History of developmental neuroethology: Early contributions from ethology

Ethology has its roots in the natural history of animal behavior. Questions of causation and function have historically been complementary, and each has rested upon a prior appreciation of the behavior of animals in nature. It is thus difficult to place a single time or place where ethology was born. Early evolutionary interests hinted at developmental constraints that continue to guide much research. It has only been relatively recently, however, that the explicit analysis of neural mechanisms in behavior has received the attention it deserves in developmental analyses. A mature developmental neuroethology will require a synthesis of the broad insights of ethology with refined neurobiological technique. Fundamental, however, is the primary focus upon behavior as it normally occurs. © 1992 John Wiley & Sons, Inc.     

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.     

Mental content and evolutionary explanation

Cognitive ethology is the comparative study of animal cognition from an evolutionary perspective. As a sub-discipline of biology it shares interest in questions concerning the immediate causes and development of behavior. As a part of ethology it is also concerned with questions about the function and evolution of behavior. I examine some recent work in cognitive ethology, and I argue that the notions of mental content and representation are important to enable researchers to answer questions and state generalizations about the function and volution of behavior.I would like to thank Dorothy Cheney, Keith Donnellan, Alan Nelson, and Robert Seyfarth for their help.     

A history of Animal Behaviour by a partial, ignorant and prejudiced ethologist

This essay looks back on the history of Animal Behaviour through a compilation of all the papers published in the journal, since it got its present name, that used sticklebacks, the white rat of ethology, as experimental subjects. This stickleback-eye view confirms the role that Animal Behaviour has played during its first 50 years in fostering and recording the important developments that have taken place in the discipline. It also speaks to its current flourishing state as a key journal for the dissemination of results in both ethology (in the sense of studies looking at causation, development and evolution as well as function) and behavioural ecology.     

Tribute to Tinbergen: Putting Niko Tinbergen's ‘Four Questions’ in Historical Context

Niko Tinbergen's (Zeit. Tier. 20, 1963, 410) paper ‘On aims and methods of ethology’ is appropriately remembered as the paper in which Tinbergen characterized ethology as ‘the biological study of behavior’ and went on to explain that to study behavior biologically is to ask four distinct questions about it: (1) How is it caused physiologically? (2) What is its survival value? (3) How has it evolved? and (4) How does it develop in the individual? Here, we consider Tinbergen's paper in its historical context by looking at it from three different perspectives: (1) a comparison of Tinbergen's formulation of ‘ethology's four questions’ with similar, but different formulations of biology's basic problems offered by Julian Huxley, Konrad Lorenz, and Ernst Mayr; (2) a survey of the roles that the four questions played in Tinbergen's own work over the course of his career; and (3) a consideration of the two explicit goals of Tinbergen's (Zeit. Tier., 20, 1963, 410) paper, namely (a.) to honor Tinbergen's friend and colleague Konrad Lorenz (as part of a Festschrift for Lorenz on the occasion of his sixtieth birthday) and (b.) to provide a sketch of ethology's scope and an evaluation of the ways the field needed to develop in the future. We suggest that just as the work of Tinbergen's Oxford research team revealed how the behavior of gulls reflected compromises worked out in the face of the diverse selective pressures of particular environments, we can identify certain conflicts that arose for Tinbergen in trying to write something that his friend Lorenz would like while also assessing ethology's current state and future prospects. That said, however, Tinbergen's enduring concern was to do all he could to ensure that ethology thrive as a field and develop a scientific understanding of animal (and human) behavior. For this to happen, he insisted, the four questions of ethology needed to be pursued in a balanced, comprehensive, and integrated fashion.     

Child social ethology and peer relations: a developmental review of methodology and findings

During the last three decades, developmental research has increasingly emphasized the relevance of peer relations in children’s socialization. However, most studies of child development still focus upon individual differences in social status, tacitly neglecting relational constraints inherent in the ecology of the peer group. In contrast, socioethological approaches have stressed that natural groups provide a variety of distinct social roles that may have a differential impact upon individual growth and development. However, ethological analyses have often been limited to aggressive relations and group dominance structures. Comparable studies of affiliative organization have been hampered by the paucity of models for the study of cohesive social structure, only recently having begun to overcome this obstacle. A potential third dimension of preschool peer group ethology, object use, has been relatively neglected in studies of peer relations. However, recent research suggests that object use is a salient and important component of the social world of the young child. This article reviews methodology and recent findings in the area of peer relations and discusses the developmental implications of this work. Received: 8 December 1998 / Received in revised form: 26 March 1999 / Accepted: 31 March 1999     

Fishing for genes influencing vertebrate behavior: zebrafish making headway

The zebrafish (Danio rerio) has been a favorite model of developmental biologists and geneticists, but only recently have investigators begun to appreciate its usefulness in behavior genetics. Papers focusing on the behavior or brain function of this species were once extremely rare, but during the past decade rapid growth has taken place. Despite the increased interest, however, the number of studies devoted to the analysis of the behavior of this species is still orders of magnitude less than those conducted on more traditional laboratory subjects including the rat and the mouse. The authors review selected literature and demonstrate that zebrafish is an excellent subject for behavior genetics research, especially in the area of forward genetics (mutagenesis).     

斑马鱼行为学实验在神经科学中的应用

斑马鱼作为新型模式动物的优势正在逐渐被人们所认识,其应用的领域也越来越宽广．斑马鱼在神经生物学中的应用,除了在发育方面比其他模式动物更具优势外,在行为学方面的应用也更加丰富．由于斑马鱼幼体在受精后前两天通体透明,眼睛大小占到大脑体积的二分之一以上,成鱼昼夜节律明显,对光反应强烈,因此斑马鱼在视觉领域应用的优势十分明显．斑马鱼的嗅觉、听觉器官都在体表可见,可以很容易地用行为学实验手段对嗅觉和听觉功能进行检测．斑马鱼习性好动,利用斑马鱼进行运动方面的行为学观察也非常便利．斑马鱼具有群聚习性,在社会生物学研究方面正得到越来越多的关注．斑马鱼行为学是一种比较简单而又有效地分析神经整合功能的方法,并形成了许多相关的实验模型．     

Nature-Nurture and the Debates Surrounding Ethology and Sociobiology

The central problem in the history of animal behavior has beenthe inability to perceive the phenotype as the result of aninteraction between genome and environment, despite the considerablelip service paid to the interaction. In North America the comparativestudy of animal behavior was overshadowed by the growth of anexperimental psychology that produced the general-process viewof learning, holding that the mechanisms underlying learningare much the same in all species. That made evolution irrelevant.During the same period ethology emerged in Europe as the studyof naturally occurring behavior in an evolutionary context.Because evolution is fundamental to ethology, the genetic basisof behavior was a central precept. Ethology and psychology collidedafter World War II. After a vigorous exchange on the issues,a synthesis by Robert Hinde materialized, one that advancedthe study of behavior and produced a sophisticated understandingof nature and nurture. A few decades later sociobiology appearedand was immediately assailed for making what were seen as unwarrantedextensions from animal to human behavior, and for emphasizinggenetic control of behavior. Much of the debate that ensuedwas distractingly political and threw little light on the scientificmerits of the issues although it moderated the stance of sociobiologists;on the other hand, the politically inspired debates may haveharmed the field of animal behavior.     

On the Problem of Man in Psychology

Very few works in our literature have been devoted to human ethology and social ethology, particularly of children. Psychological-pedagogical approaches predominate; these are based on an assessment of verbal information and on the results of experimental or educational inputs. But such studies should rest on knowledge of the distinctive features of a person's immediate behavior, and these can be ascertained solely through objective study, in which every directed input is precluded.     

Schedule-controlled behaviors as determinants of drug response.

Schedule-controlled performances began to be used in assessing the behavioral effects of drugs because of practical advantages over other techniques for studying behavior. Schedule-controlled behavior is, however, of fundamental importance in behavioral pharmacology. It has been found repeatedly that the effects of many drugs depend critically upon the patterns of responding engendered by different schedule contingencies. These dependencies of the effects of drugs on schedule-controlled behavior occur because ongoing behavior is itself an important determinant of drug action.     

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.     

Developmental programming of reproduction and fertility: what is the evidence?

The concept of the foetal/developmental origins of adult disease has been around for ~20 years and from the original epidemiological studies in human populations much more evidence has accumulated from the many studies in animal models. The majority of these have focused upon the role of early dietary intake before conception, through gestation and/or lactation and subsequent interactions with the postnatal environment, e.g. dietary and physical activity exposures. Whilst a number of theoretical models have been proposed to place the experimental data into a biological context, the underlying phenomena remain the same; developmental deficits (of single (micro) nutrients) during critical or sensitive periods of tissue growth alter the developmental pathway to ultimately constrain later functional capacity when the individual is adult. Ageing, without exception, exacerbates any programmed sequelae. Thus, adult phenotypes that have been relatively easy to characterise (e.g. blood pressure, insulin sensitivity, body fat mass) have received most attention in the literature. To date, relatively few studies have considered the effect of differential early environmental exposures on reproductive function and fecundity in predominantly mono-ovular species such as the sheep, cow and human. The available evidence suggests that prenatal insults, undernutrition for example, have little effect on lifetime reproductive capacity despite subtle effects on the hypothalamic-pituitary-gonadal axis and gonadal progenitor cell complement. The postnatal environment is clearly important, however, since neonatal/adolescent growth acceleration (itself not independent from prenatal experience) has been shown to significantly influence fecundity in farm animals. The present paper will expand these interesting areas of investigation and review the available evidence regarding developmental programming of reproduction and fertility. However, it appears there is little strong evidence to indicate that offspring fertility and reproductive senescence in the human and in farm animal species are overtly affected by prenatal nutrient exposure. Nevertheless, it is clear that the developing gonad is sensitive to its immediate environment but more detailed investigation is required to specifically test the long-term consequences of nutritional perturbations during pregnancy on adult reproductive well-being.     

The Ethological Approach to the Study of Human Behavior

Human ethology, which was established on the basis of classical zooethology, can be an inspirational contribution to the study of human behavior. The study of behavior in natural conditions is stimulating as well as the primary interest of ethologists in such behavioral patterns showing evolutionary success and benefits and which are called inborn or innate. The extensive area of human behavior, nonverbal communication, can be investigated also with the application of some ethological knowledge. Human ethology can bring significant insight to the evaluation of the pathology of human behavior in various medical disciplines. An important task of medical prediction (prognosis) can be made more reliable by considering the ethology. A specific attribute of the species Homo sapiens, his culture, is acknowledged and discussed through human ethology.     

On the optimality of coarse behavior rules

Animal behavior can be characterized by the degree of responsiveness it has to variations in the environment. Some behavior rules lead to fine-tuned responses that carefully adjust to environmental cues, while other rules fail to discriminate as carefully, and lead to more inflexible responses. In this paper we seek to explain such inflexible behavior. We show that coarse behavior, behavior which appears to be rule-bound and inflexible, and which fails to adapt to predictable changes in the environment, is an optimal response to a particular type of uncertainty we call extended uncertainty. We show that the very variability and unpredictability that arises from extended uncertainty will lead to more rigid and possibly more predictable behavior. We relate coarse behavior to the failures to meet optimality conditions in animal behavior, most notably in foraging behavior, and also address the implications of extended uncertainty and coarse behavior rules for some results in experimental versus naturalistic approaches to ethology.     

The Myth of Male Superiority: Rise and Demise

A prolific literature exists concerning the origins and functions of the institutionalization of sex-role differences. However, persistent problems and sharply divergent views remain. This study attempts to circumvent the nature/nurture controversy by utilizing a holistic-evolutionary approach to the problem. The reviews of evidence from ethology, developmental psychology, and cross-cultural investigations lead to a social-exchange model involving differences in the elasticities of male and female labor contributions in social evolution. Assumptions about future technological progress lead to prediction of the demise of the myth of male supremacy. [sex differences in social evolution, ontogeny of human behavior]     

How Elephants are Opening Doors: Developmental Neuroethology, Attachment and Social Context

Ethology's renewed interest in developmental context coincides with recent insights from neurobiology and psychology on early attachment. Attachment and social learning are understood as fundamental mechanisms in development that shape core processes responsible for informing behaviour throughout a lifetime. Each field uniquely contributes to the creation of an integrated model and encourages dialogue between Tinbergen's four analytical levels: ethology in its underscoring of social systems of behaviour and context, psychology in its emphasis on socio‐affective attachment transactions, and neuroscience in its explication of the coupled development of brain and behaviour. We review the relationship between developmental context and behaviour outcome as a topic shared by the three disciplines, with a specific focus on underlying neuroethological mechanisms. This interdisciplinary convergence is illustrated through the example of abnormal behaviour in wild African elephants (Loxodonta africana) that has been systematically observed in human‐caused altered social contexts. Such disruptions impair normative socially mediated neuroendocrinological development leading to psychobiological dysregulation that expresses as non‐normative behaviour. Aberrant behaviour in wild elephants provides a critical field example of what has been established in ex situ and clinical studies but has been largely absent in wild populations: a concrete link between effects of human disturbance on social context, and short‐ and long‐term neuroethology. By so doing, it brings attention to the significant change in theories of behaviour that has been occurring across disciplines – namely, the merging of psychobiological and ethological perspectives into common, cross‐species, human inclusive models.     

The Mechanical Properties of Rat Tail Tendon

The load-strain and stress-relaxation behavior of wet rat tail tendon has been examined with respect to the parameters strain, rate of straining, and temperature. It is found that this mechanical behavior is reproducible after resting the tendon for a few minutes after each extension so long as the strain does not exceed about 4 per cent. If this strain is exceeded, the tendon becomes progressively easier to extend but its length still returns to the original value after each extension. Extensions of over 35 per cent can be reached in this way. Temperature has no effect upon the mechanical behavior over the range 0–37°C. Just above this temperature, important changes take place in the mechanical properties of the tendon which may have biological significance. The application of the techniques used here to studies of connective tissue disorders is suggested. Some of the mechanical properties of tendon have been interpreted with a simple model.