The difference between behaviorism and ethology has always been one of human-controlled versus natural behavior. Behaviorists sought to dictate behavior by placing animals in barren environments in which they could do little else than what the experimenter wanted. If they didn’t, their behavior was classified as “misbehavior.” Raccoons, for example, are almost impossible to train to drop coins into a box, because they prefer to hold on to them and frantically rub them together—a perfectly normal foraging behavior for this species.9 Skinner had no eye for such natural proclivities, however, and preferred a language of control and domination. He spoke of behavioral engineering and manipulation, and not just in relation to animals. Later in life he sought to turn humans into happy, productive, and “maximally effective” citizens.10 While there is no doubt that operant conditioning is a solid and valuable idea and a powerful modifier of behavior, behaviorism’s big mistake was to declare it the only game in town.
Ethologists, on the other hand, are more interested in spontaneous behavior. The first ones were eighteenth-century Frenchmen, who already used the label ethology, derived from the Greek ethos, “character,” to refer to the study of species-typical characteristics. In 1902 the great American naturalist William Morton Wheeler made the English term popular as the study of “habits and instincts.”11 Ethologists did conduct experiments and were not averse to working with captive animals, but still a world of difference lay between Lorenz calling his jackdaws down from the sky or being followed by a gaggle of waddling goslings and Skinner standing before rows of cages with singly housed pigeons, firmly closing his hand around the wings of one of his birds.
Ethology developed its own specialized language about instincts, fixed action patterns (a species’ stereotypical behavior, such as the dog’s tail wagging), innate releasers (stimuli that elicit specific behavior, such as the red dot on a gull’s bill that triggers pecking by hungry chicks), displacement activities (seemingly irrelevant actions resulting from conflicting tendencies, such as scratching oneself before a decision), and so on. Without going into the details of its classical framework, ethology’s focus was on behavior that develops naturally in all members of a given species. A central question was what purpose a behavior might serve. Initially, the grand architect of ethology was Lorenz, but after he and Tinbergen met in 1936, the latter became the one to fine-tune the ideas and develop critical tests. Tinbergen was the more analytical and empirical of the two, with an excellent eye for the questions behind observable behavior; he conducted field experiments on digger wasps, sticklebacks, and gulls to pinpoint behavioral functions.12
The two men formed a complementary relationship and friendship, which was tested by World War II in which they were on opposite sides. Lorenz served as medical officer in the German army and opportunistically sympathized with Nazi doctrine; Tinbergen was imprisoned for two years by the German occupiers of the Netherlands for joining a protest against the way his Jewish colleagues at the university were treated. Remarkably, both scientists patched things up after the war for the sake of their shared love of animal behavior. Lorenz was the charismatic, flamboyant thinker—he didn’t conduct a single statistical analysis in his life—while Tinbergen did the nitty-gritty of actual data collection. I have seen both men speak and can attest to the difference. Tinbergen came across as academic, dry, and thoughtful, whereas Lorenz enthralled his audiences with his enthusiasm and intimate animal knowledge. Desmond Morris, a Tinbergen student famous for writing The Naked Ape and other popular books, got his socks knocked off by Lorenz, saying that the Austrian understood animals better than anyone he’d ever met. He described Lorenz’s 1951 lecture at Bristol University as follows:
To describe his performance as a tour de force is an understatement. Looking like a cross between God and Stalin, his presence was overpowering. “Contrary to your Shakespeare,” he boomed, “there is madness in my method.” And indeed there was. Almost all his discoveries were made by accident and his life consisted largely of a series of disasters with the menageries of animals with which he surrounded himself. His understanding of animal communication and display patterns was revelatory. When he spoke about fish, his hands became fins, when he talked about wolves his eyes were those of a predator, and when he told tales about his geese his arms became wings tucked into his sides. He wasn’t anthropomorphic, he was the opposite—theriomorphic—he became the animal he was describing.13
A journalist once recounted how she had been sent into Lorenz’s office by a receptionist with the words that he was expecting her. His office turned out to be empty. When she asked around, people assured her that he had never left. After a while, she discovered the Nobelist partially submerged in an enormous aquarium built into the office wall. This is how we like our ethologists: as close to their animals as possible. It reminds me of my own encounter with Gerard Baerends, the silverback of Dutch ethology and the very first student of Tinbergen. After my stint in the behaviorist lab, I sought to enter Baerends’s ethology course at the University of Groningen to work with the jackdaw colony that flew around the institution’s nest boxes. Everyone warned me that Baerends was very strict and did not let just anybody in. When I walked into his office, my eyes were immediately drawn to a large well-kept tank with convict cichlids. Being an avid aquarist myself, I hardly took the time to introduce myself before we launched into a discussion of how these fish raise and guard their fry, which they do extraordinarily well. Baerends must have taken my passion as a good sign, because I was admitted without a problem.
The great novelty of ethology was to bring the perspective of morphology and anatomy to bear on behavior. This was a natural step, because whereas behaviorists were mostly psychologists, ethologists were mostly zoologists. They discovered that behavior is not nearly as fluid and hard to define as it might seem. It has a structure, which can be quite stereotypical, such as the way young birds flutter their wings while begging for food with gaping mouths, or how some fish keep fertilized eggs in their mouth until they hatch. Species-typical behavior is as recognizable and measurable as any physical trait. Given their invariant structure and meaning, human facial expressions are another good example. The reason we now have software that reliably recognizes human expressions is that all members of our species contract the same facial muscles under similar emotional circumstances.
Konrad Lorenz and other ethologists wanted to know how animals behave of their own accord and how it suits their ecology. In order to understand the parent-offspring bond in waterfowl, Lorenz let goslings imprint on himself. They followed the pipe-smoking zoologist around wherever he went.
Insofar as behavior patterns are innate, Lorenz argued, they must be subject to the same rules of natural selection as physical traits and be traceable from species to species across the phylogenetic tree. This is as true for the mouth brooding of certain fish as it is for primate facial expressions. Given that the facial musculature of humans and chimpanzees is nearly identical, the laughing, grinning, and pouting of both species likely goes back to a common ancestor.14 Recognition of this parallel between anatomy and behavior was a great leap forward, which is nowadays taken for granted. We all now believe in behavioral evolution, which makes us Lorenzians. Tinbergen’s role was, as he put it himself, to act as the “conscience” of the new discipline by pushing for more precise formulations of its theories and developing ways to test them. He was overly modest saying so, though, because in the end it was he who best spelled out the ethological agenda and turned the field into a respectable science.
Keeping It Simple