this background we may briefly inspectwhat was said about the thorax and its origins bythe two greatest of the Alexandrian anatomistsand physiologists, Herophilus and Erasistratus. Itwas Herophilus who said that arteries had tunicssix times as thick as those of veins, and he wasconsequently struck by the arterial nature of the'vein' connecting the heart and lungs and by thevenous nature of the 'artery'. Rufus of Ephesus"3tells us that Herophilus introduced the terms'arterial vein' and 'venous artery' for these vessels,although Galen appears to claim them as his owninvention.14 Galen also tells us that Herophiluswrote 'carelessly' about the cardiac valves, perhapsin retaining some confusion about the 'nervous'fibres associated with the valves; we may assume atleast that Herophilus recognised the functionaldirection of the valves. Thus in the fourfoldmotion of the lungs (which uniquely in the bodyhad a motion independent of the nerves) air wasdrawn into the lungs, from the lungs to the heartand body and back again, and lastly the air wasexpelled. We can probably assume that the incom-ing air passed down the venous artery (pulmonaryvein) into the heart, and thence via the aorta tothe body. In the aorta and arteries this air orspirit was mixed with blood. Given the functionaldirection of the valves, however, we cannot seehow this air was returned to the heart and lungs;nor can we be certain that Herophilus allowed thecardiac valves to work 'properly' (in our sense).The force of traditional ideas was such that manyschemes of physiology contain allowances for themdespite the discovery of the valves. Thus theHippocratic De Corde allows that the valves onthe right side of the heart do not close completely,allowing in some air from the lungs and maintain-ing an ancient symmetry of blood and air ex-change. We shall see that Galen's ideas onbloodflow are an imperfect compromise of oldideas and new discoveries, allowing an imperfectclosure of the valves on the left of the heart.Lastly, it is possible that we see the persistence ofold ideas in Herophilus' notion that spirit entersthe body not only by way of the heart but fromall over the body.'5 This may be the idea, derivedfrom Empedocles, that tubes, or arteries, haveopen mouths at the surface of the body, drawingin air. Herophilus' study of the pulse in diagnosisand physiological theory seem to have been aninnovation. While the arteries originated in theheart, Herophilus said he could not be certain ofthe source of the veins.1'

In contrast, Herophilus' younger contemporary,

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Erasistratus, located the origin of both veins andarteries in the heart. He was in early life associ-ated with the Aristotelians, and it may be that thisanatomical suggestion is a sign of peripateticinfluence.17 On the other hand, he believed in aform of the atomic theory, involving a belief in adiscontinuous vacuum and a rejection of 'hidden'causes. Nothing could be further from theteleologically organised plenum that wasAristotle's world picture.

Erasistratus seems to have been more interestedin the working of the body than was Herophilus,whose achievements were primarily anatomical.The fragments of Erasistratus' work on thephysiology of the thoracic organs are fortunatelymore detailed than those of Herophilus. Consistentwith his anti-Aristotelian atomism was his beliefthat the body did not have (as Aristotle had said)an innate pneuma, but that the ambient air (spiritor pneuma) was drawn in during inspiration, pass-ing through the 'primary arteries'8 (the bronchi)to the lungs, from the lungs to the heart, andfrom the heart to the body. This is an elabora-tion of the traditional idea of the arteries, heart,and lungs as a single respiratory system. So takenby this idea was Erasistratus that he insisted thatthe arteries contain only pneuma, unmixed withblood.'9 Once in the body, the pneuma underwenttwo stages of concoction: in the left ventricle itbecame vital spirit, which filled the arteries, andin the arteries of the brain it became psychicspirit.20 Erasistratus simplified Herophilus' notionof the heart, regarding it as a two-chambered, notfour-chambered organ, the auricles being merelyear-shaped expansions of the vessels at their con-nection with the ventricles. This made it easier toregard one half of the heart as respiratory and theother as sanguineous or nutritive. As there wasno communication between the two systems in theheart or lungs, Erasistratus was in difficulty inexplaining how blood emerges from a woundedartery, which he considered contained pneumaonly. He thought that the finest branches of theveins, too narrow to admit blood, anastomosedwith the finest branches of the arteries, and thatin certain circumstances, as when the arteries weredamaged and their pneuma escaped, blood wasdrawn into the arteries from the veins by theresulting vacuum.

Galen21

Some time between the period of Herophilus andErasistratus and that of Galen, who was bornabout 129 AD in Pergamon, the practice of humandissection as a teaching and research practice

came to an end in Alexandria. The Ptolemy whoreigned in the second half of the second centuryBC expelled the physicians, which perhapshastened the decline of teaching. Rufus ofEphesus, writing at the time of Trajan, describedhuman dissection in retrospective terms and hadto be content to dissect the pig, although recognis-ing that apes were more similar to man.22 Al-though some Hellenistic texts23 mentionApollonius of Memphis as a dissector and a fol-lower of Herophilus (he antedated Galen) it isclear that by Galen's time the practice had ceased.Galen himself was in the midst of a revival ofGreek studies that began in the Roman worldsome 50 years before he was born.24 The partplayed by Galen in searching out the spirit ofGreek medicine was his reverence for Hippocrates,his anxious pursuit of the pupils of the famousGreek teachers of anatomy, and, it must be ad-mitted, his pretence that he, like Alexandrians,had had the opportunity to dissect the humanbody. The only human material routinely avail-able to him was the skeleton while he was learninganatomy in Alexandria.25 Otherwise, it was onlyfortuitous circumstances that furnished him withglimpses of the anatomy of the soft parts, andindeed much of his insistence on a systematictraining in comparative anatomy and the dis-section of animals reflected the need to preparethe mind for occasional access to human material.Galen's literary energy was immense, and a verylarge proportion of his numerous medical workssurvive; his influence was enormous and for amillenium and a half he was revered as 'ThePrince of Physicians'.26

Galen's system of physiology, as far as it relatesto the thorax, was an amalgam of his own dis-coveries and reasoning, and older ideas. A partialsurvival of the old idea of the body's fundamentalpair of vessels is Galen's description of the venacava as a long, continuous vessel arising from theliver. The liver was the starting point of the body'sphysiological processes, the site of the 'firstcoction' where the incoming food, transformedto chyle by the action of the stomach and in-testines and carried to the liver by the portalvein, was turned into blood, the food of the body.The body as a whole absorbed this blood slowlyby the process of assimilation, literally 'makingsimilar'-a process in which the Aristotelianqualities of the blood were so disposed to convertpart of the blood into body substance. In otherwords, each tissue or 'similar' (homogeneous) parttook from the blood what was necessary to it,leaving a small proportion of residue to be elimin-ated. The entire venous system was thus like a

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Fig. 1 Galenic cardiac physiology. For explanationsee text.

tree,27 the trunk of which was the vena cava,

planted in the liver and supplying blood to all partsof the body with a slow, unidirectional motionthrough its branches.

It follows that for Galen, unlike other writerssuch as Aristotle, the vena cava did not arise inthe heart, nor indeed did the heart interrupt thestructure of the vessels as it passed along the body,for the vena cava merely inosculated with theheart by means of the intermediary right auricle,which Galen regarded simply as a sinewy ex-

tension of the vein. This arrangement is shown inthe diagram (Fig. 1), where AB indicates the venacava arising from the liver and passing the heart,and C its opening into the right auricle. Theentire venous 'tree' is in line with the older ideaof the 'nutritive' (venous) system as opposed to the'respiratory' (arterial), and it enabled Galen toagree with the old-fashioned accounts (which we

met in the first article) of the vascular systemgiven by his idols, Hippocrates and Plato,28 and toreject those who opposed them, includingErasistratus. In fact Galen's rejection of Erasis-tratus' claim that the veins, like the arteries, arisein the liver is illuminating. Erasistratus was awareof the valves that point into the heart at the attach-ment of the vena cava (we are now considering,with Galen, the heart as a two-chambered organ)and which govern the flow of fluids into the heart.How, asks Galen, can the heart be the centre ofthe venous system when the blood, the raison

d'etre of the veins, is produced elsewhere (theliver) and carried to the heart? Galen concludedthat the vein-producing and vein-controllingfaculty must be located in the same place as thefaculty that produced and controlled the blood,for otherwise nature would be multiplyingprinciples in vain, which she never does. There islittle morphological difference between a cardiacorigin and hepatic insertion of the vena cava onthe one hand and a hepatic origin and cardiacinsertion on the other. Galen had in mind ateleological and functional conception of a liver-vena cava-heart-veins body arrangement, thevenous 'tree' that has to be translated into ana-tomical terms. This is one example of a verygeneral occurrence in the history of anatomy,namely, that physiological considerations veryoften determine anatomical statements.

Finally, there are technical reasons why Galenmay have been encouraged to think of the venacava as a long, continuous vessel. Unable to dissectthe adult human body, he was obliged to useapes. It is possible that he and Aristotle were ableto dissect human fetal material; in some apes andin the young human, the vena cava does have theappearances of a straight, continuous vessel pass-ing through the right auricle and thus inosculatingwith the heart (Fig. 2).Thus the body was supplied with blood through

the branches of the vena cava arising above thepoint B in Fig. 1. As observed, each tissue took itsproper aliment, the substance that resembledit most, from the blood. Yet there was one ex.-ception. The lung (always spoken of by Galen asa single bilobed organ),29 constructed of ex-ceptionally light and frothy substance, needed verylight and subtle blood for its nourishment.Ordinary venous blood direct from the liver wastoo heavy and coarse, and Galen therefore sup-posed that that part of it destined for the lungunderwent a second concoction in the rightventricle of the heart, which it entered throughthe opening of the vena cava into the right auricle(C in Fig. 1) and through the valve (D in Fig. 1).This preparation of the blood was the sole func-tion of the right ventricle, executed by a specific'faculty', a name Galen always used for a uniquebiological process about which little else wasknown. After concoction this blood was suppliedto the lung through the vessel E (the pulmonaryartery), which Galen considered to be a veinbecause it carried venous blood. He adopted theAlexandrian name of 'arterial vein' because hesaw clearly that the vessel had an arterialstructure. Galen did not accept that this structurewas anomalous but argued that the thicker coat

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of the vessel was designed to prevent the escapeof the highly rarefied venous blood it contained.

In this way the whole of the body was suppliedwith nourishment. Yet, in accordance with tra-ditional ideas, Galen held that the body must notonly be nourished but vivified; it had to begoverned not only by a nutritive faculty but by a

vital faculty, giving heat, life, and motion. Wemay recall the fundamental physiological observa-tions of the living and dead body with which thisseries of articles began, and also the resultanthierarchy of living principles or souls formulatedby Plato and Aristotle. The soul responsible forvivication, heat, and motion was the inspired air-or spirit-soul, and the system serving it in the bodywas the respiratory, that is, the arterial. We havedescribed above how by Galen's predecessors thelungs, heart, and arteries were considered as a

single system. Galen accepted this, but with a veryimportant reservation: he insisted that the arteriescontain blood, not simply spirit. He wrote a book30to refute Erasistratus on this point and, using the

Alexandrian vivisectional experimental method,he proved his case by experiment.31 He ligated anexposed artery of an animal in two places to pre-vent subsequent inflow of blood from the veins(the Erasis4rateans' argument) and opened theartery between the ligatures. Of course he foundit was full of blood and, like others, he was able todistinguish between arterial and venous blood bytheir colour and the force with which they flowedin their vessels.But now Galen faced a difficulty. The two

vascular systems were quite separate, one in thetraditional fashion concerned with nutrition, theother with the respiration of spirit. He insistedthat the arteries contained blood but denied trans-ference of blood from the finest branches of theveins to those of the arteries in the way describedby Erasistratus. There was for Galen no way ofgetting blood from its only place of production,the liver, into the arteries. With hindsight we cansee it would have been a simple matter for him tohave supposed that blood crossed the lungs andentered the heart through the pulmonary vein (thevenous artery), but Galen had already denied avenous-arterial exchange of blood. To havepostulated one in the lung would have been, inaddition, to obscure the traditional distinction be-tween the nutritive and respiratory systems. More-over, the lung, like all parts, absorbed the bloodcoming to it as nourishment.

In order to avoid this difficulty, Galen postulatedthe existence of pores in the interventricularseptum of the heart, so that a small quantity ofvenous blood could be transferred to the leftventricle at F (Fig. 1). The left ventricle was atraditional seat of the soul in Greek medicalthought (compare De Corde), and Galen placedit in a Vital Faculty, which transformed theincoming venous blood into arterial blood, whichcarried vital heat to the body through the arteries.The concoction that produced arterial blood neces-sarily required the inspired air or spirit of theoutside world. Galen urged that air in substance,or its insubstantial quality, passed down thevenous artery G (Fig. 1) from the lung to the leftventricle. This life-bearing pneuma was trans-formed in the left ventricle into vital spirit,pneuma zotikon, the essential component ofarterial blood. As in all concoctions, there wereresidues to be expelled, here 'fuliginous vapours',whose only route of escape was back up thearterial vein to the lung and outside world.

THE CARDIAC VALVESYet, in solving his first difficulty in this way, Galenfound himself with another. Spirit passed down

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through the venous artery into the left ventriclethrough the valve now called the mitral. It wasclear to Galen and others that the functionaldirection of this valve is into the heart, but Galenwas obliged to allow passage in the reverse direc-tion to the escaping fuliginous vapours. Moreover,the lung, like all parts of the body, needed arterialblood, which could reach it only from the leftventricle in opposition to the natural direction ofthe mitral valve.So Galen had three different substances, arterial

blood, spirit, and sooty wastes, moving in two dif-ferent directions across a valve whose naturaldirection favoured the motion of only one of themand that the least substantial-spirit, or thequality of air. This part of Galen's cardiacphysiology drew the criticisms of later naturalphilosophers and of historians. It caused Harveyto cry 'Good God! How do the mitral valveshinder the return of air, and not of blood?'32 ButHarvey knew what a valve was, and those his-torians who discuss the 'incompetence' of Galen'smitral valve are anachronistically reading back toGalen their own knowledge of valves. Galen'sattention was given to the flaps of the valve, themembranarum epiphyses, not to the whole struc-ture of the valve. Every concoction, every ex-change of materials in the body was accompaniedby some residue, some superfluity, for no substanceultimately derived from food was entirely pure. Itfollowed that all the cardiac valves allowed somereverse flow, and the mitral valve was particularlynotable in this respect because of the intensity ofthe concoction in the left ventricle, and becauseit had fewer flaps. Galen had no other notion ofwhat a valve was other than the flaps of the heart.Although he repeatedly used the analogy of abellows to explain how the heart attracted sub-stances, they were valveless bellows. Not untilshortly before the time of Harvey were valves wellenough known in other fields (in water mills, inpumps of fire engines, in the veins, and in theintestine) for the 'incompetence' of Galen's mitralvalve to stand out in contrast. Although Galenseems only to have described the mechanicalreasons for the opening and closing of the flaps,the forces impelling the passage of the fluidsthrough the valve were the purely biologicalFaculties of attraction, retention, and expulsion,which were entirely non-mechanical. So Galen'swhole idea of the movement of the fluids of theheart was sui generis, and he does not deserve allthe censure normally levelled at him.

THE SPIRITUAL AND ANIMAL ORGANS

The left side of the heart, the pulmonary vein, and

the arteries were for Galen the organs of respira-tion, drawing in air or external pneuma and con-verting it to bodily pneuma. It followed that thepulmonary vein was regarded as an artery, andbecause it looked like a vein Galen adopted thename 'arterial vein' for it. Its thin tunic, heargued, allowed it to be compressed during thecontraction of the thorax in breathing to aid thepassage of its fluids. The normal function ofthe thick arterial coat was to transmit the activewave of dilatation, which Galen considered thepulse to be; the venous artery, moved by thethorax, had no need of a pulse or thick tunic.

It is puzzling for the modern reader to under-stand why Galen did not simply allow that bloodcrossed the lungs, which would at once haveremoved the difficulties of the perforate septum,the anomalous structure of the pulmonary vessels,and the 'incompetent' mitral valve. This step wasindeed the first advance from Galenic physiology,made apparently independently three times beforeHarvey, but to ask why Galen did not take it is, aswe have seen, to ignore the power of the tra-ditional ideas which Galen took to be established.The problems which we see in his account werefor him successful modifications imposed uponthe old ideas by the recent discoveries of theAlexandrians.

In contrast, Galen's ideas on the nervous systemdid not depend on ancient ideas and on the reveredHippocrates but on comparatively recent work,much of it based on vivisectional experiments,both his own and of the Alexandrians. There wasno weight of traditional ideas to labour under, andGalen's description of the nervous system is muchmore successful in modem eyes than that of theheart. Broadly, the central nervous system forGalen represented the highest category of thetraditional hierarchy of three-fold governance ofthe body, the lower two being the nutritive andthe vital. Just as the vital faculty of the heart andarteries depended on the nutritive, so the vitalfaculty provided material for the rational or'animal' faculty of the brain. Arterial bloodascended through the carotid arteries to the retemirabile, a network of arteries which Galen (andthe Alexandrians) found above the basilar bone ofthe skull of certain domestic animals and attri-buted to man. In it the finely divided blood, saidGalen, underwent a second concoction, with airdrawn in through the nose into the ventricles ofthe brain, to produce the second of the bodilyspirits, pneuma psychikon, 'animal spirits'. Thispneuma filled the substance of the brain andnerves and served the power of motion and thefaculties of sense.

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By dissection and vivisection Galen traced outthe course of the cranial and spinal nerves andshowed how such anatomical knowledge wasessential to medicine. It is his most powerfulattack against the Empirics, and he illustrates itby the story of a patient who had fallen from hisconveyance on his way to Rome, damaged thevertebrae at the bottom of the neck, and lost theuse of three fingers. The Empirics applied theirremedies where they found the symptoms. Galentreated the origin of the nerves at the vertebraeand effected a cure.33Another story of which Galen was proud was

that of his discovery of the recurrent laryngealnerve. He tells34 how a surgeon, excising a tumourof the neck, cut too deep and rendered his patienthalf dumb. Galen was able to reproduce the situ-ation experimentally in a living pig; by tracing thenerve from the brain to the vocal organs heshowed its structure as an organ of voluntarymotion. By gently squeezing the nerve of bothsides he was able to deny the pig its voice for aslong as he pleased and then to permit its re-covery.33 His experimental analysis extended tothe control of all thoracic motions.36

In tracing out the anatomy of the recurrentlaryngeal nerve Galen was struck by the fact thatit descends to the thorax and folds itself overthoracic structures before ascending again to itsproper destination in the throat. His answer lay inthe general principle that nerves, being soft, werealways inserted into the end of the muscle thatmoved the least, in order to avoid damage. Thetop of the vocal organs was more mobile than thebottom, and therefore the nerve had to approachit from below: but a long, soft nerve needed sup-port over such a distance, and so nature contrivedto support it upon the thoracic arteries.Another experiment of Galen's,37 as striking as

that upon the recurrent laryngeal nerve, was theserial section of the spinal cord. Again, the livingpig was secured to the operating table and a scalpelwas inserted between each vertebra to destroy thelongitudinal fibres. Beginning above the sacrum,38Galen repeated the operation at every vertebraand carefully noted the results. In this way he wasable to establish the distribution of the spinalnerves and the manner of their control of theorgans: he noted the progressive loss of senseand motion as the operation proceeded throughthe thoracic vertebrae into the cervical. In parti-cular, he was able to isolate the nervous controlof the various parts of the motions of respiration,making the animal breathe by the intercostalmuscles or by the diaphragm alone. In these ex-periments Galen exhibited an acute power of

observation, and his anatomy is very detailed.40He described the rami communicantes41 of thespinal nerves but did not discover the sensory andmotor function of the different nerve roots.

In exploring the nerves of the thorax Galen paidsome attention to the vagus and sympathetictrunk on their journey through the chest. He didnot always clearly distinguish between the twonerves, sometimes describing the sympathetictrunk as a branch of the vagus, and always re-garding it as a cranial nerve, the sixth of the sevenpairs of the enumeration he adopted from histeacher, Marinos. The earliest account he gives ofthe sixth pair is in On the Use of the Parts,42 andit is largely based on the nervous anatomy ofdomestic animals. Consequently, the nerve hedescribes is single down to the level of the top ofthe thorax, the common vagosympathetic trunk ofa number of animals including the horse and thepig.43 The purpose of both nerves for Galen wasto give sense and motion to the abdominal visceraand lower organs; accordingly, both nerves wereobliged to travel over a great distance from thehead, and as it was in the nature of a nerve to besoft, they had to be defended against damage ontheir journey. This is the reason, says Galen, forthe plexuses and ganglia he saw on the sixthpair, and for the attachment of the sympathetictrunk to the spinal nerves 'at the roots of theribs'.

This account of the thoracic parts of the sixthpair shaped all anatomical opinion until, and after,Vesalius. It became formalised in the statementthat the sixth pair descended to the level of thefirst rib and divided into three branches, the'stomachic' (the vagus), the recurrent laryngeal,and the 'costal' (the sympathetic). The costal wasso called because of its connection with the spinalnerves to the muscles of the ribs, the intercostals.Later the 'costal' for the same reason becameknown as the 'intercostal', which led in the 17thcentury to confusion with the true intercostalnerves that Galen had described adequately. Inthe later On Anatomical Procedures, based onapes, Galen has a much clearer picture of theseparate paths of the vagus and sympathetic trunkin the neck and thorax.We have already seen that Galen rejected the

Aristotelian notion that the nerves arise from theheart. We have also seen that this may in fact bebased upon something of a misunderstanding ofAristotle's position, because, in his day, nerveswere not clearly distinguished from other fibres.At all events, Galen's rejection of Aristotle is partof his wider attack on the whole notion of cardio-centricity. He devoted a large part of his book on

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the opinions of Plato and Hippocrates44 to estab-lishing the supremacy of the brain as a centre ofcontrol. So strong was the idea of nervous controlof muscular motion, and so anxious was Galento deny nerves to the heart, whether leaving it or

entering it, that he denied that the heart was a

muscle. Instead, he claimed that it moved bymeans of its own unique, innate Pulsific Faculty,and its motion was natural,45 arising from the'nature' of the heart, not animal, the motion ofthe skeletal muscles arising from the will (animain the Latin tradition). This action of the heartwas one of expansion, not contraction, so thatthe active phase of the heart's motion was diastole.This was the basis of his analogy with the black-smith's bellows, which were forcibly opened by thesmith, drawing in air through the same nozzlethat also fed the fire: the heart drew in bloodfrom the vena cava, and, in turn, the activelyexpanding arteries drew in the heart's blood witha pulse derived from the heart.Here Galen is consciously disagreeing with

Alexandrian opinion, which said that the heartcontracted forcibly, expressing its contents. Hisopinion, however, agrees with a wider notion inhis physiology: voluntary muscles were composedonly of longitudinal fibres which contracted,physically pulling the tendons and bones, but thehollow organs of the viscera46 were composed ofthree different kinds of fibre, and acted by means

of Faculties that were essentially biological andnot physical. Each hollow organ was composed oflongitudinal or straight fibres, which exercised a

Faculty of Attraction, circular fibres that enabledthe Expulsive Faculty to eject the contents of theorgan, and transverse fibres, which, in a con-

certed action with the other two, retained thecontents.47 All these Faculties were subdivisions ofthe Nutritive Faculty and were primarily exercisedin the organs of nutrition. Although the left ven-

tricle was the home of the Vital Faculty, theright ventricle was part of the venous-nutritivesystem that Galen said was controlled by 'nature'.In saying that the motions of the whole heart andthe arteries were 'natural', Galen meant that theywere not controlled by the rational soul or thepsychic faculty in the brain.

THE MOVEMENT OF THE HEART AND BLOOD IN THE

FETUS

Apart from the invisibly small septal pores,Galen's description of the anatomy of the heartwas precise. He knew of the fetal ductus arteriosusand the foramen ovale and their fate at birth. Yethis physiology of fetal bloodflow contained three

major errors in modern terms: he imagined that

in both adult and fetus blood crossed the septum,that the flow of blood in the veins in the bulk ofthe body was centrifugal, not to the heart as inmodern physiology, and, lastly, in specific refer-ence to the fetal condition he imagined thatmaternal blood crossed the placenta to supply thefetus. Even with these constraints upon his inter-pretation of fetal bloodflow Galen was able toexplain in a way that fitted admirably with the restof his physiology how the particular needs of thefetus were met. The fetal lungs, deprived of air,had no need of an adult vascular supply, and theneeds of the fetus were met by the maternal blood,which completely bypassed the fetal heart, theaction of which was unnecessary in the fetus; itbeat but did not expel its contents.48Galen assumed that the maternal venous blood,

flowing through the umbilical vein, reached thefetal liver and rose to the heart through the venacava in the usual way, from A to C (Fig. 1). Sincethe fetal lung was not functioning, Galen supposedthat there was no need for the right ventricle toexercise its only function, that of preparing bloodfor the lung. The blood consequently did not enterthe right ventricle but passed directly through theopen foramen ovale (obscured in the diagramby the vessels) into the left auricle (H, Fig. 1).Since Galen considered the auricles simply assinewy expansions of the vessels, the blood in thistransit did not pass through the heart at all butascended from the left auricle to the lung throughthe venous artery (G, Fig. 1), which not onlylooked like a vein but also in the fetus carriedvenous blood.Maternal arterial blood, said Galen, entered

along the route umbilical arteries-fetal iliacarteries-fetal aorta. From the aorta it was dis-tributed to the body in the usual way, with theexception of the lungs. Since the fetus was notrespiring, the whole lungs-heart-arteries respira-tory system of the adult was fed from the lowerend, and its functional direction was reversed inthe fetus. Approaching the heart in the aorta at I(Fig. 1), the maternal arterial blood had no needto enter the fetal heart, the left ventricle of whichwas not engaged in the production of spirit; in anycase, the aortic valve would have prevented anysubstantial flow. The only route for the arterialblood to reach and vivify the lung was across thefetal ductus arteriosus J (Fig. 1) and up thearterial vein, which consequently in the fetus notonly looked like an artery but carried arterialblood. At birth the umbilical bloodflow stopped,air entered the lungs, and the foramen ovale andductus arteriosus closed, producing a normal adultGalenic bloodflow.

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Conclusion

Galen stands at a critical point in the developmentof biological science. The Hellenistic descriptiveand experimental tradition was summarised andextended in his description of the morphology ofthe heart and the changes occurring in the fetalheart at birth, which is accurate enough bymodem standards. In that sense Galen's accountof the heart looks forward to modern medicine.Yet his account of the functioning of the adultheart was almost entirely dominated by traditionalideas, which we met in the first article. In otherwords, Galen did not derive his ideas on functionfrom those on structure; nor, indeed, did anyother student of the human body, for almost in-variably in the history of anatomy and physiologyideas of function have either instituted or modifiedanatomical statements. For example, Galen's des-cription of the interventricular cardiac pores ismatched by his account of an anatomical pathwayfrom the nose to the ventricles of the brain, andfor the escape, in the reverse direction, of thewaste products of the concoction of animalspirits.48

This problem of the relationship between formand function is highlighted by the case of theheart, which is a complex structure. Indeed, themammalian heart is unnecessarily complex andarose as a four-chambered organ with a 'figure ofeight' circulation from the evolutionary accidentthat the primordial lungs that were to take overthe function of the ancient gills, as the ancestorof air-breathing animals moved out of the sea,already had their own blood supply with a venousreturn direct to the heart. This venous blood cameto be arterial and in order to provide efficient cir-culation, should have gone directly to the body,but the existing pathways could not be abandoned,so the arterial blood returned to the heart to besent out again. The four chambers of the heartevolved as a complex device to handle the doublecirculation. Further, the major changes takingplace in the chambers, vessels, and fluids of thefetal heart at birth are an additional complexityalso due to the secondary development of air-breathing in evolutionary history. They are neces-sary to adapt the already secondarily complexfour-chambered heart from an aqueous to gaseousenvironment.

It would not be difficult for a hydraulic engineerto design a simpler and more efficient organ andcircuit of circulation, and it is no surprise thatGalen's interpretation of the structure does notmatch the modern account. His interpretation isa very good example, because of the complexity

of the appearances, of the nature of his biologicalinterpretation as a whole, which has two aspects.Firstly, Galen discussed scientific procedure inmaking discoveries but, secondly, he knew in ad-vance what kind of thing he was going to discover.He wrote a book on scientific method, of whichhe was proud and which has been lost. It seemsto have included discussion on inductive general-isations from sense experience, and also much ondeductive reasoning, on the pattern of geometry,from certain irrefutable axioms. Whenever weread about 'reason' in old texts on anatomy andphysiology we may be sure it is axiomatic, deduc-tive reasoning that is being referred to. Theseaxioms reveal a great deal about the world-pictureof the writer; they are the ideas from which hisreasoning started and were themselves incapableof proof or rejection by reason. Broadly, theyrepresent the kind of thing Galen knew he wasgoing to find by applying the Aristotelian inductiveor the Alexandrian experimental method. (Galen'sexperimental method was not entirely modern.Most often his experiments are designed to provesomeone else's ideas wrong; and so are part of ageneral mode of proof by rejecting all knownalternatives. He does not use experiment to 'seewhat happens' or to establish his own uncontestedideas. An excellent possibility here would havebeen to inject water into the right ventricle todemonstrate the pores of the septum.)

So, in purely medical topics, Galen uses theaxioms of contraries-cure-contraries as the basisof his therapeutics so that medicines with thefundamental Elemental Qualities of Wet and Coolare used in Hot and Dry diseases. In anatomy andphysiology, Galen's most fundamental conceptionwas that of Nature as the Creator of the body.Unlike the Christian conception of a Creator,Galen's Nature was not omnipotent and had cer-tain recognisable characteristics. She was boundby the Necessity, such as the nature of thematerials she was working with, she was enduedwith Reason, handling the Qualities of thematerials in the best possible way, and she wasbenificent, always bearing in mind the well-beingof the animal and of the species. Being reasonable,Nature never duplicated her efforts by doingsomething unnecessarily; 'Nature does nothing invain' is Galen's most frequent axiom in reasoningabout the structures of the body and their func-tion. That the structure of the heart was unneces-sarily complex, that some mindless process ofevolution had caught up the chambers and vesselsof the heart in an irreversible complexity, and thatany competent engineer could do better, would allhave been entirely foreign to Galen's way of

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thinking. His knowledge of comparative anatomytold him that animals without lungs did not havea right ventricle, but these animals were 'imper-fect', not merely simpler. His knowledge of animalform also told him that there was a single Mindthat fashioned all of nature. The guiding principlebehind all his anatomical and physiological re-search was to discover the rationality of Nature.

Lastly, let us see how this principle worked inGalen's analysis of the thoracic organs. He sawthe invisibly small septal pores of the heart with'the eye of reason', arguing that the clearly visiblebut apparently blind pits on the wall of the rightventricle must in fact perforate into the leftventricle, otherwise the pits would be withoutfunction; but Nature does nothing in vain, andnothing in the body is without function, thereforethe pits are perforate. We have already seen thatGalen's physiological theory dictated that nervesentered the least-moving end of a muscle, and sothe recurrent laryngeal nerve had to descend tothe thorax in order to approach the vocal organfrom below. But one of the Necessities thatNature labours under is that nerves are soft eventhough they have to be stretched over long dis-tances. As we have seen, such nerves as the vagusand sympathetic trunk were consequentlyequipped by Nature with ganglia and plexuses tohold them steady on their course, and the recur-rent laryngeal was supported by Nature with theaorta on one side and the subclavian artery on theother (as an inferior substitute).Galen is very clear about the roles of Reason

and Necessity in the creation of the pulmonaryvessels. Nature's reason in creating them was totake air to the heart and blood to the lung; theNecessity that constrained Nature was the ele-mentary qualities of Moist and Dry, which Natureblended together to produce a plastic, waxy sub-stance. Drying out some parts with the Hot andstiffening others with the Cold, Nature formedthe rudiments of the vessels.50 Galen identifiedfive causes in all such changes: the material, fromwhich the organ was made; the efficient, theimmediate instrument (Hot and Cold); the formal,or shape of the product; the final, the purpose ofthe whole operation; and, lastly, the Creator orNature herself. Within this philosophical frame-work Galen had to explain the anomalous struc-ture of the pulmonary vessels. We may refer tothe account of these vessels given above and recallthe Final Cause, Nature's reason, the exchangeof fluids; the Material Cause, the constituentmaterial, with the added Necessity of the veinbeing thick-walled to retain its fine blood and theartery being thin-walled to aid the expulsion of its

contents (part of the Formal Cause); and theEfficient Cause just mentioned, Hot and Cold.With this framework Galen must have been con-fident of his position, and he could afford to bescathing about those who attempted to explainthe anomaly of the pulmonary vessels withoutReason, Necessity, or Causality. Thus he attacks,with his customary gusto, the opinions ofAsclepiades, who had argued that the venousarteries of the lung (the pulmonary veins) werethin-walled because they worked twice as hard asother arteries in having a double motion-onefrom the lungs in respiration and another from thepulse. The arteries in the rest of the body, on theother hand, move moderately with their ownmotion and so grow well-nourished and strong.The veins in the body as a whole, continuedAsclepiades, do not move and so waste away likea lazy slave who takes no exercise, but the arterialvein of the lung does move, and so grows strong.5'However much Galen's teleology disagrees withmodern evolutionary theory, it is clear that he wasphilosophically much better equipped than someof his immediate predecessors.Another form of Necessity constraining Nature

in constructing the body in Galen's system was theopposing Qualities of neighbouring parts. Ingeneral, a hard part had to be separated from asoft, lest it damage it, by a third part of middlingsoftness. Thus the comparatively soft and verynoble heart would be damaged by its proximity tothe sternum, were it not for the pericardium, amembrane of intermediate nature.52 'Nobility' wasalso a concept that played a part in Nature'sreason, the more noble organs being those thathoused the higher faculties. They were con-sequently given precedence in position in thebody, organs of less importance being displacedfrom their own ideal position.

Notes and references

1 Edelstein, L. Ancient Medicine, Baltimore, 1967:The history of anatomy in antiquity, pp. 247-301.

2 Galen, Anatomical Procedures, trans. Singer (cf.note 3), p. 31.

3 Galen tells us about Diocles' text in On Anatomi-cal Procedures, trans. Singer, C., Oxford Univer-sity Press, 1956, p. 32. For the dating andfragments of Diocles, see Jaeger, W., Dlokles vonKarystos, Berlin, 1938, summarised in Etnglish inThe Philosophical Review, 49(1940), 393-414.

4 The evidence for human dissection by the Alex-andrians has been disputed. The most importantsource is Celsus-in the proemium to his DeMedicina (there is a translation by W. G. Spencerin the Loeb series, London, 1935)-who is probably

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the source of many later comments, such as thoseof Tertullian, who called Herophilus a butcherwho cut up 600 people (see the tenth chapter ofDe Anima in the edition of J. H. Waszink, Amster-dam, 1947, p. 13). Tertullian, however, alsodepends in part upon Soranus, an opponent of dis-section and a possible source of antivivisectionopinion. The whole question may have come intoprominence as a result of its adoption as a topicin rhetoric. Another classical source is Rufus ofEphesus (it has been translated by A. J. Brockin his Greek Medicine, London, 1929, p. 126), andthese accusations were repeated in the middleages by St. Augustine and Vindicianus. The pseudo-(and post-) Galenic Finitiones Medicae speaks ofthe Rationalists' advocacy of vivisection as if thedebate were still a live issue. Later medievalsources report that the Alexandrians vivisectedfirst the nervous system, then the organs of respira-tion, including the heart, and lastly the organsof nutrition. This reflects the philosophical prece-dence of the nobility of the different organs, andso may be a medieval rationalisation of the clas-sical practice, but it is also the most profitablesequence in which to employ vivisection, leavinguntil last those organs that continue to functionthe longest.

5 Strictly, the Empirical sect best known to laterhistorians and to Galen was founded by Philinusof Cos, a pupil of the major Alexandrian figureHerophilus, but empirical attitudes were certainlydebated during the period of the formation of theAlexandrian school, and the later empiricists con-sciously compared their ideas with the empiricalHippocratic writings.

6 See Jaegar, W., Aristotle. Fundamentals of theHistory of his Development, trans. by R. Robinson,Oxford, 1948.

7 Steckerl, F., The Fragments of Praxagoras of Cosand his School, Leiden, 1958, p. 34.

8 Lloyd, G. E. R., Aristotle: the Growth and Struc-ture of his Thought, Cambridge, 1968, p. 72.

9 On the use of the technique in teaching seeKudlien, F., 'Medical education in classical an-tiquity' in The History of Medical Education, ed.C. D. O'Malley, University of California Press,1970, p. 11.

10 See Matter, M. J., Histoire de l'Ecole d'Alex-andrie, Paris, 1844, vol. 3.

11 Largely reported by Galen. They have beencollected by Dobson, who sometimes includes toomuch of the Galenic context. Dobson, J., 'Hero-philus of Alexandria', Proc. Roy. Soc. Med., 18(1924-5), Sect. Hist. Med., 19-32; Dobson, J.,'Erasistratus', Proc. Roy. Soc. Med., 20(1927),Sect. Hist. Med., 21-28. See also Jones, W. H. S.The Medical Writings of Anonymus Londinens,Cambridge, 1947, which contains a useful discus-sion of the physiology of the two Alexandrians.

12 The passage from Chalcidius is given in Diels, H.,Die Fragmente der Vorsokratiker, Berlin, 1934,3 vols.; vol. 1, p. 212. In this passage Alcmaeon,

Callisthenes, and Herophilus are jointly creditedwith discovering a number of things about the eye,including the technique of excising it and thearrangement of the optic nerves. Comparison ofthis passage with that of earlier editions, such asthat of Meursius (I. Meursius, Chalcidii V.C.Timaeus de Platonis translatus, Leiden, 1617, p.230), suggests that it cannot be interpreted to showthat Alcmaeon removed the eye and dissected outthe optic nerves; it is very likely that Alcmaeon isincluded in this list of authors on the basis of hisexcision of the eye. Chalcidius' description of theoptic nerves agrees with what Galen tells us ofHerophilus' knowledge of them, and it is verylikely that Chalcidius' description of the opticnerves derives from Herophilus. Chalcidius' ac-count, however, includes a description of the junc-tion of the optic nerves, which thus appears to be adiscovery of Herophilus, not mentioned by Galenand previously unknown to historians. (On thepossibility of Alcmaeon performing dissections, seeLloyd, G. E. R., 'Alcmaeon and the early historyof dissection', Sudhoffs Archiv, 59(1975), 113-147;Galen's description of Herophilus' account of theoptic nerves is in the edition of the Opera Omniaedited by Kuhn, Leipzig, 22 vols; vol. 3, p. 812).

13 Rufus of Ephesus, De Apellationibus PartiumCorporis Humani, in Estienne, H., Medicae ArtisPrincipes, Paris, 1567, vol. 1.

14 Ed. Kuhn, vol. 5, p. 549.15 Galen, ed. Kuhn, vol. 4, p. 731.16 Galen, ed. Kuhn, vol. 8, p. 703.17 See Lonie, I., 'Erasistratus, the Erasistrateans, and

Aristotle', Bull. Hist. Med., 38(1964), 426-443.18 Galen, ed. Kuhn, vol. 4, p. 706.19 Wilson, L., 'Erasistratus, Galen and the pneuma',

Bull. Hist. Med., 33(1959), 293-314.20 Galen, ed. Kuhn, vol. 5, p. 185.21 Good general accounts of Galen and his work are

Temkin, O., Galenism, Cornell University Press,1974, and Sarton, G., Galen of Pergamon, KansasUniversity Press, 1954. More specialised accountsare those of Walsh, J., 'Galen's studies at theAlexandrian school', Ann. Med. Hist., 9(1927),132-143; Prendergast, J., 'The background ofGalen's life and activities, and its influence on hisachievements', Proc. Roy. Soc. Med., 23(1930),Sect. Hist. Med., 1131-1148; Harris, C., The Heartand Vascular System in Ancient Greek Medicine,Oxford, 1973; and Hall, A. R., 'Studies in thehistory of the cardiovascular system', Bull. Hist.Med., 34(1960), 391-413. A great deal of Galen'ssystem of medicine and physiology has beencovered in a number of volumes by R. Siegel, whois perhaps too inclined to interpret Galen inmodern terms. (Galen's System of Physiology andMedicine, Basel, 1968; Galen on Sense Perception,Basel, 1970).

22 Rufus of Ephesus, The Names of the Parts of theHuman Body, the relevant part of which is in-cluded in English translation in Brock, A. J., GreekMedicine, London, 1929, p. 126. See also Darem-

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berg's edition of 1879, p. 149, and the renaissanceLatin version of Estienne (cf. note 13).

28 The Introductio seu Medicus and the FinitionesMedicae: both are included, as pseudo-Galenic, inthe Venice, 1625 (Junta) edition of the OperaOmnia of Galen in the class 'Isogogici Libri'.

24 Prendergast, loc. cit.25 Galen, On Anatomical Procedures, trans. C.

Singer, p. 3.26 Galen's most important surviving anatomical and

physiological works are On Anatomical Procedures,the first half of which survives in Greek and hasbeen translated by Singer (cf. note 3) and thesecond half of which survived only in Arabic. Thisis available in English translation (by WV. H. L.Duckworth), Cambridge, 1962. On the Use of theParts of the Body is Galen's major philosophicaland functional interpretation of structure; see thetranslation by M. T. May, Galen on the Usefulnessof the Parts of the Body, Cornell University Press,1968 (2 vols). These two Galenic texts contain thefundamentals of his thoracic anatomy and physi-ology, including experimental vivisection of thenerves. The biological powers that are the funda-mental principles of his physiology are discussedby Galen in On the Natural Faculties, which hasbeen translated by A. J. Brock in the Loeb series,London, 1916. Galen's specialised tracts on aspectsof thoracic structure and on thoracic medicine are

not so well served by English translation. He wroteOn the Cause of Respiration (Kiuhn's ed., vol. 4,pp. 465-469), On Difficulty of Respiration in threebooks (Kuhn, vol. 7, pp. 753-960), and On theUse of Respiration (Kuhn, vol. 4, pp. 470-51 1).

27 Kuhn's ed., vol. 5, p. 525.

28 Galen discusses the opinions of Hippocrates andPlato in De Hippocratis et Platonis Dogmatibus:Kuhn's ed., vol. 5, pp. 520-538.

29 On many occasions Galen presents animal anatomyas if he were discussing the human body, and, inparticular, On the Use of the Parts is based on thestructure of the domestic animals (with somesimian material). The book is addressed equally tophilosophers and to physicians, and Galen is anxi-ous throughout to illustrate the virtues of Natureas a Creator: the human body in this connectionis obviously of much greater interest than that ofanimals, and Galen slides easily from one to theother without always keeping his readers fullyinformed. (In contrast, the strictly practical OnAnatomical Procedures is expressly about the dis-section of apes.) A good example of Galen mis-leading his audience is his discussion of the lung:'It is not my purpose to speak of the number oflobes in each of the other animals, nor is my dis-course ever concerned with the construction of anyother instrument in animals, except perhaps to pro-vide a point of departure for the explanation ofit in man. If death does not come to me soon, Ishall one day explain construction in animals too,dissecting them in detail, just as I am doing forman'. (May's translation of On the Use of Parts,

p. 285). However, Galen describes a non-respiratory'fifth lobe' of the lung, constructed solely, he says,to support the vena cava in its passage through thethorax; this lobe is a simian feature, illustrated inSinger's tran:lation of the first half of On Ana-tomical Procedures (p. 273).

30 An in arteriis sanguis contineatur.31 See Harris (note 21) p. 268.32 Harvey, W., The Circulation of the Blood, trans.

K. J. Franklin (Everyman ed.), London, 1963, p. 17.33 Kuhn's ed., vol. 8, p. 58.34 Kuhn's ed. vol. 8, p. 55; Duckworth's translation

of On Anatomical Procedures, p. 76. See alsoWalsh, J., 'Galen's discovery and promulgation ofthe function of the recurrent laryngeal nerve'.Ann. Med. Hist., 8(1926), 176-184.

35 Duckworth's translation of On A natomical Pro-cedures, pp. 91, 106, 107, 210.

36 Book 8, Chapter 5 of On Anatomical Procedures:Singer's translation, p. 211.

3 Galen seems to have enjoyed working before anaudience, and no doubt appreciated their reactionto the spectacle of a secured and noisy pig beingdramatically silenced. He tells us how he was askedto perform the experiment before a group: Kuhn'sed. vol. 14, p. 626.

38 Duckworth's translation of On Anatomical Pro-cedures, pp. 20-24.

39 Singer's translation of On Anatomical Procedures,pp. 211ff.

40 His material for these experiments was the Barbaryape, and until very recently it has been impossibleto confirm just how precise his anatomy was be-cause of the modern rarity of this species. TheAmerican anatomist, C. M. Goss, has recentlybeen able to follow Galen's dissections on an em-balmed specimen from their limited range on theRock of Gibraltar ('where they are held sacred bythe British' in Goss's words) and has been able toestablish the great accuracy of Galen's descriptions(private communication).

41 French, R. K., 'The origins of the sympatheticnervous system from Vesalius to Riolan', Med.Hist., 15(1971), 45-54.

42 May's translation, pp. 445ff.i3 Sisson, S., The Anatomy of the Domestic Animals,

Philadelphia, 1953, pp. 852, 878.44 De Hippocratis et Platonis Dogmatibus: Kuhn's

ed., vol. 5.45 De Motu Musculorum: Kuhn's ed., vol. 1, p. 1.46 Kuhn's ed., vol. 1, p. 602.47 May's translation of On the Use of Parts, p. 294.48 Harris, op. cit., pp. 290ff.49 May's translation of On the Use of the Parts, pp.

41, 425.50 Ibid., p. 311.51 Ibid., p. 309.52 Ibid., p. 320.

Requests for reprints to: R. K. French, PhD, Director,Wellcome Unit for the History of Medicine, Univer-sity of Cambridge, Free School Lane, Cambridge.

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