# Networks for proper versus common name retrieval: insights from causal mapping and new hypotheses

Eléonor Burkhardt, Hugues Duffau, Guillaume Herbet

► To cite this version:

Eléonor Burkhardt, Hugues Duffau, Guillaume Herbet. Networks for proper versus common name retrieval: insights from causal mapping and new hypotheses. Brain Communications, 2025, 7 (4), pp.fcaf256. 10.1093/braincomms/fcaf256. hal-05173592

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<figure><img src="image_1.png" /></figure>
REVIEW ARTICLE

# Networks for proper versus common name retrieval: insights from causal mapping and new hypotheses

Eléonor Burkhardt,¹,² and Hugues Duffau²,³ and Guillaume Herbet¹,²,⁴,⁵

Over the years, the anatomical correlates of common name and proper name retrieval have been studied and discussed side by side in the field of cognitive neuroscience. Due to the particular semantic status of proper names (i.e. meaningless marker referring to unique entities with a token link), it is generally acknowledged that their lexical retrieval is cognitively more demanding than that of common names. Thus, the neural networks engaged for accessing the phonological output lexicon of common names and proper names could be partially segregated, centred on different cortical regions and underpinned by distinct structural pathways. However, this possible anatomo-functional organization has not yet been explored in an in-depth review. Hence, the objective of our review is 3-fold: (i) to highlight the cortical networks underlying common name and proper name retrieval; (ii) to summarize the findings from recent studies using causal methods of tract-oriented functional mapping (i.e. direct electrostimulation and disconnection-symptom mapping) with the aim of isolating ventral structural connections that contribute to common name and/or proper name retrieval; (iii) to develop hypotheses regarding the anatomical connectivity implicated in lexical access to common names and proper names, in relation to the input modality and semantic category (for proper names, in the latter case). Our critical review first suggests that, although both categories appear to rely on a widely distributed and overlapping left fronto-temporo-parietal cortical network, the mid-to-posterior part of the ventral temporal cortex (especially the fusiform gyrus) and the anterior temporal structures (including the temporal pole), seem more specifically involved in common name retrieval and in proper name retrieval respectively, regardless of their input modality. Second, the left inferior longitudinal fasciculus appears to play a crucial role in picture-based lexical retrieval for both common names and proper names. Based on these key findings, we propose tentative models of anatomical connections underlying lexical retrieval. We suggest that distinct components of the inferior longitudinal fasciculus are implicated in lexical retrieval for common names and proper names and that another similar segregation mediates the retrieval of face names and landmark names. We further hypothesize that the left middle longitudinal fasciculus plays a role in auditory-guided lexical retrieval, owing to its projections into the anterior temporal structures and its close relationship with the auditory associative cortex. Within the framework of the dual-language model, we discuss the heuristic value of these anatomical hypotheses to understand the mechanisms by which anomia arises in various neurological conditions, providing insights and directions for future studies.

1. Praxiling Laboratory, UMR 5267, CNRS, University Paul Valéry Montpellier 3, Montpellier 34090, France

2. Gui de Chauliac Hospital, Montpellier University Medical Centre, Montpellier 34295, France

3. CNRS, INSERM, Institute of Functional Genomics, University of Montpellier, Montpellier 34000, France

4. Department of Medicine, University of Montpellier, Montpellier 34090, France

5. Institut Universitaire de France, Paris, France

Correspondence to: Guillaume Herbet
Campus Arnaud de Villeneuve, Faculty of Medicine
641, avenue du doyen Gaston Guiraud
Occitanie, Montpellier 34090, France
E-mail: guillaume.herbet@umontpellier.fr

**Keywords:** anomia; lexical retrieval; anterior temporal lobe; basal temporal language area; inferior longitudinal fasciculus
## Graphical Abstract

<figure><img src="image_2.png" /></figure>

## Introduction

For many years, it was commonly believed that aphasia syndromes resulted from lesions affecting discrete and highly specialized areas of the cerebral cortex. An iconic example of this belief is the association between Broca's area and the impairment of speech production, as described by Paul Broca in 1861.¹ However, it gradually became evident that localizationist or simple associationist models could not fully account for both the neural organization of language and the complexity of aphasic disorders.²,³ Decades of neuropsychological studies have consistently shown that neuropsychological syndromes, in particular aphasia, stem from the disruption of brain-wide neural networks formed by distributed cortical and white matter structures.⁴⁻⁹ In the realm of language processing, contemporary network perspectives underscore the fundamental role of a predominantly left lateralized, interactive dual-stream neural system. Within this intricate neurobiological framework, the dorsal pathway mediates the precise execution of phonological and

articulatory processes, while the ventral pathway orches-
trates the complex mapping of percepts onto semantics.¹⁰⁻¹²

From a neuroanatomical standpoint, an accumulating
growing body of empirical evidence substantiates the pivotal
roles played by the arcuate fasciculus and the superior longi-
tudinal fasciculus within the dorsal stream. These white mat-
ter tracts constitute the foundational white matter
architecture through which phonological and speech articu-
latory information is seamlessly integrated. In a parallel
manner, the inferior fronto-occipital fasciculus (IFOF) (and
the so-called extreme capsule) establish critical connections
within the ventral stream for verbal and non-verbal semantic
processing.⁶,¹³⁻¹⁵ Furthermore, the complex tapestry of lan-
guage processes also encompasses the participation of the in-
ferior and middle longitudinal fasciculi (ILF and MdLF), in
addition to the uncinate fasciculus (UF), as parts of the ven-
tral stream. It is noteworthy that the precise functional role
of these tracts within the domain of language remains a sub-
ject of ongoing scientific exploration and this review endea-
vours to shed light on this particular aspect.
In the comprehensive study of the language network, the deployment of tract-oriented lesion-symptom mapping approaches offers a valuable means of exploring the relationship between surgery-induced disconnections and cognitive impairment. They are especially relevant in neurosurgical glioma patients since the anatomical connections serve as the primary conduits for tumour spreading and are generally less amenable to functional compensation, compared to associative cortical areas.¹⁶,¹⁷ In this context, these methods offer the advantage of identifying anatomo-functional associations, even in the presence of neuroplasticity.¹⁸ Thus, the disconnection-symptom mapping in patients with glioma has unveiled significant new insights regarding the language ventral stream.¹⁴ However, it is worth noting that some of these findings, especially those related to lexical retrieval, despite their importance, have yet to be comprehensively discussed and fully incorporated into existing network models of language.

In this review, we are interested in the neural networks responsible for lexical access to common names (CNs) and proper names (PNs), with a specific emphasis on ventral white matter connectivity. From a cognitive standpoint, naming an object involves a complex set of interacting processes that encompass perception, conceptual processing, word selection, phonological form retrieval, and, ultimately, verbal production through articulatory planning and execution.¹⁹⁻²¹ It also implicates supra-linguistic cognitive control functions. As a result, brain damage can disrupt naming abilities in various ways, depending on the specific process affected.²²,²³ In this neuropsychological context, pure anomia is commonly described as arising from a disconnection between preserved semantic knowledge and an intact phonological output lexicon.²⁴

On the other hand, while CNs (e.g. ‘blueberry’ or ‘table’) are meaningful and category-related, PNs (e.g. ‘John’ or ‘Geneva’) are meaningless and individuality-related.²⁵,²⁶ Thus, whereas the former are generally substitutable (with circumlocutions/synonyms), the latter are not,²⁷ which is particularly significant in the context of aphasia. Furthermore, PNs, especially when associated with faces, might involve peripheral socio-emotional processing²⁸,²⁹ and their lexical access may place a greater demand on memory resources compared to CNs, particularly in terms of episodic processes.²⁵,²⁶

Consequently, it has been hypothesized that the neural systems responsible for lexical access to PNs and CNs partly differ²⁵,²⁶ although this issue remains unresolved to date. In this context, some authors argue for a single semantic system (notably including lexical-semantic aspects), whose disruption may lead to naming impairments affecting both grammatical categories.³⁰ However, cognitive dissociations observed in neurological patients between anomia for CNs and PNs also provide some interesting insights. While there are a few reported cases of patients showing anomia for CNs without anomia for PNs,³¹⁻³⁴ albeit under particular circumstances (e.g. sparing of country names only, deficit at the semantic level, or deficit confined to the oral modality,

etc.),²⁶ the reverse pattern, especially for person names, is more frequently reported.³²,³⁵⁻³⁹ Therefore, it is somewhat expected that circumscribed damage to specialized cortical or subcortical structures can occasionally lead to selective anomia, especially for PNs and to a lesser extent, for CNs.

It is worth noting that previous reviews have thoroughly explored the cognitive (neuro)psychology of PNs, sometimes in relation to their potential neural underpinnings, compared to CNs.²⁵⁻²⁷,⁴⁰,⁴¹ However, none has attempted to dissociate the neural networks involved in the lexical access to these two grammatical categories, especially at the white matter level. This gap exists due to the only recent shift in research focus towards specifically exploring the role of ventral structural connectivity in lexical access.⁴²⁻⁴⁸ In this context, the primary aim of this critical review is to disentangle the different contributions of the ventral neural circuits to lexical retrieval, placing a particular emphasis on the ventral white matter tracts, while acknowledging the relatively limited availability of data related to the latter. Crucially, highlighting white matter damage in the context of anomia may help clarify the relationships between patients’ aphasic impairments and lesion topography, thereby advancing our understanding of the network-level anatomo-clinical mapping of naming deficits.

In the opening section of this review, we offer a comprehensive update on the cortical foundations of CN and PN retrieval, with a special focus on identifying commonalities and distinctions within the cortical systems underlying these processes. In the next section, we delve into the various anatomical connectivities implicated in CN and PN retrieval, with a special emphasis to the left ILF, for which causal evidence has accruing until recently.⁴²⁻⁴⁵,⁴⁸ Finally, in the last part of this review, we present hypothetical network models for PN/CN retrieval. These models serve as a foundational framework for developing new linguistically constrained hypotheses regarding the axonal connections involved in retrieval processes, in relation to the grammatical class of names (PNs and/or CNs), semantic subcategories within PNs (famous persons/famous landmarks) and the input modality, for both CNs and PNs (visual versus auditory stimuli).

## Cortical networks for proper name and common name retrieval: an update

As a pre-requisite, it is important to clarify that neuroscience and neuropsychological studies have generally not strictly compared the cortical bases involved in PN versus CN retrieval. While one cortical mapping study⁴⁹ administered famous face and object naming tasks to patients with brain tumours during awake surgery and highlighted a number of dissociated functional sites, it did not investigate the critical basal temporal structures. Other research has mainly focused on CN and PN retrieval separately, often using picture
naming tasks<sup>22,50,51</sup> or famous face naming (FFN) tasks<sup>52,53</sup> respectively, while controlling for several cognitive processes (e.g. recognition/access to semantic information for unique entities,<sup>54</sup> and recognition or fluency for non-unique entities<sup>50</sup>). Alternatively, some studies have investigated the effect of semantic categories on naming, including equally both common (animate and inanimate objects) and unique entities (mainly faces).<sup>28,55</sup> Lastly, a clinical group study<sup>30</sup> and numerous case studies<sup>26</sup> investigated CN and PN retrieval using comprehensive language assessments but the former only concerned patients with ATS damage while the latter did not included anatomo-clinical analyses. Of note, ATS generally correspond to the anterior aspects of all temporal gyri (including the temporal pole, the superior, middle and inferior temporal gyri together with the fusiform gyrus).<sup>56</sup>

Regarding PNs specifically, a set of behaviour-based neuropsychological<sup>57-59</sup> or neuromodulation<sup>60</sup> studies focusing on ATS, analysed the modality effect (verbal/non-verbal, mainly corresponding to naming/recognition) on performance, depending on the hemisphere laterality. Other works investigated the sensibility of the ATS to the input modality in people naming tasks, depending on whether they were presented with their voice or with their face.<sup>60,61</sup> In the same way, some studies were interested in the role of the ATS in naming unique entities depending on the semantic category (e.g. landmarks versus famous faces).<sup>52,62</sup> Importantly, our review excludes findings related to associative face/name learning<sup>63</sup> and instead focuses on lexical retrieval of PNs associated with well-known unique entities, typically assessed using overt naming paradigms (as well as of non-unique entities, generally assessed with naming tasks). Particular attention was given to distinguishing causal findings from correlational results.

# Common names

## A large-scale distributed network

Numerous studies have sought to elucidate which brain areas are responsible for concept-to-word mapping, using various methodological approaches. These includes functional<sup>22,28,64-68</sup> and structural<sup>69-72</sup> neuroimaging, lesion-symptom mapping<sup>44,50,51,73-75</sup> and electrical stimulation,<sup>65,76,77</sup> all of them involving diverse patient populations (such as those with neurodegenerative diseases, epilepsy/ temporal lobectomy, tumours or strokes) and healthy subjects.<sup>28,66,78</sup> More marginally, a few behavioural studies, notably conducted with patients affected by posterior cortical atrophy, have also provided relatively valuable insights.<sup>79-81</sup>

The generated findings have indicated that lexical retrieval/ lexical production predominantly engages a leftward cortical network, primarily located along the ventral language stream. This network encompasses key temporal structures,

particularly the inferolateral and ventro-basal regions,<sup>22,44,51,64,67,70,72,77,82,83</sup> which mostly concern their mid-to-posterior parts, occasionally involving the occipito-temporal areas.<sup>28,74,77</sup> It also includes the posterior superior<sup>67,72,76</sup> and middle temporal areas,<sup>50</sup> further extending into temporo-parietal/inferior parietal regions.<sup>22,65,67,75,82</sup> Additionally, it involves the frontal operculum<sup>22,65,66,73,78,82</sup> and, in some cases, parts of the middle frontal gyrus.<sup>66,84,85</sup> Significantly, this widely distributed fronto-temporo-parietal cortical network is robustly supported by findings from direct electrical stimulation (DES) studies conducted intraoperatively using a picture naming task,<sup>84-86</sup> which particularly underscore the causal role of the posterior part of the superior temporal gyrus. Moreover, the importance of the hippocampus in visually guided naming following anterior temporal lobectomy has been highlighted in patients with intractable epilepsy, provided that the hippocampus is not sclerotic before the surgery.<sup>87-90</sup> Consistently, meta-analyses of functional neuroimaging studies<sup>91</sup> and of lesion-symptom mapping investigations<sup>75</sup> generally align with these studies, placing an emphasis on the mid-to-posterior segments of the temporal lobe (see Fig. 1).

# Central temporal cortical areas

## The mid-to-posterior ventral temporal regions

The region commonly known as the ‘basal temporal language area’ (BTLA), whose the location is mainly assigned to the mid-to-posterior part of the fusiform (corresponding to Brodmann’s area [BA] 37),<sup>51,93,94</sup> serves as a central hub within the cortical network responsible for integrating semantic features with lexical labels. Pioneering electrostimulation studies<sup>95,96</sup> paved the way for subsequent research, with recent studies consistently confirming that disruption of the BTLA leads to anomia. This has been evidenced through other inhibitory cortical stimulation<sup>65</sup> (see also<sup>77</sup>) together with voxel-based lesion-symptom mapping (VLSM) studies related to surgical resection procedures.<sup>44,51,83,93</sup> Functional neuroimaging studies have further associated naming deficits with tissue dysfunction in this area<sup>22,94</sup> (see also<sup>65</sup>). Furthermore, Hillis et al.<sup>23</sup> demonstrated that a few days after a stroke, the restoration of blood flow in the left BA37 was linked to the recovery of lexical access.

## The anterior temporal structures

Moreover, functional neuroimaging,<sup>28,68</sup> lesion mapping<sup>28,54,68</sup> and behavioural<sup>59,97,98</sup> studies, sometimes combined with structural/morphometric MRI analyses<sup>99-102</sup> have repeatedly linked the left anterior temporal structures (ATS) with semantically driven lexical access (sometimes for a given semantic category), in various patient populations. However, naming failures resulting from ATS damage seem more frequently associated with semantic disturbances, at least evident in language production.<sup>73,99,103-107</sup> As such, the issue of the ATS’s precise involvement in lexical retrieval remains steeped in controversy, particularly within the
A

Proper name retrieval

Correlational evidence

<figure><img src="image_3.png" /></figure>

Causal evidence

<figure><img src="image_4.png" /></figure>

B

Common name retrieval

Correlational evidence

<figure><img src="image_5.png" /></figure>

Causal evidence

<figure><img src="image_6.png" /></figure>

**Figure 1** The cortical network implicated in lexical access to (A) PNs and (B) CNs. The regions reported are those the most commonly identified in studies conducted in both patients and healthy subjects. Causal evidence (right panel) is differentiated from correlational evidence (left panel), highlighting the potential greatest implication of anterior versus posterior temporal regions in PN and CN retrieval, respectively. The figure does not include exhaustive information on medial areas potentially significant for PN retrieval, which encompass the medial section of fronto-temporo-parietal regions including the orbitofrontal regions,²⁸,²⁹ the anterior cingulate cortex,²⁸,²⁹,⁹² the anterior parahippocampal gyrus²⁸ and the posterior cingulate/retrosplenial cortex.⁹² The functional central hubs are indicated in bold. ATS, anterior temporal structures; BTLA, basal temporal language area; IFG, inferior frontal gyrus; IPL, inferior parietal lobe; MTG, middle temporal gyrus; STG, superior temporal gyrus; TP, temporal pole; TPJ, temporo-parietal junction.

framework of the ‘hub-and-spoke’ model (see also⁵¹,¹⁰⁸), which preferentially emphasizes the role of bilateral ATS in modality-independent semantics,¹⁰⁹,¹¹⁰ including in the context of cortical stimulations.¹¹¹

## Synthesis

Taken together, a range of multi-modal studies has demonstrated that lexical retrieval of CNs primarily depends on the left ventral language pathway (including fronto-temporo-parietal regions), within which temporal regions play a critical role, especially with regard to the mid-to-posterior part of the ventro-basal temporal cortex and of the inferolateral one to a lesser extent (i.e. the BTLA), but also to the ATS, albeit in a likely less specifically way.

## Proper names

### A large-scale distributed network

Large scale lesion-deficit mapping study, mostly based on cerebrovascular patients, and supplemented with functional neuroimaging,²⁸,⁶⁸ have provided compelling evidence that the retrieval of PNs depends on a mainly left lateralized neural network. In particular, Damasio *et al*.²⁸ identified the temporal pole (TP) and the neighbouring anterior inferior temporal and anterior parahippocampal cortex, as well as the ‘pars orbitalis’ of the inferior frontal gyrus, as critical structures for PN retrieval. The anterior cingulate gyrus and the orbitofrontal areas may also contribute.²⁸,²⁹ The involvement of fronto-temporal regions has received
further support from DES studies⁴⁹ and PET/functional MRI (fMRI) research involving healthy subjects.⁶² Generally, the neural network of PN retrieval encompasses both medial and lateral fronto-temporal regions²⁸,²⁹,⁴⁹,⁶²,¹¹² and, occasionally, extends into the posterior middle temporal cortex and the temporo-parietal junction (see Fig. 1).¹¹³

## Central temporal cortical areas

### The key role of the anterior temporal structures in lexical-semantic processing of unique entities

Within this extensive neural network, specific sectors of the left ATS, including the TP, serve as central hubs for retrieving the names of famous individuals from faces.⁴³ According to Damasio *et al.*'s theory,²⁸,¹¹⁴ the ATS function as a 'convergence zone' for processing unique entities. This implies that they may index inputs originating from modality-dependent regions and establish connections in a bidirectional manner, with verbal labels referring to known unique entities (involving both the mapping from knowledge of unique entities to their names, and the retrieval of knowledge about these entities from their names¹¹⁵).

Overall, the ATS involvement in lexical retrieval, especially for PNs, finds robust support in an array of neuropsychological studies associated with fMRI/PET investigations.²⁸,²⁹,⁵²,⁵³,⁶⁸,¹¹⁴ These findings have been consistently substantiated by other advanced lesion-symptom mapping studies⁴³,⁵⁴ and by voxel-based morphometry¹¹⁶ together with neuromodulation⁶⁰,¹¹⁷ and electrophysiological works.⁶¹ They are also echoed in neuropsychological behavioural research, frequently conducted in patients with epilepsy/who underwent surgery to treat drug-resistant temporal epilepsy.⁵²,⁵⁷⁻⁵⁹,¹¹⁸ Overall, the role of the left-biased ATS includes lexical access for various categories such as famous faces,²⁸,⁴²,⁴³,⁶⁰,¹¹⁶,¹¹⁸ landmarks,⁶²,¹¹⁴ speaking voices⁶⁰,⁶¹,¹¹⁹ and famous melodies with or without lyrics.¹¹²,¹²⁰

### Challenging the specific role of the anterior temporal structures in unique entities processing

The 'hub-and-spoke' model challenges the understanding of the ATS's role in the processing of unique entities and PN retrieval, as particularly driven by specific semantic or grammatical categories. No special emphasis is neither placed on the left ATS's role in lexical access. Indeed, this theory rather assumes that the ATS involve bilaterally high-level semantic integration regardless of the process involved, encompassing all classes of objects. In this alternative framework, the ATS constitute a graded transmodal representational system⁵⁶,¹⁰⁹,¹¹⁰,¹²¹ enabling conceptualization process, by establishing functional connections with modality-dependent association areas.¹²² Furthermore, the ATS, especially its ventrolateral part/the anterior fusiform gyrus, is not merely conceived as a binding zone. Rather, it is suggested to support, through experience, the building of nonlinear transmodal and coherent generalizable conceptual

knowledge.¹⁰⁹,¹¹⁰ Moreover, this framework assumes that the potential functional differences between verbal and non-verbal semantically related processes, associated with the left versus the right ATS, are not attributable to a lateral specialization of the ATS *per se*.⁵⁶,¹¹⁰

It holds true that injury to the left ATS typically results in anoxia without significant semantic impairment, as evidenced by various neuropsychological studies.³⁰,¹⁰⁸,¹²³ In line with this, other works conducted in healthy subjects demonstrated that transcranial magnetic stimulation (TMS)-related virtual lesioning of the left ATS increased naming latencies.¹²²,¹²⁴,¹²⁵ Nonetheless, this pattern is not understood as linked to the ATS's hemispheric specialization but interpreted in the light of two main factors: the bilateral organization of the ATS, which confers increased resilience to unilateral neurological damage for semantic processing and an asymmetric connectivity strength leading to a left-ward hemispheric linguistic bias.¹⁰⁹,¹¹⁰,¹²⁶ In a similar vein, there may be reciprocally, a lateralized functional connectivity for nonverbal stimuli, in particular faces, into the right ATS.¹²⁷

### The hypothesis of the anterior temporal structures' sensitivity to an effect of high semantic specificity

We have previously discussed the challenges that the 'hub-and-spoke' model presents to a theory suggesting that the ATS are sensitive to naming effects, especially based on semantic/grammatical categories. Some researchers have thus rather assumed that the ATS's involvement may vary, depending on the specificity level of processing demands required by a given lexical-semantic task (i.e. greater implication with highly specific requirements).³⁰,¹²⁴,¹²⁸,¹²⁹

This proposition is grounded in few studies, which found an ATS's implication in tasks involving visual-semantic distinctions at a specific level. As such, significant results were obtained by Rogers *et al*.¹²⁸ in their category verification paradigm, where participants indicated whether the pictorial input matched a conceptual category (corresponding to either animals or vehicles) at three levels of specificity (general, basic and specific). In this PET functional imaging study, the authors found a significant relationship between the ATS activation and categorization at a specific level (e.g. 'Labrador'), as opposed to an intermediate (e.g. 'dog') and general (e.g. 'animal') levels, without category and/or domain effects. The hypothesis of the ATS's sensitivity to a specific effect has also been supported by at least one study¹²⁴ which showed a significant repetitive TMS (rTMS) slowing naming effect in healthy subjects, but only at a specific level. Moreover, neuropsychological evidence derived from semantic dementia¹²⁸,¹²⁹ further suggested that this neurological condition, which affect the ATS, impairs semantic performance by a characteristic specificity effect. Indeed, Rogers *et al*.¹²⁹ showed that performance in semantic dementia patients and to a lesser extent, in stroke patients, across several semantic tasks (picture naming, picture sorting, word-picture matching and fluency) with high specific demands, was
significantly worse than performance assessed at basic or gen-
eral levels.

Critically, Rogers et al.¹²⁸ found that the same sectors of
the ATS typically damaged in semantic dementia (impacting
their semantic performance with a specificity effect¹²⁹) and
implicated by the specific level of classification in healthy
subjects, were activated during a FFN task.⁵⁵ Accordingly,
the ATS may be involved in processing stimuli from a large
array of categories rather than unique entities exclusively,
particularly under more stringent conditions, in terms of le-
vel of classification. In this sense, the ATS activation re-
sponded selectively to famous faces/names, compared to
other categories, notably named at a basic level.⁵⁵ This sug-
gests that the unique entities may be distinguished from com-
mon objects only to the extent that the latter are generally
treated at a basic level, implying a more coarse-grained pro-
cessing than the former, treated at a specific-level. In this line,
a study showed that epileptic patients who had undergone
anterior temporal lobectomy exhibited naming impairments
for both faces and objects, although for object naming, this
impairment tended to be present only when reaction time
was measured or when the complexity of the tasks increased
(e.g. probing less familiar concepts)³⁰ (see also ¹³⁰).
Accordingly, the ‘hub-and-spoke’ theory posits that both
processing of PNs and CNs depend on the same semantic sys-
tem and are susceptible to be impaired in case of neurological
damage to the ATS. However, due to the finer distinctions re-
quired by the former, lexical-semantic processing might be
prone to deficit more salient for PNs than for CNs.

The mid-to-posterior ventral temporal regions

Interestingly, electrophysiological findings have suggested
the involvement of the posterior part of the fusiform gyrus
in naming famous faces, based on intracranial recordings,
in three cases of epileptic patients.⁶¹ Likewise, a PET study
conducted in few healthy subjects has suggested the possible
implication of the mid-posterior basal temporal region
(BA37) in lexical access for all categories (except colours), in-
cluding famous faces.⁵⁵ However, it is worth noting that re-
cent VLSM findings conducted in a relatively wide cohort of
patients affected by a low-grade glioma, did not show the
critical implication of the mid-to-posterior ventral temporal
regions in FFN retrieval, contrary to the ATS (including the
TP).⁴³ Thus, it seems that the posterior ventro-basal tem-
poral regions are generally not essential for PN retrieval (at
least for faces) though they may have a contributory role.
Finally, PN retrieval might rely in part on cortical structures
already shown to be crucial for CN retrieval, just as some
studies have reciprocally suggested an involvement of the
ATS in CN retrieval.¹⁰¹ However, further direct evidence
for this claim is actually lacking.

Synthesis

The cortical network underpinning PN retrieval mainly
comprises left fronto-temporal regions, occasionally ex-
tending into mid-posterior ventro-temporal regions and,

to a lesser extent, into the temporo-parietal junction.
Concerning key temporal regions, the ATS seem represent
a critical functional epicenter of this network while the
posterior temporal regions likely have a contributory
role. However, the specific role of the ATS in processing
unique entities, including PN retrieval, has been ques-
tioned by the ‘hub-and-spoke’ model.

The contributing roles of
cortical regions beyond the
two temporal hubs

Generally, due to the multitude of processes they imply,
naming abilities rely on a widespread neural system. As a
result, naming failures can arise due to other reasons
than lexical access impairment, which primarily involves
lesion to temporal structures.⁷⁵ As such, damage to the
fronto-parietal regions/the dorsal language stream
(mediated by the arcuate fasciculus) can lead to phono-
logical paraphasia (or anomia) whereas damage to the
fronto-temporal regions/the ventral semantic stream
(mediated by the IFOF), can result in semantic paraphasia
(or anomia),⁶ at least for CNs. Otherwise, the disruption of
the occipito-temporal network¹³¹ (including the posterior
part of the ILF)¹³² can, in turn, provoke visual recognition
impairments, possibly leading to visual paraphasias.
Likewise, damage to the neural systems involved in motor
speech output, speech initiation and control, especially
those connecting the supplementary motor area to the
caudate nucleus (via the fronto-striatal tract) or to the pos-
terior inferior frontal gyrus (via the frontal aslant tract),
may cause speech dysfluencies during naming¹³³,¹³⁴ and
articulatory errors.²² Further, damage to the inferior front-
al regions are likely to contribute to an executive deficit in
selecting information from competing lexical-semantic
alternatives.⁷³,¹³⁵⁻¹³⁸

In this vein, a PET study conducted in patients with a
probable Alzheimer disease aimed to isolate the mechanisms
underlying both picture naming and verbal fluency.¹³⁹ This
study showed that metabolic activity in inferior frontal re-
gions was associated with naming performance facilitated
by phonemic cueing (among other regions), and with phono-
logical fluency abilities (in isolation). In contrast, spontan-
eous naming performance was associated with activity in
inferior temporal structures, while semantic fluency
abilities involved both frontal and temporal regions. The
authors thus suggested that naming failures may result
from dysfunction in the inferior frontal regions, disrupting
controlled retrieval processes. On the other hand, dysfunc-
tion of the temporal structures may lead to lexical-semantic
impairments, affecting naming through a different mechan-
ism. Notably, such functional imaging findings are especially
informative in the neuropsychological context of dissociac-
tions observed in Alzheimer's patients between semantic
and phonological fluencies.¹⁴⁰
In a similar manner for PNs, the lateral prefrontal cortex/ the inferior frontal cortex may be engaged in the cognitive control aspects of lexical retrieval for PNs, as suggested by an fMRI study using a covert naming paradigm in older adults<sup>92</sup> and by another one using an associative face/name learning task in healthy subjects.<sup>141</sup> Lastly, the medial frontal regions, encompassing the orbitofrontal and anterior cingulate cortex, may be associated with the socio-emotional salience related to faces and could facilitate face naming.<sup>26,28,29</sup>

## Synthesis

We have previously shown that lexical retrieval processes primarily rely on temporal regions, although broader neural networks come into play to ensure naming efficiency. Consequently, naming impairments may arise from other causes than a pure lexical retrieval deficit. In line with this view, damage to various regions within the fronto-temporo-parietal regions can lead to naming failures.

## Summary and conclusion

At the cortical level, the retrieval processes associated with CNs and PNs appear to rely on a partially wide overlapping neural network, centred on temporal structures with extensions into frontal and temporo-parietal/inferior parietal regions (see Fig. 1 for a summary illustration of the cortical bases underpinning lexical access to CNs and PNs). Within these overlapping subnetworks, the BTLA and the ATS (especially the TP) emerge as pivotal components, likely assuming more specific roles for CN and PN retrieval, respectively. In agreement with this, evidence from VLSM studies conducted with glioma patients further showed that anoma for CNs resulted from damage to the mid-to-posterior ventral temporal cortex<sup>44</sup> (see also<sup>142,143</sup>) whereas anoma for PNs is associated with damage to mid-to-anterior ventro-temporal structures.<sup>42,43</sup>

Thus, it could be that the brain specifically distinguishes, within a unified lexical access process, two grammatical classes due to additional cognitive demands required by PNs (i.e. unique psycholinguistic features, specific-level naming, greater memory resources, socio-emotional factors), compared to CNs. Nonetheless, it is not excluded that some CNs, especially those which are highly specific, atypical, unfamiliar, infrequent and/or very abstract, might be retrieved with the same neural resources as those used for PNs.

On the other hand, the ATS encompass an array of anatomical structures, each characterized by a unique connetional architecture to modality-specific cortical areas.<sup>144</sup> Hence, it is conceivable that various subregions within the ATS may be associated with the processing of different conceptual/grammatical categories, in relation to distinct input modalities, and even distinct lexical/semantic processes.

Indeed, the ATS have been the subject of multiple investigations, revealing a multi-faceted organization. In this sense, Rice *et al.*<sup>145</sup> have notably demonstrated in an fMRI study that, beyond a localized ventral category/modality-invariant subregion, other anterior temporal areas are gradually specialized, based on person and social concepts. Accordingly, the dorsolateral and polar regions of the ATS may be specialized in some aspects of social cognition.<sup>144,146-150</sup> Further, some studies have delineated a progressive auditory/visual division within the ATS, aligned along a superior to inferior axis.<sup>149,151,152</sup> As a note of caution, it is important to recall that, before the methodological advances in fMRI data processing that enhanced signal quality in the ventral sectors of the ATS,<sup>153</sup> these regions were particularly vulnerable to magnetic susceptibility artefacts, which could hinder the detection of their activation.<sup>152,154,155</sup> This limitation should be taken into account when discussing older fMRI studies focusing on the ATS.

As a second note of caution, it should be emphasized that each neurological disease tends to affect only a specific set of brain regions. For instance, stroke injuries typically involve the territories of the middle cerebral and posterior cerebral arteries, while structures situated in the anterior temporal region or on the medial surface of the brain are rarely affected. Conversely, diseases such as semantic dementia or fronto-temporal dementia, which predominantly affect the ATS and the ventral prefrontal cortex, tend to spare the occipital, posterior temporal and parietal areas. In this context, lesion aetiologies and their distribution are crucial factors to consider when evaluating the various forms of evidence, particularly in relation to CN/PN retrieval. This also underscores the importance of conditions that frequently damage the ventral temporal cortex, especially glioma or temporal epilepsy surgery, as valuable sources of information for this topic.

In this initial section, we mainly discussed the involvement of cortical structures in lexical retrieval, which constitutes only a fraction of the narrative. A comprehensive understanding of this cognitive process at the broader cerebral scale necessitates a thorough investigation of white matter connectivity. This is of paramount significance in informing anatomical and neuro-computational models of language, particularly in unravelling the anatomical foundations of anoma, given that white matter connections are severely compromised across a large spectrum of neurological conditions.

## Exploring ventral connectivity: implications for proper name and common name retrieval

As comprehensively reviewed in the preceding sections, cortical areas within the ventral stream play a pivotal
role in the process of lexical access. Our current focus
shifts to delineating the white matter tracts that intercon-
nect these cortical areas, the anatomical connectivity
being increasingly recognized as vital building blocks
for functional network integration.

From an anatomical perspective, the ILF runs longitudin-
ally along the ventral aspect of the temporal lobe, originating
from the occipital and occipito-temporal regions and termin-
ating in various areas of the ATS.<sup>156-161</sup> Recent investiga-
tions propose a complex, multi-layered fibre organization,
encompassing three to four distinct branches, including
fusiform, dorsolateral, lingual and potentially cuneal
branches.<sup>162,163</sup> In contrast, other studies suggest the
presence of shorter connections, considered separate from
the ILF (although this distinction is debatable), between
various areas of the mid-to-posterior fusiform gyrus
and the ATS.<sup>164,165</sup> Consequently, the ILF is strategically
well-positioned to establish connections between the
mid-to-posterior ventral temporal regions (frequently asso-
ciated with the BTLA) and the ATS. Thus, it appears to be
a prime candidate for conveying lexical retrieval in this ven-
tral neural network. Compelling empirical substantiation
now firmly establishes that the left ILF plays a discernible
role in both CN<sup>44,45</sup> and PN retrieval,<sup>42,43</sup> especially in vis-
ual modality, as expounded further below. Notably, this ex-
tended functional role is not mutually exclusive with its
potential involvement in semantic processing.<sup>161</sup> A schemat-
ic illustration of all functional implications attributed to the
ILF is given in Fig. 2 and an overall view of the anatomical
position and cortical terminations of the ILF is provided
in Fig. 3.

The role of the inferior longitudinal fasciculus in common name retrieval

In accordance with its anatomical course and cortical con-
nections to areas crucial for lexical retrieval (Fig. 3), namely
the mid/posterior lexical-semantic interface and, to a lesser
extent, the ATS, there is a growing body of experimental evi-
dence supporting the central role of the left ILF in the access
to lexical representations. Thus, a large-scale neuropsycho-
logical study combining VLSM and tract-focused analyses
was conducted with a sample of 110 patients having under-
gone surgical resection for a left low-grade glioma. In this
study, it was revealed that anomia for CNs, while controlling
for semantic disturbances (phonological errors being virtual-
ly absent), was linked to damage to the mid-to-posterior por-
tion of the inferior temporal gyrus (extending to fusiform
and middle temporal structures; see also<sup>142,143</sup>), in which
the ILF is known to have cortical endpoints. Further and
most importantly, the degree of residual tumour infiltration
into the left ILF emerged as a robust predictor of this
phenomenon.<sup>44</sup>

This observation aligns with the outcomes of a DES study,
wherein it was unequivocally demonstrated that stimulation
of the left ILF resulted in pure anomia for CNs, albeit only
when the ATS remained free of tumoural infiltration.⁴⁵
Conversely, in scenarios where the ATS were impacted by
low-grade gliomas, the ILF ceased its typical function, pos-
sibly rerouting its role via alternative pathways,⁴⁵ in accord-
ance with the well-documented neuroplasticity potential of
the ATS, especially in the pathophysiological context of
slow-growing tumours.¹⁶ This overarching pattern of find-
ings has recently been reaffirmed through a longitudinal
multi-modal study that integrated DTI tractography and in-
volved neurosurgical patients affected by a glioma.⁴⁸ The
authors of this study further showed that, prior to surgery,
impaired performance in picture naming was primarily
linked to damage of the mid-to-posterior part of the left
ILF. Conversely, when the infiltration of the ILF did not re-
sult in a deficit, it was typically its anterior part that was da-
maged, often in association with the cortical ATS.
Subsequent to surgery, naming impairments exhibited a ro-
bust correlation with the extent to which the ILF had been
resected, albeit exclusively in cases where the ATS and the
ILF had not been infiltrated prior to surgery. In the same
way, it is noteworthy to mention the case reported by
Herbet et al.,⁴⁵ which detailed the experience of a single pa-
tient who developed severe and enduring anomic aphasia fol-
lowing surgical transection of the ILF (neither it nor the ATS
were significantly infiltrated preoperatively), an interven-
tion, necessitated to access a tumour located within the left
temporo-mesial structures.

Another study, mapping the relationship between neuro-
logical lesions from diverse aetiologies and picture naming
performance, has also consistently emphasized the causal
role of the left ILF in lexical access to CNs (and not to
PNs).⁵⁴ Moreover, neuropsychological studies using tract-
wise lesion analysis have unveiled causal links between the
disruption of the left ILF and impaired object naming per-
formance in individuals with brain tumours, either
preoperatively¹⁴² or postoperatively.¹⁴³,¹⁶⁶ Additional in-
direct evidence is derived from studies, which have coupled
DTI metrics with language performance assessments, across
various patient populations exhibiting difficulties in naming.
These populations include individuals with primary progres-
sive aphasia (logopenic variant),¹⁶⁷ stroke¹⁶⁸,¹⁶⁹ and tem-
poral epilepsy or epilepsy-related temporal surgery.¹⁷⁰ In
general, these studies have identified non-specific associa-
tions between ILF DTI metrics (primarily fractional
anisotropy)¹⁶⁷⁻¹⁷⁰ and object naming performance.

Synthesis

In short, the existing literature highlights the central role of
the left ILF in object naming, likely related to the process of
CN retrieval. Notably, the most direct and compelling evi-
dence originates from studies involving patients with brain
tumours. This is unsurprising given that stroke injuries
typically have a limited impact on ventro-temporal
<figure><img src="image_7.png" /><figcaption>**Figure 2** Putative functional pathways supported by the multi-layered fibre organization of the ILF. The summary illustration is derived from literature data regarding the ILF's functions, primarily based on neuropsychological findings. Thicker lines and larger circles represent functional pathways biased towards a particular hemisphere. Neuropsychological syndromes resulting from ILF disruption include: (1) pure alexia; (2) written comprehension disorder; (3) aperceptive prosopagnosia; (4) associative prosopagnosia; (5) impairments in emotion recognition or hypo-emotionality; (6) scene perception impairments; (7) visual memory impairments; (8) semantic and/or lexical retrieval impairments; and (9) visual agnosia. AMG, amygdala; ATL, anterior temporal lobe; FFA, fusiform face area; Hip, hippocampus; LOC, lateral occipital cortex; OFA, occipital face area; pITG, posterior inferior temporal gyrus; PPA, parahippocampal place area; VWFA, visual word form area. The figure is reused from Herbet et al.¹⁵⁸</figcaption></figure>

structures, in contrast to lower-grade gliomas. Interestingly, studies involving glioma patients have further shown that the lesion of the ILF can be compensated for in lexical retrieval but uniquely when its anterior part is infiltrated along with the ATS, suggesting the

recruitment of specific brain plasticity mechanisms. Finally, current evidence regarding the implication of other ventral tracts in lexical retrieval remains limited. In the following section, we will demonstrate how the ILF implication extends to PNs.
<figure><img src="image_8.png" /></figure>

**Figure 3** Associative white matter tracts supporting lexical retrieval to CNs and/or PNs. This figure highlights associative white matter tracts presumed to play a role in lexical retrieval for both CNs and PNs. The strength of evidence varies across the tracts: robust neuropsychological evidence supports the role of the ILF in picture-guided lexical retrieval for both CNs and PNs, while the role of the UF is more uncertain, with a potential implication in PN retrieval. The involvement of the MdLF in lexical retrieval remains hypothetical at this stage. The main cortical terminations of the tracts under scrutiny are indicated. ATS, anterior temporal structures; BTLA, basal temporal language area; Fusi, fusiform gyrus; IFG, inferior frontal gyrus; IOG, inferior occipital gyrus; IPL, inferior parietal lobe; ITG, inferior temporal gyrus; OFC, orbitofrontal cortex; STG, superior temporal gyrus.

# The role of the uncinate and inferior longitudinal fasciculi in proper name retrieval

The available data concerning the connectional anatomy of PN retrieval is limited. Seminal work in this regard was initially presented by Papagno et al.⁴⁶ over a decade ago, and subsequently complemented by additional neuropsychological investigations,⁵⁴,¹⁷¹ including ours.⁴²,⁴³ We delve into these reports below.

## The uncinate fasciculus

Employing a longitudinal experimental design involving patients undergoing surgical resections for gliomas, Papagno et al.⁴⁶ observed a decline in PN retrieval performance, as evaluated through a FFN task, after the removal of the left UF. This C-shaped fronto-temporal tract interconnects the ATS with orbital, ventro-lateral and anterior frontal regions¹⁵⁶,¹⁵⁹,¹⁷²-¹⁷⁴ (for a review, see Von der Heide et al.¹⁷⁵) via a multi-layer fiber organization (see Fig. 3).¹⁷²,¹⁷³ Importantly, in Papagno et al.'s⁴⁶ study, the impairment in FFN was more pronounced when the temporal part of the UF was affected, compared to its frontal part. Further analyses indicated that, in patients with UF resection, anomia for PNs did not fully recover, even 9 to 12 months post-surgery (in contrast to anomia for CNs).⁴⁷ Furthermore, a large-scale lesion-symptom study involving predominantly stroke patients demonstrated that anomia for PNs was predicted by the extent of damage to the left UF (and not to the ILF).⁵⁴ In a more anecdotal context, a progressive decline in PN retrieval ability (but not in CN retrieval) was observed during the surgical resection of the left UF in two patients operated on while awake for temporal

gliomas, and in a third patient during the UF stimulation, using stereo-electroencephalography.¹⁷¹

## The inferior longitudinal fasciculus

The involvement of the left UF in PN retrieval has been called into question by two recent studies involving patients with lower-grade gliomas.⁴²,⁴³ In the first study, a retrospective assessment was conducted in 58 patients in the chronic post-surgical phase (i.e. at least 3 months after surgery) to evaluate their ability to retrieve PNs from the mental lexicon, utilizing a FFN task.⁴³ Performance was analysed through a multi-modal approach that included location-based between-group analyses of resection areas, VLSM and disconnection-symptom mapping. These comprehensive analyses also considered various factors influencing FFN performance. Specifically, sociodemographic variables (i.e. age and educational attainment) were included in 'Model 1', associated with FFN task-related control factors (i.e. self-estimated familiarity with each seen face and biographical knowledge) in 'Model 2' or with interrelated cognitive processes (non-verbal semantic processing, verbal episodic memory and CN retrieval) in 'Model 3' while all factors were combined in 'Model 4'. In summary, the patient group showed significantly poorer performance compared to a group of 72 matched healthy participants. VLSM results revealed that damage to the mid-to-anterior part of the inferolateral and ventro-basal temporal cortex (including the TP) was particularly predictive of impaired PN retrieval. Most notably, surgery-related disruption of the left ILF was robustly associated with FFN scores, as indicated by disconnection-symptom mapping (i.e. tract-by-tract correlation analyses and spatial correlations), across all tested models. In a separate double-case study, this research⁴³ finally demonstrated that the interruption of the right ILF in a left-handed patient
resulted in a severe and long-lasting PN retrieval deficit, while the same interruption in a right-handed patient did not cause such an impairment. This finding showed that damage to the right ILF can lead to crossed anomia for PNs in left-handed individuals, reinforcing the main conclusions of this study.

It is noteworthy that the left UF did not emerge as a contributor to PN retrieval impairment, in this work.<sup>43</sup> Specifically, the resection location-based between-group analyses conducted in this study indicated that anomia for PNs could occur following damage to the ILF while sparing the UF entirely. Indeed, patients with surgical resections centered on the ATS extending to neighbouring temporal structures (i.e. both UF and ILF damaged) as well as those with resections in the mid-to-posterior inferior temporal gyrus (i.e. ILF exclusively damaged) had significantly lower FFN scores than patients who underwent resections in the dorsal frontal areas (i.e. where both ILF and UF were fully spared) or in the fronto-temporo-insular areas (i.e. with extensive UF damage and sparsely, minimal ILF damage) but no other differences have been observed between these groups.

In the second study, Burkhardt *et al.*<sup>42</sup> presented three patient cases, all affected by a low-grade glioma involving the mid-to-anterior portion of the temporal lobe. Employing the same FFN task as in the previous study,<sup>43</sup> a longitudinal assessment was conducted by the first author (E.B.) in these patients, at three time-points: 1 day pre-surgery, 4 days post-surgery and 3 months post-surgery. This evaluation was complemented by a comprehensive neuropsychological examination. Importantly, all patients exhibited an enduring decline in PN retrieval performance following the surgical procedure. This decline could not be attributed to a lack of familiarity with famous faces nor to insufficient biographical knowledge. Of note, only one patient, B.S., continued to experience persisting difficulties in retrieving CNs, thereby underscoring the specificity of the PN retrieval deficit in the two remaining patients, A.H. and C.K., in the context of their lexical access abilities. Notably, the neuropsychological examination of A.H. remained completely free for difficulties in other cognitive functions assessed, particularly verbal episodic memory. From an anatomical perspective, detailed disconnection analyses revealed that the ILF was the only white matter tract consistently disrupted across all three patients. Specifically, these analyses pinpointed the ventrolateral segment of the ILF, projecting into the implicated anterior ventrolateral temporal cortex.

## Contrasting the involvement of the inferior longitudinal fasciculus and the uncinate fasciculus

The two aforementioned studies represent a notable leap forward in the understanding of the connectional anatomy underpinning lexical access to PNs, at least for famous faces. The proposition of the central role of the ILF stands in contrast to prior reports, which had rather suggested the involvement of the left UF.<sup>46,54</sup> This discrepancy in findings

may be attributed to differences in methodological approaches. The studies by Papagno *et al.*<sup>46</sup> and Mehta *et al.*<sup>54</sup> did not consider the interplay of various cognitive processes, which may be involved in PN retrieval. Consequently, the correlations observed between UF damage and PN retrieval deficits in these studies may also reflect the influence of other participating processes.

Indeed, the integrity of the UF has been associated with various cognitive domains, including semantic processing<sup>168,176,177</sup> and episodic memory.<sup>178,179</sup> In this sense, some findings indicate that the microstructural properties of this tract correlate with associative learning performance, notably in tasks involving face-name pairing.<sup>180,181</sup> In this respect, Alm *et al.*<sup>180</sup> suggested that the UF implication in associative learning tasks may be related to its role in episodic memory, potentially under stringent conditions of retrieval competition. In this context, the faces and names employed in their experimental task were indeed designed to be closely similar and frequent respectively, in order to increase the difficulty of distinguishing between possible retrieval candidates.<sup>180</sup> This possible implication of the UF seems broadly consistent with its projections in the inferior frontal cortex, whose role in the executive component of selecting the lexical-semantic target among concurrent alternatives<sup>138,141,182</sup> is well known. Notably, competition is likely minimal in the context of retrieval labels relatively unique, necessarily specific and rather infrequent compared to CNs, named at a basic level. However, the integrity of such a control mechanism might be critical in PN retrieval, taking into account the highly precise and individualized word selection process required within a homogeneous perceptual-semantic contexts. On the other hand, several works proposed a role for the left UF in socio-emotional functions,<sup>144,175</sup> including emotional valence-based mnemonic associations,<sup>175</sup> possibly oriented towards face-name processing. Once again, that seems congruent with its cortical terminations in both the ATS (implicated in mnemonic association of person-related information)<sup>175</sup> and the face-selective region of the orbitofrontal cortex, which is implicated in social motivation processing.<sup>183</sup>

Upon closer examination, it is likewise essential to acknowledge that the study by Papagno *et al.* did not experimentally investigate the involvement of other anatomical connections, notably the ILF.<sup>46,47</sup> In contrast, anatomical analyses in Burkhardt *et al.*<sup>43</sup> found an implication of the left ILF controlled for the implication of other tracts, including the UF. Furthermore, the ILF was the unique tract damaged in common across the three patients, in Burkhardt *et al.*<sup>42</sup> Lastly, it is worth noting that in the study conducted by Mehta *et al.*,<sup>54</sup> the ILF primarily incurred damage to its anterior segment. This was attributable to the characteristics of the patient population, consisting mainly of cerebrovascular lesions and less frequently anterior temporal lobectomies for epilepsy. Disentangling the distinct contributions of the ILF and the UF in this context becomes exceedingly challenging, as both tracts terminate in the ATS (for the ILF<sup>163</sup>; for the UF<sup>173</sup>). This challenge is also evident in the study by
Koay et al.,¹⁷¹ where PN retrieval deficits following stimulation of the white matter within the ATS were attributed to a virtual lesion of the left UF.

## Synthesis

To date, only a handful of studies have examined the anatomical connectivity underlying PN retrieval. Some neuropsychological findings have suggested that the left UF may play a critical role in this process. However, more recent evidence instead highlighted the central role of the left ILF, not that of the UF. Several methodological considerations could explain these divergent results. Nonetheless, the processes potentially linking the UF to the retrieval of PNs have been discussed.

## Summary and conclusion

Taken as a whole, the most up-to-date literature corroborates the central role of the left ILF in both CN and PN retrieval processes, contrary to that of the UF. It aligns with the cortical implications discussed in the first part of this review, particularly the finding that the posterior and anterior ventral temporal cortex are predominantly involved in CN and PN retrieval, respectively. Indeed, the ILF is known to project into both of these regions, forming a coherent cortico-subcortical system for lexical retrieval with distinct specificities. However, it is important to emphasize that clearly dissociating the precise contributions of the cortex versus white matter tracts remains highly challenging, as both are often simultaneously and indistinguishably affected in most neurological conditions. Furthermore, standard lesion-symptom mapping methodologies are still suboptimal for accurately delineating the pathological variances associated with each type of structure. In this context, DES of white matter tracts represents a promising approach to dissociate the contributions of ventral tracts to the retrieval of CNs and PNs. This direction is both scientifically and clinically justified, as evidence suggests that patients undergoing inferior temporal regions resection along with ILF damage, do not recover from aphasic anomia,⁴⁴ significantly impacting their quality of life. In brief, there is a clear need for well-designed and adequately powered studies to further discern the precise functional implications of the ILF fibres in naming, especially from the cortical contributions.¹⁸⁴

In the final section of this review, we present hypothetical models outlining dissociated patterns of ventral connections, which underlie CN and PN retrieval respectively, seeking to elucidate the complex network of regions and pathways contributing to these essential language processes. Thus, we propose several promising avenues for future research, aiming to further refine our comprehension of the neural mechanisms governing naming processes, depending on input modality and/or semantic category (at least for PNs for this latter).

# Hypothetical white matter architecture supporting lexical access to common names and proper names

## Separate pathways for common name and proper name retrieval?

In light of the increasingly established role of the left ILF in both CN⁴⁴,⁴⁸ and PN⁴²,⁴³ retrieval, a significant question arises: do separate white matter components within the ILF serve to transmit psycholinguistic information necessary for naming entities belonging to each of these grammatical categories? This hypothesis gains relevance in the context of reports highlighting dissociations between anomia for CNs and PNs resulting from various brain lesions,²⁵,²⁶ and of findings from VLSM studies that have precisely located the critical cortical locus for PN retrieval as being more anterior along the ventrolateral temporal axis,⁴³ compared to CN retrieval.⁴⁴ To substantiate this hypothesis, empirical evidence is imperative, demonstrating that well-defined, selective damage to the left ILF yields either anomia for CNs or anomia for PNs, whereas more extensive lesions result in a shared lexical retrieval deficit. In support of this, recent research has provided consistent evidence showing that damage to the ventrolateral fibres of the ILF along with the antero-ventral temporal cortex, led to pure anomia for PNs without concurrent anomia for CNs (at least as assessed by a standard picture-based object naming task), in two out of the three patients studied.⁴² Notably, the third patient, who exhibited anomia for both CNs and PNs, had the largest lesion.

## Segregated fibres for semantic categories-dependent proper name retrieval?

As widely discussed above, previous studies have strongly implicated the ILF in the retrieval network of FFN.⁴²,⁴³ Remarkably, these findings parallel the involvement of the right ILF in face recognition/face memory, as demonstrated in both healthy individuals¹⁸⁵⁻¹⁸⁸ and prosopagnosic patients¹⁸⁹,¹⁹⁰ (see also¹⁹¹ for a comprehensive review). In this context, it is essential to consider whether ILF's involvement in PN retrieval is specific to certain semantic categories, possibly limited to faces. Against it, existing research has already associated the ATS—in which the rostral fibres of the ILF project—with landmark name retrieval,⁶²,¹¹⁴ extending their role beyond the processing of person-specific information,¹⁹² as part of the ‘extended system’ for face stimuli.¹⁹³ Moreover, considering the specialization of the fusiform and parahippocampal regions for face¹⁹³⁻¹⁹⁶ and spatial/ place (including landmarks, scenes and topographical features) processing¹⁹⁷⁻¹⁹⁹ respectively, the hypothesis that
distinct components of the left ILF may be selectively in-
volved in PN retrieval, depending on the semantic category,
becomes especially relevant. Indeed, given its caudal projec-
tions into the mid-to-posterior fusiform and parahippocam-
pal structures, the ILF's role in face (for the former) and
landmark (for the latter) name retrieval, warrants further in-
vestigation. Along these lines, recent studies have identified,
in the right hemisphere, distinct white matter pathways that
interconnect these face- and place-selective areas with the
ATS¹⁶⁴,²⁰⁰ (see also¹⁶⁵ for face areas), and whose micro-
structural properties have been associated with recognition
performance for each stimulus type.¹⁶⁴ It is therefore plaus-
ible that similarly segregated connections within the left ven-
tral temporal structures play a role in the retrieval of face
names and landmark names, respectively.

A putative role of the middle longitudinal fasciculus in auditory-based lexical retrieval?

As comprehensively reviewed by Herbet *et al.*<sup>158</sup> and ex-
pounded upon by Zemmoura *et al.*,<sup>161</sup> the ILF represents a
central white matter tract involved in cognitive and behav-
ioural processes, grounded in the visual modality. This align-
ment between the tract's connective properties and its
functional implications is evident in its projections to asso-
ciative visual areas and its involvement in tasks such as read-
ing, visual recognition as well as lexical access, based on
object pictures<sup>44,45</sup> and famous faces.<sup>42,43</sup> Currently, there
is no conclusive evidence that the ILF mediates lexical re-
trieval when processing information in the auditory modal-
ity. While a few isolated studies have all the same hinted at
its potential involvement in certain auditory-derived func-
tions, such as naming from aural descriptions<sup>170</sup> or auditory-
verbal comprehension,<sup>161</sup> this remains an area in need of fur-
ther investigation.

From an anatomical viewpoint, the middle longitudinal
fasciculus (MdLF) seems better placed to convey the
auditory-linguistic information critical for efficient lexical
retrieval of CNs and PNs. Through a multi-layered
organization,<sup>201-203</sup> the MdLF interconnects the ATS with
parietal and occipital regions. More specifically, it passes
through the superior temporal structures/transverse gyri,
projecting rostrally into the anterior part of the superior tem-
poral gyrus (particularly in the dorsal TP) and posteriorly
into the inferior and superior parietal lobules, as well as
occipito-parietal and occipital areas.<sup>201-208</sup> Notably, it has
terminations in the primary and associative auditory cor-
tex.<sup>201,202</sup> The MdLF is displayed in Fig. 3.

In view of its connectivity pattern, it has been hypothe-
sized to play a role in sound and auditory language process-
ing.²⁰⁹ It might also contribute to coding sublexical
representations into articulatory forms, to supporting as-
pects of acoustic-phonetic word processing²¹⁰ and acting
as an interface between auditory representation and seman-
tic/lexical access.²⁰¹ Nevertheless, the precise connective

features—including components, cortical projections, and
interindividual variability—of the MdLF have not been fully
determined, and studies assessing its possible role in aspects
of language processing are limited.²¹¹

One electrostimulation mapping study investigated the
potential implication of the MdLF in language processing
in eight patients undergoing neurosurgery for low-grade gli-
omas.²¹² No language interference was observed after stimu-
lation of the left MdLF and no permanent deficits were found
after surgical resection of its anterior part. Nonetheless, it
does not inform about the potential implication of its poster-
ior part. Additionally, the assessment of naming was only
performed in the visual modality (through picture naming),
which does not rule out a potential role of the MdLF in audi-
tory naming (based on verbal descriptions, sounds, or
voices).

Otherwise, the MdLF and the ILF share a close anatomical
relationship, characterized by their projections into the TP
and a parallel course along the temporal lobe and then
into the occipital lobe.²⁰¹,²⁰² However, the ILF follows
a more ventral and lateral trajectory compared to the
MdLF. Consequently, its anterior projections target the
ventral TP, while those of the MdLF terminate in the
dorsal TP (Fig. 3).²⁰¹⁻²⁰⁴ This spatial arrangement highlights
the possible distinct roles of the ILF and the MdLF in
mediating visual and auditory language processing,
respectively.⁵⁶

As a point of emphasis, beyond the transmodal role of the
left ATS in PN retrieval/naming people from either faces or
voices,⁶⁰,⁶¹ some of their sectors are thought to consist of dis-
tinct modality-dependent regions responsive to auditory, vis-
ual and audiovisual stimuli in the dorsal, ventral, and polar
parts, respectively.¹⁴⁹ This aligns, in a certain way, with
the connectivity-constrained ‘hub-and-spoke’ theory, which
posits that modality-specific semantic information is chan-
nelled through spokes within the graded transmodal
ATS.¹¹⁰ Consequently, visual and auditory semantic pro-
cessing may proceed independently within the inferior and
superior temporal gyri, before converging within the ATS,
through a graded integration.⁵⁶,¹⁵² In the context of naming
unique entities, electrophysiological evidence indeed indi-
cated that surrounding and more posterior modality-
dependent areas within the ventro-basal temporal cortex
and the superior temporal gyrus, exhibit selective responses
to faces and voices, respectively.⁶¹ Interestingly, this finding
mirrors the neuropsychological evidence regarding person
recognition disorders from faces and voices, associated to
the right hemisphere.²¹³ Overall, these functional implica-
tions suggest dissociated roles for the left ILF and MdLF
in lexical(-semantic) processing, depending on the input
modality.

In summary, the results presented here suggest that the left
MdLF may play a role in integrating psycholinguistic-related
information from the auditory modality into the ATS, there-
by contributing to auditory-based lexical retrieval, whether
for PNs or CNs. Damage to the left MdLF might thus result
in an auditory modality-specific anomia, potentially selective
to voices. This neuropsychological impairment, known as ‘phonoanomia’, has notably been described in a recent case study by Moretta and Trojano.²¹⁴

## Summary and conclusion

In this section, we proposed the existence of distinct ILF fibre pathways dedicated to CN and PN retrieval, respectively. We then posited that, in the context of PN retrieval, specific components of the left ILF may be selectively involved in retrieving face names and landmark names, given its caudal terminations in cortical areas specialized for these perceptual-conceptual categories. Finally, we hypothesized that the MdLF might contribute to auditory-guided naming, particularly in lexical retrieval of CNs and PNs. This final section allowed us to elaborate on several anatomical hypotheses regarding the neural organization of white matter pathways underpinning lexical access, based on grammatical and semantic categories and depending on the input modality. The ILF, MdLF (and UF) are represented in Fig. 3.

## General conclusion

In this article, we conducted a comprehensive review of the brain network responsible for CN and PN retrieval, shedding light on the cognitive processes that distinguish these two grammatical categories. While CN and PN retrieval exhibit psycholinguistic differences, primarily in terms of mapping relationships between words and their referents (i.e. a one-to-one mapping between meaningless PNs and their unique associated entities versus a one-to-many mapping between meaningful CNs and the category-level objects named), we highlighted that an overlapping cortical network, situated along the ventral language stream, supports lexical access to both CNs and PNs. Overall, this network encompasses fronto-temporo-parietal regions, though each area may not hold equal importance in the retrieval of the two grammatical categories. In this sense, the BTLA (i.e. the mid-to-posterior part of the ventral temporal structures) and the ATS, two pivotal functional hubs within this network, appear to play relatively greater roles in CN retrieval for the former, and in PN retrieval for the latter.

At the structural connectivity level, our review emphasized the critical role of the left ILF in visually guided lexical access to CNs and PNs. Disruption of this white matter tract thus leads to a persistent anomia for both grammatical categories. Recent findings have especially highlighted the ILF's paramount role in PN retrieval and challenged the long-held hypothesis of the left UF, at least as the main neural pathway implicated in PN retrieval. In light of these new findings, we proposed several anatomical hypotheses regarding the connectional architecture underlying PN and CN retrieval. These hypotheses aim to stimulate and guide empirical investigations, so future research can deepen our understanding of the complex neural mechanisms governing lexical access.

In brief, our review underscores the ILF's implication in lexical access to CNs and PNs in the visual modality, supporting an overlapping cortical network, likely centred around different functional epicentres along the temporal structures. In this way, our review significantly updates the dual-pathway model for language.⁶,¹¹,¹⁵ Furthermore, by proposing that disruption of distinct anatomical connectional networks may entail anomia for CNs or PNs —taking into account PN subcategories and input modality irrespective of the name's grammatical class— our review opens promising avenues for future research. Our hypotheses indeed aim to stimulate and guide empirical investigations in order to enrich the understanding of the complex neural mechanisms governing lexical access.

Beyond these fundamental insights, significant clinical applications are anticipated. Indeed, they contribute to improved practices in awake surgeries for brain tumours and may support functional mapping in epilepsy, enabling better identification and preservation of eloquent structures involved in PN and CN retrieval. More generally, our review paves the way for a better understanding of neuropsycholinguistic profiles of patients, affected by various neurological pathologies targeting the temporal lobe, such as epilepsy, tumours and related surgeries but also with dementia, strokes, and others. Finally, these findings have implications for language rehabilitation, as many neurologically related language disorders are predominantly associated with white matter connectivity damage, not only in patients undergoing brain tumour surgery but also in those recovering from strokes, the most common cause of aphasia.²¹⁵ Hence, identifying white matter lesions associated with anomia, in relation to the grammatical category and the input modality, may represent a promising avenue for refining clinical care related to speech-language therapy.

## Funding

Eléonor Burkhardt received a PhD fellowship from la ‘Ligue contre le cancer’ French foundation.

## Competing interests

The authors report no competing interests.

## Data availability

Data sharing is not applicable to this article as no new data were created or analysed in this study.

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