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1. Introduction: The Science Of Animal Motives
Classical ethology, the ‘biology of animal behavior’ (Eibl-Eibesfeldt 1970, Hinde 1982), compares descriptions of how diﬀerent species act in their natural environment, and in families and societies. Lorenz proposed that genetically determined motor programs or Innate Release Mechanisms (IRMs), triggered by environmental Sign Stimuli, confer ‘dispositions to learn’ that beneﬁt the species. By comparing spontaneous behavior rituals and sequences of display or response between individuals, ethologists seek evidence of innate impulses in diﬀerent species, and determine the genetic relationships and evolutionary lineage. Ethology developed from zoology and ﬁeld studies of wild animals. It accepts that animals have motives and awareness that can be understood by accurate description of their behaviors in nature, and how these develop. Darwin’s Expression of the Emotions in Man and Animals (1872) inaugurated ethological science. He related human feelings to those one can infer in animals from their expressions.
Tinbergen (1951) identiﬁed innate signaling behaviors of communication—in mating, nest building, care of young, territorial defence, hunting or escape from predators, sharing environmental resources, etc. He deﬁned four time-scales of behavioral determination: over evolutionary time; in development of the individual; in control of behavioral tendencies over hours, days, or months; and in adaptive functional response to immediate environmental events. He used ﬁeld experiments to identify that stimuli or behavior signals are ‘releasers’ of innate activities.
Scientists research animal senses, often very different from ours, and the ways animals move to investigate their environment. Communicative signals are instructive, because they have evolved to be perceived. Rhythmic light ﬂashes, visible movements of body parts, vocal or other sounds, or vibrations, convey eﬀective messages, even in primitive species. How animals react to signal patterns gives further evidence about internal processes that guide attention, movements, and emotions. In humans, innate hearing and vision has been explored by studying the vocal sounds, face expressions, and body movements that infants prefer, and the emotions infants express (Fernald 1992, Papousek and Papousek 1987, Stern 2000, Trehub 1990, Trevarthen 1998, 1999).
As in Darwin’s day, evidence for species-speciﬁc motives and feelings in humans comes from comparing spontaneous expressions of people in diﬀerent cultures, and by observing infants before they have had the chance to learn (Eibl-Eibesfeldt 1989). Newborn infants communicate and imitate, and children show systematic changes in behaviors and interests (Stern 2000, Trevarthen 1998). As children’s motives develop, they change the behavior of caregivers, stimulating ‘intuitive parenting’ and the sharing of meanings (Papousek and Papousek 1987).
2. Learning, Cognition, And Communication
Eﬀorts have been made to relate ethological theory to concepts of experimental psychology and ‘conditioning’ in simpliﬁed laboratory situations (Hinde 1982). After decades of rejection by psychologists of ‘mentalistic’ interpretations, the idea that animals may share motives, feelings, and conscious experiences comparable to those of human beings has again become scientiﬁcally acceptable. Psychologists postulate internal ‘representations’ that process perceptual information and control actions of subjects in complex circumstances. More recently, attention has turned to the evolved functions of emotional regulatory systems in the brain, and their role in self-awareness and intelligence (Damasio 1994).
Cognitive psychology acknowledges that human behavior is regulated by conscious states and intentions, and that learning integrates emotions and mental representations. Nevertheless, cognitive science remains individualistic and rational—focused on reality. Complex intelligence is analyzed in terms of perceptual categorizations and hierarchical representational systems. Diﬀerent modes of intelligence are thought to be formulated anatomically in identiﬁable brain regions. In particular, social intelligence is considered to pass, in evolution and development, through grades of cognitive representation, and levels of awareness of cognition in other individuals—from immediate, ‘thoughtless’ sensorimotor reactions, to verbalizable, rationally formulated ‘theories of mind.’ Dynamic Systems Theory, opposed to linear causal explanation, attributes behavior patterns and mental representations to ‘emergent properties’ thrown up in complex systems comprising large numbers of elements—neural connections, perceptual feature detectors, sensorimotor loops, motoric forces, representations in memory, social transactions, etc. In neither of the dominant contemporary theories of complex mental life and behavioral coordination is evolved anticipatory motivation given a determinant role.
No adaptive behavior of an animal subject is without innate ‘constraints’ due to prefunctional conﬁguration (morphogenesis) of sensory and motor structures, brain tissue, and nerve connections, which map both the body and its future engagement with the outside world in a coherent, subjective space and time. What is innate is an active neurobiological initiative that investigates the environment, applying speciﬁc anticipatory, probing processes of behavior. ‘Critical periods,’ or ages in which active investigations of experience change, correspond to brain growth periods that are generally more open to learning.
3. Behavior From Dyads To Large Social Groups
Subjective motives communicated to other individuals become intersubjective (Braten 1998). In the more intelligent, behaviorally complex species there is a rich capacity for display of mental impulses—of attention, intention, and emotion. Intersubjective relations assume central importance in reproduction, feeding, establishment of cooperative groups in families or much larger societies. Interspeciﬁc collaborations, including predator–prey interactions, control behavior in dyads and larger groups, regulating ecological relations between animals psychologically.
Sociobiology has formulated theories to explain the evolutionary advantages of various forms of conﬂict and cooperation within and between communities, and special complementary adaptations of individuals in the service of the group, or to propagate a genetic type. Reductive cost-beneﬁt analyses, of energy balance sheets for motor and metabolic investment, or of transmission and survival of genes in populations (reproductive success), have prevailed, which can be explained by an unwillingness of quantitative biologists to attribute self-regulatory motives and emotions to animals as individuals, or members of families and cooperative groups. They see behaving animals as driven by forces of competition for future success that are indiﬀerent to any aspirations for, or pleasures in, life in the present. The genetic history of humans indicates that there has been rapid adaptation to an extremely large range of environments, and new adaptability by cooperative learning of behavioral traditions (Cavalli-Sforza and Cavalli-Sforza 1995).
The beneﬁts of competition have been exaggerated. Research on the psychopathology of infanticide or mass killing by individuals indicates that direct competition by assault results from a loss of pleasure in the kinds of altruistic and aﬀectionate motives of play that lead to creative invention of new and beneﬁcial motives, ideas, or activities in a group (Bekoﬀ and Byers 1998). Communication between members of a species and between diﬀerently adapted species without concern for gain leads to intricate dynamic ecological systems and mutual dependency. Some behaviors of predator and prey in natural ecological equilibrium are cooperative, or mutually supportive. Skilled collective understanding of the environment by intelligent social species depends on positive motives for sharing enjoyment of experience and skilled action, such as are abundantly shown by the young of many species when they play together.
4. Attachment, Companionship, And Parental Support Of Early Development
Emotions linked to innate behaviors regulate parent–oﬀspring attachments and caregiving. In many species, mutual adaptations of motives in parents and young facilitate cooperative encounters that nurture the young, and parenting strategies limit the oﬀspring’s dependency. The food, care, and support that nestlings or nurslings receive sustain the developing body and brain. Human infants show attachment to the physiological or bodily beneﬁts from mothering (Schore 1994), the biological importance of which was ﬁrst demonstrated experimentally in monkeys. Infants also respond to psychological expressions of pleasure and interest carried by the voice, facial expressions, and gestures of the parent, and these lead to ‘protoconversations’ and companionship (Trevarthen 1998, 2001b; Trevarthen and Aitken 2000). Infants and parents seek contact and aﬀective communication, and deprivation of sympathetic parenting has detrimental eﬀects on both emotional and cognitive development of a child.
Pleasure and pain may be the main emotions or neurochemical systems seeking parental nurturance in mammals, including humans (Panksepp 1998), but more complex emotions, including self–other conscious states and relational emotions such as pride and shame, regulate intersubjective motives and assist the learning of a wide range of socially and culturally important skills in infants (Stern 2000, Draghi-Lorenz et al. 2001). Infants seek emotional conﬁrmation to develop knowledge and skills (Trevarthen and Aitken 2000).
5. Motives And Beneﬁts Of Play
Animal motives are revealed not only in behaviors that immediately beneﬁt the individual, or the propagation and ﬂourishing of the group or species. Their essential creative aspect is vivid in play (Bekoﬀ and Byers 1998). Ethological and psychological theories of play appear confused by a desire to ﬁnd a function outside motivation itself. However, by its nature, and by any deﬁnition, play behavior largely is determined by the anticipatory processes of behavioral control, and not any immediate adaptive consequence. Play behaviors are more apparent in the more social species that have elaborate intersubjective capacities or social awareness. By giving strong and continuous messages of exuberant motives, play invites sympathetic attention and, indeed, participation by others in changing outcomes and risks. Human children, like young of other species, can play alone, practising and shaping perceptuo-motor skills. But behaviors exhibited in self-satisfying play have a potential to be social, and they are responsive to provocative or ‘teasing’ interactions with other individuals. Play ﬁghting between young in a cooperative family group is aﬃliative, consolidating relations and training awareness of mutual intentions. Many play behaviors anticipate functions of mature ‘innate’ behavior patterns. Thus, play expresses motives for behaviors that require exercise of motor and cognitive processes for their maturation. It would seem pointless to attempt a cognitive rationalization of play if play is the prior behavior, adapted to generate intelligence and regulate cognition.
Research with infants conﬁrms that play develops relationships and skills, as well as creating rituals and ‘formats’ of cooperation. The imaginative or ‘symbolic’ play of young children foreshadows the creative eﬀorts made by young adult athletes, and the striving of outstanding craftsmen, artists and scientists throughout their lives (Bekoﬀ and Byers 1998). Art, appreciation of beauty in artefacts and ceremony in human communities may be related to universal rhythms and modes of intimate communication between infants and their partners (Dissanayake 2000).
6. ‘Communicative Musicality’ And The Pulse Of Human Thought And Imagination
Animal communication employs precisely timed rhythms, modulations of expressive activity, and accurate interaction of movements in communicating individuals (Tinbergen 1951). Infants’ expressions, and their reﬁned discrimination of temporal, prosodic, harmonic, and melodic features of vocalization or musical sounds, beginning before birth, indicate that human language communication draws on rich preverbal talents for expressive gesture and perception (Trehub 1990). Human sensitivity for the emotive rhythms, qualities, and narrative sequences of expressive song and dance, may be evolved with bipedal standing and walking, which free the erect body, and especially the hands and voice, for ﬂexible polyrhythmic gestures that can richly express dynamic emotions and their blends and changes in interpersonal engagements (Trevarthen 1999). Emotions are conveyed in regulated ‘contours’ or ‘melodies’ of muscle activity. They are held together by a coherent time-generating process in the brain that creates a hierarchy of acts and succession of perceptual events in one ‘mind time’ (Poppel 1994). Infants discriminate the pulse, expressive quality, narrative succession, and contingency of response in adult expressions.
7. The Biology Of Human Imitation, Cultural Learning, And Language
Monkeys and apes learn in social play, and imitate, but parental support for learning is limited in subhumans, and the most elaborate monkey play is between peers. Apes play over a wider range of ages, and their play assists development of family and group bonds, and they imitate how adults forage and use transformed objects as ‘tools’ for getting food. No apes show the intricate intersubjective exchanges of human vocal and gestural play with infants soon after birth, or the constructive and narrative-making fantasy games of toddlers. Donald (1991) argues from comparative cognitive psychology and the fossil record that it is likely that the ﬁrst bipedal hominids were communicating nonverbally, by gestural mimetic narratives, about nonpresent experiences. He concludes that grammar of language evolved out of the rhythmic syntax of gestural mimesis. Study of developments towards language of infants and toddlers supports this theory (Locke 1993).
Human beings have unique motives for cultural learning—the acquisition of artiﬁcial knowledge and skills. These govern progress from preverbal forms of communication and play (Trevarthen 1998). Human ethology is necessarily a science concerned with these motives. All cultural cooperation among humans, however artiﬁcial, depends on a uniquely productive capacity for ‘mirroring’ feelings, interest, and purposes of other persons, and for solving cognitive, practical, moral, and artistic problems collaboratively (Bruner 1996). Studies of imitation reveal that a neonate human has a remarkably complete image of movements in the expressive parts of another person’s body, an image that is both receptive through the constellation of all senses, and capable of forming a matching movement in communicative interactions (Braten 1998, Nadel and Butterworth 1999).
Human language learning depends upon joint attention and sympathetic understanding of intentions (Tomasello and Farrar 1986). Toddlers acquire use of words by following convergent interest to grasp meanings in other persons’ speech (Locke 1993). Long before verbal communication, an infant is proﬁcient in ‘protoconversational’ games, and in participation in rhythmic rituals of action games and songs, which parents are highly motivated to share, intuitively matching and complementing the emotions expressed by the infant (Trevarthen 1998).
8. The Evolutionary Neuroembryology Of Motives And Emotions
Discoveries in neurobiology enhance our knowledge of adaptive motive systems and emotions and their role in self awareness and communication (Schore 1994). Brain mechanisms of emotion in brainstem regions inﬂuence the higher functions of imagination, thought, and language (Panksepp 1998) and exercise an inﬂuence over the maturation of cortical cognitive systems in both prenatal and postnatal development of mammals, including human beings (Trevarthen 2001a). Cortical mechanisms for perceiving postural, facial, and vocal expressions, as well as manual gestures, for remembering experiences associated with diﬀerent expressed emotions, and for generating such expressions as elaborate narratives of internal impulses and feelings, are enlarged greatly in humans as compared to homologous brain areas of other species (Damasio 1994, Trevarthen 2001a). Play is associated with manifestations of neurochemical well-being or ‘happiness’ (Panksepp 1998). Often young animals play vigorously with body movements at a particular age, and it has been noted that in this ‘sensitive period’ the cerebellum of the hind brain is undergoing accelerated growth and acquiring reﬁnements of circuitry necessary for accurate and rapid motor control.
9. Future Directions
Only recently has over two centuries of research on the comparative anatomy of brains and their development been comprehensively integrated with psychological theories of cognition and learning. A focus on the forebrain cortex, and the eﬀects of cortical lesions on human awareness, memory, and motor skills had left the core systems that incorporate motivational and emotional controls out of the picture. Recognition that acquired skills and knowledge are directed, focused, selected, and integrated by subcortical systems leads to questions about how adaptations of behavior and awareness that anticipate experience and shape learning are generated. Ethological theory directly supports new enterprises in human genetics, neuroethology, and developmental neuroscience, and applied ﬁelds, such as education and psychiatry. We anticipate a more balanced natural science of mental life integrating the processing of empirical experience and reason with inherited emotive impulses for both individual experience and cooperative social life. Important questions will arise concerning what determines human aﬀairs and our relation to the natural world.
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