Caveat lector: This blog is where I try out new ideas. I will often be wrong, but that's the point.

Home | Personal | Entertainment | Professional | Publications | Blog

Search Archive


Santiago Ramón y Cajal

Santiago Ramón y Cajal

So this has turned out to be neuroscientific biography week for me, what with two whole biographic pieces now! But after my last post about Hans Berger I just couldn't pass up writing about my favorite neuroscientific forebear, Santiago Ramón y Cajal.

While I had come across Cajal's name prior to graduate school, I didn't really become familiar with him until the beginning of the second year of my PhD during the fall of 2005. That was the first semester where I was supposed to work as TA (or, as Berkeley refers to them, Graduate Student Instructors (GSIs)). Because I have a very strong love for neuroanatomy (after all, neuroanatomy provides the rules that constrain our brains), I opted to teach the laboratory section of MCB 163: Mammalian Neuroanatomy. This was an undergraduate course through the Department of Molecular and Cellular Biology—a field in which I had never even taken a class prior to teaching that lab—and was known as one of the most difficult courses available.

The Professor for the course, the late Jeffrey Winer, was an excellent teacher; we spent many hours together with the other GSI preparing for the upcoming lab sections (ultimately I ended up receiving the outstanding GSI award for my work in this course, which I accredit to two phenomena: Dr. Winer's fantastic guidance and the increased respect I got from my students when they found out that the reason I had ditched my first week of teaching was so that I could go to Burning Man...)

Anyway, over the course of the semester Dr. Winer and I had many interesting conversations that quickly lead him to introduce me to the work of Cajal. Dr. Winer lent me one of Cajal's autobiographies, Recollections of My Life (Recuerdos de Mi Vida). Although I delayed in reading it for several months, I was eventually drawn in (so much so that, when I eventually created the neuroscience wiki for the Berkeley PhD students, I dubbed the site "Cajal"). I later went on to read Cajal's Advice for a Young Investigator, which has advice that I believe is still relevant for any modern researcher.

Mind you, Cajal was an unrepentant misogynist and Spanish nationalist. There were many times where I would read passages out loud to my wife in amusement and astonishment. I think I was most amazed by how rarely he spoke of his wife and five children in his autobiography; just a few pages here and there.

But damn if the guy wasn't interesting and talented. As a child Cajal was kicked out of most of his schools. He was sent off to various academies and he would often sneak away. When he was 11 years old he was thrown in jail for a night because he destroyed his town's gate. He did this by building his own cannon and blowing it up.

As an adult he was drafted into the Spanish army and served as a military surgeon in Cuba where he contracted malaria and TB. He was so severely ill that he was sent back to Spain where he eventually resumed his research. His work as a neuroanatomist involved many hours spent in front of a microscope examining different types of neurons and the patterns they form. Cajal is most famous for the work that ultimately won him the 1906 Nobel Prize: his work proving the "neuron doctrine". He shared this prize with his intellectual opponent Camillo Golgi.

Golgi received the prize for discovering what is now known as Golgi's method, a technique for staining neurons, their axons, and dendrites, rendering them visible under the microscope and available for study. This method stains approximately 5-10% of the actual neurons that are prepared, though it works through a mechanism that is still unknown but widely used. This method is the one that Cajal used and improved upon to show that neurons are not actually contiguous, but have spaces (synapses) between them. This is the neuron doctrine.

What's amusing is that Golgi still believed that neurons were contiguous and all physically connected (known as reticular theory now, but previously as link theory, among other names) and said as much in his acceptance speech. Golgi gave his speech first, then Cajal went up to give his. In his Nobel acceptance speech Cajal took several jabs at Golgi's theory, making for what was apparently quite an amusing lecture. For example, he says:

...[l]ike many scientific errors professed in good faith by distinguished scientists the link theory is the result of two conditions: one subjective, and the other objective. The first is the regrettable but inevitable tendency of certain impatient minds, to reject the use of elective methods, such as those of Golgi and of Ehrlich which do not lend themselves easily to improvisation...

He flair for the intellectual smack-down aside, Cajal had quite a way with words. Another example from his Nobel speech: "[u]nfortunately, nature seems unaware of our intellectual need for convenience and unity, and very often takes delight in complication and diversity."

Santiago Ramón y Cajal retinaAs I said, the Golgi method for staining neurons is still used today (among many other methods, of course). What's striking to me is how accurate and intricate Cajal's drawings were. The example at the left is a drawing he made of the chick retina. In modern neuroanatomy such drawings are captured digitally or the images are projected onto the table next to the microscope for manual tracing. In Cajal's day such methods weren't available. In order to create such accurate, intricate representations of the neuronal fields, Cajal would stare into the microscope for long stretches of time. Once he felt he had the images well memorized he would head down to a local cafe where he would drink absinthe while drawing the cells from memory. His reproductions were so accurate that they are hardly discernible from the images used in modern texts (aside from their artistic superiority).

I suppose my reasons for liking Cajal are many-fold. Misogyny and nationalism aside, the man expertly melded art and science, both in his drawings and in his writings. In reading over older scientific manuscripts I can't help but want a little more of a narrative in modern scientific writing. The dry, technical, academic style that pervades the peer-review system hides the actual process of science: one which is often messy, difficult, and frustrating, but ultimately a wonderfully rewarding experience. I think that adding back a little bit of that narrative focus on the process would benefit not only the readability of the science, but perhaps even engage the public more as well.


  1. Michele Insanally14:36

    Golgi made a very important observation that has fueled research endeavors for over 100 years now. He described and characterized what we now call the "perineuronal net." These are net-like like structures comprised of extracellular matrix molecules and proteins that surround the soma and dendrites of many neurons and are contiguous with the more diffuse extracellular matrix. Cajal erroneously thought these nets were artefacts of capricious staining techniques. Unfortunately, Cajal's opinion ended the first surge of interest in these structures. Later, with the advancement of cellular and molecular techniques scientists would go on to discover that perineuronal nets play an important role in regulating plasticity. Here's a link to an exciting, high-impact paper involving these nets (Pizzorusso et al., Science, 2002):


  2. Michele:

    That's super cool! I didn't know about that. I wasn't trying to downplay Golgi at all. I just think that Cajal's views on him were fascinating.

    And apparently scientifically unfortunate...

  3. And while Cajal’s neuron doctrine no doubt moved the field forward, it is illuminating to see how well Golgi’s reticularist ideas have held up in the face of modern evidence. Direct electrical coupling through gap junctions – still too small for even Cajal’s absinthe-enhanced eyes to see in 1906 – create a version of Golgi’s syncytium whereby “[s]erially interconnected neurons could be traced laterally in a boundless manner through multiple gap junctions.” (Fukuda et al, DOI:10.1523/JNEUROSCI.4076-05.2006). Maybe their Nobel debate remains ongoing, over a century later?

  4. True! But like you say, given the evidence and tools available at the time, there would not have been any way for Golgi to know about gap junctions! He was correct, but not based upon the scientific evidence available to him!

  5. Thank you for featuring the lovely artwork by Santiago Ramon y Cajal. This post would make a nice starting point for an art history lecture or essay. That is probably not the sort of praise you want, but my knowledge of art history is a lot stronger than my knowledge of neuroscience (in the latter, nearly zero).

    I also appreciate that you found something in the public domain to post. That required effort, I'm certain. It is great to view this, and determine that I can reproduce it (referring to the art, with attribution, not your post!) on my own blog or elsewhere.

  6. Thanks Ellie! Wasn't that difficult, actually. Just happenstance. Cajal's a great neuroscientist, and a mentor gave me one of his books to read.