Cerebral cortex
The cerebral cortex, also known as the cerebral mantle, is the outer layer of neural tissue of the cerebrum of the brain in humans and other mammals. It is the largest site of neural integration in the central nervous system, and plays a key role in attention, perception, awareness, thought, memory, language, and consciousness.
The six-layered neocortex makes up approximately 90% of the cortex, with the allocortex making up the remainder. The cortex is divided into left and right parts by the longitudinal fissure, which separates the two cerebral hemispheres that are joined beneath the cortex by the corpus callosum and other commissural fibers. In most mammals, apart from small mammals that have small brains, the cerebral cortex is folded, providing a greater surface area in the confined volume of the cranium. Apart from minimising brain and cranial volume, cortical folding is crucial for the brain circuitry and its functional organisation. In mammals with small brains, there is no folding and the cortex is smooth.
A fold or ridge in the cortex is termed a gyrus and a groove is termed a sulcus. These surface convolutions appear during fetal development and continue to mature after birth through the process of gyrification. In the human brain, the majority of the cerebral cortex is not visible from the outside, but buried in the sulci. The major sulci and gyri mark the divisions of the cerebrum into the lobes of the brain. The four major lobes are the frontal, parietal, occipital and temporal lobes. Other lobes are the limbic lobe, and the insular cortex often referred to as the insular lobe.
There are between 14 and 16 billion neurons in the human cerebral cortex. These are organised into horizontal cortical layers, and radially into cortical columns and minicolumns. Cortical areas have specific functions such as movement in the motor cortex, and sight in the visual cortex. The motor cortex is primarily located in the precentral gyrus, and the visual cortex is located in the occipital lobe.
Structure
The cerebral cortex is the outer covering of the surfaces of the cerebral hemispheres and is folded into peaks called gyri, and grooves called sulci. In the human brain, it is between 2 and 3-4 mm. thick, and makes up 40% of the brain's mass. 90% of the cerebral cortex is the six-layered neocortex whilst the other 10% is made up of the three/four-layered allocortex. There are between 14 and 16 billion neurons in the cortex. These cortical neurons are organized radially in cortical columns, and minicolumns, in the horizontally organized layers of the cortex.The neocortex is separable into different regions of cortex known in the plural as cortices, and include the motor cortex and visual cortex. About two thirds of the cortical surface is buried in the sulci and the insular cortex is completely hidden. The cortex is thickest over the top of a gyrus and thinnest at the bottom of a sulcus.
Folds
The cerebral cortex is folded in a way that allows a large surface area of neural tissue to fit within the confines of the neurocranium. When unfolded in the human, each hemispheric cortex has a total surface area of about. The folding is inward away from the surface of the brain, and is also present on the medial surface of each hemisphere within the longitudinal fissure. Most mammals have a cerebral cortex that is convoluted with the peaks known as gyri and the troughs or grooves known as sulci. Some small mammals including some small rodents have smooth cerebral surfaces without gyrification.Lobes
The larger sulci and gyri mark the divisions of the cortex of the cerebrum into the lobes of the brain. There are four main lobes: the frontal lobe, parietal lobe, temporal lobe, and occipital lobe. The insular cortex is often included as the insular lobe. The limbic lobe is a rim of cortex on the medial side of each hemisphere and is also often included. There are also three lobules of the brain described: the paracentral lobule, the superior parietal lobule, and the inferior parietal lobule.Thickness
For species of mammals, larger brains tend to have thicker cortices. The smallest mammals, such as shrews, have a neocortical thickness of about 0.5 mm; the ones with the largest brains, such as humans and fin whales, have thicknesses of 2–4 mm. There is an approximately logarithmic relationship between brain weight and cortical thickness.Magnetic resonance imaging of the brain makes it possible to get a measure for the thickness of the human cerebral cortex and relate it to other measures. The thickness of different cortical areas varies but in general, sensory cortex is thinner than motor cortex. One study has found some positive association between the cortical thickness and intelligence.
Another study has found that the somatosensory cortex is thicker in migraine patients, though it is not known if this is the result of migraine attacks, the cause of them or if both are the result of a shared cause.
A later study using a larger patient population reports no change in the cortical thickness in patients with migraine.
A genetic disorder of the cerebral cortex, whereby decreased folding in certain areas results in a microgyrus, where there are four layers instead of six, is in some instances seen to be related to dyslexia.
Layers of neocortex
The neocortex is formed of six layers, numbered I to VI, from the outermost layer I – near to the pia mater, to the innermost layer VI – near to the underlying white matter. Each cortical layer has a characteristic distribution of different neurons and their connections with other cortical and subcortical regions. There are direct connections between different cortical areas and indirect connections via the thalamus.One of the clearest examples of cortical layering is the line of Gennari in the primary visual cortex. This is a band of whiter tissue that can be observed with the naked eye in the calcarine sulcus of the occipital lobe. The line of Gennari is composed of axons bringing visual information from the thalamus into layer IV of the visual cortex.
Staining cross-sections of the cortex to reveal the position of neuronal cell bodies and the intracortical axon tracts allowed neuroanatomists in the early 20th century to produce a detailed description of the laminar structure of the cortex in different species. The work of Korbinian Brodmann established that the mammalian neocortex is consistently divided into six layers.
Layer I
Layer I is the molecular layer, and contains few scattered neurons, including GABAergic rosehip neurons. Layer I consists largely of extensions of apical dendritic tufts of pyramidal neurons and horizontally oriented axons, as well as glial cells. During development, Cajal–Retzius cells and subpial granular layer cells are present in this layer. Also, some spiny stellate cells can be found here. Inputs to the apical tufts are thought to be crucial for the feedback interactions in the cerebral cortex involved in associative learning and attention.While it was once thought that the input to layer I came from the cortex itself, it is now known that layer I across the cerebral cortex receives substantial input from matrix or M-type thalamus cells, as opposed to core or C-type that go to layer IV.
It is thought that layer I serves as a central hub for collecting and processing widespread information. It integrates ascending sensory inputs with top-down expectations, regulating how sensory perceptions align with anticipated outcomes. Further, layer I sorts, directs, and combines excitatory inputs, integrating them with neuromodulatory signals. Inhibitory interneurons, both within layer I and from other cortical layers, gate these signals. Together, these interactions dynamically calibrate information flow throughout the neocortex, shaping perceptions and experiences.