Physiology or Medicine, 2nd Place
Large Shoes to Fill: The Legacy of Camillo Golgi


Horace Mann School, The Bronx

Nobel laureate Camillo Golgi was an extraordinary scientist. A man of many talents, Golgi was an innovative researcher and an inspiring teacher and mentor. His breakthroughs changed the face of science forever. Across the planet, biology students are learning about his discoveries, and researchers are employing his concepts and techniques. As famed advertiser and art director George Lois once said, “Creativity can solve almost any problem.” Golgi exemplified this idea throughout his life and work.

Golgi was born in 1843 in Coteno, Italy, studied medicine in Pavia, and spent most of his career on the faculty of its university (Altamura, 1996). A man of brilliant new ideas and concepts, Golgi dedicated himself to the study of cellular biology; he wanted to discover more about the functions of the human body and the causes of disease (Newkirk). The Nobel Foundation honored Golgi’s ground-breaking research in 1906 by awarding him one half of the Nobel Prize for Medicine or Physiology. Today, nearly a century after Golgi became a Nobel laureate, his findings are used in laboratories and biology classrooms around the world; and his discoveries and techniques will be utilized in future research.

Golgi began his psychiatric and research career after receiving a degree in Medicine from the University of Pavia in 1865 (Newkirk). He soon began work at the Ospedale San Matteo Psychiatric Clinic under the direction of pre-eminent psychiatrist Cesare Lombroso, who inspired his neurological studies (Mazzarello, 1999). Working in collaboration with Lombroso, Golgi became one of the first scientists to investigate the etiology of mental and neurological illness from an anti-metaphysical point of view. During this period in history, psychiatry was still embedded in a medieval and nearly religious atmosphere. Golgi, in fact, was one of the first to form a link between neuroscience and psychiatry (Altamura, 1996). Convinced that psychiatric disease theory had to be supported by facts, Golgi began to concentrate on the experimental study of the structure of the nervous system (Bentivoglio).

In studying the nervous system, Golgi became a member of an elite class of scientists who not only develop new instruments, but construct new theories based on their inventions. Histological techniques, such as fixation procedures and tissue staining had been introduced in the middle of the 19th century. However, due to the complex and peculiar nature of the nervous system’s organization, these techniques were ill-suited for research (Bentivoglio). Golgi solved this problem in 1872 with his invention of the “black reaction,” a revolutionary new staining technique for nervous tissue. Now known as Golgi staining or Golgi impregnation, this technique involves hardening nervous tissue in potassium bichromate and impregnating it with silver nitrate. Golgi’s invention allowed, for the first time, a clear visualization of a nerve cell body with all its processes in its entirety (Bentivoglio). The technique was a ground-breaking discovery: once the staining method was used in other countries, it stirred a flurry of new hypotheses about the anatomical organization and eventually the function of the nervous system (Van Gijn, 2001). However, it was Golgi’s discovery of the black reaction and his subsequent investigations that provided a substantial contribution to the advancement of the knowledge on the structural organization of the nervous tissue (Bentivoglio).

Golgi made many important discoveries about the nervous system and contributed greatly to the modern knowledge of its structure. From Golgi-impregnated nervous tissue samples, Golgi described the morphological features of glial cells and the relationships between their cell processes and blood vessels. He also identified two fundamental types of nerve cells: “Golgi type I,” which are known today as “projection neurons,” and “Golgi type II,” which correspond to “interneurons” and “local circuit neurons” (Bentivoglio). Additionally, Golgi made advancements on intracellular nervous system structure using the black reaction, officially identifying the “Golgi internal reticular apparatus,” also known as the “Golgi complex.” The discovery of this cell organelle was a breakthrough in cytology and cell biology (Bentivoglio). It contributed to the foundation of modern neuroscience, as well, providing the spark to a truly scientific revolution that allowed the morphology and the basic architecture of the cerebral tissue to be displayed in all its complexity (Mazzarello, 1999). Although Golgi misinterpreted the overall view of the organization of the nervous system, theorizing that the nervous system is a syncytial system and that the nervous impulse propagates along a diffuse network, his rival and Co-Nobel prize recipient Ramon y Cajal correctly interpreted the structure of the nervous system by using Golgi impregnation techniques (Bentivoglio). Golgi’s work in neurology and psychiatry is clearly both advanced and awe-inspiring.

However, while he is probably best known for his work in the fields of neurology and psychiatry, Golgi made important discoveries in other areas of research as well. In addition to his contributions to general medicine with regard to intestinal worm infections and Bright’s disease of the kidney, Golgi also made advancements in studying malaria (Van Gijn, 2001). Golgi was able to determine the entire intraerythrocytic cycle of development of the malaria parasite Plasmodium; as a result, the cycle is now called the Golgi cycle. He also defined what is now known as the Golgi law, which defines the sequential relation between the intermittent feverish bout and the segmentation of the parasite (Mazzarello, 1999). Finally, Golgi studied kidney histology, histopathology, and histogenesis and discovered the important relationship between the vascular pole of the Malpighian glomerulus and the distal tubule, which plays an important role in the regulation of blood pressure (Mazzarello, 1999). Undoubtedly, Golgi’s contributions to the world of science were remarkable; his techniques and discoveries were ground-breaking in the scientific community. But, while Golgi’s Nobel Prize may be nearly a century old, his ideas are being built on, today.

The scientific community is still benefiting from Golgi’s stimulating teaching techniques and experimental discoveries. After being appointed to the chair of General Pathology at the University of Pavia in 1881, Golgi established a very active laboratory in the Institute of General Pathology (Bentivoglio). Golgi was an inspiring mentor: many of his students made notable discoveries while working in his lab. For example, in Golgi’s laboratory, Emilio Veratti became the first to describe the sarcoplasmic reticulum in skeletal muscle fibers, (Bentivoglio), and Aldo Perroncito described the phases of regeneration in the nerves. Another student, Adelchi Negri, discovered the intraneuronal inclusions, also known as the Negri bodies, in animals and humans infected with the rabies virus (Mazzarello, 1999). Many other distinguished scientists were inspired by Golgi while spending periods of study and specialization in his lab, including Giovanni Battista Grasse, the discoverer of the Anopheles that transmit human malaria, and Fritjof Nansen, the Norwegian zoologist and Nobel Peace Prize winner in 1922 (Mazzarello, 1999).

In addition to motivating his students to make significant new insights in research, Golgi left a legacy of his own experimental discoveries, which scientists have been using in their own research for the past century. Medical treatment for malaria has been developed, for example, based on his definitions of the Golgi cycle and the Golgi law that he developed for this disease. His discovery of the Golgi apparatus, additionally, led to a flurry of studies on the organelle’s function, thus contributing to the knowledge of intracellular structure. Due to Golgi’s discovery, scientists found that the Golgi apparatus plays a key role in the intracellular sorting, trafficking, and targeting or proteins (Bentivoglio). Additionally, as aforementioned, Golgi’s invention of the black reaction led to the correct interpretation of the structure of the nervous system, and scientists still use this technique today in research.

Golgi’s impregnation technique is still used by researchers today, and will be in the future. Golgi was one of the first scientists to connect the areas of neuroscience and psychiatry, believing that experimental findings should be the basis of theories surrounding psychiatric disorders. Today, the research fields of neuroscience and psychiatry are, and will continue to be intertwined because of Golgi’s contributions. There has been a great deal of speculation, for example, that signal transmission is the underlying abnormality in the neuropsychiatric disorder called schizophrenia (Arnold et al, 1991; Dean, 2000; Harrison, 1999; Kalus et al, 2000). In studying this disorder recently, researchers have been following in Golgi’s footsteps: using Golgi impregnation technique, neuroscientists are using experimental evidence to support theories about the disorder’s pathology. One theory currently held by neuroscientists is that the prefrontal cortex may be a site of pathological changes in schizophrenia (Beasley and Reynolds, 1997; Benes et al., 1986, 1991; Honer et al., 1997; Jones et al., 2002; Jones, 2004; Kalus et al, 2000; Pierri et al, 2001). To test this theory, scientists have been employing Golgi’s staining methods to derive experimental evidence. For example, Kalus et al, 2000 studied Golgi-impregnated prefrontal pyramidal neurons in schizophrenia patients and controls to investigate the differences in dendritic architecture. Additionally, Broadbelt et al, 2002 provided evidence for a decrease in basilar dendrites of pyramidal cells in schizophrenic medial prefrontal cortex by studying Golgi-stained material. These two studies are clear examples of current research being carried out using both Golgi’s techniques and concepts. However, there is a great deal still to learn about neuropsychiatric disorders like schizophrenia. Hence, in the future, the research community will continue to employ the discoveries and techniques left behind by Golgi.

Camillo Golgi was an extraordinary scientist. Clearly a man of great talent, Golgi was an innovative researcher and an inspiring mentor and teacher; his breakthroughs transformed the world of science. Around the world, Golgi’s legacy continues to live on in classrooms and research facilities. In fact, Golgi is currently the most frequently cited scientist in cell and molecular biology (Bentivoglio). As Golgi once said, “We, who have grown up in the faith of infinite progress, but who have been educated by experience and disciplined by scientific logic, look confidently upon you [students], on whom rests the task of consolidating and widening acquired knowledge, to participate, by spreading intellectual education with a mind free from political prejudice, in transforming society,” (Mazzarello, 1999). The students of both today and tomorrow can only hope that they will be able to transform society in the way that Golgi did; as a scientist, he left large shoes to fill.

References
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