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Developmental plasticity enables organisms to cope with new environmental challenges. If deploying such plasticity is costly in terms of time or energy, the same adaptive behavior could subsequently evolve through piecemeal genomic reorganisation that replaces the requirement to acquire that adaptation by individual plasticity. Here we report a new dimension to the way in which plasticity can drive evolutionary change leading to ever greater complexity in biological organization. Our model deploys the concept of partially overlapping functional systems. We found that plasticity accelerated dramatically the evolutionary accumulation of adaptive systems in model organisms with relatively low rates of mutation. The effect of plasticity on the evolutionary growth of complexity was even greater when the number of elements needed to construct a functional system was increased. These results suggest that as the difficulty of challenges from the environment become greater, so plasticity exerts an ever more powerful role in meeting those challenges and in opening up new avenues for the subsequent evolution of complex adaptations.

Robert Hinde led the study of animal behaviour for much of his long working life, drawing on his expertise in zoology and ethology to build bridges with psychology, psychiatry and the social sciences. He pioneered the application of a quantitative scientific approach to observations of behaviour, making important contributions to understanding underlying mechanisms, behavioural development and the dynamics of relationships between individuals. His influence on the field of animal behaviour was deep and pervasive, he was held in exceptionally high regard by others and his students and their students occupy many key positions in behavioural studies today. He went on to develop a science of personal relationships in humans, ranging from child development to the nature of aggression. Robert's dedication to research was combined with a commitment to promoting peace, including key roles in the Pugwash Conferences in Science and World Affairs. In retirement, he applied his ethological perspective to writing on the origins of religion and morality, the bases of war, and our culture of greed. Throughout his academic life he was a devoted member of St John's College, Cambridge—serving as teaching fellow, tutor and eventually Master.

Robert Hinde led the study of animal behaviour for much of his long working life, drawing on his expertise in zoology and ethology to build bridges with psychology, psychiatry and the social sciences. He pioneered the application of a quantitative scientific approach to observations of behaviour, making important contributions to understanding underlying mechanisms, behavioural development and the dynamics of relationships between individuals. His influence on the field of animal behaviour was deep and pervasive, he was held in exceptionally high regard by others and his students and their students occupy many key positions in behavioural studies today. He went on to develop a science of personal relationships in humans, ranging from child development to the nature of aggression. Robert's dedication to research was combined with a commitment to promoting peace, including key roles in the Pugwash Conferences in Science and World Affairs. In retirement, he applied his ethological perspective to writing on the origins of religion and morality, the bases of war, and our culture of greed. Throughout his academic life he was a devoted member of St John's College, Cambridge—serving as teaching fellow, tutor and eventually master.

The formation of new species was thought to result from a slow process of Darwinian evolution, but evidence indicates it can occur suddenly. The organism was thought to be passive, playing no role in evolution, but it can affect the evolution of its descendants because of its mobility, choices, control of the environment, and adaptability. Developmental processes were thought to be irrelevant to an understanding of evolution, but the enormous growth of epigenetics suggests that these processes can play an important role in evolutionary change. Acquired information can be passed to progeny without changing DNA sequences, and information can be inherited for a period in the absence of the initial environmental trigger. All this evidence suggests that evolutionary theory is evolving.

This issue of Interface Focus publishes articles based on presentations to a joint Discussion Meeting of the British Academy and the Royal Society held at Carlton House Terrace on 7–9 November 2016. Why was it a joint meeting? The reason is that both sides have much to offer and much to learn

Many forms of developmental plasticity have been observed and these are usually beneficial to the organism. The Predictive Adaptive Response (PAR) hypothesis refers to a form of developmental plasticity in which cues received in early life influence the development of a phenotype that is normally adapted to the environmental conditions of later life. When the predicted and actual environments differ, the mismatch between the individual's phenotype and the conditions in which it finds itself can have adverse consequences for Darwinian fitness and, later, for health. Numerous examples exist of the long-term effects of cues indicating a threatening environment affecting the subsequent phenotype of the individual organism. Other examples consist of the long-term effects of variations in environment within a normal range, particularly in the individual's nutritional environment. In mammals the cues to developing offspring are often provided by the mother's plane of nutrition, her body composition or stress levels. This hypothetical effect in humans is thought to be important by some scientists and controversial by others. In resolving the conflict, distinctions should be drawn between PARs induced by normative variations in the developmental environment and the ill effects on development of extremes in environment such as a very poor or very rich nutritional environment. Tests to distinguish between different developmental processes impacting on adult characteristics are proposed. Many of the mechanisms underlying developmental plasticity involve molecular epigenetic processes, and their elucidation in the context of PARs and more widely has implications for the revision of classical evolutionary theory.

Explanations for biological evolution in terms of changes in gene frequencies refer to outcomes rather than process. Integrating epigenetic studies with older evolutionary theories has drawn attention to the ways in which evolution occurs. Adaptation at the level of the gene is giving way to adaptation at the level of the organism and higher-order assemblages of organisms. These ideas impact on the theories of how cooperation might have evolved. Two of the theories, i.e. that cooperating individuals are genetically related or that they cooperate for self-interested reasons, have been accepted for a long time. The idea that adaptation takes place at the level of groups is much more controversial. However, bringing together studies of development with those of evolution is taking away much of the heat in the debate about the evolution of group behaviour.

This article investigates whether self-reports about playfulness are related to self-reports about creativity and the alternate uses of objects. An on-line survey was conducted of how people think about themselves. One thousand, five hundred and thirty-six people completed the survey. They were asked whether a variety of statements were very characteristic of themselves through to whether they were very uncharacteristic. Respondents were then asked to offer alternative uses for 2 different objects. Those people who characterized themselves as being playful clearly thought of themselves as being creative. The self-reports of their playfulness, creativity, and innovation were positively correlated with each other and were validated with their suggested uses for 2 different objects. Personality measures were derived from the respondents' self-assessments. On the openness scale, the measure was positively correlated with the respondents' assessments of their own playfulness and with the number of alternative ...

This year is the 50th anniversary of Tinbergen’s (1963) article ‘On aims and methods of ethology’, where he first outlined the four ‘major problems of biology’. The classification of the four problems, or questions, is one of Tinbergen’s most enduring legacies, and it remains as valuable today as 50 years ago in highlighting the value of a comprehensive, multifaceted understanding of a characteristic, with answers to each question providing complementary insights. Nonetheless, much has changed in the intervening years, and new data call for a more nuanced application of Tinbergen’s framework. The anniversary would seem a suitable opportunity to reflect on the four questions and evaluate the scientific work that they encourage. Origins of Tinbergen’s questions

The article focuses on the active role of the organism in the subsequent evolution of its descendants. Choice, control of the environment, adaptability, and mobility all play their part. This growth area in biology and other active centres of research on epigenetics and different forms of inheritance are re-invigorating evolutionary biology. Many evolutionary biologists have taken the view that an understanding of development is irrelevant to theories of evolution. However, the integration of several disciplines now suggests that the orthodoxy is misplaced. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112, 268–275.

Behavioural biologists have typically combined interests in the control, function, development and evolution of behaviour. They have used observational and experimental methods, and their findings have been both attractive and scientifically invigorating. A future to be hoped for is that they will continue to combine an understanding of behaviour with studies carried out at other levels but that they will not become too locked into a purely analytical framework. Methodologies are needed that enable scientists to deal with all the principal factors that influence behaviour. In so doing, behavioural biologists should be able to retain a grasp of what is to be an intact, freely moving animal. I believe that the late Gunther Tembrock, to whom this paper is dedicated, would have approved of such a systems approach to behaviour.

The concept of innateness is often used in explanations and classifications of biological and cognitive traits. But does this concept have a legitimate role to play in contemporary scientific discourse? Empirical studies and theoretical developments have revealed that simple and intuitively appealing ways of classifying traits (e.g. genetically specified versus owing to the environment) are inadequate. They have also revealed a variety of scientifically interesting ways of classifying traits each of which captures some aspect of the innate/non-innate distinction. These include things such as whether a trait is canalized, whether it has a history of natural selection, whether it developed without learning or without a specific set of environmental triggers, whether it is causally correlated with the action of certain specific genes, etc. We offer an analogy: the term ‘jade’ was once thought to refer to a single natural kind; it was then discovered that it refers to two different chemical compounds, jadeite and nephrite. In the same way, we argue, researchers should recognize that ‘innateness’ refers not to a single natural kind but to a set of (possibly related) natural kinds. When this happens, it will be easier to progress in the field of biological and cognitive sciences.

I greatly admire the breadth of Geoffrey Lloyd‟s knowledge and the critical mind that he brings to subjects not his own. He highlights the problems that arise when those who believe in human universals confront those who focus on human variation. However, for many the issue is not whether all behaviour is of one kind or the other, but whether or not a particular instance of behaviour depends on an individual‟s experience. This is certainly the debate that arises persistently in my area of behavioural biology (see Johnston 1987). Lloyd rightly criticizes the extremes in the debate over the universal character of human cognition. The target of my critique will be the view that two types of behaviour can be found in any one human being. One of these types supposedly is innate and the other is not. The dichotomy is sometimes presented at the underlying structure of behaviour. For example, while humans obviously differ in the surface characteristics of their language, the “deep structure” is universal. I shall argue that even here the suggestion that two kinds of development, innate and not innate, is false.

Charles Darwin has had an extraordinary impact on many aspects of human affairs apart from revolutionizing biology. On the 200th anniversary of his birth, the Cambridge Darwin Festival in July 2009 celebrated these contributions to the humanities, philosophy and religion and the approach to medicine, economics and the social sciences. He is a man to revere. It is no discredit to him that the science of evolutionary biology should continue to evolve. In this article I shall consider some of the ways in which this has happened since his day.

Japanese quail (Coturnix coturnix japonica) have finely-tuned kin-recognition abilities that may serve a role in mate choice. To investigate whether these abilities are expressed in other life stages, we examined patterns of association among quail chicks in the laboratory four days after hatching. Hatchlings were reared either in mixed-family groups, consisting of siblings and nonsiblings, or in pure-family groups, consisting just of siblings. Chicks were highly social in testing sessions, as reflected in their clumped spatial distributions, and interacted more with their siblings than with nonsiblings. Siblings in each of four mixed-family groups preferentially associated with one another over nonsiblings. No discrimination was evident among differently marked siblings in four purefamily groups. Thus, at an early age, quail discriminate between siblings and nonsiblings despite social experience of both. Recognition abilities that develop prior to or soon after hatching may facilitate cooperation among chicks, may enable siblings to track each others' traits as they mature, or both.

Adult male Japanese quail and, to a lesser extent, adult females were likely to approach a member of the opposite sex which they had not seen before in preference to one with which they grew up. However, they only did this if the group in which they were reared when young contained a particular number of the opposite sex: two females in the case of males and three males in the case of females. The result is explained in terms of the likelihood of the novel member of the opposite sex being slightly different from those with which the birds grew up. If the number of individuals of the opposite sex is small, a novel one is likely to be unacceptably strange, and if the number is high any novel bird from a limited laboratory stock is likely to resemble one or other of the familiar birds. A functional explanation for the quail's behaviour in terms of optical outbreeding is discussed.

1focusing mainly on short-term outcomes such as infant survival and stunting. 2 However, the longer term eff ects on adult health 3 of a poor start to life suggest a further perspective. Developmental eff ects have been viewed traditionally in the context of major disruptions such as caused by teratogens, prematurity and growth retardation, but there is increasing appreciation of the role of developmental plasticity, which provides individuals with the fl exibility to adjust their trajectory of development to match their environment. Plasticity operates across the entire range of environment, from undernutrition to excessive nutritional environments associated with gestational diabetes or maternal obesity, 4,5

Cues received from mothers may have important effects on early development in mammals. We examined the behavioural development of genetically wild-type mice, Mus musculus, offspring born to wild-type or mutant Peg3+/− (paternally expressed gene 3) mothers who are impaired in various aspects of maternal care during both the pre- and the postnatal periods. We demonstrate that impaired maternal care leads to offspring exhibiting increased neophobia and decreased exploratory behaviour. However, these effects were limited to female offspring only, with male offspring being indistinguishable from wild types in both the open-field and the novel object tests. Due to the breeding design of this study we were able to show that this was due not to the inheritance of a genetic mutation but to an epigenetic inheritance. Consistent with this, we also observed a nongenomic inheritance of impairments in maternal care. Wild-type daughters born to mutant mothers were impaired in their ability to retrieve pups to a nest in a retrieval test. Furthermore, the reduced exploration and neophobic phenotypes were transmitted to a third generation, with the granddaughters of mutant females exhibiting increased neophobia even though they had genetically wild-type mothers. We therefore demonstrate a nongenomic transmission of behavioural traits across successive generations operating via the matriline.

Gilbert Gottlieb, a pioneer in developmental psychobiology, died on July 13, 2006. We pay tribute to the man, to his career, and to his dedication to developmental science. © 2007 Wiley Periodicals, Inc. Dev Psychobiol 49: 446–449, 2007.

Ethologists have focused on behavior that is characteristic of the species and adapted to its biological requirements. They have attempted to integrate the four separate problems identified by Tinbergen, namely those to do with current processes, current utility, evolution and development. Studies of development have brought ethologists together with those working in many other fields of biology, psychology, psychiatry, and epigenetics. In doing so, issues such as unitary innateness, nature versus nurture, and heritability have proved to be misleading. Attention is now focused on how an individual develops and the interplay between the processes generating the robust features of an individual's behavior and the many processes involved in plasticity. Individuals make choices and control their environment. Their adaptability is crucial. All these activities have an impact on the evolution of their descendants. Keywords: adaptability; current utility; epigenetics; ethology; evolution; innateness; nature/nurture; plasticity; robustness

Summary Playful play is undoubtedly fun. Even so, many people think, incorrectly, that as they get older, they are no longer capable of such frivolous activity. They should heed George Bernard Shaw's advice: "We don't stop playing because we grow old, we grow old because we stop playing." The motivation to be playful comes from within. No external bribes are needed. In fact attempting to encourage such activity with food or money is likely to be counterproductive.

If temperature does not vary from one generation from to the next but its value is crucial for the development of particular phenotypic characteristics, a long-term change in its value may trigger major evolutionary changes of the organism. If a bird's nest maintains the critical temperature, then a statement that the bird is the nest's way of making another nest is as helpful as accounts couched in terms of genes' intentions. However, the language of intentions rests on different evidence and assumptions from causal language and the languages are not interchangeable. Understanding ontogeny in causal terms requires explanations that are made as simple as possible but not so simple that they become completely unrealistic.

The concept of innateness is a part of folk wisdom but is also used by biologists and cognitive scientists. This concept has a legitimate role to play in science only if the colloquial usage relates to a coherent body of evidence. We examine many different candidates for the post of scientific successor of the folk concept of innateness. We argue that none of these candidates is entirely satisfactory. Some of the candidates are more interesting and useful than others, but the interesting candidates are not equivalent to each other and the empirical and evidential relations between them are far from clear. Researchers have treated the various scientific notions that capture some aspect of the folk concept of innateness as equivalent to each other or at least as tracking properties that are strongly correlated with each other. But whether these correlations exist is an empirical issue. This empirical issue has not been thoroughly investigated because in the attempt to create a bridge between the folk view and their theories, researchers have often assumed that the properties must somehow cluster. Rather than making further attempts to import the folk concept of innateness into the sciences, efforts should now be made to focus on the empirical questions raised by the debates and pave the way to a better way of studying the development of living organisms. Such empirical questions must be answered before it can be decided whether a good scientific successor – in the form of a concept that refers to a collection of biologically significant properties that tend to co-occur – can be identified or whether the concept of innateness deserves no place in science.