Vernon Benjamin Mountcastle, Jr
“The Father of Neuroscience”
It is a sad day today as I learned about the passing away of Vernon Mountcastle last Sunday. Most of my research in regards to how the cerebral cortex works, its modular organization and how it is involved in different psychiatric conditions, is based on Vernon’s groundbreaking research (for its relevance to autism see: http://bit.ly/1B5vvdY). He was my mentor, friend and confidant.
Mountcastle was born in Shelbyville, Kentucky but soon moved to Roanoke, Virginia where he attended primary and secondary school. He was a precocious learner, skipping several grades during his early education. He earned his B.S. degree in Chemistry from Roanoke College in 3 years with honors. He was accepted to study medicine at Johns Hopkins and interned in Surgery before serving in the U.S. Naval Amphibious Forces for 3 years. During this time he participated in campaigns in Anzio, Italy and Normandy, France. After the war ended he married his wife to be for seven decades, Nancy Clayton and had three children.
After his military duties were over in 1946, he joined the Department of Physiology at Johns Hopkins as a postdoctoral fellow under the supervision of Elwood Henneman. He worked alongside Stephen Kuffler and some of his fellows (Hubel and Wiesel), Jerzy Rose, and others. Mountcastle would eventually rise through the academic ranks to become a full professor and Director of the Department of Physiology for seventeen years.
Together with Henneman he mapped the tactile representation of the body as a “figurine” in the ventrolateral thalamus of cats and monkeys. The close collaboration was brought to an end when Henneman departed in 1955 to establish a department of Neurobiology at Harvard University. His initial studies set the stage for Mountcastle’s discovery of the functional cortical columns in the somatosensory cortex of cats and monkeys. His studies were based on recordings from over twenty-three hundred neurons for which, in many cases, he collected the latency and spiking frequencies. These responses were analyzed in terms of their receptive field and modality, i.e., the nature of the external stimuli.
A funny story in regard to Mountcastle’s awe-impressive cumulus of work was provided by Hubel in his Nobel lecture of 1981:
“…we [Hubel and Wiesel] had gone to a lecture by Vernon (this was a few years after his discovery of cortical columns in which he had amazed us by reporting on the results of recording from some 900 somatosensory cortical cells, for those days an astronomical number. We knew we could never catch up, so we catapulted ourselves to respectability by calling our first cell No. 2000 and numbering subsequent ones from there. When Vernon visited our circus tent we were in the middle of a S-unit recording, cell Nos. 3007, 3008, and 3009. We made sure that we mentioned their identification numbers. All three cells had the same receptive-field orientation but neither Vernon nor we realized, then want that implied”.
Mountcastle’s view on the columnar organization of the cortex went far beyond its putative role in regards to information processing. In fact the columnar research led him to describe several physiological principles of cortical function. Mountcastle was the first person to describe how the neurons within each column responded to one specific submodality and dermatomal locale. The columns extended throughout the cortex from layers II through VI and each cell responded with approximately the same latency to stimulation, at least early in the response. Within proprioceptive columns, Mountcastle described how cells in adjacent columns exhibited activation to alternating flexion and extension of joints. Because activity in one cell led to the inactivation of the cell in the opposing column, Mountcastle inferred the mechanism of reciprocal lateral inhibition. Finally, Mountcastle described a pattern of surround inhibition where stimulation of areas in the peripheral receptive field actually inhibited cell activity. (Note: I have proposed a deficit of surround inhibition in autism and supporting evidence has been derived from EEG and flutter stimuli studies).
All of Mountcastle’s discoveries were done in a field that was antagonistic to his way of thinking and provided mounting criticism. Among his 2 publications in 1957 the columnarity principle was elaborated upon in the one having Mountcastle as the sole author. This was done at the request of his coauthors Drs. Davis and Berman, both of whom believed that columnarity was a radical departure from accepted tenets! In fact, Mountcastle was subjected to a great deal of criticism, even by his good friend Jerzy Rose. Back then the perspective of classical anatomists permeated neurosciences and cytoarchitecture was primarily about laminae, not columns.
Vernon was a towering figure within the field of neuroscience. Upon winning his Nobel Prize David Hubel placed Vernon’s achievements in perspective: “[To] Vernon Mountcastle, whose discovery of columns in the somatosensory cortex was surely the single most important contribution to the understanding of the cerebral cortex since Cajal”. Nobody now disputes his findings.
Mountcastle retired from academic duties in 2005 at the age of 87 years. During his long career he won many accolades including the Schmitt Prize and Medal from MIT, the U.S. National Medal of Science, and the Fyssen Foundation Prize from France. In 1978 he was awarded the Louisa Gross Horwits Prize from Columbia University and in 1883 the Albert Lasker Award for Basic Medical Research. He held 6 honorary degrees. He was a member of the Royal Society, Academie des Sciences, and the National Academy of Sciences. From the latter he received the United States National Medal of Science in 1986 for his lifetime of work in the neurosciences.
Mountcastle’s had forty eight postdoctoral fellows during his career. Some of the ones known to me include in alphabetical order: Carlos Acuna, Richard Andersen, Sven Anderson, Pradep Atluri, Frank Baker, Alvin Berman, James Campbell, Giancarlo Carii, Mirko Carreras, Ian Darian-Smith, John Downer, Charles Duffy, Robert Dykes, Solomon Erulkar, Apostolos Georgopoulos, Edward Glaser, Gundez Gucer, Thomas Harrington, Clinton Harrison, Juhani Hyvarinen, Kenneth Johnson, Cecil Kidd, Hans Kornhuber, Robert LaMotte, James Lane, Randall Long, James Lynch, Michael Merzenich, Mark Molliver, Brad Motter, Hiroshi Nakahama, Edwardo Oswaldo-Cruz, Edward Perl (his first fellow in 1950), Gian Poggio, Thomas Powell, Barbara Renkin, Rodolfo Romo, Sten Skoglund, Michael Steinmetz, Tadaaki Sumi, William Talbot, James Taylor, and Thomas Yin.
Despite all of these awards Mountcastle remained a humble individual deferential to his modest upbringing in Virginia. We kept in touch during the last few decades. He once told me that he used to enjoy horseback riding until age 82 years. “Falls now hurt, and bones do not heal quickly”, he said. My biggest achievement was when I wrote a book on cortical modularity and dedicated the same to him. Vernon said he had bought several copies and distributed them to members of his family.
It seems fitting to conclude this blog on Vernon Mountcastle with a paragraph from my introduction to Neocortical Modularity and the Cell Minicolumn: “Mountcastle is an outsized character that marched to his own beat. Never the person he “ought” to be, he was never boxed into preconceived ideas. This character trait was of immense help throughout his research career: Having no shoulders to stand on, he set his own goals and built upon his strengths. He aspired to an objective truth, the Holy Grail of neurosciences. In his research he overcame narrowness of scientific vision and became an heir to Socrates. While others went for easily accessible and exploitable prizes, Mountcastle claimed no glory, no acclaim. Personally he treated others by the categorical imperative: with respect and dignity as ends themselves. He is a unique individual: We can’t exchange Mountcastle for someone else and have an equal”.