Abstract
Language has come to be commonly understood as something accomplished by the brain. Much scientific investigation has consequently focused in looking for language in the brain. Although it may sound intuitive, this approach suggests that language is an object located inside another object. This spatial metaphor has generated important insights into the brain sites important for language, from nineteenth century studies of brain-damaged patients to more recent and refined evidence from brain imaging.
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- 1.
Chomsky’s forays into dualistic rationalism were blended with and at least partly motivated by his political views, which eclectically (and bizarrely) equated empiricism with US imperialism of the Vietnam era and rationalism with progressive enlightenment (Chomsky 1972b).
- 2.
Interestingly, this scientific debate had political and ideological undertones in mid-nineteenth century France of the second empire, as holistic views were considered a conservative and theologically warranted reflection of the unity of the soul, whereas localizationism was associated with antimonarchic and anticlerical left-wing views at the time (Harrington 1987).
- 3.
Modern neurology has provided evidence in support of diaschisis. For example, in patients with damage in cerebral cortex, reduced glucose metabolism is often seen far away from the site of damage, in the opposite hemisphere of the cerebellum. This phenomenon, called ‘crossed cerebellar diaschisis’, is attributed to contralateral (“crossed”) connections between forebrain and cerebellum (Gold and Lauritzen 2002; Pantano et al. 1986).
- 4.
In more technical verbatim terms: Broca’s area “handles only intrasentential dependency relations” (Grodzinsky 2000: 21) and “is the neural home to receptive mechanisms involved in the computation of the relation between transformationally moved phrasal constituents and their extraction sites...” (ibid.: 1).
- 5.
Corresponding issues have been debated in artificial intelligence for decades. Computational models informed by cognitive theories may approximate the behavior of a human being, but their internal organization in no way resembles functional brain organization. The cognitive model-driven approach in AI (e.g., Newell and Simon 1963) was countered in the 1980s by parallel-distributed (“connectionist”) models (McClelland and Rumelhart 1986). These were inspired ‘bottom-up’ by simplified features of neurons and their connections and aspired to demonstrate complex input-output behaviors mimicking human cognition as emergent properties of such very simple neural networks.
- 6.
Since conventional fMRI statistics test for activation effects separately in each voxel, the large number of voxels creates a need to correct for multiple comparisons. For example, a given voxel shows an effect that is significant at the 95 % confidence level (i.e., the probability of obtaining the finding by chance alone is \(\le 5\,\%\)). If comparisons are performed in 10 voxels, the probability of obtaining the finding by chance increases to 50 %. A correction for multiple comparisons is therefore necessary. In its simplest form (the “Bonferroni correction”), the p-value is multiplied by the number of comparisons. Such a correction would require that an effect in a single voxel reaches a p-value of 0.0000005 in order to “survive” a Bonferroni correction for 100,000 comparisons (assuming that an fMRI study includes this number of brain voxels).
- 7.
In technical terms, the region of interest approach drastically reduces the need to correct for multiple comparisons, as only a small part of the brain is considered, thus increasing the probability of a statistically significant finding.
- 8.
Among these are statistical power (number of participants), analysis type (whole-brain vs. region of interest), signal to noise ratio (reflecting head motion and other sources of noise), and the investigator’s selection of significance thresholds and method of multiple comparison correction.
- 9.
Participants had to judge whether sentences were the same or different. For some sentences, this required syntactic processing (e.g., “The policeman arrested the thief” is the same as “The thief was arrested by the policeman”), whereas in other sentence pairs it required semantic processing (“East of the city is the lake” is different from “East of the city is the river”).
- 10.
Function words (e.g., “if” or “by”) are primarily grammatical with relatively little meaning, whereas content words (e.g., nouns such as “house” or “love”) are meaningful (semantically rich).
- 11.
Garden-path sentences have misleading syntactic structure. A classic example is the sentence “The horse raced past the barn fell”, where the final word prompts reanalysis of the initially expected syntactic structure (“The horse raced past the barn”).
- 12.
Churchland’s eliminative materialism claims that psychology will be eventually replaced by neuroscience (1986).
- 13.
The grammatical structure of sentences is not fully determined by word order, but also by insertion of small and relatively meaningless grammatical words (such as prepositions), changes in word endings etc. This is reflected in the term “morphosyntax”. For example, interpretation of the sentence “The Rottweiler was bitten by the mailman” is not possible based on word order alone, but requires decoding of morphological cues (passive voice indicated by the auxiliary verb “be” and the ending of the verb “bite”, which makes it a participle) and the preposition “by” indicating that “mailman” is the agent of the sentence, i.e. the one who (unexpectedly) does the biting.
- 14.
A minicolumn includes about 100 neurons with strong excitatory interconnections. These neurons are vertically arranged, giving the impression of a ‘column’. Minicolumns are considered basic functional units of mammalian cortex (Mountcastle 1997).
- 15.
For an example, consider the length of the vowel /i/ in the word “fit”, which is around 60 ms. With length of > 100 ms, a phonemic boundary will be crossed and the vowel will be perceived as a long /i:/, resulting in the word “feet”. On the other hand, a change in vowel length from 100 to 150 ms will not be perceived as a different phoneme (and the perceived word will remain “feet”). Whereas acoustic length of the vowel can thus vary in graded fashion, some relatively small changes in vowel length around the category boundary will be perceived as distinct phonemes that can change the meaning of a word.
- 16.
The mismatch negativity (MMN) is an EEG response that follows about 150–250 ms after a deviant stimulus (e.g., a high tone among a series of low tones, or an omitted click in an otherwise regular sequence of clicks; Cheour et al. 2000). It can be reliably measured even in infants and therefore provides a technique for demonstrating perceptual discrimination at an age when verbal responses cannot be obtained. If an infant shows an MMN to stimulus A in a series of presentations of stimulus B, one can infer that the infant brain is able to distinguish A from B.
- 17.
Greenough et al. (1987, p. 540) define experience-expectant plasticity as “designed to utilize the sort of environmental information that is ubiquitous and has been so throughout much of the evolutionary history of a species.” An example is the development of ocular dominance columns, which are small vertically organized units in visual cortex that respond to information coming from one eye only.
- 18.
For a more comprehensive review of the literature on developmental change in language networks see Müller and Palmer (2008).
- 19.
Meyer and colleagues (2000) instructed some of their participants to repair sentence violations. For example, when presented with the sentence “The spy was in the caught”, participants would not only indicate that the sentence was ungrammatical, but also covertly fix the violation, generating “The spy was caught” in their minds (without speaking). Activation for this repair condition was particularly strong in right IFG and STG.
- 20.
Unlike English, German morphosyntax (cf. Footnote 13) relies heavily on grammatical gender since every noun has one of three genders (feminine, masculine, neuter) and articles and pronouns change in agreement with the gender of the corresponding noun. For example, in “Die Tüte riss und ihr Inhalt fiel heraus” (“The bag tore and its contents fell down”), the article “die” and the pronoun “ihr” have the feminine form, in agreement with the grammatical gender of “Tüte”.
- 21.
Donald Hebb (1949) hypothesized that a synapse is strengthened as a result of synchronous firing of the two neurons the synapse connects. This idea, popularized in the phrase “What fires together wires together”, has proven basically accurate.
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Preparation of this review was supported by the National Institutes of Health, grants R01-NS43999, R01-DC006155 and R01-MH081023. Thanks to Patricia Shih for help in editing the manuscript.
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Müller, RA. (2013). Neurobiology: Language By, In, Through and Across the Brain. In: Binder, PM., Smith, K. (eds) The Language Phenomenon. The Frontiers Collection. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3-642-36086-2_2
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