Animal Mind & Evolutionary Psychology: Two sides of the same coin

1. AnimalMind & EvolutionaryPsychology: Two sides of the same coin • Animal Mind: How much are animals like us? • Evolutionary Psychology: How much are we like animals? • Both try to reconcile biology and psychology. • Concepts that prove central to one, turn out to be central to the other: • Learning • Prosociality • Theory of mind • Personality (‘behavioral syndromes’)

2. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 • Definitions of emotions typically describe proximate aspects such as • Physiology • Subjective experience • Facial expression • An evolutionary approach defines what emotions are in terms of how they came to exist. • Emotions are modes of functioning, shaped by natural selection, that coordinate physiological, cognitive, motivational, behavioral and subjective responses in patterns that increase the ability to meet the adaptive challenges of situations that have recurred over evolutionary time (Nesse, 1990).

3. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 but only in certain situations. • They are adaptations that are useful • Like pain and sweating, they remain latent until an evolved mechanism detects cues associated with the situations in which they are advantageous. • Unlike simpler adaptations, however, emotions are not unimodal responses to specific situations, like sweating in response to overheating. Instead, emotions adjust multiple component processes to create an organized response to the adaptive challenges of a given situation. • Example: appraisals that indicate a nearby predator arouse an emergency response that adjusts and coordinates many aspects of physiology, cognition and behavior.

4. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 • Questions about an emotional trait: • What trait is like, how it works; • How it develops over the individual’s lifetime; • How it developed over time in the history of the species; • What evolutionary factors shaped the trait. • i.e., Tinbergen’s Four Questions

5. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 • How many emotions exist, and what are they? • Some theories: just two basic states – positive and negative • Other theories: small set of basic emotions • Still other theories: potentially infinite number • Agreement: valence is a necessary quality of emotion: pleasure & pain, approach & avoidance fundamental • Many models array the emotions on these dimensions, e.g., the circumplex model (Posner et al 2005)

6. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139

7. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139 • A phylogeny of emotions: specific emotions partially differentiated from more primal generic states because they improved ability to copy with specific kinds of threats and opportunities. • New emotional variations are useless unless they are expressed in situations where they are advantageous. • Emotions do not have clear boundaries. No clear taxonomy of emotions possible.

8. Nesse & Ellsworth (2009) Evolution, Emotions & Mental Disorders American Psychologist 64: 129-139

9. Animal Emotion

10. Inside the Animal Mind: Do Animals have Emotions? Jane Goodall versus Marian Dawkins Goodall just asserts that animals have emotions – its obvious! – and says that emotional similarity is most likely in animals closely related to us (like chimps).

11. Inside the Animal Mind: Do Animals have Emotions? Jane Goodall versus Marian Dawkins Dawkins emphasizes that emotions are likely to be different in different species – and so we should not anthropomorphize. But she also says that we should give animals the benefit of the doubt and assume they do have emotions unless we somehow get evidence otherwise. ☞Issue important because of concerns about animal welfare. General point: absence of evidence ≠ evidence of absence

12. Animal Emotion • Dawkins: Facial expressions are not a reliable signal of emotions in most other species. • Hard for us to see this because we expect animals to express emotions in the same way as we do. • Not expressing emotions for communication purposes is not the same thing as not having emotions.

13. Animal Emotion 1. attention2. excitement3. narrow smile4. broad inviting grin5. laughing6. crying

14. Parr, L. A. & Waller, B. M. (2006) Understanding chimpanzee facial expression: insights into the evolution of communication. Social, Cognitive and Affective Neuroscience 1: 221-228 Chimpanzees display a complex, flexible facial expression repertoire with many physical and functional similarities to humans. To understand the evolution of emotional communication, comparative research on facial expression similarities between humans and related species is essential. This paper reviews what is known about these facial expression repertoires and introduces a new coding system: ChimpFACS. • It describes how ChimpFACS can be used to determine homologies between human and chimpanzee facial expressions. It reviews previous studies on the categorization of facial expressions by chimpanzees using computerized tasks, and discusses the importance of configural processing for this skill in both humans and chimpanzees.

15. Parr, L. A. & Waller, B. M. (2006) Understanding chimpanzee facial expression: insights into the evolution of communication. Social, Cognitive and Affective Neuroscience 1: 221-228 Researchers have developed an objective, standardized method for measuring facial movement in the chimpanzee that is directly comparable to humans, thus facilitating research on facial expression homologues (www.chimpfacs.com). This system, referred to as ChimpFACS, is based on the well-known human Facial Action Coding System, or FACS, developed by Ekman and colleagues to provide an objective tool for studying the biological basis for human expressions and emotion. Both of these systems identify the most minimal units of facial movement according to the function of the underlying musculature. Therefore, both systems enable the objective description of facial appearance changes in the human face and the chimpanzee face that are associated with movements of the underlying facial musculature. Each movement is described using a numeric code, referred to as an Action Unit (AU), and researchers are trained to use these codes reliably through a standardized testing process, making all users of the system statistically reliable with one another (http://face-and-emotion.com).

16. Parr, L. A. & Waller, B. M. (2006) Understanding chimpanzee facial expression: insights into the evolution of communication. Social, Cognitive and Affective Neuroscience 1: 221-228 Because the facial musculature of chimps and humans is highly comparable, the two systems provide a basis for understanding homologous facial expression structure in the two species. Because the FACS and ChimpFACS systems are both standardized, researchers from different groups are now able to compare their results directly using a common nomenclature. Thus researchers wanting to determine whether expressions like the bared-teeth display (chimp) and the smile (human) are homologous, can begin by using subjective ratings of their similarity, but then utilize the more objective, standardized assessment of their anatomical similarity by comparing individual action units in both species. To test whether this bottom-up approach (action unit to expression configuration) could actually validate the existing categories of chimpanzee facial expressions made previously by expert observers of chimpanzee communicative behavior, the authors used a discriminate functions analysis (DFA) to determine whether the a prioricategories of chimp expressions (made by Parr) categories could be predicted by common patterns of underlying facial movements, or patterns of AUs, coded from each expression using ChimpFACS (by Waller, “a certified ChimpFACS expert”).

17. Parr, L. A. & Waller, B. M. (2006) Understanding chimpanzee facial expression: insights into the evolution of communication. Social, Cognitive and Affective Neuroscience 1: 221-228 Computer-generated faces in the left columns were produced with the software Poser (“the world’s easiest way to create art and animation using 3D characters”). The Poser animated expression, on the left is compared with the natural chimp expression on the right. Under the Poser expression is the prototypical AU configuration as identified by the DFA. Under the natural photo is the % of category agreement between the AU configuration and the aprioriclassification. (Pout is the only one that doesn’t work well!)

18. Parr, L. A. & Waller, B. M. (2006) Understanding chimpanzee facial expression: insights into the evolution of communication. Social, Cognitive and Affective Neuroscience 1: 221-228 Prototypical chimpanzee facial expressions and homologous facial movements in a human

19. Parr, L. A. & Waller, B. M. (2006) Understanding chimpanzee facial expression: insights into the evolution of communication. Social, Cognitive and Affective Neuroscience 1: 221-228 An alternative approach is to first match human and chimpanzee expressions based on the muscular components, and then make inferences about the emotional quality of human faces that are structurally homologous to chimpanzee expressions. This figure shows such a comparison where the human images are taken directly from the FACS manual (Ekman et al2002) and show facial movements that are structurally homologous to the prototypical chimpanzee expressions. The identical AUs shared by the two expression examples are highlighted in bold italics. Previous researchers have suggested a homology between the chimpanzee bared-teeth display and the human smile. However, the human bared-teeth expression in the figure resembles a grimace, or a forced smile, and it does not suggest happiness. The chimp play face suggests happiness and has been said to be homologous with laughter in humans.

20. Continuity of facial expressions in primates? From Jolly 1972

21. Continuity of facial expressions in primates? From Jolly 1972

22. Central Issues • Do we need to posit emotion to explain response (e.g., fear to explain response to danger)? • What actually is the question we’re asking? • Do animals have emotions? • Do they have emotions similar to ours? • What does it take to answer these questions?

23. What is Emotion? Do we need “fear” to avoid the predator? Is “fear” just our perception of our internal physiological changes accompanying the innate flight or fight response? James-Lange theory

24. James-Lange Theory of Emotion “My theory ... is that the bodily changes follow directly the perception of the exciting fact, and that our feeling of the same changes as they occur is the emotion. Common sense says, we lose our fortune, are sorry and weep; we meet a bear, are frightened and run; we are insulted by a rival, are angry and strike. The hypothesis here to be defended says that this order of sequence is incorrect ... and that the more rational statement is that we feel sorry because we cry, angry because we strike, afraid because we tremble ... Without the bodily states following on the perception, the latter would be purely cognitive in form, pale, colorless, destitute of emotional warmth.” William James

25. Animal Emotion – a Potpourri Pet behavior problems Neurobiological perspectives Love, lust – hormonal mechanisms Do animals fall in love? Pleasure centers in the brain Baby faces Animal welfare and Mark Bekoff

26. Pet Behavior Problems: Can we just assume they had unhappy childhoods?

27. Pet Behavior Problems: or that they were just swayed by Devil Dog?

28. Lust, Attraction & Attachment in Mammalian Reproduction Helen Fisher, Arthur Aron & Lucy Brown Principle sources and targets of hormones and neurotransmitters implicated in emotions of lust(androgens & estrogens, gonads), attraction (catecholamines, LC, VTA, SN), and attachment (oxytocin & vasopressin, PVN/SON of hypothalamus).

29. Do Animals Fall in Love? • Peacock no • Prairie vole yes – life-long pair bonds • Oxytocin (& vasopressin) • “Their chemistry may be, well, just chemistry” • Oxytocin involved • in parent-child bond too

30. Primary Emotions • fear, sex • primitive brain, limbic system (esp amygdala) • Secondary (self conscious) Emotions • shame, guilt, embarrassment • require self-awareness (test: mirror recognition test) • Neocortex • Affection, Jealousy – primary or secondary? • Same chemistry = same emotions (e.g., the Prozac test)

31. Pleasure Centers of the Brain James Olds and Peter Milner in the 1950s probing the limbic systems of rats looking for pain centers, found instead “pleasure centers”: rats would respond at high rates for stimulation of their septal areas, located near the brain stem and among the oldest areas of the brain. Rats would cross electrified floors to press a self-stimulation switch, and would press it 1000s of times to the exclusion of food or water. Female rats would even abandon their unweaned pups to self-stimulate. Key centers were medial forebrain bundle (MFB) and ventral tegmentum (VT)Dopamine appears to be the most important neurotransmitter involved.

32. Pleasure Centers of the Brain Right here!

33. Pleasure Centers of the Brain Only a few experiments have been conducted involving the electrical stimulation of human pleasure centers. Generally these investigations are considered taboo. In the 1970s, Robert Heath, who believed he could "cure" homosexuality, wired up gay volunteers to an electrical apparatus that directly stimulated their nucleus accumbens, producing feelings of extreme pleasure. Given the choice, one man, code-named B-19, electrically self-stimulated his reward circuitry some 1,500 times in a 3-hour session (“he had to be disconnected, despite his vigorous protests”). Few experiments directly stimulating the human pleasure centers have been conducted since.

34. Baby schema modulates the brain reward system in nulliparous women Glocker et al PNAS 2009 Ethologist Konrad Lorenz defined the baby schema as a set of infantile physical features, such as round face, high forehead and big eyes, that is perceived as cute and motivates caretaking behavior in animals including humans, with the evolutionary function of enhancing offspring survival. “Using functional magnetic resonance imaging and controlled manipulation of the baby schema in infant faces, we found that the baby schema activates the nucleus accumbens, a key structure of the mesocorticolimbic system mediating reward processing and appetitive motivation, in nulliparous women. Our findings suggest that engagement of the mesocorticolimbic system is the neurophysiologic mechanism by which baby schema promotes human caregiving, regardless of kinship.”

35. Baby schema modulates the brain reward system in nulliparous women Glocker et al PNAS 2009

36. Baby schema modulates the brain reward system in nulliparous women Glocker et al PNAS 2009

37. Marc Bekoff

38. Animal Emotions: Exploring Passionate Natures Marc Bekoff “Emotions can be broadly defined as psychological phenomena that help in behavioral management and control”. Well that’s a nice definition, but how exactly do the emotions do this? Sometimes they seem disruptive, not particularly helpful. For example, Spock→ seems to do better without emotions!

39. Animal Emotions: Exploring Passionate Natures Marc Bekoff Much of Bekoff’s argument seems to rely on a combination of an appeal to authority (Jane Goodall, Joyce Poole, et al) and compelling anecdotes or commentaries involving charismatic animals (e.g., chimps, elephants). e.g.:

40. Animal Emotions: Exploring Passionate Natures Marc Bekoff The study of play has always been an occasion to talk about animal emotions, perhaps because it seems to be an activity engaged in just for the fun of it. (Though contemporary thinking emphasizes its role in the development of necessary adult skills that may be used in hunting, fighting, etc.)

41. Animal Emotions: Exploring Passionate Natures Marc Bekoff Sadness & Grief Many of examples have accumulated showing apparently sad or grieving animals after loss of child or parent or mate, e.g., Goodall’s story of the chimp Flint’s (terminal) depression following the death of his mother Flo.

42. Animal Emotions: Exploring Passionate Natures Marc Bekoff Developing comparative evidence Taking off from the maxim “absence of evidence is not evidence of absence”, I would agree that the burden of proof is equally on skeptics (animals do not have emotions) and true believers (animals have human-like emotions). But the key will be to develop methods that can convincingly reveal the nature of emotional states (similarities and differences) in different animals.

46. Lust, Attraction & Attachment in Mammalian Reproduction Helen Fisher, Arthur Aron & Lucy Brown Principle sources and targets of hormones and neurotransmitters implicated in emotions of lust(androgens & estrogens, gonads), attraction (catecholamines, LC, VTA, SN), and attachment (oxytocin & vasopressin, PVN/SON of hypothalamus).

50. Animal Emotions: Exploring Passionate Natures Marc Bekoff