Evolution of nervous systems
Encyclopedia
The evolution of nervous systems dates back to the first development of nervous systems in animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...

s (or metazoans). Neurons developed as specialized electrical signaling cells in multicellular animals, adapting the mechanism of action potential
Action potential
In physiology, an action potential is a short-lasting event in which the electrical membrane potential of a cell rapidly rises and falls, following a consistent trajectory. Action potentials occur in several types of animal cells, called excitable cells, which include neurons, muscle cells, and...

s present in motile single-celled and colonial eukaryote
Eukaryote
A eukaryote is an organism whose cells contain complex structures enclosed within membranes. Eukaryotes may more formally be referred to as the taxon Eukarya or Eukaryota. The defining membrane-bound structure that sets eukaryotic cells apart from prokaryotic cells is the nucleus, or nuclear...

s. Simple nerve net
Nerve net
A nerve net is a type of simple nervous system that is found in members of the cnidaria, ctenophora, and echinodermata phyla. Nerve nets consist of interconnected neurons lacking a brain or any form of cephalization. This nervous system allows cnidarians to respond to physical contact. They may...

s seen in animals like cnidaria
Cnidaria
Cnidaria is a phylum containing over 9,000 species of animals found exclusively in aquatic and mostly marine environments. Their distinguishing feature is cnidocytes, specialized cells that they use mainly for capturing prey. Their bodies consist of mesoglea, a non-living jelly-like substance,...

 evolved first, followed by nerve cords in bilateral animals - dorsal nerve cord
Dorsal nerve cord
The dorsal nerve cord is one of the embryonic features unique to chordates, along with a notochord, a post-anal tail and pharyngeal slits. The dorsal nerve cord is a hollow cord dorsal to the notochord.It is formed from a part of the ectoderm that rolls, forming the hollow tube, compared to other...

s in invertebrates and ventral nerve cord
Ventral nerve cord
The ventral nerve cord makes up the nervous system of some phyla of the invertebrates, particularly within the nematodes, annelids and the arthropods. It usually consists of cerebral ganglia anteriorly with the nerve cords running down the ventral plane of the organism...

s surrounded by a notochord
Notochord
The notochord is a flexible, rod-shaped body found in embryos of all chordates. It is composed of cells derived from the mesoderm and defines the primitive axis of the embryo. In some chordates, it persists throughout life as the main axial support of the body, while in most vertebrates it becomes...

 in chordates. Bilateralization led to the evolution of brains, a process called cephalization
Cephalization
Cephalization is considered an evolutionary trend, whereby nervous tissue, over many generations, becomes concentrated toward one end of an organism. This process eventually produces a head region with sensory organs....

.

Neural precursors

Action potentials, which are necessary for neural activity, evolved in single-celled eukaryotes. These use calcium rather than sodium action potentials, but the mechanism was probably adapted into neural electrical signaling in multicellular animals. Colonial eukaryotes such as Obelia
Obelia
Obelia is a genus in the class Hydrozoa, which consists of mainly marine and some freshwater animal species and have both the polyp and medusa stages in their life cycle...

 lack a nervous system, but electrical signals do propagate through epithelial cells in the shared digestive system of the colony.

Sponges

Sponges have no cells connected to each other by synaptic junctions
Synapse
In the nervous system, a synapse is a structure that permits a neuron to pass an electrical or chemical signal to another cell...

, that is, no neurons, and therefore no nervous system. They do, however, have homologs
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...

 of many genes that play key roles in synaptic function. Recent studies have shown that sponge cells express a group of proteins that cluster together to form a structure resembling a postsynaptic density
Postsynaptic density
The postsynaptic density is a protein dense specialization attached to the postsynaptic membrane. PSDs were originally identified by electron microscopy as an electron-dense region at the membrane of a postsynaptic neuron...

 (the signal-receiving part of a synapse). However, the function of this structure is currently unclear. Although sponge cells do not show synaptic transmission, they do communicate with each other via calcium waves and other impulses, which mediate some simple actions such as whole-body contraction.

Nerve nets

Jellyfish
Cnidaria
Cnidaria is a phylum containing over 9,000 species of animals found exclusively in aquatic and mostly marine environments. Their distinguishing feature is cnidocytes, specialized cells that they use mainly for capturing prey. Their bodies consist of mesoglea, a non-living jelly-like substance,...

, comb jellies
Ctenophore
The Ctenophora are a phylum of animals that live in marine waters worldwide. Their most distinctive feature is the "combs", groups of cilia that they use for swimming, and they are the largest animals that swim by means of cilia – adults of various species range from a few millimeters to in size...

, and related animals have diffuse nerve nets rather than a central nervous system. In most jellyfish the nerve net is spread more or less evenly across the body; in comb jellies it is concentrated near the mouth. The nerve nets consist of sensory neurons that pick up chemical, tactile, and visual signals, motor neurons that can activate contractions of the body wall, and intermediate neurons that detect patterns of activity in the sensory neurons and send signals to groups of motor neurons as a result. In some cases groups of intermediate neurons are clustered into discrete ganglia
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....

.

The development of the nervous system in radiata
Radiata
The Radiata are the radially symmetric animals of the Eumetazoa subkingdom. The term Radiata has had various meanings in the history of classification...

 is relatively unstructured. Unlike bilaterians, radiata only have two primordial cell layers, endoderm
Endoderm
Endoderm is one of the three primary germ cell layers in the very early embryo. The other two layers are the ectoderm and mesoderm , with the endoderm as the intermost layer...

 and ectoderm
Ectoderm
The "ectoderm" is one of the three primary germ cell layers in the very early embryo. The other two layers are the mesoderm and endoderm , with the ectoderm as the most exterior layer...

. Neurons are generated from a special set of ectodermal precursor cells, which also serve as precursors for every other ectodermal cell type.

Nerve cords

The vast majority of existing animals are bilateria
Bilateria
The bilateria are all animals having a bilateral symmetry, i.e. they have a front and a back end, as well as an upside and downside. Radially symmetrical animals like jellyfish have a topside and downside, but no front and back...

ns, meaning animals with left and right sides that are approximate mirror images of each other. All bilateria are thought to have descended from a common wormlike ancestor that appeared in the Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...

 period, 550–600 million years ago. The fundamental bilaterian body form is a tube with a hollow gut cavity running from mouth to anus, and a nerve cord with an enlargement (a "ganglion") for each body segment, with an especially large ganglion at the front, called the "brain".

Even mammals, including humans, show the segmented bilaterian body plan at the level of the nervous system. The spinal cord contains a series of segmental ganglia, each giving rise to motor and sensory nerves that innervate a portion of the body surface and underlying musculature. On the limbs, the layout of the innervation pattern is complex, but on the trunk it gives rise to a series of narrow bands. The top three segments belong to the brain, giving rise to the forebrain, midbrain, and hindbrain.

Bilaterians can be divided, based on events that occur very early in embryonic development, into two groups (superphyla) called protostomes and deuterostome
Deuterostome
Deuterostomes are a superphylum of animals. They are a subtaxon of the Bilateria branch of the subregnum Eumetazoa, and are opposed to the protostomes...

s. Deuterostomes include vertebrates as well as echinoderm
Echinoderm
Echinoderms are a phylum of marine animals. Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone....

s, hemichordates
Hemichordata
Hemichordata is a phylum of marine deuterostome animals, generally considered the sister group of the echinoderms. They date back to the Lower or Middle Cambrian and include two main classes: Enteropneusta , and Pterobranchia. A third class, Planctosphaeroidea, is known only from the larva of a...

 (mainly acorn worms), and Xenoturbellidans. Protostomes, the more diverse group, include arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

s, molluscs, and numerous types of worms. There is a basic difference between the two groups in the placement of the nervous system within the body: protostomes possess a nerve cord on the ventral (usually bottom) side of the body, whereas in deuterostomes the nerve cord is on the dorsal (usually top) side. In fact, numerous aspects of the body are inverted between the two groups, including the expression patterns of several genes that show dorsal-to-ventral gradients. Most anatomists now consider that the bodies of protostomes and deuterostomes are "flipped over" with respect to each other, a hypothesis that was first proposed by Geoffroy Saint-Hilaire
Étienne Geoffroy Saint-Hilaire
Étienne Geoffroy Saint-Hilaire was a French naturalist who established the principle of "unity of composition". He was a colleague of Jean-Baptiste Lamarck and expanded and defended Lamarck's evolutionary theories...

 for insects in comparison to vertebrates. Thus insects, for example, have nerve cords that run along the ventral midline of the body, while all vertebrates have spinal cords that run along the dorsal midline.

Worms

Worm
Worm
The term worm refers to an obsolete taxon used by Carolus Linnaeus and Jean-Baptiste Lamarck for all non-arthropod invertebrate animals, and stems from the Old English word wyrm. Currently it is used to describe many different distantly-related animals that typically have a long cylindrical...

s are the simplest bilaterian animals, and reveal the basic structure of the bilaterian nervous system in the most straightforward way. As an example, earthworm
Earthworm
Earthworm is the common name for the largest members of Oligochaeta in the phylum Annelida. In classical systems they were placed in the order Opisthopora, on the basis of the male pores opening posterior to the female pores, even though the internal male segments are anterior to the female...

s have dual nerve cord
Ventral nerve cord
The ventral nerve cord makes up the nervous system of some phyla of the invertebrates, particularly within the nematodes, annelids and the arthropods. It usually consists of cerebral ganglia anteriorly with the nerve cords running down the ventral plane of the organism...

s running along the length of the body and merging at the tail and the mouth. These nerve cords are connected by transverse
Transverse plane
The transverse plane is an imaginary plane that divides the body into superior and inferior parts. It is perpendicular to the coronal and sagittal planes....

 nerves like the rungs of a ladder. These transverse nerves help coordinate the two sides of the animal. Two ganglia
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....

 at the head end function similar to a simple brain. Photoreceptors on the animal's eyespots provide sensory information on light and dark.

The nervous system of one very small worm, the roundworm Caenorhabditis elegans
Caenorhabditis elegans
Caenorhabditis elegans is a free-living, transparent nematode , about 1 mm in length, which lives in temperate soil environments. Research into the molecular and developmental biology of C. elegans was begun in 1974 by Sydney Brenner and it has since been used extensively as a model...

, has been mapped out down to the synaptic level. Every neuron and its cellular lineage
Fate mapping
Introduction=Fate mapping is a method of understanding the embryonic origin of various tissues in the adult organism by establishing the correspondence between individual cells or groups of cells at one stage of development and their progeny at later stages of development...

 has been recorded and most, if not all, of the neural connections are known. In this species, the nervous system is sexually dimorphic; the nervous systems of the two sexes, males and hermaphrodites, have different numbers of neurons and groups of neurons that perform sex-specific functions. In C. elegans, males have exactly 383 neurons, while hermaphrodites have exactly 302 neurons.

Arthropods

Arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

s, such as insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s and crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...

s, have a nervous system made up of a series of ganglia
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....

, connected by a ventral nerve cord
Ventral nerve cord
The ventral nerve cord makes up the nervous system of some phyla of the invertebrates, particularly within the nematodes, annelids and the arthropods. It usually consists of cerebral ganglia anteriorly with the nerve cords running down the ventral plane of the organism...

 made up of two parallel connectives running along the length of the belly
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...

. Typically, each body segment has one ganglion
Ganglion
In anatomy, a ganglion is a biological tissue mass, most commonly a mass of nerve cell bodies. Cells found in a ganglion are called ganglion cells, though this term is also sometimes used to refer specifically to retinal ganglion cells....

 on each side, though some ganglia are fused to form the brain and other large ganglia. The head segment contains the brain, also known as the supraesophageal ganglion
Supraesophageal ganglion
The supraesophageal ganglion is the first part of the insect and fish central nervous system. It receives and processes information from the first, second, and third metameres. The supraesophageal ganglion lies dorsal to the esophagus and consists of three parts: the protocerebrum , the...

. In the insect nervous system, the brain is anatomically divided into the protocerebrum, deutocerebrum, and tritocerebrum. Immediately behind the brain is the subesophageal ganglion
Subesophageal ganglion
The subesophageal ganglion of insects is composed of three pairs of fused ganglia. It controls the mouthparts, the salivary glands and certain muscles....

, which is composed of three pairs of fused ganglia. It controls the mouthparts, the salivary glands and certain muscle
Muscle
Muscle is a contractile tissue of animals and is derived from the mesodermal layer of embryonic germ cells. Muscle cells contain contractile filaments that move past each other and change the size of the cell. They are classified as skeletal, cardiac, or smooth muscles. Their function is to...

s. Many arthropods have well-developed sensory
Sense
Senses are physiological capacities of organisms that provide inputs for perception. The senses and their operation, classification, and theory are overlapping topics studied by a variety of fields, most notably neuroscience, cognitive psychology , and philosophy of perception...

 organs, including compound eyes for vision and antennae
Antenna (biology)
Antennae in biology have historically been paired appendages used for sensing in arthropods. More recently, the term has also been applied to cilium structures present in most cell types of eukaryotes....

 for olfaction
Olfaction
Olfaction is the sense of smell. This sense is mediated by specialized sensory cells of the nasal cavity of vertebrates, and, by analogy, sensory cells of the antennae of invertebrates...

 and pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...

 sensation. The sensory information from these organs is processed by the brain.

In insects, many neurons have cell bodies that are positioned at the edge of the brain and are electrically passive—the cell bodies serve only to provide metabolic support and do not participate in signalling. A protoplasmic fiber runs from the cell body and branches profusely, with some parts transmitting signals and other parts receiving signals. Thus, most parts of the insect brain have passive cell bodies arranged around the periphery, while the neural signal processing takes place in a tangle of protoplasmic fibers called neuropil
Neuropil
In neuroanatomy, a neuropil, which is sometimes referred to as a neuropile, is a region between neuronal cell bodies in the gray matter of the brain and blood-brain barrier . It consists of a dense tangle of axon terminals, dendrites and glial cell processes...

, in the interior.

Evolution of innate behaviors

Behaviors such as the "tail-flip" escape reaction
Caridoid escape reaction
The Caridoid Escape Reaction, also known as lobstering or tail-flipping, refers to an innate escape mechanism in marine and freshwater crustaceans such as lobsters, krill, shrimp and crayfish....

 in crustacea such as crayfish and lobsters are fixed action pattern
Fixed action pattern
In ethology, a fixed action pattern , or modal action pattern, is an instinctive behavioral sequence that is indivisible and runs to completion...

s that may have evolved from earlier ancestral patterns.

Evolution of central nervous systems

Evolution of the human brain

There has been a gradual increase in brain volume as the ancestors of modern humans progressed along the human timeline of evolution
Timeline of human evolution
The timeline of human evolution outlines the major events in the development of human species, and the evolution of humans' ancestors. It includes a brief explanation of some animals, species or genera, which are possible ancestors of Homo sapiens...

 (see Homininae
Homininae
Homininae is a subfamily of Hominidae, which includes humans, gorillas and chimpanzees, and some extinct relatives; it comprises all those hominids, such as Australopithecus, that arose after the split from orangutans . Our family tree, which has 3 main branches leading to chimpanzees, humans and...

), starting from about 600 cm3 in Homo habilis
Homo habilis
Homo habilis is a species of the genus Homo, which lived from approximately at the beginning of the Pleistocene period. The discovery and description of this species is credited to both Mary and Louis Leakey, who found fossils in Tanzania, East Africa, between 1962 and 1964. Homo habilis Homo...

up to 1500 cm3 in Homo sapiens neanderthalensis
Neanderthal
The Neanderthal is an extinct member of the Homo genus known from Pleistocene specimens found in Europe and parts of western and central Asia...

. Thus, in general there is a correlation between brain volume and intelligence. However, modern Homo sapiens have a brain volume slightly smaller (brain size
Brain size
Brain size is one aspect of animal anatomy and evolution. Both overall brain size and the size of substructures have been analysed, and the question of links between size and functioning - particularly intelligence - has often proved controversial...

 1250 cm3) than neanderthals, women have a brain volume slightly smaller than men and the Flores hominids (Homo floresiensis
Homo floresiensis
Homo floresiensis is a possible species, now extinct, in the genus Homo. The remains were discovered in 2003 on the island of Flores in Indonesia. Partial skeletons of nine individuals have been recovered, including one complete cranium...

), nicknamed "hobbits", had a cranial capacity of about 380 cm3, about a third of the Homo erectus
Homo erectus
Homo erectus is an extinct species of hominid that lived from the end of the Pliocene epoch to the later Pleistocene, about . The species originated in Africa and spread as far as India, China and Java. There is still disagreement on the subject of the classification, ancestry, and progeny of H...

average and considered small for a chimpanzee
Chimpanzee
Chimpanzee, sometimes colloquially chimp, is the common name for the two extant species of ape in the genus Pan. The Congo River forms the boundary between the native habitat of the two species:...

. It is proposed that they evolved from H. erectus as a case of insular dwarfism. In spite of their threefold smaller brain there is evidence that H. floresiensis used fire and made stone tools as sophisticated as those of their proposed ancestor, H. erectus. "As large as you need and as small as you can" as Iain Davidson summarizes the opposite evolutionary constraints on human brain size.

Brain evolution is can be studied using endocast
Endocast
An endocast is the internal cast of a hollow object, often specifically used for an endocasts of the cranial vault. Endocasts can be man-made for examining the properties of a hollow, inaccessible space, or occur naturally through fossilisation....

s, a branch of neurology and paleontology called paleoneurology
Paleoneurology
Paleoneurology is the study of brain evolution by analysis of brain endocasts to determine endocranial traits and volumes. The cranium is unique in that it grows in response to the growth of brain tissue rather than genetic guidance, as is the case with bones that support movement...

.
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