Neuropsychology Option Week 5

1. Neuropsychology Option Week 5 2nd Lecture

2. T.H. Huxley (1863, 1874) “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, 1871/1901, p. 312) ‘Every principal gyrus and sulcus of a chimpanzee’s brain is clearly represented in that of a man’.

3. Human brain

4. Chimpanzee brain The chimpanzee brain is less than a third the size of a typical human brain, but superficially looks similar

5. 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)

6. Huxley (1871) • ‘Every principal gyrus and sulcus of a chimpanzee’s brain is clearly represented in that of a man’ • This second claim is probably misleading, since the reverse is certainly not true, and it maybe that the 21st century conclusion is radically different from the 19th century one, when more is known about the genetics of the difference between human and chimpanzee brains (Culotta, 2005).

7. Mikkelsen et al, (2005)

8. 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.

9. 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. • Taken together, these findings suggest that 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.”

10. A minor confirmation by genetic analyses is that the human sense of smell is reduced by comparison to the chimpanzee both humans and chimpanzees have less sense of smell that dogs or mice This can be assessed by counting olfactory receptor genes and the proportion of these which are inactive (pseudogenes). Humans have a significantly higher proportion of these than chimpanzees (Gilad et al., 2005). Sense of Smell

11. Olfactory receptor genes: Linda Buck Nobel 2004

12. Although further detailed distinctive features of the human brain may be expected, it is also the case that there are some features of the human brain, in particular the organization of the visual system, where the details in the human brain differ very little from those in chimpanzees (e.g. Cola et al., 2005 – bottom of p 2 of handout) • “Thus, in the aspects of organization we examined, the inferior pulvinar of chimpanzees closely resembles that of humans and monkeys”

13. Functional differences: language • Apart from looking at neurophysiological or genetic details, it is possible to give hypothetical answers to both the how? and why? questions by making assumptions about the new psychological capacities subserved by human brain evolution, • “The origin of humans was accompanied by the emergence of new behavioural and cognitive functions, including language and specialized forms of abstract representation.” (Caceres et al., 2003).

14. The two most frequently appealed to candidates for new psychological capacities are language and an enhanced capacity for social cognition. • Language remains a strong candidate for a human specialization because chimpanzee abilities appear to be so limited (see weeks 10 and 11 Psychobiology II). • There are however several very different suggestions as to how this human specialization arose.

15. Chomsky pic in guardian

18. Hauser et al. (2002) for instance, although considering other possibilities, favour the notion that recursion is the key uniquely human component of language, • but paradoxically from a Darwinian point of view, they argue that recursion probably evolved “for reasons other than language”. • Recursion is illustrated by Chomsky’s well-known sentence “colourless green ideas furiously sleep”, which we understand as a grammatically correct sentence even though it makes little sense. • The sequence “I will say a very, very long sentence”; I will say a very, very, very long sentence” and so on is a simpler illustration that a principle for combining just a few words can generate an infinite number of possible utterances.

19. Gentner, T. Q., et al. (2006). Recursive syntactic pattern learning by songbirds. Nature, 440(7088), 1204-1207 Humans regularly produce new utterances that are understood by other members of the same language community…. The recursive, hierarchical embedding of language units (for example, words or phrases within shorter sentences) that is part of the ability to construct new utterances minimally requires a 'context-free' grammar. Recent hypotheses make the central claim that the capacity for syntactic recursion forms the computational core of a uniquely human language faculty Here we show that European starlings (Sturnus vulgaris) accurately recognize acoustic patterns defined by a recursive, self-embedding, context-free grammar…Thus, the capacity to classify sequences from recursive, centre-embedded grammars is not uniquely human.

20. They used 8 recorded starling ‘rattles’ and 8 ‘warbles’ to make up a total of 4096 stimuli

21. ii) Darwinian evolution of all parts of the human language system • The Hauser et al. (2002) suggestion tends to minimize the role of human evolution because a large part of the language system is held to be shared with other species, while the uniquely human part, recursion, they would prefer not to be an adaptation (i.e. not evolved by Darwinian selection). • Pinker and Jakendoff (2005) supply a lengthy argument against the Hauser et al. (2002) position, since they had both previously put forward the position that human language is “a system of co-adapted traits that evolved by natural selection” for the purpose of communicating ideas

22. ii) Darwinian evolution of all parts of the human language system • Pinker and Jakendoff believe that the capacity of the human brain to handle recursion evolved by natural selection, but that many other aspects of brain capacity necessary for language, particularly for conceptual structure, speech perception and speech production, needed to be shaped by natural selection as well. • Pinker and Jakendoff (2005) are also able to appeal to genetic evidence that was not available 10 years ago.

23. Genes and Language: BBC October 01

24. Pinker comments October 2001

25. Wellcome foxp2

26. National Geographic

27. No sign of Foxp2

28. Liegeois 03

29. Nature 2002 • Language is a uniquely human trait likely to have been a prerequisite for the development of human culture. The ability to develop articulate speech relies on capabilities, such as fine control of the larynx and mouth, that are absent in chimpanzees and other great apes. FOXP2 is the first gene relevant to the human ability to develop language. A point mutation in FOXP2 co-segregates with a disorder in a family in which half of the members have severe articulation difficulties accompanied by linguistic and grammatical impairment. This gene is disrupted by translocation in an unrelated individual who has a similar disorder. Thus, two functional copies of FOXP2 seem to be required for acquisition of normal spoken language. We sequenced the complementary DNAs that encode the FOXP2 protein in the chimpanzee, gorilla, orang-utan, rhesus macaque and mouse, and compared them with the human cDNA. We also investigated intraspecific variation of the human FOXP2 gene. Here we show that human FOXP2 contains changes in amino-acid coding and a pattern of nucleotide polymorphism, which strongly suggest that this gene has been the target of selection during recent human evolution.

30. FOXP2 — a) • Pinker and Jakendoff (p. 218) are able to make the point that “The possibility that the affected people are impaired only in recursion is a non-starter.” • Instead the expression pattern of FOXP2 in both mice and humans suggests that it is involved in the development of circuits for motor control necessary in vocalization (Lai et al., 2003; Shu et al., 2005). • The fact that there are homologies between humans and mice in this respect could be taken to support Hauser et al.’s point about some aspects of human language having a long evolutionary history,

31. FOXP2 — b) • the more conventional Darwinian position would be to say that motor control for vocalization has a long evolutionary history, but that the uniquely human capacity for speech will have necessitated special brain mechanisms for the co-ordination of articulatory organs which are not shared with other species.

32. Lai et al. (2003). FOXP2 expression during brain development coincides with adult sites of pathology in a severe speech and language disorder. Brain, 126, 2455-2462. • the homologous pattern of FOXP2/Foxp2 expression in human and mouse argues for a role for this gene in development of motor-related circuits throughout mammalian species. Overall, this study provides support for the hypothesis that impairments in sequencing of movement and procedural learning might be central to the FOXP2-related speech and language disorder

33. Shu et al. (2005). Altered ultrasonic vocalization in mice with a disruption in the Foxp2 gene. Proceedings of the National Academy of Sciences of the United States of America, 102(27), 9643-9648 shu • our findings support a role for Foxp2 in cerebellar development and in a developmental process that subsumes social communication functions in diverse organisms.

34. Teramitsu, I., & White, S. A. (2006). FoxP2 regulation during undirected singing in adult songbirds. Journal of Neuroscience, 26(28), 7390-7394. (not on handout) “Our data suggest that FoxP2 is important not only for the formation but also for the function of vocal control circuitry”. (In zebra finches)

35. iii) Emergence of language from a number of “language ingredients” • Elman (1999, 2005) provides yet another account of language, which is Darwinian in that it emphasises that • “species-specific biological factors play a critical role in the ability of humans to acquire and process language (1999, p. 1) • but which differs from the account given by Pinker in predicting a lack of any genetic control of specific cortical micro-circuitry for language. Instead • language “is simply the result of a number of tweaks and twiddles” which produce changes in humans in such things as vocal tract control, sociality, imitation and shared attention.

36. Emergence of language from a number of “language ingredients”-b) • These traits then interact to produce the unique human capacity for language. • this account comes from the connectionist tradition and the “emergentist” aspect leads to the expectation that there will be complex developmental trajectories • but “tweaks and twiddles” are entirely consistent with Darwinian processes, and changes in vocal tract control presumably would need to be brought about by something like changes to FOXP2.

37. Elman, 1999, 2005

38. Functional differences: social cognition and theory of mind • An alternative function role for the large brains of primates has been suggested to be social cognition (Jolly 1966; Humphrey, 1976; Barrett & Henzi, 2005). • This has the advantage of applying to the large brains of non-human primates, as well as, putatively, to humans, but the disadvantage that it does not by itself explain human tool using, • although the social transmission of tool using skills may have been a crucial component of the success of this strategy.

39. Functional differences: social cognition and theory of mind • Current work on social cognition includes evidence that chimpanzees and other great apes appear to have skills which might be regarded as precursors to a theory of mind, • for instance the ability to understand both human (Call et al., 2004) and conspecific (Tomasello et al., 2003) psychological states, • but also points to the severe limitations of chimpanzees’ social cognition by comparison with the human case. • In particular, chimpanzees can show understanding of what a conspecific has or has not seen when competing for contested food, but the often “show surprisingly weak social-cognitive skills” in tasks which require social co-operation (Hare & Tomasello, 2004).

40. Blindfolds picture

41. Box 1 of tomasello, call and hare Tomasello, Call and Hare (2003)

42. Box 1 text Tomasello, Call and Hare (2003)

43. Box 2 diagram Tomasello, Call and Hare (2003)

44. Bod 2 text Tomasello, Call and Hare (2003)

45. Warneken, F., Chen, F., & Tomasello, M. (2006). Cooperative activities in young children and chimpanzees. Child Development, 77(3), 640-663. Human children 18 or 24 months of age and 3 young chimpanzees interacted in 4 cooperative activities with a human adult partner. The human children successfully participated in cooperative problem-solving activities and social games, whereas the chimpanzees were uninterested in the social games. As an experimental manipulation, in each task the adult partner stopped participating at a specific point during the activity. All children produced at least one communicative attempt to reengage him, perhaps suggesting that they were trying to reinstate a shared goal. No chimpanzee ever made any communicative attempt to reengage the partner. These results are interpreted as evidence for a uniquely human form of cooperative activity involving shared intentionality that emerges in the second year of life.

46. Warneken et al., 2006. Top row, problem solving tasks, bottom row ‘social tasks’ Above, alternative trapdoor task for chimps

47. Functional differences: cultural learning and invention • Tomasello & Rakoczy (2003) have argued that there are 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 form the basis enable the “cultural ratchet” of social and technological innovation (Tomasello et al., 1993; Tomasello, 1999)

48. T and rakoczy “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 those of other great apes.” (121)

49. Tomasello, M., Carpenter, M., Call, J., Behne, T., & Moll, H. (2005). In search of the uniquely human - Response. Behavioral and Brain Sciences, 28(5), 721-735 Human beings are the world’s experts at mind reading. As compared with other species, humans are much more skillful at discerning what others are perceiving, intending, desiring, knowing, and believing. Our hypothesis for this “something additional” is shared intentionality. We propose that human beings, and only human beings, are biologically adapted for participating in collaborative activities involving shared goals and socially coordinated action plans

50. Tomasello, M., Carpenter, M., Call, J., Behne, T., & Moll, H. (2005). In search of the uniquely human - Response. Behavioral and Brain Sciences, 28(5), 721-735 Our attempt was to propose a theory of the social-cognitive and social-motivational bases of humans’ ability and propensity to live in this local, that is, this cultural, way – which no other species does – focusing on such things as the ability to collaborate and to create shared material and symbolic artifacts Some…. provided alternative “magic bullets” for explaining the key features of human cognitive and social uniqueness….But, in nearly all of these cases, we find that these alternative accounts basically sneak in through the back door one or another form of shared intentionality as a kind of hidden premise.