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Gyrus

A gyrus (plural: gyri) is a ridge-like elevation of gray matter on the surface of the cerebral cortex, surrounded by depressions known as sulci, which together form the folded structure of the brain's outer layer.^[1] These structures are essential components of the cerebral cortex, the outermost layer of gray matter that covers the cerebrum and constitutes about half of the brain's total mass.^[2] Gyri play a critical role in maximizing the brain's computational capacity by dramatically increasing the surface area of the cerebral cortex—up to approximately 2,000 square centimeters in humans—without requiring a proportional enlargement of the skull, an adaptation that has evolved to support advanced cognition in mammals.^[1] This folding allows for a greater number of neurons, estimated at 14 to 16 billion in the cortex, enabling complex processing of information related to sensory perception, motor control, language, memory, reasoning, and consciousness.^[2] Structurally, each gyrus consists primarily of layers of nerve cell bodies and dendrites within the gray matter, with variations in size, shape, and precise layout occurring between individuals, contributing to unique brain architectures.^[1] Developmentally, gyri form through a process of cortical folding during fetal brain growth, driven by mechanical forces and genetic factors, which can sometimes lead to abnormalities associated with neurological conditions such as epilepsy or developmental delays if folding is excessive or disrupted.^[3] Notable examples include the precentral gyrus in the frontal lobe, which serves as the primary motor cortex responsible for initiating voluntary movements; the postcentral gyrus in the parietal lobe, functioning as the primary somatosensory cortex for processing touch and proprioception; and the superior temporal gyrus in the temporal lobe, which houses Wernicke's area critical for language comprehension.^[1] These specialized gyri, along with sulci, divide the cerebral cortex into functional lobes—frontal, parietal, temporal, and occipital—each contributing to distinct aspects of human behavior and cognition.^[2]

Anatomy

Definition and gross features

A gyrus is a ridge-like elevation or convolution on the surface of the cerebral cortex, representing the raised folds formed by the infolding of the brain's outer gray matter layer.^[1] These structures are integral to the macroscopic appearance of the cerebrum, where they constitute the prominent features visible upon gross examination of the brain.^[4] The term "gyrus" derives from the Latin word gyrus, meaning "circle" or "ring," which aptly describes the curved, encircling pattern these ridges often form in the cortical landscape.^[5] Gyri are predominantly found on the cerebral hemispheres, with notable examples including the precentral gyrus located along the posterior margin of the frontal lobe.^[6] They are separated by sulci, the shallower or deeper grooves that delineate their boundaries.^[7] In terms of gross morphology, gyri vary in size, rising as elevations between adjacent sulci, and their convoluted arrangement significantly expands the cortical surface area—by approximately 2.5 times relative to a hypothetical smooth cortex—as quantified by the gyrification index in human brains.^[8] This folding accommodates a total cortical surface of about 2,000 cm², about 60-65% of which (approximately 1,200–1,300 cm²) is concealed within sulcal depths.^[1]

Microscopic structure

The gyri of the neocortex are characterized by a distinct six-layered histological architecture, comprising layers I (molecular), II (external granular), III (external pyramidal), IV (internal granular), V (internal pyramidal), and VI (multiform).^[4] This layered organization is a hallmark of neocortical tissue, differentiating it from allocortical regions with fewer layers. Layer IV, in particular, functions as the primary zone for receiving thalamic afferent projections, integrating sensory and other inputs into the cortical circuitry.^[9] The cellular composition of gyral neocortex features a predominance of pyramidal neurons in layers III and V, which serve as the principal excitatory output elements projecting to other cortical areas and subcortical structures. Layer IV contains abundant granule cells, including stellate variants, that facilitate local processing of incoming signals. Glial cells, such as astrocytes and oligodendrocytes, are distributed across all layers, providing essential structural support, metabolic maintenance, and myelination of axons.^[4]^[10] Vascularization in gyral tissue includes dense capillary networks, particularly concentrated in the crowns of gyri relative to sulcal depths, to meet the high metabolic demands of neuronal activity; these capillaries arise primarily from branches of the cerebral arteries.^[11]^[12] The average thickness of the gyral cortex ranges from 2.5 to 3 mm, forming a thin but densely packed gray matter mantle overlying the thicker subcortical white matter.^[13] These microscopic attributes underlie the macroscopic appearance of gyri as elevated ridges on the cerebral surface.

Relation to sulci and fissures

Gyri represent the elevated ridges of the cerebral cortex, bounded laterally by shallower sulci and more deeply separated by fissures, which collectively form the convoluted surface of the brain. Sulci are narrow grooves that delineate the margins of individual gyri, while fissures, such as the central sulcus (also known as the fissure of Rolando), are deeper clefts that divide larger cortical regions, like the frontal and parietal lobes. This spatial arrangement allows gyri to protrude between these indentations, creating a folded topography that maximizes cortical packing within the cranial vault.^[4]^[14] The folding patterns of gyri exhibit diverse orientations, including transverse (e.g., Heschl's gyrus in the temporal lobe), longitudinal (e.g., superior frontal gyrus running parallel to the midline), and oblique (e.g., angular gyrus at an angle to the main axes), which interdigitate with corresponding sulci to produce the brain's characteristic convolutions. The ratio of gyral to sulcal surface area contributes significantly to cortical expansion, with about 60-65% of the total cortical surface area (around 1,200–1,300 cm² out of 2,000 cm² in adults) buried within sulci, effectively increasing the exposed surface by a factor reflected in the gyrification index of about 2.56. This configuration enhances the brain's computational capacity by accommodating more neural tissue without proportionally enlarging the skull.^[4]^[15] Functionally, gyri often encompass relatively homogeneous regions of cortical processing, such as the precentral gyrus dedicated to motor control, while sulci serve as natural boundaries demarcating transitions between distinct areas, exemplified by the central sulcus separating primary motor and sensory cortices. This organization facilitates modular neural architecture, where sulci not only provide physical separation but also align with cytoarchitectonic borders in many cases. In neuroimaging, gyri manifest as prominent ridges of intermediate signal intensity on T1-weighted MRI scans, attributable to the dense packing of gray matter, contrasting with the brighter underlying white matter and darker cerebrospinal fluid in sulci and fissures.^[16]^[4]^[17]

Development and variation

Embryonic and fetal development

The development of gyri in the human cerebral cortex begins during the embryonic period and intensifies in the fetal stage, marking the transition from a smooth lissencephalic surface to a folded gyrencephalic structure. Initial signs of cortical folding emerge around gestational weeks 10 to 15, as the first primary sulci indent the cortical surface, setting the stage for gyrus formation.^[3] By gestational weeks 24 to 28, primary gyri become more defined, with the cortex expanding rapidly to accommodate increased neuronal numbers and establish the foundational folding patterns observed in the mature brain.^[18] This timeline reflects a hierarchical process where early folds precede more complex secondary and tertiary structures later in gestation.^[19] Key mechanisms driving gyrification involve the tangential expansion of the cortical surface through proliferation of neural progenitors and the radial migration of neurons to their laminar positions. Tangential expansion arises from symmetric divisions of apical radial glial cells, increasing the number of radial units in the ventricular zone and thereby enlarging the cortical plate.^[20] Radial migration occurs along radial glial scaffolds, positioning postmitotic neurons in an inside-out manner to build cortical layers, which contributes to surface buckling under mechanical stress.^[21] Fibroblast growth factor (FGF) signaling plays a pivotal role in regulating progenitor proliferation, particularly in the outer subventricular zone, where it promotes the expansion of basal progenitors essential for gyrencephalic folding.^[22] Disruption of FGF pathways leads to reduced cortical surface area and impaired gyrus formation, underscoring its upstream influence on these processes.^[23] Genetic factors critically influence gyrification, with mutations in specific genes disrupting neuronal migration and resulting in abnormal folding. Mutations in the LIS1 gene (also known as PAFAH1B1), which encodes a regulator of microtubule dynamics, cause classical lissencephaly characterized by a near absence of gyri due to arrested radial migration of neurons.^[24] Similarly, mutations in the DCX gene, encoding doublecortin—a microtubule-associated protein—lead to subcortical band heterotopia and pachygyria, where gyri are broadly simplified or lacking, reflecting defective neuronal positioning during corticogenesis.^[25] These genetic disruptions highlight the dependence of gyrus formation on precise cytoskeletal regulation during the fetal period.^[26] During mid-gestation, specific fetal milestones illustrate the progressive emergence of major gyri, often driven by axonal tension and connectivity patterns. The calcarine sulcus, delineating the future cuneus and lingual gyri, first appears around gestational weeks 16 to 22, becoming fully formed as one of the earliest primary folds.^[27] This development is influenced by axonal tension from growing thalamocortical and callosal fibers, which exert mechanical forces to indent the cortex and promote localized gyrus elevation.^[28] Such milestones establish the basic architecture of visual and association cortices before late-gestational refinements.^[29]

Postnatal maturation and individual differences

The postnatal maturation of cerebral gyri involves dynamic structural changes that continue well after birth, building on embryonic foundations to refine cortical folding patterns. In the first two years of life, gyri undergo rapid expansion, driven by intense increases in cortical surface area and volume, alongside progressive myelination that thickens underlying white matter tracts and supports enhanced connectivity.^[30] This phase is marked by a surge in gyrification, with a notable discontinuity at birth contributing to nearly two-thirds of postnatal folding growth, as observed in MRI studies of preterm and term infants.^[31] By the end of this period, white matter maturation patterns, including denser axon packing and reduced extra-axonal water, align with gyral development, particularly in superficial layers corresponding to subplate remnants.^[32] These changes stabilize progressively through childhood, with cortical thinning shifting from primary sensory-motor regions to multimodal areas, reaching relative stability by early adolescence.^[30] Individual differences in gyral maturation manifest prominently in hemispheric asymmetry, particularly in language-related regions. The left hemisphere typically exhibits larger gyri in areas such as the planum temporale and Heschl's gyrus, reflecting structural bases for language lateralization, with leftward surface area asymmetries averaging +0.176 in the planum temporale.^[33] Sex differences further contribute to variability, with males often displaying more pronounced folding through greater sulcal depth and overall cortical surface area, while females show higher gyral complexity in regions like the superior temporal cortex.^[34] These patterns emerge early and persist, influencing developmental trajectories without altering the core timeline of maturation.^[35] Population-level variations in gyral patterns are evident across ethnic groups, as revealed by MRI morphometry. For instance, East Asian individuals demonstrate greater cortical thickness in the bilateral superior frontal gyrus compared to Caucasians, potentially reflecting differences in gyral architecture adjusted for age and sex.^[36] Such ethnic disparities in frontal gyral metrics, including thickness and surface area, highlight genetic and environmental influences on postnatal refinement.^[37] Aging introduces gradual degenerative changes to gyri, beginning in the fourth decade with subtle neuronal loss of 2–4% that accelerates thereafter. This leads to cortical atrophy at rates of 0.2–0.5% per year in brain volume post-35, manifesting as gyral thinning and sulcal widening—approximately 0.7 mm per decade—due to expanded cerebrospinal fluid spaces and reduced gray matter density.^[38] By the 50s and beyond, gyrification declines by 0.035 per decade, with pronounced effects in multimodal association areas, underscoring the impact of cumulative neuronal attrition on cortical morphology.^[38]

Function

General physiological role

Gyri play a crucial role in expanding the surface area of the cerebral cortex, enabling a greater number of neurons to be accommodated within the confines of the skull. This folding mechanism results in approximately a 2.5-fold increase in cortical surface area compared to an unfolded state, supporting around 16 billion neurons in the human cerebral cortex.^[39]^[40] By forming ridges that interdigitate with sulci, gyri maximize neural packing density while maintaining a compact overall brain volume, which is essential for efficient information processing.^[41] In terms of signal integration, gyri facilitate parallel processing by segregating neural inputs across their elevated surfaces, allowing distinct functional modules to operate concurrently without excessive interference. This structural arrangement promotes the exchange of information between remote gyral regions and adjacent sulcal areas, enhancing the brain's capacity for multifaceted neural computations.^[42] Such segregation supports the cortex's ability to handle diverse sensory and cognitive inputs simultaneously, contributing to overall physiological efficiency. The physiological demands of gyri are heightened due to their dense synaptic connections, which drive elevated metabolic activity and oxygen consumption in gyral tissue. Synapses, concentrated in these folded regions, account for a significant portion of the brain's energy expenditure, with oxygen utilization reflecting the high energetic cost of maintaining and signaling through these neural junctions.^[43] This increased metabolic rate underscores the gyri's role in sustaining the cortex's intensive computational workload. Evolutionarily, gyri confer an advantage by correlating with enhanced cognitive complexity across mammals, evolving from minimal folds in rodents to intricate patterns in primates. This progression in gyrification parallels expansions in brain size and neural capacity, enabling more sophisticated behaviors and problem-solving abilities in higher mammals, including humans.^[44]^[45]

Contributions to neural processing

Gyri facilitate neural computation through distinct patterns of connectivity that integrate local and distant cortical regions. Intra-gyral short-range connections are primarily mediated by U-fibers, which are superficial association fibers linking adjacent gyri to support fine-grained cortico-cortical networks.^[46] Inter-gyral long-range connections, in contrast, rely on major white matter tracts such as the arcuate fasciculus, which links regions across gyri, including the superior and middle temporal gyri to perisylvian frontal areas, enabling broader integration of neural signals.^[47] A key principle in gyral contributions to neural processing is the localization of functional areas onto specific gyral structures, as delineated by Brodmann's cytoarchitectonic mapping. For instance, Brodmann area 4, corresponding to the primary motor cortex, occupies the precentral gyrus and is dedicated to motor execution through its projection to spinal motor neurons.^[48] This mapping ensures that gyri serve as modular platforms for specialized neural operations, aligning cytoarchitecture with anatomical folds. Signal propagation within gyri is optimized by their structural features, particularly the crowns, which receive thalamocortical connections that span sensorimotor to association regions and mature hierarchically during development.^[49] Cortical folding into gyri further enhances processing efficiency by minimizing the average path lengths between interconnected sites, thereby reducing signal propagation times compared to unfolded cortices.^[50] These attributes collectively support rapid and reliable neural transmission. Gyral circuits undergo refinement through synaptic pruning during childhood, a process that eliminates excess connections to streamline network efficiency and consolidate active pathways.^[51] This pruning aligns with gyrification changes, potentially influencing fold maturation by releasing tension in neuronal fibers and optimizing circuit specificity within gyral domains.^[52] Overall, gyri expand cortical surface area, thereby increasing the capacity for complex neural computations.

Clinical and research aspects

Associated disorders and imaging

Focal cortical dysplasia (FCD), characterized by abnormal gyral formation and scarring, represents the most common cause of drug-resistant focal epilepsy in children and young adults.^[53] This malformation often manifests as localized disruptions in cortical layering and neuronal migration within affected gyri, leading to epileptogenic foci that require precise identification for management.^[54] In Alzheimer's disease (AD), gyral atrophy begins prominently in the temporal regions, including the medial occipitotemporal, inferior, and middle temporal gyri, serving as an early predictor of progression from mild cognitive impairment to dementia.^[55] These volumetric reductions correlate with cognitive decline and are among the initial pathological changes observed in the disease course.^[56] Schizophrenia is associated with gyral simplification, evidenced by reduced cortical gyrification, particularly in the left medial superior frontal gyrus, reflecting underlying neurodevelopmental alterations.^[57] Diffusion tensor imaging (DTI) in these patients reveals reduced fractional anisotropy in gray and white matter microstructures, indicating disrupted fiber integrity and connectivity that may contribute to symptom severity.^[58] Magnetic resonance imaging (MRI) techniques, such as voxel-based morphometry (VBM), enable quantitative measurement of gyral volumes by analyzing gray matter density across the cortex, aiding in the detection of atrophy patterns in disorders like epilepsy and AD.^[59] Functional MRI (fMRI) further assesses activation patterns within specific gyri during cognitive tasks, revealing altered hemodynamic responses in regions like the prefrontal and temporal gyri that correlate with impaired executive function and memory.^[60] DTI complements these by quantifying fractional anisotropy to highlight microstructural changes beneath simplified gyri in schizophrenia.^[61] Recent advances in 7T MRI have improved resolution of sub-gyral layers, allowing visualization of early biomarkers such as tau accumulation in the entorhinal and temporal cortices of AD patients.^[62] These high-field scans detect disturbed cortical lamination and correlate iron deposition with tau pathology, facilitating earlier diagnostic interventions.^[63] Such imaging supports targeted surgical planning by delineating pathological gyral boundaries.^[64]

Surgical and evolutionary considerations

In epilepsy surgery, intraoperative electrocorticography (ECoG) is employed to map epileptogenic zones across gyral surfaces, guiding precise resection while minimizing disruption to surrounding cortical tissue.^[65] This technique involves placing electrode grids directly on the exposed cortex to record electrical activity, thereby delineating functional boundaries within gyri to optimize seizure control outcomes.^[66] Stereotactic navigation systems further enhance this process by integrating preoperative imaging with real-time intraoperative guidance, allowing surgeons to preserve critical gyral regions associated with motor, sensory, or language functions during resections.^[67] Awake craniotomy represents a key technique for identifying gyral boundaries in eloquent brain areas, where patients perform cognitive or motor tasks under local anesthesia to enable direct electrical stimulation mapping.^[68] This approach helps delineate the limits of functional cortex on gyral crests and margins, reducing postoperative deficits by confirming safe resection margins.^[69] Since 2015, advancements in neuronavigation, including integration with intraoperative MRI, have improved surgical precision in epilepsy procedures, leading to enhanced seizure freedom rates and fewer neurological complications compared to earlier methods.^[70]^[71] Cortical gyrification first emerged in early mammals around 200 million years ago, with ancestral species exhibiting relatively simple folding patterns to accommodate expanding neural tissue within a confined skull.^[72] This foundational complexity increased over evolutionary time, reaching human-like intricacy by the appearance of Homo erectus approximately 1.8 million years ago, coinciding with advanced cognitive demands such as sophisticated tool use and social behaviors.^[73]^[74] Endocasts from H. erectus fossils reveal prominent gyri and sulci, indicating a brain reorganization that supported enhanced manual dexterity and planning capabilities.^[73] Comparative anatomy highlights gyrification's variability across mammals, correlating with the encephalization quotient (EQ), a measure of relative brain size. Monotremes such as the platypus display lissencephalic (smooth) brains with minimal folding and low EQ values, reflecting simpler neural architectures suited to their ecological niches.^[75] In contrast, cetaceans like dolphins exhibit highly gyrified cortices with EQs approaching those of humans (often exceeding 4), enabling complex echolocation, social structures, and problem-solving through expanded cortical surface area.^[76]^[77] This gyrification-EQ link underscores how folding facilitates cognitive expansion in large-brained species.^[78]

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Awake brain mapping is considered the gold-standard for identifying and preserving those important areas of the brain during surgery.Missing: gyral | Show results with:gyral
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Awake Craniotomy for Gliomas in the Non-Dominant Right ...
Mar 15, 2024 · This article aims to comprehensively review the current literature on the benefits of awake craniotomy in gliomas of the non-dominant right hemisphere.
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Intraoperative MR imaging and neuronavigation during resection of ...
Aug 6, 2025 · Intraoperative MR imaging and neuronavigation during resection of FCD Type II in adult epilepsy surgery offers better seizure outcome.<|separator|>
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Epilepsy Surgery in the Last 10 Years: Advancements and ... - NIH
Jul 9, 2025 · One of the most notable advancements in epilepsy surgery over the last decade has been the rise of less invasive techniques, which have been ...
[72]
The Formation of Folds on the Surface of the Brain
May 5, 2017 · It is likely that the ancestral mammal, which lived around 200 million years ago, also had a folded brain. ... cortical folding. Mice with ...
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The primitive brain of early Homo - Science
Apr 10, 2021 · ... complex cognitive tasks, such as social cognition, tool use, and language. When these differences arose during human evolution is a question ...
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First 'Homo' species left Africa with ape-like brains | Live Science
Apr 8, 2021 · Researchers have since learned "that stone tools were made as early as 3.3 million years ago (well before the expected origin of Homo) and ...
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Distinct Development of the Cerebral Cortex in Platypus and Echidna
Dec 6, 2011 · Our findings indicate that gyrencephaly begins to emerge in the echidna brain shortly before birth (crown-rump length 12.5 mm), whereas the ...
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Cetacean brains: How aquatic are they? - Marino - 2007
May 21, 2007 · More specifically, brain size is often expressed as an encephalization quotient, or EQ (Jerison, 1973), which is a value that represents how ...Abstract · LARGE BRAINS AND HIGH... · NEUROANATOMICAL...
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Phylogeny and adaptive evolution of the brain-development gene ...
Apr 14, 2011 · Variation in absolute brain size, relative brain size (EQ = encephalization quotient), and body size in cetaceans in a phylogenetic context.
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Genetic Mechanisms Underlying Cortical Evolution in Mammals - PMC
The remarkable sensory, motor, and cognitive abilities of mammals mainly depend on the neocortex. Thus, the emergence of the six-layered neocortex in reptilian ...