Marcello Malpighi and the discovery of the pulmonary ... · London inviting him to send manuscripts...

Marcello Malpighi and the discovery of the pulmonary capillaries and alveoli

John B. WestDepartment of Medicine, University of California San Diego, La Jolla, California

Submitted 17 January 2013; accepted in final form 25 January 2013

West JB. Marcello Malpighi and the discovery of the pulmonary capil-laries and alveoli. Am J Physiol Lung Cell Mol Physiol 304: L383–L390,2013. First published February 1, 2013; doi:10.1152/ajplung.00016.2013.—Marcello Malpighi (1628–1694) was an Italian scientist who madeoutstanding contributions in many areas, including the anatomicalbasis of respiration in amphibia, mammals, and insects and also in thevery different fields of embryology and botany. He was one of the firstbiologists to make use of the newly invented microscope and is bestknown as the discoverer of the pulmonary capillaries and alveoli.However, he also discovered the spiracles and tracheae that enablerespiration in insects. His studies of the embryology of the chickenwere far ahead of his time; he then turned to the anatomy of plants,where he made important contributions. Indeed, in some articlesMalpighi is referred to as the father of embryology and in otherpublications as one of the fathers of plant anatomy. His work on thelung was chiefly carried out on the frog; he referred to this animal asthe “microscope of nature” because it allowed him to see structuresthat were not visible in larger animals such as mammals. He alsoargued that nature undertakes its great works in larger animals after aseries of attempts in lower animals. For breadth of interest, innova-tion, and productivity, it is not easy to think of his equal in the fieldof life sciences.

comparative physiology; insect respiration; embryology; plant anat-omy; silkworm

MARCELLO MALPIGHI (1628–1694) was born in Crevalcore nearBologna into a family that was comfortably off (Fig. 1). Aninteresting tidbit about his date of birth is that this was the yearof publication of William Harvey’s De motu cordis describingthe circulation of the blood, and in a sense Malpighi completedHarvey’s missing link on the pulmonary circulation. Little isknown about Malpighi’s childhood and youth except that hewas schooled in “grammatical studies” in Bologna. Readerswho become interested in Malpighi are fortunate that Adel-mann (1) has written an exhaustive study in five large volumesthat is not only extremely detailed but also highly readable.

Malpighi studied philosophy for a few years but in 1653 heturned his attention to anatomy at the University of Bologna,and this was the beginning of an extraordinarily productivecareer in this science. In 1656 he was invited to be professor oftheoretical medicine at the University of Pisa, where he begana long friendship with Giovanni Borelli (1608–1679). Thisman was an eminent mathematician and naturalist who wasactive in the Accademia del Cimento, one of the earliestscientific societies, and Malpighi became a member. However,in 1659 Malpighi decided to return to Bologna. The subsequenttwo years were very productive, with extensive discoveriesabout the lung. He moved again in 1662 to a professorship inmedicine at the University of Messina, Sicily. In fact, through-out his life he spent periods at both Pisa and Messina, although

he always regarded Bologna as his home. His time in Messinaalso was very productive but he returned to Bologna in 1667.

In 1668 Malpighi received a letter from the Royal Society inLondon inviting him to send manuscripts to this prestigiousgroup. Malpighi was honored by the invitation; as a result hecarried out a classical study on the anatomy of the silkworm.He sent the manuscript to the Society, which published it in1669 (7) and made him a member. He maintained strong linkswith that group throughout the rest of his life, and most of hisanatomical studies were published by the Society. The admi-ration was mutual. On one occasion the secretary wrote to him“ . . . to no one . . . observing the structure of the human bodydoes Nature seem to have revealed her secrets as fully as to herbeloved Malpighi . . . So too our Royal Society embraces noone with greater affection” (1).

Unhappily, Malpighi was involved in a number of bitterdisputes about his scientific discoveries throughout his life, andhe also suffered from ill health in his later years. In 1684 therewas a disastrous fire at his home that destroyed many of hismanuscripts and much of his equipment. In 1691 he wasinvited to Rome by Pope Innocent XII to be the Pope’spersonal physician, which was a high honor. Malpighi died ofa stroke in 1694 and is buried in the church of Santi Gregorioe Siro in Bologna where there is a memorial.

Malpighi was an extraordinarily productive scientist. As weshall see he was the first person to describe the pulmonarycapillaries and the alveoli. In addition he was the first todescribe the anatomical basis of insect respiration as a result ofhis studies on the silkworm. Many of his discoveries initiallybore his name and some still do. He discovered the renalglomeruli (initially called the Malpighian corpuscles), the Mal-pighian corpuscles in the spleen, the Malpighi layer in the skin,and the Malpighian tubules in the excretory system of insects.He then went on to make extensive botanical studies and finallydid such extensive work on morphogenesis that he is some-times called the father of embryology on the basis of his workon the chicken embryo.

Discovery of the Pulmonary Capillaries

Much of Malpighi’s research was made possible by therecent invention of the compound microscope. Magnifyingspectacles using one lens go back a long way and were in usein the 13th century. However, the compound microscope, thatis one with both an objective and an eyepiece lens, appearedmuch later. Some authorities attribute the invention to Hansand Zacharias Janssen in the Netherlands at the end of the 16thcentury. Galileo developed a compound microscope in 1609,but it was some time before this was exploited for scientificresearch. Certainly Malpighi was one of the first to use acompound microscope in 1660, and Robert Hooke’s Mi-crographia published in 1665 contained beautiful illustrations.Nehemiah Grew (1641–1712) and Antonie van Leeuwenhoek(1632–1723) made further important advances. However,

Address for reprint requests and other correspondence: J. B. West, UCSDDept. of Medicine 0623A, 9500 Gilman Dr., La Jolla, CA 92093-0623 (e-mail:[email protected]).

Am J Physiol Lung Cell Mol Physiol 304: L383–L390, 2013.First published February 1, 2013; doi:10.1152/ajplung.00016.2013. Perspectives

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many of Malpighi’s discoveries were apparently made by useof a single magnifying lens.

Malpighi’s historic description of the pulmonary capillarieswas made in his second epistle to Borelli published in 1661with the title De pulmonibus (5). Early in this letter Malpighibeautifully described how he came to use the frog for hisdissections. He first studied sheep and other mammals butdespite enormous efforts the results were disappointing. In factMalpighi frequently emphasized the tediousness and inade-quate results of most of his dissections.

However, when he eventually used the frog he was jubilantand in a striking passage referred to the frog as the “microscopeof nature.” By this he meant that he was able to visualize, witha relatively small magnification, features as minute as thecapillary network that had eluded him in mammals because thestructures were so small that they could not be seen under hismicroscope. He went on to say that nature is accustomed “toundertake its great works only after a series of attempts atlower levels, and to outline in imperfect animals the plan ofperfect animals.” In other words he is alluding to the fact that,as he sees it, evolution has tried out its advances in “imperfectanimals,” by which he means frogs, before using the samestructures in a more advanced form in the so-called “perfectanimals,” by which he means mammals. He goes on, “ForNature is accustomed to rehearse with certain large, perhapsbaser, and all classes of wild [animals], and to place in theimperfect the rudiments of the perfect animals.”

A little later Malpighi makes a droll statement about theamount of labor the work has taken him. “For the unloosing of

these knots [that is, elucidating these problems] I have de-stroyed almost the whole race of frogs, which does not happenin the savage Batrachomyomachia of Homer.” Here he isreferring to the imaginary fierce battle between frogs and micethat is a parody of the Iliad and that, incidentally, was probablynot written by Homer.

Now turning to Malpighi’s actual observations, after hisabortive efforts on mammals such as sheep he first looked atthe living lung in the frog. However, although he could clearlysee the blood moving rapidly through small arteries, he couldnot determine what eventually happened to it. He then dried thelung of a frog and wrote as follows: “I could not extend thepower of the eye any further in the living animal, hence Ibelieve that the mass of blood poured into an empty space andwas recollected by the outgoing vessels and the structure oftheir walls . . . however, the dried lung of a frog resolved mydoubts. In a very small portion of it . . . there may be seen, witha perfect glass no broader than the eye, the points which arecalled ‘Sagrino’ [dark spots on the surface of the lung of a frog]forming the membrane, but mixed with looped vessels. Sogreat is the branching of these vessels, after they extend outhither and thither from the vein and artery, that no largersystem of vessels will be served, but a network appears, formedby the offshoots of the two vessels. This network not onlyoccupies the whole floor [of the air space] but is extended tothe walls and adheres to the outgoing vessel, just as I couldobserve more abundantly, but with greater difficulty in theoblong lung of a tortoise, which is likewise membranous andtranslucent. Here it lies revealed to the senses that, as the bloodpasses out through these twisting divided vessels, it is notpoured into spaces, but is always passed through tubules and isdistributed by the many windings of the vessels” (translationfrom Ref. 12).

The two letters to Borelli contained two illustrations shownin Figs. 2 and 3. Malpighi’s drawing of the capillary mesh isshown in the bottom part of Fig. 2. This depicts one of thevesicles (alveoli) with its base and various sides that has beenopened up so that the dense network of capillaries can be seenin all the walls. The upper part of Fig. 2 shows the two lungsof the frog. On the left we can see the alveoli and on the rightis another depiction of the capillaries.

Malpighi’s discovery of the pulmonary capillaries was mo-mentous. In fact it was the first description of capillaries in anycirculation. Harvey in De motu cordis in 1628 had supposed, ashad Galen and Columbus before him, that the blood found itsway from the right ventricle through the parenchyma of thelungs into the pulmonary vein and left ventricle. However,Harvey could not see the capillary vessels but called them“pulmonum caecas porositates et vasorum eorum oscilla,” thatis “the invisible porosity of the lungs and the minute cavities oftheir vessels.” It was Malpighi’s great triumph that he was thefirst person to see and describe them.

Discovery of the Alveoli

Malpighi’s description of the alveoli may not have beenquite so momentous as his discovery of the capillaries, but infact it completely changed perceptions of the structure of thelung. Prior to his time the structure and function of the lungwere a mystery. Vesalius, following the writings of Galen, heldthat the lungs had been formed from solidified bloody foam.

Fig. 1. Marcello Malpighi (1628–1694).

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Harvey compared the substance of the lung with that of thekidney or liver and argued that one of its principal purposeswas to cool the animal.

Malpighi refers to these earlier notions near the beginning ofhis Epistle I of De pulmonibus addressed to Borelli. He stated“The substance of the lungs is commonly supposed to be fleshybecause it owes its origin to the blood, and it is believed to benot unlike the liver or the spleen . . . ” [This and the subsequentquotations from Malpighi are from the translation by Young(14)]. But then in the same paragraph Malpighi goes on to drophis bombshell: “By diligent investigation I have found thewhole mass of the lungs, with the vessels going out of itattached, to be an aggregate of very light and very thinmembranes, which, tense and sinuous, form an almost infinitenumber of orbicular vesicles and cavities, such as we see in thehoney-comb alveoli of bees, formed of wax spread out intopartitions. These [vesicles and cavities] have situations andconnection as if there is an entrance into them from the trachea,directly from the one into the other; and at last they end in thecontaining membrane.” Malpighi goes on to refer to one of theillustrations at the end of the Epistles, shown in Fig. 3. Inreferring to the left-hand part of the figure he states that “withgreatest diligence I have been able to make out, those mem-branous vesicles seem to be formed out of the endings of thetrachea, which goes away at the extremities and sides intoampulus cavities.” Parts I and II of the figure show the vesicles(alveoli), and Part III is a diagrammatic representation of thefinal branching of the prolongations of the trachea. He adds,

“Seeing that the air which rushes from the trachea into thelungs requires a continuous path for easy and rapid ingress andegress, whence possible this internal tunic of the trachea, endsin sinuses and vesicles, makes a mass of vesicles like animperfect sponge so to speak.” Although the illustrations arefrom the frog lung, Malpighi also refers to somewhat similarobservations in a dog’s lung.

So Malpighi had discovered that air entering the lung isconducted down a series of what we now call airways into thetiny alveoli, and also that the surface of the alveoli is coveredwith a rich network of blood vessels as shown in image II at thebottom of Fig. 2. He seems to be very close to an understandingof the primary function of the lungs, which is the exchange ofgases between the alveolar space and the blood in the capil-laries. However, this eludes him. Further on in the first Epistlehe states “Concerning the use of the lungs I know that manyviews are held from the ancients onwards, and about themthere is very much dispute—especially about the cooling,which is taken to be the principal purpose, when it strives withthe imagined excessive heat of the heart which may requireeventation; wherefore these things have made me diligentlyinquisitive in the investigations of another purpose and fromthese things which I subjoin I can believe that the lungs aremade by Nature for mixing the mass of blood.” So Malpighi ispoised to understand the gas exchange function of the lung butat the last moment steps away.

Insect Respiration

It is fascinating that, in addition to discovering the pulmo-nary capillaries and alveoli in amphibia and by implicationmammals since he made similar observations in the dog,

Fig. 3. Malpighi’s drawing of the alveoli. I and II: 2 different views of alveoli.III: diagram showing the branching of the trachea ending up in the alveoli.From Ref. 5.

Fig. 2. Malpighi’s drawing of the pulmonary capillaries and alveoli. I: 2 lungswith the alveoli on the left and the capillaries on the right. II: pulmonarycapillaries in a diagram of an alveolus that has been opened up. From Ref. 5.

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Malpighi also was the first person to describe the mode ofrespiration in insects. Admittedly this was not his primarypurpose in his study of the silkworm. Nevertheless he was thefirst person to clearly describe the spiracles and tracheae thatallow oxygen to reach the body tissues of insects.

Malpighi’s work on the silkworm was stimulated by a letterfrom Henry Oldenburg, who was the secretary of the RoyalSociety in London. We have already seen that Malpighi’sreputation as a scientist was so great in 1667 that he wasinvited to become a correspondent of the Society and send hisscientific findings to them for publication. Oldenburg told Mal-pighi that the Society was particularly interested in the silkworm,and indeed Malpighi had previously done some work on thisanimal. However, stimulated by the invitation, he embarked ona major project to describe the anatomy of the silkworm. This

was presented to the Society in a manuscript titled Dissertatioepistolica de bombyce, which appeared in the PhilosophicalTransactions of the Royal Society (7).

Adelmann quotes Cole (2), who described the contents of Debombyce; excerpts are summarized here. Malpighi anatomizedall phases of the species, but, apart from his very remarkableand accurate observations on the genitalia of the moth, thelarva claimed the greater part of his attention. The head of thecaterpillar is described in detail and the arrangement of the legsand their function in crawling is analyzed. However, the mostimportant part from our point of view is that Malpighi de-scribed for the first time the spiracles of insects, and the systemof vessels associated with them (Fig. 4). There are nine pairs ofspiracles in the larva of bombyces, and each spiracle has itsown bundle of vessels (airways), two of which anastomose

Fig. 4. Malpighi’s drawing of the spiracles, tracheae, andnerve trunk of the silkworm. Fig. I: small tracheoles endingnear a nerve ganglion. Fig. II: nerve trunk in the center andthe 9 pairs of spiracles that form the openings of thetracheae on the body surface. From Ref. 7.

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with corresponding vessels in front and behind. The result is alongitudinal spiracular trunk on each side that stretches fromhead to tail. There also transverse anastomoses across the body.Like the arteries, the tracheae from the spiracles go on dividingand diminishing in size until they can hardly be seen even witha microscope. Malpighi described these air tubes as tracheaeand compared them with the lungs of vertebrates. He carriedout additional studies in the cicada, stag beetle, locust, wasp,and bee and identified expansions of the trachea that heidentified with the air chambers of lungs. He combined phys-iological studies with his anatomical observations. For exam-ple, he occluded the spiracles with oil and immediately theanimal developed convulsions and died “intra Dominicae ora-tionis spatium” [during the time it takes to recite the Lord’sPrayer]. He went on to show that if the anterior spiracles areblocked only the corresponding part of the body is affected,and the animal subsequently recovers. Malpighi also describedthe nerve trunk and ganglia, the silk glands, the pulsating bloodvessel extending the whole length of the worm that is theprimitive heart, and the urinary tubules that are still referred toby his name. Figure 4 shows Malpighi’s illustration of asilkworm larva with the spiracles clearly identified. The nervetrunk is also shown with a ganglion on the left and its supplyof tracheae.

This brilliant project enhanced Malpighi’s reputation evenmore. Adelmann (1) lists some of the accolades, which beginwith the statement that Malpighi’s book was generally admit-ted to be the first really great contribution to insect morphol-ogy. One scientist referred to it as a “tissue of discoveries” andwent on to describe it as “a treatise from which one can learnmore about the wonderful internal structure of insects thanfrom all earlier works combined.” Another stated that “Despitethe few months at his disposal for carrying out his anatomicalresearches, Malpighi obtained prodigious results and saw morewith eyes armed with a simple lens than many others who camelater, thought they possessed all the means subsequently of-fered by the perfected physical sciences.”

Embryological Studies

Malpighi’s skills with the microscope prompted him toembark on another major scientific enterprise and the resultwas a description of the development of the chicken embryo infar more detail than had been previously possible. In fact someauthorities argue that his morphogenesis studies were his mostimportant scientific contributions. For example, four of the fivelarge volumes in Adelmann’s exhaustive study of Malpighi aredevoted to his embryological work.

Malpighi was influenced by a philosophical movement at thetime of emphasizing the similarity of biological functions tothose of machines. For example, the French philosopher RenéDescartes (1596–1650) had referred to man as an earthymachine (machine de terre). This was also part of the attitudeof Galileo (1564–1642), whom Malpighi never met but forwhom he had an enormous regard. Malpighi’s friend Borellishared the same notions, and his famous book De motu ani-malium contains many examples. Malpighi’s most importantbook on morphogenesis was Dissertatio epistolica de forma-tione pulli in ovo (Discourse letter on the formation of thechicken in the egg) (8) in which Malpighi draws an analogywith an artisan who “in building machines must first manufac-

ture the individual parts, so that the pieces are first seenseparately, which must then be fitted together.” Elsewhere hewrites “Nature in order to carry out the marvelous operations[that occur] in animals and plants has been pleased to constructtheir organized bodies with a very large number of machines,which are necessarily made up of extremely minute parts soshaped and situated as to form a marvelous organ, the structureand composition of which are usually invisible to the naked eyewithout the aid of the microscope. . . . ”

Malpighi’s philosophy here has some similarities with thatdescribed earlier where he stated that Nature is accustomed “toundertake its great works only after a series of attempts atlower levels, and to outline in imperfect animals the plan ofperfect animals.” Now in De formatione pulli in ovo he statesthat the study “of the first unelaborated outlines of animals inthe course of development” is particularly useful because theartisan Nature forms them separately before combining themwith one another. He gives as an example the miliary glands(embryonic hepatocytes) of the embryo, which will merge toform the liver but are still distinguishable as individual groupsof cells which in crustaceans remain distinct.

Malpighi’s studies of the chicken embryo were very exten-sive. He was particularly interested in the early development ofthe heart including the primitive cardiac tube and its segmen-tation, the aortic arches, and the somites. He also worked onthe development of the nervous system including the neuralfolds, the neural tube, the cerebral vesicles, and the opticvesicles. He studied the heart within 30 h of incubation andnoticed that it began to beat before the blood became red.Malpighi’s drawings of the heart in an embryo 2 days old areshown in Fig. 5. Incidentally, he was also apparently the firstperson to identify red blood cells although he did not under-stand their significance. In an observation that became contro-versial, he stated that he had seen the complete form of theembryo in an unincubated egg. This notion of “preformation”was taken up by subsequent scientists and of course was anerror but Malpighi only alluded to it tangentially.

Botanical Studies

Malpighi’s enormous energy and scientific curiosity areevidenced by the fact that, as we have seen, he not only madefundamental advances on the anatomy of the lungs, but inaddition he carried out extensive studies on the silkworm andembryology of the chicken. His boundless enthusiasm is fur-ther indicated by the following quotation when he referred tohis interest in the investigation of higher animals and added“ . . . but these, enveloped in their own shadows, remain inobscurity; hence it is necessary to study them through theanalogs provided by simple animals. I was therefore attractedto the investigation of insects; but this too has its difficulties.So, in the end, I turned to the investigation of plants so that byan extensive study of this kingdom I might find a way to returnto earlier studies, beginning with vegetant Nature. But perhapsnot even this will be enough, since the simpler kingdom ofminerals and elements should take precedence. At this pointthe undertaking becomes immense, and absolutely out of allproportion to my strength” (1).

Fortunately for the progress of science he resisted the temp-tation to take on minerals and elements. But Malpighi turned tothe study of plants with extraordinary success. The results were

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published in Anatomes plantarum pars prima (9) followed byAnatomes plantarum pars altera (10). In fact, Malpighi iscommonly regarded as the founder of the microscopic study ofplant anatomy along with his English contemporary NehemiahGrew (1641–1712).

Malpighi’s interest in the anatomy of plants began in Mes-sina when he was walking in the garden of his patron VisconteRuffo. He noticed that when a branch of a chestnut tree wassnapped, what appeared to be vascular bundles projected fromthe open end. These were the tubes of the xylem that transportwater and soluble mineral nutrients from the roots to theleaves. They are known as tracheids and the nomenclature isinteresting: trachea in mammals, tracheae in insects, and trac-heids in plants.

Malpighi was immediately reminded of the tracheae ofinsects and this aroused his interest in the nutrition of plants.He then took up the topic with his customary enthusiasm andsystematically studied all aspects of plant anatomy includingthe stem, leaf, root, bud, flower, seed, and seedling. When heexamined the fine tubes in the xylem of the stem that remindedhim of the insect tracheae, he found that these formed annularrings in some plants but scattered bundles in others. Here hewas inadvertently referring to a fundamental difference be-tween the two great families of plants, the dicotyledons and themonocotyledons, although these were not described until muchlater.

Malpighi recognized that the leaves of a plant constituted alaboratory that produced sap, which was essential for plantnutrition. He showed, for example, that when he removed oneof the two leaves of a germinating seedling its growth was

stunted and if he removed both leaves the plant died. He alsodescribed that when the bark of a tree was removed in a ringaround the trunk, a swelling appeared above the ring becauseof obstruction to the sap descending from the leaves in thephloem, which is located in the periphery of the trunk justunder the bark, and the periphery of a stem just under theepithelium.

Malpighi was a talented artist and sketched beautiful sec-tions of plants such as Nigella that he noticed produced honeyin the depths of the flowers. Examples of his drawings areshown in Fig. 6. In keeping with his interest in morphogenesisthat he had demonstrated in his studies of the chick embryo, hestudied the early stages of development of plants such as thebean (Leguminosae) with elegant drawings. Another interestwas galls, which are abnormal growths on the trunks orbranches of some trees. He demonstrated that these werecaused by the deposition of insect eggs.

With Nehemiah Grew, Malpighi is known as a founder ofthe microscopic study of plant anatomy. Both of them com-municated their publications to the Royal Society in the sameperiod and they were on cordial terms. Grew became thesecretary of the Society after the retirement of Oldenburg.

Other Studies

It is impossible to summarize all the scientific contributionsof Malpighi. However, some others should be briefly men-tioned. He described the papillae of the tongue and postulatedtheir role in taste. In studies of the skin he described a layer ofcells that now bears his name. Additional studies were made on

Fig. 5. Drawings of the developing heart in the chickembryo. The blood comes from the veins A and passesinto the auricle (atrium) B. Sometimes there is a briefintermediate channel as the blood is pushed into theright ventricle C and from there into the left ventricle D.Finally the blood enters the arteries E and from themmoves either to the head F or to the umbilical vessels G.From Ref. 8.

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Fig. 6. Botanical illustrations by Malpighi. He was particularly interested in the reproduction of plants and many drawings show the sex organs including thestamens, anthers, pistils, stigmas, and styles. From Ref. 9.

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the brain with particular reference to the white matter that heshowed consisted of bundles of fibers.

His book De viscerum structura exercitatio anatomica (6)describes the histology of the kidney, spleen, and liver. He sawthe glomeruli in the kidney and for a time these were namedafter him. Small groups of lymphatic bodies in the spleen wereknown as Malpighian corpuscles. In the liver he saw smalllobules that he concluded represented the fundamental unit ofthat organ.

Malpighi’s Difficulties

It would be natural to think that a man as original, industri-ous, and productive as Malpighi would live a satisfying life.Unhappily this was not the case. First he was unfortunate inthat he suffered from ill health for most of his life. Moreimportant was the savage criticism of much of his scientificwork. In part this was the result of professional jealousy.However, in addition there was criticism of Malpighi’s scien-tific approach. Although we marvel at how much he achievedwith an early microscope, many of his detractors argued thatthis type of research was useless and it would not lead toimprovements in medical treatment.

A tragedy in 1659 had a lasting influence on Malpighi. Hisbrother Bartholomew got into an argument with the eldest sonof Durolamo Sbiraglia and fatally stabbed him. The result wasa smoldering feud between the two families and a lifelongbitter criticism of Malpighi’s work by Sbiraglia, who was alsoa scientist. Perhaps there are echoes of the feud of the twofamilies in Shakespeare’s Romeo and Juliet. Malpighi wasfinally driven to publish a long bitter response to his critics inhis Opera posthuma published in 1697 (11). However, historywill always regard Malpighi as one of the most outstandingbiologists.

NOTE ON SOURCES

By far the best source of information is the five-volume work byAdelmann, Marcello Malpighi and the Evolution of Embryology (1).The first volume is a general account of Malpighi’s life and work anddespite its length makes excellent reading. A review of the book by

Wilson (13) is a good introduction. Also useful are E. J. Cole’sHistory of Comparative Anatomy (2) and the chapter on Malpighi inMichael Foster’s Lectures on the History of Physiology (4). The lattercontains another translation of part of De pulmonibus.

Access to Malpighi’s publications has been greatly improvedbecause of the Internet. Publications with reference numbers 5, 7, 8,9, and 10 are on the Web. Some of the original publications are inspecialized libraries, especially that of the Royal Society in London.There are English translations on the Web of De pulmonibus byYoung (14), De polypo cordis by Forrester (3), which forms part ofRef. 6, and Pulli in ovo by E. Corti.

DISCLOSURES

No conflicts of interest, financial or otherwise, are declared by the author(s).

AUTHOR CONTRIBUTIONS

J.B.W. conceived and designed research, drafted manuscript, edited andrevised manuscript, approved final version of manuscript.

REFERENCES

1. Adelmann HB. Marcello Malpighi and the Evolution of Embryology (5vols). Ithaca, NY: Cornell University Press, 1966.

2. Cole FJ. A history of comparative anatomy. London: Macmillan, 1949.3. Forrester JM. Malpighi’s De polypo cordis: an annotated translation.

Med Hist 39: 477–492, 1995.4. Foster M. Lectures on the History of Physiology. Cambridge, UK;

Cambridge Univ. Press, 1924.5. Malpighi M. De pulmonibus. London: Philosophical Transactions of the

Royal Society, 1661.6. Malpighi M. De viscerum structura exercitatio anatomica. Bologna:

Jacopo Monti, 1666.7. Malpighi M. Dissertatio epistolica de bombyce. London: Philosophical

Transactions of the Royal Society, 1669.8. Malpighi M. Dissertatio epistolica de formatione pulli in ovo. London:

Philosophical Transactions of the Royal Society, 1673.9. Malpighi M. Anatomes plantarum pars prima. London: Philosophical

Transactions of the Royal Society, 1675.10. Malpighi M. Anatomes plantarum pars altera. London: Philosophical

Transactions of the Royal Society, 1679.11. Malpighi M. Opera posthuma. London: Churchill, 1697.12. Wilson LG. The transformation of ancient concepts of respiration in the

seventeenth century. Isis 51: 161–172, 1960.13. Wilson LG. Malpighi and seventeenth-century embryology: an essay

review. J Hist Med Allied Sci 22: 190–198, 1967.14. Young J. Malpighi’s “De pulmonibus.” Proc R Soc Med 23: 1–11, 1929.

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