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1 For a history of the identification of sensory systems see M. R. Bennett, S. Hatton, D. F. Hermens and J. Lagopoulos, ‘Behaviour, neuropsychology and fMRI’, Progress in Neurobiology, 145–6 (2016), pp. 1–25.

2 Aristotle, De Anima 412a20. Subsequent references to this treatise in the text will be flagged ‘DA’.

3 It should be noted that Aristotle held that a sightless eye is no more an eye than a painted eye, just as a corpse is no more an animal than a statue.

4 Note that when Aristotle says that we do these things with our soul, this is not like doing something with our hands or eyes, but rather like doing something with our talents and abilities.

5 We disregard here the complexities, and incoherences, that arise with regard to Aristotle’ s distinction between the active and passive intellect and the intimation that the active intellect may be capable of existing without a body (DA 429a18–29, 430a18–25). These passages were crucial for the later scholastic synthesis of Aristotelian philosophy of mind with Christian doctrine concerning the immortality of the soul.

6 For his reasoning, see De Partibus Animalium 647a22–34. In this respect he differed from the Hippocratic tradition. The Hippocratic lecture on epilepsy noted that ‘It ought to be generally known that the source of our pleasure, merriment, laughter and amusement, as of our grief, pain, anxiety and tears, is none other than the brain. It is especially the organ which enables us to think, see and hear, and to distinguish the ugly and the beautiful, the bad and the good, pleasant and unpleasant… . It is the brain too which is the seat of madness and delirium, of the fears and frights which assail us, often by night, but sometimes even by day; it is there where lies the cause of insomnia and sleepwalking’ (‘The sacred disease’, §17, in G. E. R. Lloyd (ed.), Hippocratic Writings (Penguin Books, Harmondsworth, 1978). The Hippocratic insight is wonderful; the physiological reasoning is, however, no less erroneous than Aristotle’ s reasoning in support of his different hypothesis.

7 Aristotle, De Somno 455a21. This is the Barnes translation; an alternative translation of the sentence in which this term appears is ‘For there exists a single sense-faculty, and the master organ is single’.

8 His term is aisthe¯sis koine¯, which occurs only in De Anima 425a27, De Memoria 450a10 and De Partibus Animalium 686a27.

9 Aristotle, De Sensu 449a5–11.

10 10 See, e.g., F. Crick, The Astonishing Hypothesis (Touchstone, London, 1995), p. 22; A. Damasio, The Feeling of What Happens (Heinemann, London, 1999), p. 320; E. Kandel and R. Wurtz, ‘Constructing the visual image’, in E. R. Kandel, J. H. Schwartz and T. M. Jessell (eds), Principles of Neural Science (Elsevier, New York, 2001), p. 492. For a discussion of the binding problem, see §5.2.3.

11 11 Aristotle, De Somno 455a15–20.

12 12 e.g. L. Weiskrantz, Blindsight: A Case Study and Implications (Oxford University Press, Oxford, 1986); Weiskrantz, ‘Varieties of residual experiences’, Quarterly Journal of Experimental Psychology, 32 (1980), pp. 365–86. See §17.3.1.

13 13 The argument is curious, inasmuch as it is unclear in what sense he thinks we discriminate white from sweet. To be sure, we possess the faculties to see white things and distinguish them from other coloured things, and to taste sweet things and distinguish them from things with other tastes, and we (language-users) also possess the concepts of white (and other colours) and sweet (and other gustatory qualities). But we do not discriminate white things from sweet things; nor do we need any further organ to differentiate white from sweet (for what would it be to confuse them?).

14 14 Aristotle, De Somno 455a.

15 15 Aristotle, De Memoria 450a9–14.

16 16 Cicero, Tusculan Disputations 1.10.22, quoted by D. Furley, ‘Aristotle the philosopher of nature’, in D. Furley (ed.), From Aristotle to Augustine, vol. iv of Routledge History of Philosophy (Routledge, London, 1999), p. 16. Note that Cicero must surely be mistaken in ascribing to Aristotle the view that the soul is made of anything.

17 17 Galenus, Hippocratis de natura hominis commentaria III, In Hippocratis de victu acutorum commentaria IV, De diaeta Hippocratis in morbis acutis, ed. J. Mewaldt (Teubner, Berlin/Leipzig, 1914), p. 70, 5–6.

18 18 H. Von Staden, Herophilus (Cambridge University Press, Cambridge, 1989), pp. 155–6.

19 19 Ibid. T 77a & 77b.

20 20 Galen, ‘On the use of the parts ’ 8.11 (III 665.7K = I. pp. 482–4. Helmrich = Herophilus Frs 77a & 78. Von Staden).

21 21 Ibid.

22 22 ‘De partibus corporis humani ’, p. 185, 5–6. Daremberg – Ruells; T81 Herophilus. Fr.125 Von Steden.

23 23 F. D. Retief and L. Colliers, ‘The nervous system in antiquity’, The South African Medical Journal, 98, no. 10 (2008), pp. 768–72. H. Von Staden, Herophilus (Cambridge University Press, Cambridge, 1989), pp. 159–206, 248, 314.

24 24 M. R. Bennett, History of the Synapse (Taylor, London, 2001).

25 25 C. Galen, Du Movement des muscles, sect. I, ch. 1, French translation by C. Daremberg, in Oeuvres anatomiques, physiologiques et médicales de Galen (Ballière, Paris, 1854–6), vol. 2, p. 323.

26 26 For further detail, see M. R. Bennett, ‘The early history of the synapse: from Plato to Sherrington’, Brain Research Bulletin, 50 (1999), pp. 95–118.

27 27 C. Galen, Des Lieux affectés, sect. IV, ch. 3, tr. Daremberg in Oeuvres, vol. 2, p. 590; C. Galen, Utilité de parties du corps, sect. IX, chs 13–14, tr. Daremberg in Oeuvres, vol. 1, pp. 593–7; see also W. H. L. Duckworth, Galen on Anatomical Procedures, ed. M. C. Lyons and B. Towers (Cambridge University Press, Cambridge, 1962), pp. 22–6.

28 28 C. Galen, Utilité de parties du corps, sect. IX, ch. 14, tr. Daremberg in Oeuvres, vol. 1, pp. 597f.

29 29 C. Galen, Hippocrates librum de alimento commentarius, sect. III, ch.1, in K. G. Kühn (ed.), Opera Omnia Claudii Galeni (Cnobloch, Leipzig, 1821–33), vol. 15, p. 257.

30 30 C. Galen, De Symptomatum Differentis, sect. VII, in Kühn (ed.), Opera Omnia, vol. 7, pp. 55–6.

31 31 C. Galen, Utilité de parties du corps, sect. VIII, ch. 6, tr. Daremberg in Oeuvres, vol. 1, pp. 541–3.

32 32 C. Galen, Des Lieux affectés, sect. IV, ch. 3, tr. Daremberg in Oeuvres, vol. 2, p. 590.

33 33 Nemesius, ‘The nature of man’, in Cyril of Jerusalem and Nemesius of Emesa, tr. and ed. William Telfer (Westminster Press, Philadelphia, 1955), pp. 341–2.

34 34 Presumably by ‘imagination’ here Nemesius means sensibility.

35 35 Nemesius, ‘Nature of man’, pp. 321 and 331f.

36 36 F. Rahman, Avicenna’ s Psychology (Oxford University Press, Oxford, 1952), p. 31.

37 37 A. L. Benton and R. Joynt, ‘Early descriptions of aphasia’, Archives of Neurology, 3 (1960), pp. 205–22. See also Antonio Guainerio’ s Opera medica (Antonio de Carcano, Pavia, 1481).

38 38 A. Vesalius, De humani corporis fabrica (Basel, 1543), bk. VII, ch. i, p. 623.

39 39 W. Singer, Vesalius on the Human Brain (Oxford University Press, Oxford, 1952), p. 40.

40 40 J. Fernel, De naturali parte medicinae (Simon de Colines, Paris, 1542); see Physiologia, bk. II, Praefatio.

41 41 Aquinas, capitalizing on Aristotle’s obscure remarks about the active intellect, argued that ‘the intellectual principle which is called the mind or intellect has an operation through itself (per se) in which the body does not participate. Nothing, however, can operate through itself (per se) unless it subsists through itself, for activity belongs to a being in act … Consequently, the human soul, which is called the intellect or mind, is something incorporeal and subsisting’ (Summa Theologiae I, 76, 1).

42 42 For a discussion of Aquinas’s philosophy of psychology, see A. J. P. Kenny, Aquinas on Mind (Routledge, London, 1993).

43 43 Fernel, Physiologia, bk. VI, ch. 13.

44 44 Ibid., bk. IX, ch. 8, p. 109a.

45 45 Descartes, Principles of Philosophy, I-9. Repr. in The Philosophical Writings of Descartes, vol. 1, tr. J. Cottingham, R. Stoothoff and D. Murdoch (Cambridge University Press, Cambridge, 1985), p. 195. Subsequent page references to this translation will be abbreviated ‘CSM’. References to the canonical Oeuvres de Descartes, ed. Ch. Adam and P. Tannery, rev. edn (Paris: Vrin/C. N. R. S., 1964–76) will be given in the form ‘AT’ followed by volume and page numbers – here AT VIII A, 7. Other references are given by section number.

46 46 Descartes, Optics, CSM I, pp. 152–75; AT VI, 81–146.

47 47 Bennett, ‘Early history of the synapse’.

48 48 Descartes, Treatise on Man, CSM I, p. 100; AT XI, 129.

49 49 Descartes, Passions of the Soul, I-7.

50 50 Ibid., I-10.

51 51 C. S. Sherrington, Man on his Nature, 2nd edn (Cambridge University Press, Cambridge, 1953), p. 151.

52 52 Descartes, Passions of the Soul, I-31.

53 53 Ibid., I-32, italics added.

54 54 Descartes, Treatise on Man, CSM I, p. 106; AT XI, 119.

55 55 Descartes, Optics, CSM I, p. 167; AT VI, 130.

56 56 There is some controversy as to whether Descartes considered the soul, a res cogitans, to be a part, i.e. the immortal part, of a human being or only a constituent substance. For discussion, see Appendix 3 below, pp. 511–12.

57 57 T. Willis, De anima brutorum (Thomas Dring, London, 1683). English translation by S. Pordage: Two Discourses Concerning the Soul of Brutes, which is that of the Vital and Sensitive of Man (Scholars’ Facsimiles & Reprints, Gainesville, FL, 1971).

58 58 T. Willis, Cerebri anatome, cui accessit nervorum descriptio et usus (Thomas Dring, London, 1681); for translation, see Tercentenary Facsimile Edition, The Anatomy of the Brain and Nerves, tr. S. Pordage, ed. William Feidel (McGill University Press, Montreal, 1965). Subsequent references in the text to this volume are flagged ‘ABN ’ followed by the page number.

59 59 Willis, Two Discourses Concerning the Soul of Brutes, pp. 43f.

60 60 Ibid.

61 61 J. Prochàska, ‘De functionibus systemis nervosi, et observationes anatomico-pathologicae’, in Adnotationum Academicarum (W. Gerle, Prague, 1784), tr. T. Laycock, as ‘A dissertation on the functions of the nervous system’, in Unzer and Procháska on the Nervous System (Sydenham Society, London, 1851), pp. 141–3.

62 62 D. Mistichelli, ‘Trattato dell’Apoplessia’ (Roma, A de Rossi alla Piazza di Ceri), tr. C. D. O’Malley, in E. Clarke and C. D. O’Malley, The Human Brain and Spinal Cord (University of California Press, Berkeley, 1968), pp. 282–3.

63 63 A. Stuart, Lecture III of the Croonian Lectures, Proceedings of the Royal Society, 40 (1739), p. 36.

64 64 R. Whytt, ‘An essay on the vital and other, involuntary, motions of animals’ (1751), repr. in A. Walker, Documents and Dates of Modern Discoveries in the Nervous System (1839), pp. 112– 22; facsimile ed. P. Cranfield (Scarecrow Reprint Corp., Metuchen, NJ, 1973).

65 65 Ibid., p. 120.

66 66 Procháska, ‘A dissertation’, p. 123.

67 67 Ibid., pp. 127–9.

68 68 For more detail, see Bennett, ‘Early history of the synapse’, pp. 103–5.

69 69 L. Galvani, ‘De viribus electricitatis in motu musculari commentarius’, De Bononiensi Scientiarum et Atrium Instituto atque Academia commentarii, 7 (1791), pp. 363–418.

70 70 C. Bell, ‘Idea of a new anatomy of the brain; submitted for the observations of his friends’, repr. in G. Gordon-Taylor and E. W. Walls, Sir Charles Bell, His Life and Times (Livingstone, Edinburgh, 1958), pp. 218–31; idem, ‘On the nerves; giving an account of some experiments on their structure and functions, which lead to a new arrangement of the system’, Philosophical Transactions of the Royal Society, 111 (1821), p. 398.

71 71 C. Bell, ‘On the functions of some parts of the brain, and on the relations between the brain and nerves of motion and sensation’, Philosophical Transactions of the Royal Society, 124 (1834), pp. 471–83; idem, ‘Continuation of the paper on the relations between the nerves of motion and of sensation, and the brain; more particularly on the structure of the medulla oblongata and the spinal marrow’, Philosophical Transactions of the Royal Society, 125 (1835), pp. 255–62.

72 72 F. Magendie, ‘Expériences sur les fonctions des racines des nerfs rachidiens’, Journal Physiologie expérimentale ct de pathologie, 3 (1822), pp. 276–9; repr. with trans. in Walker, Documents and Dates, pp. 88, 95.

73 73 Ibid., p. 91.

74 74 M. Hall, ‘On the reflex function of the medulla oblongata and medulla spinalis’, Philosophical Transactions of the Royal Society, 123 (1833), pp. 635–65; idem, ‘These motions independent of sensation and volition’, Proceedings of the Committee of Science, Zoological Society, 27 Nov. 1832, repr. in Walker, Documents and Dates, p. 138.

75 75 M. Hall, ‘Synopsis of the diastaltic nervous system or the system of the spinal marrow and its reflex arcs, as the nervous agent in all the functions of ingestion and of egestion in the animal economy’, Croonian Lectures (Mallett, London, 1850).

76 76 M. Foster, A Textbook of Physiology (Macmillan, London, 1890), p. 912. The Cartesian roots of this conception of the spinal soul are here evident in its association with consciousness.

77 77 P. Broca, ‘Remarques sur le siège de la faculté du language articulé, suivies d’une observation d’aphémie (perte de la parole)’, Bulletins de la Société Anatomique (Paris), 6 (1861), pp. 330–57, 398–407; (tr. as ‘Remarks on the seat of the faculty of articulate language, followed by an observation of aphemia’, in G. von Bonin, Some Papers on the Cerebral Cortex (Charles C. Thomas, Springfield, IL, 1960), pp. 49–72.

78 78 M. J. P. Flourens, Recherches expérimentales sur les propriétés et les fonctions du système nerveux dans les animaux vertébrés (Ballière, Paris, 1823).

79 79 G. Fritsch and E. Hitzig, ‘Über die elektrische Erregbarkeit des Grosshirns’, Archiv für Anatomie, Physiologie and wissenschaftliche Medicin, Leipzig, 37 (1870), pp. 300–32; tr. as ‘On the electrical excitability of the cerebrum’ in von Bonin, Some Papers on the Cerebral Cortex, pp. 73–96.

80 80 J. H. Jackson, ‘Convulsive spasms of the right hand and arm preceding epileptic seizures’, Medical Times and Gazette, 2 (1863), pp. 110–11.

81 81 D. Ferrier, ‘The localization of function in the brain’, Proceedings of the Royal Society, 22 (1873–4),pp. 228–32; idem, ‘Experiments on the brain of monkeys’, Croonian Lecture (2nd ser.), Philosophical Transactions of the Royal Society, 165 (1876), pp. 433–88; idem, The Function of the Brain (Smith Elder and Company, London, 1876).

82 82 C. E. Beevor and V. Horsley, ‘A minute analysis (experimental) of the various movements produced by stimulating in the monkey different regions of the cortical centre for the upper limb, as defined by Professor Ferrier’, Philosophical Transactions of the Royal Society, 178 (1887), pp. 153–67; idem, ‘A further minute analysis by electrical stimulation of the so called motor regions (facial area) of the cortex cerebri in the monkey (Macacus sinicus)’, Philosophical Transactions of the Royal Society, 185 (1894), pp. 39–81; idem, ‘A record of the results obtained by electrical excitation of the so-called motor cortex and internal capsule in an orang-outang (Simia satyrus)’, Philosophical Transactions of the Royal Society, 181 (1890), pp. 129–58.

83 83 R. Caton, ‘The electrical currents of the brain’, British Medical Journal, 2 (1875), p. 278; idem, ‘Interim report on investigation of the electric currents of the brain’, British Medical Journal, 1 (1877), Suppl. L, pp. 62–5; idem, ‘Researches on electrical phenomena of cerebral grey matter’, Transactions of the Ninth Intemational Medical Congress, 3 (1887), pp. 246–9.

84 84 A. Beck, ‘Die Bestimmung der Localisation der Gehirn- und Rückenmarkfunktionen vermittelst der elektrischen Erscheinungen’, Centralblatt für Physiologie, 4 (1890), pp. 473–6.

85 85 C. S. Sherrington, ‘Notes on the arrangement of some motor fibres in the lumbo-sacral plexus’, Journal of Physiology, 13 (1892), pp. 621–772.

86 86 C. S. Sherrington, ‘On reciprocal innervation of antagonistic muscles: Seventh note’, Proceedings of the Royal Society, B 76 (1905), pp. 160–3; idem, ‘On reciprocal innervation of antagonistic muscles: Eighth note’, Proceedings on the Royal Society, B 76 (1905), pp. 269–97.

87 87 C. S. Sherrington, ‘Flexion-reflex of the limb, crossed extension-reflex, and reflex stepping and standing’, Journal of Physiology, 40 (1910), pp. 28–121.

88 88 A. S. F. Grünbaum and C. S. Sherrington, ‘Observations on the physiology of the cerebral cortex of some of the higher apes (preliminary communication)’, Proceedings of the Royal Sociey, 69 (1902), pp. 206–9.

89 89 Ibid.

90 90 Ibid.

91 91 C. S. Sherrington and C. S. Roy, ‘On the regulation of the blood-supply of the brain’, Journal of Physiology, 11 (1890), p. 106.

92 92 Ibid., p. 105.

93 93 S. Ogawa, T. M. Lee, A. S. Nayak and P. Glynn, ‘Oxygen-sensitive contrast in magnetic resonance image of rodent brain at high magnetic fields’, Magnetic Resonance Medicine, 14, no. 1 (1990), pp. 68–78.

94 94 S. Ogawa, T. M. Lee, A. R. Ray and D. W. Tank, ‘Brain magnetic resonance imaging with contrast dependent on blood oxygenation’, Proceedings of the National Academy of Sciences of the United States of America, 87 (1990), pp. 9868–72.

95 95 S. Ogawa, D. W. Tank, R. Menon, J. M. Ellermann, S-G Kim, H. Merkle and K. Ugurbil, ‘Intrinsic signal changes accompanying sensory stimulation: Functional brain mapping with magnetic resonance imaging’, Proceedings of the National Academy of Sciences of the United States of America, 89 (1992), pp. 5951–5; see also K. K. Kwong, J. W. Belliveau, D. A. Chesler et al., ‘Dynamic magnetic resonance imaging of human brain activity during primary sensory stimulation’, Proceedings of the National Academy of Sciences of the United States of America, 89 (1992), pp. 5675–9.

96 96 B. Biswal, F. Z. Yetkin, V. M. Haughton and J. S. Hyde, ‘Functional connectivity in the motor cortex of resting human brain using echo-planar MRI’, Magnetic Resonance Medicine, 34, no. 4 (1995), pp. 537–41.

97 97 M. E. Raichle, A. M. Macleod, A. Z. Snyder, W. J. Powers, D. A. Gusnard and G. L. Shulman, ‘A default mode of brain function’, Proceedings of the National Academy of Sciences of the United States of America, 98 (2001), pp. 676–82.

98 98 M. D. Grecius, V. Kiviniemi, O. Tervonen, V. Vainionpaa, S. Alahuhta, A. L. Reiss and V. Menon, ‘Persistent default-mode network connectivity during light sedation’, Human Brain Mapping, 29, no. 7 (2008), pp. 839–47.

99 99 C. Kennedy, M. H. Des Rosierd, O. Sakurada, M. Reivich, J. W. Jehle and L. Sokoloff, ‘Metabolic mapping of the primary visual system of the monkey by means of the autoradiographic [14C]deoxyglucose technique’, Proceedings of the National Academy of Sciences of the United States of America, 73, no. 11 (1976), pp. 4230–4.

100 100 M. Reivich, D. Kuhl, A. Wolf, J. Greenberg, M. Phelps, T. Ido, V. Casella, J. Fowler, E. Hoffman, A. Alavi, P. Som and L. Sokoloff, ‘The [18F]fluorodeoxyglucose method for the measurement of local cerebral glucose utilization in man’, Circulation Research 44, no. 1 (1979), pp. 127–37,; see also M. E. Phelps, S. C. Huang, E. J. Hoffman, C. Selin and D. E. Kuhl, ‘Tomographic measurement of local cerebral glucose metabolic rate in humans (F-18)2-fluoro-2-deoxy-D-glucose: validation of method’, Annals of Neurology 6, no. 5 (1979), pp. 371–86.

101 101 L. Elliott, A. R. Knodt, D. Ireland, M. L. Morris, R. Poulton, S. Ramrakha, M. L. Sison, T. E. Moffitt, A. Caspi and R. Ahmad, ‘Hariri: Poor test-retest reliability of task-fMRI: new empirical evidence and a meta-analysis’, bioRxiv (2019 June). doi: https://doi.org/10.1101/681700; S. Noble, D. Scheinost and R. T. Constable, ‘A decade of test-retest reliability of functional connectivity: A systematic review and meta-analysis’, Neuroimage, 203 (2019), p. 116157.

102 102 F. Cauda, T. Costa, M. Diano, K. Sacco, S. Duca, G. Geminiani and D. M. Torta, ‘Massive modulation of brain areas after mechanical pain stimulation: a time-resolved FMRI study’, Cerebral Cortex, 24, no. 11 (2014), pp. 2991–3005; J. Gonzalez-Castillo, Z. S. Saad, D. A. Handwerker, S. J. Inati, N. Brenowitz and P. A. Bandettini, ‘Whole-brain, time-locked activation with simple tasks revealed using massive averaging and model-free analysis’, Proceedings of the National Academy of Sciences of the United States of America, 109, no. 14 (2012), pp. 5487–92; J. W. Ibinson and K. M. Vogt, ‘Pain does not follow the boxcar model: temporal dynamics of the BOLD fMRI signal during constant current painful electric nerve stimulation’, Journal of Pain, 14, no. 12 (2013), pp. 1611–19.

103 103 M. R. Bennett, L. Farnell and W. G. Gibson, ‘Quantitative relations between transient BOLD responses, cortical energetics, and impulse firing in different cortical regions’, Journal of Neurophysiology, 122, no. 3 (2019), pp. 1226–37.

104 104 N. J. Maandag, D. Coman, B. G. Sanganahalli, P. Herman, A. J. Smith, H. Blumenfeld, R. G. Shulman and F. Hyder, ‘Energetics of neuronal signaling and fMRI activity’, Proceedings of the National Academy of Sciences of the United States of America, 104, no. 51 (2007), pp. 20546–51; A. J. Smith, H. Blumenfeld, K. L. Behar, D. L. Rothman, R. G. Shulman and F. Hyder, ‘Cerebral energetics and spiking frequency: the neurophysiological basis of fMRI’, Proceedings of the National Academy of Sciences of the United States of America, 99, no. 16 (2002), pp. 10765–70.

105 105 M. R. Bennett, L. Farnell and W. G. Gibson, ‘Quantitative relations between BOLD responses, cortical energetics, and impulse firing’, Journal of Neurophysiology, 119, no. 3 (2018), pp. 979–89.

106 106 F. Hyder, R. K. Fulbright, R. G. Shulman and D. L. Rothman, ‘Glutamatergic function in the resting awake human brain is supported by uniformly high oxidative energy’, Journal of Cerebral Blood Flow & Metabolism, 33, no. 3 (2013), pp. 339–47; F. Hyder, D. L. Rothman and M. R. Bennett, ‘Cortical energy demands of signaling and non-signaling components in brain are conserved across mammalian species and activity levels’, Proceedings of the National Academy of Sciences of the United States of America, 110, no. 9 (2013), pp. 3549–54.

107 107 M. T. Alkire, ‘Loss of effective connectivity during general anesthesia’, International Anesthesiology Clinics, 46, no. 3 (2008), pp. 55–73; idem, ‘Probing the mind: anesthesia and neuroimaging’, Clinical Pharmacology & Therapeutics, 84, no. 1 (2008), pp. 149–52; Hyder, Fulbright, Shulman and Rothman, ‘Glutamatergic function in the resting awake human brain is supported by uniformly high oxidative energy’.

108 108 R. M. Birn, J. B. Diamond, M. A. Smith and P. A. Bandettini, ‘Separating respiratory-variation-related fluctuations from neuronal-activity-related fluctuations in fMRI’, Neuroimage, 31, no. 4 (2006), pp. 1536–48; R. G. Wise, K. Ide, M. J. Poulin and I. Tracey, ‘Resting fluctuations in arterial carbon dioxide induce significant low frequency variations in BOLD signal’, Neuroimage, 21, no. 4 (2004), pp. 1652–64.

109 109 G. K. Aguirre, E. Zarahn and M. D’Esposito, ‘The inferential impact of global signal covariates in functional neuroimaging analyses’, Neuroimage, 8, no. 3 (1998), pp. 302–6; P. M. Macey, K. E. Macey, R. Kumar and R. M. Harper, ‘A method for removal of global effects from fMRI time series’, Neuroimage, 22, no. 1 (2004), pp. 360–6.

110 110 K. Murphy, R. M. Birn, D. A. Handwerker, T. B. Jones and P. A. Bandettini, ‘The impact of global signal regression on resting state correlations: are anti-correlated networks introduced?’ Neuroimage, 44, no. 3 (2009), pp. 893–905;M. D. Fox, D. Zhang, A. Z. Snyder and M. E. Raichle, ‘The global signal and observed anticorrelated resting state brain networks’, Journal of Neurophysiology, 101, no. 6 (2009), pp. 3270–83.

111 111 K. Murphy and M. D. Fox, ‘Towards a consensus regarding global signal regression for resting state functional connectivity MRI’, Neuroimage, 154 (2017), pp. 169–73.

112 112 M. L. Schölvinck, A Maier, F. Q. Ye, J. H. Duyn and D. A. Leopold, ‘Neural basis of global resting-state fMRI activity’, Proceedings of the National Academy of Sciences of the United States of America, 107, no. 22 (2010), pp. 10238–43; C. W. Wong, P. N. DeYoung and T. T. Liu, ‘Differences in the resting-state fMRI global signal amplitude between the eyes open and eyes closed states are related to changes in EEG vigilance’, Neuroimage, 124, Pt A (2016), pp. 24–31; C. W. Wong, V. Olafsson, O. Tal and T. T. Liu, ‘The amplitude of the resting-state fMRI global signal is related to EEG vigilance measures’, Neuroimage, 83 (2013), pp. 983–90; C. W. Wong, V. Olafsson, O. Tal and T. T. Liu, ‘Anti-correlated networks, global signal regression, and the effects of caffeine in resting-state functional MRI’, Neuroimage, 63, no. 1 (2012), pp. 356–64.

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