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CBE 2018 - A02 1 The ecological, social and cultural origins of primate intellect The evolution of intellect Hypotheses on the evolution of intellgence in human and non-human primates The “Technological” Hypothesis Oakley (Hominins only...) The Foraging Hypothesis Milton, Clutton-Brock The “Social Brain” Hypotheses (Dunbar et al) The “Machiavellian Intelligence” Hypothesis Byrne & Whiten, Cheney & Seyfarth The “Cultural Intelligence” Hypotheses van Schaik, Tomasello 1 2
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CBE 2018 - A02

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The ecological, social and cultural origins of primate intellect

The evolution of intellectHypotheses on the evolution of intellgencein human and non-human primates

The “Technological” Hypothesis Oakley (Hominins only...)

The Foraging HypothesisMilton, Clutton-Brock

The “Social Brain” Hypotheses (Dunbar et al)

The “Machiavellian Intelligence” HypothesisByrne & Whiten, Cheney & Seyfarth

The “Cultural Intelligence” Hypothesesvan Schaik, Tomasello

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The “Ecological” Hypotheses

The “Technological” HypothesisKenneth Oakley (1959): “Man, the toolmaker”

Jerison (1973):predators x prey

Ungulates’ and Carnivores’ brain sizes covaried through Paleogene / Neogene

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2001: A Space Odyssey (Stanley Kubrick, 1968)

Tools and hominization

Forest X savannaBipedalismFreeing of handsOpposition of thumbsEncefalization...

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“A mere 12,000 years separate the first bow and arrow from the International Space Station” (Ambrose 2001)

Tool use in nonhuman primates

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Tool use in non-primate vertebratesBottlenose dolphins (Krützen et al 2005)

Tool use in non-primate vertebrates

Egiptian vulture cracking ostrich eggOtter cracking oyster with stone

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Tool use in non-primate vertebrates

New Caledonian crows(Hunt 1996)

Tool use in non-primate vertebrates

Hyacynth macaws(Anodorhynchus hyacintinus)Borsari & Ottoni 2005

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Tool use in invertebrates

Octopus (Amphioctopus marginatus) with coconut shell (Finn & al 2009)

H

Ariadna spiders (Henschel 1995) Chimpanzee

Tools and behavioral plasticity

Köhler 1917“Species-specific” behavior (~”innate”)

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The “Technological” HypothesisKenneth Oakley (1959): “Man, the toolmaker”

Hominins:Changes in brain volume X Technological changes: LOW CORRELATION (Wynn 1988)

•Australopitecines x chimpanzees;•Technology and H. sapiens sapiens;•Exception: H. erectus ?

Primates: spontaneous and customary use of tools in the wild: FEW SPECIES

Homo spp, Pan troglodytes, Pongo pigmaeus, Sapajus spp, Macacafascicularis

X all primates’ encefalizaton

Environmental pressures in primates’ life: foragingNutritional needs and foraging efficiency:

Efficiency in food processing x Memory and cognitive maps

Evolutionary “Option”: investment in INTESTINES X BRAINS

Spider monkey (Ateles geoffroyi) frugivore, lone foraging, previous exploration XHowler monkey (Alouatta paliatta) folivore (low carbohidrate), symbionts

Spider monkey: twice the Howlers’ brain volume

Australopithecus x Homo: Brains ↑↑, Teeth ↓↓

Folivores x frugivoresCorrelation between home ranges and neocortex size (...?)

The Foraging HypothesisKatherine Milton (1993)

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R. Wrangham (2010): cooking

Raw x cooked food:Digestive tract ↓ x brains ↑More free timePair bonding, sexual division of labor

Encefalizationandsocial group size(Dunbar 1992)

Neocortical Ratio XHome range residue

disconting the effect of bodily mass

Neocortical Ratio XAverage group size

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The “Social Brain” Hypotheses

Primatology:Long-term field studies, research with captive groups:

Social complexity

AlliancesWarReconciliationStatus seekingDeception

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Frans de Waal (1982, 1989)

Primate sociality

de Waal:

primates are “prisioners” of societyconflict and reconciliation

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The “Machiavellian Intelligence” Hypothesis

Byrne & Whiten 1989 Whiten & Byrne 1997

Selection pressures from complex social life:All primates (except prosimians) are socialexception of the exception: social lemuroids from Madagascar (Jolly 1966)

Effects of social deprivation (Harlow)

The “Machiavellian Intelligence” Hypothesis(Humphrey, Jolly, Chance & Mead, Byrne & Whiten, Cheney & Seyfarth)

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Harlow, H. F. (1959) “Love in infant monkeys”

Harlow: social deprivation (Rhesus) → Bowlby: Attachment Theory (children)

Social Isolation inRhesus monkeys

Harlow, H. F. (1970)

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Selection pressures from complex social life:All primates (except prosimians) are socialexception of the exception: social lemuroids from Madagascar (Jolly 1966)

Effects of social deprivation (Harlow)

Social complexity:Hierarchies, alliances, reconciliation, grooming, food sharing...

“Neocortical suppression of autonomic functions”

(Chance & Mead 1953)

“Tripartite relationships” in baboons (Kummer 1967)

Conspecifics as “social tools” (Humphrey 1976)

The “Machiavellian Intelligence” Hypothesis(Humphrey, Jolly, Chance & Mead, Byrne & Whiten, Cheney & Seyfarth)

The “Machiavellian Intelligence” Hypothesis

Social cognition x cognition about physical worldIndividual (and relational) recognition of conspecifics:

Dasser 1988 (Macaca fascicularis) [match-to-sample]

Cheney & Seyfarth 1990 (Chlorocebus pygerithrus)

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[functionally]referentialcommunication

Alarm calls in vervet monkeys

(Cheney & Seyfarth 1990)

Chlorocebus pygerithrusOLD Cercopithecus aethiops

The “Machiavellian Intelligence” Hypothesis

Social cognition x cognition about physical world

Humans:

Anthropomorphism and animism(Piaget)

Children understand social causality before physical causality(Laurendeau & Pinard 1962)

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Cognitive adaptations for social life

PREDISPOSITIONS of the human infant for social exchange:Faces/voice discrimination (“motherese”)/ human movementsEarly “dialog” interaction (Papousek & Papousek).Newborns: “imitation” of facial expressions (Meltzoff & Moore)

[but see criticisms…]

Evolution of cooperation and detection of social rules’ violatorsCosmides & Tooby (1992): the ‘Wason Task”

Tactical deception (1)

Juvenile baboon cries near male (TARGET) with food, attracting his mother’s (SOCIAL TOOL) attention.

Juvenile pursued by adult (TARGET) adopts “predator monitoring” posture

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Dominant chimp stalks and exposes a cheating subordinate (F. Plooij)

Young gorila ‘building’ daily “nest” to get closer to baby without atracting mother attention (D. Fossey)

Tactical deception (2)

a: Grooming another animal to steal food (babboon);b: Harass adult male to obter gain maternal care (Theropithecus gelada)

Tactical deception (3)

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c: Neutral expression favoring approach to actually attack;d: Lead the march, but only to go back and seize food (chimpanzee)

Tactical deception (4)

Tactical deception X Neocortical Ratio

(Byrne 1995)

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Criticisms to the “Machiavellian Intelligence” Hypothesis

Deception episodes in nonhuman primates: alternative non-mentalist (associative) explanations more parcimonious

a - attribution of visual perspective (?)b - attribution of belief (??)

(Hamadryas baboon – after an observation by H. Kummer)

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Criticisms to the “Machiavellian Intelligence” Hypothesis

Complex sociality impliescomplex individual cognition?

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“Situated action”/ “Socially distributed cognition”(Strum et al 1997, Hutchins 1991, Johnson 2001)

Encefalization and social group size(Dunbar 2003)

Neocortex X social variables

• social group size• grooming clique size• social skill in male mating strategies• frequency of tactical deception• frequency of play

ALL SIGNIFICANTBUT

Neocortex X ecological variables

• fruit in diet• home range size• day journey length• foraging style (categories of extractive foraging)

NOT SIGNIFICANT(when ecological vars. corrected for relative body size)

(of course, correlation ≠ causality...)

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Criticism: Reader & Laland (2002)

Frequencies of behavioral innovation / social learning / tool use: allsignificantly correlated with absolute and relative volume ofexecutive brain (neocortex plus striate cortex) in primates, whenappropriate controls are made for phylogeny and research effort.

Innovation and social learning co-vary across species: underminesthe claim of an evolutionary trade-off between reliance on socialexperience and on individual experience.Reader & Lefebvre (2001): no correlation between the sociallearning index and social group size once brain volume has beentaken into account.

Ecological factors may have been as (or more) important thansocial factors in primate brain evolution.

Encefalization and social group size(Dunbar 2003)

Dunbar replies to R&L criticisms:

Behaviors analized by R&L all refer to foraging situations: they do not comparecognitive demands in ecological problem solving (including social learning tosolve these problems) X management of social relationships.

Lack of correlation between social learning index used by R&L and group size:

(1) Cognitive demands for purely social tasks not required for foraging (ToM?)(2) Social tasks involve components not included in the executive brain

(Limbic System?)

Dunbar (2003): frontal lobe: even better fit to social group sizeFrontal lobe + Amygdala: social emotions, moral judgement etcSize of basolateral complex of the amygdala correlates w group size

Does not discard a role for ecological cognition in the start of theencefalization process: ex: color vision

Encefalization and social group size(Dunbar 2003)

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Evolutionary encefalization and proportionof taxa in stablesocial groups(Shultz & Dunbar 2010)

Temporal trend in brain sizechanges: estimated andmeasured brain size andbody size for 511 extant andfossil species of mammals(six orders; only taxa whereboth fossil and extantspecies were available).

Encefalization and social group size(David-Barrett & Dunbar 2013)

Processing power limits social group size: computational evidencefor the cognitive costs of sociality

Sociality is primarily a coordination problem.D-B&D 2013: an agent-based model to test the hypothesis that thecomplexity of information processed influences the computational demandsinvolved.The social (or communication) complexity hypothesis: the kinds ofinformation that can be acquired and processed may limit the size and/orcomplexity of social groups that a species can maintain.Larger groups can only be achieved at the cost of more sophisticated kindsof information processing that are disadvantageous when optimal groupsize is small.For animals in smaller social groups, the cost of having a large brainoutweighs the benefits.These results simultaneously support both the social brain and the socialcomplexity hypotheses.

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The “Communication complexity” hypothesis(Dunbar)

Origins of languageMusic: common origin or “cheesecake” (Pinker)?

“Hmmmm” (Mithen)

The “Cultural Intelligence” hypothesisvan Schaik & Pradhan 2003Whiten & van Schaik 2007

Positive selection of characters associated to social learning

Evolution of social tolerance

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Social cognition and “Theory of Mind”:anecdotes x experiments

Premack & Woodruff 1978

Povinelli & Vonk 2003

Tomasello, Call & Hare 2003

Do primates have a “Theory of Mind”?Understanding the other

as a causal agency

X understanding mental states

“Theory of Mind” (ToM): attribution of mental states to outher individuals, using them to predict and explain their behavior.

Premack & Woodruff (1978). Does a chimpanzee have a “Theory of Mind”?

A “limited” ToMin chimpanzees?

Povinelli et al 1994

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Mirror self-recognition

Chimpanzees,

(Gallup & Povinelli)

bonobos,

orangutans.

Gorillas?

but also

elephants,

dolphins,

corvids...

Povinelli et al 1998

Understanding the difference between intention and accident?

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Povinelli, Nelson & Boysen,1990

X

Povinelli & Eddy, 1996

Understanding the relation between seeing and possession of information?

Esta é a SallySally põe suabola na caixa

Sally vai embora

Aonde a Sally vaiprocurar pela bola? Esta é a AnneAnne move a bolapara o seu cesto

Sally & Anne

© e.b.ottoni 2007

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“Theory of Mind” (ToM)and child development

The Sally-Anne task (Baron-Cohen,Leslie, & Frith, 1985)

ToM e Autism/AspergerBaron-Cohen 1995“Mindblindness”

Reality-biased responses: verbal tasks?“Where” question prematurely interpretedby young children as referring to thelocation of the hidden object?(Southgate et al 2007) The “Sally & Anne” [or “false belief”] test

Onishi & Baillargeon 2005Do 15-Month-Old Infants Understand False Beliefs?

Nonverbal task: violation-of-expectation

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The “Cultural Intelligence” hypothesis

Michael Tomasello (1999):

Animal “culture” based in distinct/simpler processes (“stimulus enhancement”)

Theory of MindJoint AttentionImitationTeaching

The “ratchet effect”: cumulative cultureTomasello (2016): [imitation + conformity] + teaching

Subordinate X Dominant:visual barrier

Understanding the relation between seeing and possesion of information?M. Tomasello & J. Call

Call 2001

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Call 2001

Informed dominant replaced byanother, uniformed one

Uninformed / misinformed dominant?

Understanding the relation between seeing and possesion of information?M. Tomasello & J. Call

Krupenye, Kano, Hirata, Call & Tomasello 2016

Great apes anticipate that other individuals will act according to false beliefs

Chimpanzees (Pan troglodytes), bonobos (Pan paniscus) & orangutans (Pongo abelii).Familiariz. FB1 FB2Familiariz. FB1 FB2

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Great apes distinguish true from false beliefs in an interactive helping taskButtelmann et al 2017 (16/18-mo children: Buttelmann et al 2009)

Helping behavior: true x false belief x ignorance conditions

“Niveis” de Teoria da Mente?

Leslie 1994:

ToBy (“Theory of Body”): agentes x não-agentes (início do 1º ano)

ToM I: pessoas como entes que percebem o ambiente e buscam certos propósitos (“agentes intencionais”) (final do 1º ano)

ToMII: mecanismo para lidar com os agentes como seres capazes de atitudes proposicionais (acreditar/ imaginar/ fingir) falsas ou verdadeiras (2º ano).

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Teoria da Mente: competências distintas?

Compreensão de intenções, desejos, pensamentos, conhecimento, crenças... Provavelmente não se desenvolvem ao mesmo tempo.

Bartsch & Wellman (1995)

Sequência de desenvolvimento:

1. 2 anos: compreensão de estados como atenção, emoções e percepçõessimples (não-representacional, sem interpretação das conexões daspessoas com as coisas em termos de “estados mentais”);

2. 3 anos: “pensamentos” e “desejos” como representações falsas ouverdadeiras, que podem variar de pessoa para pessoa;

3. 4 anos: ToM adulta, onde desejos e crenças interagem na causação docomportamento.

ToM and Levels of Intentionality(Dennett, DC)

Dunbar 2003:

Monkeys: 1st orderChimpanzees: 2nd orderAdult humans: 4th order

Ottoni, Rodriguez & Barreto 2006

3-13 yo:

• Understanding graphic representation (“thought bubbles”): 4/5 to 7 yo

• Understanding of “false belief”: 4/5 - 9 yo• 2nd / 3rd order (“nested”) representations: 9 yo.

La Frenière 1988The ontogeny of tactical deception in humans

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Evolutionary hypotheses Nature of organism [genotype / phenotipe] / environment interaction

“Technological Intelligence”

Inheritable individual variation + Natural selection1 information channel (genetics)

Humans

“Foraging Hypothesis”(Brains x Guts)

Inheritable Individual variation + Natural selection(with 2 hypothetical evolutionary “trajectories”)1 information channel (genetics)

Primates

“Machiavellian/Social Intelligence”

Inheritable individual variation + Natural selectionIntraspecific competitionSexual selection (Miller 1999)1 information channel (genetics)

Primates

“Cultural Intelligence” 2 information channels•Inheritable individual variation + Nat. selection•Non-genetic Social Information TransferNiche Construction: phenotypes as AGENTS of the evolutionary process (change selective pressures acting on genes)Gene-Culture Coevolution

Mammals and birds?

(... not necessarily mutually exclusive)

Evolutionary approachesto human behavior

Next:

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Adaptações cognitivas para a socialidade

PREDISPOSIÇÕES do bebê humano para a interação:Discriminação de rostos/ voz (“motherese”)/ movimento humanosInteração “dialogada” precoce (Papousek & Papousek).Recém-nascidos: “imitação” da expressão facial (Meltzoff & Moore)

Michael Tomasello: cultural learning and joint attention

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“Teoria da Mente”: Modulos Inatos x Construção de Teorias?

Autismo (Asperger)X

Sindrome de Williams

Prosopagnosia

“Estágios” no Desenvolvimentoda Teoria da MenteX

Flavell (1999): uma teoria adequada terá de integrar ambas as perspectivas...

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