Evolution, Psychology, Evolutionary Psychology and Human Uniqueness Stephen Walker MSc Psychological Research Methods PSYC073P Epistemology and Philosophy of Science December 12th 2006 Overview • There will inevitably be some overlap with nature/nurture issues (Dec 5th) • There is also a a contrast between evolutionary or biological psychology and social constructionist or psychosocial approaches (Nov 28th) • At least two kinds of reductionism will be illustrated: explaining behaviours as adaptations, and explaining behaviours in terms of neural circuits that control them. Outline • I will start off by looking at animal behaviour, where evolution has relevance. • “Evolutionary Psychology” is based on the claim that human psychology is strongly determined by human evolution. • The evidence for these claims is often extremely weak, but I will look at a few recent examples. • I will also look briefly at the evidence concerning the course of human evolution. Paper Handout • *NB* the paper handout includes only a small fraction of the slides in this presentation: a pdf of most of the slides will be available on the intranet • The paper handout includes a list of alternative books on human evolution which are in BK library on page 6 • Any pieces of work mentioned in the presentation should have its citation listed on pages 7 & 8 of the handout. Fossils versus Genomics • There are epistemological difficulties in studying human evolution. • The difficulties are in inferring the past course of human evolution from a limited number of fossil finds. • But in the last 10 or 15 years technologies have become available allowing geneticists to pinpoint where and when the human genome has undergone significant changes. Evolution and DNA There is a huge amount of information pertinent to evolution produced by recent technologies: these are citations for the paper by Altschul et al. (1990) about a search tool for DNA and protein sequence databases Wikipedia says this was the most widely cited of all scientific papers published in the 1990s Genomics and Bioinformatics • DNA sequencing, inc human and chimpanzee • Gene splicing and genetic engineering • Gene expression data from ‘Microarrays’ The Darwinian Theory of Evolution − “Descent with Modification” There are inherited differences between individuals These include random variations Resources are not unlimited Some individuals will flourish more than others and produce more offspring Natural selection occurs if a population changes over generations because of this Evolution — II • The first point about evolution is that it connects the human species with the rest of the animal kingdom, • However, it is also possible and indeed likely that the course of human evolution has led to humans being uniquely different from all other currently living species Human Uniqueness on behavioural grounds Almost all animal behaviour is genetically pre-programmed (by evolution) to fit an ecological niche (Darwin, 1859; Tinbergen, 1951; Manoli and Baker, 2004) Almost all human behaviour involves cultural learning (Tomasello and Razoksky, 2003; Tomasello et al., 2005) Darwin (1859) The Origin of Species • Chap VII “Instinct”. Not defined, but three main examples • The instinct of the female cuckoo to lay small eggs in other bird’s nests, and the egg-ejection behavior of the newly hatched cuckoo chick; • Slave-making instincts in some species of ant • The cell-making instinct of the honeybee • The behaviors were seen by Darwin as not necessarily dependent on anatomical characteristics Darwin on Honey bees • Darwin thought that the honey comb was “absolutely perfect in economising labour and wax” on the grounds of geometry. • But it also “can be explained by natural selection having taken advantage of numerous, successive, slight modifications of simpler instincts”. These were based on spheres as in bumble bees, with S. American stingless bees intermediate. NB thousands of solitary bees which do not store honey Another of Darwin’s examples:cuckoos Another of Darwin’s examples:cuckoos Darwin (1859) page 185 function and form http://darwin-online.org.uk • “…. the acutest observer by examining the dead body of the water-ouzel would never have suspected its sub-aquatic habits; yet this anomalous member of the strictly terrestrial thrush family wholly subsists by diving,—grasping the stones with its feet and using its wings under water.” • Actually has some anatomical adaptations: 3rd eyelid, nostril flaps and oil gland 10 times larger than non-aquatic perching birds. • But Voelker (2002) agrees that the thrush is the closest relative and suggest that dippers diverged only 4m years bp Darwin (1859) page 185 http://darwin-online.org.uk • And the behavioural adaptation would have come first − • Thrushes foraging in streams instead of solid ground would then find enlarged oil glands useful Dipper diet includes aquatic insect larvae • Caddis fly larvae build themselves cases, in various ways depending on the species. • The Darwinian position would be that they inherit the behaviours required for this task. • But in fact Stuart and Currie (2002) found that there was little relation between the types of behaviours and the structural end-product across a variety of species. • Do species need a ‘genetic blueprint’ for the endproduct? Evolution connects the human species with the rest of the animal kingdom Era Cenozoic Period, begins (Million Years) Quaternary Holocene 0,01 Pleistocene 1.6 Tertiary Pliocene 5 Miocene 23 Oligocene 35 Eocene 56 Paleocene 65 145 210 250 Dinosaurs Mesozoic Cretaceous Jura Trias 290 360 410 440 505 545 Reptiles, birds, mammals (Amniotes) Paleozoic Permian Carboniferous Devonian Silurian Ordovician Cambrian Proterozoic Archean 2500 3800 Precambrian Man Primitive fish Oldest fossils of complex animals Creation of extant phyla First multicellular organisms First bacteria Lappin et al., 2006; standard texts EVOLUTIONARILY CONSERVED MOLECULAR GENETIC MECHANISMS FOR PATTERNING THE EMBRYONIC BRAIN . Reichert, H., & Simeone, A. (2001) Fly mutant restored with human gene Fly mutant restored with mouse gene Mouse mutant restored with fly gene Bishop et al., 2002 Ethological analyses of animal behaviour Vitalists believed in the instincts as mystical…. and behaviourists were preoccupied with learning…the way out was focussing on the survival value of behaviour patterns. “Behaviour patterns become explicable when interpreted as the result of natural selection, analogous with anatomical and physiological characteristics.” Tinbergen 1951, title and frontispiece Genome news network Zig-zag dance Experimental stimulus variation • There is no entry for gene in the 1951 index • But instinct and “innate” are the themes, and refers to genetics and mutations in the chapter on “The Evolution of Behaviour” • Tinbergen included some ill-advised evolutionary psychology • But there is a case that explanations for stickleback and human behaviour should be fundamentally different. A Study of Instinct 1951 Instinct - 2 • Tinbergen (1951) stressed two key concepts • 1. “sign stimuli” e.g. redness for sticklebacks. • 2. The “innate releasing mechanism”, by which particular sign stimuli release particular instinctive behaviour patterns • He believed these concepts applied to mammals • But the evidence is much clearer with lower vertebrates and invertebrates E.g. Spiders The orb web A first attempt e.g. spiders 2 Darwin’s comment on spiders “Thus everywhere in nature are battle, craft, and ingenuity, all following the merciless law of egoism, in order to maintain their own lives and to destroy those of others” Charles Darwin writing in Animal Intelligence by G.J. Romanes (1882), commenting on wolf and trapdoor spiders, p. 213 Innate behaviours e.g. fruitfly courtship Kimura, K. I., Ote, M., Tazawa, T., & Yamamoto, D. (2005). Fruitless specifies sexually dimorphic neural circuitry in the Drosophila brain. Nature, 438(7065), 229-233. “…..we identify a subset of fru-expressing interneurons in the brain that show marked sexual dimorphism in their number and projection pattern……. Fru expression can produce a male-specific neural circuit,” “Throughout the animal kingdom the innate nature of basic behaviour routines suggests that the underlying neuronal substrates necessary for their execution are genetically determined and developmentally programmed” Manoli & Baker (2004). e.g. fruitfly aggression Vrontou et al., (2006) fruitless regulates aggression and dominance in Drosophila Nature Neuroscience, 9,(01 Dec 2006), 1469 - 1471 When competing for resources, two flies of the same sex fight each other. Males and females fight with distinctly different styles, and males but not females establish dominance relationships. Here we show that sex-specific splicing of the fruitless gene plays a critical role in determining who and how a fly fights, and whether a dominance relationship forms. “our data indicate that aggressive behaviors are hardwired into the fly’s nervous system” also vertebrates e.g. zebrafish Gahtan, E., Tanger, P., & Baier, H. (2005). Visual prey capture in larval zebrafish is controlled by identified (four of them) reticulospinal neurons downstream of the tectum. Journal of Neuroscience, 25(40), 9294-9303. “Many vertebrates are efficient hunters and recognize their prey by innate neural mechanisms. During prey capture, the internal representation of the prey's location must be constantly updated and made available to premotor neurons that convey the information to spinal motor circuits.” “Seven-day-old zebrafish oriented toward, chased, and consumed paramecia with high accuracy.” Also birds e.g. Dilger (1961) Also for mammals Choi et al. (2005) Lhx6 delineates a pathway mediating innate reproductive behaviors from the amygdala to the hypothalamus, Neuron, 46(4), 647-660 (in embryonic and adult mice) “Virtually all metazoan organisms exhibit innate reproductive and defensive behaviors that are triggered by signals sensed from conspecifics or predators. …. The stereotypical nature of these behaviors suggests that their underlying neural circuits are likely to be genetically ‘hard-wired’.” “In mammals, innate reproductive and defensive behaviors are mediated by anatomically segregated connections between the amygdala and hypothalamus.” Slide 4 from the 2004 Nobel Lecture: Linda Buck Contrasts • there remain very contrasting positions within contemporary psychology particularly relating to how much emphasis is given to broadly biological as opposed to psychosocial evidence and theory. • The contrasts are less stark if we use “horses for courses”: explaining how the olfactory system works in mice is different from theorising about voting intentions in the Ukraine (2004) or the Tory leadership election (2005) or political events in Lebanon (2006) Evolutionary Psychology • Attempts to explain human psychology as a series of specialized adaptations • Driven in part by Chomskyan linguistics (see last week’s lectures): Pinker has written several more general books, most recently “The Blank Slate” (2002) as well as “The Language Instinct (1994) Adaptations • Williams (1966) defined an adaptation as “a characteristic that has arisen through and been shaped by natural and/or sexual selection. • It regularly develops in members of the same species because it helped to solve problems of survival and reproduction in the evolutionary ancestry of the organism. • Consequently it can be expected to have a genetic basis ensuring that the adaptation is passed through the generations”. Spandrels Gould and Lewontin, 1979; Gould, 1997 “An adaptationist programme has dominated evolutionary thought in England and the United States during the past forty years. It is based on faith in the power of natural selection as an optimizing agent.” We fault the adaptationist programme for its unwillingness to consider alternatives to spandrels and adaptive stories……..” exaptations are side effects of natural selection Evolutionary Psychology Bjorklund, D. F., & Smith, P. K. (2003). Evolutionary developmental psychology: Introduction to the special issue. Journal of Experimental Child Psychology, 85(3), 195-198 Evolutionary psychology, originally on the fringe of academic psychology.., has gained respectability within the last decade. Articles written from an evolutionary psychological perspective are found in the field’s most prestigious outlets; it has professional societies and journals of its own; college courses and textbooks are devoted to it; and there are academic positions specifically designated for evolutionary psychologists. Tooby and Cosmides • Cosmides, L. (1989). The logic of social exchange: has natural selection shaped how humans reason? Studies with the Wason selection task. Cognition, 31: 187 - 276. • Duchaine, B., Cosmides, L., & Tooby, J. (2001). Evolutionary psychology and the brain. Current Opinion in Neurobiology, 11(2), 225230. Swiss Army Knife Cosmides: The Swiss Army knife is a flexible tool. Its flexibility is not the result of having just one tool that is applied to all problems. Instead, it is a bundle of tools, each well-designed for solving a different problem –. Similarly, the human mind does not have just one blunt tool for solving all problems – and if it did, we would be very limited indeed.. Cosmides (1989) • Logic: If P then Q is only violated by P & ~Q • Typically participants do not use this logic in the Wason 4 card test (if vowel, theneven number on back) • A. if you have a bus pass, then you travel by bus • B. if you travel by bus, then you have a bus pass • Cosmides used many more elaborate scenarios, and found a strong bias towards “detecting cheaters” in that participants get A wrong, but B right. • But there is widespread disagreement, both with the specific claims made about reasoning and the Wason card-turning test, • And with the general claims about highly specialized mental tools for solving specific problems Examples • However, a few examples follow of papers firmly in the field of evolutionary psychology in recent issues of reputable journals. • Catatonia, Anorexia Nervosa and depression are proposed as adaptations, • there is a fairly general theory about individual decision rules interacting with group dynamics, • a paper proposing an evolutionary account of human facial expression of pain, • and a paper arguing that a human “innate releasing mechanism” for understanding agency is a key feature of religious concepts of the supernatural. Moskowitz, A. K. (2004). “Scared stiff”: Catatonia as an evolutionary-based fear response. Psychological Review, 111(4), 9841002. Guisinger, S. (2003). Adapted to flee famine: Adding an evolutionary perspective on anorexia nervosa. Psychological Review, 110(4), 745-761. Anorexia nervosa (AN) is ..attributed to psychological conflicts, attempts to be fashionably slender, neuroendocrine dysfunction, ……. Considerable research reveals these theories to be incomplete….. This article presents evidence that AN's distinctive symptoms of restricting food, denial of starvation, and hyperactivity are likely to be evolved adaptive mechanisms that facilitated ancestral nomadic foragers leaving depleted environments; genetically susceptible individuals who lose too much weight may trigger these archaic adaptations. This hypothesis accounts for the occurrence of AN- like syndromes in both humans and animals and is consistent with changes observed in the physiology, cognitions, and behavior of patients with AN. Guisinger table Depression • Allen, N. B., & Badcock, P. B. T. (2006). Darwinian models of depression: A review of evolutionary accounts of mood and mood disorders. Progress in Neuro-Psychopharmacology & Biological Psychiatry, 30(5), 815-826. • According to the social risk hypothesis, depression represents an adaptive response to the threat of exclusion from social relationships that, over the course of evolution, have been critical to maintaining an individual's fitness prospects • in the ancestral environment, depression induced: (i) sensitivity to social risk/threat; (ii) signaling behaviours that elicit social support; and (iii) a reduction in risky behaviours Kenrick, D. T., Li, N. P., & Butner, J. (2003). Dynamical evolutionary psychology: Individual decision rules and emergent social norms. Psychological Review, 110(1), 3-28. • Following evolutionary models, psychological mechanisms are conceived as conditional decision rules designed to address fundamental problems confronted by human ancestors, • A new theory integrating evolutionary and dynamical approaches is proposed. • Three series of simulations examining trade-offs in cooperation and mating decisions illustrate how individual decision mechanisms and group dynamics mutually constrain one another, and offer insights about gene-culture interactions. Kenrick and Butner, 2004 “At the most general level, evolutionary psychology can be defined as the study of cognitive, affective, and behavioral mechanisms as the solutions to recurrent adaptive problems.” “Along with the morphological features designed by natural selection, organisms also inherit central nervous systems……The behavioural inclinations of a bat would not work well in the body of a dolphin or giraffe and viceversa.” Kenrick and Buttner wrong • over a period of 35 years in Sweden (1965- 1999), there was no overall overrepresentation of stepchildren as victims. • Temrin, Nordlund, & Sterner, H. (2004) • In families with both stepchildren and children genetically related to the offender, genetic children tended to be more likely to be victims. Williams, A. C. D. (2002). Facial expression of pain: An evolutionary account. Behavioral and Brain Sciences, 25(4), 439-+. • This paper proposes that human expression of pain ..,, arises from evolved propensities. • The function of pain is to demand attention and prioritise escape, recovery, and healing; where others can help …, a distinct and specific facial expression of pain from infancy to old age, consistent across stimuli, and recognizable as pain by observers. • ……..there has been skepticism about the presence or extent of pain, judgments of malingering, and sometimes the withholding of caregiving and help. • … an evolutionary account can generate improved assessment of pain and reactions to it. Darwin’s “The expression of the emotions in man and animals” (1872) Peleg et al. (2006). Hereditary family signature of facial expression. PNAS 103(43), 15921-15926 • Correlated facial expressions in congenitally blind subjects and their seeing relatives, anticipates genes. Further examples • Rhodes, G. (2006). The evolutionary psychology of facial beauty. Annual Review of Psychology, 57, 199-226. • face preferences may be adaptations for mate choice because attractive traits signal important aspects of mate quality, such as health • Averageness, symmetry, and sexual dimorphism are good candidates for biologically based standards of beauty Atran, S., & Norenzayan, A. (2004). Religion's evolutionary landscape: Counterintuition, commitment, compassion, communion. Behavioral and Brain Sciences, 27(06), 713-730. • Religion is not an evolutionary adaptation per se, • but a recurring cultural by-product of the complex evolutionary landscape • A key feature of the supernatural agent concepts common to all religions is the triggering of an “Innate Releasing Mechanism,” or “agency detector,” • whose proper (naturally selected) domain encompasses objects relevant to hominid survival – such as predators, – but which actually extends to moving dots on computer screens, voices in wind, and faces on clouds. Domain General and Domain Specific • The “Swiss army knife” idea is more formally expressed in terms of domain specific or modular capacities. • The next paper argues that general intelligence is also domain-specific • This is dangerous for Evolutionary Psychology, since it opens the door to the idea that human evolution ended up by providing us with very open-ended and general purpose psychological capacities. Kanazawa, S. (2004). General intelligence as a domainspecific adaptation. Psychological Review, 111(2), 512-523 • General intelligence (g) poses a problem for evolutionary psychology's modular view of the human brain. The author …. argues that general intelligence evolved as a domain-specific adaptation for the originally limited sphere of evolutionary novelty in the ancestral environment… • It has accidentally become universally important merely because we now live in an evolutionarily novel world Kanisawa, Psych Review 2004 More Domain General ideas Atkinson, A. P., & Wheeler, M. (2004). The grain of domains: The evolutionary-psychological case against domain-general cognition. Mind & Language, 19(2), 147-176. ..evolutionary psychologists have argued that our innate psychological endowment consists of numerous domain- specific cognitive resources, rather than a few domain-general ones. … We conclude (a) that the fundamental logic of Darwinism,….. does not entail that the innate mind consists exclusively, or even massively, of domainspecific features, and (b) that a mixed innate cognitive economy of domain-specific and domaingeneral resources remains a genuine conceptual possibility. Human Ancestors • “psychological mechanisms are conceived as conditional decision rules designed to address fundamental problems confronted by human ancestors” (Kenrick and Butner, 2004) • This is a typical claim in evolutionary psychology, but there seems little detailed interest in what human ancestors might have done. Human Evolution • The epistemology of human evolution is necessarily difficult, since it relies on fossils, but fossil evidence has a reasonable track record in other areas, and human artefacts, in particular stone tools, provide another source of evidence Human evolution, summarised on p. 6 of handout Millions top Family Tree A new early fossil (2006) Alemseged, Z., et al. (2006). A juvenile early hominin skeleton from Dikika, Ethiopia. Nature, 443(7109), 296-301 Dikika is only 4km from where ‘Lucy’ was found (Australopithecus afarensis ) The Dikika specimen, from 3.3m yrs ago was about 3yrs old and probably female. The legs were human-like for bi-pedal walking, but the arms and hands ape-like. The hyoid bone (for the larynx) was also ape-like New Neanderthal data: Green et al., (2006) • Suggests common ancestor ~450,000 yrs ago Human uniqueness • The human brain is uniquely large • It is also functionally lateralized in a way which differs from chimpanzees • It may be metabolically enhanced Cacares et al (2003) • It may include different physiological components (Allman et al., 2005) • It may be organized uniquely, e.g. large frontal lobes (Deacon, 1997) Number of neurons in the nervous system 1,000,000,000,000 350,000,000,000 100,000,000,000 500,000,000 300,000,000 50,000,000 850,000 250,000 20,000 381 302 • • • • • • • • • • • Homo sapiens (maybe 1014) Chimpanzee Rhesus monkey Mouse Octopus Stickleback Honey bee Fruitfly Sea slug Thread worm male Thread worm In Striedter, G. F. (2006). Precis of Principles of brain evolution. Striedter brain szie Dorus, S., et al. (2004). Accelerated evolution of nervous system genes in the origin of Homo sapiens. Cell, 119(7), 1027-1040 Human, chimp, organg, Rhesus primary microcephaly Ponting and Jackson, 2005 • …….recent advances from the cloning of two human disease genes promise to make inroads in the area .. of brain size evolution. • Microcephalin (MCPH1) and Abnormal spindle-like microcephaly associated (ASPM) are genes mutated in primary microcephaly. • In this, the brain is of a size comparable with that of early hominids. • It has been proposed that these genes evolved adaptively with increasing primate brain size. ….both genes have undergone positive selection during great ape evolution. • the evolutionary patterns of all four presently known primary microcephaly genes are consistent with the hypothesis that genes regulating brain size during development might also play a role in brain evolution in primates and especially humans (Evans, 2006) More on brain size genes • Evans et al., (2005) claim the microcephalin has continued to evolve adaptively in modern humans • They say one genetic variant of microcephalin appeared as recently as 37,000 years ago. • The same team (Merkel-Bobrov et al, 2005) say that APSM, another gene regulating brain size, had a new variant only 5,800 years ago Huxley’s comparisons “So far as cerebral structure goes therefore, it is clear that man differs less from the Chimpanzee or the Orang, than these do even from the monkeys, and that the difference between the brains of the Chimpanzee and of Man is almost insignificant, when compared with that between the Chimpanzee brain and that of a Lemur. “(Darwin, 1874/1901, p. 312) semendeferi1 The Semendeferi et al., (2002) table Semendeferi table Brain re-organization: expansion of the frontal lobes The sherwood 2005 • Schoenemann et al.(2005) recently suggested that prefrontal white matter is disproportionately larger in humans than in other primates • but Sherwood et al. (2005) countered that a) the boundary between prefrontal and other cortex is not well defined; and b) that in any case, although the data showed humans having more white matter than the average primate, they did not show a difference between humans and great apes. • • • • Allman et al 2005 Von Economo neurons (VENs) are a recently evolved cell type which may be involved in the fast intuitive assessment of complex situations. As such, they could be part of the circuitry supporting human social networks. We propose that the VENs relay an output of fronto-insular and anterior cingulate cortex to the parts of frontal and temporal cortex associated with theory-of-mind We propose that in autism spectrum disorders the VENs fail to develop normally Allman et al., 2005 Human Brain Asymmetries Hutsler, 2003 7 autopsies: 50-97 yrs of age. No chimpanzees Sun, T., & Walsh, C. A. (2006). Molecular approaches to brain asymmetry and handedness. Nature Reviews Neuroscience, 7(8), 655-662 Sun, T., & Walsh, C. A. (2006). Molecular approaches to brain asymmetry and handedness. Elevated neuronal activity? • Caceres et al. (2003) applied a variety of genetic techniques to the cortical tissue (removed post-mortem) of humans, chimpanzees and rhesus macaques. • These suggested that humans and chimpanzees are more similar to each other than to the macaques, which is as expected, • but also that there were dozens of genes that were expressed very differently in human and chimpanzee cortex, with 90% of these being expressed more actively in humans than in chimpanzees, which suggested that • The human is brain is characterized by “elevated levels of neuronal activity”. • As a contrast, comparing gene expressing in the human and chimpanzee heart and liver revealed very little difference of this kind. Sherwood, C. C., et al. (2006). Evolution of increased glia-neuron ratios in the human frontal cortex. PNAS, 103(37), 13606-13611. • the human glia-neuron ratio in the prefrontal region did not differ significantly from predictions based on brain size. • Further analyses of glia-neuron ratios across frontal areas in a humans, chimpanzees, and macaque monkeys showed that regions involved in specialized human cognitive functions, such as "theory of mind" (area 32) and language (area 44) have not evolved differentially higher requirements for metabolic support. • …greater metabolic consumption of human neocortical neurons relates to the energetic costs of maintaining expansive dendritic arbors and long-range projecting axons in the context of an enlarged brain. • “Sherwood et al. (1) provide support for the idea that the human brain is more or less a large hominoid (ape) brain and can be understood in that context.” • Enard et al., (2002) did cross-species comparisons of the DNA for FoxP2, which when mutated gives rise to articulatory disorders in humans. Although it is “highly conserved” their data suggested that “this gene has been the target of selection during recent human evolution. • Watakabe et al., (2006) review gene expression profiling of postnatal rhesus neocortex. Although there is overall homogeneity of gene expression across different cortical areas, a few genes show marked area-specific patterns, e.g. genes specific to visual cortex and to association cortex. More genetic suggestions for human uniqueness • Prabhakar et al, (2006) found many humanspecific changes in regulatory sequences of DNA with almost no overlap with chimpanzee equivalents and suggest that these may have contributed to uniquely human features of brain development. • Pollard et al., (2006) found a particular regulatory gene expressed especially in certain neurons in human neocortex from 7 to 9 gestational weeks and say that this and similar “human accelerated regions provide new candidates in the search for uniquely human biology” Paleontological evidence from human evolution • Stone tools provide the main currently available clues to human evolution • But partly because they survive only periods of geological time • Other artifacts made from wood and bone may have been important, even if nothing now survives Psychologist cover Oldowan tools >2m years Acheulian tools 1,4 m – 0.5 m yrs, mainly Homo erectus Size of Handaxes Neanderthal tools, 400k yrs ago – 100k (Mousterian) Approx 25 k years ago, modern homo sapiens Delagnes & Roche (2005) • Even 2.34m years ago there was a highly controlled technology for producing stone flakes following constant technical rules and resulting in high productivity. • Their data consists of reconstructions of cobble reduction sequences --- putting the flakes back together, e.g. Delagnes & Roche (2005) 2.34m years ago The hand and tools The hand and tools • Some apes from around the time of the last common ancestor seemed to have hands like early hominids (Moya-Sola et al, 2005; Alba et al., 2003) • Since modern apes show some evidence of tool use it is likely that the hands became adaptively useful from the very earliest stages of bipedalism. Chimpanzee tools The hand and tools: Castiello 2005 Findings from patients with brain damage who have difficulty in grasping objects are difficult to reconcile with neurophysiological findings, ………. ……as the patients' lesions are confined to regions that, in monkeys, do not seem to be involved in graspingrelated visuomotor transformations --- Napier (1980) Napier infants Napier, ape hands Napier, power and precision grips Napier – Screwtop The hand and tools: conclusion • The consequence of evolution is that humans have domain-general potential for manual skill • The hands can be used for anything anatomically possible • There is no evidence for completely steretotyped movements, even for the precision grip (Wong and Whishaw, 2004) Wong, Y. J., & Whishaw, I. Q. (2004). Precision grasps of children and young and old adults: individual differences in digit contact strategy, purchase pattern, and digit posture. Behavioural Brain Research, 154(1), 113-123. The grasping patterns of male and female young adults, older adults and children were examined as they reached (with both left and right hand) for five small beads (3-16 mm diameter). Frame-by-frame analysis of grasping indicated a high degree of variability in digit contact strategies, purchase patterns and digit posture both within and between subjects. Other technologies: Neanderthal hut Mammoth remains Other technologies: Cave paintings, Southern France and Spain, 31,000 to 8,000 year before present Chauvet (31k) bison with active legs Detail of horses at Chauvet (31k) Hand at Chauvet (31k) Lamp at Lascaux (13k) Bull at Lascaux (13k) Functional differences: cultural learning • Tomasello & Rakoczy (2003) have argued that there are and invention • • • • two (initial) stages of uniquely human social cognition. The first stage is observable in one year olds, who have an understanding of other persons as intentional agents, This enables them to take part in pretend play, and is important as a prerequisite for shared attention and early social and linguistic learning. The second stage is the “Theory of Mind” belief-desire psychology which normally starts around 4 years of age, but which is dependent on several years of linguistic communication. These early stages of uniquely human social cognition enable the cultural “ratchet” of social and technological innovation (Tomasello et al., 2005) “And so if we imagine a human child born onto a desert island, somehow magically kept alive by itself until adulthood, it is possible that this adult’s cognitive skills would not differ very much – perhaps a little – but not very much, from T and rakoczy those of other great apes.” (p. 121) Understanding and sharing intentions Tomasello et al., 2005 • a species-unique motivation to share emotions, experience, and activities with other persons.. Leading to .. • “species-unique forms of cultural cognition and evolution” Understanding and sharing intentions 2 Tomasello et al., 2005 • “But then about 2 million years ago, when the manufacturing of tools began to be especially important .., individuals who could analyze intentional actions even more deeply, … were at a selective advantage.” • But prefer not to end with .. evolutionary scenarios, but with our ontogenetic model, of which more components are directly testable Human Uniqueness on behavioural grounds Almost all animal behaviour is genetically pre-programmed (by evolution) to fit an ecological niche (Darwin, 1859; Tinbergen, 1951; Manoli & Baker, 2004; Choi et al. 2005) Almost all human behaviour involves cultural learning (Tomasello and Razoksky, 2003; Tomasello et al., 2005) Reading • As last week for nature/nurture • Any of the papers quoted. • Or a debate initiated by Lickliter, R., & Honeycutt, H. (2003). Developmental dynamics: Toward a biologically plausible evolutionary psychology. Psychological Bulletin, 129(6), 819-835. • Or take a brief look at one of the books on human evolution listed in the handout. Books on Human Evolution (alternatives) on page 6 of handout Bradshaw, J. L. (1997). Human Evolution: A Neuropsychological Perspective. Hove: Psychology Press. BK lib 599.935BRA. Johanson, Donald C., and Edgar, Blake (2001) From Lucy to Language. London: Cassell paperbacks. 2 copies in Main Birkbeck Library, classmark=599.938 JOH Jones, S., Martin, R. D., & Pilbeam, D. R. (1992). The Cambridge encyclopedia of human evolution. Cambridge [England] ; New York, NY, USA: Cambridge University Press, BK lib 599.9 CAM, 3 copies Richards, G. (1987) Human Evolution. Routledge: London. (Bk Lib GYW, N [Ric]) Conclusions • Evolutionary theory is essential for many areas of animal behaviour, and rapid advances in molecular genetics may impinge on knowledge of the physiological underpinnings of human capacities • But a crucial outcome of human evolution was a fairly open aptitude for cultural and technological invention • The human brain may not be equivalent to a blank slate, but it has large areas of free space for cultural and historical changes — the blank parts may be the most important.