Instinct

The Instinct Concept of the Early Konrad Lorenz
Ingo Brigandt
Department of History and Philosophy of Science
1017 Cathedral of Learning
University of Pittsburgh
Pittsburgh, PA 15260
E-mail: [email protected]
July 30, 2004 (penultimate draft)

Forthcoming in the Journal of the History of Biology
Abstract. Peculiar to Konrad Lorenz’s view of instinctive behavior is his strong innate-learned
dichotomy. He claimed that there are neither ontogenetic nor phylogenetic transitions between
instinctive and experience-based behavior components, thus contradicting all former accounts of
instinct. The present study discusses how Lorenz came to hold this controversial position by
examining the history of Lorenz’s early theoretical development in the crucial period from 1931
to 1937, taking relevant influences into account. Lorenz’s intellectual development is viewed as
being guided by four theoretical and practical commitments as to how to study and explain
behavior. These four factors, which were part of the general approach of Lorenz but not of other
animal psychologists, were crucial in bringing about his specific position on instinctive behavior.
Keywords: Konrad Lorenz, ethology, instinct, innateness

THE INSTINCT CONCEPT OF THE EARLY KONRAD LORENZ 2
Konrad Lorenz was undoubtedly one of the main founders of ethology as a biological discipline.
In fact, the conceptual and theoretical framework of classical ethology was developed to a large
extent by him. Crucial for Lorenz’s view was that behavioral patterns have to be analyzed into
sequences of innate and learned behavior components. Only the innate components qualify as
instinctive behavior. Peculiar to Lorenz’s position from early on is the strong dichotomy between
the innate and the learned: there are neither ontogenetic nor phylogenetic transitions between
innate and learned components of behavior. Instinctive behavior patterns are rigid and do not get
modified or become more flexible due to experience in the course of ontogeny. In addition,
flexible or intelligent behavior does not evolve from instinctive behavior—a tenet that might
sound un-Darwinian and might be initially surprising given Lorenz’s commitment to largely
gradual evolution by natural selection. When formulating this position Lorenz contradicted
virtually all former and contemporary assumptions about instinctive behavior. His innate-learned
distinction became subject to vigorous criticism by psychologists in the 50s and 60s (most
prominently Daniel Lehrman),1 who argued that this approach was conceptually problematic and
fruitless as a means to understanding behavior and its development. Despite this critique Lorenz
never abandoned his strong innate-learned dichotomy.2
The aim of the present paper is to discuss how Lorenz came to hold this view of instinct. I
view Lorenz’s early theoretical development as being guided by four theoretical and practical
commitments. These are views as to how to study and explain behavior that emerged very early
on in Lorenz’s career. Taken together, these four factors guided Lorenz’s further development.
The four commitments are 1) Lorenz’s focus on innate, rather than learned, behavior, 2) the idea
that behavior has to be explained by physiological rather than psychological means, 3) the
comparative and taxonomic approach to behavior, and 4) the use of ideas from embryology to
account for the development of instincts. These four aspects of Lorenz’s general approach to
behavior emerged very early and were stable features of his perspective. The four commitments
constrained and drove his intellectual development, and they make intelligible why he ended up
with his strong tenet that instinct and experience are exclusive and that instinctive behavior does
not evolve into more flexible behavior. Other approaches in animal psychology did not endorse
these four components, and this difference in perspective explains why it came for instance to a
clash between the Lorenzian ethologists and the Dutch purposivists tradition in animal
psychology. Even though Lorenz developed his theory until 1935 on his own, I will suggest that
there are important influences on Lorenz. As I will explain later in more detail, several of the
intellectual influences on Lorenz are best viewed not as providing direct theoretical contributions
to Lorenz’s novel ideas but as supporting his general approach as embodied in the four aspects of
his framework.
Lorenz’s intellectual background and influences
Lorenz’s own education was important in determining his early theoretical development. Konrad
Lorenz (1903–1989) was the son of the internationally reputed orthopedic surgeon Adolf Lorenz.
Konrad’s elder brother Albert was a physician, as was Konrad’s wife Margarethe. Adolf Lorenz
was a dominant father figure, and even though Konrad would have preferred to study zoology, he
compelled his son to go to medical school first. From 1923 onwards Konrad studied at the
Second Anatomical Institute in Vienna and received his medical doctorate in 1928. Lorenz was a
student of the comparative anatomist Ferdinand Hochstetter, and his work focused on the
comparative anatomy of vertebrates. Once passing his doctorate in medicine, Lorenz enrolled at
the Zoological Institute, where he received his Ph.D. in zoology in 1933. The director of
Zoological Institute was Jan Versluys, a comparative anatomist as well. In fact, zoology at the
University of Vienna was dominated by comparative morphologists and embryologists. While

Lorenz was working towards his Ph.D. in zoology, he still worked as Hochstetter’s assistant at
the anatomical institute and he was teaching as a demonstrator (laboratory instructor) in anatomy,
which was his primary income apart from his wife’s. For a several years Lorenz could work in
anatomy while at the same pursuing his main academic interest—the time-consuming study of
bird behavior. While Lorenz was working on the ‘Companion’, the monograph-long essay on
animal behavior that secured him wide recognition among ornithologists,3 he was still teaching
anatomy. This situation, however, became problematic once Hochstetter retired in 1933. His
follower was unsympathetic to Lorenz’s work on animal behavior. In 1934 Lorenz made up his
mind and left the field of anatomy, pursuing a career as an animal psychologists. The upshot of
this brief overview of Lorenz’s education is clear—Lorenz’s intellectual background was medical
and his primary competence was the field of comparative anatomy. The discussion below will
reveal why this is so crucial for Lorenz’s theoretical development.4
Despite the anatomical work with Hochstetter in Vienna, Lorenz’s primary scientific mentor
was the ornithologist Oskar Heinroth (1871–1945).5 Heinroth was assistant director of the Berlin
Zoo and dedicated his life—as did his first wife Magdalena—to the study of birds and especially
their behavior. Out of years of observations grew detailed comparative studies of bird behavior,
in particular a work on the birds of middle Europe encompassing four volumes.6 In fact, Heinroth
was one of the few before Lorenz who realized the taxonomic importance of behavioral patterns,
i.e., the idea that behavior patterns are characteristic of taxonomic groups and can be used to
classify organisms in the same manner as morphological characters are. From 1930 onwards
Lorenz had an intense correspondence with Heinroth.7 Despite detailed comparative studies of
instinctive patterns, Heinroth’s work was not of a theoretical nature. Indeed, Heinroth largely
avoided scientific terminology and wrote in a style accessible even to lay readers. Explicit
theoretical discussions of the nature of instinct or the literature on animal psychology were
absent from Heinroth’s work. Instead, theoretical assumptions were only implicit in the
perspective that Heinroth took. The Heinroth–Lorenz correspondence dealt primarily with reports
about new animals obtained, their keeping, some observations on them, and the loss of animals.
Discussion about instincts did not take place, and Heinroth seemingly had no explicit intellectual
influence on Lorenz developing his new theoretical ideas on instinctive behavior (published from
1932 onwards). Due to Lorenz’s ambition to found ethology as a distinct biological discipline, he
was trying to develop a theory of instincts. It appears that Lorenz developed his novel theoretical
ideas until 1935 primarily on his own. Lorenz sent his early theoretical manuscripts to the editor
of the Journal für Ornithologie, Erwin Stresemann, and only later gave them to Heinroth,
because he was afraid that Heinroth would view Lorenz’s account as mere speculation.8 Heinroth
seemingly liked Lorenz’s account,9 but the point is that he did not directly suggest any of these
ideas about the definition of instinct to Lorenz. Despite the fact that Heinroth had no specifically
theoretical contribution to Lorenz’s conceptual innovations, he nonetheless influenced and
supported Lorenz as regards the general scientific perspective on and approach to instinctive
behavior, as I will discuss in detail below.
Now I want to lay out the intellectual scene that the young Lorenz entered. It is beyond the
scope of this paper to offer an overview of the different approaches to and theories of animal
behavior that existed before the emergence of ethology. I will restrict my discussion to a few
issues that help to understand how Lorenz’s approach differed from most former traditions and
why his views were novel. Focusing on those aspects of the tradition that Lorenz rejected makes
sense for the purposes of understanding his theoretical development because Lorenz’s early
theory was based on his personal ideas and practical needs, rather than on a thorough
consideration and reflection of former accounts on the nature of instinct. Lorenz developed his
own approach, ignoring a good deal of prior debates and insight, and selectively addressed and
adopted a few theoretical considerations, which he deemed useful for his purposes at the time.
This holds to some extent for early ethology as a new discipline and research tradition in
general. Even though traditional animal psychology did not exist as a discipline distinct from
psychology or zoology, research in animal psychology was institutionally and theoretically well
established. In particular in the United States, research in animal psychology was carried out in
universities. Whereas animal psychologist often worked in laboratories, the early ethologists
instead observed animals in relatively natural conditions—as field naturalists like Tinbergen, or
as an animal keeper like Lorenz. There were surely ‘forerunners’ of this ethological practice,
such as the British field naturalists, or American ornithologists such as C. O. Whitman and
Wallace Craig. But as Richard Burkhardt has documented, it is far from obvious that something
like ethology would have developed out of these small traditions in Great Britain or America.10
The emergence of ethology was by no means an event to be expected; and the early ethologists
largely created their own research tradition and theoretical approach.
Before the emergence of ethology there were a variety of theories on instinct and behavior.
Different approaches disagreed on basic issues such as the relation between animate and
inanimate nature, the relation between humans and animals, or the question as to whether and
how evolutionary considerations have any bearing on the explanation of instinctive behavior. One
theory of instincts that was quite popular among biologists and in particular physiologists in the
first decades of the 20th century was the chain reflex theory of instinct. This theory viewed
instincts as complex sequences of reflexes and was inspired by physiological studies of
invertebrate behavior and reflexes in vertebrates (as conducted by Jacques Loeb and Heinrich
Ernst Ziegler in Germany, as well as Charles Scott Sherrington, Ivan Pavlov, and Vladimir
Bechterev). Many animal psychologists and proponents of more traditional theories rejected the
chain reflex theory because it could not account for the spontaneity of behavior. They argued that
whereas a reflex goes off in the presence of a stimulus only, animals are active and behave
spontaneously even without particular stimuli. Many of the approaches that rejected the reflex
theory as inadequate coincided with approaches that Lorenz would later label ‘purposivist’. This
name surely lumped together quite different approaches, but Lorenz were not particularly
interested in the details that distinguished them. Instead Lorenz’s own theory focused on a limited
set of issues and his views on these matters clearly distinguished his approach from those of
others—the purposivists thus became Lorenz’s primary target.11 These were approaches that
emphasized the purposiveness or goal-directedness of behavior. The idea is that behavior changes
flexibly depending on the situation and that we can understand behavior only if we view it as
directed towards a goal that persists despite variation in the behavior performed. Sometimes goal-
directedness was taken as a mere descriptive category in that it was stressed that one has to
individuate different types of behaviors according to the different goals that are pursued. Often
teleology and goal-directedness were taken as explanatory categories. For instance, it was
assumed that animals have a representation of the goal and act accordingly.12 Many animal
psychologists stressed consciousness, an animal psyche or mental states. Subjective states and
experiences of the individual were invoked to account for the motivation, spontaneity, and goal-
directedness of behavior. An influential case is the psychology of William McDougall. On his
picture, instincts were viewed as psychological drives or dispositions that motivate behavior.
McDougall offered a taxonomy of basic and subordinated instincts in accordance with the goals
they pursue. Some researchers endorsed vitalism in that they claimed that instincts and the goal-
directed-ness of behavior cannot be explained by physiological means only; instead specific and
irreducible psychological factors need to be taken into account.13

This selective exposition of the intellectual scene that Lorenz entered serves the purpose of
explaining the novelty of his position and understanding his intellectual development. Lorenz
(and later ethology to a large extent as well) broke with this broad purposivist tradition as
follows. Lorenz urged the need to break down behavior into relatively small components, and an
instinct is only a small part of an overall behavioral sequence. Lorenz viewed instincts as innate
motor patterns or what ethologists later called fixed action patterns. An example is the movement
of the neck and the beak a bird of a particular species makes when it catches an insect. Thus,
instincts on this account are bodily movements that can be cinematographically recorded. They
are not certain subjective states such as mental drives or desires that cannot be directly observed.
The final purpose of a behavior sequence or the animal’s awareness of this purpose is not the
feature that explains behavior; in fact, Lorenz assumed that birds do not have any awareness of
the purpose of their behavior (as we conceive it). On his account, the final goal such as the
building of a bird nest is reached because it is the outcome of a sequence of individual behavioral
components, where the performance of one leads to the next one. When a bird perceives nest-
building material, it carries out the movement to grasp this object with its beak, and this in turn
causes the next behavioral element that makes the bird to integrate the object into the nest. Once
this action is completed, the bird is ready to react to further stimuli from building material. This is
how the building of a nest is explained without assuming that the bird has a representation of the
nest to come into being or an insight into the goal of its actions. Lorenz viewed the purposivists
as constantly conflating proximate and ultimate causes (to use terms later introduced by Ernst
Mayr). Lorenz acknowledged that we need to recognize the adaptedness and goal-directedness of
behavior in that we give an evolutionary explanation of it. But these evolutionary causes of
behavioral adaptations are not to be confused with proximate causes such as physiological and
environmental triggers of behavior. The latter have nothing to do with the biological purpose of
behavior. An important argument for Lorenz’s position was the existence of so-called vacuum
activities. This refers to the situation when a behavior pattern is carried out while the normal
releaser is absent. For instance, a bird may carry out the insect-catching movement (an instinct in
Lorenz’s sense) even though there is no insect present. Vacuum activities show that instinctive
behavior is performed even though it is not adaptive and does not meet its usual goal. In
particular, the goal cannot explain instinctive behavior.
Lorenz developed his early views in a sequence of publications. His first important paper
‘Contributions to the study of the ethology of social Corvidae’ (1931) reported detailed
observations; in fact, it was very much a paper like the ones Lorenz’s scientific mentor Oskar
Heinroth wrote.14 Despite the absence of theoretical discussions, the paper assumed that a certain
type of behavior is instinctive if it is always performed in the same manner even though in certain
situations a different behavior may be more adaptive. This foreshadowed Lorenz later explicit
conviction that instincts are rigid and stereotypical behavior patterns that may occur independent
of the purpose they serve. Lorenz’s 1932 paper ‘A consideration of methods of identification of
species-specific behavior patterns in birds’ can be viewed as his first theoretical step. The notion
of vacuum activities was introduced in this publication, thus explicitly separating instinctive
behavior and insightful-purposive behavior. The famous essay ‘Companions as factors in the
bird's environment: The conspecific as the eliciting factor for social behavior patterns’, the so-
called ‘Companion’ paper from 1935, brought about Lorenz’s intellectual breakthrough in the
ornithology community. This paper contained Lorenz’s complete early theory of instinct,
introducing notions such as the innate releasing mechanism. The well-known first instinct paper
‘The establishment of the instinct concept’,15 published in May 1937, did not introduce new
concepts or novel theoretical insights. But now Lorenz gave for the first time a detailed critique
of former theoretical positions on instincts, clearly contrasting his theory with others. This paper
secured Lorenz wide recognition among biologists and animal psychologists beyond the
ornithological community. The second instinct paper ‘On the concept of instinctive behavior’,16
published a few months after the first instinct paper, restated Lorenz’s position with
modifications.
A main focus of my discussion is Lorenz’s strong innate-learned dichotomy. Apart from neo-
Lamarckists and behaviorists, most animal psychologists emphasized the clear difference
between innate and learned behavior, and took instincts to be innate features. However, virtually
everyone assumed that instincts could be modified, shaped and fine-tuned by the influence of
learning and experience. (In 1935 Lorenz stated that the British zoologists Elliot Howard was the
only one besides him who denied the influence of experience on instincts.)17 Animal
psychologists who viewed instincts as innate drives assumed that these innate factors motivate
behavior and guide it in a certain direction, but that the actual performance of behavior, its
repetition and the experiencing of a particular environment modify instinctive behavior. Even
researchers such as Conwy Lloyd Morgan, who viewed instincts as being based on inherited
rather than individually acquired neural features, assumed that they can be influenced by
learning. Lorenz, however, maintained that instinctive behavior patterns are rigid and cannot be
modified by experience—a conviction he clearly expressed in his publications from 1932
onwards. To reject arguments to the contrary, Lorenz introduced in his 1932 paper the notion of
instinct-conditioning intercalation. The idea is that overall behavior can be analyzed into a
sequence of components. Some of them are innate and are instincts in Lorenz’s sense, i.e., fully
inflexible behavior patterns. But other components of the overall behavior sequence are variable
as they involve prior learning, predominantly conditioning and in a few cases insightful behavior.
Thus the overall flexibility of behavior is compatible with Lorenz’s tenet that instinctive behavior
components are inflexible. Lorenz’s 1935 ‘Companion’ offered a further argument for his
position by viewing instincts in analogy to organs and stressing the idea of maturation. Even
though the performance of some instinctive patterns may improve during ontogeny, this does not
mean that this is due to the influence of experience. Instead, one has to consider the possibility
that the behavior pattern is fully innate but needs to mature in the course of development—like
organs, which are nothing but innate structures that have to develop during ontogeny until the
adult performance is reached. A maturation process is not necessarily a learning process. On
Lorenz’s arguments, standard observations and arguments of animal psychologists regards the
flexibility of behavior need not show that innate instincts are modified by learning.
Lorenz rejected not only the idea that there are ontogenetic transitions and influences between
innate and learned behavior components, he also maintained that instincts do no gradually evolve
into more flexible behavior. One traditional view about the evolution of instinct was the theory
that instincts are lapsed intelligence. The idea is that instincts are derived from acquired habits.
Due to the intelligence or the flexibility of animals, habits are formed that become part of the
hereditary equipment of an animal and thus turn into inflexible instincts. This theory stemmed
from the work of Charles Darwin and was endorsed prominently by George John Romanes and
Herbert Spencer. As this view assumes use-inheritance or other Lamarckist mechanisms of
inheritance, it was not very often endorsed by Lorenz’s contemporaries. But instead everyone
else assumed that flexible or insightful behavior is gradually derived from instincts. The
evolution of higher animals and intelligent behavior was viewed as a process whereby primitive
instincts become less and less rigid and instead the influence of ontogenetic adaptation and
learning increases.18 In the ‘Companion’ from 1935, Lorenz claimed for the first time that there
are no phylogenetic transitions between instinctive and flexible behavior. He emphasized that he
did “not regard the instinctive behaviour patterns as homologous with all acquired or insight-
based behaviour patterns.”19 On Lorenz’s account, flexible behavior does not gradually evolve
out of instinctive behavior. Instead, instinctive behavior components are lost and later replaced by
(non-homologous) insight-based behavior components. I shall explain Lorenz’s evolutionary
views in more detail below when we are trying to understand why he endorsed this rival view.
After the war Lorenz claimed that his early theoretical papers, such as his first important
publication from 1931 or the 1932 theoretical discussion “was written without any knowledge of
the theories held on the subject by purposivistic and behaviouristic psychologists”.20 This bold
claim was not quite right. Some of Lorenz’s later statements about his ignorance of other
theoretical accounts are probably to be viewed as part of the rhetoric used to justify his new
discipline.21 For Lorenz, good empirical science and the development of theories had to start with
extensive ‘theory-free’ observation, so that theoretical prejudices do not affect later
interpretation.22 Thus, Lorenz tried to represent his personal theoretical development in
accordance with his own philosophy of science. In actuality, Lorenz’s relatively theoretical paper
from 1932 on the criteria of instinct was partially influenced by Heinrich Ernst Ziegler’s book
The Instinct Concept in Past and Present Times.23 Ziegler’s treatise gave a historical and
systematic overview of different approaches and accounts of instincts before defending his own
theory, including a discussion of purposivist, vitalist, and Lamarckist approaches. So when
working on his first papers Lorenz knew about some of the possible methodological and
theoretical possibilities for studying instinctive behavior. Even though Lorenz was not totally
ignorant of some of the literature on animal psychology, he still was at best a very selective
reader, in fact, he actually was a highly unsystematic reader. He read some of the main figures in
animal psychology such Conwy Lloyd Morgan and William McDougall not very long before
working on the famous ‘Companion’ paper, probably in 1933.24 But at this time some of Lorenz’s
early theoretical views were already in place. This holds for his basic view of what instincts are
and his tenet that innate instincts are not influenced by learning. Thus, when we shall later be
trying to understand Lorenz’s strong innate-learned dichotomy and his position on the evolution
if instincts, we have to keep in mind that these novel and controversial ideas were not at all based
on a thorough reflection on the existing literature in animal psychology. Instead, Lorenz was
unaware of large parts of the literature, he came up with his ideas primarily on his own and
disagreed with other traditional accounts once he became aware of them.
The four commitments guiding Lorenz’s approach
As is well-known, Lorenz’s papers on instinctive behavior from 1935 onwards had an enormous
impact. Established scientists such as the American ornithologists Wallace Craig, who had done
substantial work on animal behavior and influenced Lorenz himself, were impressed.25 The early
theoretical framework of classical ethology was largely created by Lorenz. Even his collaborator
Niko Tinbergen, who played a crucial role in bringing ethology as an accepted discipline into
existence, was clear about the fact that the first fundamental theoretical steps were due to
Lorenz.26 This section of my study will discuss what I perceive as the four key conceptual and
methodological commitments as to how to study and explain behavior that Lorenz—unlike
several other researchers—endorsed. Not all of these aspects of Lorenz’s general approach were
fully present from the beginning of his intellectual development. But they emerged relatively
soon; and once they were fully formed and spelled out, they strongly guided Lorenz’s further
theoretical development and guided it into a certain direction. In the following section I will show
how these four factors help us to understand why Lorenz ended up with the strong tenet that
instinct and experience are exclusive and that instinctive behavior does not evolve into more
flexible behavior. Because of the crucial importance of these four aspects, the biographical and
intellectual influences that relate to this level of Lorenz’s theoretical development are the most
interesting ones and will be highlighted in the following discussion.

1. The primacy of the innate
As is well-known, Lorenz approached the study of behavior from a zoological rather than a
psychological point of view. On his view, animal behavior was the model for human behavior. In
fact, Lorenz assumed that we do not need a human psychology that works independently of
zoology and animal psychology. But many animal psychologists did not necessarily disagree with
this. What was more peculiar to Lorenz is his emphasis on innateness. While it was standard
practice among animal psychologists to make the distinction between innate and learned behavior
and to recognize the existence of instincts, most animal psychologists stressed the importance of
learned and intelligent behavior.27 Lorenz disagreed with many animal psychologists in that he
claimed the innate rather then learned as the core of behavior. He argued that the focus in the
study of animal behavior has to be on the instinctive instead of the insightful. Lorenz claimed that
the study of intelligence makes sense insofar as the set of innate behavior patterns is known and
previously understood. A consequence of this focus is that while Lorenz’s early theory was good
at analyzing innate behavior and what he termed instincts, it left him without a real framework
for conceptualizing and explaining the various behavioral phenomena that involved learning. The
Dutch animal psychologist Bierens de Haan—whose approach I will discuss in more detail
later—complained about Lorenz’s notion of appetitive behavior that it lumped together the many
quite distinct behavioral processes in which organisms react flexibly to their environment, from
orienting movements in amoebae to insightful behavior in primates.28
If we take a look at Lorenz’s views of human behavior and morality, we can understand that
his commitment to the primacy of the innate was not just a methodological preference but
fundamental to his views about the nature of behavior. From 1939 onwards a new topic appeared
in his publications. Lorenz started to discuss human behavior, in particular human moral
behavior, from the point of view of instincts, and he famously—or notoriously—warned against
the genetic degradation of moral behavior. Lorenz crucial assumptions was that important
features of human behavior such as ethical and aesthetical values are based on instincts. He
viewed the process of human civilization in analogy to the domestication of animals, and argued
that in both types of processes the original functional instincts get lost, leading to a step-wise
disruption of behavior and a degradation of social behavior in the case of humans. During the
Nazi period, Lorenz used this account in order to receive popularity and further his career.29 As
Lorenz’s theory stressed the dangers of the accumulation of morally and genetically inferior
people within a society or ethnic group, his account diverged somewhat from Nazi racial theories,
which focused on alleged differences between ethnic groups. But Lorenz still tried to appeal to
eugenic thinking in Nazi Germany and Austria. Even after the war, Lorenz continued to warn
against genetically based moral degradation (whereas nowadays even the assumptions about
genetics and evolution underlying this view are taken to be fundamentally misguided).
The issue of the social consequences of domestication was addressed in the correspondence
with Heinroth not before March 1938,30 and stressed in Lorenz’s publications and public talks
only from 1938 onwards.31 But one ingredient of this view was present in Lorenz’s thinking at an
earlier stage—namely the idea that human moral and aesthetic values are inherited features
encoded in instincts. Already in a letter from 1933 Lorenz clearly expressed the conviction that
an action counts as moral only if it is done instinctively, but not if it is based on reasoning. He
points to Oskar Heinroth as another person who views morality as an instinct:
“If something good and decent is not done for pleasure … and purely instinctively, then it immediately
loses the characteristic of a ‘moral’ act, … That is, my social reaction of gratitude responds only if ‘the
other’ showed accommodating in an instinctive fashion (acted ‘on his own’, ‘for love’). As soon as he
considers or reflects about it, he will at best receive a letter of thanks, but no gratitude. …
Unfortunately, Heinroth’s doctrine of ‘morality’ as a real complex of instincts goes against the grain
with so many people; for them it appears to be cynical and to degrade the nature of morality.”32
Thus, Lorenz viewed instincts as the basis of human behavior, in particular human moral
behavior. He was convinced that the innate is the core of behavior.
Heinroth was a crucial influence as far as this first factor of Lorenz’s perspective is
concerned. I already pointed out that Heinroth’s work was not really of a theoretical nature, and
that he—unlike Lorenz—did not attempt at offering his definition or theory of instinct. But
Heinroth’s work clearly exhibits a certain approach to the study animals and their behavior.
Heinroth was a supporter of Lorenz in that his research focused on instincts and innate behavior
rather than on learning. In fact, his hand-rearing of young birds often occurred in isolation from
conspecifics, so that Heinroth systematically conducted deprivation experiments that help to
discern innate features of bird behavior.33 Apart from the focus on the innate and the instinctive,
Heinroth was convinced—just like Lorenz—that humans have many more instincts than is
usually acknowledged and that the study of human instincts is crucial for understanding human
social behavior.34 As the above quoted letter from Lorenz shows, Heinroth viewed morality as
based on instincts.
2. The need for physiological explanations
The second factor guiding Lorenz’s approach was the idea that a biological rather than
psychological explanation of instinctive behavior was needed. According to Lorenz, the
physiological, not the mental is the core of behavior. Behavior is to be explained by means of its
underlying physiological causal basis, but not by means of invoking psychological drives and
subjective motivations of behavior. Animal psychologists with a purposivist approach such as
McDougall or Bierens de Haan emphasized the goal-directedness of behavior and made recourse
to subjective experiences and desires that motivate behavior. Even a zoologist such as Julian
Huxley assumed that an understanding of animal minds is a crucial factor in behavior studies; for
instance, he argued that we need to appeal to psychological factors to explain courtship behavior
in birds. For Lorenz, however, the goal-directedness was not an explanation of behavior, it itself
needs to be explained by causal factors.35 It is the focus on causal-physiological explanation that
Tinbergen appreciated so much about Lorenz’s approach.36 Lorenz’s and Tinbergen’s emphasis
on a causal-physiological study and explanation of behavior proved to be important for the
success of ethology as a biological approach to behavior.
Until 1937 Lorenz had certain sympathies with the chain reflex theory of instinct, which
viewed instincts as sequences of reflexes. One reason for this was Lorenz’s physiological
approach to the explanation of instinctive behavior. For a reflex is caused by an external stimulus
based on clear-cut neurological pathways and physiological factors—mental states of the animal
or not needed to explain reflexes. On Lorenz’s physiological approach, physiological triggers are
more of a model for explaining instinctive behavior than psychological factors. This is why
Lorenz deemed the chain reflex theory as a potentially useful way to think about instinctive
behavior. In the second instinct paper from 1937, Lorenz fully abandoned the chain reflex
theory.37 The change of view taking place in this paper was the shift from regarding instinctive
behavior as elicited by external factors to a position that includes endogenous factors in the
production of instinctive behavior. Lorenz’s tenet that the performance of an innate behavior
pattern is carried out in a stereotypic fashion and thus looks like a reflex fits with the chain reflex
theory. But the reflex theory assumes that an instinct—like any reflex—is fully caused by an
external stimulus. Now Lorenz for the first time argued that factors internal to the organisms play
an important role in the production of instinctive behavior. The idea is that if an instinctive
behavior pattern is not released for some while, then nervous energy can build up within the
organisms and lead to appetitive behavior (the animal actively searching for the releasing
stimulus) and finally to vacuum activities. It is well-known that this important theoretical shift
was solely due to the influence of the neurophysiologist Erich von Holst, a student of Albrecht
Bethe.
Theodora Kalikow has discussed Lorenz’s commitment to the chain reflex theory.38 Her
account that Lorenz endorsed the reflex theory, in particular Ziegler’s version, is probably right.
However, Kalikow does not have real evidence for her claims and instead relies too much on
Lorenz’s post-war account according to which he “was a firm adherent of classical
Sherringtonian reflex theory.”39 In addition, Kalikow’s brief discussion simply portrays Lorenz
as endorsing the chain reflex theory until he abandoned it in 1937 under the influence of von
Holst, ignoring any previous changes in Lorenz’s stance towards the reflex theory. It is true that
Lorenz himself acknowledged in 1942 that he “still adhered to the Ziegler theory of instinctive
motor patterns in its customary form in 1937”,40 but my impression is that Lorenz’s relation to
the chain reflex theory was not always as clear and unambiguous as he later asserted. In any case,
it is admittedly hard to settle this issue because in his early papers Lorenz’s does not mention the
chain reflex theory, and when he addresses it from 1935 onwards, he already acknowledges that it
cannot account for all behavioral phenomena.
Let us take a closer look at this matter. Lorenz’s first theoretical paper from 1932 was clearly
influenced by the work of Heinrich Ernst Ziegler (1858–1925), a German physiologist endorsing
the view that instincts are sequences of reflexes. Lorenz explicitly stated that he uses Ziegler’s
basic definition of instincts: “I have defined the difference between instinctive and intelligent
activities in the following way: the former are based upon inherited pathways and the latter on
individually-acquired pathways. The psychological definition is thus replaced by a histological
definition.”41 Whereas some animal psychologists characterized instincts by the absence of

insight or other subjective factors, using Ziegler’s physiological definition of instincts Lorenz
could define instinctive as well as intelligent behavior without any reference to mental factors.
However, apart from this definition of instincts, Lorenz did not endorse the chain reflex theory of
instincts. In fact, in the 1932 paper he did not even mention this theory or other aspects of
Ziegler’s work.42 Despite Ziegler’s partial influence on Lorenz, the latter did definitely not
endorse Ziegler’s whole theory. For instance, Ziegler assumed the existence of incomplete
instincts, which need some practice or even experience. Incomplete instincts are a kind of drive
or innate disposition for a type of behavior rather than a rigid reflex that is performed in the same
manner from the very beginning.43 This was surely incompatible with Lorenz strong innate-
learned dichotomy, and already in his 1932 paper Lorenz heavily emphasized the inflexibility and
rigidity of instincts and states at a few places that an innate behavior pattern is performed in a
complete manner on its first performance.44
Lorenz’s 1932 paper actually did not mention the chain reflex theory of instincts, but it was
addressed in subsequent publications. However, while the ‘Companion’ stated that instincts are
chain reflexes,45 it also mentioned limitations of Ziegler’s definition of instinct.46 In his first
instinct paper Lorenz said that even though he presents himself “with some reservation” as a
supporter of the reflex theory, “this does not mean that [he] regard[s] this as a particularly
promising working hypothesis”.47 The reasons for this is that a reflex approach cannot account
for certain phenomena. Vacuum activities were very important for Lorenz because they show that
the same stereotypical behavior pattern is executed independent of its purpose. But as a vacuum
activity occurs without the usual releasing stimulus, it is not a like a reflex. Craig’s notion of
appetites, that Lorenz acknowledged, refers to behavior without a clear-cut external stimulus as
well. To be sure, the fact that Lorenz never explicitly endorsed the reflex theory does not mean
that he did not take it serious. Lorenz probably viewed it as the best available starting point,
despite many unsolved issues.
Kalikow views Lorenz commitment to the chain reflex theory stemming from his
mechanistic, anti-vitalist view of nature.48 ‘Mechanism’, however, is a problematic label for
Lorenz’s point of view. In the early 30s, Lorenz might have been a mechanist in the sense that he
was reluctant to accept the relevance of subjective experience for the explanation of behavior.49
But later he seemed to accept the relevance of emotions and subjective states in animals, thus
agreeing more with his mentor Heinroth. Heinroth countered the claim from his lay readers that
he was viewing animals as mere reflex machines by stating that in fact he likened animals to
human persons that are extremely emotional and of low intelligence. Independent of what Lorenz
thought about emotions in animals, he was definitely not a mechanist as some proponents of the
chain reflex theory were. The latter did neurophysiological work in laboratories and assumed that
reflexes and instincts are performed by certain organs only. Lorenz, as an observer of animals in
a relatively natural environment, was aware of the fact that instinctive behavior involves the
whole organism. Lorenz emphasized that we have to break down and analyze phenomena, but
unlike mechanists he still had the whole organism and its environment in view.50 Moreover,
proponents of the chain reflex theory were usually not interested in the purposiveness and
adaptedness of behavior, which they associated with vitalist schemes of explaining behavior by
reference to unanalyzable forces that make animals develop and behave goal-directed. Lorenz,
however, was an evolutionary thinker and explicitly viewed the adaptedness as something that
has to be studied and explained, albeit by evolutionary principles such as natural selection. This
difference from strong mechanistic positions was due to Lorenz’s research practice as a whole
organism biologist, and because of this research practice and different perspective the orthodox
chain reflex theory could not have been fully congenial to Lorenz.51
I offered some challenges to the standard assumption that Lorenz was a firm and
unambiguous adherent of the chain-reflex theory of instinct until 1937. What I want to
emphasize, however, is that the real point of my discussion of Lorenz’s commitment to the reflex
theory is actually to shift the historiographic focus away from the reflex theory issue. Based on
the work of von Holst, Lorenz ultimately abandoned the reflex theory as a whole. But my
discussion is about stable and thus more telling features of Lorenz’s perspective. An example of
this is his strong innate-learned dichotomy. Even though von Holst’s work brought about a
change in Lorenz’s overall theory, Lorenz’s views about innateness and the inflexibility of
instincts were unaffected by this. Lorenz never wavered regards the innate-learned dichotomy or
abandoned this tenet; this is the reason why I view it as a much more important and influential
feature of Lorenz’s thought. This also applies to my four factors as they are stable features
guiding Lorenz’s theoretical development. Relevant for this context is Lorenz’s second
commitment, i.e., the idea that we need a neurobiological or physiological explanation of
behavior rather than a psychological explanation based on subjective factors. Lorenz’s preference
for the reflex theory was obviously driven by this commitment, but the later endorsement of von
Holst’s theory as well. Now Lorenz assumed that endogenous factors play an important role in
the production of behavior. But Lorenz is clear about the fact that these internal factors are not
psychological drives but neurobiological stimuli—von Holst’s work was purely
neurophysiological. The psychohydraulic model that was proposed in this period relies on a
mechanical analogy (gas, water and pressure) to illustrate the causation of behavior. Lorenz’s
commitment to the second factor—the physiological approach—was unaffected by the shift
towards von Holst’s ideas.
There is a received account as to how Erich von Holst convinced Lorenz that the reflex theory
has to be fully abandoned. Lorenz met von Holst the first time in February 1936, when he gave a
talk about an early version of the first instinct paper. After the war, Lorenz stated that von Holst
convinced him immediately at this meeting that the reflex theory was wrong.52 Even though the
history is often recounted in this manner,53 Lorenz remembered incorrectly. Actually, it took
Lorenz more than a year to fully appreciate von Holst’s ideas and figure how to modify his
instinct theory so as to neatly integrate von Holst’s insights. Even though Lorenz and von Holst
did meet in February 1936, the first instinct paper did not mention von Holst’s ideas, while the
final version of this paper went to the publisher in September 1936.54 As late as April 1937 von
Holst criticizes Lorenz’s draft of the second instinct paper, because it still uses ideas and
terminology of the reflex theory, such as Lorenz’s formulation that an instinct is an intended
reflex. Von Holst points out that vacuum activities are much better explained by the endogenous
production of behavior rather than Lorenz’s idea that they are reflexes that can be caused by an
unspecific external stimulus.55 Based on von Holst’s critique, Lorenz is revising his draft of the
second instinct paper for several weeks. Indeed, the editor of Folia Biotheoretica, van der
Klaauw, complains about Lorenz constantly reworking the manuscript rather than sending him
the latest version, because the journal issue has to be printed.56 But after receiving the final
version at the end of May 1937, van der Klaauw states that he is glad that Lorenz was able to
include his new perspective based on von Holst’s results.57 Already while reworking his
manuscript, Lorenz tells the ornithologist Erwin Stresemann about the profound changes in his
views that von Holst’s ideas made necessary.58 Given the fact that Lorenz met von Holst in early
1936, one may wonder whether von Holst attempted to convince Lorenz of the falsehood of the
reflex approach to instincts at this meeting already. If yes (but this is unclear),59 then it is not easy
to answer why it took Lorenz more than one year to appreciate von Holst’s points.60
Let us return to the main topic of this section—the need for physiological explanations of
behavior as the second factor of Lorenz’s perspective. An obvious influence on the early Lorenz
was the neurophysiological tradition stemming from the work of Sherrington that viewed
instincts as reflex chains. The representative of this tradition with the largest influence on Lorenz
was the already mentioned Heinrich Ziegler, by virtue of his 1910 book which deals explicitly
with theories of instinctive behavior and defending a reflex approach of instincts. But the aspect
of Ziegler’s influence that is of interest for me is not the fact that Lorenz’s 1932 paper took over
Ziegler’s definition of instinct (which characterizes instincts as based on innate rather than
acquired neural pathways). The for my purposes crucial influence attaches to the second aspect of
Lorenz’s perspective. What Ziegler’s work showed to Lorenz was that it is possible to have a
potentially fruitful research approach to instinctive behavior that is based solely on
neurophysiological factors and does not make recourse to psychological explanations.61 Another
likely intellectual influence was Lorenz’s university education. Lorenz started out studying
medicine and he passed his first doctoral degree in medicine before studying zoology. Due to his
medical training Lorenz learned about physiology and probably got acquainted with the reflex
physiology tradition in this manner. The medical training helped Lorenz adopting a physiological
approach to the causation of behavior.
3. The taxonomic and phylogenetic approach to behavior
The third and most important feature of Lorenz’s approach was his deep commitment to a
comparative, taxonomic or phylogenetic approach to behavior that drove both his research
practice and his theoretical development. Sometimes classical ethology is characterized as the
comparative study of behavior. But the label ‘comparative’ can fail to express what was peculiar
to Lorenz’s approach. Traditional comparative psychology was comparative in that animal
psychologists studied the behavior of different species from a variety of taxonomic groups. But
Lorenz’s approach meant much more than dealing with different groups of organisms. Lorenz
viewed instinctive behavioral patterns like morphological characters that can be identified in
different species. In other words, Lorenz homologized behavioral features; in fact, the notion of
behavioral homology is a type of homology stemming from classical ethology. Instincts as
taxonomic characters can be used to classify taxonomic groups and study their phylogeny and
evolution. This is the reason that I prefer the label taxonomic or phylogenetic approach rather
than comparative approach. Lorenz made the same point in a post-war paper. In the section that
has the telling title “Comparative anatomy of behavior”, Lorenz states that he does not need to
explain to biologists what the concepts of ‘comparative’ anatomy and physiology refer to, while
psychologists use the term ‘comparative’ in a “very loose sense to all behavior studies concerned
with different forms of life”. Lorenz resents “when an American journal masquerades under the
title of ‘comparative’ psychology, although … no really comparative paper has ever been
published in it.”62
The taxonomic-phylogenetic perspective was a crucial part of Lorenz’s approach from the
very beginning. His publications clearly show that Lorenz followed this research method. His
first larger publication from 1931 contained detailed observational studies of different species
from one taxonomic family of birds. Later publications offered an explicit theoretical account of
the nature and importance of this method. The second instinct paper, for instance, started out with
the taxonomic relevance of instincts, before discussing their development and physiology.63 The
taxonomic approach is the reason why Lorenz viewed instincts in analogy to morphological
structures. Lorenz emphasized that both are inherited and often have a long phylogenetic history.
They can be evolutionarily conservative, so that they serve as taxonomic characters that allow to
characterize and classify biological taxa.64 In fact, Lorenz claimed that instincts are sometimes
more conservative than morphological features.65 Tinbergen learned about this taxonomic (rather
than just comparative) approach to behavior from Lorenz.66

The reason why the taxonomic-phylogenetic approach was such a crucial feature of Lorenz’s
approach is that it was strongly embedded in his research practice. The research practice of
Lorenz and other behavioral researchers has been discussed in detail by Richard Burkhardt.67
Lorenz’s practice resembled that of Charles Otis Whitman. Neither Whitman nor Lorenz were
field naturalists. Like Lorenz, Whitman kept many birds—primarily pigeons in the case of
Whitman—in close proximity to his home so that he could observe them.68 Both made
comparative observations on closely related species that allowed them to gain insights into the
phylogeny and evolution of behavior. Compare the taxonomic practice of Lorenz with some of
his theoretical claims such as the definition of instinct taken over from Ziegler. Ziegler’s
definition is based on the difference between innate and acquired neural pathways. However,
while Lorenz endorsed this neurobiological definition for some time, it never had an impact on
his research practice. Ziegler and other neurophysiologists made neurobiological experiments and
observations. Lorenz, on the other hand, stuck to observing living animals in a relatively natural
environment rather than investigating their neurophysiology. The taxonomic-phylogenetic
approach, in contrast, was part and parcel of Lorenz’s research practice. What Lorenz did in his
ethological research every day was to observe animal behavior from a comparative, taxonomic,
and phylogenetic point of view.
A crucial influence on Lorenz in this respect was his university education. As already
mentioned, Lorenz studied medicine, focusing on comparative anatomy, and his first doctoral
degree was in comparative anatomy. Apart from anatomy at the medical school, zoology at the
University of Vienna was dominated by comparative morphologists and embryologists. Lorenz
was still teaching anatomy classes when writing on the ‘Companion’. Lorenz’s primary
competence was in comparative anatomy above all other traditional zoological fields. Another
influence or support came from Oskar Heinroth. Even before Lorenz, Heinroth homologized
behavioral patterns and used them as taxonomic characters.69 Lorenz probably developed his
research practice on his own, but he soon realized that Heinroth’s research had made use of a
similar practice. Heinroth’s published work and the correspondence with him encouraged Lorenz
in pursuing the taxonomic perspective on behavior. Lorenz acknowledged this support by
Heinroth, in particular his comparative-taxonomic approach, on many occasions. In a paper after
the war, for instance, Lorenz cites Heinroth and Whitman as forerunners of the taxonomic
approach, and of ethology in the first place. Expressing in fact his own point of view, Lorenz
states that given Whitman’s and Heinroth’s perspective (which differed from the approach of
their contemporaries), it was actually no surprise that they avoided the pitfalls of former
approaches and developed some early insights of ethology. He continues:
“No very great tribute either is due to them for applying the phyletic method to the study of innate
behaviour. Being comparative morphologists, it was only natural for them to do so. But by doing it,
they discovered a fact which could not be discovered from any but the phyletic point of view; …”70
Thus my suggestion is that we have to view Lorenz primarily as a comparative anatomist and the
taxonomic approach as the main factor guiding his theoretical development.
4. The embryological approach to behavior
The last factor concerns the way in which Lorenz approached the development of behavior.
Lorenz used ideas from embryology and the developmental biology of morphological structures
to think about the ontogeny of behavioral features. However, this feature was not as influential as
the other theoretical and practical commitments driving Lorenz’s perspective, and it was not
always clearly and consistently present in Lorenz’s account. This is due to the fact that Lorenz
focused on the phylogeny and the evolutionary adaptation of behavioral characters, but was never
really interested in studying and explaining the development of behavior—unlike behavioral

biologists such as Daniel Lehrman. But when Lorenz had to address the development of
behavior, he used ideas from experimental morphology.
The embryological approach fits with Lorenz’s general idea that instincts are like
morphological structures and that instincts are to be studied from a causal-physiological point of
view. Lorenz was not the first to use the analogy of instincts and organs.71 But he pushed the
analogy further by using the idea of maturation to support the claim that instincts are not
modified in ontogeny under the influence of experience. On his account, even though there are
cases where the performance of instincts changes during early ontogeny, this just reflects the fact
that instincts as well as their underlying morphological and physiological basis have to maturate,
i.e., instinct have to fully develop in the first place—just like organs. This line of thought
permitted Lorenz to maintain that change in instinctive behavior during ontogeny need not be due
the influence of learning or experience; a change in a behavioral character is compatible with it
being completely innate. Lorenz stressed this point to rebut alleged evidence that was invoked to
show that innate instincts can be modified by learning.72
Sometimes Lorenz used more specific embryological ideas, such as theoretical concepts from
developmental mechanics. Lorenz’s account of imprinting shows this. The notion of imprinting
was introduced in the 1935 ‘Companion’ paper. A bird ‘recognizes’ its parents as follows. When
a young bird hatches from its egg it will be imprinted to the animals it encounters immediately
after hatching (normally its parents), i.e., it will subsequently show any behavior that is normally
directed at its parents towards the animal to which it was imprinted. If for example Lorenz was
the first living being that the hatched bird encountered, then the bird would behave towards
Lorenz as if he was one of its parents. A bird has to be imprinted to its parents and thus needs to
acquire the external information (to ‘learn’) what its parents are. Lorenz, however, denies that
imprinting is actually a learning process . He points out that imprinting can be carried out only in
a very short period of development and that it is irreversible—unlike standard learning processes.
In fact, Lorenz assumes that imprinting is like the process of embryological induction or
determination.73 This is a well-known concept from the German tradition of developmental
mechanics (the Spemann school) used to account for the development of specific morphological
structures by means of stimuli from adjacent tissues. As a nice illustration of how Lorenz used
the framework of experimental embryology to think about the development of behavior, consider
a margin note that Lorenz made in his copy of an article by Otto Koehler (Koehler, 1933).
Koehler’s essay deals with the idea of wholeness in biology; and at one point Koehler describes
an instance of embryological induction by means of a dead organizer, i.e., non-living matter
derived from animal tissue (p. 182). Lorenz annotates on the margin of his reprint “Prägung!”
(Imprinting!). It is intriguing that this margin note is about a dead organizer, because a bird can
be imprinted not just to other animals, but to inanimate objects as well.
In sum, even though Lorenz was not primarily interested in the development of behavior and
did not have a coherent theory of the ontogeny of behavioral features, in order to defend his
views of instincts Lorenz thought along the lines of a kind of behavioral embryogenesis, using
ideas from developmental mechanics.
Lorenz’s early views on instinct and innateness
The four aspects of Lorenz’s perspective are quite constant features of his approach. Taken
together, these factors guided and constrained his early theoretical development. This holds in
particular for the formation of Lorenz’s instinct concept and his innate-learned dichotomy, so that
these features that were so characteristic about Lorenz’s theoretical position and that
distinguished him from other animal psychologists can be understood based on the four factors.74
Before discussing in detail how Lorenz’s perspective furthered his innate-learned dichotomy, I
want to illustrate the explanatory relevance of the four factors by comparing Lorenz with a quite
different tradition—the Dutch purposivists. From the late 30s onwards, the proponents of the
ethological approach (Lorenz, Tinbergen, and their supporters) and the Dutch animal
psychologists working with the more traditional purposivist perspective criticized each other
vehemently.75 In my view, one important factor for this development was institutional. Lorenz
and Tinbergen attempted to create a new biological discipline, which required emphasizing the
difference between their discipline and other ‘non-biological’ approaches to animal behavior. But
the fundamental disagreement was based on differences in perspective as well.
I want to focus on a brief comparison between Lorenz and Johannes Abraham Bierens de
Haan (1883–1958), the most influential animal psychologists of the Dutch purposivists tradition
at the time. When Lorenz’s first publications appeared, Bierens de Haan was already a well
established researcher and theorist. Bierens de Haan—a student of F.J.J. Buytendijk—was
fundamentally influenced by William McDougall’s psychology, adopting and developing his idea
of instincts as innate drives motivating behavior.76 Bierens de Haan viewed an instinct as the
drive or striving to behave in a certain manner given certain subjective perceptions.77 This was in
direct contrast to Lorenz, who defined instincts as particular motor patterns, i.e., bodily
movements that can be filmed, rather than subjective features of the psyche such as mental
propensities. This disagreement as to how to use the term ‘instincts’ was not simply a
terminological matter. It was a difference in view as to what is the important about animal
behavior, what is the nature of behavior, and how to explain it.
Bierens de Haan stressed the way in which instincts are modified in ontogeny under the
influence of experience. In fact, one of his books, which states his mature theory, is entitled
Animal Instincts and their Rebuilding by Experience.78 On his account, the inflexibility that the
performance of some instincts exhibits is not due to the innateness of instincts—as Lorenz would
have it. Quite on the contrary, habit and the repeated use of certain behavior patterns brings about
stereotypic and potentially unadaptive behavior patterns.79 It is interesting to note that Bierens de
Haan very well agreed with some of the important observations and arguments of Lorenz, such as
Lorenz’s point that we need to distinguish change of behavioral performance due to maturation
and the influence of learning. But Lorenz and Bierens de Haan still offered very different
interpretations because the point of agreements are embedded in very different theoretical
perspectives. Bierens de Haan accepted the idea that there is a morphological as well as
physiological aspect of behavior. However, he stated that he focuses on the psychological study
of behavior only,80 whereas for Lorenz psychological aspects such as subjective feelings only
make sense as part of a biological approach to behavior. Bierens de Haan acknowledged the
existence of reflexes and reflex-like behavior. But his focus was on intelligence, learning, and the
goal-directedness of behavior.81 In fact, he claimed that the reflex is about causation, while the
instinct is about teleology (the goal-directedness), so that instincts and reflexes cannot possibly
be identified.82 Directly criticizing Lorenz, Bierens de Haan argued that we cannot just equate
instincts and instinctive behavior patterns. In his view, motor patterns are only the expression of
instincts, while instinct is an explanatory and psychological concept referring to subjective
factors motivating behavior. The intention of the individual is the fundamental feature of
instinct.83 Bierens de Haan stressed the continuity between the human and animal psyche. The
mental is a feature that emerged with the origin of animals. Instincts were viewed as a
specifically psychological properties of an individual, which were contrasted with biological
properties.84 A fundamental tenet of Bierens de Haan was that objective (biological) issues such
as behavior patterns have to be distinguished from subjective (psychological) issues such as
instincts.85
Overall, Bierens de Haan disagreed with each of the four aspects of Lorenz perspective. He
acknowledged the existence and importance of instincts and innate behavioral characters, but his
theoretical focus was on learning, intelligence, and the way in which innate features are
embedded in and modified by subjective experience. And he did not view innate instincts as
something that is to be explained by evolution and natural selection. Bierens de Haan vehemently
rejected Lorenz’s preference for causal-physiological explanations of behavior by stressing the
necessity of psychological explanations and the existence of innate drives and subjective desires
motivating behavior. I emphasized that I view the taxonomic-phylogenetic approach to behavior
as the main factor driving Lorenz’s theoretical development. Bierens de Haan, however, did not
think much of Lorenz’s taxonomic approach to behavior.86 He also did not use models from
embryology or experimental morphology to explain the development of behavior. Due to the
fundamental difference of perspective the debate between Lorenz and Bierens de Haan led to a
clash. In 1942, Bierens de Haan restates the critique of Lorenz and makes some polemical
remarks. Lorenz strikes back, vigorously attacking Bierens de Haan’s general purposivist and
vitalist approach rather than addressing the interesting and critical points that the latter raised.87
Now that I have illustrated the difference between the perspective of Lorenz and more
traditional approaches in animal psychology, I finally turn to the development of Lorenz’s
instinct concept. The four aspects of Lorenz’s perspective make intelligible why he came to hold
this particular instinct concept and the strong innate-learned dichotomy. Lorenz’s views on these
matters emerged very soon in Lorenz’s intellectual development, and remained non-negotiable
claims. Lorenz never wavered on his tenet that innate instincts and learned behavior components
are different in nature and are clearly separate. In my view, the main reason for this is that
Lorenz’s taxonomic approach his the main factor guiding Lorenz’s views on instincts, and the
taxonomic approach is part of Lorenz’s research practice from the very beginning.
First, there was Lorenz’s view that the concept of ‘instinct’ ought to refer to motor patterns—
observable bodily movements rather than drives or disposition to behavior or even certain
subjective states as many other researchers assumed.88 Lorenz’s taxonomic approach precisely
suggested his account or definition of instinct. Once one views instincts as taxonomic characters
or homologues, instincts have to be features that we can identify when observing the behavior of
an organism. In addition, they have to be features that we can find in different species; thus they
have to be relative small units of the overall behavior of an individual. As in the case of
morphological structures, we have to break down an organisms into smaller modules or
homologues that we can compare with corresponding features in other species. In his daily
observations of birds from closely related species, Lorenz could observe and study instincts in his
sense of motor or behavior patterns. Even though Lorenz originally developed this taxonomic
research practice on his own, before him Lorenz’s scientific mentor Oskar Heinroth had dealt
with behavior based on a taxonomic-phylogenetic framework. In fact, just like Lorenz he focused
on characteristic movements of certain parts of the body.89 Lorenz obtained crucial support in
pursuing this approach to the study of instinctive behavior. Apart from the taxonomic-
phylogenetic aspect of Lorenz’s perspective, his view that instincts are in fact motor patterns fits
perfectly with his focus on innate (as opposed to learned) behavior. Motor patterns are also
amenable to experimental study and a within the scope of physiological theories, so that instincts
in Lorenz’s sense are likely candidates for successful physiological explanation, in accordance
with his views as to how behavior has to be explained.
Second, apart from Lorenz’s definition of instincts, there was his peculiar and controversial
claim that there are no ontogenetic transitions between innate and learned behavior components.
Unlike virtually anyone else, Lorenz assumed that instincts are not modified by experience at all.
Lorenz’s view of instincts as motor patterns—driven by his four commitments and in particular
the taxonomic approach—actually supported the idea that innate instincts are stereotypic and
inflexible. The behavior patterns that are characteristic of a species are quite rigid and do not
appear to be much influenced by learning. Based on the instinct-organ analogy and
embryological ideas Lorenz could put forward his notion of maturation. Pointing out that
maturation processes are not to be confused with real learning processes, Lorenz could argue that
instincts in his sense show full performance once they are developed and maturated, and are not
modified by later learning. Using the notion of instinct-conditioning intercalation, he could
account for the overall flexibility of behavior without abandoning the idea that the particulate
components he calls instincts are rigid.
Lorenz’s taxonomic-phylogenetic approach was the main force driving his notion of instinct,
and thus an indirect cause of the idea that instincts are independent of experience. But there is a
more direct bearing of the taxonomic point of view on Lorenz’s strong innate-learned dichotomy.
Lorenz assumed that the fact that instincts can serve as taxonomic characters and are often
evolutionarily conservative is incompatible with them being modifiable. Linguist who currently
use methods similar to the ones from taxonomy and molecular phylogeny to reconstruct the
relationship and evolution of different human languages may be surprised about the claim that
anything that is inherited over many generations and can serve as a taxonomic character cannot
be learned. But Lorenz clearly made this assumption, and it was an important idea driving his
innate-learned dichotomy. He made this point in several publications. It appeared for the first
time in the ‘Companion’:
“Nobody can deny that the phylogenetic mutability of an innate behaviour pattern possesses exactly
the same characteristics as an organ and does not resemble that of a learning function. Its mutability is
so similar to that of a particularly ‘conservative’ organ that the instinctive behaviour pattern actually
carries greater weight as a taxonomic feature, …”90

Both the first and the second instinct paper repeated this argument in a more explicit form.91 The
clearest expression of this idea can be found in a book manuscript that Lorenz wrote during the
war:
“If highly complicated behavior patterns are reliable, phylogenetically interpretable characters of
species, genera, and orders, like any morphological characters, then this fact alone is enough to
demonstrate that these behavior patterns cannot undergo substantial modification through individual
experience, as has been assumed by Spencer, Lloyd Morgan, and others.”92
Finally, Lorenz had a bold view about how instinctive behavior changes in the course of
evolution. He maintained that it is impossible for an instinct to evolve gradually into experience-
based and intelligent behavior. More precisely, an instinct is never homologous to a flexible
behavior component. This position contradicted all former and contemporary accounts. An
interesting proponent of the more traditional view was the zoologist Charles Otis Whitman.93 In
his influential 1898 lecture, Whitman discussed animal behavior and its evolution based on
detailed observations.94 He argued in particular against the view that instincts are lapsed
intelligence or derived from acquired habits, a position on animal behavior quite popular at that
time and sometimes based on Lamarckist assumptions. He rejected the idea that instincts are
relatively recent features, which originated much later than morphological structures. Rather than
becoming later integrated into the given morphology and hereditary make-up of organisms,
instincts evolved form the very beginning together with their anatomical structures. But even
though Whitman—like Lorenz—had an explicitly biological and comparative-phylogenetic
approach to behavior, he assumed that instincts can evolve into more flexible behavior, agreeing
with Herbert Spencer and Conwy Lloyd Morgan in this respect.95 In fact, Whitman’s lecture
starts with a quote from Darwin’s Origin, emphasizing the continuity between animal and human
cognitive features: “Natura non facit saltum, is applicable to instincts as well as to corporeal
structure.”96
How did Lorenz come to hold his peculiar view about the evolution of instincts? He offers a
different scenario on the evolution of behavior than other zoologists. On Lorenz’s account, the
importance of intelligence can increase only when the role of instinctive behavior diminishes.
This may happen if in an overall behavior sequence, consisting of several components, new
components that are based on learning are inserted. Thus instinctive behavior is broken up by
newly introduced flexible behavior components, reducing the amount of rigid instincts that are
part of the overall behavior. Alternatively, innate behavior pattern may be completely lost and
experience-based components fill the corresponding functional positions. The overall behavioral
sequence involving different components is modified, by the addition or deletion of specific
instinctive or experience-based components. Lorenz’s general picture is that instead of evolving
into more flexible components, instincts atrophy, are lost, and replaced by new and distinct
experience-based behavior components.97 This is the reason why Lorenz can claim that instincts
are not homologous to novel components that are based on learning. Lorenz described as early as
1931 the loss of instinctive patterns.98 This shows how important his early focus on particular
observations is for his later evolutionary account. Whitman, in contrast, used comparative studies
of wild, semi-domesticated, and domesticated pigeon species to support his view that instincts
become more plastic in the course of evolution. Lorenz does not really challenge Whitman’s
basic observations, but he offers a different interpretation of his results. Rather than becoming
more flexible, behavioral sequences involving instinctive components are broken up and
appetitive and condition-based components may be added.99 When Whitman states that
“instinctive action is gradually superseded by intelligent action”,100 Lorenz objects by annotating
his copy of Whitman 1899: “no! step by step”.

Apart from the fact that Lorenz has a different preferred hypothesis, why does he think that
the received scenario about the evolution of instincts is impossible? One consideration he offers
in support of his view is that instincts and insight-based behavior components appear to be
mutually exclusive, so that only one can fill the same functional role.101 The presence of an
instinctive component inhibits the development of intelligent components with the same
function.102 This idea is rather about the development of instincts, and it alone cannot establish
Lorenz’s evolutionary position. For this argument does show that an inflexible component cannot
correspond in the next generation to a less inflexible homologous pattern, which need not
presuppose that an instinct gets more flexible during ontogeny. More interesting is Lorenz’s idea
that instincts do not get replaced by more flexible components because they do the adaptive job
better than the intelligence of the corresponding animal permits.103 An annotation puts this idea
more strongly than we know from Lorenz’s publications. Whitman quotes a passage from Lloyd
Morgan expressing the idea that instinct evolve into more flexible patterns: “And this, if
continued, would tend to convert what had been a stereotyped instinct into innate capacity; that
is, a general tendency to certain activities (mental or bodily), the exact form and direction of
which are not fixed, …”104 Lorenz counters on the margin: “This would convert instinct into
nonsensical action.” So Lorenz seems to be convinced that instinctive behavior patterns are so
adaptive that softening them up would make them less adaptive. This claim can actually support
the idea that it is very unlikely that a flexible behavior pattern replacing a rigid instincts would be
maintained by natural selection. Nevertheless, the idea that rigid instincts hardly ever evolve into
more variable behavior components does not entail that such a thing never occurs, i.e., that an
instinct is never homologous to a flexible behavior component in the descendant.
Overall, Lorenz does not give a conclusive argument for his rejection of the idea that non-
instinctive behavior could be homologous to instinctive behavior patterns. As far as his own rival
evolutionary scenario is concerned, Lorenz does not offer a theory of the evolution of instincts
that is spelled out in detail and defended. This is probably due to the fact Lorenz was not really
an evolutionary theorist. He was an evolutionary thinker in that he defended ethology as the
biological, phylogenetic, and evolutionary approach to behavior (and proposed his evolutionary
epistemology), but he was not much engaged in discussions about the mechanisms of
evolutionary change. Lorenz surely addressed instinctive behavior from the point of view of
phylogeny and adaptation, but still the absence of substantial discussions about evolutionary
mechanisms in the early work of Lorenz is quite striking. This is on the one hand due to the fact
that Lorenz contrasted his theoretical ideas with other approaches in animal psychology, but did
not relate them much to contemporary debates about evolution. On the other hand, Lorenz
primary specialty was comparative anatomy, not evolutionary theory.105 Lorenz’s emphasis on
ethology as the biological, in fact, evolutionary science of behavior was to some extent part of his
rhetoric to create a new biological field.106 In this respect Lorenz was very much unlike
Whitman, who at end of the nineteenth century was at the forefront of discussions about
Darwinism and evolutionary theory. While Whitman was fundamentally concerned with the
relationship of evolution, development and heredity, Lorenz hardly ever acknowledged the fact
that characters (including morphological structures) show variation within a species; at any rate,
he definitely does not view the possibility of intra-specific variation as theoretically significant.
Instead, he stressed the idea that an innate character is either present or that it is completely lost.
This might strike the contemporary reader as an instance of what Ernst Mayr calls a ‘typological’
approach to species. Lorenz’s taxonomic approach was the main reason why he emphasized that
instincts are rigid and species-specific.
The four aspects of Lorenz’s approach help us somewhat to understand how Lorenz came to
hold his specific position on the evolution of instincts. For they promoted Lorenz’s views on how
behavior is composed and how it develops, which indirectly suggest his evolutionary position. On
Lorenz’s account, behavior is composed out of distinct units, some of which are instinctive
behavior patterns, others are conditioned reflexes, taxes, or appetites. Each component is either
innate or it involves experience. Given this framework, it is theoretically hard to see how an
instinctive behavior pattern could gradually evolve into a more flexible component. Innate motor
patterns obviously are not homologous to conditioned reflexes in the next generation. Thus
Lorenz’s overall conceptual framework suggests that instincts to not evolve into flexible
behavior.
Ultimately, it is quite unclear what actually motivated Lorenz’s views on the evolution of
instincts—whether it were certain arguments that Lorenz used to defend his position or whether it
were deeper factors such as his taxonomic approach. Lorenz does not have an elaborated theory
of the evolution of instinctive behavior and it was not important for his research practice to have
elaborated ideas about the mechanisms of evolutionary change; which makes it hard to assess the
roots of his position on the evolution of instincts. But independent of whether we can really
understand his claim that there are no phylogenetic transitions between instinct and learning, we
saw that what I called Lorenz’s four theoretical and practical commitments explain some crucial
aspects of his early theoretical development. The four factors make intelligible why Lorenz chose
to view instincts as nothing but innate motor patterns, and why he maintained that instincts are
inflexible and not influenced by learning. As I tried to show, the main driving force behind this
position was Lorenz’s use of instinct as taxonomic characters. If we want to understand why
Lorenz assumed that his research program might break down if he abandoned the view that there
are no transitions between instinctive and learned behavior, then we have to look at his
taxonomic-phylogenetic approach that was so deeply embedded in his research practice. The
early Konrad Lorenz has to be viewed as a comparative anatomist much more than as an
evolutionary theorist, which fits with his academic training. These features explain Lorenz’s
strong innate-learned dichotomy.
Acknowledgements
The research on this paper was funded by a Junior Fellowship of the Konrad Lorenz Institute for
Evolution and Cognition Research, Altenberg (Austria). I would like to thank the Konrad Lorenz
Institute and especially Werner Callebaut for the opportunity to work at the KLI and for the
access to the annotated books in Lorenz’s library. I am greatly indebted to Lorenz’s daughter
Agnes von Cranach for showing me some of his letters and permitting me to quote from them. I
would like to thank Chip Burkhardt, Paul Griffiths, Tania Munz, Robert Olby, and Jim Lennox
for comments on this paper.
Note on archival sources
The correspondence between Konrad Lorenz and Erwin Stresemann is stored at the
Staatsbibliothek zu Berlin, Preußischer Kulturbesitz (Germany) as ‘Nachlaß 150 (E. Stresemann),
Kasten 40’. Any letter from Lorenz that is part of this source is referred to in the paper as
‘Lorenz, letter to Stresemann’. Some of Lorenz’s correspondence is in the possession of his
daughter Agnes von Cranach and is referred to in the notes as the ‘Konrad Lorenz Family
Papers’. Annotations by Lorenz referred to in the paper are from the books in Lorenz’s library,
which is preserved in the library of the Konrad Lorenz Institute for Evolution and Cognition
Research, Altenberg (Austria).

Notes
1
Lehrman, 1953.
2
Lorenz, 1981, p. 261.
3
Lorenz, 1935.
4
For more details about Lorenz’s life see Taschwer and Föger, 2003; Burkhardt, 2004.
5
For a biography see Heinroth, 1971.
6
Heinroth and Heinroth, 1924–1933; see also Heinroth, 1911.
7
Heinroth and Lorenz, 1988 provides a posthumous publication of the whole correspondence from
1930 until 1940, even though some letters were lost (and political statements were removed from the
edited version).
8
This holds for Lorenz’s first theoretical paper from 1932 (Heinroth and Lorenz, 1988, p. 71; Lorenz,
letter to Stresemann, July 30, 1931) as well as the ‘Companion’ from 1935 (Lorenz, letters to Stresemann,
November 21, 1934 and August 3, 1935).
9
Heinroth’s response to Lorenz’s 1932 paper is lost, but a reply from Lorenz suggests that Heinroth
approved of Lorenz’s account (Heinroth and Lorenz, 1988, p. 77).
10
Burkhardt, 2004.
11
Behaviorism became an explicit enemy of Lorenz only after the war.
12
E. S. Russell is a zoologist who emphasizes the goal-directedness as a purely descriptive category,
but sometimes slides into a view that seems to invoke these goal as an explanation of the behavior
(Russell, 1934).
13
An example from the university of Vienna (but a generation before Lorenz studied there) was the
zoologist Karl Camillo Scheider. Schneider argued that in order to account for the behavior of any animal
(including amoebae) we need to assume an animal psyche and mental factors such as desires and drives
(Schneider, 1909).
14
Virtually all of Lorenz’s relevant publications in the period discussed are translated and reprinted in
the collection Lorenz, 1970.
15
Lorenz, 1937a.
16
Lorenz, 1937b.
17
Lorenz, letter to Stresemann, August 3, 1935.
18
Friedrich Hempelmann was a German zoologist writing on animal psychology at the time Lorenz
started his studies. Hempelmann was not a ‘purposivists’, he maintained that instincts grow out of
reflexes. But he still assumed that they are influenced by learning and that in the course of evolution there
is a gradual increase of flexibility and intelligence (Hempelmann, 1926).
19
Quoted from the translation Lorenz, 1970, pp. 116–117.
20
Lorenz, 1970, p. xviii.
21
Burkhardt, 1981.
22
Lorenz, 1996; Brigandt, 2003.
23
Ziegler, 1910.
24
On Lorenz’s account (which is credible), the Vienna psychologist Karl Bühler, with whom Lorenz
worked for some while when he decided to leave anatomy and pursue a career in animal psychology,
made him read this literature (Lorenz, 1970, p. xviii).

25
Lorenz’s ‘Companion’ made use of Craig’s notion of ‘appetite’ (An appetite is the state of unrest
that characterizes an animal as long as a certain desired stimulus is absent. This state of agitation makes
the individual search for the appeted stimulus. Craig, 1918). Craig wrote to Lorenz about his ‘Companion’
and first instinct paper: “Your paper, ‘Der Kumpan in der Umwelt des Vogels’ is by far the greatest paper
on instinctive behavior of birds I ever read in my life.” (Craig, letter to Lorenz, April 27, 1937, Konrad
Lorenz Family Papers). “You have solved so many problems while other observers (including myself)
have merely hacked at them.” (August 16, 1937). “To a great extent, your papers have persuaded me that
your views are correct and that my former views were wrong.” (September 19, 1937). So Craig’s work on
appetites did not only have impact on Lorenz, but conversely Craig adopted some of Lorenz’s ideas.
However, based on empirical examples Craig did not accept Lorenz’s absolute distinction of appetitive
and instinctive behavior (as he says in the letter from September 19, 1937).
26
A note on the first page of Lorenz and Tinbergen, 1938 specifies that it is based on Lorenz’s instinct
concept and that the theoretical part of the paper is mainly due to Lorenz. Tinbergen himself suggested to
Lorenz to add this note (Tinbergen, letter to Lorenz, October 12, 1937, Konrad Lorenz Family Papers).
27
See Morgan, 1896; McDougall, 1933; Bierens de Haan, 1933.
28
Bierens de Haan, 1942, p. 100.
29
Kalikow, 1976; Föger and Taschwer, 2001; Taschwer and Föger, 2003.
30
Heinroth and Lorenz, 1988, p. 244.
31
Lorenz, 1939a, 1940, 1943.
32
Lorenz, letter to Stresemann, March 5, 1933; my translation.
33
Heinroth and Heinroth, 1924–1933.
34
“Die Sauropsiden-Reihe hat hier ganz ähnliche Affekte, Gebräuche und Motive entwickelt, wie wir
sie bei uns Menschen gewöhnlich für verdienstvoll, moralisch und dem Verstande entsprungen halten. Das
Studium der Ethologie der höheren Tiere—leider ein noch sehr unbeackertes Feld—wird uns immer mehr
zu der Erkennnis bringen, daß es sich bei unserem Benehmen gegen Familie und Fremde, beim
Liebeswerben und ähnlichem um rein angeborene, viel primitivere Vorgänge handelt, als wir gemeinhin
glauben.” (Heinroth, 1911, p. 702)
35
E. S. Russell stated that it is unlikely that the migration of fish can be explained by a purely
physiological theory (Russell, 1934, p. 42). In his copy of this monograph, Lorenz annotated this passage:
“why in hell??”
36
Tinbergen termed this an ‘objectivistic’ in contrast to Bierens de Haan’s ‘subjectivistic’ approach to
behavior (Tinbergen, 1942).
37
Lorenz, 1937b.
38
Kalikow, 1975.
39
Lorenz, 1970, p. xvii.
40
Lorenz, 1942; quoted from the translation Lorenz, 1970, p. 362.
41
Lorenz 1932, quoted from the translation Lorenz, 1970, p. 57. Compare Ziegler, 1910, p. 46.
42
Kalikow gives a list of five criteria of innateness “which Lorenz first justified, in 1932, by means of
the chain reflex theory” (1975, p. 334). It is true that these criteria are consistent with a chain reflex
approach and that Lorenz might have had the chain reflex theory in mind when formulating these criteria.
But it is not the case that Lorenz justified them by means of the chain reflex theory in his paper—he did
not even explicitly mention this theory.
43
Ziegler, 1910, p. 45.
44
Lorenz [1932] 1970, p. 90. Ziegler’s work was actually influenced by Herbert Spencer and Conwy
Lloyd Morgan. Ziegler stated explicitly that he follows Spencer in taking instincts as complex reflexes
(1910, p. 44) and that the instinct concept he uses agrees with the one employed by Morgan in Habit and
Instinct (p. 47). Spencer and Morgan, however, became one of the main targets of Lorenz’s later critique
(in particular Lorenz, 1937a).
It also is interesting to note that Ziegler mentioned C. O. Whitman and gave a full-paragraph quote
from his seminal 1898 lecture on animal behavior (Ziegler, 1910, p. 49; compare Whitman, 1899, p. 328).
Lorenz seemingly overlooked this passage, because he effectively learned about Whitman’s work only
from his student Wallace Craig and read Whitman’s lecture not before 1934. This is peculiar because the
passage that Ziegler quotes starts with “Instinct and structures are to be studied from the common
standpoint of phyletic descent”, the famous statement that Lorenz will later on quote several times in
support of his approach (Lorenz, 1937b, p. 21; 1939b, p. 71; 1996, p. 238). Even though using instincts as
taxonomic characters was one of the earliest theoretical ideas of Lorenz and a very important one, he
apparently did not pay attention to this quote given by Ziegler
45
Lorenz [1935] 1970, pp. 116, 249.
46
Lorenz [1935] 1970, pp. 121, 250.
47
Lorenz 1937a; quoted from the translation Lorenz, 1970, p. 305.
48
Kalikow, 1975, pp. 331f.
49
In a letter to Heinroth from October 1931, Lorenz complained about a paper that Heinroth sent him.
(It is very likely that Lorenz referred to Loeser, 1930, who had argued against the instinct concept,
claiming that it is non-explanatory. To put Loeser’s account very roughly, apart from real reflexes that
involve only a part of an organism, animal behavior has to be explained by principles of gaining pleasure
and avoiding discomfort.) Lorenz mentioned that he and Heinroth had talked several times about pleasure
accompanying instinctive behavior. Lorenz, it seems, was more skeptical than Heinroth about this issue,
stating that he rather agrees with Ziegler that pleasure and discomfort exist only where corresponding
associations can be formed, i.e., where they have a biological meaning. Lorenz seemed to admit subjective
experience, but not in every case and not as an explanatory principle (Heinroth and Lorenz, 1988, p. 79).
50
In a letter from 1936 Lorenz complained that so many physiological chemists are incapable of
viewing animals as an organic entireties: “Ich kenne aber so viele Physiologische Chemiker, die alle ganz
unfähig sind, im Tier eine organische Ganzheit zu sehen.” (Lorenz, letter to Stresemann, October 4, 1936)
51
One of Lorenz’s marginalia on McDougall shows additionally that Lorenz did not view himself as a
mechanist. In a passge where McDougall points out that animals have the capacity to learn (McDougall,
1933, p. 94), Lorenz annotated on the margin: “Who denies this. McD is always argumenting as if the
assumption of an unintelligent instinct, not to be influenced by experience implies that the man who
makes it is a mechanist.”
52
Lorenz, 1985.
53
Wuketits, 1990.
54
F. Süffert, letter to Lorenz, September 5, 1936, Konrad Lorenz Family Papers.
55
“… so finde ich ihre Schlussdefinition: ‘angestrebter Reflexablauf’ nicht völlig ideal …
‘Angestrebt’ ist natürlich ausgezeichnet, aber gegen ‘Reflex’ (Reflexkette) werden sich alle wenden, die in
den letzten Jahren, besonders an den Insekten, gerade die Undulationstätigkeit hervorgehoben haben. …
Ihre Leerlaufreaktion hat Sie ja selbst dazu geführt zum mindesten Reflexe mit u. U. unendlich niedriger
Reizschwelle anzunehmen und damit das wesentliche (für die Reflexlehre wesentliche) des Reflexes, dass
er Hineinströmendes herausspiegelt, aufzuheben. Wenn Sie annehmen, dass jeder Instinkthandlung ein an
sich schon aktives Ganglienmaterial zugrundeliegt … dann erübrigt sich natürlich auch die Suche nach der
speziellen Ursache der Leerlaufreaktion vollkommen.” (von Holst, letter to Lorenz, April 1, 1937, Konrad
Lorenz Family Papers)
56
Van der Klaauw, letter to Lorenz, Mai 16, 1937, Konrad Lorenz Family Papers.
57
“Nicht gerne hätte ich die Arbeit ohne diese Neuorientierung auf Grund der Ergebnisse von Holst in
Druck gegeben.” (van der Klaauw, letter to Lorenz, Mai 25, 1937, Konrad Lorenz Family Papers)
58
“Sachlich ist zunächst festzustellen, dass die Dinge von Holst eine ganz unglaubliche Förderung
meines eigenen Verständnisses der von Lorenz beschriebenen Erscheinungen zu Folge haben. Vieles wird
falsch, z. B. die Definition des ‘angestrebten Kettenreflexes’. Es spricht eine überwiegende und
überzeugende Zahl von Tatsachen, Schwellenerniedrigung, Leerlaufreaktion, Intensitätsabnahme nach ein
oder mehrmaligem Ablauf, unabhängig von Zielerreichung usw. usw. dafür, dass wirklich alle
eigentlichen Instinkthandlungen Holstsche automatisch-rhythmische Prozesse sind. Wirklich unbedingte
Reflexe sind die angeborenen Auslösemechanismen. Damit kommt die Notwendigkeit, analytisch viel
schärfer zwischen dem Ablauf der Instinkthandlung und ihrer Auslösung zu unterscheiden usw. usw. usw.
Dass die Reflexlehre einfach falsch ist, ist nicht meine Schuld, sondern die der Physiologen und eine
unglaubliche Zahl von aus der Reflextheorie heraus unverständlicher Dinge werden auf einnmal einer
überzeugend einfachen Erklärung zugänglich.” (Lorenz, letter to Stresemann, April 16, 1937)
59
In January 1937, long after Lorenz met von Holst, Otto Koehler commented on Lorenz’s first
instinct paper (that was to appear in a few months) and recommended him to read some articles of von
Holst on the neural coordination of rhythmic movements, because these experimentally founded ideas
fitted with Lorenz’s gas-pressure model of the causation of instinctive behavior: “Lesen Sie ferner
v.Holsts 4 Arbeiten im Pflüger 1935/6 über die Koordination der Rhytmen der Flossenschläge, so finden
Sie dort Vorstellungen über deren Zustandekommen, die Ihrem Gasdruckkessel weitgehend parallellaufen
und zudem physiologisch experimentell gut begründet sind.” (Otto Koehler, letter to Lorenz, January 9,
1937, Konrad Lorenz Family Papers)
60
Taschwer and Föger have already pointed to the fact that von Holst did not convince Lorenz before
April 1937, but they did not lay out the chronology and evidence in detail (Taschwer and Föger, 2003).
61
Lorenz says in 1931 about the draft of 1932 paper that it is the attempt to deal in a physiological
manner with features that are usually approached from a psychological point of view: “Es ist nur ein
Versuch, Erscheinungen, die meist nur von der psychologischen Seite betrachtet wurden, einmal
möglichst physiologisch zu fassen.” (Lorenz, letter to Stresemann, July 30, 1931)
62
Lorenz, 1950, pp. 240, 241.
63
Lorenz, 1937b.
64
Lorenz, [1931] 1970, p. 21.
65
Lorenz [1932] 1970, p. 96; [1935] 1970, p. 249; [1937a] 1970, p. 282.
66
Röell, 2000.
67
Burkhardt 1999, 2003, 2004.
68
Whitman, 1919.
69
Heinroth, 1911; Heinroth and Heinroth, 1924–1933.
70
Lorenz, 1950, p. 222, my emphasis. From the perspective of Burkhardt it becomes clear that the
frequent citation of Heinroth and Whitman had the function of justifying the existence of an emerging
discipline (Burkhardt, 1981).
71
The German biologists Friedrich Alverdes and Friedrich Hempelmann used an instinct-organ
analogy before Lorenz, saying that instincts are given to the organism the same way organs are, or
inherited like morphological structures (Alverdes, 1925; Hempelmann, 1926).
72
Lorenz [1935] 1970, pp. 117, 250; [1937a] 1970, pp. 262, 314; 1937b, p. 23.
73
Lorenz, 1935.
74
My account can only show why Lorenz was more likely to have the unusual phylogenetic position
than another one. In contrast, Kalikow is too bold when she claims that the concept of innate releasing
mechanism “could not have been invented without an over-all optic such as that provided by the chain-
reflex theory” (Kalikow, 1975, p. 334). McDougall, for instance, talked about organisms being “endowed
by nature with a lock”, to be unlocked by “a key”, i.e., “a specific object that evokes an instinctive
response” (McDougall, 1933, pp. 98f). So McDougall’s work contained an idea that has some of the main
features of the innate releasing mechanism concept. The difference is not that Lorenz had some ideas with
which other authors could not have come up. Other authors might have had similar ideas, albeit embedded
in a completely different theory. The difference is that Lorenz was more likely to take certain steps given
his observations and theoretical assumptions. (Lorenz actually read this passage of McDougall’s, probably
when working on the ‘Companion’. His annotations show that he was very sympathetic to McDougall’s
key-lock metaphor.)
75
In his discussion of the Dutch tradition in animal psychology and ethology, Röell addresses the
interesting question as to why the Lorenzian ethologists and the representatives of the traditional animal
psychological approach did not want to collaborate or at least to coexist (Röell, 2000). Among other
things, he points out that National Socialism and anti-German sentiments played a role for the attitude of
several Dutch animal towards ethologists such as Konrad Lorenz and Otto Koehler.
76
His research practice was also similar to that of American animal psychologists.
77
Bierens de Haan 1935, 1937. He actually took a vitalist stance by assuming that instincts are brute
facts that we cannot explain. Instincts and the goal-directedness of behavior are fundamental explanatory
principles that cannot even be explained by the fact of evolution (1935, 1940).
78
Bierens de Haan, 1940.
79
80
Bierens de Haan, 1940, p. X.
81
Bierens de Haan 1933, 1935.
82
83
Bierens de Haan, 1942.
84
85
86
87
Bierens de Haan, 1942; Lorenz, 1942.
88
In fact, Lorenz did not talk about instincts because this term is too vague. Instead, he preferred the
term ‘instinctive behavior pattern’.
89
Heinroth 1918, 1930.
90
Lorenz, 1935; quoted from the translation Lorenz, 1970, p. 249
91
Lorenz [1937a] 1970, p. 314; Lorenz, 1937b, p. 23.
92
Lorenz, 1996, p. 239.
93
On Whitman see Burkhardt 1988, 2004.
94
Whitman, 1899.
95
Whitman, 1899, pp. 331ff, 338.
96
Whitman, 1899, p. 285
97
Lorenz [1935] 1970, p. 120; [1937a] 1970, p. 291; 1996, pp. 245f.
98
In 1931, Lorenz described the absence or loss of a specific innate behavioral pattern and its sudden
reappearance, depending on the physical condition or the social developmental environment of an
organism. He remarked that intelligence cannot compensate for the loss of these patterns (Lorenz [1931]
1970, pp. 55f). One year later, he observed that environmental and physiological disturbances can result in
the loss of instinctive behavior patterns, claiming that “these disturbances always result in the absence of
components of such patterns and never in the performance of novel ones; there is always breakdown
rather than an alteration.” (Lorenz [1932] 1970, p. 85)

99
When Whitman states that “intelligence would have a tendency to break up and render plastic a
previously stereotyped instinct” (1899, p. 337), Lorenz annotates on the margin: “break up and render
plastic are two very different things!!!” Whitman emphasizes the continuity of instinct and intelligence:
“Plasticity of instinct is not intelligence, but it is the open door through which the great educator,
experience, comes in and works every wonder of intelligence” (p. 338), while Lorenz disagrees by
annotating: “Error! unles Plasticity=Breaking up”. Lorenz finds Whitman’s observation very useful, but
thinks that they are erroneously interpreted. At the end of his copy of Whitman 1899, Lorenz writes: “If
Prof. Whitman wasnt influenced by Spencer and Lloyd Morgan he’d be much nearer the truth.”
100
Whitman, 1899, p. 329.
101
Lorenz [1935] 1970, p. 120.
102
Lorenz [1937a] 1970, p. 290.
103
Lorenz [1937a] 1970, p. 290.
104
Whitman, 1899, p. 337.
105
It is for instance not clear whether he ever read Darwin’s book on emotions. Burkhardt points out
that the preface that Lorenz wrote for the Chicago edition (Darwin, 1965) does not demonstrate any
knowledge of Darwin’s account (Burkhardt, 2004).
106
See Burkhardt 1983, 2004
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The Instinct Concept of the Early Konrad Lorenz

July 30, 2004 (penultimate draft)

Keywords: Konrad Lorenz, ethology, instinct, innateness

never abandoned his strong innate-learned dichotomy.2

Lorenz’s intellectual background and influences

University of Vienna was dominated by comparative morphologists and embryologists. While

reveal why this is so crucial for Lorenz’s theoretical development.4

theoretical contribution to Lorenz’s conceptual innovations, he nonetheless influenced and

behavior, as I will discuss in detail below.

largely created their own research tradition and theoretical approach.

instincts as complex sequences of reflexes and was inspired by physiological studies of

directedness of behavior. An influential case is the psychology of William McDougall. On his

irreducible psychological factors need to be taken into account.13

particular, the goal cannot explain instinctive behavior.

modified by experience—a conviction he clearly expressed in his publications from 1932

adult performance is reached. A maturation process is not necessarily a learning process. On

Herbert Spencer. As this view assumes use-inheritance or other Lamarckist mechanisms of

(non-homologous) insight-based behavior components. I shall explain Lorenz’s evolutionary

extensive ‘theory-free’ observation, so that theoretical prejudices do not affect later

interpretation.22 Thus, Lorenz tried to represent his personal theoretical development in

theory, including a discussion of purposivist, vitalist, and Lamarckist approaches. So when

disagreed with other traditional accounts once he became aware of them.

The four commitments guiding Lorenz’s approach

methodological commitments as to how to study and explain behavior that Lorenz—unlike

interesting ones and will be highlighted in the following discussion.

1. The primacy of the innate

orienting movements in amoebae to insightful behavior in primates.28

points to Oskar Heinroth as another person who views morality as an instinct:

behavior. He was convinced that the innate is the core of behavior.

conspecifics, so that Heinroth systematically conducted deprivation experiments that help to

2. The need for physiological explanations

psychological explanation of instinctive behavior was needed. According to Lorenz, the

subjective motivations of behavior. Animal psychologists with a purposivist approach such as

needs to be explained by causal factors.35 It is the focus on causal-physiological explanation that

on a causal-physiological study and explanation of behavior proved to be important for the

success of ethology as a biological approach to behavior.

Lorenz’s post-war account according to which he “was a firm adherent of classical

cannot account for all behavioral phenomena.

individually-acquired pathways. The psychological definition is thus replaced by a histological

definition.”41 Whereas some animal psychologists characterized instincts by the absence of

complete manner on its first performance.44

despite many unsolved issues.

mechanistic, anti-vitalist view of nature.48 ‘Mechanism’, however, is a problematic label for

commitment, i.e., the idea that we need a neurobiological or physiological explanation of

psychological drives but neurobiological stimuli—von Holst’s work was purely

commitment to the second factor—the physiological approach—was unaffected by the shift

towards von Holst’s ideas.

potentially fruitful research approach to instinctive behavior that is based solely on

approach to the causation of behavior.

3. The taxonomic and phylogenetic approach to behavior

to Lorenz’s approach. Traditional comparative psychology was comparative in that animal

behavioral homology is a type of homology stemming from classical ethology. Instincts as

than just comparative) approach to behavior from Lorenz.66

environment rather than investigating their neurophysiology. The taxonomic-phylogenetic

and phylogenetic point of view.

University of Vienna was dominated by comparative morphologists and embryologists. Lorenz

in pursuing the taxonomic perspective on behavior. Lorenz acknowledged this support by

Heinroth, in particular his comparative-taxonomic approach, on many occasions. In a paper after

approaches and developed some early insights of ethology. He continues: