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Review
The diffuse nervous network of Camillo Golgi: Facts and fiction
Elio Raviolaa,⁎, Paolo Mazzarellob
aDepartment of Neurobiology, Harvard Medical School, Boston, MA, USAbMuseum for the History of the University of Pavia and Department of Experimental Medicine, University of Pavia, Italy
Please cite this article as: Raviola, E., MazRev. (2010), doi:10.1016/j.brainresrev.201
A B S T R A C T
Article history:Accepted 8 September 2010
The name of Camillo Golgi is inextricably associated, in the mind of most neuroscientists,with the theory that nerve cells communicate with one another by means of an intricatenetwork of anastomosing axonal branches contained in the neuropil intervening betweencell bodies in the gray matter of the brain and spinal cord. Examination, however, of Golgi'sdrawings in the papers published in the decade intervening between publication of hismethod (1873) and the beginning of his studies on malaria (1885) shows that axonalarborization in the cerebellar cortex and olfactory bulb are depicted as independent of oneother. This is in striking contrast with the drawings included by Golgi in his 1906 Nobellecture where the entire granular layer of the cerebellar cortex is occupied by a network ofbranching and anastomosing nerve processes. Thus, Golgi in his original papers on thecerebellum represents nerve cells as discrete units and only later in life merges axonalarborizations in the context of a lecture in defense of the reticular theory.
For most biomedical scientists the name of Camillo Golgi(1843–1926) is inevitably eponymous with the cytoplasmicorganelle he discovered. Among neuroscientists, he is chieflyremembered because of his dichromate-silver staining meth-od for nerve cells and the fact that he proposed and staunchlydefended a reticularist theory of the connections betweenneurons. The extent of his contribution to our understandingof the anatomy of the nervous system is generally ignored, asare his fundamental discoveries on malaria. It is therefore of
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some interest to reassess with the eye of a modern neurosci-entist Golgi's evidence during the decade that followed thediscovery of the black reaction and compare it with the imageshe presented in his 1906 Nobel prize lecture. It should be notedthat – for what we know – Golgi himself made all hispreparations and drawings of the nervous system, perhapswith some help for the cerebellum by his pupil R. Fusari. Wewere therefore surprised that some of the illustrations in hisoriginal papers are at odds with his ideas of a “diffuse nervous
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network”. In the present paper, we limit our discussion to thecerebellar cortex, because comparison of drawings drafted indifferent moments of his career and for different purposesuncover an episode in which Golgi's strongly held persuasionled him to betray his devotion to the microscopic evidence.
It is difficult to imagine today the excitement that Golgimust have felt in observing for the first time nerve cells intheir entirety and in extraordinary detail. One can guess theimpact of a Golgi specimen on an observer of the 19th centuryfrom the reaction of Cajal in 1887 when slides impregnatedwith the chromoargentic technique were shown to him byLuis Simarro Lacabra (Ramón y Cajal, 1989): they changed thecourse of his entire scientific life. Nothing of the kind had beenseen before. In 1865, Deiters (his manuscript was publishedposthumously by his mentor Max Schultze) had reported thatnerve cells had two types of processes: the “protoplasmicprocesses” (the term “dendrites” was introduced by W. His in1889), so called because their cytoplasm had the samestructure as that of the cell body, and the “nerve process”,one per cell and light-refracting, that became the axis of amyelinated nerve fiber and was therefore named “Achsency-linder” by Rosenthal in 1839 (for all above references seeShepherd, 1991) and “cylindraxis” by English-speakingauthors. The origin of the term “axon” was attributed toKölliker by Lenhossék (Lenhossék, 1895). Nonmyelinatedaxons and their arborizations, however, had never been seenin the gray matter of the central nervous system and theprotoplasmic processes soon lost their individuality as they
Fig. 1 – Human cerebellar cortex: dendrites and cell bodies are blacgranular layer are marked by black dots. No anastomoses betwee
Please cite this article as: Raviola, E., Mazzarello, P., The diffuse nRev. (2010), doi:10.1016/j.brainresrev.2010.09.005
merged into the structureless substance that seemed to fill thespaces between the neuronal cell bodies
Yet, anybody who has tried the Golgi method is aware ofthe difficulty in obtaining a well-impregnated section ofnervous tissue. Most likely, Golgi's discovery began as anaccidental observation, possibly during his studies of theperivascular connective tissue of the brain or Virchow–Robin'sspaces (1869–1870; Golgi, 1870), after immersing in silvernitrate a specimen stored in potassium dichromate (Müller'sfluid) to stain the intercellular boundaries between endothe-lial cells (see Mazzarello, 2010). This initial observation wasanything but trivial, because a neuron had never been seenbefore in its entirety, and was probably followed by endlessmodifications of the technique, which is notoriously fickle.Furthermore, the very notion that only a few neurons werestained was counterintuitive, for the dyestuffs known at thetime, such as carmine, would stain all cell nuclei. Golgi musthave been concerned by the scarce reproducibility of the“black reaction” and, during his initial trials, must havedeveloped the preoccupation with technical artifacts thattormented him throughout his life, became an obsession withhis pupils and a trait of the entire biomedical establishment ofthe University of Pavia during the first half of the past century.
Artifacts were routine with the chromoargentic impregna-tion: with immersion fixation, because of the slow penetrationof dichromate and osmium (about 1 mm per hour), only themost superficial region of the tissue block was well preserved.Dendrites and axons deep in the tissue would acquire a
k, axons are red. Inset: Free endings of axonal branches in then axonal branches are present in this area of the drawing.
ervous network of Camillo Golgi: Facts and fiction, Brain Res.
Fig. 3 – Pyramidal cells in the posterior end of the humansuperior occipital gyrus, an extrastriate region that is beyondknown topographically organized areas. Note the absence ofdendritic spines.
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beaded appearance because mitochondria would swell as aresult of anoxia: this was especially true for the humanspecimens that Golgi obtained after autopsy. Thus, he musthave compared the appearance of neurons in brains fixed indichromate by immersion with those fixed by perfusion,because he never published drawings of beaded neurons. Inthis respect, we know that Golgi was the first to use perfusionfixation by arterial injection of dichromate (Golgi, 1885).
With his staining method, in the years between 1873 and1886, Golgi laid the cornerstone of modern neuroanatomy: hedefined the precisemorphology of neurons and glia, dendritesand axons, projection neurons and interneurons. He providedthe first description of the structure of the cerebellum,olfactory bulb, hippocampus and cerebral cortex. In thecerebellum (Golgi, 1874, 1885), he described Purkinje cells,
Please cite this article as: Raviola, E., Mazzarello, P., The diffuse nRev. (2010), doi:10.1016/j.brainresrev.2010.09.005
stellate and basket cells, Golgi cells and granule cells (Fig. 1).He discovered the recurrent collaterals of the axon of Purkinjecells, but ignored their dendritic spines. Hemissed the basketssurrounding the soma and initial segment of Purkinje cellsand the continuity of the axons of granules with the parallelfibers. He saw, but did not identify, climbing andmossy fibers.In his masterful description of the olfactory bulb (Golgi, 1875,1885, 1903), he identified mitral, tufted, periglomerular andgranule cells (Fig. 2). He provided an accurate description ofthe composition of the glomeruli and noted the absence of an
ervous network of Camillo Golgi: Facts and fiction, Brain Res.
Fig. 5 – Olfactory bulb (dog): incoming olfactory fibers andcentrifugal fibers are black, whereas the arborizations of theaxons of periglomerular cells and tufted cells are blue. Axonalbranchesend freely inboth the glomeruli (A) and themolecularlayer (B); there is no “diffuse nervous network”. Curiously, inone glomerulus, three olfactory axons anastomose with oneanother (arrowheads).
Fig. 4 – Pyramidal cells of the hippocampus (rabbit). Again,note the absence of dendritic spines.
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axon in granule cells. In the cerebral cortex, he publishedstunning images of the pyramidal cells in the precentral gyrusand occipital cortex (Fig. 3; Golgi, 1882), but in the precentralgyrus, he missed the giant pyramidal cells of Betz.
Golgi, however, made a crucial error of interpretation in hisfirst paper (Golgi, 1873), when he attributed to the dendritesthe role of “organs of nutrition” based on the hypotheticalrelationship of their branches with the “cells of the interstitialtissue” and blood vessels. Throughout his life, he stuckuncritically to this unproven idea, a surprising attitude insuch a careful scientist who was so preoccupied with themicroscopic evidence. This perhaps explains why Golgiomitted the dendritic spines in his drawings of the Purkinjecells (Fig. 1) and pyramidal cells of the cortex (Fig. 3) andhippocampus (Fig. 4; Golgi, 1883b). He undoubtedly saw thespines, but at this stage of his discoveries, he probablyconsidered them an irrelevant detail, unrelated to thecomputations carried out by neurons.
If dendrites, however, and the cell body had a trophicfunction, Golgi was left with the alternative that neurons hadto communicate with one another through their axons. At thetime, the simplest assumption was the existence of “anasto-mosis between the nervous filaments which originate fromthe continuation of the cylinder axis of different nerve cells”:in a nutshell, the “diffuse nervous network” that Golgi woulddevelop in successive papers. Thus, Golgi did not seem to bepreoccupied by the fact that nerve cells would be joined in asyncytium and therefore would represent an exception to thetheory that cells were elementary, independent constituentsof animal and plants, the most powerful biological general-ization of the first half of the 19th century, originally stated bySchleiden and Schwann in 1838–1839 and developed by Remakand Virchow in the 1850's (Harris, 2000).
However, upon close inspection of the drawings publishedby Golgi during the decade that followed the invention of hismethod, we noted that the diffuse nervous network is absent.In a drawing of the cerebellar cortex, made in collaborationwith Fusari (Fig. 1, corresponding to Table XI of Golgi, 1883a,1885), the recurrent collaterals of the axons of Purkinje cellsascend toward the deep boundary of the molecular layer, butdo not mingle with the plexus formed by the axons of basket
Please cite this article as: Raviola, E., Mazzarello, P., The diffuse nRev. (2010), doi:10.1016/j.brainresrev.2010.09.005
cells. Some of the ascending axons originating from the whitematter (probably mossy fibers) branch and freely terminate inthe granular layer. Other ascending axons (probably climbingfibers) reach the molecular layer and intersect but do notmerge with the tangential plexus of the basket cell axons.Axonal branches running a tangential course in the deep partof the molecular layer apparently give rise to collateralbranches at a right angle which descend into and freelyterminate in the granular layer: but these are probably parallelfibers intermingled with the axons of basket cells and theirdescending collaterals are actually the parent axon emanatingfrom the granule cells. In the drawing by Golgi of the granularlayer of the cerebellar cortex, we counted approximately 140free endings of axonal branches over an area of 105 μm2 (Fig. 1,inset). Excluding intersections, in only three cases does abranch of an axon merge with that of another axon. But theseapparent anastomoses could just represent noise in thedrawing, because Golgi does not attract the reader's attentionto them. Even in the paper of Golgi's student Fusari (1883)bridges between “arcuate fibers” (a mixture of basket cellaxons and parallel fibers) are very few.
In Golgi' illustration of the anatomy of the olfactory bulbmost of the branches of the olfactory axons appear toterminate freely in the glomeruli (Fig. 5A) and there is not a
ervous network of Camillo Golgi: Facts and fiction, Brain Res.
Fig. 6 – Cerebellar cortex in the drawing made by Golgi toillustrate his Nobel lecture. In contrast with the drawing inFig. 1, axonal branches extensively anastomose with oneanother in the granular layer. Baskets surrounding Purkinjecell bodies are now present in the drawing.
Fig. 7 – In this drawing, the branches of the axonalarborizations of basket cells extensively anastomose as theysurround the perikarya of Purkinje cells. Pinceaux nowsurround the initial segment of the Purkinje cell axon, butsome of their fibrils continue into the diffuse nervousnetwork of the granular layer.
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single case of anastomosis between axonal branches in anyother layer of the bulb (Fig. 5B). Curiously, Golgi drewanastomoses between the dendrites of mitral and periglo-merular cells and at least one clear-cut case of anastomosisbetween branches of centrifugal axons and dendrites of aperiglomerular cell. We assume that these connections – inGolgi's mind – were meaningless from the point of view ofneural processing because of his belief that dendrites had atrophic function. Finally, it is of note that incoming olfactoryaxons merge into one another in two glomeruli, giving rise tocurving loops.
Thus, Golgi drawings show that anastomoses betweenaxonal branches of different nerve cells are uncommon. Onthe other hand, Golgi's writings, in these initial years of hisresearch, after he excluded dendrites' participation in neuro-nal communication, betray a concern that the cases ofcontinuity between the axonal branches comprising thediffuse nervous network were too few to account for themassive number of computations carried out by the nervoussystem, as we would say today.
By the mid-eighties, Golgi's interest in the nervous systemhad waned. This is not surprising, for he had cornered himselfinto an awkward predicament: by excluding dendrites fromcomputations and postulating a “diffuse” network as thesubstrate for neuronal communication, there was nothingreally interesting left to discover in the brain. It is surprising,however, that this brilliant histologist never expressed inwriting two critical questions: Why do neurons have suchdifferent shapes? And why are different morphological types ofneurons so rigorously stratified in cortices? Is it likely that henever considered the possibility that neuronal diversity was thevisible expression of the specificity of their connections.
Meanwhile, as a general pathologist, he could not beindifferent to the discoveries by the great bacteriologists of
Please cite this article as: Raviola, E., Mazzarello, P., The diffuse nRev. (2010), doi:10.1016/j.brainresrev.2010.09.005
the time who had just identified the microorganisms respon-sible for anthrax, tuberculosis and cholera. Drawn into thecontroversy about the agent of malaria by a colleague in hismedical school, he spent the next decade of his life studyingthe pathophysiology of the disease, which was at the timequite common in the countryside around Pavia (Mazzarello,2010). His fundamental discoveries were the observation thatthe febrile attacks coincided with cell division of the Plasmo-dium inside red blood cells, hemolysis and release of theparasites into the blood stream (Golgi, 1886a,b). Furthermore,he showed that different species of Plasmodia were respon-sible for the tertian and quartan fevers, which recur atapproximately two- and three-day intervals respectively(Golgi, 1889a). These discoveries, based on careful microscopicobservations of red blood cells, made Golgi instantaneouslyfamous internationally, in contrast to the slow acceptance ofhis findings on nerve cells. As a further evidence of hisdiminished interest in the nervous system, during this periodof time Golgi also published interesting studies on thedevelopment of the kidney (Golgi, 1888, 1889b).
While Golgi's mind was occupied with malaria, a revolu-tion was happening in neurobiology: as W. His (1886), Forel(1887) and Waldeyer (1891) formulated the neuron theory,Santiago Ramón y Cajal in a burst of superhuman activity (48papers published between1888 and 1892) provided incontro-vertible anatomical evidence in favor of this theory by usingGolgi's technique. Golgi's did not accept the new discoveriesand criticized Cajal's results on the grounds that he did notknow the black reaction well, his impregnations wereincomplete, he did not repeat the method a sufficient number
ervous network of Camillo Golgi: Facts and fiction, Brain Res.
Fig. 9 –Anearlier versionof thedrawing in Fig. 6, sent byGolgito L. Luciani. Both drawings contain the samebasket cells, butdifferences in their dendritic trees (e.g. the dendrites labeledwith asterisks are absent in Fig. 6) suggest that the cells weredrawn at the microscope in two different occasions and atslightlydifferent depths of focus. Furthermore, both drawingscontain nine Purkinje cells and therefore have roughly thesame tangential width. In this figure, however, the granularlayer is approximately half the radial thickness of that inFig. 6. Clearly, at least in Fig. 6, this layerwasnot drawnwith acamera lucida and does not represent a faithful illustration ofthe specimen.
Fig. 8 – The diffuse nervous network in the granular layersurrounds empty spaceswhose diameter ismuch larger thanthe perikaryon of a granule cell (compare with the outline ofthe cell body of a Purkinje cell at the arrowhead and the sizeof the granules in Fig. 6). These spaces do not correspond tocerebellar glomeruli, for these are clearly represented asregions of higher complexity of the network. Clearly, Golgihas overestimated the diameter of a granule, a surprisingmistake for such an astute microscopist.
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of times, in many different animal species, at different ages;and, finally, that his images did not reflect the complexity ofthe microscopic evidence.
Golgi's scientific activity, however, had slowed down be-cause of his commitments as rector of the University of Pavia(1893–1896) and a member of the municipal legislative body ofthe city. He was back in the laboratory between 1896 and 1899,during which he reported the presence of a new cytoplasmicorganelle in nerve cells, the internal reticular apparatus, knowntodayas theGolgi complex (Golgi, 1898). But after this three-yearstint, he permanently abandoned major scientific projectsbecause of his numerous academic and government commit-ments: he was in fact appointed a member of the senate by theking (1900), became dean of themedical school (1899–1901) and,again, rector of the university (1901–1909).
In 1906, Golgi and Cajal were awarded the Nobel Prize inPhysiology or Medicine in recognition of their work on thestructure of the nervous system. Golgi's criticism of theneuron theory and his defense of the diffuse nervous networkin his Nobel lecture are well known and have been discussedin detail elsewhere (Mazzarello, 2010). Here, however, we areconcerned with three drawings of the cerebellar cortex thatillustrate Golgi's Nobel lecture (Golgi, 1907). These drawingsare dramatically different from those published in the 1870's:the granular layer is occupied by branching and anastomos-ing, sinuous process, continuous with the baskets thatsurround the perikarya of Purkinje cells (Figs. 6–8). Freeendings are few and far apart and arborizations of individual
Please cite this article as: Raviola, E., Mazzarello, P., The diffuse nRev. (2010), doi:10.1016/j.brainresrev.2010.09.005
axons cannot be distinguished from one another. As a whole,the processes give rise to a network that intercepts openingsof varying diameter and irregular shape, in which arecontained the cell bodies of the cerebellar granules. In patches(Fig. 8), this network is denser, possibly Golgi's rendition of thecerebellar glomeruli.
Any observer, who is familiar with the appearance of aplexus of dendritic and axonal branches in a Golgi-impreg-nated specimen, would immediately recognize that thesedrawings do not reflect reality. Equally unrealistic is thedepiction in Fig. 7 of the axonal baskets that surround theperikarya of Purkinje cells, whereas those in Fig. 6 are moreauthentic. Finally, the openings of the network in the granularlayer aremuchwider in Fig. 8 than in Fig. 6, ostensibly becausethemagnification is higher. However, judging from the outlineof two Purkinje cell bodies, the size of the openings is too largeto enclose the perikaryon of an individual granule cell(alternatively, the Purkinje cells are drawn too small). Howcould Golgi overlook such an obvious anatomical point?
Howmuch of these drawings depicted actual data and howmuch represented an idealized rendition of his diffusenervous network? A useful hint at the answer is obtained bycomparing Fig. 6 with a drawing of the cerebellar cortex (Fig. 9)that Golgi sent to L. Luciani, the chairman of physiology in theUniversity of Rome four years before he received the prize(May 30, 1902). Both the drawing and an excerpt of theaccompanying letter were published by Luciani in his textbookof human physiology (Golgi, 1905). The two figures exhibit
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significant differences: both of them contain the same twobasket cells, but – most likely – they were drawn at themicroscope on separate occasions at different depths of focus,because their dendritic arbors are not identical. But, if one usesthe dendrites of the basket cells as a yardstick, the granularlayer in the 1906 drawing is about 2 times thicker than that of1902. Thus, there is no doubt that, at least in one of the twofigures, the granular layer was not drawnwith a camera lucidaand its thickness does not reflect reality.
We cannot judge Golgi's distortion of the microscopicevidence using today's ethical criteria. Combining fact withfiction in a drawing was common at the turn of the 19thcentury. At a time in which photography of microscopicobjects was in its infancy, drawingswere themain tool used toillustrate data in a paper and investigators were not specifi-cally required by journals to use the camera lucida inrepresenting the details of the specimen. Modern day obses-sion with image manipulation is the consequence of theintroduction of computer graphics: still in the 1970's, electronmicroscopists removed stain precipitates from their micro-graphs and, in the era before patch clamp, electrophysiologistswould retouch the upstroke of action potentials. It issurprising, however, that Golgi, a skilled microscopist and apositivist who preached the most rigorous respect for themicroscopic detail, abandoned his usual caution in the Nobellecture and displayed images so profoundly different fromthose he had published in the 1870's.
Speculating on the psychological underpinning of thebehavior of an individual who lived a century ago is fraughtwith uncertainty and may be an exercise of arguable value,but it may be useful to list a series of circumstances thatcould have contributed to such incongruous conduct: anas-tomosis between the axonal branches of nerve cells was aplausible hypothesis in the scientific world of the 1870s andin keeping with popular holistic views of nerve function; in1906, the hypothesis of the diffuse nervous network had beensuccessfully challenged by the neuron theory and thereforeneeded vigorous defense; Golgi's reputation and that of his“school” were in jeopardy, including the future academiccareer of his students; he never took into serious consider-ation Cajal's data and, anyhow, clashes between schools ofthought spiced up 19th century biology; he perhaps felt angertoward a younger, inexperienced competitor – he wasn'teven German! – who had misused the technique he hadinvented; he was no longer personally active in the labora-tory, although he continued to supervise his pupils; stubbornand self-assured, he never abandoned the idea that dendriteswere trophic organs for the cell body, although he reluctantlyadmitted in the Nobel lecture that they could “share in theparticular function which we attribute to the nerve cell”; orhe was simply aging and out of touch with the progress of afield he had long abandoned.
If, however, we focus our attention on his early contribu-tions, if we consider the episode of the Nobel prize lecture asan anomaly in an otherwise stellar career, if we disregard thecircumstance that Golgi's prestige had validated the papers ofa number of reticularists who were employing reduced silvertechniques, then this extraordinary microscopist can berightfully restored to his pedestal as one of the foundingfathers of modern neuroscience.
Please cite this article as: Raviola, E., Mazzarello, P., The diffuse nRev. (2010), doi:10.1016/j.brainresrev.2010.09.005
Acknowledgments
The authors thank Dr. Giovanni Berlucchi for pointing out toone of us (E.R.) Golgi's letter and drawing published inLuciani's textbook and both Drs. Berlucchi and Richard T.Born for reading the manuscript.
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