Cortex Refresher

The surface of the brain, also known as the cerebral cortex, has a unique and intricate structure characterized by convolutions, or folds, that significantly increase its surface area (Azevedo et al., 2009). The simplest way to divide the cortex is into the frontal and posterior cortex. The frontal cortex (frontal lobe) specializes in action, ranging from cognition, emotion, and autonomic control to movements and speech. The posterior cortex (parietal, temporal, and occipital lobes) is concerned with perception and memory. The frontal and posterior cortex, subcortical structures and the peripheral nervous system, provide the hierarchically arranged feedback loops that allow us to interact with our environment to achieve goals successfully. Graphic © Gordenkoff/Shutterstock.com.

This post covers anatomical Orientations, Directional Terms, Cortical Features, The Unfixed Brain, Dissecting Brains, and the Cortical Lobes.

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Orientations

Three customary planes for viewing the body and brain are sagittal, coronal, and horizontal. The sagittal plane divides the body into right and left halves. The coronal plane separates the body into front and back parts. Finally, the horizontal (transverse) plane divides the brain into upper and lower parts (Breedlove & Watson, 2023). Graphic courtesy of Blausen.com staff "Blausen gallery 2014," Wikiversity Journal of Medicine.

Directional Terms

Important directional terms include medial (toward the middle) and lateral (toward the side), ipsilateral (same side) and contralateral (opposite side), superior (above) and inferior (below), anterior/rostral (toward the head), and caudal (toward the tail), proximal (near the center) and distal (toward the periphery, and dorsal (toward or at the back) and ventral (toward the belly) (Breedlove & Watson, 2023).

Cortex Features

The adult human brain has a volume of about 1100 square cm, weighs about 3 pounds, and requires convolutions to fit within the skull (Bear, Connors, & Paradiso, 2020). Placing your fists together roughly approximates the two cortical hemispheres' deceptively small size. Two-thirds of the cortical surface lies within these folds (Breedlove & Watson, 2023).

The complex structure of the human brain, characterized by its wrinkled, uneven surface, arises from the intricate folding of a thick layer of tissue known as the cerebral cortex. The cortex largely comprises the dendrites, cell bodies, and axons of neurons. Brain tissue is commonly identified as gray matter. When a cross-section of the brain is examined, the outermost layers of the cortex exhibit a darker shade due to a higher concentration of neuronal cell bodies and dendrites. In comparison, the underlying white matter is primarily made up of axons. It exhibits a lighter shade due to the presence of the white, fatty myelin that insulates many neuronal axons. Gray matter primarily receives and processes information, while white matter conveys information to other regions.

Anatomists distinguish three topographical features of the cerebral cortex: gyrus, sulcus, and fissure. Surface landmark graphic © Snapgalleria/Dreamstime.com.

A gyrus is a ridged area of the brain. The precentral gyrus, anterior to the central sulcus, is the primary motor cortex (controls muscles and movements). The postcentral gyrus, posterior to the central sulcus, is the primary somatosensory cortex (which receives somatosensory information).

A sulcus is a groove in the cortical surface. As we observed, the central sulcus separates the primary motor cortex from the primary somatosensory cortex. A fissure is a deep groove. The Sylvian fissure (also called the lateral fissure or lateral sulcus) is the upper boundary of the temporal lobe (Breedlove & Watson, 2023). Graphic courtesy of Blausen.com staff "Blausen gallery 2014," Wikiversity Journal of Medicine.

The Unfixed Brain

This video was produced by Suzanne Stensaas, PhD, Department of Neurobiology and Anatomy, and the Spencer S. Eccles Health Sciences Library, University of Utah.

Dissecting Brains

Cortical Lobes

The cortical lobes are named for the overlying skull bones (Breedlove & Watson, 2023). Graphic © Sebastian Kaulitzki/Shutterstock.com.

The cortex is required for executive functions like attention, planning, and problem-solving.

Without a cerebral cortex, a person would be blind, deaf, dumb, and unable to initiate voluntary movement (Bear, Connors, & Paradiso, 2016, p. 205).

The five major cortical regions include the frontal, parietal, temporal, and occipital lobes, and the insula (not shown). Cortical lobes graphic © Madrock24/Shutterstock.com.

Frontal Lobes

The frontal lobes (F7, F3, Fz, F8, F4) consist of the cortex anterior to the central sulcus and consist of the primary motor cortex, motor association cortex, Broca's area, and prefrontal cortex. Frontal lobe graphic Graphic © Kateryna Kon/Shutterstock.com.

Essential left frontal lobe functions include working memory, concentration, planning, and positive emotion. The main clinical concern is Major Depressive Disorder (MDD).

Critical right frontal lobe functions include declarative memory, social awareness, and negative emotions. The main clinical concerns include Generalized Anxiety Disorder (GAD), fear, and impaired executive functioning.

Frontal lobe damage may result in impaired flexibility and problem-solving, increased risk-taking, changes in social behavior, an inability to use external cues, and deficits in emotional self-regulation.

The primary motor cortex (precentral gyrus) is located in the precentral gyrus (Brodmann area 4, BA 4). It organizes the opposite side of the body's muscles and movements required for the fine motor coordination required by tasks like writing. Lesions can result in loss of motor control, including rigid paralysis. Graphic © Kateryna Kon/Dreamstime.com.

The motor association cortex (premotor cortex) is rostral to the primary motor cortex (BA 6) and helps program and execute movements. The motor association cortex is the piano player, and the primary motor cortex is the piano keyboard (Carlson & Birkett, 2016). The primary and motor association cortex collectively appear to map behaviors instead of specific muscles or movements (Breedlove & Watson, 2023).

Three interconnected primary motor cortex (M1) regions participate in the somato-cognitive action network (SCAN), which controls the integrated movement of multiple body parts (Gordon et al., 2023).

Broca's area, which is located in the inferior frontal gyrus (BA 44 and 45) of the dominant hemisphere (F7-T3 in the left hemisphere), is concerned with speech production, grammar, language comprehension, and sequencing (Caplan, 2006). Lesions to Broca's area can produce dyslexia, grammar, spelling, and reading deficits, and Broca's aphasia. Broca's area receives input from Wernicke's area via the arcuate fasciculus (Breedlove & Watson, 2023). Graphic courtesy of Blausen.com staff "Blausen gallery 2014," Wikiversity Journal of Medicine.

The prefrontal cortex (PFC) is rostral to the motor association area (BA 9, 10, 11, 12, 46, 47). It is responsible for executive functions, which involve attention, working memory, prediction of the outcomes of current and hypothetical actions, working toward goals, problem-solving, planning, and the ability to suppress actions that could lead to unwanted outcomes (Diamond, 2013). The PFC integrates emotion and reward in decision-making (Fuster, 2015).

Important subdivisions of the PFC include the orbitofrontal cortex (OFC), ventromedial prefrontal cortex (VMPFC), and dorsolateral PFC (DLPFC). They play important roles in reward and addiction.

The orbitofrontal cortex (OFC) comprises Brodmann areas 10, 11, and 47 (Kringelbach, 2005). The OFC may aid planning by evaluating our actions' consequences (rewards and punishments) and helping to motivate us to ingest drugs. Phineas Gage's profound personality changes were produced by damage to this subdivision and the VMPFC. The OFC appears to adjust decision-making based on the stakes involved, enabling us to switch between significant (investments) and trivial (snacks) choices. Finally, the OFC compares our current options with recent ones, while the anterior cingulate cortex registers our predictions and prediction errors (Kennerley et al., 2011).

The ventromedial prefrontal cortex (VMPFC) corresponds to the ventromedial reward network (Ongür & Price, 2000) and includes BA 10, 14, 25, 32, and parts of 11, 12, and 13. The VMPFC is implicated in making decisions where the outcomes are uncertain and moral values must be applied to actual situations. Patients with damage to the VMPFC choose outputs that lead to immediate reward, regardless of their future cost. They do not learn from their mistakes. Since they have difficulty understanding social cues, they may not recognize deception, irony, or sarcasm (Zald & Andreotti, 2010). Likewise, they may not control their emotional reactions in social situations, particularly anger, and violence (Carlson & Birkett, 2016).

The dorsolateral prefrontal cortex (DLPFC) is located in the middle frontal gyrus and includes BA 9 and 46. The DLPFC is a critical component of the executive network and shares responsibility with cortical and subcortical networks for functions like abstract reasoning, cognitive flexibility, decision-making, inhibition, planning, and working memory (Miller & Cummings, 2007). It exercises the highest cortical level of motor control (Hale & Fiorello, 2004).

The left DLPFC is concerned with approach behavior and positive affect. It helps us select positive goals and organizes and implements behavior to achieve them. The right DLPFC organizes withdrawal-related behavior and negative affect and mediates threat-related vigilance. It plays a role in working memory for object location. In unipolar depression and premenstrual dysphoric disorder, the right DLPFC may be more active than the left (alpha asymmetry).

The cingulate cortex has reciprocal connections with the parahippocampal gyri, integrates limbic functions, and is part of the salience network. Cingulate cortical functions include child nurturing, grooming, play, organization, and managing input/output functions.

The anterior cingulate cortex (ACC) (Fpz, Fz, Cz, Pz) lies above the corpus callosum (BA 24, 32, 33). The dorsal ACC is connected to both the PFC and parietal cortex. The ACC plays a vital role in attention and is activated during working memory. The ACC mediates emotional and physical pain, and has cognitive (dorsal anterior cingulate) and affective (ventral anterior cingulate) conflict-monitoring components.

The anterior cingulate is part of the affective network and is triggered when we make mistakes (Arnsten, 2009).

The Stroop test illustrates a cognitive monitoring task where color and names conflict. Discrepancies between facial and vocal cues show an affective conflict. The anterior cingulate recruits other brain areas to resolve these conflicts.

The anterior cingulate helps us allocate attention to focusing on a target and then disengaging, perceiving options, and making adaptive choices. The anterior cingulate gyrus, the prefrontal cortex, and the caudate function abnormally in children diagnosed with ADHD during selective attention tasks. fMRI evaluation showed that neurofeedback could teach children to normalize activity in these structures (Beauregard & Levesque, 2006).

The anterior cingulate gyrus is involved in motivation and perceiving emotional and physical pain. Eisenberger, Lieberman, and Williams (2003) used an fMRI to study the brains of participants who believed that two companions playing an online baseball simulation suddenly dropped them from the game. Their emotional distress activated the anterior cingulate cortex, which evaluates the unpleasantness of physical pain. de Charms and colleagues (2005) provided real-time fMRI feedback from the anterior cingulate to subjects. They learned to reduce its metabolism and the intensity of experimental pain.

Lesions to the cingulate can produce akinetic mutism, in which a person cannot produce orienting responses. Cingulate malfunction can result in addictive behaviors (alcohol or drug abuse, eating disorders, chronic pain), obsessive-compulsive disorder and OCD spectrum disorders, and “road rage.”

Parietal Lobes

The parietal lobes (Pz, P3, P4) are posterior to the frontal lobes (BA 1, 2, 3, 5, 7, 39, 40) and are divided into the primary somatosensory cortex and secondary somatosensory cortex. Their main function is to process somatosensory information like pain and touch. Parietal lobe graphic © Kateryna Kon/Dreamstime.

Major left parietal lobe functions include problem-solving, math, complex grammar, attention, and association. Essential right parietal lobe functions include spatial awareness and geometry.

The primary somatosensory cortex (S1) is located in the parietal lobe's postcentral gyrus posterior to the central sulcus (BA 3, 1, and 2). S1 maps touch and pain information from the opposite side of the body. The secondary somatosensory cortex (S2) is adjacent to S1 (BA 40 and 43), receives projections from it, and maps touch and pain from both sides of the body (Breedlove & Watson, 2023). Graphic by Paskari from Wikimedia Commons.

The primary function of the parietal lobes is to process somatosensory information like pain and touch. The parietal cortex monitors our preparation for a movement and is responsible for our subjective feeling of intending to move (Sirigu et al., 2004).

The angular gyrus near the superior temporal lobe (BA 39) is involved in reading, math, and copying writing. The graphic below highlights the angular gyrus in red © Kateryna Kon/Shutterstock.com.

Major left hemisphere functions include attention, association, complex grammar, math, object names, and somatosensation.

Major right hemisphere functions include body boundary, geometry, guiding reaching with the hands, somatosensation, and spatial perception (Demos, 2019).

Temporal Lobes

The temporal lobes (T3, T4, T5, T6) are separated from the rest of the cortical lobes by the Sylvian fissure (BA 15, 20, 21, 22, 37, 38, 39, 40, 52). The temporal lobes process hearing, smell, and taste information and help us understand spoken language and recognize visual objects and faces (Breedlove & Watson, 2023). Temporal lobe graphic © Kateryna Kon/Shutterstock.com.

Wernicke's area, located in the dominant hemisphere's temporoparietal cortex (BA 22), is specialized for speech perception and production. Damage can result in an inability to understand speech's meaning and construct intelligible sentences. Graphic courtesy of Blausen.com staff "Blausen gallery 2014," Wikiversity Journal of Medicine.

Major left hemisphere functions include affect, declarative memories, language comprehension, perception of movement, reading, and word recognition.

Important right hemisphere functions include face and object recognition, music, and social cues (Demos, 2019).

The parahippocampal gyri are located within the medial temporal lobe. The parahippocampal gyri form spatial and nonspatial contextual associations, which serve as building blocks for contextual processing, episodic memory, navigation, and scene processing (Aminoff, Kveraga, & Bar, 2013). They may also play a role in emotional responsiveness.

Occipital Lobes

The occipital lobes (Oz, O1, O2) are posterior to the parietal lobes. The primary visual cortex (VI) is located within the calcarine sulcus (BA 17). The occipital lobes process visual information from the eyes in collaboration with the frontal, parietal, and temporal lobes. Occipital lobe graphic © Kateryna Kon/Shutterstock.com.

Their primary functions are visual, including the analysis of orientation, color, spatial frequency, illusory contours, and complex patterns like concentric and radial stimuli (Breedlove & Watson, 2023).

Insular Cortex

The insular cortex lies deep within the lateral sulcus that divides the temporal and parietal lobes (BA 13).

The insula is involved in emotional and autonomic responses to external stimuli and is part of the salience network. The insula detects salient events via afferent pathways and switches between other large-scale networks when such events are identified, affecting attention and working memory. The anterior and poster insulae interact to regulate autonomic responses to salient stimuli. Interactive communication between the insula and anterior cingulate cortex facilitates motor control (Menon & Uddin, 2010). The right insula mediates awareness of our body, empathy, and understanding others’ points of view (Khazan, 2019).

Increased heart rate variability strengthens the connectivity between the ACC and the insula for empathy and the ability to understand others’ emotions, feel gratitude, socially connect, understand our own psychophysiological states, and restore nervous system balance. Mindfulness meditation increases insula gray matter and activation.

The insula functions as an integrative and organizational hub for the salience network. The insula integrates interoceptive awareness, emotional experience, and external perception to facilitate our global perception of the world and our relationship with it. The insula directs specific networks in processing salient stimuli and in generating appropriate behavioral responses to these stimuli (Wiebking & Northoff, 2014).

The insula is the primary taste cortex and is activated when you see something that disgusts you (a fraternity bathroom) or observe another person’s expression of disgust. Pictures of lovers also activate the anterior insula as opposed to friends. When the anterior insula was activated in neuroeconomic studies, subjects chose risk-avoidant financial strategies (choosing bonds instead of stocks). In the Prisoner’s Dilemma game, mutually cooperative decisions also resulted in the activation of this region.

Antonio Damasio has proposed that this region helps map visceral states associated with emotional experience and generate conscious feelings. This could provide the basis of somatic markers like the discomfort produced by a risky decision.

The insular cortex has been implicated in the experience of pain and basic emotions, including anger, disgust, fear, happiness, and sadness. The insular cortex receives reports of internal states, like hunger and drug craving, and motivates individuals to engage in consummatory behavior. The insular cortex plays a crucial role in craving and impulse control. Drug-related cues stimulate this region and may activate memories of pleasurable drug-related experiences. Stroke damage to the insular cortex (see red area below) can eliminate nicotine addiction. Graphic from National Institute of Drug Abuse found in Wikimedia Commons.

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Glossary

affective network: a network triggered when we make mistakes and monitors cognitive activity to predict when errors are likely, and greater executive control may be needed. The affective network includes the anterior cingulate cortex, hippocampal cortex, entorhinal cortex, superior temporal gyrus, inferior temporal gyrus, posterior parietal cortex, globus pallidus internal segment, substantia nigra, pars reticulata, and medial dorsal nucleus of the thalamus. angular gyrus: a part of the parietal lobe implicated in several processes related to language, number processing and spatial cognition, memory retrieval, attention, and theory of mind.

anterior: toward the front end or front of the body.

anterior cingulate cortex: a part of the cingulate cortex involved in decision-making, emotion, and reward anticipation.

Broca’s area is a region in the brain's frontal lobe linked to speech production.

caudal: toward the back or tail end of the body.

central sulcus: a fold in the brain's cerebral cortex, separating the frontal and parietal lobes.

cingulate cortex: a part of the brain situated in the medial aspect of the cerebral cortex, involved in processing emotions and behavior regulation.

contralateral: on the opposite side of the body.

coronal plane: an anatomical plane vertical to the ground, which divides the body into anterior (front) and posterior (back) halves.

distal: further from the center of the body or point of attachment.

dorsal: at the back or toward the back of the body.

dorsolateral prefrontal cortex (DLPFC): a part of the frontal lobes involved in working memory, cognitive flexibility, planning, inhibition, and abstract reasoning. The DLPFC is divided into left and right regions, each potentially playing different roles.

executive network: a network responsible for allocating attention, cognitive inhibition, behavioral inhibition, working memory, and cognitive flexibility. The executive network includes the dorsolateral prefrontal cortex, posterior parietal cortex, arcuate premotor area, globus pallidus internal segment, substantia nigra, pars reticulata, ventral anterior nucleus of the thalamus, and medial dorsal nucleus of the thalamus.

fissure: a deep sulcus or groove in the brain.

frontal lobes: the largest of the four main lobes of the brain, involved in various higher cognitive functions like decision-making, problem-solving, and planning.

gray matter: areas of the brain and spinal cord mainly made up of cell bodies of neurons.

horizontal (transverse) plane: an anatomical plane that is perpendicular to the coronal and sagittal planes, dividing the body into superior (upper) and inferior (lower) parts.

inferior: below or towards the feet.

insular cortex: a part of the cerebral cortex folded deep within the lateral sulcus, implicated in various functions ranging from pain perception to social emotions.

ipsilateral: on the same side of the body.

lateral: away from the midline of the body.

left dorsolateral prefrontal cortex: the left part of the dorsolateral prefrontal cortex, implicated in language, memory, and cognitive processing.

medial: toward the midline of the body.

occipital lobe: a region at the back of the brain that processes visual information.

orbitofrontal cortex (OFC): a part of the prefrontal cortex located at the front of the brain, involved in decision-making and reward anticipation.

parietal lobe: a brain region involved in sensory perception and integration, including spatial sense and navigation.

parahippocampal gyri: gray matter ridges located in the medial part of the temporal lobes, involved in memory encoding and retrieval.

posterior: toward the back end or rear of the body.

posterior cingulate cortex: a part of the cingulate cortex involved in memory and visuospatial processing.

precentral gyrus: located in the frontal lobe, it's the primary site of the motor cortex, which is responsible for voluntary movement.

prefrontal cortex: the anterior part of the frontal lobes, involved in planning complex cognitive behaviors, decision-making, and moderating social behavior.

premotor cortex: a brain region located in the frontal lobe, responsible for helping plan movements.

primary motor cortex: located in the precentral gyrus, it's the main contributor to generating neural impulses that pass down to the spinal cord and control the execution of movement.

primary somatosensory cortex (S1): located in the postcentral gyrus of the parietal lobe, it's the main sensory receptive area for the sense of touch.

Prisoner’s Dilemma: a theoretical situation in game theory where individuals, who could benefit from cooperating, often get the worse possible outcome due to mistrust and an unwillingness to cooperate.

proximal: closer to the center of the body or point of attachment.

right dorsolateral prefrontal cortex: the right part of the dorsolateral prefrontal cortex, implicated in task management and memory processes.

rostral: toward the front or head of the body.

sagittal plane: an anatomical plane that divides the body into right and left parts.

salience network: structures including the insula and anterior cingulate cortex that seek to monitor our external and internal environments to determine which inputs are salient and require further processing and attention.

secondary somatosensory cortex (S2): a part of the brain that processes sensory information. It is located in the parietal lobe. somato-cognitive action network (SCAN): a network comprising M1, the SMA, Centromedian nucleus (CM) and Ventral Intermediate nucleus (VIM) of the thalamus, posterior putamen, and vermis and the flocculonodular lobe of the cerebellum, mediating posture and balance. The SCAN controls the integrated movement of multiple body parts.

Stroop Test: a psychological test of mental speed and flexibility, requiring participants to say the word's color and not the word's name.

sulcus: a groove, crevice, or furrow structure in the brain, smaller than a fissure.

superior: above or towards the head.

Sylvian fissure: also known as the lateral sulcus, it separates the frontal and parietal lobes from the temporal lobe.

temporal lobe: a part of the brain involved in processing auditory information and encoding memory.

ventral: at the front or towards the belly side of the body.

ventromedial prefrontal cortex (VMPFC): a part of the prefrontal cortex involved in processing risk and fear and regulating social and emotional behaviors. ventromedial reward network: a network of brain regions that process reward, motivation, and decision-making. This network primarily includes the ventromedial prefrontal cortex (VMPFC) and interconnected regions like the ventral striatum (including the nucleus accumbens), the amygdala, and the hypothalamus. Wernicke’s area: an area of the brain located in the temporal lobe, crucial for language comprehension.

white matter: a part of the brain and spinal cord consisting mainly of myelinated nerve fibers. It plays a crucial role in learning and brain functions.

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