4.36 Nuclear Schizophrenic Symptoms As The Key To The Evolution of The Human Brain

4.
36 Nuclear Schizophrenic Symptoms as the Key to the Evolution of the Human Brain
T J Crow, University of Oxford, Oxford, UK
2007 Elsevier Inc. All rights reserved.
4.36.1 4.36.2 4.36.3 4.36.4 4.36.5 4.36.6
The Nuclear Symptoms of Schizophrenia and the Central Paradox The Problem of Language for Evolutionary Theory Darwins Intuition on Sexual Selection Paul Broca and Cerebral Asymmetry The Torque and Related Asymmetries in Psychosis The Structure of Language and Its Decomposition in Psychosis 4.36.6.1 The Linguistic Sign Is Bihemispheric 4.36.6.2 Deixis and the Significance of the Indexical 4.36.6.3 The Human Brain as a Four-Chambered Organ 4.36.6.4 The Deictic Origin and the Performative Hypothesis 4.36.7 XY Homology and the Xq21.3/Yp Translocation 4.36.8 Implications for Evolutionary Theory 4.36.8.1 The Case for Saltation 4.36.8.2 Sexual Selection and the Mate Recognition Principle 4.36.8.3 Species-Specific Variation Is Epigenetic 4.36.8.4 Speciation Events Occur on the Heterogametic Chromosome 4.36.9 Conclusions
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Glossary
Brocas area A region of the frontal lobe of the left cerebral hemisphere associated with articulate language. The concept that the two cerebral hemispheres differ in function. This is reflected in differences in the anatomical organization of the two hemispheres. A psychopathological disorder characterized by delusions, hallucinations, disorganized thought, and a poverty of emotion, speech, and intention. The process where some individuals have greater reproductive success than others because of competitive advantages in mating.
cerebral asymmetry
schizophrenia
sexual selection
2. Voices commenting: a voice or voices heard by the subject speaking about him and therefore referring to him in the third person. 3. Passivity (delusions of control): the subject experiences his will as replaced by that of some other force or agency. 4. Thought insertion: the subject experiences thoughts which are not his own intruding into his mind. In the most typical case, the alien thoughts are said to have been inserted into the mind from outside, by means of radar or telepathy or some other means. 5. Thought withdrawal: the subject says that his thoughts have been removed from his head so that he has no thoughts. 6. Thought broadcast: the subject experiences his thoughts actually being shared with others. 7. Primary delusions: based upon sensory experiences (delusional perceptions) the patient suddenly becomes convinced that a particular set of events has a special meaning.
4.36.1 The Nuclear Symptoms of Schizophrenia and the Central Paradox

The core nuclear symptoms of schizophrenia according to Kurt Schneider, as defined by the glossary of the Present State Examination (Wing et al., 1974), are:
1. Thought echo or commentary: the subject experiences his own thought as repeated or echoed with very little interval between the original and the echo.
Why do the core symptoms have this form? What primary function is disturbed? What neural structure is the focus of the disturbance? Through the presence of these features the authors of the WHO Ten Country Study of the Incidence and Manifestations of Schizophrenia (Jablensky et al., 1992) reached their conclusion that:
schizophrenic illnesses are ubiquitous, appear with similar incidence in different cultures, and have clinical features that are more remarkable by their similarity across cultures than by their difference.
Thus, schizophrenia is constant across populations that differ widely in geographic, climatic, industrial, and social environment; it seems it is a characteristic of human populations. It is a disease (perhaps the disease) of humanity. If the core syndrome is a characteristic of populations it must somehow be intrinsic, i.e., genetic in origin. This raises the central paradox: why if the disease is associated with a biological disadvantage is this genetic variation not selected out? About the existence of the fecundity deficit there is little doubt: it is of the order of 70% in males and 30% in females (Essen-Moller, 1959; MacSorley, 1964; Vogel, 1979; Haverkamp et al., 1982; Penrose 1991). To balance such a disadvantage, a substantial and universal advantage must be invoked. But what could such an advantage be, and by whom might it be carried? These questions were first clearly formulated in a paper (Huxley et al., 1964) notable by an authorship that includes J. S. Huxley and E. Mayr, two progenitors of the modern evolutionary synthesis of Mendelian genetics and Darwinian theory. The first paragraph identifies these two as the originators of the notion. Yet the theory they proposed that the balance lay in resistance to wound shock and stress was clearly mistaken, as Kuttner et al. (1967) pointed out soon after. It makes no sense to suppose that the advantage of a particular genetic variation lies in a field that is unrelated to the disadvantage. Kuttner et al. considered three advantages intelligence, language, and complex social ability and favored the last. But these three are clearly related and one language is both of more obvious adaptive significance and is more readily defined in terms of neural function than the other two. What is striking about the nuclear symptoms is that they can hardly be conceived except within the framework of language. Auditory hallucinations (items 1 and 2 above) are self-evidently an anomaly of the perception of the spoken word. Thought insertion, withdrawal, and broadcast (items 4, 5, and 6) are disturbances of the experience of thought and of the transition from thought to speech production. Primary delusions (item 7) represent a deviation in the attachment of meaning to symbolic representations, that is to say, they are a disturbance of semantics. Only delusions of control (item 3) are not immediately recognizable as a disturbance of speech, but these cannot be described except with the use of language. They can perhaps be understood as anomalies of identification of the self in relation to the rest of the universe of symbols.
4.36.2 The Problem of Language for Evolutionary Theory

The concept of language as the defining characteristic of humanity has an ancient origin:
In most of our abilities we differ not at all from the animals; we are in fact behind many in swiftness and strength and other resources. But because there is born in us the power to persuade each other and to show ourselves whatever we wish, we not only have escaped from living as brutes, but also by coming together have founded cities and set up laws and invented arts, and speech has helped us attain practically all of the things we have devised (Isocrates, 436338 BC, quoted in Harris and Talbot, 1997, p. xiii).
Darwin can be quoted in agreement with this view. On p. 53 of The Descent of Man, he writes that language has justly been considered as one of the chief distinctions between man and the lower animals and he seems not to have regarded this as a particular difficulty. But in 1873, within 2 years of the publication of The Descent of Man, Mueller (1873), who held the chair of Philology in the University of Oxford, delivered a series of three lectures at the Royal Institution in which he drew attention to the problems that language raises for Darwins theory:
My object is simply to point out a strange omission, and to call attention to one kind of evidence I mean the evidence of language which has been most unaccountably neglected, both in studying the development of the human intellect, and determining the position which man holds in the system of the world.
In the second lecture Mueller addresses the problem:

There is one difficulty which Mr Darwin has not sufficiently appreciated . . . There is between the whole animal kingdom on the one side, and man, even in his lowest state, on the other, a barrier which no animal has ever crossed, and that barrier is Language . . . If anything has a right to the name of specific difference, it is language, as we find it in man, and in man only . . . If we removed the name of specific difference from our philosophic dictionaries, I should still hold that nothing deserves the name of man except what is able to speak . . . a speaking elephant or an elephantine speaker could never be called an elephant. Professor Schleicher, though an enthusiastic admirer of Darwin, observed once jokingly, but not without a deep meaning, If a pig were ever to say to me, I am a pig it would ipso facto cease to be a pig.
Mueller thus raised for Darwin the problem for evolutionary theory raised 86 years later by Chomsky (1959) in his critique of Skinners claim to have an explanation of language in operant principles derived form studies in the rat. The strong implication was that there were principles underlying language that were human-specific. The concept of universal
Nuclear Schizophrenic Symptoms as the Key to the Evolution of the Human Brain
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grammar as a defining human characteristic and of its generativity have implications for speciation theory. The question raised by Chomsky, as by Mueller, is: what is the nature of a species? In the last chapter entitled Recapitulation and Conclusion of The Origin of Species by Means of Natural Selection, Darwin had written:
In the distant future I see open fields for far more important researches. Psychology will be based on a new foundation, that of the necessary acquirement of each mental power and capacity by gradation. Light will be thrown on the origin of man and his history.
The form of words that of the necessary acquirement of each mental power and capacity by gradation is of note because it clearly expresses Darwins predilection for a gradualist account of the origin of human faculties along with other evolutionary innovations. The paradoxical point has been made that an issue on which The Origin of Species is weak is the origin of species. One figure appears in the book (chapter 4) to show how within one genus different varieties and species emerge over eons of time. The implication of the figure is that variation within species is qualitatively the same as variation between species, hence the absence of a clear differentiation of varieties and species. In the same chapter Darwin wrote:
Hereafter we shall be compelled to acknowledge that the only distinction between species and well-marked varieties is, that the latter are known, or believed, to be connected by intermediate gradations, whereas species were formerly thus connected.
was no greater than that between any pair of the great apes compared on their own. Huxley (1863) thus fit humans into the framework of the Darwinian theory of natural selection of which he was so powerful an advocate. But on the issue of speciation, he did not see eye to eye with his mentor. After the publication of The Origin, he wrote to Darwin that he was ready to go to the stake for the theory but added that he thought that Darwin had loaded himself with an unnecessary difficulty in adopting Natura non facit saltum so unreservedly. Huxley was thus the first in a line of evolutionists including Bateson (1894), De Vries (1901), Goldschmidt (1940), and most recently Gould (2002), who have held that species transitions were more discontinuous, and the characteristics of species more stable, than appeared to follow from Darwins formulation (see The Evolution of Language Systems in the Human Brain, The Interpreter in Human Psychology).
4.36.3 Darwins Intuition on Sexual Selection

When Darwin (1871) addressed himself to the problem of human origins in The Descent of Man this publication linked in a single volume The Descent of Man and Selection in Relation to Sex the anatomical and paleontological case with Darwins theory of sexual selection. In his introduction, Darwin writes that:
During many years it has seemed to me highly probable that sexual selection has played an important part in differentiating the races of man; but in my Origin of Species I contented myself by merely alluding to this belief. When I came to apply this belief to man, I found it indispensable to treat the whole subject in full detail (Darwin, 1871, The Descent of Man, pp. 45).
His aim was to establish that species have a common origin, they are not the subjects of independent creations. Species can arise, subject to favorable environmental circumstance, out of the variation that is present in the natural world at any point in time. It is a gradualist theory. That gradualism was preserved into the evolutionary synthesis of the Darwinian theory of natural selection with Mendels laws of genetic inheritance in the 1940s (Mayr and Provine, 1998). The test comes in the application of the theory to actual speciation events. Such events are not generally observed, but somewhere in the figure, if it were to be applied to the case of humans, we must suppose there is a separation that leads on the one hand to the chimpanzee and bonobo and on the other to Australopithecus, Homo erectus, and modern Homo sapiens. The branch point and its sequelae is of theoretical and specific interest. In 1863, T. H. Huxley addressed the question in Mans Place in Nature. He mounted a powerful case that on a series of anatomical comparisons the distance between humans and any one of the great apes
Some passages in the second part of the book indicate that Darwin considered the two arguments to be related in a more fundamental way. Thus:
. . .Sexual selection has apparently acted on both the male and the female side, causing the two sexes of man to differ in body and mind . . . [and] has indirectly influenced the progressive development of various bodily structures and of certain mental qualities. Courage, pugnacity, perseverance, strength and size of body . . . have all been indirectly gained by one sex or the other, through the influence of love or jealousy, through the appreciation of the beautiful in sound, color and form, and through the exertion of choice. . . (Darwin, 1871, Selection in Relation to Sex, p. 402).
Although Darwin believed that sexual selection and the descent of man are related, he nowhere specifies exactly how this is the case and the fact remains that these are separate books. Thus, on the one hand we have Charles Darwin committed to the
view that humans are descended from the great apes by the process of natural selection with a strong intuition that the ancillary process of sexual selection also has something to do with it but unable to integrate these processes. On the other hand, we have Friedrich Max Mueller complaining that language has characteristics that are present in the communicative abilities of no animal other than humans and that Darwin has given no account of its origins. The solution to this dispute that I have offered (Crow, 1998a, 1998b, 2000a, 2002a, 2002b, 2002c) is that the Darwinian gradualist account indeed has to give ground to a saltational version, as argued strongly for example by Goldschmidt and Gould. Speciation events have a reality that is obscured in The Origin of Species. But that there is a relationship between speciation and sexual selection, as Kaneshiro (1980) and Carson (1997), among others, have argued; species are distinguished by characteristics that are often sexually dimorphic. The possibility is that the first change in the process of speciation occurs in one sex, generally the male, and that this change is then subject to mate choice to define what Paterson has called a specific mate recognition system. My proposal is that changes on the sex chromosomes, including chromosomal rearrangements and subsequent epigenetic modifications of gene control, play a critical role in these transitions. This concept of the nature of speciation comes from a consideration of the problem that vexed Max Mueller the origins of language and its relationship to the origin of humans and its relevance to the central paradox of psychosis the universal persistence in human populations of a genetic predisposition in the face of a biological disadvantage (Crow, 2000b).
presumably unrelated to those expressed in the cortex that are associated with language. Such species specificity would have been no surprise to Broca (see Harrington, 1987, pp. 4951, for his views on the essence of human nature), but the assumption that directional asymmetry on a population level is present in other primates, perhaps based upon Darwinian gradualist principles, has been widespread in the literature. Annett introduced a discussion of the issue in her earlier volume (Annett, 1985, pp. 169173) with reference to the work of Finch (1941), who found no evidence of directional handedness in a group of 30 chimpanzees. Subsequent studies, for example, Annett and Annett (1991), of 31 lowland gorillas in European zoos and Byrne and Byrne (1991), of 38 mountain gorillas in Rwanda, have reinforced this conclusion. Recently, Marchant and McGrew (1996) systematically studied 42 chimpanzees in the Gombe National Park and reviewed the primate literature to conclude that non-human primate hand function has not been shown to be lateralized at the species level it is not the norm for any species, task, or setting, and so offers no easy model for the evolution of human handedness (McGrew and Marchant, 1997; see also Holder, 1999 for congruent conclusions from a field survey of primates in Africa). Thus, we have evidence that a putative correlate of the capacity for language that Mueller and Chomsky identify as the defining characteristic of H. sapiens demonstrates a discontinuity in the primate phylogenetic tree (Figure 1). Thus, at some point in the course of hominin evolution, the dimension of asymmetry was introduced in the sequence of brain development, and this dimension, or some modification of it, is the obvious correlate of language.
4.36.4 Paul Broca and Cerebral Asymmetry

The key to the solution lies in the asymmetry or torque that appears to be characteristic of the human brain. After he had convinced himself (Broca, 1861) of the reality of the earlier observations of Marc Dax that language in the frontal lobes is localized on the left side, Broca (1877) came to the conclusion that:
Man is, of all the animals, the one whose brain in the normal state is the most asymmetrical. He is also the one who possesses the most acquired faculties. Among these faculties . . . the faculty of articulate language holds pride of place. It is this that distinguishes us most clearly from the animals.
Chimpanzee Human 100 L 50 0
Marchant and McGrew Provins et al. 50 100 R
Number of Activities Individuals Chimpanzee Human 38 1960 46 75
While it is clear that there are directional asymmetries (e.g., of the habenular nucleus) that are ancient in vertebrate phylogeny, these are
Figure 1 Hand preference for everyday activities in chimpanzees and H. sapiens compared. Data for chimpanzees referred to a community of 38 animals (Pan troglodytes schweinfurthii) observed by Marchant and McGrew (1996). Data for H. sapiens were collected by questionnaire by populations of undergraduates by Provins et al. (1982). Medians and boundary values for 95% have been extracted from graphs in the original publications.
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But why should the capacity for language be variable between individuals? Perhaps only when we understand the nature of the genetic mechanism will we have a clear answer (Crow, 2002c). The important point is that Brocas hypothesis that language is lateralized in the brain provides an indication of the neurophysiological basis and an approach to its pathophysiology. Buxhoeveden et al. (2001) have recently documented the anatomical correlate of population bias to right-handedness. Through a statistical analysis of the minicolumn structure of the cerebral cortex, they have demonstrated asymmetries, for example, of minicolumn width and separation that are present in the planum temporale of human but absent in those of the chimpanzee and rhesus monkey.
4.36.5 The Torque and Related Asymmetries in Psychosis

The form of the asymmetry from right frontal to left occipital, described as cerebral torque has implications for function, as the anatomy has for pathophysiology. Aspects of anatomical asymmetry are deviant in individuals who suffer from psychosis (see Crow, 1990b, 1997; Petty, 1999; Esiri and Crow, 2002) and there is evidence from a study of handedness in childhood that they are lateralizing less, or more slowly (Crow et al., 1996; Leask and Crow, 2005) than the population as a whole. In postmortem studies, the anatomical changes appear to be more posterior in the brain; losses or reversals of asymmetry have been detected in fusiform, parahippocampal (McDonald et al., 2000), and superior temporal (Highley et al., 1999b) gyri. A curious feature of these findings is that although the loss or reversal was present in both sexes, the relationship to age of onset was different: greater anomaly related to earlier age of onset in females but to later age of onset in males. Other sex differences have been detected in postmortem brain. Density of fibers in the corpus callosum was greater in females than in males, consistent with the generalization that connectivity is inversely related to degree of asymmetry (Highley et al., 1999a). In patients with psychosis, fiber density was reduced relative to female controls, while in males it was increased relative to male controls. Consistent with these findings in an magnetic resonance imaging (MRI) study (Highley et al., 2003), white matter in the occipitotemporo-occipital regions was greater in females than in males, while in female patients it was reduced and in male patients increased relative to same sex controls.
Thus, there are morphological changes in the brain in psychosis that are sex-dependent and may relate to the sex difference in age of onset: onsets are earlier in males, and earlier onsets are generally associated with a poorer outcome and a preponderance of negative symptoms (affective flattening and poverty of speech). Conversely, with increasing age of onset the proportion of females increases and the form of the illness is more likely to be paranoid, i.e., delusional. This sex difference has been identified by two evolutionary theorists (Gould and Gould, 1989) as evidence that sexual selection operates in humans. This attractive argument encounters the difficulty that on a simple developmental interpretation, the sex difference is in the wrong direction: brain size (Kretschmann et al., 1979) and verbal ability (Crow et al., 1998) develop faster in females than in males, yet onset of psychosis is earlier in males (Figure 2). A solution to this problem may unravel the evolution of the human brain. Some recent data are relevant. In a postmortem study (Chance et al., 2006), planum temporale asymmetry of surface area to the left was found to be greater in males than females, consistent with the sex difference in the torque. But there was also an asymmetry of Heschls gyrus (primary auditory cortex) and this, as has been found with the asymmetry of the paracingulate sulcus in the frontal lobes (Clark et al., 2006) was greater in females. A possible generalization therefore is that the brain growth is faster and the plateau is earlier in females and that asymmetries, for example of primary sensory cortex, that are formed earlier are greater in females, whereas those, for example of posterior heteromodal association cortex that have the opportunity of developing longer, are greater in males. The principle that interhemispheric connections are inversely related to degree of asymmetry together with the hypothesis that there is a relative loss of fibers in one direction (Witelson and Nowakowski, 1991) provides a possible solution to the age of onset paradox. Myelination continues into the third and fourth decades of life in the corpus callosum and apparently continues longer in females than males (Cowell et al., 1992; Pujol et al., 1993). This sex difference may reflect the longer time required in females to myelinate the larger body of interhemispheric axons. If we assume that the first symptoms of psychosis emerge as a consequence of some limit imposed on interhemispheric transmission by these late myelinations (for example, that those predisposed to psychosis fail to differentiate connections in the two directions), then owing to the faster myelinization, this may be apparent earlier in males. In
Superior temporal
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L R L R Control Schizo n = 14 n = 13 Female
L R L R Control Schizo n = 15 n = 17 Male
Figure 2 Volumes of cortical gyri (superior temporal, fusiform, parahippocampal) in patients with schizophrenia and controls assessed by stereology on postmortem brain. There are subtle asymmetries to the left in controls, and to the right in patients in each case. Reproduced from Esiri, M. M. and Crow, T. J. 2002. The neuropathology of psychiatric disorder. In: Greenfields Neuropathology (eds. D. I. Graham and P. L. Lantos), 7th edn., vol. II, pp. 431470. Arnold.
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In frontal regions, no gross asymmetry and no change in gyral volume was detected in schizophrenia (Highley et al., 2001), but in the density of cells in the cortex (in area 9) there was an asymmetry (greater cell density) to the left in controls and loss or reversal of this asymmetry in patients (Cullen et al., 2006).
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4.36.6 The Structure of Language and Its Decomposition in Psychosis

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Figure 3 Pyramidal cell density in layer 3 of the left and right hemispheres of schizophrenic and control brains. An asymmetry to the left is present in controls and to the right in patients. Reproduced from Cullen, T. J., Walker, M. A., Eastwood, S. L., Esiri, M. M., Harrison, P. J., and Crow, T. J. 2006. Anomalies of asymmetry of pyramidal cell density and structure in dorsolateral prefrontal cortex in schizophrenia. Br. J. Psychiatry 188, 2631, Elsevier.
How are the symptoms to be explained? On the basis that the anatomical changes reflect an alteration in connectivity and that the asymmetry of the human brain (the torque) is the foundation of the faculty of language, one can construct a theory of nuclear symptoms that these are primary disorders of the structure of language, and that they reveal its constituent elements and the way in which the elements are segregated within and relate to each other between the four quadrants of heteromodal association cortex.
4.36.6.1 The Linguistic Sign Is Bihemispheric
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this way, age of onset of psychosis may reveal to us the operation of sexual selection in humans in determining the limits of species-specific phenotypic variation (Crow, 1993, 1998a, 2002c) (Figure 3).
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de Saussure (1916) maintained that the linguistic sign (the word) was characteristically bipartite,
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comprising a signal (the sound pattern or phonological engram) and a signaled (the associated concept or meanings). The association between the sound pattern and its meanings, according to de Saussure, is arbitrary any sound pattern can be associated with any concept or meaning (the first principle). This is what is distinctive about the human use of words and what makes language so flexible. There is a two-way relationship between the components, with movement from the sound pattern to the meanings in speech reception, and from the concepts to sound patterns in speech production. One can ask, what is the neural basis of the separation of the two components? If asymmetry is what is characteristic of the human brain, it seems that there must be a relationship between specialization of function of the hemispheres and the feature that de Saussure identifies as the key to language. The most parsimonious hypothesis is that the components are (at least in part) segregated to the two hemispheres. From Brocas observations, it is clear that what is localized in the hemisphere that is labeled as dominant is the phonological engram. It follows that some part of the signifieds must be assumed to be located in the nondominant hemisphere. For each phonological engram, there must be a corresponding engram a mirror image in the nondominant hemisphere, but one that is systematically transformed by the differing terminations of the interhemispheric connections in that hemisphere. de Saussures second principle is that speech is linear, just a ribbon of sound. Allied to this is the notion that there must be a speaker and a hearer speech is necessarily communicative and the ribbon of sound is what travels between them (Figure 4). One envisages therefore that speech is encoded, and this is a bihemispheric process, by the speaker from his concepts or thoughts into phonological engrams that are then transformed into the ribbon of sound, and that this is received by the hearer, and decoded into his own meanings or concepts, and that this decodification takes place partly by interhemispheric interactions. Communication depends upon the hearer sharing at least some of the
speakers signifiersignified associations, in other words, that they speak the same language.
4.36.6.2 Deixis and the Significance of the Indexical
The system works well so long as the speaker refers to the world outside himself and the hearer. But a complication arises when he refers to himself. As Hurford (1992) points out, such a referral necessitates further decoding on the part of the hearer that the I that the speaker refers to, relates not to the I of the hearer but to the you (to him) of the speaker. This class of symbols the indexicals referring to the speaker or hearer belongs to the wider class of deictic symbols that include reference to the here of the present place and the now of the present moment in time. The interest is that deixis the necessity to define this class of symbols by pointing has a special status in philosophy and in the structure of language. According to Buehler (1934), this triad of terms defining the present moment and the location and identity of the speaker is the coordinate origin around which language is structured this place, at the present moment in time, defined by the I of the speaker. Without this, language has no point of reference and loses its capacity to convey meaning. The concept of the indexical also relates to the nature of psychotic symptoms. While it is true that there is no general misuse of the first person pronoun in individuals with psychosis, those with earlyonset developmental disorders such as autism and Aspergers syndrome sometimes have difficulty in acquiring the distinction between the use of I and you. The more general significance arises in relation to Hurfords point that these symbols relate to what is self-generated in speech and what is othergenerated that there is a fundamental dichotomy between speech production and speech perception and that this dichotomy can be understood in terms of the brain torque (Figure 5).
4.36.6.3 The Human Brain as a Four-Chambered Organ
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Figure 4 The relationship between the speaker and the hearer according to de Saussure (1916).
It is often overlooked that the asymmetry of the human brain is not a simple leftright difference but a deviation across the fronto-occipital axis that transforms the human brain from the standard primate and vertebrate pattern of two-chambers (anterior and posterior corresponding to motor and sensory compartments) into a four-chambered organ in which motor and sensory compartments are distinguished on the left and the right sides. The torque has the effect of differentiating the two
Anterior Anterior
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Figure 5 The Yakovlevian torque from right frontal to left occipital. Figure 6 The implications of the torque for the relationship between the four areas of association cortex. Interhemispheric fibers can merge from right to left anteriorly and from left to right posteriorly. The figure also illustrates the segregation of function that has allowed the separation of thoughts from speech production and meaning from speech perception.
sides of the brain by influencing the relative surface area of the cerebral cortex, but it does so in different directions in the anterior (motor) and posterior (sensory) halves. It is a remarkable fact that the volumes of the two hemispheres are closely similar. What has changed relative to other primates is either the distribution of tissue between the two sides along the anteroposterior axis or perhaps even more subtly the sulcogyral folding of the cortical surface. The effect must be assumed to be that the distribution of interhemispheric connections differs on the two sides and it is this that allows the spread of neural activity to be systematically different on one side compared to the other. But the direction of the difference is opposite in the motor and sensory halves of the brain converging from right to left anteriorly and from left to right posteriorly (Figure 6). If the torque is what is characteristic of the human brain, these changes, and no others, are critical to the evolution of language. The interconnections between areas of heteromodal cortex have changed and these changes alone must account for what is characteristic of the capacity for language. Two features are suggested the arbitrariness of the association between the signifier and the signifieds, the phonological engram and its meanings (de Saussure, 1916), and universal grammar, a mechanism that generates the structure of the sentence (Chomsky, 1988). But the striking fact about the evolution of language is that it occurred relatively abruptly, in the transition to modern H. sapiens, and all of a piece (Bickerton, 1990, 1995) along with the capacity for symbolic representation (Mellars, 2002). It does not make sense to suppose
that there were two sequential innovations, each contributing a revolution to brain function. These two features must reflect different aspects of the same change and that change must be either the introduction of the torque or some modification of a torque that was introduced earlier in hominin evolution. The key change is the separation of a phonological engram from its associations. The human cortex is not qualitatively different from that of the chimpanzee or any other primate. It must presumably have the same capacity to receive inputs and to transform them into outputs. What is different about the human brain is the interaction between areas of (particularly heteromodal) cortex on the two sides. Thus, if we assume that what is segregated in the dominant hemisphere is the phonological engram a collection of simple but heavily interconnected motor sequences this leaves open the possibility that each of these motor patterns has connections with engrams that are systematically different (either more diffuse or more restricted) in the nondominant hemisphere. But here is the key consequence of the torque whether the connections are more diffuse or more restricted depends upon whether they are in the motor or sensory halves of the brain. The convergences are in different directions in the anterior and posterior heteromodal association cortices, and this has an obvious implication for the organization of the capacity for language. Whatever transformation takes place from right to left in dorsolateral prefrontal cortex is reversed in sense in the transition from
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left to right in occipitoparietotemporal cortex. This can be regarded as the first principle in the neural organization of language. The second principle is that the sensory phonological engram is distinct from the motor engram. This must obviously be the case and these engrams are presumably located in Wernickes and Brocas areas, respectively. But it is not so obvious that the form of the engram must be different the motor engram can be relatively directly associated with motor neurones and with the output, but the sensory engram is one step removed from the acoustic input word traces need to be filtered out from the totality of incoming sensory information. Nonetheless, there is a relationship between the two. Words that are heard are recognized as related if not identical to those that are spoken. The form of the relationship presumably is what is established in the course of language acquisition. Conversely, some aspect of the distinction between the two is what is lost in the case of auditory hallucinations; what is clearly intrinsically generated (whether in the process of thought or in motor planning) has activated engrams that are normally accessible only to incoming acoustic stimuli. In this distinction, and in the association of the signifier and the signifieds, lies a solution to the conundrum of psychosis. The four chambers of the human brain are the framework within which the problem can be solved, in the separation of the motor from the sensory by the central sulcus, and in the segregation of the signifier (the phonological engram) in its two forms in the dominant hemisphere from its primary associations in the nondominant, by the bias of the torque. The key to the solution is to specify the nature of the difference between what is motor and what is sensory and to identify what is intrinsic to the signifier and what is arbitrarily associated with it, on the one hand in its sensory form, and on the other in its motor configuration. Speech is versatile what has to be accounted for is the infinity of sentences that can be generated and the diversity of meanings that can be extracted, and the fact that these two processes are integrated with each other and yet separated in the brain. The simplest approach is to assume that contiguous functions are dealt with in anatomically related areas. Thus, if the phonological engrams of the output are assembled in the association areas of cortex that are focused on Brocas area in the left hemisphere, one must assume that the processes that are related to this assembly, but differ from it in the crucial respect that they confer upon it its arbitrariness and flexibility, are located in the homologous regions in the right hemisphere. In functional terms,
these engrams can be loosely referred to as thought, the precursor of, or the plans for, speech. It has a relationship to speech in that each element has an associated phonological engram in the left hemisphere, but it is not speech in that without the linear sequence that forms the output on the left, the association between elements is, to a certain degree, random and arbitrary. This presumably is related to the fact that each phonological engram on the left is associated with a number of less well-structured engrams on the right. The relationship from right to left is many-to-one. In the parietotemporo-occipital junction association areas, the convergence is in the opposite direction. From the phonological engrams that have been extracted from the primary acoustic signal in the auditory association areas on the left, there is a convergence to a smaller area of homologous cortex on the right. Thus, the many-to-one transition in this case is from left to right, and the process of simplification may be identified as the distillation of meaning from the linear sequence of phonology on the left. According to this concept, the transition from thought to speech takes place from right to left in dorsolateral prefrontal cortex, and from perceived speech to meaning from left to right in parietotemporo-occipital association cortex. But what is the relationship between perceived meaning and the thought that is the plan for speech? Both are concepts or associations (signifieds) removed from the phonological engram, but one is sensory and one is motor. There is a connection between them and that connection is mediated by the uncinate and arcuate bundles in the right hemisphere. The patterns of activity in the right parietotemporo-occipital cortex is thus accessible to the thought processes in right dorsolateral prefrontal cortex, but the activities are distinct, and the distinction differs from that between the activity in the association cortices around Brocas and Wernickes areas.
4.36.6.4 The Deictic Origin and the Performative Hypothesis
The key here is syntax the ability to relate the self to the outside world and to use words to do it. According to Buehler, as noted above, language is built around a coordinate framework with an origin in the self, the present location and the present moment in time defined by the deictic symbols I, here, and now. Without this, language loses its structure. Language mediates the individuals relationship with the outside world and other individuals. In every linguistic interaction, there is
a speaker and a hearer, and sentences are the substance of the negotiation between them. The distinction between what is self- and what is othergenerated and the meanings of these two classes of symbol is critical. This distinction clearly relates to the division between the motor and sensory association cortices in the right hemisphere. The nuclear symptoms of schizophrenia tell us what happens when the distinction breaks down. Thought, the precursor of speech, loses its characteristic of independence from the outside world thoughts are inserted into or removed from the individuals mind while retaining the features of thought, they have lost the relationship to self-generated acts, which is a defining feature of thought as a precursor of speech. The obvious interpretation is that they are influenced by activity in posterior association cortex in a way that differs from the normal exchange between posterior and anterior regions. Conversely, auditory hallucinations such as thoughts spoken aloud or running commentary presumably represent self-generated neural activity (thoughts or plans for action) that activates phonological engrams (perhaps in the superior temporal cortex on the left side) that are normally activated by speech from another individual. In each case, there is a loss of the boundary in symbolic representation (words) between what is self- and what is other-generated. In each case, we can see that the boundary has something to do with what is anterior and what is posterior in association cortex. But we cannot suppose that what is abnormal is simply that some neural activity crosses this boundary. Activity must normally be transmitted between posterior and anterior along, for example, the arcuate and uncinate bundles. The problem is to specify what is normal and what is abnormal. The nature of nuclear symptoms is the clue that we have (Figure 7).
The possibility must be considered that Buehlers formulation that language is intrinsically organized around a deictic core and the distinction between what is self- and what is other-generated that is illustrated by the nature of the nuclear symptoms are both necessary to a solution of the problem of syntax. In his thesis of How to Do Things with Words, Austin (1962) made the case that many uses of language are not simply to convey information but to have an effect, that is to say to bring about a state of affairs according to the speakers intention. Such utterances Austin referred to as performatives to distinguish them from the constatives, the utterances, more usually the subject of linguistic and philosophic analyses, that convey information, generally about the external world. Later he generalized the concept of the performative into the notion that all utterances have an illocutionary force in the sense that they are formulated toward some objective of the speaker. An interesting parallel to Austins concept in the performative hypothesis has been formulated in different forms by a small number of theoretical linguists to account for certain features of the use of indexicals (symbols that refer to the speaker or hearer). Ross (1970) defines the theory as that declarative sentences (constatives) must also be analysed as being implicit performatives, and must be derived from deep structures containing an explicitly represented performative main verb. Performative sentences, he says:
. . . must have first person subjects and usually have second person direct or indirect objects in deep structure. They must be affirmative and non-negative, they must be in the present tense, and . . . their main verb must belong to the class that includes verbs such as advise, answer, ask, beg, command, declare, implore, inform, pronounce, say, write, in other words the class that designates the transmission of information, instructions, orders etc. The implication of the performative hypothesis is that the declarative sentence has an implicit (unstated) superordinate clause of the form I say unto you . . . in the first person and the present tense.
Thoughts spoken aloud Running commentary
Alien thoughts
Figure 7 The origin of nuclear symptoms forging the concept of the compartmentation of language. According to this concept, symptoms arise because leakage occurs of neural activity normally segregated within one or more of the chambers.
Austins concept and the performative hypothesis bear a relationship to Buehlers notion of a deictic origin to the coordinate frame of language, and to de Saussures insistence that language can only be understood in terms of the relationship between the speaker and the hearer. Without the deictic frame, Buehler insists the structure of language dissolves. This may be what happens in the case of thought disorder the determining focus is lost. What the nuclear symptoms of schizophrenia are telling us is what happens when the distinction between the indexicals I of the speaker and you of the hearer begins to dissolve. If the performative hypothesis is
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right, thus conceived they are disorders of the foundations of syntax. They tell us that the phonological engram for the perception of speech is quite separate from the phonological engram for speech production. They tell us that thoughts as the precursor to speech are distinct from the meanings that are extracted from perceived speech in the nondominant hemisphere. They draw attention to the obscure process of the motor and sensory elements of the associations (the signifieds) that takes place in the nondominant hemisphere (Mitchell and Crow, 2005). In each case, the phenomena of psychosis provide evidence on the neural organization of language through what happens when the mechanism goes wrong. We begin to understand this through the structure of the torque.
4.36.7 XY Homology and the Xq21.3/Yp Translocation

The most fundamental prediction of the asymmetry hypothesis is that the genetics of psychosis is the genetics of the speciation of H. sapiens (Crow, 2004); in other words, the genetics of asymmetry is conceived as the species-defining characteristic. To approach such predictions, it is necessary that the cerebral dominance gene or right shift factor be identified. An important clue comes from sex chromosome aneuploidies. Individuals who lack an X chromosome (XO, Turners syndrome) have nondominant hemisphere (spatial) deficits on cognitive testing. Individuals with an extra X (XXY, Klinefelters and XXX syndromes) have verbal or dominant hemisphere deficits (Table 1). A possible explanation is that an asymmetry determinant is present on the X chromosome. But then the question arises of why males, who only have one X chromosome do not have spatial deficits such as are seen in Turners syndrome. The answer must be that the copy of the gene on the X chromosome is complemented by a copy on the Y, i.e., that the gene is in the XY homologous class (Crow, 1993). A hormonal explanation will not account for the similarity of the changes in XXY individuals, who are male, and XXX individuals, who are female. The case that the
gene is present also on the Y chromosome is strongly reinforced by the verbal deficits/delays that are observed in XYY individuals (Geerts et al., 2003). The hypothesis is further strengthened by evidence that Turners and Klinefelters syndrome individuals have corresponding deviations in anatomical asymmetry (Rezaie et al., 2004) and by the demonstration of a same sex concordance effect the tendency for handedness and sex to be associated above chance expectation the hallmark of XY linkage (Corballis et al., 1996). A role for an XY homologous gene is consistent with the presence of a sex difference brain growth is faster (Kretschmann et al., 1979) and lateralization is stronger (Crow et al., 1998) in females. When we come to consider where such a gene might be located, there is an important lead. A major chromosomal rearrangement took place in the course of hominin evolution. Two regions on the human Y chromosome short arm share homology with a single region on the human X chromosome long arm (Xq21.3) (Page et al., 1984; Lambson et al., 1992; Sargent et al., 1996). These homologies were created by the translocation of a 3.5Mb contiguous block of sequences from a chimpanzee hominin precursor X chromosome to the Y chromosome short arm that was subsequently split by a paracentric inversion (by a recombination, presently undated, of LINE-1 elements (Schwartz et al., 1998; Skaletsky et al., 2003) to give two blocks of homology in Yp11.2 (Figure 8). Genes within this region are therefore present on both the X and Y chromosomes in H. sapiens but on the X alone in other great apes and primates, and an explanation for the retention of the sequences on Yp presumably lies in the gene content of this block. Three genes are known to be expressed within this region; PABPC5, a poly (A) -binding protein whose Y gametologue has been lost during hominin evolution; TGIF2LX and Y, (homeobox-containing genes with testis-specific expression), and ProtocadherinX (PCDH11X) and ProtocadherinY (PCDH11Y). PCDH11X and Y (each comprising seven extracellular cadherin motifs, a short transmembrane region, and an intracellular cytoplasmic tail) that code for cell surface adhesion molecules of the cadherin
Table 1 Neuropsychological impairments associated with sex chromosome aneuploidies XX Normal female No. of sex chromosomes Verbal ability Spatial ability 2 Normal Normal XY Normal male 2 Normal Normal XO Turners syndrome 1 Normal Decreased XXY Klinefelters syndrome 3 Delayed Normal XXX 3 Delayed Normal XYY 3 Delayed Normal
a 11.3 11.2 11.1 11.1 11.21 11.22 11.23 12 k 11.1 11.1 13 21.1 21.3
a b c d e f g h i j k Yp PAR1 RPS4Y/RPS4X ZFY/ZFX Distal Yp11.2/Xq21.3 AMGY/AMGX Proximal Yp11.2/Xq21.3 Yq11.21/Xp22.3 SMCY/SMCX-HY antigen Yq11.22/Xq28 Yq11.23/Xq22.2 Yq PAR2
22.3 22.2 21.3
11.3
22.2
28
Figure 8 The regions of homology between the X and the Y chromosomes including PAR1 at the telomeres of the short arms and PAR2 at the telomeres of the long arms. Blocks of homology are labeled from a to k on the Y chromosome. Some blocks are identified by their gene content, e.g., RPS4Y/RSP4X, AMGY/AMGX, SMCY/SMCX. Reproduced from Affara, N., Bishop, C., Brown, W., et al. 1996. Report on the second international workshop on Y chromosome mapping 1995. Cytogenet. Cell Genet. 73, 3376, Elsevier.
superfamily are of note because both forms of the gene have been retained and are highly expressed both in fetal and adult brain (Yoshida and Sugano 1999; Blanco et al., 2000), including the germinal layer of the cortex (T. H. Priddle, personal communication). The protein products of this gene pair are thus expected to play a role in intercellular communication, perhaps acting as axonal guidance factors and influencing the connectivity of the cerebral cortex. These genes may thus have been subject to selective pressure relating to one or more brain characteristics during hominin evolution.
to consider concepts of evolutionary change other than Darwinian gradualism and the biological or isolation species concept. The theory of punctuated equilibria (Eldredge and Gould, 1972) was preceded by a long history of challenges to the gradualist version (Bateson, 1894; De Vries, 1901; Goldschmidt, 1940), and some of these have had an explicit genetic basis. Thus, White (1978) and King (1993) have argued strongly for a role for chromosomal change in speciation, but their arguments have not overwhelmed the established view. Against chromosomal change, it is argued (e.g., Coyne and Orr, 1998) that radical rearrangements, for example chromosomal fusions, may apparently have few phenotypic effects, and in some cases alternative chromosomal configurations persist, as it were, as a polymorphism within a species. The case for saltation in species transitions has been argued at a macroevolutionary level (Stanley, 1998), stating that the amounts of change seen within species and other taxa are simply insufficient to account for the overall pattern of evolutionary change that is seen over time, and at the level of morphology (Mellars, 1998), claiming that the intermediate states in the transition between species that are required by the gradualist theory are absent. But all such general arguments come up against the difficulty that Goldschmidts (1940) hopeful monster ran into: the greater the magnitude of the saltational change, the less likely it is to have survival value, and the greater the difficulty the hopeful monster will have in identifying a mate. The difficulty is particularly great if the change has the reproductive consequence of reducing fertility in the hybrid state. The possibility that the monster can identify an individual with the same mutation is clearly dependent on reproduction already having taken place, and even then the new mutation is at a severe statistical disadvantage with respect to the existing population.
4.36.8.2 Sexual Selection and the Mate Recognition Principle
4.36.8 Implications for Evolutionary Theory

4.36.8.1 The Case for Saltation
To take seriously a structural discontinuity as an explanation for a species difference, it is necessary
However, here Darwins (1871) juxtaposition of The Descent of Man and the theory of sexual selection offers a way out. If sexual selection and speciation were in some way interdependent this might solve the problem of discontinuity and that of mate selection. The case of cerebral lateralization in modern H. sapiens illustrates the possibility. All authorities on the genetics of lateralization (Annett, 1985, 2002; McManus, 1985; Corballis, 1997; Crow, 1998a, 1998b; Mckeever, 2000) agree that there is a sex difference: females are more
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right-handed than males (although the adult male brain is more asymmetrical than that of the female; Bear et al., 1986; Barrick et al., 2001). The female brain grows faster than that of the male (Kretschmann et al., 1979) and females have greater mean verbal fluency and acquire words earlier (Maccoby and Jacklin, 1975; McGlone, 1980; Halpern, 2000) than males. If language is the species-defining characteristic and lateralization is the process by which it evolved, these facts are related, and they tell us about the nature of the genetic mechanism. Only two explanations of the sex difference in lateralization are conceivable, that it is hormonal in origin (Geschwind and Galaburda, 1985) or that it reflects a sex chromosomal locus (Crow, 1993, 1994), and the facts of sex chromosomal aneuploidies (XXY and XXX individuals who differ in hormonal status have similar hemispheric deviations in development) speak decisively in favor of the latter interpretation. The hypothesis that the asymmetry factor is present on both X and Y chromosomes (Crow, 1993; Corballis et al., 1996) can explain the transmission of handedness within families and apparent dosage effects in the aneuploidies. That there are problems (Corballis, 1997) in accounting for persisting variation in males and females in terms of conventional polymorphisms and heterozygote advantage explanations should not dissuade us from pursuing the line of thought. The genetic principles involved may not be those on which we have hitherto relied. The paradigm of H. sapiens therefore suggests a new version of saltational speciation that it is not chromosomal changes in general that play a role in speciation but changes on the sex chromosomes, and perhaps particularly changes in regions of XY homology that are involved. These regions have a special status because they can account (as in the case of lateralization in humans) for quantitative differences in a characteristic in males and females, and such quantitative differences are a potential substrate for sexual selection. The Y chromosome itself has a unique role, because it is not necessary for survival. There are interindividual differences on the Y (reviewed by Tyler-Smith, 2002), but there are also large interspecific differences. While the X is the most stable chromosome across species, the Y is by far the most variable. The mammalian Y therefore can be seen as a testbed of evolutionary change. One possibility is that the primary change in speciation takes place on the Y, and when it is located in a region of homology with the X, there is the possibility of correlated but independent change in the two sexes. Such correlated but quantitatively differing ranges of variation have
the potential to explain the type of runaway sexual selection envisaged by Fisher (1930), and this may be what occurred with respect to cerebral asymmetry at some point in hominin evolution (Crow, 1998a, 1998b); the introduction of the dimension of symmetryasymmetry allowed brain growth to equilibrate at a new point of plateau, and this equilibration took place around successive modifications of genes on the Y and then on the X chromosome. There is thus a potential three-way relationship between sexual selection, sex linkage, and speciation, and the pattern suggested by hominin evolution is backed up in the recent literature relating to other species. A role for sexual selection in modifying a primary change in a sexually dimorphic feature to establish a new species boundary has been argued in relation to Hawaiian Drosophilid species by Kaneshiro (1980) and Carson (1997). Similar arguments apply in the case of the prolific speciation of cichlid fishes in the lakes of East Africa (Dominey, 1984; McKaye, 1991) and may also apply in birds (Price, 1998). Some putative speciation loci, for example, the Odysseus homoeobox (Ting et al., 1998) and the per gene (Ritchie and Kyriacou, 1994) that have been identified in Drosophila species, are X-linked. In discussing the relationship between the X chromosome and speciation that she finds in Lepidoptera, Prowell (1998) offers three explanations: (1) that X-linked traits evolve more quickly, (2) that traits related to speciation tend to be sex-limited, and that sex-limited traits tend to be on the sex chromosomes, and (3) that female-limited X-linked traits undergo faster rates of evolution when, as in the case of Lepidoptera, the female is heterogametic. These explanations are not mutually exclusive. Prowell asks whether the X chromosome bias is unique to Lepidoptera and concludes that this is unlikely. Haldanes rule is that when, in a species hybrid, one sex is sterile or inviable it is the heterogametic sex. Coyne and Orr (1998) consider various explanations, including faster evolution and recessivity of genes on the X chromosome. While each of these observations and hypotheses is consistent with a relationship between speciation and the sex chromosomes, none of the authors considers the more restrictive formulation suggested by the sequence of events (Sargent et al., 1996, 2001, 2002) on the mammalian Y chromosome, that it is the interaction between the sex chromosomes, particularly the possibility of transfer of material between them, that is critical in speciation.
4.36.8.3 Species-Specific Variation Is Epigenetic
Reinhold considers the case that sexual selection acts selectively on sexual dimorphisms that relate
to sex-linked genes, as suggested by Rice (1984). The sequence of events, including a translocation and a paracentric inversion, suggested by the work summarized by Sargent et al. (2002) and by the XY hypothesis as relevant to the course of hominin evolution, carries the further implication that epigenetic modification is involved in the process of sexual selection and speciation. In mammals, genes on one X chromosome are subject to the process of X inactivation, but gene sequences that are also represented on the Y chromosome are protected from this influence. Such genes are expressed from both X and Y in males and from both Xs in females, a similar dosage thus being maintained in each sex. The mechanism by which this protection is achieved is unknown (Burgoyne and McLaren, 1985; Crow, 1991). Gene sequences that have been transferred from the X to the Y are in a new situation; whatever the mechanism, a phase of epigenetic equilibration must be assumed (Jegalian and Page, 1998). If XY pairing in male meiosis plays a role, the orientation of the sequence on the Y is also relevant. The paracentric inversion on the Y short arm could be critical. An MRI investigation in monozygotic twins of handedness and asymmetry of the planum temporale (Steinmetz et al., 1995) indicates that there is room for an epigenetic influence on cerebral asymmetry, and this may account for the stochastic element incorporated in genetic theories (Annett, 1985, 2002; McManus, 1985). There is a possibility, therefore, that the genetic mechanisms underlying the development of cerebral asymmetry in humans are a paradigm for a more general interaction between genetic and epigenetic mechanisms in sexual selection and speciation. Perhaps sexual selection and natural selection are mediated by distinct but complementary genetic processes: natural selection ensures the organisms survival through response of any part of the genome to environmental change, whereas sexual selection reflects the sequential response of the female genome (for example, the mammalian XX complement) to change on the Y chromosome, and that this process involves particularly the epigenetic modulation of genes on the X. According to this concept, speciation follows the history of the nonrecombining sex chromosome, in mammals the Y.
4.36.8.4 Speciation Events Occur on the Heterogametic Chromosome
Birdsong has relevance to the genetics of speciesspecific characteristics (see The Evolution of Vocal Learning Systems in Birds, The Evolution of Vocal
Learning Systems in Birds and Humans). The neural capacity is lateralized, season-dependent, and strongly sexually dimorphic, being generally confined to males. The lateralization is a parallel and independent evolutionary development to the cerebral torque of the human brain, such lateralization being absent in intervening vertebrate families such as rodents and primates, and the neural mechanisms are quite distinct. But the individual songs of birds, like the capacity for human language, are speciesspecific. In that they are a component of mating behavior, they can be regarded as specific mate recognition systems as described by Paterson. Whereas there are elements, for example the season-dependence that is hormone-sensitive, there are also components that are determined by genes and these are located on the Z and W chromosomes (Arnold, 2004). The hormone-sensitive elements are generally those that cross species, whereas the components that are species-specific are mostly genetically determined. Thus, there is the possibility that the speciesdefining characteristic in song birds, which can reasonably be described as the mate recognition system, is dependent upon genes present on both Z and W chromosomes. Moreover, it appears that the differences between species in this salient characteristic must be determined by genetic differences at the relevant loci on the sex chromosomes. This theory attributes general significance to XY (or WZ) homologous genes in relation to a mate recognition system in Patersons sense, and thereby accommodates the role for an XY homologous determinant of cerebral asymmetry and language in humans, as suggested above. Perhaps one can propose that it is events on the heterogametic chromosome (the Y in mammals) that have particular significance in relation to speciation, and that it is the interaction between those sequences on the Y that have changed and the pre-existing sequences on the X that is of particular interest. For while the X is remarkably stable in structure across species (the gross structure of the mouse X closely resembling that of the chromosome in humans) the structure of the Y is highly variable even between quite closely related species. Thus, one can postulate that the primary change in speciation takes place on the Y (more generally the heterogametic chromosome), and the subsequent changes on the X establish the stable sexual dimorphisms that distinguish species. Ellegren has pointed to a possible molecular correlate of such a sequence. Considered in relation to the rest of the genome and when relative population size in terms of the two sexes is taken into account, the genetic variation on the heterogametic chromosome (the Y in mammals, the W in birds) is
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considerably less than would be expected. The reduction in polymorphic loci may reflect a selective sweep at the origin of the species. The hypothesis emerges that the primary event that leads to a new species is a change (maybe a duplicative translocation as in the hominin lineage, but deletions or insertions are also possible) that occurs on the heterogametic chromosome. The change occurs in one individual (a male in mammals, a female in birds) that represents the rare case in which such a chromosomal change is associated with a reproductive advantage in that such individuals are preferentially selected as mates. Thus, the new chromosome increases in frequency in the population along with its associated phenotypic innovation, but in the first generation is confined to the heterogametic sex. But in the next generation there is the possibility that there will be an impact on the homogametic chromosome because the altered structure of the Y (or the W) has the potential to influence the epigenetic status of the X (or the Z) because regions of the sex chromosomes that are paired are protected from the process of inactivation (meiotic suppression of unpaired DNA) that otherwise leads to the inactivation of one of the two Xs or Zs in the daughters or sons, respectively, of the founder individual and his or her immediate same sex progeny. Such epigenetic change will be particularly relevant if the primary change occurs in or generates a region of X/Y (or Z/W) homology. Thus, homologies may perhaps be preferentially selected by the speciation process. If the primary event occurs within the nonrecombining region of the heterogametic chromosome (the mammalian Y or the avian W), this allows genes relating to the speciation characteristic and its homogametic homologue and related genes to undergo sequence variation that is independent in the two sexes. The variability in this genetic process could account for the diversity of sexual dimorphisms across species. These proposals deriving from the case of hominin evolution bear a relationship to concepts of speciation in Drosophila developed by R. S. Singh and colleagues as well as to the literature on sexual selection and speciation referred to above. Civetta and Singh (1999) distinguished between sex-related and non-sex-related genes and found a higher rate of Ka/Ks ratios among the former. They suggested that directional sexual selection had shaped the evolution of sex-related genes (defined in a broad sense to include all components of sexuality not confined to courtship and mating) and that these changes were more likely to have occurred in the early stages of speciation. Developing these ideas, Singh and
Kulathinal (2005) emphasize the rapid evolution of sex-related traits in a wide variety of taxa, the faster rate of DNA sequence divergence in genes affecting sexual function and fertility, and the evidence for involvement of novel traits and genes in sexual function.
4.36.9 Conclusions
1. Language and the paradox of psychosis: The faculty of language as a defining feature of H. sapiens with characteristics absent in the communicative systems of the great apes challenges Darwinian gradualism. It is more readily assimilated to a saltational account of speciation as suggested by T. H. Huxley and developed by R. Goldschmidt. The paradox is that interindividual variation apparently genetic in origin persists at approximately the same frequency in all populations in the face of a fecundity disadvantage. This variation represents a component of variation associated with the capacity for language; it is argued that the phenomena of psychosis are the key to an understanding of the neural organization of language. Thus, psychosis and language have a common evolutionary origin in the speciation event. 2. The cerebral torque and its implications: Brocas hypothesis that cerebral asymmetry is the characteristic that defines H. sapiens is supported by recent cross-species comparisons for directional handedness and anatomical asymmetry. The cerebral torque (the bias from right frontal to left occipital across the anteroposterior axis) defines the human brain as a four-chambered organ by comparison with the two chambers (anterior motor and posterior sensory) of the brains of other primates, and dictates a reversal of sign of the convergence of interhemispheric connections (from left to right posteriorly and from right to left anteriorly). 3. The quadripartite structure of language: These transitions are critical to the separation of the sensory and motor phonological engrams in the dominant left hemisphere from some of their associated signifiers (the sensory meanings and the motor thoughts) in the nondominant hemisphere. Critical to the distinction between the speaker and the hearer and to what is motor and what is sensory in the neural representation of speech is the notion (Buehlers) of a deictic origin (I,
here, now) to the coordinate system of language. The nuclear symptoms of schizophrenia (e.g., thoughts spoken aloud, running commentary, thought insertion) are seen as the ectopic presence (leakage) of one or more of the components of language outside the relevant compartment of association cortex. The deictic origin is located in Brocas area and defined by its interaction through the uncinate and arcuate bundles with Wernickes area. 4. The genetic basis of the speciation event: There is a strong case from the phenomena of sex chromosome aneuploidies and a family study of handedness that the cerebral dominance gene is in a region of homology between X and Y chromosomes. A duplicative translocation 6Mya created a sapiens-specific region of homology between Xq21.3 and Yp11, within which there have been subsequent rearrangements including a paracentric inversion. Within the region of homology, the ProtocadherinXY gene pair coding for cell surface adhesion proteins has been subject to accelerated evolution. 5. Implications for evolutionary theory: The sequence of events in the speciation of H. sapiens was saltational in form. It suggests the general theory of speciation that rearrangements on the heterogametic chromosome play a role by initiating a phase of sexual selection that, on account of sequences shared between the sex chromosomes, involves related characteristics in the two sexes. The interaction establishes a mate recognition system (in Patersons sense) that defines the new species.
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Further Reading
Annett, M. 2002. Handedness and Brain Asymmetry: The Right Shift Theory. Psychology Press. Bickerton, D. 1995. Language and Human Behavior. University of Washington. Chaika, E. 1990. Understanding Psychotic Speech: Beyond Freud and Chomsky. C. C. Thomas. Crow, T. J. (ed.) 2002. The Speciation of Modern Homo sapiens. Oxford University Press.
Crow, T. J. 2005. Who forgot Paul Broca? The origin of language as test case for speciation theory. J. Ling. 41, 133156. DeLisi, L. E. 2001. Speech disorder in schizophrenia. A review of the literature and exploration of its relation to the uniquely human capacity for language. Schizoph. Bull. 27, 481496. Huxley, T. H. 1863. Evidence of Mans Place in Nature. Williams and Norgate. Lambert, D. M. and Spencer, H. G. (eds.) 1994. Speciation and the Recognition Concept: Theory and Application. Johns Hopkins Press. Penner, J. G. 2000. Evolution Challenged by Language and Speech. Minerva. Sims, A. C. P. (ed.) 1995. Speech and Language Disorders in Psychiatry. Gaskell.

4.36 Nuclear Schizophrenic Symptoms As The Key To The Evolution of The Human Brain

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4.36.1 4.36.2 4.36.3 4.36.4 4.36.5 4.36.6

4.36.1 The Nuclear Symptoms of Schizophrenia and the Central Paradox

4.36.2 The Problem of Language for Evolutionary Theory

In the second lecture Mueller addresses the problem:

4.36.3 Darwins Intuition on Sexual Selection

4.36.4 Paul Broca and Cerebral Asymmetry

Chimpanzee Human 100 L 50 0

Marchant and McGrew Provins et al. 50 100 R

Number of Activities Individuals Chimpanzee Human 38 1960 46 75

4.36.5 The Torque and Related Asymmetries in Psychosis

Gyrus volume (cm3)

Gyrus volume (cm3)

L R L R Control Schizo n = 14 n = 13 Female

L R L R Control Schizo n = 15 n = 17 Male

Neurons per mm3

4.36.6 The Structure of Language and Its Decomposition in Psychosis

Speech generation (phonology)

Thought plans for action

Thoughts spoken aloud Running commentary

4.36.7 XY Homology and the Xq21.3/Yp Translocation

22.3 22.2 21.3

4.36.8 Implications for Evolutionary Theory

To take seriously a structural discontinuity as an explanation for a species difference, it is necessary

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