Studies in Animal and Human Behaviour

The papers argue that ethology must operate as a biological discipline, not a branch of psychology. Lorenz contends that species-specific behavior patterns—innate motor sequences, releasers, and companion-schemas—are evolutionarily conserved structures whose comparative study reveals phylogenetic relationships, just as comparative anatomy does. This requires classifying behavior by homology rather than by superficial similarity, and rejecting both the reflex-chain model of behavior and anthropomorphic interpretation. The 1937 essay “The establishment of the instinct concept” systematically dismantles the Spencer/Lloyd Morgan and McDougall theories of instinct, while the 1932 paper on identification methods sets out the criteria for recognizing species-specific patterns in birds.

This paper introduces the companion-schema (Kumpan-Schema), the innate perceptual template through which birds recognize and respond to specific social partners: parental, sibling, sexual, and social companions. Lorenz argues that these schemas are species-specific and innate, though they become individually refined through experience—most dramatically through imprinting, the process by which a young bird irreversibly learns the characteristics of its species or surrogate parent. The companion-schema concept influenced subsequent research on social recognition in animals and contributed to attachment theory through its emphasis on innate social predispositions. Contemporary ethology and developmental psychology retain the insight that social bonding involves species-specific perceptual biases, though the strict innateness claims have been qualified.

The 1938 paper on egg-rolling is among the most cited demonstrations of the fixed action pattern concept. When a greylag goose sees an egg outside its nest, it performs a stereotyped movement: extending its neck, rolling the egg back with the underside of its bill, and tucking it beneath the body. Lorenz showed that the motor sequence is endogenously organized and relatively inflexible: if the egg is removed mid-roll, the goose completes the movement as if the egg were still present. He distinguished this fixed pattern from the taxis—the orienting response to the egg’s position—and demonstrated that the two components can be experimentally separated. The study became a model for distinguishing instinctive motor coordination from orienting behavior.

Reviews were largely positive but noted limitations. Eckhard Hess, writing in Contemporary Psychology in 1971, praised the volumes for making Lorenz’s foundational papers accessible to Anglophone readers and for providing “a real closeup of evolutionary, taxonomical, and other biological concepts in application to behavior.” Hess acknowledged Lorenz’s contributions to research problems while noting the work’s “entertaining” quality. A review in Science praised the collection as evidence of ethology evolving into a mature science. However, Book Review Digest noted Lorenz’s “fondness for analogical reasoning and assertion,” observing that his excursions into human psychology and theory of knowledge were less successful than his animal behavior work. The translator, Robert Martin, provided a foreword contextualizing the papers.

The Foundations of Ethology (1978/1981) is Lorenz’s systematic restatement of the theoretical positions first articulated in these papers, updated to respond to three decades of criticism. Whereas Studies in Animal and Human Behaviour presents the original arguments in their historical context—including the 1937 “establishment of the instinct concept,” the 1938 egg-rolling study, and the 1942 essay on inductive and teleological psychology—The Foundations reorganizes and defends those commitments against critics like Robert Hinde and Daniel Lehrman. The later book revises the psycho-hydraulic model of motivation and introduces the “open program” of learning. Together they trace Lorenz’s career from founding polemics to retrospective systematization.

Lorenz argues that behaviorism’s reflex-chain model cannot account for the endogenous production of behavior. In the 1937 essay, he contends that instinctive acts are not concatenations of reflexes but centrally coordinated patterns that occur spontaneously, even in the absence of external stimuli, once the organism’s action-specific potential accumulates. This position rejects the stimulus-response framework that dominated American psychology. The 1942 paper “Inductive and teleological psychology” further distinguishes ethology’s biological method from both behaviorist reductionism and introspective psychology, arguing that the comparative study of species-specific behavior provides a causal, evolutionary account without recourse to subjective experience.