Thursday, July 5, 2012

Spat out of Nature by Nature: Konrad Lorenz and the Rise and Fall of Ethology

Jackdaws (Corvus monedula), widespread crows throughout Ireland, make delightful pets.  A school friend of mine in Dublin, Sean Farrell, kept one for a few months back in the late 1970s when we were both in our early teens.  The bird had broken its wing and Sean nursed it back to health.  The jackdaw was a noisy fellow and had his species' penchant for shiny things.  Sometime later I was happy to read that Konrad Lorenz (1903-1989), the Austrian ethologist, published a behavioral study of a colony of Jackdaws that he maintained and included an account of the birds in his charming book King Solomon’s Ring.  King Solomon’s Ring, (the title is taken from the legend that Solomon had a ring that allowed him to talk to the animals), was a popularization of the emerging science of ethology, that is, the biological study of behaviour, and was perhaps the first book that confirmed to me that there was a to make a living out of what I happened to like to do.

I progressed from the lighter but nonetheless delightful accounts of animal behavior in King Solomon’s Ring and in Niko Tinbergen’s Curious Naturalists to the greater heft of Lorenz’s classic On Aggression (1966) which I read as a zoology undergrad at University College Dublin in the 1980s.  In this book, published later in his career when he was in his sixties, Lorenz shared his mature analysis of how insights from the study of the instinctual behaviour of animals can be helpful in thinking about the human condition.  As we shall see, as a good Darwinian he regarded human conduct as revealing much about our essentially animal nature, but unlike other animals we posses, he argued, a characteristic of being able to overcome this legacy.  In fact, he deemed it critical to our species survival that we simultaneously evaluate the evolution of aggressiveness in a clear-headed way while we find cultural solutions to discharging these unavoidable tendencies in a harmless way.  More harmless than war, that is.  Lorenz had lived through a war and he was committed to helping humanity avoid another one on that scale.

Before looking at the details of Lorenz’s analysis of human nature (in the post to follow this one) a word or two about the science of ethology which emerged as biological subdiscipline in the 20th Century under Lorenz’s and Tinbergen’s influence.  Together with Karl von Frisch, who worked out the details on the so-called “waggle-dance” of honeybees that allowed the hive to share information on the location of a food source, Lorenz and Tinbergen shared the Nobel Prize (in medicine) in 1973.  A primary task of ethology was to place questions about the behaviour of animals in an evolutionary context.  How does the behavior function to increase the success of the animal; what are the triggers for the expression of that behavior, how does the behavior develop in the life of that individual; and, finally, what was the pattern in the evolutionary development of the behavior?[1],[2]  Some regarded it as a strength of the discipline that it simultaneously asked questions about the adaptive nature of the behavior, the mechanism by which the behavior is displayed, and how functions in the ecology of the organism. Critics, however, saw in this the danger that teleological thinking could creep into the analysis: the assumption that behavior developed to the point of ever increasing perfection as the behavior reached a preordained goal.

Characteristic of early ethological studies are investigations of the behavior of animals in the field conditions or otherwise in circumstances where the full and natural expression of behaviour can be observed.  A research theme that was especially important in ethological investigations in the mid-twentieth century centered on unveiling so-called “fixed action patterns”, an sequence of behavioral component that are set in motion in response to a sign stimulus, also known as the innate release mechanism.  The fixed action pattern is instinctive, predictable and invariable.  Tinbergen showed a variety of such behaviors in animals as diverse as fish, especially stickleback, a favorite organism among early ethologists, and birds.  The sign stimulus can be a relatively narrowly defined environmental signal that elicits a fixed response from an animal.  So, in the case of a red flag to a raging bull, the redness of the flag is the signal.  Now, the color red does not, in fact, serve as a releaser for bull behavior; any fluttering material regardless of color can serve the purpose, but for sticklebacks red is actually the signal.

One of Lorenz’s early contributions to the development of the discipline was his discovery of imprinting, where, in one of the earliest observations of it, a newly hatched flock of Graylag goose goslings would follow the first organism they were introduced to after hatching, including Lorenz himself.  Oftentimes, Lorenz was pictured ambling in front of his little brood of goslings.  As the birds grew they could apparently have confused species identities, even at times attempting to mate with people.  Imprinting can come in a variety of forms in different circumstances – sexual imprinting which determine that appropriate potential mate is distinct from filial imprinting where goslings follow their parent or parent surrogate, although the two can be connected.  Elements of imprinting appear to be at play in the processes whereby birds learn their song.  Another Lorenzian proposal concerns the accumulation of energy directed at a specific energy – his hydraulic model of behavioral motivation.  He called this “action specific energy”.  Each behavior has its own associated internal force and the behavior is regulated by an innate releasing mechanism.  As the force builds, ultimately it is discharged in response to the appropriate sign stimulus.  In places Lorenz discussed the “unexpected correspondences” between his understanding of motivation and that emerging from Freudian psychoanalysis. 

Between the Dutch-born Niko Tinbergen and the Austrian Konrad Lorenz the discipline of ethology developed as an influential evolutionary account of animal behaviour during the middle years of the 20th Century.  By all accounts these men, who rarely met in person, developed a synergetic research program that was highly influential not only because both men were persuasive advocates of their discipline, but also because they did not shy away from translating their insights from the study of birds and fishes into prescriptions about our human predicament.  During the cold-war years, where fears of nuclear annihilation and environmental destruction were mounting ethology pronounced on those aspects of behavior that were hard-wired and those that could be redirected.  Despite the eminence and appeal of the discipline, ethology has largely disappeared, at least in its classical form (the theoretical constructs of Tinbergen and Lorenz are sometimes dubbed “classical ethology”).  The reasons for the demise are summarized below.  I remark on them, not just because it is an interesting case study in the process whereby an academic discipline recedes from power, but because it serves to remind us that the clout that accompanies disciplines instructing us on the best path forward need to be tempered by humility and some caution is needed in regarding their findings.    


Some species go extinct when their entire population diminishes to zero and they are supplanted by a new species; others disappear from the record when the population simple evolves into a new species over time.  Although ethology lingers as a discipline, to a large extent ethology as a discipline was eclipsed and absorbed by other developments in behavioral disciplines.  Ethology, it seems, conforms to the second extinction pattern. 

In 1975 E. O. Wilson published his groundbreaking and controversial book Sociobiology: The New Synthesis which drew upon ethological insights.  However, sociobiology defined questions about the emergence of social behavior in a way that relied more on population biology rather than on the methods of ethology.  Wilson predicted that ethology would simply be subsumed by behavioral ecology, neurophysiology, and psychology.  Shortly afterwards Richard Dawkins, a student of Tinbergen’s, published The Selfish Gene (1976) which popularized the gene-centric approach to understanding social behavior that had been developing among evolutionary biologists since the 1960s.  Though Dawkins defined himself as an ethologist in The Selfish Gene nevertheless he had some pointed criticisms of the evolutionary approach of the early ethologists.  “The trouble with these books [the books of Lorenz and some other ethologists]”, Dawkins fulminated, “is that their authors got it totally and utterly wrong because they misunderstood how evolution works”.  Wrong because in attempting to locate an adaptive explanation for a behavior ethologists looked to that behavior’s species preserving function, rather that asking how it increased individual fitness, or fitness at the level of the gene.  Lorenz, for example, when querying the role of aggression between individual of the same species enumerated a number of ways in which “intra-specific aggression assists the preservation of an animal species.” 

A yet more recent outgrowth of ethology, evolutionary psychology continues some of the research program inaugurated by both ethology and sociobiology.  A major emphasis in evolutionary psychology and building upon foundational books by Donald Symons The Evolution of Human Sexuality (1979) and The Adapted Mind by husband and wife team John Tooby and Leda Cosmides (1992) are questions concerning the influence of pre-historic environments of the evolution of the human mind.

Finally, behavioral ecology reinvigorated the evolutionary focus of classical ethology and true to its ethological roots examines a wide range of organisms, often in the wild.  This is in contrast to evolutionary psychology, for instance, that is primarily concerned with the human animal.  Behavioral ecology has its own questions on habitat selection, foraging behavior, group behavior, reproductive and life history strategies, and so forth that are distinct from its ethological antecedents.  John Krebs, an influential behavioral ecologist was also one of Tinbergen’s students.[3]
It is clear that the ethology of Lorenz and Tinbergen got absorbed into a number of derivate disciplines.  Part of this seems like ordinary academic development.  For example, my academic training is as a zoologist.  These days zoology departments have become rarer, and those who can claim to be zoologists oftentimes identify by their specialty: ecologist, physiologist and so forth.  However, some of the reasons for the decline of ethology may relate to broader politics.  There has always been a level of discomfort over Lorenz’s relationship with the Third Reich and this may have tarnished the reputation of ethology (see an interesting comment on this from Michael Ruse).  Certainly when I become aware of this when I was in college, I found my infatuation with Lorenz waned rather quickly, though I still regarded his findings to be relevant and interesting.  

The facts are these: Lorenz applied to join the Nazi in June of 1938 shortly after the Anschluss, Gemany’s annexing of Austria in March 1938.  Furthermore, he claimed at the time that his work was compatible with the aims of National Socialism.[4] In his writing at that time he stressed some points of convergence between his work and Nazi ideology.  It appears that Lorenz’s enthusiasm was rewarded.  In 1940 he was appointed as professor at Königsberg University (the professorial chair that had been Immanuel Kant’s).  From 1942–44 he was a physician in the German army, serving in Poland.  His work at that time included efforts to psychologically distinguish Poles from Germans.[5] [6]  At the time of receiving the Nobel Prize in 1973 Lorenz apologetically claimed that he had been naïve about the regime and had cooperated in order to maintain his career.  For Tinbergen, who had been interned in the Netherlands by the Nazis in 1942, Lorenz’s apologies were enough, and they continued to collaborate after the war.  For some this was not apology enough.  However, questions about the political implications of ethology and its descendents continue to be raised.  Dawkins, for instance, addressed what might seem like “disagreeable social, political or economic implications of The Selfish Gene” in prefaces to more recent reissues of his book.  Although the cruder versions of all of these disciplines seems to open to charges of determinism, it is a change that is resisted forcefully by most behavioral scientists.

Whether or not ethology has declined or has been transformed into a set of new disciplines its influence has been undeniable.  One of its distinctive qualities was the degree to which its founders devoted themselves to issues of environmental protection.  Late in his life Lorenz was supportive of the Austrian Green Party.  In his writing a significant concern was that we apply an understanding of animal behavior to humans in an effort to solve our environmental problems.  This can be done, he and Tinbergen imagined, by recognizing what in humans is instinctive and what could be culturally modified.  If these days we no longer maintain quite as crisp a distinction between what seemed fixed and what seemed flexible in human behavior, nonetheless their work provides us with a significant model of the human as being both part of nature, and in some ways set apart from the rest of nature.   The details of this model I’ll address in my next post.

[1] Tinbergen, N. 1963. “On aims and methods of ethology.” Zeitschrift für Tierpsychologie 20:410-433.
[2] Tinbergen, N  On War and Peace in Animals and Man  Science, Volume 160, Issue 3835, pp. 1411-1418. 1968.
[3] For an excellent introduction to the themes, methods, and results of this discipline see Nicholas B. Davies and John R. Krebs An Introduction to Behavioural Ecology. Wiley-Blackwell (original edition 1981)
[4] A good account of Lorenz’s politics and his behavior during WWII is provided by Richard W. Burkhardt's Patterns of Behavior: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology (University of Chicago Press, 2005).
[5] Reader's Guide to the History of Science, s.v. "Lorenz, Konrad 1903-1989," accessed July 05, 2012,
[6] The Hutchinson Dictionary of Scientific Biography, s.v. "Lorenz, Konrad Zacharias (1903–1989), accessed July 05, 2012,

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