Abstract
Abstract. Reading is an example of complex learning specific to human beings. In readers, an area of the brain is dedicated to the visual processing of letters and words, referred to as the visual word form area (VWFA). The existence of this brain area is paradoxical. Reading is too recent to be a phylogenic product of Darwinian evolution. It likely develops with intense school training via a neuroplastic ontogenic process of neuronal recycling: neurons in the lateral occipitotemporal lobe originally tuned to the visual recognition of stimuli, such as faces, objects, and animals, will be recycled for the visual recognition of letters and words. Thus, the VWFA inherits the intrinsic properties of these neurons, notably, mirror generalization, a process (or heuristic) applied to all visual stimuli that enables the recognition of a stimulus irrespective of its left-right orientation. On its own, this inherited property is not adapted to reading because it makes children confuse mirror letters, such as b and d in the Latin alphabet. In this article, we present evidence that inhibitory control is critical to avoid mirror errors inherited from the neuronal recycling process by blocking the mirror generalization heuristic in the context of reading. We subsequently argue that the “neuronal recycling + inhibitory control” law constitutes a general law of the learning brain by demonstrating that it may also account for the development of numeracy.
References
2016). Inhibition of the mirror generalization process in reading in school-aged children. Journal of Experimental Child Psychology, 145, 1–9. doi: 10.1016/j.jecp.2015.12.009
(2012). Culture and education: new frontiers in brain plasticity. Trends in Cognitive Sciences, 16, 93–95. doi: 10.1016/j.tics.2011.11.016
(2004). Inhibition and the right inferior frontal cortex. Trends in Cognitive Sciences, 8, 170–177. doi: 10.1016/j.tics.2004.02.010
(1971). Discrimination of spatially confusable letters by young children. Journal of Experimental Child Psychology, 11, 11–20. doi: 10.1016/0022-0965(71)90059-2
(2012). Hierarchical processing of face viewpoint in human visual cortex. The Journal of Neuroscience, 32, 2442–2452. doi: 10.1523/JNEUROSCI.4770-11.2012
(2007). Visual word processing and experiential origins of functional selectivity in human extrastriate cortex. Proceedings of the National Academy of Sciences of the United States of America, 104, 9087–9092. doi: 10.1073/pnas.0703300104
(2001). Shape-coding in IT cells generalizes over contrast and mirror reversal, but not figure-ground reversal. Nature Neuroscience, 4, 937–942. doi: 10.1038/nn0901-937
(1987). Recognition-by-components: A theory of human image understanding. Psychological Review, 94, 115–147. doi: 10.1037/0033-295x.94.2.115
(2005). Cross-cultural effect on the brain revisited: Universal structures plus writing system variation. Human Brain Mapping, 25, 92–104. doi: 10.1002/hbm.20124
(1978). Perceptual similarity of mirror images in infancy. Cognition, 6, 89–116. doi: 10.1016/0010-0277(78)90017-3
(2015). The cost of blocking the mirror generalization process in reading: Evidence for the role of inhibitory control in discriminating letters with lateral mirror-image counterparts. Psychonomic Bulletin & Review, 22, 228–234. doi: 10.3758/s13423-014-0663-9
(2016). Early cerebral constraints on reading skills in school-age children: An MRI study. Mind, Brain, and Education, 10, 47–54. doi: 10.1111/mbe.12098
(2013).
(Negative priming in logicomathematical reasoning: The cost of blocking your intuition . In W. De NeysM. OsmanEds., New approaches in reasoning research (pp. 34–50). New York, NY: Psychology Press. doi: 10.4324/97813158798572013). Inhibitory control efficiency in a Piaget-like class-inclusion task in school-age children and adults: A developmental negative priming study. Developmental Psychology, 49, 1366–1374. doi: 10.1037/a0029622
(2005). Character complexity and redundancy in writing systems over human history. Proceedings of the Royal Society B: Biological Sciences, 272, 267–275. doi: 10.1098/rspb.2004.2942
(2006). The structures of letters and symbols throughout human history are selected to match those found in objects in natural scenes. The American Naturalist, 167, E117–E139. doi: 10.1086/502806
(2000). The visual word form area: Spatial and temporal characterization of an initial stage of reading in normal subjects and posterior split-brain patients. Brain : A Journal of Neurology, 123, 291–307. doi: 10.1093/brain/123.2.291
(1985). Spontaneous mirror-writing in children. Canadian Journal of Psychology/Revue Canadienne de Psychologie, 39, 174–179. doi: 10.1037/h0080122
(2009). Mirror writing in pre-school children: A pilot study. Cognitive Processing, 10, 101–104. doi: 10.1007/s10339-008-0233-z
(1935). A study of the confusing letters b, d, p, and q. Pedagogical Seminary and Journal of Genetic Psychology, 42, 458–468. doi: 10.1080/08856559.1935.10534056
(2004).
(Evolution of human cortical circuits for reading and arithmetic: The “neuronal recycling” hypothesis . In S. DehaeneJ. DuhamelM. HauserG. RizzolattiEds., From monkey brain to human brain Cambridge, MA: MIT Press.2013). Inside the letterbox: How literacy transforms the human brain. Cerebrum, 7, 1–16.
(2011). The unique role of the visual word form area in reading. Trends in Cognitive Sciences, 15, 254–262. doi: 10.1016/j.tics.2011.04.003
(2015). Illiterate to literate: behavioural and cerebral changes induced by reading acquisition. Nature Reviews Neuroscience, 16, 234–244. doi: 10.1038/nrn3924
(2005). The neural code for written words: a proposal. Trends in Cognitive Sciences, 9, 335–341. doi: 10.1016/j.tics.2005.05.004
(2002). The visual word form area: a prelexical representation of visual words in the fusiform gyrus. Neuroreport, 13, 321–325. doi: 10.1097/00001756-200203040-00015
(2010). Why do children make mirror errors in reading? Neural correlates of mirror invariance in the visual word form area. NeuroImage, 49, 1837–1848. doi: 10.1016/j.neuroimage.2009.09.024
(2010). How learning to read changes the cortical networks for vision and language. Science, 330, 1359–1364. doi: 10.1126/science.1194140
(2007). Preschool program improves cognitive control. Science, 318, 1387–1388. doi: 10.1126/science.1151148
(1989). Comparison of human infants and rhesus monkeys on Piaget’s AB task: Evidence for dependence on dorsolateral prefrontal cortex. Experimental Brain Research, 74, 24–40. doi: 10.1007/BF00248277
(2011). Mirror-image sensitivity and invariance in object and scene processing pathways. The Journal of Neuroscience, 31, 11305–11312. doi: 10.1523/JNEUROSCI.1935-11.2011
(2009). Common and specific contributions of the intraparietal sulci to numerosity and length processing. Human Brain Mapping, 30, 2466–2476. doi: 10.1002/hbm.20677
(2011). Through the looking-glass: Mirror reading. NeuroImage, 54, 3004–3009. doi: 10.1016/j.neuroimage.2010.10.079
(2013). The joint development of hemispheric lateralization for words and faces. Journal of Experimental Psychology. General, 142, 348–358. doi: 10.1037/a0029503
(2012). Unraveling the mystery of mirror writing in typically developing children. Journal of Educational Psychology, 104, 193–205. doi: 10.1037/a0025735
(2010). Functional compartmentalization and viewpoint generalization within the macaque face-processing system. Science, 330, 845–851. doi: 10.1126/science.1194908.Functional
(1991). Exaptation: A crucial tool for an evolutionary psychology. Journal of Social Issues, 47, 43–65. doi: 10.1111/j.1540-4560.1991.tb01822.x
(1982). Exaptation-A missing term in the science of form. Paleobiology, 8, 4–15. doi: 10.2307/2400563
(1992).
(Tools – The Paleolithic record . In S. JonesR. MartinD. PilbeamEds., The Cambridge encyclopedia of human evolution (pp. 350–360). New York, NY: Cambridge University Press.1978). Left and right in science and art. Leonardo, 11, 29–38. doi: 10.2307/1573500
(2015). The direct segment of the arcuate fasciculus is predictive of longitudinal reading change. Developmental Cognitive Neuroscience, 13, 68–74. doi: 10.1016/j.dcn.2015.05.002
(2003). Large-scale mirror-symmetry organization of human occipito-temporal object areas. Neuron, 37, 1027–1041. doi: 10.1016/s0896-6273(03)00144-2
(1934). Reversals in reading and writing. Journal of Educational Psychology, 25, 1–20. doi: 10.1037/h0074907
(2000). Inhibition and cognitive development: Object, number, categorization, and reasoning. Cognitive Development, 15(1), 63–73. doi: 10.1016/s0885-2014(00)00015-0
(2014). Measuring inhibitory control in children and adults: Brain imaging and mental chronometry. Frontiers in Psychology, 5, 1–7. doi: 10.3389/fpsyg.2014.00616
(2001). Negative priming effect after inhibition of number/length interference in a Piaget-like task. Developmental Science, 4, 119–123. doi: 10.1111/1467-7687.00156
(2011). Functional magnetic resonance imaging study of Piaget’s conservation-of-number task in preschool and school-age children: A neo-Piagetian approach. Journal of Experimental Child Psychology, 110, 332–346. doi: 10.1016/j.jecp.2011.04.008
(2010). Mapping numerical processing, reading, and executive functions in the developing brain: An fMRI meta-analysis of 52 studies including 842 children. Developmental Science, 13, 876–885. doi: 10.1111/j.1467-7687.2009.00938.x
(1977). Evolution and tinkering. Science, 196, 1161–1166. doi: 10.1126/science.860134
(2013). Automatic inhibition and habitual control: Alternative views in neuroscience research on response inhibition and inhibitory control. Frontiers in Behavioral Neuroscience, 7(April), 1–4. doi: 10.3389/fnbeh.2013.00025
(2003). Evaluation of the dual route theory of reading: A metanalysis of 35 neuroimaging studies. NeuroImage, 20, 693–712. doi: 10.1016/S1053-8119(03)00343-4
(2012). Prevalence of selectivity for mirror-symmetric views of faces in the ventral and dorsal visual pathways. The Journal of Neuroscience, 32, 11763–11772. doi: 10.1523/JNEUROSCI.0126-12.2012
(2014). Searching for the principles of brain plasticity and behavior. Cortex, 58, 1–10. doi: 10.1016/j.cortex.2013.11.012
(2002).
(Reading disability as a deficit in functional coordination . In E. WitrukA. D. FriedericiT. LachmannEds., Basic functions of language, reading and reading disability (Vol. 20, pp. 165–198). Boston, MA: Springer. doi: 10.1007/978-1-4615-1011-6_112003). Letter reversals in dyslexia: Is the case really closed? A critical review and conclusions. Psychology Science, 45, 50–72. doi: 10.1016/s0010-9452(71)80009-6
(2016). Inhibitory control is needed to overcome written verb inflection errors: Evidence from a developmental negative priming study. Cognitive Development, 37, 18–27. doi: 10.1016/j.cogdev.2015.10.005
(2012). fMRI repetition suppression: Neuronal adaptation or stimulus expectation? Cerebral Cortex, 22, 567–576. doi: 10.1093/cercor/bhr119
(2006). Age differences in memory control: Evidence from updating and retrieval-practice tasks. Acta Psychologica, 123, 279–298. doi: 10.1016/j.actpsy.2006.01.006
(2011). Old dogs learning new tricks: Neuroplasticity beyond the juvenile period. Developmental Review: DR, 31, 207–239. doi: 10.1016/j.dr.2011.07.008
(2013). Inhibitory control is needed for the resolution of arithmetic word problems: A developmental negative priming study. Journal of Educational Psychology, 105, 701–708. doi: 10.1037/a0032625
(2015). Reading in the brain of children and adults: A meta-analysis of 40 functional magnetic resonance imaging studies. Human Brain Mapping, 36, 1963–1981. doi: 10.1002/hbm.22749
(1981). An interactive activation model of context effects in letter perception: I. An account of basic findings. Psychological Review, 88, 375–407. doi: 10.1037/0033-295X.88.5.375
(1986). The archaic texts from Uruk. World Archaeology, 17, 317–334. doi: 10.1080/00438243.1986.9979973
(1925). “Word-blindness” in school children. Archives of Neurology and Psychiatry, 14, 581–615. doi: 10.1001/archneurpsyc.1925.02200170002001
(2014). Timing the impact of literacy on visual processing. Proceedings of the National Academy of Sciences of the United States of America, 111(49), E5233–E5242. doi: 10.1073/pnas.1417347111
(2011). Breaking the symmetry: Mirror discrimination for single letters but not for pictures in the Visual Word Form Area. NeuroImage, 55, 742–749. doi: 10.1016/j.neuroimage.2010.11.043
(1988). Positron emission tomographic studies of the cortical anatomy of single-word processing. Nature, 331, 585–589. doi: 10.1038/331585a0
(1952). The child’s conception of number. London, UK: Routledge & Kegan Paul.
(1983). Piaget’s theory. Handbook of child psychology (pp. 103–128). New York, NY: Wiley.
(2012). Number conservation is related to children’s prefrontal inhibitory control: An fMRI study of a piagetian task. PLoS One, 7, e40802. doi: 10.1371/journal.pone.0040802
(1992). Lexical access and the brain: Anatomical constraints on cognitive models of word recognition. The American Journal of Psychology, 105, 1–26. doi: 10.2307/1422979
(2011). The interactive account of ventral occipitotemporal contributions to reading. Trends in Cognitive Sciences, 15, 246–253. doi: 10.1016/j.tics.2011.04.001
(2000). Mirror-image confusion in single neurons of the macaque inferotemporal cortex. Science, 287, 1506–1508. doi: 10.1126/science.287.5457.1506
(2006). Neural representation of language: Activation versus long-range connectivity. Trends in Cognitive Sciences, 10, 519–525. doi: 10.1016/j.tics.2006.09.007
(1971). Discrimination of orientation by Zambian children. Journal of Comparative and Physiological Psychology, 75, 312–316. doi: 10.1037/h0030832
(1960). Visual discrimination of orientation by octopus: Mirror images. British Journal of Psychology, 51, 9–18. doi: 10.1016/0003-3472(65)90109-0
(1985). The negative priming effect: Inhibitory priming by ignored objects. The Quarterly Journal of Experimental Psychology, 37, 571–590. doi: 10.1080/14640748508400920
(1983). The discrimination of mirror-image forms by pigeons. Perception & Psychophysics, 34, 397–402. doi: 10.3758/BF03203053
(2015). Auditory processing in noise: A preschool biomarker for literacy. PLoS Biology, 13, 1–17. doi: 10.1371/journal.pbio.1002196
(2012). Emergence of the neural network for reading in five-year old beginning readers of different levels of pre-literacy abilities: An fMRI study. NeuroImage, 57, 704–713. doi: 10.1016/j.neuroimage.2010.10.057.Emergence
(2004). Face perception: Domain specific, not process specific. Neuron, 44, 889–898. doi: 10.3923/itj.2008.105.111
(