Santiago Ramón y Cajal ForMemRS (Spanish: [sanˈtjaɣo raˈmon i kaˈxal]; 1 May 1852 – 18 October 1934) was a Spanish pathologist, histologist, neuroscientist and Nobel laureate. His original pioneering investigations of the microscopic structure of the brain have led him to be designated by many as the father of modern neuroscience. His medical artistry was legendary, and hundreds of his drawings illustrating the delicate arborizations of brain cells are still in use for educational and training purposes.
Ramón y Cajal's early work was accomplished at the Universities of Zaragoza and Valencia, where he focused on the pathology of inflammation, the microbiology of cholera, and the structure of Epithelial cells and tissues. It was not until he moved to the University of Barcelona in 1887 that he learned Golgi's silver nitrate preparation and turned his attention to the central nervous system. During this period he made extensive studies of neural material covering many species and most major regions of the brain.
Ramón y Cajal made several major contributions to neuroanatomy. He discovered the axonal growth cone, and experimentally demonstrated that the relationship between nerve cells was not continuous but contiguous. This provided definitive evidence for what would later be known as "neuron doctrine", now widely considered the foundation of modern neuroscience. In debating neural network theories (e.g. neuron theory, reticular theory), Ramón y Cajal was a fierce defender of the neuron theory.
He provided detailed descriptions of cell types associated with neural structures, and produced excellent depictions of structures and their connectivity.
He was an advocate of the existence of dendritic spines, although he did not recognize them as the site of contact from presynaptic cells. He was a proponent of polarization of nerve cell function and his student Rafael Lorente de Nó would continue this study of input/output systems into cable theory and some of the earliest circuit analysis of neural structures.
He discovered a new type of cell, to be named after him: the interstitial cell of Cajal (ICC). This cell is found interleaved among neurons embedded within the smooth muscles lining the gut, serving as the generator and pacemaker of the slow waves of contraction that move material along the gastrointestine, vitally mediating neurotransmission from motor nerves to smooth muscle cells.
In his 1894 Croonian Lecture, he suggested in an extended metaphor that cortical pyramidal cells may become more elaborate with time, as a tree grows and extends its branches. He also devoted a considerable amount of his time to studying hypnosis (which he used to help his wife with birth labor) and parapsychological phenomena, but a book he had written on these areas got lost during the Spanish Civil War.
Cajal received many prizes, distinctions and societal memberships along his scientific career including and honorary Doctorates in Medicine of the Universities of Cambridge and Würzburg and an honorary Doctorate in Philosophy of the Clark University. Nevertheless the most famous distinction he was awarded was the Nobel Prize in Physiology or Medicine in 1906 together with Italian Camillo Golgi "in recognition of their work on the structure of the nervous system". This was seen as quite controversial because Golgi, a stout reticularist, disagreed with Cajal in his view of the neuron doctrine.
The asteroid 117413 Ramonycajal is named in his honor. The Spanish public television filmed a biopic series to commemorate his life.