A possible contribution of phenomenology to ethology: Application to a behaviour pattern in the mouse

Classical ethology encourages a causal approach to animal behaviour, using Tinbergen's four questions concerning evolution, function, mechanism and development of behaviour. It sets aside the study of mental processes, which could otherwise help to unify our picture of the relationships between animal and environment. Here the steps in research focused on the psychological meaning of a peculiar behaviour in the mouse — carrying its tail — and what this implies regarding the mouse's cognitive world are given. Initial empirical observations suggested epistemic choices concerning space and object notions in the mouse; this led us to go beyond the first stage in exploring the significance of this behaviour. Later experiments showed the limitations of an explanation based on a cause-effect relationship. An interpretative model integrating a phenomenological conceptual framework is proposed.     

A textbook history of animal behaviour

The history of animal behaviour research over the last 50 years has been marked by a decline in traditional ethology and an increase in the prominence of behavioural ecology. Here I examine these changes in the discipline of animal behaviour during the past 50 years as reflected in specialized behaviour textbooks and introductory textbooks for general biology courses. Prior to 1975, all the behaviour texts devoted far more coverage to ethological research on the proximate causes of behaviour than to research on the adaptive value of behavioural traits. After 1975, ultimate questions about adaptation received much more attention than they had previously. The change took place because of events in the mid-1960s, including W. D. Hamilton's solution to the evolutionary puzzle of altruism and the group selection debate involving V. C. Wynne-Edwards, G. C. Williams and D. Lack. The effect of these events was to encourage researchers to use sound adaptationist theory untainted by species-benefit thinking. The theory helped identify many new questions about the adaptive value of behavioural traits, stimulating workers to give as much weight to these issues as had previously been devoted to research into the physiology and development of behaviour. Although pure ethological research appeared to decline after 1975, in fact the major proximate concerns of ethology were simply taken on by cell biologists and neurophysiologists who used tools not available to the classic ethologist. The result was the development of neuroethology, which has flourished over the years. However, classic (pre-1975) ethological research still receives considerable coverage in many introductory biology textbooks, possibly because of the weight given these studies by W. T. Keeton in his influential textbook. None the less, ever since the 1980s, most biology textbooks have also discussed some of the fruits of adaptationist theory, especially Hamilton's explanation for altruism, thereby providing a more balanced treatment of proximate and ultimate issues than was once the case. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Behavioural ecology's ethological roots

Since Krebs and Davies's (1978) landmark publication, it is acknowledged that behavioural ecology owes much to the ethological tradition in the study of animal behaviour. Although this assumption seems to be right-many of the first behavioural ecologists were trained in departments where ethology developed and matured-it still to be properly assessed. In this paper, I undertake to identify the approaches used by ethologists that contributed to behavioural ecology's constitution as a field of inquiry. It is my contention that the current practices in behavioural biology owe ethology something much subtler than the simple transposition of Tinbergen's Four Problems for heuristic purposes. Demonstrating what ethology inherited from the long naturalist tradition shows the tensions that strained the field and that later led to the loss of both its unity and its specificity. It also allows for a precise delineating of what behavioural ecology picked up from the ethological practice, and it helps to cast some light on the introduction of economical thinking in behavioural sciences.     

A microcomputer method for recording and analysing behavioural elements

The trend in ethology, psychopharmacology and behaviour genetics is to get increasingly higher resolution of the behaviour of animals, since it increases the sensitivity of the tests used. Consequently the higher resolution requires an increased data-logging efficiency, and a decreased time-investment for data handling. In this paper a method for inexpensive microcomputers is presented, which enables an individual experimenter to obtain frequency, relative duration and its variance from as many behavioural elements as desired.     

Sex and boldness explain individual differences in spatial learning in a lizard

Understanding individual differences in cognitive performance is a major challenge to animal behaviour and cognition studies. We used the Eastern water skink (Eulamprus quoyii) to examine associations between exploration, boldness and individual variability in spatial learning, a dimension of lizard cognition with important bearing on fitness. We show that males perform better than females in a biologically relevant spatial learning task. This is the first evidence for sex differences in learning in a reptile, and we argue that it is probably owing to sex-specific selective pressures that may be widespread in lizards. Across the sexes, we found a clear association between boldness after a simulated predatory attack and the probability of learning the spatial task. In contrast to previous studies, we found a nonlinear association between boldness and learning: both ‘bold’ and ‘shy’ behavioural types were more successful learners than intermediate males. Our results do not fit with recent predictions suggesting that individual differences in learning may be linked with behavioural types via high–low-risk/reward trade-offs. We suggest the possibility that differences in spatial cognitive performance may arise in lizards as a consequence of the distinct environmental variability and complexity experienced by individuals as a result of their sex and social tactics.     

What Capabilities for the Animal?

In this essay, I defend a bi-constructivist approach to ethology—a constructivist ethology assuming that each animal adopts constructivist strategies. I put it in opposition to what I call a realist-Cartesian approach, which is currently the dominant approach to ethology and comparative psychology. The starting point of the bi-constructivist approach can be formulated as a shift from the classical Aristotelian question “What is an animal?” to the Spinozean question, which is much less classical but which seems to me to be much stronger: “What are the capacities of the animal?”. Is it possible to conceptualize an ethology which insists on interpretation and therefore on invention, innovation and creativity, rather than on causality, the monotony of behavioural routines, and/or genetic or environmental determination? Such an ethology would be based not on the fiction of an absent observer but on fully recognizing the necessity of an observer, who is effectively present in order to get an observation. A pluralistic ethology does not dissociate itself from the marginal epistemologies of practitioners like animal trainers, hunters, stockbreeders etc., or, moreover, non-western experts. An ethology of this kind is not clamped within the boundaries of purely academic epistemology, obsessed by demarcation lines between the human and the animal. My work on the bi-constructivist approach represents a contribution towards the elaboration of an authentically biosemiotic ethology, one which is significantly different from the mechanical ethology of today.     

Optimal behaviour can violate the principle of regularity

Understanding decisions is a fundamental aim of behavioural ecology, psychology and economics. The regularity axiom of utility theory holds that a preference between options should be maintained when other options are made available. Empirical studies have shown that animals violate regularity but this has not been understood from a theoretical perspective, such decisions have therefore been labelled as irrational. Here, I use models of state-dependent behaviour to demonstrate that choices can violate regularity even when behavioural strategies are optimal. I also show that the range of conditions over which regularity should be violated can be larger when options do not always persist into the future. Consequently, utility theory—based on axioms, including transitivity, regularity and the independence of irrelevant alternatives—is undermined, because even alternatives that are never chosen by an animal (in its current state) can be relevant to a decision.     

Behavioural and physiological stress responses to handling in wild guanacos

Both ethology and physiology can provide insights about stress conditions in wild animals during handling. Lama guanicoe (guanaco) is the major large herbivore inhabiting arid zones in Argentina. Since 2005, the Payún Matrú Cooperative, a business established and run by native people, has captured and handled guanacos for shearing at Payunia Reserve, Mendoza Province, Argentina, to market the species’ fibre for clothing and other products. The aim of our study was to evaluate the behavioural and physiological stress responses to handling of wild guanacos during these shearing activities. We evaluated stress conditions in guanacos by monitoring the frequency of agonistic behaviour in the holding corrals, following capture, the rate of vocalisation and agonistic behaviour during handling, and by estimating cortisol levels in handled animals. Crowding increased agonistic behaviour; this appeared to be related to dominance conflicts between males in the corrals. Time of corralling increased rate of agonistic behaviour and vocalisations during handling. The rate of behavioural stress responses was negatively related to cortisol levels in males; which may be due to animal fatigue or because behavioural stress responses may help mitigate physiological stress responses. Incorporating our results into animal welfare practices could help reduce stress conditions in wild guanacos. Our results suggest that both behavioural and physiological responses are important measures of stress in handled animals, and that behavioural stress responses cannot be used as a simple surrogate for glucocorticoid levels.     

Behaviour in captivity predicts some aspects of natural behaviour,but not others,in a wild cricket population

Examining the relevance of ‘animal personality’ involves linking consistent among- and within-individual behavioural variation to fitness in the wild. Studies aiming to do this typically assay personality in captivity and rely on the assumption that measures of traits in the laboratory reflect their expression in nature. We examined this rarely tested assumption by comparing laboratory and field measurements of the behaviour of wild field crickets (Gryllus campestris) by continuously monitoring individual behaviour in nature, and repeatedly capturing the same individuals and measuring their behaviour in captivity. We focused on three traits that are frequently examined in personality studies: shyness, activity and exploration. All of them showed repeatability in the laboratory. Laboratory activity and exploration predicted the expression of their equivalent behaviours in the wild, but shyness did not. Traits in the wild were predictably influenced by environmental factors such as temperature and sunlight, but only activity showed appreciable within-individual repeatability. This suggests that some behaviours typically studied as personality traits can be accurately assayed in captivity, but the expression of others may be highly context-specific. Our results highlight the importance of validating the relevance of laboratory behavioural assays to analogous traits measured in the wild.     

Swarm Cognition: an interdisciplinary approach to?the?study of self-organising biological collectives

Basic elements of cognition have been identified in the behaviour displayed by animal collectives, ranging from honeybee swarms to human societies. For example, an insect swarm is often considered a ??super-organism?? that appears to exhibit cognitive behaviour as a result of the interactions among the individual insects and between the insects and the environment. Progress in disciplines such as neurosciences, cognitive psychology, social ethology and swarm intelligence has allowed researchers to recognise and model the distributed basis of cognition and to draw parallels between the behaviour of social insects and brain dynamics. In this paper, we discuss the theoretical premises and the biological basis of Swarm Cognition, a novel approach to the study of cognition as a distributed self-organising phenomenon, and we point to novel fascinating directions for future work.     

Three-Dimensional Tracking of Small Aquatic Organisms Using Fluorescent Nanoparticles

Tracking techniques are vital for the understanding of the biology and ecology of organisms. While such techniques have provided important information on the movement and migration of large animals, such as mammals and birds, scientific advances in understanding the individual behaviour and interactions of small (mm-scale) organisms have been hampered by constraints, such as the sizes of existing tracking devices, in existing tracking methods. By combining biology, chemistry and physics we here present a method that allows three-dimensional (3D) tracking of individual mm-sized aquatic organisms. The method is based on in-vivo labelling of the organisms with fluorescent nanoparticles, so-called quantum dots, and tracking of the organisms in 3D via the quantum-dot fluorescence using a synchronized multiple camera system. It allows for the efficient and simultaneous study of the behaviour of one as well as multiple individuals in large volumes of observation, thus enabling the study of behavioural interactions at the community scale. The method is non-perturbing – we demonstrate that the labelling is not affecting the behavioural response of the organisms – and is applicable over a wide range of taxa, including cladocerans as well as insects, suggesting that our methodological concept opens up for new research fields on individual behaviour of small animals. Hence, this offers opportunities to focus on important biological, ecological and behavioural questions never before possible to address.     

Conservation needs ethologv

With the number of species and near-to-natural habitats rapidly dwindling, conservation has become an undebatable necessity. There have been some laudable, successful species conservation projects but there have also been many deplorable failures. The failures are exacerbated by limited funding. Conservationists depend on funding by national government organizations (NG0s) and by private sponsors, more than other practitioners of organismic biology do. To maximize their success, conservationists would be well advised to heed the messages resulting from animal behaviour study (i.e. ethology) and/or to involve ethologists in their projects. Here, I illustrate how ethology can benefit both in situ and ex situ conservation measures; the need for conservation-oriented behaviour research is paramount.     

A cognitive behavioural intervention to reduce sexually transmitted infections among gay men: randomised trial

ObjectiveTo determine the effectiveness of a brief cognitive behavioural intervention in reducing the incidence of sexually transmitted infections among gay men.DesignRandomised controlled trial with 12 months'' follow up.SettingSexual health clinic in London.Participants343 gay men with an acute sexually transmitted infection or who reported having had unprotected anal intercourse in the past year.Results72% (361/499) of men invited to enter the study did so. 90% (308/343) of participants returned at least one follow up questionnaire or re-attended the clinic and requested a check up for sexually transmitted infections during follow up. At baseline, 37% (63/172) of the intervention group and 30% (50/166) of the control group reported having had unprotected anal intercourse in the past month. At 12 months, the proportions were 27% (31/114) and 32% ( 39/124) respectively (P=0.56). However, 31% (38/123) of the intervention group and 21% (35/168) of controls had had at least one new infection diagnosed at the clinic (adjusted odds ratio 1.66, 95% confidence interval 1.00 to 2.74). Considering only men who requested a check up for sexually transmitted infections, the proportion diagnosed with a new infection was 58% (53/91) for men in the intervention group and 43% (35/81) for men in the control group (adjusted odds ratio 1.84, 0.99 to 3.40). Using a regional database that includes information from 23 sexual health clinics in London, we determined that few participants had attended other sexual health clinics.ConclusionsThis behavioural intervention was acceptable and feasible to deliver, but it did not reduce the risk of acquiring a new sexually transmitted infection among these gay men at high risk. Even carefully designed interventions should not be assumed to bring benefit. It is important to evaluate their effects in randomised trials with objective clinical end points.

What is already known on this topic

The need for effective HIV prevention strategies based on reducing sexual risk behaviour remains importantFew interventions to reduce sexual risk behaviour have been rigorously evaluated using randomised controlled trials

What this study adds

This is the first randomised controlled trial of an intervention addressing sexual behaviour in homosexual men that uses sexually transmitted infections and self reported behaviour as end pointsThe intervention was brief and feasible to use in a busy clinic, but it did not reduce the risk of participants acquiring new infectionsThe potential for behavioural interventions to do more harm than good needs to be taken seriously     

Immunogenetic polymorphism of lipoproteins in swine

Five new antigenic markers (allotypes) of swine serum lipoproteins are described. Specific antiallotype reagents were obtained from alloimmune precipitating sera. Identification studies and genetic analysis indicate that the five serum alloantigens—designated Lpp6, Lpp11, Lpp12, Lpp13, and Lpp14—are markers of low-density lipoproteins (LDL),d 1.002–1.075 g/ml, and are members of a previously described Lpp system. The Lpp6 allotype belongs to the group of individual markers and is determined by a new codominant allelic gene,Lpp ⁶, whereas the remaining four antigens—Lpp11, Lpp12, Lpp13, and Lpp 14—named common specificities, behave as alternative variants to Lpp1, Lpp2, Lpp3, and Lpp4, respectively, forming pairs of mutually exclusive alloantigens. Each Lpp gene in a heterozygous animal expresses itself independently on separate molecules and each haplotype carries one individual and at least four common specificities. The relationship between common and individual specificities, together with their number in the complex haplotypes, seems to shed some light on evolution of Lpp genes. It is proposed in this concept that the original gene for low-density lipoproteins in swine, and also in rhesus monkeys and the human, consisted of genetic information for common specificities only, the individual specificities evolving later as a result of point mutations.     

Dimensions of Animal Personalities in Guinea Pigs

Behavioural phenotypes can be studied from a variety of perspectives. Recent developments have focused on the individual, seeking patterns of behaviour that are stable over time and/or across different contexts (animal personalities). This study applied this method of understanding individual behavioural variability to domestic guinea pigs. Two behavioural domains were investigated: emotionality and social behaviour. Additionally, individual cortisol–stress reactivity and dominance status were examined. Adult male domestic guinea pigs living in large mixed‐sex colonies were subjected to a series of behavioural and physiological tests twice with an intertest interval of 8 wk. Individual consistency over time was found regarding social behaviour, cortisol reactivity and dominance status, whereas no stability regarding emotional behaviour was detected. Furthermore, no stability over contexts was found. Our results suggest that the concept of animal personality is applicable to domestic guinea pigs. The ecological relevance of these data is underscored by the fact that they were obtained in animals from a very rich, socially complex scenario. Moreover, our study highlights that behaviour alone is not sufficient to describe individual phenotypic consistency. Physiological parameters such as stress reactivity should be included in animal personality research. Furthermore dominance – a relative measure which is not an absolute attribute of individuals – proved to be stable over time and thus also shed light on individuality.     

Identification of Behaviour in Freely Moving Dogs (Canis familiaris) Using Inertial Sensors

Monitoring and describing the physical movements and body postures of animals is one of the most fundamental tasks of ethology. The more precise the observations are the more sophisticated the interpretations can be about the biology of a certain individual or species. Animal-borne data loggers have recently contributed much to the collection of motion-data from individuals, however, the problem of translating these measurements to distinct behavioural categories to create an ethogram is not overcome yet. The objective of the present study was to develop a “behaviour tracker”: a system composed of a multiple sensor data-logger device (with a tri-axial accelerometer and a tri-axial gyroscope) and a supervised learning algorithm as means of automated identification of the behaviour of freely moving dogs. We collected parallel sensor measurements and video recordings of each of our subjects (Belgian Malinois, N=12; Labrador Retrievers, N=12) that were guided through a predetermined series of standard activities. Seven behavioural categories (lay, sit, stand, walk, trot, gallop, canter) were pre-defined and each video recording was tagged accordingly. Evaluation of the measurements was performed by support vector machine (SVM) classification. During the analysis we used different combinations of independent measurements for training and validation (belonging to the same or different individuals or using different training data size) to determine the robustness of the application. We reached an overall accuracy of above 90% perfect identification of all the defined seven categories of behaviour when both training and validation data belonged to the same individual, and over 80% perfect recognition rate using a generalized training data set of multiple subjects. Our results indicate that the present method provides a good model for an easily applicable, fast, automatic behaviour classification system that can be trained with arbitrary motion patterns and potentially be applied to a wide range of species and situations.     

Why Was Thomas A. Sebeok Not a Cognitive Ethologist? From “Animal Mind” to “Semiotic Self”

In the current debates about zoosemiotics its relations with the neighbouring disciplines are a relevant topic. The present article aims to analyse the complex relations between zoosemiotics and cognitive ethology with special attention to their establishers: Thomas A. Sebeok and Donald R. Griffin. It is argued that zoosemiotics and cognitive ethology have common roots in comparative studies of animal communication in the early 1960s. For supporting this claim Sebeok’s works are analysed, the classical and philosophical periods of his zoosemiotic views are distinguished and the changing relations between zoosemiotics and cognitive ethology are described. The animal language controversy can be interpreted as the explicit point of divergence of the two paradigms, which, however, is a mere symptom of a deeper cleavage. The analysis brings out later critical differences between Sebeok’s and Griffin’s views on animal cognition and language. This disagreement has been the main reason for the critical reception and later neglect of Sebeok’s works in cognitive ethology. Sebeok’s position in this debate remains, however, paradigmatic, i.e. it proceeds from understanding of the contextualisation of semiotic processes that do not allow treating the animal mind as a distinct entity. As a peculiar parallel to Griffin’s metaphor of “animal mind”, Sebeok develops his understanding of “semiotic self” as a layered structure, characterised by an ability to make distinctions, foremost between itself and the surrounding environment. It appears that the history of zoosemiotics has two layers: in addition to the chronological history starting in 1963, when Sebeok proposed a name for the field, zoosemiotics is also philosophically rooted in Peircean semiotics and German biological philosophy. It is argued that the confrontation between zoosemiotics and cognitive ethology is related to different epistemological approaches and at least partly induced by underlying philosophical traditions.     

Impedance traces of copepod appendage movements illustrating sensory feeding behaviour

An experimental system incorporating a computerized micro-impedance unit has been used to make direct measurements of the activity of copepod cephalic appendages. As the appendages are used to both propel the copepod through water and handle particles, it follows that appendage activity reflects feeding behaviour.To investigate the sensory feeding behaviour of copepods, their activity was recorded with food stimuli varying in size and chemical composition. Sample impedance traces are given for the appendage movements of Temora longicornis in the presence of: 1 — filtered seawater; 2 — beads; 3 — phytoplankton cells; 4 — dissolved free amino acids. The normal appendage movements shown in filtered seawater were modified when copepods were offered particles and dissolved chemicals. Results show that chemical and mechanical stimuli are responsible for the recognition and selection of food. Impedance traces distinguish between behavioural responses such as: antennule flicks, leg kicks, combing, handling and rejection of particles. Spectral analyses of traces have demonstrated that differences in beat pattern are significant.Paper presentation, at the Third International Conference on Copepoda. British Museum (Natural History), London, U. K., 10–14 August, 1987.     

An ant’s eye view of culture: propagation of new traditions through triggering dormant behavioural patterns

We consider a previously unknown way of propagation of behavioural traditions in animal communities using hunting in ants as an example. We experimentally revealed that common litter dwelling ants Myrmica rubra effectively hunt jumping prey and the way the hunting behavioural pattern is distributed within ant colonies is rather sophisticated. Comparison of our results with those obtained on vertebrates enables us to suggest that “distributed social learning” plays an important role in spreading new traditions in animal communities: initial performances by a few carriers of an “at once and entirely” available behavioural pattern propagate this pattern among specimens which have only dormant “sketches” of it. Spread of these behaviours in populations is based on relatively simple forms of social learning such as social facilitation which underlies species’ predisposition to learn certain sequences of behavioural acts. To be triggered, carriers of dormant “sketches” of a relevant behavioural pattern should encounter performances of this pattern with sufficient frequency. We call this strategy triggering of dormant behavioural patterns. Integration of behaviour thus takes place not only at the individual level but at the population level as well.     

Reproduction and the central project of evolutionary theory

In much of the discourse of evolutionary theory, reproduction is treated as an autonomous function of the individual organism — even in discussions of sexually reproducing organisms. In this paper, I examine some of the functions and consequences of such manifestly peculiar language. In particular, I suggest that it provides crucial support for the central project of evolutionary theory — namely that of locating causal efficacy in intrinsic properties of the individual organism. Furthermore, I argue that the language of individual reproduction is maintained by certain methodological conventions that both obscure many of the problems it generates and serve to actively impede attempts to redress those difficulties that can be identified. Finally, I suggest that inclusion of the complexities introduced by sexual reproduction — in both language and methodology — may radically undermine the individualist focus of evolutionary theory.I am indepted to the Rockefeller Foundation for a Humanities Fellowship that supported this research during the spring of 1986. I am also grateful to Richard Lewontin, Diane Paul, and Lisa Lloyd for many extremely helpful conversations.