Contribution of the language network to the comprehension of Python programming code.

Does the perisylvian language network contribute to comprehension of programming languages, like Python? Univariate neuroimaging studies find high responses to code in fronto-parietal executive areas but not in fronto-temporal language areas, suggesting the language network does little. We used multivariate-pattern-analysis to test whether the language network encodes Python functions. Python programmers read functions while undergoing fMRI. A linear SVM decoded for-loops from if-conditionals based on activity in lateral temporal (LT) language cortex. In searchlight analysis, decoding accuracy was higher in LT language cortex than anywhere else. Follow up analysis showed that decoding was not driven by presence of different words across functions, "for" vs "if," but by compositional program properties. Finally, univariate responses to code peaked earlier in LT language-cortex than in the fronto-parietal network. We propose that the language system forms initial "surface meaning" representations of programs, which input to the reasoning network for processing of algorithms.

Developing cortex is functionally pluripotent: Evidence from blindness.

How rigidly does innate architecture constrain function of developing cortex? What is the contribution of early experience? We review insights into these questions from visual cortex function in people born blind. In blindness, occipital cortices are active during auditory and tactile tasks. What 'cross-modal' plasticity tells us about cortical flexibility is debated. On the one hand, visual networks of blind people respond to higher cognitive information, such as sentence grammar, suggesting drastic repurposing. On the other, in line with 'metamodal' accounts, sighted and blind populations show shared domain preferences in ventral occipito-temporal cortex (vOTC), suggesting visual areas switch input modality but perform the same or similar perceptual functions (e.g., face recognition) in blindness. Here we bring these disparate literatures together, reviewing and synthesizing evidence that speaks to whether visual cortices have similar or different functions in blind and sighted people. Together, the evidence suggests that in blindness, visual cortices are incorporated into higher-cognitive (e.g., fronto-parietal) networks, which are a major source long-range input to the visual system. We propose the connectivity-constrained experience-dependent account. Functional development is constrained by innate anatomical connectivity, experience and behavioral needs. Infant cortex is pluripotent, the same anatomical constraints develop into different functional outcomes.

Neural specialization for 'visual' concepts emerges in the absence of vision.

Vision provides a key source of information about many concepts, including 'living things' (e.g., tiger) and visual events (e.g., sparkle). According to a prominent theoretical framework, neural specialization for different conceptual categories is shaped by sensory features, e.g., living things are neurally dissociable from navigable places because living things concepts depend more on visual features. We tested this framework by comparing the neural basis of 'visual' concepts across sighted (n=22) and congenitally blind (n=21) adults. Participants judged the similarity of words varying in their reliance on vision while undergoing fMRI. We compared neural responses to living things nouns (birds, mammals) and place nouns (natural, manmade). In addition, we compared visual event verbs (e.g., 'sparkle') to non-visual events (sound emission, hand motion, mouth motion). People born blind exhibited distinctive univariate and multivariate responses to living things in a temporo-parietal semantic network activated by nouns, including the precuneus (PC). To our knowledge, this is the first demonstration that neural selectivity for living things does not require vision. We additionally observed preserved neural signatures of 'visual' light events in the left middle temporal gyrus (LMTG+). Across a wide range of semantic types, neural representations of sensory concepts develop independent of sensory experience.

Reading Braille by Touch Recruits Posterior Parietal Cortex.

Blind readers use a tactile reading system consisting of raised dot arrays: braille/⠃⠗⠇. How do human brains implement reading by touch? The current study looked for signatures of reading-specific orthographic processes in braille, separate from low-level somatosensory responses and semantic processes. Of specific interest were responses in posterior parietal cortices (PPCs), because of their role in high-level tactile perception. Congenitally blind, proficient braille readers read real words and pseudowords by touch while undergoing fMRI. We leveraged the system of contractions in English braille, where one braille cell can represent multiple English print letters (e.g., "ing" ⠬, "one" ⠐⠕), making it possible to separate physical and orthographic word length. All words in the study consisted of four braille cells, but their corresponding Roman letter spellings varied from four to seven letters (e.g., "con-c-er-t" ⠒⠉⠻⠞. contracted: four cells; uncontracted: seven letters). We found that the bilateral supramarginal gyrus in the PPC increased its activity as the uncontracted word length increased. By contrast, in the hand region of primary somatosensory cortex (S1), activity increased as a function of a low-level somatosensory feature: dot-number per word. The PPC also showed greater response to pseudowords than real words and distinguished between real and pseudowords in multivariate-pattern analysis. Parieto-occipital, early visual and ventral occipito-temporal, as well as prefrontal cortices also showed sensitivity to the real-versus-pseudoword distinction. We conclude that PPC is involved in orthographic processing for braille, that is, braille character and word recognition, possibly because of braille's tactile modality.

Visual experience shapes functional connectivity between occipital and non-visual networks.

Comparisons across adults with different sensory histories (blind vs. sighted) have uncovered effects of experience on human brain function. In people born blind visual cortices are responsive to non-visual tasks and show altered functional connectivity at rest. Since almost all research has been done with adults, little is known about the developmental origins of this plasticity. Are infant visual cortices initially functionally like those of sighted adults and blindness causes reorganization? Alternatively, do infants start like blind adults, with vision required to set up the sighted pattern? To distinguish between these possibilities, we compare resting state functional connectivity across blind (n = 30) and blindfolded sighted (n = 50) adults to a large cohort of sighted infants (Developing Human Connectome Project, n = 475). Remarkably, we find that infant secondary visual cortices functionally resemble those of blind more than sighted adults, consistent with the idea that visual experience is required to set up long-range functional connectivity. Primary visual cortices show a mixture of instructive effects of vision and reorganizing effects of blindness. Specifically, in sighted adults, visual cortices show stronger functional coupling with nonvisual sensory-motor networks (i.e., auditory, somatosensory/motor) than with higher-cognitive prefrontal cortices (PFC). In blind adults, visual cortices show stronger coupling with PFC. In infants, connectivity of secondary visual cortices is stronger with PFC, while V1 shows equal sensory-motor/PFC connectivity. In contrast, lateralization of occipital-to-frontal connectivity resembles the sighted adults at birth and is reorganized by blindness, possibly due to recruitment of occipital networks for lateralized cognitive functions, such as language.

Sensory modality and spoken language shape reading network in blind readers of Braille.

The neural basis of reading is highly consistent across many languages and scripts. Are there alternative neural routes to reading? How does the sensory modality of symbols (tactile vs. visual) influence their neural representations? We examined these questions by comparing reading of visual print (sighted group, n = 19) and tactile Braille (congenitally blind group, n = 19). Blind and sighted readers were presented with written (words, consonant strings, non-letter shapes) and spoken stimuli (words, backward speech) that varied in word-likeness. Consistent with prior work, the ventral occipitotemporal cortex (vOTC) was active during Braille and visual reading. A posterior/anterior vOTC word-form gradient was observed only in sighted readers with more anterior regions preferring larger orthographic units (words). No such gradient was observed in blind readers. Consistent with connectivity predictions, in blind compared to sighted readers, posterior parietal cortices were recruited to a greater degree and contained word-preferring patches. Lateralization of Braille in blind readers was predicted by laterality of spoken language and reading hand. The effect of spoken language increased along a cortical hierarchy, whereas effect of reading hand waned. These results suggested that the neural basis of reading is influenced by symbol modality and spoken language and support connectivity-based views of cortical function.

Naturalistic stimuli reveal a sensitive period in cross modal responses of visual cortex: Evidence from adult-onset blindness.

How do life experiences impact cortical function? In people who are born blind, the "visual" cortices are recruited during nonvisual tasks, such as Braille reading and sound localization. Do visual cortices have a latent capacity to respond to nonvisual information throughout the lifespan? Alternatively, is there a sensitive period of heightened plasticity that makes visual cortex repurposing especially possible during childhood? To gain insight into these questions, we leveraged meaningful naturalistic auditory stimuli to simultaneously engage a broad range of cognitive domains and quantify cross-modal responses across congenitally blind (n = 22), adult-onset blind (vision loss >18 years-of-age, n = 14) and sighted (n = 22) individuals. During fMRI scanning, participants listened to two types of meaningful naturalistic auditory stimuli: excerpts from movies and a spoken narrative. As controls, participants heard the same narrative with the sentences shuffled and the narrative played backwards (i.e., meaningless sounds). We correlated the voxel-wise timecourses of different participants within condition and group. For all groups, all stimulus conditions induced synchrony in auditory cortex while only the narrative stimuli synchronized responses in higher-cognitive fronto-parietal and temporal regions. As previously reported, inter-subject synchrony in visual cortices was higher in congenitally blind than sighted blindfolded participants and this between-group difference was particularly pronounced for meaningful stimuli (movies and narrative). Critically, visual cortex synchrony was no higher in adult-onset blind than sighted blindfolded participants and did not increase with blindness duration. Sensitive period plasticity enables cross-modal repurposing in visual cortices.

Naturalistic Audio-Movies reveal common spatial organization across "visual" cortices of different blind individuals.

Occipital cortices of different sighted people contain analogous maps of visual information (e.g. foveal vs. peripheral). In congenital blindness, "visual" cortices respond to nonvisual stimuli. Do visual cortices of different blind people represent common informational maps? We leverage naturalistic stimuli and inter-subject pattern similarity analysis to address this question. Blindfolded sighted (n = 22) and congenitally blind (n = 22) participants listened to 6 sound clips (5-7 min each): 3 auditory excerpts from movies; a naturalistic spoken narrative; and matched degraded auditory stimuli (Backwards Speech, scrambled sentences), during functional magnetic resonance imaging scanning. We compared the spatial activity patterns evoked by each unique 10-s segment of the different auditory excerpts across blind and sighted people. Segments of meaningful naturalistic stimuli produced distinctive activity patterns in frontotemporal networks that were shared across blind and across sighted individuals. In the blind group only, segment-specific, cross-subject patterns emerged in visual cortex, but only for meaningful naturalistic stimuli and not Backwards Speech. Spatial patterns of activity within visual cortices are sensitive to time-varying information in meaningful naturalistic auditory stimuli in a broadly similar manner across blind individuals.

Superior verbal but not nonverbal memory in congenital blindness.

Previous studies suggest that people who are congenitally blind outperform sighted people on some memory tasks. Whether blindness-associated memory advantages are specific to verbal materials or are also observed with nonverbal sounds has not been determined. Congenitally blind individuals (n = 20) and age and education matched blindfolded sighted controls (n = 22) performed a series of auditory memory tasks. These included: verbal forward and backward letter spans, a complex letter span with intervening equations, as well as two matched recognition tasks: one with verbal stimuli (i.e., letters) and one with nonverbal complex meaningless sounds. Replicating previously observed findings, blind participants outperformed sighted people on forward and backward letter span tasks. Blind participants also recalled more letters on the complex letter span task despite the interference of intervening equations. Critically, the same blind participants showed larger advantages on the verbal as compared to the nonverbal recognition task. These results suggest that blindness selectively enhances memory for verbal material. Possible explanations for blindness-related verbal memory advantages include blindness-induced memory practice and 'visual' cortex recruitment for verbal processing.

Neural basis of approximate number in congenital blindness.

Although humans are unique among animals in their ability to manipulate symbolic numbers, we share with other species an approximate number sense that allows us to estimate and compare the number of objects or events in a set, such as the number of apples in a tree. Our ability to discriminate the numerosity of two sets decreases as the ratio between them becomes smaller (e.g., 8 vs 16 items is harder to discriminate than 8 vs 32 items). The intraparietal sulcus (IPS) plays a key role in this numerical approximation. Neuronal populations within the IPS code for numerosity, with stimuli of different numerosities eliciting discriminable spatial patterns of activity. The developmental origins of these IPS number representations are not known. Here, we tested the hypothesis that representations of number in the IPS require visual experience with object sets, by working with individuals blind from birth. While undergoing fMRI, congenitally blind (n = 17) and blindfolded sighted (n = 25) participants judged which of two sequences of beeps was more numerous. In both sighted and blind individuals, patterns of activity in the IPS discriminated among different numerosities (4, 8, 16 vs 32), with better discrimination in the IPS of the blind group. In both groups, decoding performance decreased as the ratio between numerosities decreased (e.g., 8 vs 16 was less discriminable than 8 vs 32). These findings suggest that number representations in the IPS either have innate precursors, or that auditory or tactile experience with sets is sufficient for typical development.

Shared understanding of color among sighted and blind adults.

Empiricist philosophers such as Locke famously argued that people born blind might learn arbitrary color facts (e.g., marigolds are yellow) but would lack color understanding. Contrary to this intuition, we find that blind and sighted adults share causal understanding of color, despite not always agreeing about arbitrary color facts. Relative to sighted people, blind individuals are less likely to generate "yellow" for banana and "red" for stop sign but make similar generative inferences about real and novel objects' colors, and provide similar causal explanations. For example, people infer that two natural kinds (e.g., bananas) and two artifacts with functional colors (e.g., stop signs) are more likely to have the same color than two artifacts with nonfunctional colors (e.g., cars). People develop intuitive and inferentially rich "theories" of color regardless of visual experience. Linguistic communication is more effective at aligning intuitive theories than knowledge of arbitrary facts.

'Visual' cortices of congenitally blind adults are sensitive to response selection demands in a go/no-go task.

Studies of occipital cortex plasticity in blindness provide insight into how intrinsic constraints interact with experience to determine cortical specialization. We tested the cognitive nature and anatomical origins of occipital responses during non-verbal, non-spatial auditory tasks. In a go/no-go task, congenitally blind (N=23) and sighted (N=24) individuals heard rapidly occurring (<1/s) non-verbal sounds and made one of two button presses (frequent-go 50%, infrequent-go 25%) or withheld a response (no-go, 25%). Rapid and frequent button presses heighten response selection/inhibition demands on the no-go trials: In sighted and blind adults a right-lateralized prefrontal (PFC) network responded most to no-go trials, followed by infrequent-go and finally frequent-go trials. In the blind group only, a right-lateralized occipital network showed the same response profile and the laterality of occipital and PFC responses was correlated across blind individuals. A second experiment with spoken sentences and equations (N=16) found that no-go responses in occipital cortex are distinct from previously identified occipital responses to spoken language. Finally, in resting-state data (N=30 blind, N=31 blindfolded sighted), no-go responsive 'visual' cortex of blind relative to sighted participants was more synchronized with PFC and less synchronized with primary auditory and sensory-motor cortices. No-go responsive occipital cortex showed higher resting-state correlations with no-go responsive PFC than language responsive inferior frontal cortex. We conclude that in blindness, a right-lateralized occipital network responds to non-verbal executive processes, including response selection. These results suggest that connectivity with fronto-parietal executive networks is a key mechanism for plasticity in blindness.

How does a blind person see? Developmental change in applying visual verbs to agents with disabilities.

Classic theories emphasize the primacy of first-person sensory experience for learning meanings of words: to know what "see" means, one must be able to use the eyes to perceive. Contrary to this idea, blind adults and children acquire normative meanings of "visual" verbs, e.g., interpreting "see" and "look" to mean with the eyes for sighted agents. Here we ask the flip side of this question: how easily do sighted children acquire the meanings of "visual" verbs as they apply to blind agents? We asked sighted 4-, 6- and 9-year-olds to tell us what part of the body a blind or a sighted agent would use to "see", "look" (and other visual verbs, n = 5), vs. "listen", "smell" (and other non-visual verbs, n = 10). Even the youngest children consistently reported the correct body parts for sighted agents (eyes for "look", ears for "listen"). By contrast, there was striking developmental change in applying "visual" verbs to blind agents. Adults, 9- and 6-year-olds, either extended visual verbs to other modalities for blind agents (e.g., "seeing" with hands or a cane) or stated that the blind agent "cannot" "look" or "see". By contrast, 4-year-olds said that a blind agent would use her eyes to "see", "look", etc., even while explicitly acknowledging that the agent's "eyes don't work". Young children also endorsed "she is looking at the dax" descriptions of photographs where the blind agent had the object in her "line of sight", irrespective of whether she had physical contact with the object. This pattern held for leg-motion verbs ("walk", "run") applied to wheelchair users. The ability to modify verb modality for agents with disabilities undergoes developmental change between 4 and 6. Despite this, we find that 4-year-olds are sensitive to the semantic distinction between active ("look") and stative ("see"), even when applied to blind agents. These results challenge the primacy of first-person sensory experience and highlight the importance of linguistic input and social interaction in the acquisition of verb meaning.

Computer code comprehension shares neural resources with formal logical inference in the fronto-parietal network.

Despite the importance of programming to modern society, the cognitive and neural bases of code comprehension are largely unknown. Programming languages might 'recycle' neurocognitive mechanisms originally developed for natural languages. Alternatively, comprehension of code could depend on fronto-parietal networks shared with other culturally-invented symbol systems, such as formal logic and symbolic math such as algebra. Expert programmers (average 11 years of programming experience) performed code comprehension and memory control tasks while undergoing fMRI. The same participants also performed formal logic, symbolic math, executive control, and language localizer tasks. A left-lateralized fronto-parietal network was recruited for code comprehension. Patterns of activity within this network distinguish between 'for' loops and 'if' conditional code functions. In terms of the underlying neural basis, code comprehension overlapped extensively with formal logic and to a lesser degree math. Overlap with executive processes and language was low, but laterality of language and code covaried across individuals. Cultural symbol systems, including code, depend on a distinctive fronto-parietal cortical network.

Enhanced performance on a sentence comprehension task in congenitally blind adults.

People born blind habitually experience linguistic utterances in the absence of visual cues and activate "visual" cortices during sentence comprehension. Do blind individuals show superior performance on sentence processing tasks? Congenitally blind (n=25) and age and education matched sighted (n=52) participants answered yes/no who-did-what-to-whom questions for auditorily-presented sentences, some of which contained a grammatical complexity manipulation (long-distance movement dependency or garden path). Short-term memory was measured with a forward and backward letter-spans. A battery of control tasks included two speeded math tasks and vocabulary and reading tasks from Woodcock Johnson III. The blind group outperformed the sighted on the sentence comprehension task, particularly for garden-path sentences, and on short-term memory span tasks, but performed similar to the sighted on control tasks. Sentence comprehension performance was not correlated with span performance, suggesting independent enhancements.

A sensitive period in the neural phenotype of language in blind individuals.

Congenital blindness modifies the neural basis of language: "visual" cortices respond to linguistic information, and fronto-temporal language networks are less left-lateralized. We tested the hypothesis that this plasticity follows a sensitive period by comparing the neural basis of sentence processing between adult-onset blind (AB, n = 16), congenitally blind (CB, n = 22) and blindfolded sighted adults (n = 18). In Experiment 1, participants made semantic judgments for spoken sentences and, in a control condition, solved math equations. In Experiment 2, participants answered "who did what to whom" yes/no questions for grammatically complex (with syntactic movement) and simpler sentences. In a control condition, participants performed a memory task with non-words. In both experiments, visual cortices of CB and AB but not sighted participants responded more to sentences than control conditions, but the effect was much larger in the CB group. Only the "visual" cortex of CB participants responded to grammatical complexity. Unlike the CB group, the AB group showed no reduction in left-lateralization of fronto-temporal language network, relative to the sighted. These results suggest that congenital blindness modifies the neural basis of language differently from adult-onset blindness, consistent with a developmental sensitive period hypothesis.

Sensitive periods in cortical specialization for language: insights from studies with Deaf and blind individuals.

Studies with Deaf and blind individuals demonstrate that linguistic and sensory experiences during sensitive periods have potent effects on neurocognitive basis of language. Native users of sign and spoken languages recruit similar fronto-temporal systems during language processing. By contrast, delays in sign language access impact proficiency and the neural basis of language. Analogously, early but not late-onset blindness modifies the neural basis of language. People born blind recruit 'visual' areas during language processing, show reduced left-lateralization of language and enhanced performance on some language tasks. Sensitive period plasticity in and outside fronto-temporal language systems shapes the neural basis of language.