Telencephalon
Encyclopedia
The cerebrum or telencephalon, together with the diencephalon
Diencephalon
The diencephalon is the region of the vertebrate neural tube which gives rise to posterior forebrain structures. In development, the forebrain develops from the prosencephalon, the most anterior vesicle of the neural tube which later forms both the diencephalon and the...

, constitutes the forebrain. The cerebrum is the most anterior (or, in humans, most superior) region of the vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

 central nervous system
Central nervous system
The central nervous system is the part of the nervous system that integrates the information that it receives from, and coordinates the activity of, all parts of the bodies of bilaterian animals—that is, all multicellular animals except sponges and radially symmetric animals such as jellyfish...

. Telencephalon refers to the embryonic structure, from which the mature cerebrum develops. In mammals, the dorsal
Dorsum (biology)
In anatomy, the dorsum is the upper side of animals that typically run, fly, or swim in a horizontal position, and the back side of animals that walk upright. In vertebrates the dorsum contains the backbone. The term dorsal refers to anatomical structures that are either situated toward or grow...

 telencephalon, or pallium
Pallium (neuroanatomy)
In a neuroanatomy context, the word pallium refers to the layers of gray and white matter that cover the upper surface of the telencephalon in vertebrates. The non-pallial part of the telencephalon builds the subpallium. In basal vertebrates the pallium is a relatively simple three-layered...

, develops into the cerebral cortex
Cerebral cortex
The cerebral cortex is a sheet of neural tissue that is outermost to the cerebrum of the mammalian brain. It plays a key role in memory, attention, perceptual awareness, thought, language, and consciousness. It is constituted of up to six horizontal layers, each of which has a different...

, and the ventral telencephalon, or subpallium, becomes the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

. The cerebrum is also divided into approximately symmetric left and right cerebral hemispheres.
Lateralization of brain function
A longitudinal fissure separates the human brain into two distinct cerebral hemispheres, connected by the corpus callosum. The sides resemble each other and each hemisphere's structure is generally mirrored by the other side. Yet despite the strong anatomical similarities, the functions of each...



With the assistance of the cerebellum
Cerebellum
The cerebellum is a region of the brain that plays an important role in motor control. It may also be involved in some cognitive functions such as attention and language, and in regulating fear and pleasure responses, but its movement-related functions are the most solidly established...

, the cerebrum controls all voluntary actions in the body.

Development

During vertebrate embryonic development, the prosencephalon
Prosencephalon
In the anatomy of the brain of vertebrates, the prosencephalon is the rostral-most portion of the brain. The prosencephalon, the mesencephalon , and rhombencephalon are the three primary portions of the brain during early development of the central nervous system...

, the most anterior of three vesicle
Vesicle (biology)
A vesicle is a bubble of liquid within another liquid, a supramolecular assembly made up of many different molecules. More technically, a vesicle is a small membrane-enclosed sack that can store or transport substances. Vesicles can form naturally because of the properties of lipid membranes , or...

s that form from the embryo
Embryo
An embryo is a multicellular diploid eukaryote in its earliest stage of development, from the time of first cell division until birth, hatching, or germination...

nic neural tube
Neural tube
In the developing vertebrate, the neural tube is the embryo's precursor to the central nervous system, which comprises the brain and spinal cord...

, is further subdivided into the telencephalon and diencephalon
Diencephalon
The diencephalon is the region of the vertebrate neural tube which gives rise to posterior forebrain structures. In development, the forebrain develops from the prosencephalon, the most anterior vesicle of the neural tube which later forms both the diencephalon and the...

. The telencephalon then forms two lateral telencephalic vesicles which develop into the left and right cerebral hemispheres.

Structure

The cerebrum is composed of the following sub-regions:
  • Cerebral cortex
    Cerebral cortex
    The cerebral cortex is a sheet of neural tissue that is outermost to the cerebrum of the mammalian brain. It plays a key role in memory, attention, perceptual awareness, thought, language, and consciousness. It is constituted of up to six horizontal layers, each of which has a different...

    , or cortices of the cerebral hemispheres
  • Basal ganglia
    Basal ganglia
    The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

    , or basal nuclei
  • Limbic System
    Limbic system
    The limbic system is a set of brain structures including the hippocampus, amygdala, anterior thalamic nuclei, septum, limbic cortex and fornix, which seemingly support a variety of functions including emotion, behavior, long term memory, and olfaction. The term "limbic" comes from the Latin...


Composition

The cerebrum comprises what most people think of as the "brain
Brain
The brain is the center of the nervous system in all vertebrate and most invertebrate animals—only a few primitive invertebrates such as sponges, jellyfish, sea squirts and starfishes do not have one. It is located in the head, usually close to primary sensory apparatus such as vision, hearing,...

." It lies in front or on top of the brainstem and in humans is the largest and most well-developed of the five major divisions of the brain. The cerebrum is the newest structure in the phylogenetic sense, with mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s having the largest and most well-developed among all species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

. In larger mammals, the cerebral cortex is folded into many gyri and sulci, which has allowed the cortex to expand in surface area without taking up much greater volume.

In human
Human
Humans are the only living species in the Homo genus...

s, the cerebrum surrounds older parts of the brain. Limbic, olfactory, and motor systems project fibers from the cerebrum to the brainstem and spinal cord
Spinal cord
The spinal cord is a long, thin, tubular bundle of nervous tissue and support cells that extends from the brain . The brain and spinal cord together make up the central nervous system...

. Cognitive
Cognition
In science, cognition refers to mental processes. These processes include attention, remembering, producing and understanding language, solving problems, and making decisions. Cognition is studied in various disciplines such as psychology, philosophy, linguistics, and computer science...

 and volitive
Volition (psychology)
Volition or will is the cognitive process by which an individual decides on and commits to a particular course of action. It is defined as purposive striving, and is one of the primary human psychological functions...

 systems project fibers from the cerebrum to the thalamus
Thalamus
The thalamus is a midline paired symmetrical structure within the brains of vertebrates, including humans. It is situated between the cerebral cortex and midbrain, both in terms of location and neurological connections...

 and to specific regions of the midbrain. The neural networks of the cerebrum facilitate complex behaviors such as social interactions, thought, judgement, learning, working memory
Working memory
Working memory has been defined as the system which actively holds information in the mind to do verbal and nonverbal tasks such as reasoning and comprehension, and to make it available for further information processing...

, and in humans, speech and language
Language
Language may refer either to the specifically human capacity for acquiring and using complex systems of communication, or to a specific instance of such a system of complex communication...

.

Functions

Note: As the cerebrum is a gross division with many subdivisions and sub-regions, it is important to state that this section lists the functions that the cerebrum as a whole serves. See main articles on cerebral cortex
Cerebral cortex
The cerebral cortex is a sheet of neural tissue that is outermost to the cerebrum of the mammalian brain. It plays a key role in memory, attention, perceptual awareness, thought, language, and consciousness. It is constituted of up to six horizontal layers, each of which has a different...

 and basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

 for more information.

Movement

The cerebrum directs the conscious or volitional motor functions of the body. These functions originate within the primary motor cortex
Primary motor cortex
The primary motor cortex is a brain region that in humans is located in the posterior portion of the frontal lobe. Itworks in association with pre-motor areas to plan and execute movements. M1 contains large neurons known as Betz cells, which send long axons down the spinal cord to synapse onto...

 and other frontal lobe motor areas where actions are planned. Upper motor neuron
Upper motor neuron
Upper motor neurons are motor neurons that originate in the motor region of the cerebral cortex or the brain stem and carry motor information down to the final common pathway, that is, any motor neurons that are not directly responsible for stimulating the target muscle...

s in the primary motor cortex send their axon
Axon
An axon is a long, slender projection of a nerve cell, or neuron, that conducts electrical impulses away from the neuron's cell body or soma....

s to the brainstem and spinal cord to synapse
Synapse
In the nervous system, a synapse is a structure that permits a neuron to pass an electrical or chemical signal to another cell...

 on the lower motor neurons, which innervate the muscles. Damage to motor areas of cortex can lead to certain types of motor neuron disease. This kind of damage results in loss of muscular power and precision rather than total paralysis
Paralysis
Paralysis is loss of muscle function for one or more muscles. Paralysis can be accompanied by a loss of feeling in the affected area if there is sensory damage as well as motor. A study conducted by the Christopher & Dana Reeve Foundation, suggests that about 1 in 50 people have been diagnosed...

.

It functions as the center of sensory perception, memory, thoughts and judgement; also functions as the center of voluntary motor activity.

Sensory processing

The primary sensory areas of the cerebral cortex
Cerebral cortex
The cerebral cortex is a sheet of neural tissue that is outermost to the cerebrum of the mammalian brain. It plays a key role in memory, attention, perceptual awareness, thought, language, and consciousness. It is constituted of up to six horizontal layers, each of which has a different...

 receive and process visual, auditory, somatosensory, gustatory, and olfactory information. Together with association cortical areas, these brain regions synthesize sensory information into our perceptions of the world around us.

Olfaction

The olfactory bulb
Olfactory bulb
The olfactory bulb is a structure of the vertebrate forebrain involved in olfaction, the perception of odors.-Anatomy:In most vertebrates, the olfactory bulb is the most rostral part of the brain. In humans, however, the olfactory bulb is on the inferior side of the brain...

 in most vertebrates is the most anterior portion of the cerebrum, and makes up a relatively large proportion of the telencephalon. However, in humans, this part of the brain is much smaller, and lies underneath the frontal lobe. The olfactory sensory system is unique in the sense that neurons in the olfactory bulb send their axons directly to the olfactory cortex
Piriform cortex
In anatomy of animals, the piriform cortex, or pyriform cortex is a region in the brain.-Anatomy and function:The piriform cortex is part of the rhinencephalon situated in the telencephalon....

, rather than to the thalamus
Thalamus
The thalamus is a midline paired symmetrical structure within the brains of vertebrates, including humans. It is situated between the cerebral cortex and midbrain, both in terms of location and neurological connections...

 first. Damage to the olfactory bulb results in a loss of the sense of smell.

Language and communication

Speech and language are mainly attributed to parts of the cerebral cortex. Motor portions of language are attributed to Broca's area
Broca's area
Broca's area is a region of the hominid brain with functions linked to speech production.The production of language has been linked to the Broca’s area since Pierre Paul Broca reported impairments in two patients. They had lost the ability to speak after injury to the posterior inferior frontal...

 within the frontal lobe. Speech comprehension is attributed to Wernicke's area
Wernicke's area
Wernicke's area is one of the two parts of the cerebral cortex linked since the late nineteenth century to speech . It is involved in the understanding of written and spoken language...

, at the temporal-parietal lobe junction. These two regions are interconnected by a large white matter
White matter
White matter is one of the two components of the central nervous system and consists mostly of myelinated axons. White matter tissue of the freshly cut brain appears pinkish white to the naked eye because myelin is composed largely of lipid tissue veined with capillaries. Its white color is due to...

 tract, the arcuate fasciculus
Arcuate fasciculus
The arcuate fasciculus is the neural pathway connecting the posterior part of the temporoparietal junction with the frontal cortex in the brain and is now considered as part of the superior longitudinal fasciculus..-Neuroanatomy:...

. Damage to the Broca's area results in expressive aphasia
Expressive aphasia
Expressive aphasia , also known as Broca's aphasia in clinical neuropsychology and agrammatic aphasia in cognitive neuropsychology, is caused by damage to or developmental issues in anterior regions of the brain, including the left posterior inferior frontal gyrus known as Broca's area...

 (non-fluent aphasia) while damage to Wernicke's area results in receptive aphasia
Receptive aphasia
Receptive aphasia, also known as Wernicke’s aphasia, fluent aphasia, or sensory aphasia, is a type of aphasia traditionally associated with neurological damage to Wernicke’s area in the brain,...

 (also called fluent aphasia).

Learning and memory

Explicit or declarative (factual) memory formation is attributed to the hippocampus
Hippocampus
The hippocampus is a major component of the brains of humans and other vertebrates. It belongs to the limbic system and plays important roles in the consolidation of information from short-term memory to long-term memory and spatial navigation. Humans and other mammals have two hippocampi, one in...

 and associated regions of the medial temporal lobe. This association was originally described after a patient known as HM
HM (patient)
Henry Gustav Molaison , famously known as HM or H.M., was an American memory disorder patient who was widely studied from late 1957 until his death...

 had both his hippocampuses (left and right) surgically removed to treat severe epilepsy. After surgery, HM had anterograde amnesia
Anterograde amnesia
Anterograde amnesia is a loss of the ability to create new memories after the event that caused the amnesia, leading to a partial or complete inability to recall the recent past, while long-term memories from before the event remain intact. This is in contrast to retrograde amnesia, where memories...

, or the inability to form new memories.

Implicit or procedural memory, such as complex motor behaviors, involves the basal ganglia.

Short-term or working memory involves association areas of the cortex, especially the dorsolateral prefrontal cortex
Dorsolateral prefrontal cortex
The dorsolateral prefrontal cortex , according to a more restricted definition, is roughly equivalent to Brodmann areas 9 and 46. According to a broader definition DL-PFC consists of the lateral portions of Brodmann areas 9 – 12, of areas 45, 46, and the superior part of area 47. These regions...

, as well as the hippocampus.

Variation among species

In the most primitive living vertebrates, the hagfish
Hagfish
Hagfish, the clade Myxini , are eel-shaped slime-producing marine animals . They are the only living animals that have a skull but not a vertebral column. Along with lampreys, hagfish are jawless and are living fossils whose next nearest relatives include all vertebrates...

es and lamprey
Lamprey
Lampreys are a family of jawless fish, whose adults are characterized by a toothed, funnel-like sucking mouth. Translated from an admixture of Latin and Greek, lamprey means stone lickers...

s, the cerebrum is a relatively simple structure receiving nerve impulses from the olfactory bulb
Olfactory bulb
The olfactory bulb is a structure of the vertebrate forebrain involved in olfaction, the perception of odors.-Anatomy:In most vertebrates, the olfactory bulb is the most rostral part of the brain. In humans, however, the olfactory bulb is on the inferior side of the brain...

. In cartilaginous and lobe-finned fishes, and also in amphibian
Amphibian
Amphibians , are a class of vertebrate animals including animals such as toads, frogs, caecilians, and salamanders. They are characterized as non-amniote ectothermic tetrapods...

s, a more complex structure is present, with the cerebrum being divided into three distinct regions. The lowermost (or ventral) region forms the basal nuclei, and contains fibres connecting the rest of the cerebrum to the thalamus
Thalamus
The thalamus is a midline paired symmetrical structure within the brains of vertebrates, including humans. It is situated between the cerebral cortex and midbrain, both in terms of location and neurological connections...

. Above this, and forming the lateral part of the cerebrum, is the paleopallium, while the uppermost (or dorsal) part is referred to as the archipallium. The cerebrum remains largely devoted to olfactory sensation in these animals, despite its much wider range of functions in amniote
Amniote
The amniotes are a group of tetrapods that have a terrestrially adapted egg. They include synapsids and sauropsids , as well as their fossil ancestors. Amniote embryos, whether laid as eggs or carried by the female, are protected and aided by several extensive membranes...

s.

In ray-finned fishes, the structure is somewhat different. The inner surfaces of the lateral and ventral regions of the cerebrum bulge up into the ventricles; these include both the basal nuclei and the various parts of the pallium, and may be complex in structure, especially in teleosts. The dorsal surface of the cerebrum is membranous, and does not contain any nervous tissue.

In the amniotes, the cerebrum becomes increasingly large and complex. In reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...

s, the paleopallium is much larger than in amphibians, and its growth has pushed the basal nuclei into the central regions of the cerebrum. As in the lower vertebrates, the grey matter is generally located beneath the white matter, but in some reptiles, it spreads out to the surface to form a primitive cortex, especially in the anterior part of the brain.

In mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s, this development proceeds further, so that the cortex covers almost the whole of the cerebral hemispheres, especially in more "advanced" species, such as primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

s. The paleopallium is pushed to the ventral surface of the brain, where it becomes the olfactory lobes, while the archipallium becomes rolled over at the medial dorsal edge to form the hippocampus
Hippocampus
The hippocampus is a major component of the brains of humans and other vertebrates. It belongs to the limbic system and plays important roles in the consolidation of information from short-term memory to long-term memory and spatial navigation. Humans and other mammals have two hippocampi, one in...

. In placental mammals, a corpus callosum
Corpus callosum
The corpus callosum , also known as the colossal commissure, is a wide, flat bundle of neural fibers beneath the cortex in the eutherian brain at the longitudinal fissure. It connects the left and right cerebral hemispheres and facilitates interhemispheric communication...

 also develops, further connecting the two hemispheres. The complex convolutions of the cerebral surface are also found only in higher mammals.

The cerebrum of bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s has evolved along different lines to that of mammals, although they are similarly enlarged, by comparison with reptiles. However, this enlargement is largely due to the basal ganglia, with the other areas remaining relatively primitive in structure. For example, there is no great expansion of the cerebral cortex, as there is in mammals. Instead, an HVC
High vocal center
The HVC is a nucleus in the brain of the songbirds necessary for both the learning and the production of bird song...

 develops just above the basal ganglia, and this appears to be the area of the bird brain most concerned with learning complex tasks.

External links

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