Stephanie Forkel

Publications

Displaying 1 - 8 of 8
  • Dulyan, L., Guzmán Chacón, E. G., & Forkel, S. J. (2025). Navigating neuroanatomy. In J. H. Grafman (Ed.), Encyclopedia of the human brain (2nd ed.). Amsterdam: Elsevier.

    Abstract

    This chapter introduces the origins and development of our current anatomical terminology. It scrutinizes the historical evolution and etymological significance of the over 1900 official anatomical terms in the current nomenclature, underscoring their impact on the contemporary comprehension of cognitive neuroanatomy. The chapter traces unification efforts from the Basel Nomina Anatomica in 1895 to the 1998 Terminologia Anatomica, noting challenges arising from outdated terminology in light of recent anatomical advancements.

    Highlighting the influence of terminologies on interpretations of brain anatomy, the chapter explores several anatomical mapping methods such as surface, sectional, connectional, and functional anatomy. It illuminates discrepancies and controversies, exemplified by divergent interpretations of the number of brain lobes and the definitions of 'Broca' and 'Wernicke' areas.

    The chapter explores anatomical terms' historical and cultural underpinnings, encompassing mythonyms, eponyms, and cultural influences on nomenclature. It critically examines the implications of these terminologies on contemporary research and shows that Large Language Models mirror these discrepancies. It underscores the need for more inclusive and culturally sensitive approaches in anatomical education.

    Lastly, we advocate for updating anatomical nomenclature, suggesting that a deeper understanding of these terminologies could provide insights and aid in resolving ongoing debates in the field. This examination sheds light on historical knowledge and emphasizes the dynamic interplay between language, culture, and anatomy in shaping our comprehension of the neurobiology of the brain and how we navigate neuroanatomy in the 21st century.
  • Dulyan, L., Bortolami, C., & Forkel, S. J. (2025). Asymmetries in the human brain. In C. Papagno, & P. Corballis (Eds.), Cerebral Asymmetries: Handbook of Clinical Neurology (pp. 15-36). Amsterdam: Elsevier.

    Abstract

    The human brain is an intricate network of cortical regions interconnected by white matter pathways, dynamically supporting cognitive functions. While cortical asymmetries have been consistently reported, the asymmetry of white matter connections remains less explored. This chapter provides a brief overview of asymmetries observed at the cortical, subcortical, cytoarchitectural, and receptor levels before exploring the detailed connectional anatomy of the human brain. It thoroughly examines the lateralization and interindividual variability of 56 distinct white matter tracts, offering a comprehensive review of their structural characteristics and interindividual variability. Additionally, we provide an extensive update on the asymmetry of a wide range of white matter tracts using high-resolution data from the Human Connectome Project (7T HCP www.humanconnectome.org). Future research and advanced quantitative analyses are crucial to understanding fully how asymmetry contributes to interindividual variability. This comprehensive exploration enhances our understanding of white matter organization and its potential implications for brain function.
  • Forkel, S. J., Bortolami, C., Dulyan, L., Barrett, R. L. C., & Beyh, A. (2025). Dissecting white matter pathways: A neuroanatomical approach. In F. Dell'Acqua, M. Descoteaux, & A. Leemans (Eds.), Handbook of Diffusion MR Tractography (pp. 397-421). Amsterdam: Elsevier.

    Abstract

    The brain is the most magnificent structure, and we are only at the cusp of unraveling some of its complexity. Neuroanatomy is the best tool to map the brain's structural complexity. As such, neuroanatomy is not just an academic exercise; it serves our fundamental understanding of the neurobiology of cognition and improves clinical practice. A deepened anatomical understanding has advanced our conceptual grasp of the evolution of the brain, interindividual variability of cognition in health and disease, and the conceptual shift toward the emergence of cognition. For the past 20 years, diffusion imaging tractography has dramatically facilitated these advances by enabling the study of the delicate networks that orchestrate brain processes (for review, see Thiebaut de Schotten and Forkel, 2022). Several steps are consistent across all studied populations and brain states (health/disease) when analyzing tractography data. We discuss various considerations for dissections across populations and give practical tips on common pitfalls and features to improve the visualization of the dissections. We briefly discuss specific considerations for manual dissections in nonhuman primates. Lastly, we provide an atlas of regions of interest (ROIs) for the most commonly delineated white matter connections in the human brain.
  • Satoer, D., Dulyan, L., & Forkel, S. J. (2025). Oncology: Brain asymmetries in language-relevant brain tumors. In C. Papagno, & P. Corballis (Eds.), Cerebral Asymmetries: Handbook of Clinical Neurology (pp. 65-87). Amsterdam: Elsevier.

    Abstract

    Brain tumors are classified as rare diseases, with an annual occurrence of 300,000 cases and account for an annual loss of 241,000 lives, highlighting their devastating nature. Recent advancements in diagnosis and treatment have significantly improved the management and care of brain tumors. This chapter provides an overview of the common types of primary brain tumors affecting language functions—gliomas and meningiomas. Techniques for identifying and mapping critical language areas, including the white matter language system, such as awake brain tumor surgery and diffusion-weighted tractography, are pivotal for understanding language localization and informing personalized treatment approaches. Numerous studies have demonstrated that gliomas in the dominant hemisphere can lead to (often subtle) impairments across various cognitive domains, with a particular emphasis on language. Recently, increased attention has been directed toward (nonverbal) cognitive deficits in patients with gliomas in the nondominant hemisphere, as well as cognitive outcomes in patients with meningiomas, a group historically overlooked. A patient-tailored approach to language and cognitive functions across the pre-, intra-, and postoperative phases is mandatory for brain tumor patients to preserve quality of life. Continued follow-up studies, in conjunction with advanced imaging techniques, are crucial for understanding the brain's potential for neuroplasticity and optimizing patient outcomes.
  • Nozais, V., Forkel, S. J., Petit, L., Talozzi, L., Corbetta, M., Thiebaut de Schotten, M., & Joliot, M. (2023). Atlasing white matter and grey matter joint contributions to resting-state networks in the human brain. Communications Biology, 6: 726. doi:10.1038/s42003-023-05107-3.

    Abstract

    Over the past two decades, the study of resting-state functional magnetic resonance imaging has revealed that functional connectivity within and between networks is linked to cognitive states and pathologies. However, the white matter connections supporting this connectivity remain only partially described. We developed a method to jointly map the white and grey matter contributing to each resting-state network (RSN). Using the Human Connectome Project, we generated an atlas of 30 RSNs. The method also highlighted the overlap between networks, which revealed that most of the brain’s white matter (89%) is shared between multiple RSNs, with 16% shared by at least 7 RSNs. These overlaps, especially the existence of regions shared by numerous networks, suggest that white matter lesions in these areas might strongly impact the communication within networks. We provide an atlas and an open-source software to explore the joint contribution of white and grey matter to RSNs and facilitate the study of the impact of white matter damage to these networks. In a first application of the software with clinical data, we were able to link stroke patients and impacted RSNs, showing that their symptoms aligned well with the estimated functions of the networks.
  • Parlatini, V., Itahashi, T., Lee, Y., Liu, S., Nguyen, T. T., Aoki, Y. Y., Forkel, S. J., Catani, M., Rubia, K., Zhou, J. H., Murphy, D. G., & Cortese, S. (2023). White matter alterations in Attention-Deficit/Hyperactivity Disorder (ADHD): a systematic review of 129 diffusion imaging studies with meta-analysis. Molecular Psychiatry, 28, 4098-4123. doi:10.1038/s41380-023-02173-1.

    Abstract

    Aberrant anatomical brain connections in attention-deficit/hyperactivity disorder (ADHD) are reported inconsistently across
    diffusion weighted imaging (DWI) studies. Based on a pre-registered protocol (Prospero: CRD42021259192), we searched PubMed,
    Ovid, and Web of Knowledge until 26/03/2022 to conduct a systematic review of DWI studies. We performed a quality assessment
    based on imaging acquisition, preprocessing, and analysis. Using signed differential mapping, we meta-analyzed a subset of the
    retrieved studies amenable to quantitative evidence synthesis, i.e., tract-based spatial statistics (TBSS) studies, in individuals of any
    age and, separately, in children, adults, and high-quality datasets. Finally, we conducted meta-regressions to test the effect of age,
    sex, and medication-naïvety. We included 129 studies (6739 ADHD participants and 6476 controls), of which 25 TBSS studies
    provided peak coordinates for case-control differences in fractional anisotropy (FA)(32 datasets) and 18 in mean diffusivity (MD)(23
    datasets). The systematic review highlighted white matter alterations (especially reduced FA) in projection, commissural and
    association pathways of individuals with ADHD, which were associated with symptom severity and cognitive deficits. The meta-
    analysis showed a consistent reduced FA in the splenium and body of the corpus callosum, extending to the cingulum. Lower FA
    was related to older age, and case-control differences did not survive in the pediatric meta-analysis. About 68% of studies were of
    low quality, mainly due to acquisitions with non-isotropic voxels or lack of motion correction; and the sensitivity analysis in high-
    quality datasets yielded no significant results. Findings suggest prominent alterations in posterior interhemispheric connections
    subserving cognitive and motor functions affected in ADHD, although these might be influenced by non-optimal acquisition
    parameters/preprocessing. Absence of findings in children may be related to the late development of callosal fibers, which may
    enhance case-control differences in adulthood. Clinicodemographic and methodological differences were major barriers to
    consistency and comparability among studies, and should be addressed in future investigations.
  • Forkel, S. J., & Catani, M. (2019). Diffusion imaging methods in language sciences. In G. I. De Zubicaray, & N. O. Schiller (Eds.), The Oxford Handbook of Neurolinguistics (pp. 212-228). Oxford: Oxford University Press.

    Abstract

    The field of neuroanatomy of language is moving forward at a fast pace. This
    progression is partially due to the development of diffusion tractography, which
    has been used to describe white matter connections in the living human brain.
    For the field of neurolinguistics this advancement is timely and important for
    two reasons. First, it allows clinical researchers to liberate themselves from
    neuroanatomical models of language derived from animal studies. Second, for
    the first time, it offers the possibility of testing network correlates of
    neurolinguistic models directly in the human brain. This chapter introduces the
    reader to general principles of diffusion imaging and tractography. Examples of
    its applications, such as tract analysis, will be used to explicate its potentials and
    limitations.
  • Thiebaut de Schotten, M., Friedrich, P., & Forkel, S. J. (2019). One size fits all does not apply to brain lateralisation. Physics of Life Reviews, 30, 30-33. doi:10.1016/j.plrev.2019.07.007.

    Abstract

    Our understanding of the functioning of the brain is primarily based on an average model of the brain's functional organisation, and any deviation from the standard is considered as random noise or a pathological appearance. Studying pathologies has, however, greatly contributed to our understanding of brain functions. For instance, the study of naturally-occurring or surgically-induced brain lesions revealed that language is predominantly lateralised to the left hemisphere while perception/action and emotion are commonly lateralised to the right hemisphere. The lateralisation of function was subsequently replicated by task-related functional neuroimaging in the healthy population. Despite its high significance and reproducibility, this pattern of lateralisation of function is true for most, but not all participants. Bilateral and flipped representations of classically lateralised functions have been reported during development and in the healthy adult population for language, perception/action and emotion. Understanding these different functional representations at an individual level is crucial to improve the sophistication of our models and account for the variance in developmental trajectories, cognitive performance differences and clinical recovery. With the availability of in vivo neuroimaging, it has become feasible to study large numbers of participants and reliably characterise individual differences, also referred to as phenotypes. Yet, we are at the beginning of inter-individual variability modelling, and new theories of brain function will have to account for these differences across participants.

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