Humans are the only known species where much of knowledge learning happens symbolically through language, in addition to information received directly from the senses. For example, humans can learn about the color of some rose flowers from the popular expression “roses are red” without needing to see any red roses – allowing them to accumulate knowledge beyond the constraints of their own senses.

Recent work suggests that a region of the brain known as the dorsal anterior temporal lobe represents knowledge acquired from language instead of sensory experiences. However, these studies were based on volunteers deprived of sensory experiences rather than those with reduced language exposure. Therefore, it was not clear whether this brain structure represents knowledge derived specifically from language and the importance of language in shaping non-sensory knowledge.

To address this question, Wang et al. studied the brain activity of deaf adult volunteers in a word meaning judgement task. Volunteers were either born deaf or lost their hearing as toddlers, and all primarily used Chinese Sign Language for communication. One group of volunteers had been exposed to sign language from birth, giving them similar exposure to language as hearing individuals. The other group had less exposure to language in their early years and only learned sign language later in childhood.

The task included 90 written words that were familiar to the volunteers. They included a mixture of object words – related to material objects – such as “shoulder” and “hammer” and abstract words – which are not linked to physical objects – such as “cause” and “violence”. The volunteers were shown each word in turn and asked to think about the word’s meaning. Brain scans revealed that the left dorsal anterior temporal lobes of the volunteers with reduced early language exposure were less sensitive to the meaning of the words compared with those of the other volunteers.

The findings demonstrate that the dorsal anterior temporal lobe specifically supports meaning derived from a person’s experience of language as opposed to sensory experience, providing a new angle to understand the mechanism of knowledge representations. Increased understanding of how language supports knowledge will help to uncover the human-specific ways of representing and creating knowledge in the brain.

Humans are believed to be the only species in the animal kingdom where knowledge learning can be achieved symbolically, mostly through language (‘roses are red’), in addition to sensory channels (visually perceiving roses in color red) (Gelman and Roberts, 2017; Perszyk and Waxman, 2018). A common view shared by the modern neurocognitive theories of semantics is that semantic knowledge, even those acquired through language, is ultimately grounded in (nonlinguistic) sensory/motor experiences, encoded in the distributed brain areas encompassing high-level sensorimotor areas, and potential hub regions that bind such sensory-derived representations (Barsalou, 2016; Binder et al., 2009; Fernandino et al., 2022; Ralph et al., 2017; Martin, 2016). However, language experiences may contribute to semantic development beyond sensory experiences by facilitating or modulating categorizations by the nature of labeling (i.e. words) to the sensory experiences, and/or by constructing semantic relations based on various types of word relations (Gelman and Roberts, 2017; Perszyk and Waxman, 2018; Unger and Fisher, 2021). Are such cognitive contributions manifested by modulating the neural representations of the sensory-derived semantic spaces, or by also formulating neural representations that specialize in representing knowledge derived from language experience (i.e. not nonlinguistic sensory experience)?

Neural representations of fully nonsensory, language-derived knowledge have only recently been inferred based on studies with sensory-deprived individuals (Bottini et al., 2020; Striem-Amit et al., 2018; Wang et al., 2020, see Bi, 2021, for a review). Congenitally blind individuals, who cannot acquire visual-specific knowledge (e.g. color) through sensory experiences, can nevertheless acquire semantic structures about such knowledge behaviorally similar to those in the sighted, presumably derived from language. Such nonsensory semantic structures in the blind (and also sighted) are represented in the dorsal anterior temporal lobe (dATL), with sighted individuals additionally representing the highly similar knowledge structure (presumably derived from sensory experience) in the visual cortex (Wang et al., 2020). This dATL cluster is distinct from the central ‘amodal’ semantic hub proposed to bind together multiple sensory attributes, which is located in the more ventral-medial territory of the ATL (Ralph et al., 2017; Patterson et al., 2007; see discussions in Striem-Amit et al., 2018). The left dATL is also more strongly activated by abstract than concrete words in typically developed individuals (Binder et al., 2009; Bucur and Papagno, 2021; Wang et al., 2010; Wang et al., 2019). It was thus proposed that this area represents knowledge derived from language, in addition to those from sensory experiences in various perceptual (association) regions (Bi, 2021). These lines of evidence, while highly suggestive, are based on manipulating sensory experience by examining individuals deprived of a sensory modality or by contrasting concepts with rich sensory experiences versus those without, rather than the positive manipulation of language experience. Thus, the positive evidence for the necessity of language experiences for the neural semantic representation here is still lacking.

Here, we address this issue in a special human model that varies in early language exposure: individuals who were born profoundly deaf in hearing families and had limited natural language exposure (speech or sign) during early childhood (Goldin-Meadow and Feldman, 1977; Mayberry et al., 2002). These individuals often acquired their first language (sign language) around school age, which is much later than the age of first language (L1) acquisition in typically developed children or in native deaf signers who were born in deaf families and acquired sign language from birth. Previous research has shown that such early language exposure limitation led to long-lasting effects on various aspects of language processing. Behaviorally, these delayed deaf signers, even during adulthood with many years of sign language usage, have lower proficiency in phonological, morphological, and syntactic processing of sign language (Bogliotti et al., 2020; Caselli et al., 2021; Cheng and Mayberry, 2021; Lieberman et al., 2015; Mayberry et al., 2002; Mayberry and Fischer, 1989; Newport, 1990; Tomaszewski et al., 2022). Neurally, brain functional imaging studies have reported decreased activation magnitude in the left inferior frontal and posterior temporal regions in tasks of sign sentence judgments (Mayberry et al., 2011; Richardson et al., 2020; Twomey et al., 2020). Anatomical alterations in regions typically recruited in language tasks – reduced cortical volume in the left inferior frontal region, reduced cortical thickness in the left posterior middle temporal region, and reduced fractional anisotropy values in the left arcuate fasciculus – were also reported in signers with delayed L1 acquisition (Cheng et al., 2023; Cheng et al., 2019).

Despite these documented effects of delayed L1 acquisition on phonology, morphology, and syntax (Curtiss, 1977; Lenneberg, 1967; Lillo-Martin and Henner, 2021; Mayberry and Kluender, 2018), studies have reported little effects on semantic behaviors, including semantic interference effects in the picture-sign paradigm (Baus et al., 2008), scalar implicature (Davidson and Mayberry, 2015), or accuracy scores of several written word semantic tasks (e.g. synonym judgment) (Choubsaz and Gheitury, 2017). However, as shown by the color knowledge in the congenitally blind studies (e.g. Wang et al., 2020), similar semantic behaviors may arise from (partly) different neural representations. Semantic processing is supported by a multifaceted cognitive system and a complex neural network entailing distributed brain regions (Bi, 2021; Binder and Desai, 2011; Ralph et al., 2017; Martin, 2016), and thus focal neural changes may not necessarily lead to semantic behavioral changes. Neurally, neurophysiological signatures assumed to reflect semantic processes showed incongruent effects across studies: N400 effects in the semantic violation of written sentences were not affected (Skotara et al., 2012), whereas M400 in the picture-sign matching task showed atypical activation patterns (reduced recruitment of the left fronto-temporal regions and involvement of the right parietal and occipital regions) (Ferjan Ramirez et al., 2016; Ferjan Ramirez et al., 2014; Mayberry et al., 2018). It remains to be tested whether and where delayed L1 acquisition affects how semantics are neurally represented, using imaging techniques with higher spatial resolutions.

In this study, by a rare opportunity of manipulation of early language exposure offered by nature, we aim to test the neural system representing the language-derived semantic representations, beyond the sensory-derived semantic representations (Bi, 2021; Wang et al., 2020). With fMRI experiments on semantic processing of familiar words, we compared the neural semantic structures between congenitally deaf signers with delayed sign language acquisition (delayed deaf signers) and congenitally deaf signers with native sign language acquisition (native deaf signers), that is, two groups that are matched on their nonlinguistic sensory experiences but varied in early language experience. Significant group differences would provide positive evidence for the role of language experience in the formation of the identified neural semantic structures.

By comparing fMRI BOLD responses to the word semantic structures between native and delayed early deaf signers, we aimed to examine how early-life language exposure affects the semantic neural representations. Our results demonstrated that semantic information in the left dATL was significantly reduced in delayed signers compared with native signers in both multivariate (representation of rich semantic space) and univariate (preference to abstract/nonobject words) analyses. That is, early-life language acquisition – critical to language system neurodevelopment – is necessary for the left dATL to exhibit the typical semantic organization in adulthood. These results provide the first positive evidence for the effect of language experience on driving the semantic functional development of a particular brain region.

Two types of specificity are to be clarified – anatomical specificity and information specificity. First, is this group effect specific to the dATL, relative to other brain regions? We do not have evidence for such a strong region specificity. We did not observe a significant group-by-ROI interaction in either the RSA or the univariate analyses. That is, the group differences were not significantly stronger in the dATL than in the other semantic regions being analyzed (IFG and pMTG). We are thus not claiming that the dATL is the only region that derives semantic representation from language experience, but choose to focus the following discussion on this region because of the robust positive effects here. Second, is the dATL semantic representation specifically derived from language experience and not other (nonlinguistic) sensory experiences? The manipulation of the current study – the two groups of deaf signers – varies on the language exposure while matching on the sensory experiences (see below), and thus did not test the presence or absence of sensory-derived semantic representations. Inferences could be drawn in combination with the previous studies that focused on the manipulation of sensory experiences by studying visual knowledge in congenitally blind subjects. There, it was reported that the blind and sighted had comparable semantic information encoding in the RSA (Wang et al., 2020). In terms of univariate effects (abstractness or color concept adaptation), deprivation of sensory experiences did not reduce, but actually tended to enhance the effects here (Bottini et al., 2020; Striem-Amit et al., 2018). That is, evidence from congenitally blind studies does not support the additional sensory-derived semantic encoding here. While drawing negative conclusions is always difficult, we reason that it is parsimonious, based on available data, to propose that the dATL’s contribution to semantic encoding is specific to language, and not sensory-derived representations.

What kind of variables best account for the observed group differences in the dATL? The two deaf groups were matched on sensory experiences (both similarly profoundly deaf), a wide range of nonlinguistic environmental variables including the socioeconomic status (except for parents’ education backgrounds, which were better for the delayed signer group), the length and level of formal education, through which largely comparable sign language and word reading skills were achieved. The salient difference was the early-life language experience (before 6.9 years of age – the mean age of acquisition in our delayed signer group). Notably, deaf children with limited access to natural language often spontaneously develop homesign systems, which share some aspects with natural language (Goldin-Meadow, 2003). For instance, they could use homesigns to produce generic utterances in a similar manner to typically developed children (Goldin-Meadow et al., 2005). Notably, routine nation-wide neonate hearing screening in China did not start until 2009, years after the early childhood of our participants (born before 2000), and some hearing parents may nonetheless try to give deaf children additional aids of exposure to signs (via preschool special education programs) or speech (via hearing aids). Critically, our positive results of the robust group differences in the dATL suggest that early homesign/aid measures and later formal education for sign and written language experiences are insufficient for typical dATL neurodevelopment; the full-fledged language experience during early infancy and childhood (before school age) plays a necessary role in this process. A further variable to consider is that a lack of early language exposure may lead to alterations beyond the language system, affecting non-language cognitive domains such as working memory or theory of mind in children (Marshall et al., 2015; Richardson et al., 2020). While we cannot rule out the effects of these potential intermediate cognitive processes, the positive evidence of these variables in dATL functionality is lacking. The most direct manipulation and parsimonious account for the dATL effects is language-related.

The effects of language experience in shaping the semantic representation in the dATL observed here are corroborated by several previous lines of indirect evidence for the relevance of language in its functionality. It shows a functional preference for abstract words, which are less salient in sensory attributes than concrete words and are assumed to entail more language processes such as context diversity (e.g. Binder et al., 2009; Bucur and Papagno, 2021; Wang et al., 2010). In congenitally blind individuals, it encodes visual knowledge such as color and the activity strength is negatively modulated by perceptibility (Bottini et al., 2020; Striem-Amit et al., 2018; Wang et al., 2020). It is sensitive to semantic composition, such as in two-word phrases or sentences (Pallier et al., 2011; Pylkkänen, 2019). This region is functionally and structurally connected with other language-sensitive regions (Fan et al., 2014; Friederici et al., 2017; Ralph et al., 2017; Pascual et al., 2015; Wang et al., 2019). Finally, a recent neurodevelopmental study shows that the early-life resting-state functional connectivity patterns of the left temporal pole significantly predict performances in tasks entailing semantic processes, not in tasks of phonological skills or rapid automatized naming, at about 6.5 years of age (Yu et al., 2021). These lines of findings are often correlational evidence for the language effect on their own. Together with the positive evidence of early language experience in developing typical semantic representation in the dATL reported here, the parsimonious proposal accounting for them together would be that the dATL represents knowledge derived from language (e.g. from higher-order word relations). Without early language exposure, the semantic structure representation and the abstractness preference here are reduced.

Together, this package of findings extends beyond the majority of the neurocognitive semantic theories, which mainly focus on the representing, binding, and controlling of knowledge derived from nonlinguistic multisensory experiences (Binder, 2016; Ralph et al., 2017; Martin, 2016; but see Mahon and Caramazza, 2008). The recently developed dual-neural-coding framework for semantic knowledge (Bi, 2021), which explicitly proposes that the human brain has developed regions (the dATL) for language-derived semantic representations, in addition to the sensory-derived knowledge representations, naturally accommodates these observations. Language, as a symbolic system, allows us to convey and manipulate meanings free from sensorimotor experiences and plays a key role in knowledge transmission and accumulation across individuals and generations (Gelman and Roberts, 2017). The existence of the language-derived knowledge neural system may lay the foundation for the efficient storage and manipulation of semantic knowledge, particularly abstract knowledge without tangible and consistent sensorimotor referents (Bi, 2021).

The current results also have implications for the role of early language exposure in neurodevelopment more generally. Previous studies have reported that delayed L1 acquisition associates with reduced cortical volume in the left inferior frontal region, reduced cortical thickness in the left posterior middle temporal region, and the reduced fractional anisotropy values in the left arcuate fasciculus that structurally connects the two regions, and not ATL or its related white matter tracts (Cheng et al., 2023; Cheng et al., 2019). Our findings of semantic functional alterations in the dATL are thus not easily attributable to broad anatomical changes associated with late L1 acquisition. Notably, different from phonological and syntactic processes, where both visible behavioral underdevelopment (e.g. Caselli et al., 2021; Cheng and Mayberry, 2021; Mayberry et al., 2002) and brain functional changes (Mayberry et al., 2011; Richardson et al., 2020; Twomey et al., 2020) were observed, for semantics we only observed brain functional changes in the dATL but no visible behavioral effects. Consistent with the literature where deaf delayed signers did not show differences to controls in semantic interference effects in the picture-sign paradigm (Baus et al., 2008), scalar implicature (Davidson and Mayberry, 2015), or N400 measures (Skotara et al., 2012), we did not observe visible differences in terms of semantic distance structures (Figure 1a) or reaction time of lexical decision and word-triplet semantic judgment (Figure 1—figure supplement 1). As reasoned in the Introduction, this seeming neuro-behavior discrepancy might be related to the multifaceted, distributed nature of the cognitive and neural basis of semantics more broadly. The general semantic behavioral tasks we employed could be achieved with representations derived from multiple types of experiences, supported by highly distributed neural systems (e.g. Bi, 2021; Binder and Desai, 2011; Ralph et al., 2017; Martin, 2016), including those not affected by delayed L1 acquisition in regions beyond the dATL. This finding invites future studies to specify the exact developmental mechanisms in the left dATL (Fu et al., 2023; Unger and Fisher, 2021) and to uncover semantic behavioral consequences related to the functionality of this area.

To conclude, by manipulation of early language experience by nature we were able to positively identify a human brain region – dATL – robustly supporting word meanings derived from language experience, in contrast to those grounding nonlinguistic, sensory experiences. These findings highlight the role of language in forming specific neural semantic representations in the human brain, and raise new questions about the ontogenetic mechanisms of this intriguing neural structure.

Source data files have been provided for Table 1 and all the figures. Additional behavioral and neural data have been made available on OSF at the link https://osf.io/wz6q9/. The whole-brain unthresholded statistical maps have also been made available on NeuroVault at the link https://neurovault.org/collections/13705/. Deidentified Nifti files are not shared openly because of ethics constraints, but are available from the corresponding authors upon reasonable request.

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   Early language exposure affects neural mechanisms of semantic representations.

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Author details

1. Xiaosha Wang

   1. State Key Laboratory of Cognitive Neuroscience and Learning & IDG/McGovern Institute for Brain Research, Beijing Normal University, Beijing, China
   2. Beijing Key Laboratory of Brain Imaging and Connectomics, Beijing Normal University, Beijing, China

   Contribution

   Conceptualization, Data curation, Software, Formal analysis, Funding acquisition, Validation, Investigation, Visualization, Methodology, Writing – original draft, Project administration, Writing – review and editing

   For correspondence

   wangxiaosha@bnu.edu.cn

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-2133-8161

2. Bijun Wang

   1. State Key Laboratory of Cognitive Neuroscience and Learning & IDG/McGovern Institute for Brain Research, Beijing Normal University, Beijing, China
   2. Beijing Key Laboratory of Brain Imaging and Connectomics, Beijing Normal University, Beijing, China

   Contribution

   Data curation, Formal analysis, Methodology, Project administration

   Competing interests

   No competing interests declared

3. Yanchao Bi

   1. State Key Laboratory of Cognitive Neuroscience and Learning & IDG/McGovern Institute for Brain Research, Beijing Normal University, Beijing, China
   2. Beijing Key Laboratory of Brain Imaging and Connectomics, Beijing Normal University, Beijing, China
   3. Chinese Institute for Brain Research, Beijing, China

   Contribution

   Conceptualization, Resources, Supervision, Funding acquisition, Investigation, Writing – original draft, Project administration, Writing – review and editing

   For correspondence

   ybi@bnu.edu.cn

   Competing interests

   Senior Editor, eLife

   "This ORCID iD identifies the author of this article:" 0000-0002-0522-3372

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