The topic of my talk, as I hope to show you, fits very nicely into a conference on "NMR and Biomolecular Signaling". True, this is not going to be a scientific presentation, even though I've paid my dues and I have done affectionate work on calcium-binding in my time.
We have heard today about the interaction of metallic ions with biological molecules exquisitely tuned for that purpose. We are now going to look at biological systems that differ by a few additional steps in complexity -- we shall thus position ourselves downstream from the previous talks if you will accept the metaphor -- but still applying the laws of physics and of chemistry, regarding in particular ionic transfer across cell compartments , to those higher-level phenomena observable as patterns of animal behavior. This was my first reason for choosing this topic.
The second reason is Sture Forsén's scientific personality. Sture has had the immense merit of steering clear of fashions and bandwagons. Sture has charted his own original and creative course. For this reason, I have elected to tell you about a man and a piece of work from Twentieth Century biology. The man was brilliant and his work is superb. I have no doubt that historians of science, in a few decades, will place him among the giants of Twentieth Century science.
The third and last reason is more personal. Sture and I have been friends for nearly 40 years. I don't know who started it, but I am very fond of this little tradition: we recommend to one another musical pieces and book titles. Accordingly, today I am making public the title of a book which I am submitting for Sture's attention. It is Kenneth D. Roeder's Nerve Cells and Insect Behavior, published by Harvard University Press. I was able to order a copy three or four years ago, and I would not be surprised if the book continues to be available. It is truly superb.
The background
First, a word about the historical context: Roeder's book was published during the period which came to be known, after its brutal disappearance, as the Golden Sixties. It was indeed a Golden Age for science. We should feel privileged to have experienced it. We now know how Adam and Eve must have felt upon being expelled by God from the Garden of Eden.
Scientific research then underwent a big expansion. Governments of industrialized countries funded public research, civilian and military both, at unheard-of levels. They seemed to have grasped its key role in the economy. This was also the time for a spectacular increase in university demographics. As you will know, the bulge of academic staff hired in the 60s is gradually being retired at present, thus promising to offer a glut of new positions to be filled by much younger people. Furthermore, the Sixties was the time when the general public felt very positive about science. As a consequence, science studies could recruit students among The Best and the Brightest, to borrow the title of a book that chronicled American politics of this era.
What about biology? From the viewpoint of the balance between the various sub- disciplines, the aftermath to World War II saw a big development of neurobiology. Its rise was made possible, in part, by technological innovations, by advances in electronics especially, which accompanied the development of radar in particular. As you know, they led to the discovery of nmr in 1945. More relevant to my topic, Andrew Hodgkin who, during the war, had been involved in radar development, made use of his knowledge of electronics to establish the physicochemical mechanism for transmission of the nerve impulse -- by way of transmembrane fluxes of sodium and potassium ions -- a fine piece of work which led to his being awarded the Nobel prize for physiology or medicine in 1963, the very year in which Kenneth D. Roeder's book first appeared.
The landscape of biology in the Sixties, besides the entry of a vigorous neurobiology, shows another newcomer. For being less delocalized and less imperialistic than neurobiology, ethology (the study of animal behavior) nevertheless made its impact felt. Furthermore, books written for the general public by such pioneers as Nikolaas Tinbergen, Konrad Lorenz and Karl von Frisch made a strong impression and they became instant worldwide best-sellers. Tinbergen had published The Herring Gull's World in 1960. He published another more popular book, Animal Behavior, in 1965. Konrad Lorenz enjoyed comparable success with On Aggression in 1966. These three practitioners of experimental science, to whom we are indebted for discoveries such as the imprinting of the mother on the infant (and other seeds for sociobiology), or the dance of the honeybees communicating topographical information about flowers to reap pollen from, would share another Nobel Prize in physiology ten years later, in 1973.
Let me also mention, as part of the intellectual landscape for Roeder's book appearance, the invention of sociobiology by Edward O. Wilson: also an entomologist by training, also an outstanding writer of popular science, he set about during the period contemporary to publication of Nerve Cells and Insect Behavior, to re-naturalize the human species. This book Insect Societies was published in 1971. Wilson's aim was to replace our behavioral patterns in their proper place as instances of more general animal patterns. There was an outcry: publication of Wilson's major book in the 1970s would launch a major controversy, more political than scientific. Biologists such as Stephen Jay Gould and Richard Lewontin, or a linguist such as Noam Chomsky, went up in arms; they attacked Wilson savagely, making a caricature of his ideas in order to ridicule and destroy them. This may count as the opening salvo from militants of a scientific Left in the ensuing and drawn-out fight for political correctness.
The landscape of biology is also revolutionized, let me remind you, during the same period, by the forceful emergence of a molecular biology, at long last one might comment. Watson's and Crick's discovery of the DNA double helix was almost instantly greeted with a Nobel Prize, again for physiology or medicine: they shared with Wilkins the 1962 Prize.
A new element in the overall picture of biology at the time, which one would be remiss in not mentioning, even though it, too, borders on the political, is the appearance of ecology as a generalized concern about mankind predating upon all other species and destroying biodiversity. Here again the landmark was publication of a book. In 1962, the year preceding that of Roeder's book, Rachel Carson's Silent Spring appeared. It documented the ill effects of DDT on numerous animal species. This pesticide had caused havoc since it had been used massively and indiscriminately in the fight against malaria, predominantly. This also led to a major controversy.
Shall we note too, outside the scope of our camera with its focus on science history but as yet another pertinent element, because it is part and parcel of the same Zeitgeist or of the same epistémé (to borrow Michel Foucault's term) the mediatic event of the publication of the novel Lolita by an amateur entomologist, Vladimir Nabokov. Much closer to our concern his memoir Speak Memory, also published in the Sixties, told of a childhood lived in bliss, in an environment of happiness and luxury reminiscent of scenes by Impressionist painters when in particular, together with his father, this happy child was roaming Central Asia in the search for rare butterfly species.
We are dealing with a small book of 238 pages including the bibliography and the index with overall dimensions of 14 cm by 21.5 cm. It is cut into 13 chapters of no more than 10 to 20 pages each. Conversely, it sports abundant illustration, with a total of 63 figures, divided between photographs and line drawings.
The first three chapters serve as a reminder of the general principles of biology. They deal with the encoding of information, with methods for study of animal behavior, and with the physicochemical mechanism for the nerve impulse. The three following chapters tell how bats feed on night moths. Then comes a chapter on the escape behavior by cockroaches, and next a chapter on postural reflexes at the level of the knee, in higher vertebrates
The ninth chapter, again with a general topic, covers endogenous neuronal activity. This leads to the next chapter linking this endogenous activity to behavioral traits in both the cockroach and the praying mantis. Chapter 11 examines what neuronal signals become in the central nervous system of a moth. The penultimate chapter takes for its topic the brain of insects. And the last chapter presents a synthesis, based on evolutionary theory, on the parcimony with which regulation of behavior among insects relies on inhibition mechanisms.
The quality of the writing is outstanding, from beginning to end. The book is a gem. It reads as a whodunit. Roeder is a born science communicator. Memorable formulas spring forth just at the right place, the very place where such analogies are most illuminating. He is being constantly very careful to qualify the inferences he makes from his data, whether observational or experimental and he does not sweep under the rug what remains uncertain or doubtful. One can and does admire also what an astute experimentalist he is, putting together ingenious devices to suit his goals.
Whenever a well-recognized and experienced scientist publishes a book, the aim
differs from that of the publications. He may want to reach out to a larger
public than the international peer group he belongs to and usually addresses.
He may also want, if he has reached the stage when, he can review the field
authoritatively and if, in his judgment, he can offer furthermore a novel and
synthetic view, to give it a combination of depth and brilliancy, and thus
reach a wide audience through the writing of such a book
Roeder's
one-upmanship is striking in his book. From the very first sentences in the
preface, the author assumes a magisterial tone. The conventional "we" of
primary publications is swept aside and replaced by a self-referring "I" which
is given pride of place from the very first sentence; in which Roeder refers to
(I quote) "aspects of the neural activity and of the behavior of insect of
particular interest to me".
Such an assertive mode shows little respect for the readership. The reader is abruptly confronted with a clear-cut choice, either to totally espouse the views of the author, or to turn down the invitation to let himself be led through the nose, in a narrative in which the rights of the readership, such as skepticism toward reception of the author's ideas, are being systematically ignored.
The empirical evidence in support of this observation is a simple word count: the pronouns "I" and "me" occur four times in the very first paragraph, and 10 times in the second (not including the possessive "my"). There are quite a few sentences asserting scientific authority such as (I quote) "my own scientific experiences "(end of quote) in which "my own" yet reinforces "my" which many a writer might have deemed sufficient.
I turn now to
If one continues to read his Preface, the author drops quite a few epistemological hints. Since they are a little weird, they are worth a closer look. A first such note, and it shows such a light touch that one might doubt its very existence, but it comes in the very first paragraph, is the contrast between the two expressions "of particular interest to me" and "mainstream of research" in the first and in the last sentence, respectively.
Mainstream of research rings an all too familiar bell to many of us: it evokes the notion of paradigmatic science which Thomas Kuhn introduced in his famous book, almost exactly the contemporary to Roader's: The Structure of Scientific Revolutions was published in 1962.
And the contrast I have drawn your attention to is not unexpected. One is not surprised to find such a claim of originality on the part of an extremely intellectual biologist, of an essay-writer who is obviously convinced of his worth , and who combines being an innovator and an iconoclast.
However, further on, Roeder writes a truly bizarre sentence. So far so good, we did understand that he was placing himself apart from the norm and above it. This other sentence occurs at the onset of the third paragraph. It pits as opposite two types of scientific fields: on one hand well-established fields and, on the other fields valuable for their internal consistency.
If, for argument's sake, one returns to the kuhnian paradigm, the distinction posed by Roeder splits the paradigm into two rival camps. Allow my insistence that we find here not a stylistic and rhetorical ambiguity, but a genuine opposition. With this sentence, Roeder announces an attempt, his attempt to launch bridges between these two camps, that he sets as rival and as antagonistic. Clearly, their existence is not a matter of doubt, in his mind. Thus, we should attempt at identifying these two variaties of science. Let us give a look, for this purpose, at the examples that Roeder gives further on, in his Preface.
He mentions, in his first paragraph, the possibility that the various chapters of his study would not form a coherent whole, that they lack an overall plan. In the absolute, such a statement may look like a mere rhetorical flourish. After the opposition Roeder sets, it becomes a different matter. Furthermore, he tells us in the very next sentence that he has chosen to be a conformist, to place himself in a paradigmatic fold so to say: he will use neurophysiology as a backdrop. But who is to believe him, after this weird sentence of his? And indeed most of his results will run counter to the tradition of neurophysiology.
Pitting together those two assertions does not help, to the contrary: we can only observe that Roeder, at least at this stage, nowhere finds coherence. His own results according to him lack coherence. And neurophysiology, as a whole, also lacks coherence: it is a well-established field, therefore pluralistic and admitting of quite a few incommensurable interpretations.
Even though we are not in a position to make a strong assertion of it yet, it appears nevertheless that Roeder is setting-up an opposition, not merely rhetorical, and its trace is apparent already in the title of his book (Nerve Cells and Insect Behavior), between the two disciplines, neurophysiology on one hand, the study of animal behavior, ethology if you will, on the other hand. The text he wrote is unambiguous: Roeder implies that internal coherence is to be found in the then brand-new field of ethology; whereas incoherence is the price that neurophysiology has to pay in exchange for its comfortable position as a well-established scientific discipline.
Such a provisional conclusion however is simplistic. It is too clear-cut. Roeder's thought and its expression are quite a bit elusive. To hold him to firm and explicit expression of what he means is a tough proposition. For instance, those two disciplines, at a distance from one another but complementary, neurophysiology and ethology, have each their orthodox practitioners. Roeder is an heretic, he wants quite obviously to appear as an outsider to both sides.
Let us note in passing that his view of epistemology turns Kuhn's analysis upside down. Let me remind you that one of the criteria for a paradigm, according to Kuhn, is its becoming rigidified and formalized in the transmission of acquired knowledge, in textbooks in particular.
Roeder's perception differs from Kuhn's. It is no less mythical. "Brand new is beautiful" could become his motto. The emergent discipline, that of ethology, has the strength stemming from being a unitary whole. It presents itself in the shape of a coherent discourse. By contrast, the existing and well-established discipline, neurophysiology, suffers -- I am verbalizing here what remains implicit in the text of Roeder's Preface and is never expressed in such a guise -- it suffers from polyphony or polyglottism. Such an interpretation is borne out by the last sentences in the Preface. Kenneth Roeder sets himself not only as a referee between the two jousting camps, he states explicitly the interdisciplinary transfers which, in his opinion, ought to connect the two fields.
Indeed Roeder's book aims at being bidisciplinary. The conclusion to its Preface is worded in a familiar rhetoric, that of the understatement. But Roeder is nevertheless quite clear. He tells ethologists to learn some neurophysiology, they will gain in their understanding of some behavioral patterns. Likewise, he invites neurophysiologists, in symmetrical manner, to engage into some observation of their experimental subjects, to look upon them not only as pieces of meat to be inserted between electrodes but to promote them to the full dignity of animals observed in the wild.
Allow me a brief summation before I go on. The Preface to Roeder's book puts forward the necessity of a cross-linking of disciplines, of a successful hybridization between ethology and neurophysiology. Roeder is quite explicit as to how to achieve success in such hybridization: the transfer he advocates is an infusion from ethology into physiology.
Roeder makes a strong case for attentive observation of the normal behavior of animals in their natural habitat, by contrast to the artificial conditions prevailing in a laboratory. From such time-consuming data gathering, ethologists should be able to put forward hypotheses and concepts. This is step number one. Step number two is to look for analogies to these concepts in the results from neurophysiology. Once these analogies are identified, the scientist should look for cause-and-effect relations responsible for a phenomenon observed in the framework of neurophysiology to express itself in behavior thtat the ethologist can watch.
There is a model, there is a precursor as well for such bridge-building between the two fields. It is the child. The child who happens to be a future scientist, the scientist-to-be is crucial here. The recollections by a scientist of his awakening to science questions deserves scrutiny. The conventional view would have us cast away such testimonies from introspection and autobiography as tainted. I would argue conversely that they provide us with subtle insights into epistemology.
Does Roeder have anything to say about himself, as a child? He tells us that, confronted with the unforgettable wonder of seeing metamorphosis from a larva into an adult insect, he realized then that form and function are tightly correlated.
The generalization is for neuronal activity, if it is considered in turn as a morphological type, to have its functional correspondent, viz. an observable behavior. The intuition of the child for Roeder links with the experience of the scientist nearing the end of his career. Roeder suggests that morphogenesis anticipates upon the behavioral traits of the adult animal, that behavior recapitulates development, in a reformulation of Haeckel's Law, according to which "embryogenesis recapitulates phylogenesis".
Roeder tells us, still in his Preface, that "Most of the chapters have been written as self-contained essays". His book is seen thus to conform to a genre, that of the essay on a scientific theme. At the same time of the early sixties, Lewis Thomas was starting to write his essays for the New England Journal of Medicine, entitled 'Notes of a Biology Watcher"; and Stephen Jay Gould was also starting his monthly column in the magazine Natural History. Thus the reader of Roeder's book expects to find in it recent scientific work, chosen for its impact in biology, chosen also for what it will change in our ideas about both neurophysiology and animal behavior.
The reader has other expectations too. He has probably read already books such as The Origin of Species, or Maeterlinck's La vie des abeilles, or Jean-Henri Fabrés Promenades entomologiques. The reader expects scenes from the life of insects: he is looking forward to a narrative from natural history.
And indeed Roeder answers this yearning. The very next sentence in the Preface promises "adventures": the author tells the reader that he is about to read a thriller, to share astounding adventures, out of a novel. The promise is fulfilled. For instance, Roeder provides gripping descriptions of air combat between a moth and a bat, the astute truly clever maneuvering by the moth to try and escape from its predator. There is the occasional eye-catching expression or wording. To take an example, Roeder uses the word "resifting", with the meaning of pouring a mixture, that has already undergone separation, again through a sieve. To paraphrase the sentence in which it occurs, what Roeder tells us is that: "In a sense, this book reconsiders my whole experience as a scientist, in order to establish which of my perceived analogies stem from genuine cause-and-effect relationships".
Kenneth Roeder is intent upon describing his concept of bidisciplinarity in the
third paragraph. He does so with an image, very graphic, which provides us with
an emblem of his views:
"threads are set to connect only at their periphery
each of the fields, so that their edges tend to distort and to lose their
proportion with respect to the central core in each". The image is that of a
piece of fabric, stretching under the application of a tension. One cannot help
being reminded at the same time, when reading this description by a biologist,
of the preparation for a dissection: an animal has been sacrificed and opened
up, pins are stuck into the flesh to pull it and to stretch it for inspection,
to display the anatomy, the arrangement of the internal organs in the way that
will be the clearest for scrutiny .
I shall content myself pinning down the rhetorical ploy most often resorted to: it is the modesty of the object of study. The Preface states it and is even repetitive in this respect. Insects belong to a class of animals apparently inferior to more noble species, such as fish, birds, frogs, squids or cats. Insects are a 'boring topic" to the neurophysiologist. But Roeder finds it his duty to turn it around and to make it fascinating.
Furthermore, within the myriad insect species, which does Roeder elect to study if not the most modest, the most despised surely, the cockroach. Wittily , the acknowledgment part of this Preface expresses Roeder's gratitude to (I quote) this "alert, elegant and most misunderstood insect".
He sets up in this manner a handsome antithesis, by which he will turn this most misunderstood insect into the subject matter for the advancement of knowledge.
Roeder is keen upon resetting biology outside of the laboratory, within nature, and to link again in this manner with natural history. His interest is animal behavior. According to him, this is the only way in wich one can make sense of laboratory experiments on neurones.
According to Roeder, traditional physico-chemical reductionism breaks down biological facts into fragments that no longer make any sense. His approach is the exact opposite. Where the usual reductionist procedure is top down, Roeder's is bottom up: he starts from the very bases in order to understand the super structure.
And I must say that he is highly successful in so doing. (And so was Sture, in the handsome work he has presented us with throughout his career).