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Historical Entry #1: Early Theories of the Nervous System (1600s-1700s)

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Historical Entry #1: Early Theories of the Nervous System (1600s-1700s)

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In the 1600s-1700s, the understanding of the nervous system was limited to what researchers knew about philosophy and very simple anatomical ideas. Rene Descartes proposed the idea that the body is a machine and the nervous system functioned as tubes that were filled with fluid. Giovanni Aldini, an Italian scientist, conducted some of the first electrical experiments in hopes to understand more about the role of electricity in the nervous system. These early thoughts were important for the later work of scientists who focused on investigating the nervous system. Though technology was not developed to what it is today, the philosophical debates about the body and the mind and electricity of nerves were huge when advancing the scientific understanding that led to Camillio Golgi's silver staining technique in the 19th century.

Source:

https://web.stanford.edu/class/history13/earlysciencelab/body/nervespages/nerves.html

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"By means of nerves, the pathways of the senses are distributed like the roots and fibers of a tree." --Alessandro Benedetti, 1497

Nerves proved to be a fairly difficult part of the body to categorize. The origins of the word "nerve" -- initially a Greek word meaning tendon or sinew -- suggests a certain confusion between connective tissues and other, more subtle types of physical connections within the body. As late as the twelfth century, the Jewish philosopher Moses Maimonides observed: "One who is not knowledgeable in anatomy may mistake ligaments, tendons and chords for nerves." Nonetheless, ancient medical practitioners did understand that nerves served roughly two functions: movement and sensation. The question was how they operated and under the direction of which principal organ.

In the fourth century B. C., the Greek philosopher Aristotle believed firmly that the nerves were controlled by and originated in the heart because it was, in his interpretation, the first organ of the body and the seat of all motion and sensation. Not surprisingly, he was misled by his confusion between ligaments and nerves in drawing this conclusion. Six centuries later, the Roman physician Galen contradicted him, disparaging those "who know nothing of what is to be seen in dissection." Instead he concluded that the brain was the most important organ of the body, with the nerves emanating from it:

"I have shown in my book On the Teachings of Hippocrates and Plato that the source of the nerves, of all sensation, and of voluntary motion is the encephalon [the brain] and that the source of the arteries and of the innate heat is the heart."

Galen saw the spinal cord as an extension of the brain which carried sensation to the limbs. He believed th at the nerves controlled the actions of muscles in the limbs, and that the two principal functions of the nervous system, sensation and motion, were governed by two different types of nerves: respectively soft and hard. He further insisted on a curious anatomical feature of the nerves, imagining them to be hollow tubes. Quite logically, he reasoned that this must be so in order for the animal spiritus, the body's principal source of vitality in his system, to circulate throughout the body. As the Renaissance illustration here indicates, the investigation of the nerves after Galen also became an inquiry into the effect of the brain on the body.

Medieval physicians, in agreement with Galen, believed nerves were offshoots of and controlled by the brain. The Islamic medical philosopher Avicenna wrote in the early eleventh century that "Nerves are one of the 'simple members' -- homogeneous, indivisible, the 'elementary tissues' (others include the bone, cartilage, tendons, ligaments, arteries, veins, membranes, and flesh)." He offered a more precise physical description of them -- "white, soft, pliant, difficult to tear." He and his contemporaries began to describe the complex and varied arrangements of nerves throughout the body, attempting to differentiate further their functions. In the Canon of Medicine, he observed: "Dryness in the nerves is the state which follows anger." Such statements suggest that Avicenna also believed the nerves to be entangled with and responsive to the emotions, yet another sign of their strong connections to the brain.

Slightly more than a century later, Master Nicolaus offered a more precise vocabulary to express the new complexity of the nervous system, discarding the terms "soft" and "hard" for the more familiar idea of sensory and motor nerves. He considered nerves subservient members of the brain that carried out the animal spirits to all members, endowing them with sensation and motion. He further differentiated their points of origin and termination:

"According to some authorities, all the sensory nerves originate from the cellula phantastica, the motor from the cellula memorialis. There are also five kinds of sensory nerves, which are classified according to the operations of the five senses, namely, sight, hearing, smell, taste, and touch. Two nerves arise from the cellula phantastica and cross in the middle of the forehead, one of them passing to the pupil of the right eye, the other to the left. Through these nerves visual spirits are conveyed to the pupils." Leonardo da Vinci's famous image of the human head makes these cells quite apparent in a drawing that clearly is based on medieval theories of the body rather than actual dissection. Many medieval anatomists devised elaborate theories of how the nerves created emotions and sensations in different parts of the body through their attachment to principal organs and members.

The writings of the Renaissance physicians reflect a mixture of ancient and medieval beliefs about the nervous system as well as new knowledge from anatomical dissections. In 1520, Alessandro Achillini could still write, much in the spirit of Galen, that "the nerves are light to receive the spirit and thin in order to offer swift and easy passage to the spirit and flexible to serve the members." Discussions of the nervous system increasingly reflected a mixture of close reading of ancient texts and knowledge culled from dissections. In the late fifteenth century, for example, Alessandro Be nedetti might still subscribe to the idea of spiritus, but his physical description of the nervous system was considerably more precise and detailed because of his examinations of cadavers. Similarly, his contemporary Leonardo da Vinci initially relied quite heavily on the writings of Avicenna and others in describing the nervous system and believed the heart to be the first organ. But this did not prevent him from recognizing that the nerves emanated from a different source: "The medulla is the source of the nerves which give voluntary motion to the members." Leonardo grew increasingly more confident of his own observations, ingeniously using a wax injection technique to obtain casts of the cerebral ventricles and used frogs for physiological experiments on the nervous system. From his studies he concluded that the spinal cord was the center of life. Look at his drawing of the spinal cord. How is he trying to convey the positioning of the nerves?

Throughout the sixteenth and early seventeenth centuries, ancient views of the nervous system existed in harmony with new findings. Vesalius, and even Descartes in his Treatise on Man (written between 1629 and 1633 and published in 1664), continued to speak of the place for animal spirits to roam throughout the body. In fact, Descartes continued to belief that the nerves were still hollow. Helkiah Crooke's Microcosmographia (1631) put it quite eloquently:

"The Nerves are nothing else but productions of the marrowy and slimy substance of the Brain, through which the Animal spirits do rather beam than are transported. And this substance is indeed more fit for irradiation then a conspicuous or open cavity, which would have made our motions and sensations more sudden, commotive, violent and disturbed, whereas now the members receiving a gentle and successive illumination are better commanded by our will and moderated by our reason."

Yet the space for animal spirits grew smaller and smaller as the anatomy and physiology of the nervous system became more elaborate. By 1653, William Harvey still might cite medieval authorities: "According to Avicenna, the nerves are like plantings of the brain, and provide ready intelligence for the organs of sensation; like the fingers of the hand; wherefore the brain neither sees or hears, yet knows all things." But he also took great pride in his century's extensive anatomical studies that had identified the optic, auditory, and olfactory nerves. And despite Descartes' insistence on the hollow nerve, the Scottish medical student John Moir record in his 1620 lecture notes: "nerves have no perceptible cavity internally, as the veins and arteries have."

Both Harvey and Descartes represent a transitional moment in which the animate powers of the nervous system were in doubt but no fully satisfactory alternative, that would explain how unseen messages were transmitted through the body, was yet in place. Harvey wrote in his Lectures on the Whole of Anatomy: "For the nerves only carry down, they do not act, move, nor are they sentient by a faculty, but are organs." Descartes offered an important revision of the nervous system as responding to "different kinds of particle motion (we should term these stimuli)" which stimulated the nerve endings, producing a series of successive motions that connected nervous fibers -- a new and much more physically precise description of the structure of the nerve -- to the brain. He concluded: "The essence of motor control is, then, the direction of animal spirits into the proper interfilamentous channels for transmission to the proper nerve."

As of 1681, a new word had appeared in English -- "neurologie" -- coined by Thomas Willis to described the study of neuroanatomy. Not until the late eighteenth century did physiologists begin to make the connection between recent work on electricity and theories of the nervous system.
QUESTIONS: WHAT DIFFERENCE DOES IT MAKE IF THE NERVE IS A HOLLOW TUBE OR A FIBER? WHY WAS THE THEORY OF ANIMAL SPIRITUS SO LONG-LIVED? Return to History of the Body Home Page Some Additional Readings

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"By means of nerves, the pathways of the senses are distributed like the roots and fibers of a tree." --Alessandro Benedetti, 1497 Nerves proved to be a fairly difficult part of the body to categorize.

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Historical Entry #2: The Discovery of Nerve Fibers (1800s)

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Historical Entry #2: The Discovery of Nerve Fibers (1800s)

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In the early 1800s, researchers like Charles Bell and Francois Magendie began to discover groundbreaking information about the nervous system. Bell and Magendie's work discovered the direction of nerve impulses. Bell showed that the sensory nerves transmitted signals to the brain and the motor nerves carried signals from the brain to the muscles. This discovery was critical for advancing the idea that nerves has a specific function and their own pathways. This thought also led to developments with neuroanatomy, like detailed studies of individual neurons but Golgi.

Source:

https://www.med.uio.no/klinmed/english/research/news-and-events/news/2015/nerve-fibres-secrets-revealed.html

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Nerve fibres’ secrets revealed - Institute of Clinical Medicine

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Nerve fibres, which transmit impulses from neurons, play a key role in the nervous system. Until now, no-one knew how they formed.

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Historical Entry #3: The Birth of Cellular Theory in Biology (Mid 1800s)

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Historical Entry #3: The Birth of Cellular Theory in Biology (Mid 1800s)

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The development of cellular theory in the mid-1800s by botanist Matthias Schleiden and zoologist Theodor Schwann revolutionized thinking in biology. They discovered that all living organisms were composed of cells. This was a fundamental idea that included the nervous system. Before this, the structure of the nervous system was pretty unclear. There were various theories that suggested the nervous system was made up of continuous networks rather than individual cells. The idea that the nervous system was made of cells instead was a huge breaking point for scientists including Golgi and Santiago Ramon y Cajal. It changed how they viewed the brain. Cellular theory was essential for understanding that neurons were the building blocks of the nervous system. This concept was a HUGE stepping stone in the early stages and throughout Golgi's work.

Source:

https://www.britannica.com/science/cell-theory

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cell theory, fundamental scientific theory of biology according to which cells are held to be the basic units of all living tissues. First proposed by German scientists Theodor Schwann and Matthias Jakob Schleiden in 1838, the theory that all plants and animals are made up of cells marked a great conceptual advance in biology and resulted in renewed attention to the living processes that go on in cells.

The history of cell theory is a history of the actual observation of cells, because early prediction and speculation about the nature of the cell were generally unsuccessful. The decisive event that allowed the observation of cells was the invention of the microscope in the 16th century, after which interest in the "invisible" world was stimulated. English physicist Robert Hooke, who described cork and other plant tissues in 1665, introduced the term cell because the cellulose walls of dead cork cells reminded him of the blocks of cells occupied by monks. Even after the publication in 1672 of excellent pictures of plant tissues, no significance was attached to the contents within the cell walls. The magnifying powers of the microscope and the inadequacy of techniques for preparing cells for observation precluded a study of the intimate details of the cell contents. The inspired Dutch microscopist Antonie van Leeuwenhoek, beginning in 1673, discovered blood cells, spermatozoa, and a lively world of "animalcules." A new world of unicellular organisms was opened up. Such discoveries extended the known variety of living things but did not bring insight into their basic uniformity. Moreover, when Leeuwenhoek observed the swarming of his animalcules but failed to observe their division, he could reinforce only the idea that they arose spontaneously.

Cell theory was not formulated for nearly 200 years after the introduction of microscopy. Explanations for this delay range from the poor quality of the microscopes to the persistence of ancient ideas concerning the definition of a fundamental living unit. Many observations of cells were made, but apparently none of the observers was able to assert forcefully that cells are the units of biological structure and function. Three critical discoveries made during the 1830s, when improved microscopes with suitable lenses, higher powers of magnification without aberration, and more satisfactory illumination became available, were decisive events in the early development of cell theory. First, the nucleus was observed by Scottish botanist Robert Brown in 1833 as a constant component of plant cells. Next, nuclei were also observed and recognized as such in some animal cells. Finally, a living substance called protoplasm was recognized within cells, its vitality made evident by its active streaming, or flowing, movements, especially in plant cells. After these three discoveries, cells, previously considered as mere pores in plant tissue, could no longer be thought of as empty, because they contained living material.

It was not until 1838 that the botanist Matthias Jakob Schleiden, interested in plant anatomy, stated that "the lower plants all consist of one cell, while the higher ones are composed of (many) individual cells." When the physiologist Theodor Schwann, Schleiden's friend, extended the cellular theory to include animals, he thereby brought about a rapprochement between botany and zoology. The two scientists clearly stated in 1839 that cells are the "elementary particles of organisms" in both plants and animals and recognized that some organisms are unicellular and others multicellular. This statement was made in Schwann's Mikroskopische Untersuchungen über die Übereinstimmung in der Struktur und dem Wachstume der Tiere und Pflanzen (1839; Microscopical Researches into the Accordance in the Structure and Growth of Animals and Plants). Schleiden's contributions on plants were acknowledged by Schwann as the basis for his comparison of animal and plant structure.

Schleiden and Schwann's descriptive statements concerning the cellular basis of biologic structure are straightforward and acceptable to modern thought. They recognized the common features of cells to be the membrane, nucleus, and cell body and described them in comparisons of various animal and plant tissues. A statement by Schleiden pointed toward the future direction of cell studies:

Each cell leads a double life: an independent one, pertaining to its own development alone; and another incidental, insofar as it has become an integral part of a plant. It is, however, easy to perceive that the vital process of the individual cells must form the first, absolutely indispensable fundamental basis, both as regards vegetable physiology and comparative physiology in general.

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Cell theory | Definition, History, Importance, Scientists, First Proposed, & Facts

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Cell theory, fundamental scientific theory of biology according to which cells are held to be the basic units of all living tissues. First proposed by German scientists Theodor Schwann and Matthias Jakob Schleiden in 1838, the theory that all plants and animals are made up of cells marked a great

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Historical Entry #4: Golgi's Development of Silver Staining (1870s)

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Historical Entry #4: Golgi's Development of Silver Staining (1870s)

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In the 1870s, Camillo Golgi invented silver chromate method, which was a groundbreaking technique that involved staining nerve tissue. Golgi's method allowed scientists to see individual neurons very clearly for the first time by making their complex structures easier to see. The staining technique was so important because it enabled the visualization of neurons in their natural state rather than isolated. Golgi's technique became widely known in neuroanatomy research allowed scientists to get a more accurate idea of how the nervous system was actually constructed.

Source:

https://doi.org/10.3390/cells11244112

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An Important Step in Neuroscience: Camillo Golgi and His Discoveries

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Camillo Golgi (Figure 1) is one of the most prestigious personalities of modern medicine [...]

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Historical Entry #5: The Neuron Doctrine vs. Reticular Theory (20th Century)

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Historical Entry #5: The Neuron Doctrine vs. Reticular Theory (20th Century)

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Once the 20th century began, a major debate emerged between two neuroanatomist, Camillo Golgi and Santiago Ramon y Cajal. Golgi supported the reticular theory, which suggested that the nervous system was composed of a continuous network of fibers. While Cajal used Golgi's staining method, he concluded that the nervous system actually consisted of individual distinct cells called neurons. This debate was known as the neuron doctrine vs. the reticular theory. Although Golgi was awarded a Nobel Prize for his work, Cajal's findings proved to be more accurate. This dispute was such a significant moment in the history of neuroscience because it really solidified the idea that neurons were made up of individual cells.

Source:

https://en.wikipedia.org/wiki/Reticular_theory

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Historical Entry #6: Golgi's Nobel Prize (1906)

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Historical Entry #6: Golgi's Nobel Prize (1906)

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In 1906, Camillo Golgi was awarded the Nobel Prize in Physiology, along with Santiago Ramon y Cajal. This award was for their work in neuroanatomy. Golgi's silver staining method was a fundamental part in revealing the structure of the nervous system. Although Golgi believed in the reticular theory originally, his method helped Cajal illustrate the neuron doctrine. This prize recognized Golgi's innovative contributions to neuroscience. The Nobel Prize solidified Golgi's place as a key historical figure in neuroscience and showed the true importance of how his findings advanced research in the field of neuroscience for many years to come.

Source:

https://www.nobelprize.org/prizes/medicine/1906/summary/

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Thirteen laureates were awarded a Nobel Prize in 2021, for achievements that have conferred the greatest benefit to humankind.

Their work and discoveries range from the Earth's climate and our sense of touch to efforts to safeguard freedom of expression.

See them all presented here.

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The Nobel Prize in Physiology or Medicine 1906

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Thirteen laureates were awarded a Nobel Prize in 2021, for achievements that have conferred the greatest benefit to humankind. Their work and discoveries range from the Earth's climate and our sense of touch to efforts to safeguard freedom of expression. See them all presented here.

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Historical Entry #7: Early Contributions of Santiago Ramon y Cajal

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Historical Entry #7: Early Contributions of Santiago Ramon y Cajal

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Santiago Ramon y Cajal used Golgi's silver staining method to conduct research into the structure of the nervous system. Cajal came up with tons of detailed drawings of neurons and showed their individuality and how they were complexly connected. This helped him refute the reticular theory that was backed by Golgi. His work showed that neurons were separate entities that communicated through synapses. This was an important step for the future of modern neuroanatomy. Cajal's contributions were important for shaping the neuron doctrine. He challenged Golgi's views and also advanced the understanding of the nervous system at the same time.

Source:

https://www.nobelprize.org/prizes/medicine/1906/cajal/facts/

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Santiago Ramón y Cajal
The Nobel Prize in Physiology or Medicine 1906

Born: 1 May 1852, Petilla de Aragón, Spain

Died: 17 October 1934, Madrid, Spain

Affiliation at the time of the award: Madrid University, Madrid, Spain

Prize motivation: "in recognition of their work on the structure of the nervous system."

Prize share: 1/2

Life

Santiago Ramón y Cajal was born in Petilla de Aragón in Navarra, Spain, as the son of a doctor. Though he was talented artistically, he chose to study medicine at the University of Zaragoza, where his father lectured in Applied Anatomy. After serving as an army medic, Cajal received his PhD in Madrid in 1877 and became a professor of anatomy at the University of Valencia in 1883. He also held professorships at universities in Barcelona and Madrid as well as serving as Director of the Zaragoza Museum. Santiago Ramón y Cajal was married with seven children.

Work

Our bodies are controlled by the interaction between the brain and the nervous system, which extends throughout the body. Camillo Golgi's discovery in the 1870s that nerve cells could be colored using silver nitrate opened up new opportunities for their study. Santiago Ramón y Cajal began using this method in 1887 and achieved many groundbreaking results in the years that followed. This included proving that each nerve cell is an independent entity and nerve synapses transfer nerve impulses from one cell to another.

To cite this section
MLA style: Santiago Ramón y Cajal - Facts. NobelPrize.org. Nobel Prize Outreach AB 2022. Wed. 20 Apr 2022. <https://www.nobelprize.org/prizes/medicine/1906/cajal/facts/>

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The Nobel Prize in Physiology or Medicine 1906

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Santiago Ramón y Cajal The Nobel Prize in Physiology or Medicine 1906 Born: 1 May 1852, Petilla de Aragón, Spain Died: 17 October 1934, Madrid, Spain Affiliation at the time of the award: Madrid University, Madrid, Spain Prize motivation: "in recognition of their work on the structure of the nervous system."

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Historical Entry #8: The Development of Modern Microscopy (1800s)

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Historical Entry #8: The Development of Modern Microscopy (1800s)

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In the 1800s, there were advancements in microscopy that played an important role in the study of cells and tissues, including the nervous system. Early microscope technology enabled scientists to visualize the intricate details in each and every individual cell. Golgi's silver staining technique, along with advancements in optical technology, allowed him to reveal the complex structures that each neuron has. These thoughts led to more modern imaging techniques that can now be seen in modern neuroscience. The combination of enhanced microscopy technology gave us more insight on how complex the nervous system actually can be.

Sources:

https://www.alzforum.org/news/research-news/modern-microscopy-plumbs-depths-brain-tissue

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Botcherby EJ, Juskaitis R, Booth MJ, Wilson T

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Modern Microscopy Plumbs the Depths of Brain Tissue | ALZFORUM

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Historical Entry #9: The Path to Golgi's Silver Staining

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Historical Entry #9: The Path to Golgi's Silver Staining

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Before Camillo Golgi's development of the silver staining technique, scientists struggled to visualize nerve cells in their natural setting. Earlier staining methods, such as basic dyes, could not differentiate between individual neurons or their own complex structures. Golgi's work with the silver staining chromate method was a powerful tool for neuroanatomists to use. His technique allowed for better visualization and distinction in their structures. This technology is still an essential tool that is used for studying the nervous system today.

Sources:

https://doi.org/10.1007/s00401-007-0200-2

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Silver diagnosis in neuropathology: principles, practice and revised interpretation - Acta Neuropathologica

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Silver-staining methods are helpful for histological identification of pathological deposits. In spite of some ambiguities regarding their mechanism and interpretation, they are widely used for histopathological diagnosis. In this review, four major silver-staining methods, modified Bielschowsky, Bodian, Gallyas (GAL) and Campbell–Switzer (CS) methods, are outlined with respect to their principles, basic protocols and interpretations, thereby providing neuropathologists, technicians and neuroscientists with a common basis for comparing findings and identifying the issues that still need to be clarified. Some consider “argyrophilia” to be a homogeneous phenomenon irrespective of the lesion and the method. Thus, they seek to explain the differences among the methods by pointing to their different sensitivities in detecting lesions (quantitative difference). Comparative studies, however, have demonstrated that argyrophilia is heterogeneous and dependent not only on the method but also on the lesion (qualitative difference). Each staining method has its own lesion-dependent specificity and, within this specificity, its own sensitivity. This “method- and lesion-dependent” nature of argyrophilia enables operational sorting of disease-specific lesions based on their silver-staining profiles, which may potentially represent some disease-specific aspects. Furthermore, comparisons between immunohistochemical and biochemical data have revealed an empirical correlation between GAL+/CS-deposits and 4-repeat (4R) tau (corticobasal degeneration, progressive supranuclear palsy and argyrophilic grains) and its complementary reversal between GAL-/CS+deposits and 3-repeat (3R) tau (Pick bodies). Deposits containing both 3R and 4R tau (neurofibrillary tangles of Alzheimer type) are GAL+/CS+. Although no molecular explanations, other than these empiric correlations, are currently available, these distinctive features, especially when combined with immunohistochemistry, are useful because silver-staining methods and immunoreactions are complementary to each other.

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Historical Entry #10: The Impact of Golgi's Work on Neuroanatomy

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Historical Entry #10: The Impact of Golgi's Work on Neuroanatomy

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Golgi's contributions to neuroscience continue to resonate with the modern community of researchers. His staining method provided a clear and detailed way to look at images of neurons. This was revolutionary in the field of neuroscience and set down a good sturdy foundation for future discoveries. Today, modern neuroanatomists use different variations of Golgi's staining technique to study brain structure and function. Golgi's influence extends beyond that as well. His legacy is going to carry on for many, many years to come. A lot of modern groundbreaking work and technologies all stem from the basis of his work.

Source:

https://neuroanatomy.ca

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Contemporary Entry: Golgi's Legacy Applied to Modern Neuroscience

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Contemporary Entry: Golgi's Legacy Applied to Modern Neuroscience

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The legacy of Golgi's staining technique is evident in today's contemporary neuroscience. Researchers still use advanced versions of his staining method to visualize the intricacies of neurons and their connections to each other. Techniques such as Golgi staining are used in research on neurodegenerative diseases, brain mapping and synaptic connectivity. Recent advancements in imaging technologies and microscopy have enhanced and strung Golgi's technique further along. This allows for even more precise studies of the brain and its complex structure.

Source:

https://doi.org/10.4103/1673-5374.156950

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Baloyannis, Stavros J. M.D., Ph.D.,* Author Information

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Staining neurons with Golgi techniques in degenerative... : Neural Regeneration Research

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Individual Entry: The Life and Legacy of Camillo Golgi (1843-1926)

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Individual Entry: The Life and Legacy of Camillo Golgi (1843-1926)

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Camillo Golgi (1843-1926) was an Italian neuroscientist who revolutionized the field of neuroanatomy with his development of silver staining chromate method. He was born in a small village in northern Italy. He studied medicine and became a professor in histology. His work on the nervous system was so important because it allowed for clearer visualization of individual neurons. Golgi's silver staining technique became a very essential tool in neuroanatomy despite his support of the reticular theory. His legacy is reflected everyday in the work of modern neuroscientists. His professional life was dedicated to scientific discovery in neuroscience and a passion for developing the understanding of the nervous system for the greater good.

Source:

https://en.wikipedia.org/wiki/Camillo_Golgi

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Camillo Golgi ( Italian: [kaˈmillo ˈɡɔldʒi]; 7 July 1843 - 21 January 1926) was an Italian biologist and pathologist known for his works on the central nervous system. He studied medicine at the University of Pavia (where he later spent most of his professional career) between 1860 and 1868 under the tutelage of Cesare Lombroso. Inspired by pathologist Giulio Bizzozero, he pursued research in the nervous system. His discovery of a staining technique called black reaction (sometimes called Golgi's method or Golgi's staining in his honour) in 1873 was a major breakthrough in neuroscience. Several structures and phenomena in anatomy and physiology are named for him, including the Golgi apparatus, the Golgi tendon organ and the Golgi tendon reflex.

Golgi and the Spanish biologist Santiago Ramón y Cajal were jointly given the Nobel Prize in Physiology or Medicine 1906 "in recognition of their work on the structure of the nervous system".

Camillo Golgi was born on 7 July 1843 in the village of Corteno near Brescia, in the province of Brescia ( Lombardy), at the time Kingdom of Lombardy-Venetia, today Italy. The village is now named Corteno Golgi in his honour. His father Alessandro Golgi was a physician and district medical officer, originally from Pavia. In 1860, he entered the University of Pavia to study medicine, and earned his medical degree in 1865. He did an internship at the San Matteo Hospital (now IRCCS Policlinico San Matteo Foundation). During his internship he briefly worked as a civil physician in the Italian Army, and as assistant surgeon at the Novara Hospital (now Azienda Ospedaliero Universitaria Maggiore della Carità di Novara). At the same time he was also involved in the medical team for investigating cholera epidemic in villages around Pavia.

In 1867, he resumed his academic study under the supervision of Cesare Lombroso. Lombroso was a renowned scientist in medical psychology such as genius, madness and criminality. Inspired by Lombroso, Golgi wrote a thesis on the etiology of mental disorders, from which he obtained his M.D. in 1868. He became more interested in experimental medicine, and started attending the Institute of General Pathology headed by Giulio Bizzozero. Three years his junior, Bizzozero was an eloquent teacher and experimenter, who specialised in histology of the nervous system and the properties of bone marrow. The most important research publications of Golgi were directly or indirectly influenced by Bizzozero. The two became so close that they lived in the same building; and Golgi later married Bizzozero's niece, Lina Aletti. By 1872, Golgi was an established clinician and histopathologist. He, however, had no opportunity as a tenured professor in Pavia to pursue teaching and research in neurology.

Financial pressure prompted him to join the Hospital of the Chronically Ill (Pio Luogo degli Incurabili) in Abbiategrasso, near Milan, as Chief Medical Officer in 1872. To continue research, he set up a simple laboratory on his own in a refurbished hospital kitchen, and it was there that he started making his most notable discoveries. His major achievement was the development of staining technique for nerve tissue called the black reaction (later the Golgi's method). He published his major works between 1875 and 1885 in the journal Rivista sperimentale di Freniatria e di medicina legale. In 1875, he joined the faculty of histology at the University of Pavia. In 1879, he was appointed Chair of Anatomy at the University of Siena. But the next year, he returned to the University of Pavia as full Professor of histology. From 1879 he also became Professor of General Pathology as well as Honorary Chief ( Primario ad honorarem) at the San Matteo Hospital. He served as Rector of the University of Pavia twice, first between 1893 and 1896, and second between 1901 and 1909. During the First World War (1914-1917), he directed the military hospital Collegio Borrmeo at Pavia. He retired in 1918 and continued to research in his private laboratory till 1923. He died on 21 January 1926.

Golgi and his wife Lina Aletti had no children, and they adopted Golgi's niece Carolina.

Golgi was irreligious in his later life and became an agnostic atheist. One of his former students attempted an unsuccessful deathbed conversion on him.

Black reaction or Golgi's staining

The Central nervous system was difficult to study during Golgi's time because the cells were hard to identify. The available tissue staining techniques were useless for studying nervous tissue. While working as chief medical officer at the Hospital of the Chronically Ill, he experimented with metal impregnation of nervous tissue, using mainly silver ( silver staining). In early 1873, he discovered a method of staining nervous tissue that would stain a limited number of cells at random in their entirety. He first treated the tissue with potassium dichromate to harden it, and then with silver nitrate. Under the microscope, the outline of the neuron became distinct from the surrounding tissue and cells. The silver chromate precipitate, as a reaction product, selectively stains only some cellular components randomly, sparing other cell parts. The silver chromate particles create a stark black deposit on the soma (nerve cell body) as well as on the axon and all dendrites, providing an exceedingly clear and well-contrasted picture of neuron against a yellow background. This makes it easier to trace the structure of the nerve cells in the brain for the first time. Since cells are selective stained in black, he called the process la reazione nera ("the black reaction"), but today it is called Golgi's method or the Golgi stain. On 16 February 1873, he wrote to his friend Niccolò Manfredi:

I am delighted that I have found a new reaction to demonstrate, even to the blind, the structure of the interstitial stroma of the cerebral cortex.

His discovery was published in the Gazzeta Medica Italiani on 2 August 1873.

In 1871, a German anatomist Joseph von Gerlach postulated that the brain is a complex "protoplasmic network", in the form of a continuous network called the reticulum. Using his black reaction, Golgi could trace various regions of the cerebro-spinal axis, clearly distinguishing the different nervous projections, namely axon from the dendrites. He drew up a new classification of cells on the basis of the structure of their nervous prolongation. He described an extremely dense and intricate network, composed of a web of intertwined branches of axons coming from different cell layers ("diffuse nervous network"). This network structure, which emerges from the axons, is essentially different from that hypothesized by Gerlach. It was the main organ of the central nervous system according to Golgi. Thus, Golgi presented the reticular theory which states that the brain is a single network of nerve fibres, and not of discrete cells. Although Golgi's earlier works between 1873 and 1885 clearly depicted the axonal connections of cerebellar cortex and olfactory bulb as independent of one another, his later works including the Nobel Lecture showed the entire granular layer of the cerebellar cortex occupied by a network of branching and anastomosing nerve processes. This was due to his strong conviction in the reticular theory. Golgi's theory was challenged by Ramón y Cajal, who used the same technique developed by Golgi. According to Ramón y Cajal's neurone theory, the nervous system is but a collection of individual cells, the neurones, which are interconnected to form a network.

In addition to this, Golgi was the first to give clear descriptions of the structure of the cerebellum, hippocampus, spinal cord, olfactory lobe, as well as striatal and cortical lesions in a case of chorea. In 1878, he also discovered a receptor organ that senses changes in muscle tension, and is now known as Golgi tendon organ or Golgi receptor; and Golgi-Mazzoni corpuscles (pressure transductors). He further developed a stain specific for myelin (a specialised membrane which wraps around the axon) using potassium dichromate and mercuric chloride. Using this he discovered the myelin annular apparatus, often called the horny funnel of Golgi-Rezzonico.

Golgi studied kidney function during 1882 to 1889. In 1882, he published his observations on the mechanism of renal hypertrophy, which he understood to be due to renal cell proliferation. In 1884, he described tubular cell mitoses in the kidney of a person suffering from tubulointerstitial nephritis, and he noted that the process was an essential part of repairing the kidney tissue. He was the first to dissect out intact nephrons, and show that the distal tubulus ( loop of Henle) of the nephron returns to its originating glomerulus, a finding that he published in 1889 ("Annotazioni intorno all'Istologia dei reni dell'uomo e di altri mammifieri e sull'istogenesi dei canalicoli oriniferi". Rendiconti R. Acad. Lincei 5: 545-557, 1889).

A French Army physician Charles Louis Alphonse Laveran discovered that malaria was caused by microscopic parasite (now called Plasmodium falciparum) in 1880. But scientists were sceptical until Golgi intervened. It was Golgi who helped him prove that malarial parasite was a microscopic protozoan. From 1885, Golgi studied the malarial parasite and its transmission. He established two types of malaria, tertian and quartan fevers caused by Plasmodium vivax and Plasmodium malariae respectively. In 1886, he discovered that malarial fever ( paroxysm) was produced by the asexual stage in the human blood (called erythocytic cycle, or Golgi cycle). In 1889-1890, Golgi and Ettore Marchiafava described the differences between benign tertian malaria and malignant tertian malaria (the latter caused by P. falciparum). By 1898, along with Giovanni Battista Grassi, Amico Bignami, Giuseppe Bastianelli, Angelo Celli and Marchiafava, he confirmed that malaria was transmitted by Anopheline mosquito.

An organelle in eukaryotice cells now known as Golgi apparatus or Golgi complex, or sometimes simply as Golgi, was discovered by Camillo Golgi. Golgi modified his black reaction using osmium dichromate solution with which he stained the nerve cells ( Purkinje cells) of the cerebellum of an owl. He noticed thread-like networks inside the cells and named them apparato reticolare interno (internal reticular apparatus). Recognising them to be unique cellular components, he presented his discovery before the Medical-Surgical Society of Pavia in April 1898. After the same was confirmed by his assistant Emilio Veratti, he published it in the Bollettino della Società medico-chirurgica di Pavia. However, most scientists disputed his discovery as nothing but a staining artefact. Their microscopes were not powerful enough to identify the organelles. By the 1930s, Golgi's description was largely rejected. It was only firmly established 50 years after its discovery, when electron microscopes were developed.

Golgi, together with Santiago Ramón y Cajal, received the Nobel Prize in Physiology or Medicine in 1906 for his studies of the structure of the nervous system. In 1900 he was named senator by King Umberto I. In 1913 he became foreign member of the Royal Netherlands Academy of Arts and Sciences. He received honorary doctorates from the University of Cambridge, University of Geneva, Kristiania University College, National and Kapodistrian University of Athens, and Paris-Sorbonne University. In 1994, the European Community commemorated him with postage stamps.

Monuments in Pavia

In Pavia several landmarks stand as Golgi's memory.

A marble statue, in a yard of the old buildings of the University of Pavia, at N.65 of the central "Strada Nuova". On the basement, there is the following inscription in Italian language: "Camillo Golgi / patologo sommo / della scienza istologica / antesignano e maestro / la segreta struttura / del tessuto nervoso / con intenta vigilia / sorprese e descrisse / qui operò / qui vive / guida e luce ai venturi / MDCCCXLIII - MCMXXVI" (Camillo Golgi / outstanding pathologist / of histological science / precursor and master / the secret structure / of the nervous tissue / with strenuous effort / discovered and described / here he worked / here he lives / here he guides and enlightens future scholars / 1843 - 1926).

"Golgi's home", also in Strada Nuova, at N.77, a few hundreds meters away from the University, just in front to the historical "Teatro Fraschini". It is the home in which Golgi spent the most of his family life, with his wife Lina.

Golgi's tomb is in the Monumental Cemetery of Pavia (viale San Giovannino), along the central lane, just before the big monument to the fallen of the First World War. It is a very simple granite grave, with a bronze medallion representing the scientist's profile. Near Golgi's tomb, apart from his wife, two other important Italian medical scientists are buried: Bartolomeo Panizza and Adelchi Negri.

Golgi's museum was created in 2012, in the ancient Palazzo Botta of the University of Pavia at N.10 of Piazza Antoniotto Botta reconstructs the study of Camillo Golgi and its laboratories with furniture and original instruments.

Mazzarello, Paolo (2010), Golgi: A Biography of the Founder of Modern Neuroscience, translated by Badiani, Aldo; Buchtel, Henry A., New York: Oxford University Press, ISBN 978-0-19-533784-6

Mironov, Alexander A.; Margit, Pavelka (2006). The Golgi Apparatus State of Art After 110 Years of Camillo's Discovery. Dordrecht: Springer. ISBN 3-211-76310-4.

Morré, D. James; Mollenhauer, Hilton H. (2009). The Golgi Apparatus: The First 100 Years. New York: Springer. ISBN 978-0-387-74347-9.

De Carlos, Juan A; Borrell, José (2007), "A historical reflection of the contributions of Cajal and Golgi to the foundations of neuroscience.", Brain Research Reviews (published August 2007), vol. 55, no. 1, pp. 8-16, doi:10.1016/j.brainresrev.2007.03.010, hdl:10261/62299, PMID 17490748, S2CID 7266966

Muscatello, Umberto (2007), "Golgi's contribution to medicine.", Brain Research Reviews (published August 2007), vol. 55, no. 1, pp. 3-7, doi:10.1016/j.brainresrev.2007.03.007, PMID 17462742, S2CID 41680914

Kruger, Lawrence (2007), "The sensory neuron and the triumph of Camillo Golgi", Brain Research Reviews (published October 2007), vol. 55, no. 2, pp. 406-10, doi:10.1016/j.brainresrev.2007.01.008, PMID 17408565, S2CID 32486297

Fabene, P F; Bentivoglio, M (1998), "1898-1998: Camillo Golgi and "the Golgi": one hundred years of terminological clones.", Brain Res. Bull. (published October 1998), vol. 47, no. 3, pp. 195-8, doi:10.1016/S0361-9230(98)00079-3, PMID 9865849

Mironov, A A; Komissarchik, Ia Iu; Mironov, A A; Snigirevskaia, E S; Luini, A (1998), "[Current concept of structure and function of the Golgi apparatus. On the 100-anniversary of the discovery by Camillo Golgi]", Tsitologiia, vol. 40, no. 6, pp. 483-96, PMID 9778732

Farquhar, M G; Palade, G E (1998), "The Golgi apparatus: 100 years of progress and controversy.", Trends Cell Biol. (published January 1998), vol. 8, no. 1, pp. 2-10, doi:10.1016/S0962-8924(97)01187-2, PMC , PMID 9695800

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Camillo Golgi ( Italian: [kaˈmillo ˈɡɔldʒi]; 7 July 1843 - 21 January 1926) was an Italian biologist and pathologist known for his works on the central nervous system. He studied medicine at the University of Pavia (where he later spent most of his professional career) between 1860 and 1868 under the tutelage of Cesare Lombroso.