Evolution of nervous systems

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The evolution of nervous systems dates back to the first development of nervous systems in animals (or metazoans). Neurons developed as specialized electrical signaling cells in multicellular animals, adapting the mechanism of action potentials present in motile single-celled and colonial eukaryotes. Primitive systems, like those found in protists, use chemical signalling for movement and sensitivity; data suggests these were precursors to modern neural cell types and their synapses. [1] When some animals started living a mobile lifestyle and eating larger food particles externally, they developed ciliated epithelia, contractile muscles and coordinating & sensitive neurons for it in their outer layer. [2] [3]

Contents

Simple nerve nets seen in acoels (basal bilaterians) and cnidarians are thought to be the ancestral condition for the Planulozoa (bilaterians plus cnidarians and, perhaps, placozoans). A more complex nerve net with simple nerve cords is present in ancient animals called ctenophores [4] but no nerves, thus no nervous systems, are present in another group of ancient animals, the sponges (Porifera). Due to the common presence and similarity of some neural genes in these ancient animals and their protist relatives, the controversy of whether ctenophores or sponges diverged earlier, and the recent discovery of "neuroid" cells specialized in coordination of digestive choanocytes in Spongilla , the origin of neurons in the phylogenetic tree of life is still disputed. [5] [6] [7] Further cephalization and nerve cord (ventral and dorsal) evolution occurred many times independently in bilaterians. [5]

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. In some colonial eukaryotes, such as Obelia, electrical signals propagate not only through neural nets, but also through epithelial cells in the shared digestive system of the colony. [8] Several non-metazoan phyla, including choanoflagellates, filasterea, and mesomycetozoea, have been found to have synaptic protein homologs, including secretory SNAREs, Shank, and Homer. In choanoflagellates and mesomycetozoea, these proteins are upregulated during colonial phases, suggesting the importance of these proto-synaptic proteins for cell to cell communication. [9] The history of ideas on how neurons and the first nervous systems emerged in evolution has been discussed in a 2015 book by Michel Antcil. [10] In 2022 two proteins SMIM20 and NUCB2, that are precursors of the neuropeptides phoenixin and nesfatin-1 respectively have been found to have deep homology across all lineages that preceded creatures with central nervous systems, bilaterians, cnidarians, ctenophores, and sponges as well as in choanoflagellates. [11] [12]

Sponges

Sponges have no cells connected to each other by synaptic junctions, that is, no neurons, and therefore no nervous system. They do, however, have homologs 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 (the signal-receiving part of a synapse). [13] 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. [14] Other ways sponge cells communicate with neighboring cells is through vesicular transport across highly dense regions of the cell membranes. These vesicles carry ions and other signaling molecules, but contain no true synaptic function. [15]

Nerve nets

Jellyfish, comb jellies, 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. [16]

The development of the nervous system in radiata is relatively unstructured. Unlike bilaterians, radiata only have two primordial cell layers, endoderm and ectoderm. Neurons are generated from a special set of ectodermal precursor cells, which also serve as precursors for every other ectodermal cell type. [17]

Nerve cords

Nervous system of a bilaterian animal, in the form of a nerve cord with a "brain" at the front Bilaterian-plan.svg
Nervous system of a bilaterian animal, in the form of a nerve cord with a "brain" at the front

The vast majority of existing animals are bilaterians, 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 Cryogenian period, 700–650 million years ago. [18] The fundamental bilaterian body form is a tube with a hollow gut cavity running from mouth to anus, and a nerve cord with an especially large ganglion at the front, called the "brain".

Area of the human body surface innervated by each spinal nerve Gray797.png
Area of the human body surface innervated by each spinal nerve

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. [19]

Bilaterians can be divided, based on events that occur very early in embryonic development, into two groups (superphyla) called protostomes and deuterostomes. [20] Deuterostomes include vertebrates as well as echinoderms and hemichordates (mainly acorn worms). Protostomes, the more diverse group, include arthropods, 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. Some 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 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. [21] But recent molecular data from different protostomes and deuterostomes reject this scenario and suggest that nerve cords independently evolved in both. [22]

Annelida

Earthworms have dual nerve cords running along the length of the body and merging at the tail and the mouth. These nerve cords are connected by transverse nerves like the rungs of a ladder. These transverse nerves help coordinate the two sides of the animal. Two ganglia at the head end function similar to a simple brain. Photoreceptors on the animal's eyespots provide sensory information on light and dark. [23]

Nematoda

The nervous system of one very small worm, the roundworm Caenorhabditis elegans , has been mapped out down to the synaptic level. Every neuron and its cellular lineage 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. [24]

Arthropods

Internal anatomy of a spider, showing the nervous system in blue Spider internal anatomy-en.svg
Internal anatomy of a spider, showing the nervous system in blue

Arthropods, such as insects and crustaceans, have a nervous system made up of a series of ganglia, connected by a ventral nerve cord made up of two parallel connectives running along the length of the belly. [25] Typically, each body segment has one ganglion 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. In the insect nervous system, the brain is anatomically divided into the protocerebrum, deutocerebrum, and tritocerebrum. Immediately behind the brain is the subesophageal ganglion, which is composed of three pairs of fused ganglia. It controls the mouthparts, the salivary glands and certain muscles. Many arthropods have well-developed sensory organs, including compound eyes for vision and antennae for olfaction and pheromone 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, in the interior. [26]

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 (see Homininae), starting from about 600 cm3 in Homo habilis up to 1736 cm3 in Homo neanderthalensis . Thus, in general there is a correlation between brain volume and intelligence. [27] However, modern Homo sapiens have a smaller brain volume (brain size 1250 cm3) than neanderthals; women have a brain volume slightly smaller than men, and the Flores hominids ( Homo floresiensis ), nicknamed "hobbits", had a cranial capacity of about 380 cm3, about a third of the Homo erectus average and considered small for a chimpanzee. 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. [28] Iain Davidson summarizes the opposite evolutionary constraints on human brain size as "As large as you need and as small as you can". [29] The human brain has evolved around the metabolic, environmental, and social needs that the species has dealt with throughout its existence. As hominid species evolved with increased brain size and processing power, the overall metabolic need increased. Compared to chimpanzees, humans consume more calories from animals than from plants. While not certain, studies have shown that this shift in diet is due to the increased need for the fatty acids more readily found in animal products[ citation needed ]. These fatty acids are essential for brain maintenance and development. Other factors to consider are the need for social interaction and how hominids have interacted with their environments over time. [30]

Brain evolution can be studied using endocasts, a branch of neurology and paleontology called paleoneurology.

See also

Related Research Articles

<span class="mw-page-title-main">Axon</span> Long projection on a neuron that conducts signals to other neurons

An axon or nerve fiber is a long, slender projection of a nerve cell, or neuron, in vertebrates, that typically conducts electrical impulses known as action potentials away from the nerve cell body. The function of the axon is to transmit information to different neurons, muscles, and glands. In certain sensory neurons, such as those for touch and warmth, the axons are called afferent nerve fibers and the electrical impulse travels along these from the periphery to the cell body and from the cell body to the spinal cord along another branch of the same axon. Axon dysfunction can be the cause of many inherited and acquired neurological disorders that affect both the peripheral and central neurons. Nerve fibers are classed into three types – group A nerve fibers, group B nerve fibers, and group C nerve fibers. Groups A and B are myelinated, and group C are unmyelinated. These groups include both sensory fibers and motor fibers. Another classification groups only the sensory fibers as Type I, Type II, Type III, and Type IV.

<span class="mw-page-title-main">Brain</span> Organ that controls the nervous system in vertebrates and most invertebrates

The brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. In vertebrates, a small part of the brain called the hypothalamus is the neural control center for all endocrine systems. The brain is the largest cluster of neurons in the body and is typically located in the head, usually near organs for special senses such as vision, hearing and olfaction. It is the most energy-consuming organ of the body, and the most specialized, responsible for endocrine regulation, sensory perception, motor control, and the development of intelligence.

<span class="mw-page-title-main">Central nervous system</span> Brain and spinal cord

The central nervous system (CNS) is the part of the nervous system consisting of the brain and spinal cord, the retina and optic nerve, and the olfactory nerve and epithelia. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all parts of the bodies of bilaterally symmetric and triploblastic animals—that is, all multicellular animals except sponges and diploblasts. It is a structure composed of nervous tissue positioned along the rostral to caudal axis of the body and may have an enlarged section at the rostral end which is a brain. Only arthropods, cephalopods and vertebrates have a true brain, though precursor structures exist in onychophorans, gastropods and lancelets.

<span class="mw-page-title-main">Neuron</span> Electrically excitable cell found in the nervous system of animals

Within a nervous system, a neuron, neurone, or nerve cell is an electrically excitable cell that fires electric signals called action potentials across a neural network. Neurons communicate with other cells via synapses, which are specialized connections that commonly use minute amounts of chemical neurotransmitters to pass the electric signal from the presynaptic neuron to the target cell through the synaptic gap.

<span class="mw-page-title-main">Nervous system</span> Part of an animal that coordinates actions and senses

In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes that impact the body, then works in tandem with the endocrine system to respond to such events. Nervous tissue first arose in wormlike organisms about 550 to 600 million years ago. In vertebrates, it consists of two main parts, the central nervous system (CNS) and the peripheral nervous system (PNS). The CNS consists of the brain and spinal cord. The PNS consists mainly of nerves, which are enclosed bundles of the long fibers, or axons, that connect the CNS to every other part of the body. Nerves that transmit signals from the brain are called motor nerves or efferent nerves, while those nerves that transmit information from the body to the CNS are called sensory nerves or afferent. Spinal nerves are mixed nerves that serve both functions. The PNS is divided into three separate subsystems, the somatic, autonomic, and enteric nervous systems. Somatic nerves mediate voluntary movement. The autonomic nervous system is further subdivided into the sympathetic and the parasympathetic nervous systems. The sympathetic nervous system is activated in cases of emergencies to mobilize energy, while the parasympathetic nervous system is activated when organisms are in a relaxed state. The enteric nervous system functions to control the gastrointestinal system. Both autonomic and enteric nervous systems function involuntarily. Nerves that exit from the cranium are called cranial nerves while those exiting from the spinal cord are called spinal nerves.

<span class="mw-page-title-main">Bilateria</span> Animals with embryonic bilateral symmetry

Bilateria is a large clade or infrakingdom of animals called bilaterians, characterized by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front and a rear end, as well as a left–right–symmetrical belly (ventral) and back (dorsal) surface. Nearly all bilaterians maintain a bilaterally symmetrical body as adults; the most notable exception is the echinoderms, which achieve secondary pentaradial symmetry as adults, but are bilaterally symmetrical as an embryo. Cephalization is also a characteristic feature among most bilaterians, where the special sense organs and central nerve ganglia become concentrated at the front/rostral end.

<span class="mw-page-title-main">Ctenophora</span> Phylum of gelatinous marine animals

Ctenophora comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming, and they are the largest animals to swim with the help of cilia.

<span class="mw-page-title-main">Nervous tissue</span> Main component of the nervous system

Nervous tissue, also called neural tissue, is the main tissue component of the nervous system. The nervous system regulates and controls body functions and activity. It consists of two parts: the central nervous system (CNS) comprising the brain and spinal cord, and the peripheral nervous system (PNS) comprising the branching peripheral nerves. It is composed of neurons, also known as nerve cells, which receive and transmit impulses, and neuroglia, also known as glial cells or glia, which assist the propagation of the nerve impulse as well as provide nutrients to the neurons.

<span class="mw-page-title-main">Nerve net</span> Nervous systems lacking a brain

A nerve net consists of interconnected neurons lacking a brain or any form of cephalization. While organisms with bilateral body symmetry are normally associated with a condensation of neurons or, in more advanced forms, a central nervous system, organisms with radial symmetry are associated with nerve nets, and are found in members of the Ctenophora, Cnidaria, and Echinodermata phyla, all of which are found in marine environments. In the Xenacoelomorpha, a phylum of bilaterally symmetrical animals, members of the subphylum Xenoturbellida also possess a nerve net. Nerve nets can provide animals with the ability to sense objects through the use of the sensory neurons within the nerve net.

<span class="mw-page-title-main">Neuropeptide</span> Peptides released by neurons as intercellular messengers

Neuropeptides are chemical messengers made up of small chains of amino acids that are synthesized and released by neurons. Neuropeptides typically bind to G protein-coupled receptors (GPCRs) to modulate neural activity and other tissues like the gut, muscles, and heart.

<span class="mw-page-title-main">Cephalization</span> Evolutionary trend of a head region developing

Cephalization is an evolutionary trend in animals that, over many generations, the special sense organs and nerve ganglia become concentrated towards the rostral end of the body where the mouth is located, often producing an enlarged head. This is associated with the animal's movement direction and bilateral symmetry, and cephalization of the nervous system led to the formation of a functional centralized brain in three groups of bilaterian animals, namely the arthropods, cephalopod molluscs, and vertebrates (craniates).

<span class="mw-page-title-main">Dorsal root ganglion</span> Cluster of neurons in a dorsal root of a spinal nerve

A dorsal root ganglion is a cluster of neurons in a dorsal root of a spinal nerve. The cell bodies of sensory neurons known as first-order neurons are located in the dorsal root ganglia.

<span class="mw-page-title-main">Ventral nerve cord</span>

The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice versa, integrating sensory input and locomotor output. Because arthropods have an open circulatory system, decapitated insects can still walk, groom, and mate — illustrating that the circuitry of the ventral nerve cord is sufficient to perform complex motor programs without brain input.

<span class="mw-page-title-main">Evolution of the brain</span> Overview of the evolution of the brain

There is much to be discovered about the evolution of the brain and the principles that govern it. While much has been discovered, not everything currently known is well understood. The evolution of the brain has appeared to exhibit diverging adaptations within taxonomic classes such as Mammalia and more vastly diverse adaptations across other taxonomic classes. Brain to body size scales allometrically. This means as body size changes, so do other physiological, anatomical, and biochemical constructs connecting the brain to the body. Small bodied mammals have relatively large brains compared to their bodies whereas large mammals have a smaller brain to body ratios. If brain weight is plotted against body weight for primates, the regression line of the sample points can indicate the brain power of a primate species. Lemurs for example fall below this line which means that for a primate of equivalent size, we would expect a larger brain size. Humans lie well above the line indicating that humans are more encephalized than lemurs. In fact, humans are more encephalized compared to all other primates. This means that human brains have exhibited a larger evolutionary increase in its complexity relative to its size. Some of these evolutionary changes have been found to be linked to multiple genetic factors, such as proteins and other organelles.

The urbilaterian is the hypothetical last common ancestor of the bilaterian clade, i.e., all animals having a bilateral symmetry.

<span class="mw-page-title-main">Nervous system of gastropods</span>

The nervous system of gastropods consists of a series of paired ganglia connected by major nerve cords, and a number of smaller branching nerves. It is sometimes called ganglionic.

In evolutionary developmental biology, inversion refers to the hypothesis that during the course of animal evolution, the structures along the dorsoventral (DV) axis have taken on an orientation opposite that of the ancestral form.

<span class="mw-page-title-main">Xenacoelomorpha</span> A deep-branching bilaterian clade of animals with a simple body plan

Xenacoelomorpha is a small phylum of bilaterian invertebrate animals, consisting of two sister groups: xenoturbellids and acoelomorphs. This new phylum was named in February 2011 and suggested based on morphological synapomorphies, which was then confirmed by phylogenomic analyses of molecular data.

<span class="mw-page-title-main">Outline of the human nervous system</span> Overview of and topical guide to the human nervous system

The following diagram is provided as an overview of and topical guide to the human nervous system:

Segmentation is the physical characteristic by which the human body is divided into repeating subunits called segments arranged along a longitudinal axis. In humans, the segmentation characteristic observed in the nervous system is of biological and evolutionary significance. Segmentation is a crucial developmental process involved in the patterning and segregation of groups of cells with different features, generating regional properties for such cell groups and organizing them both within the tissues as well as along the embryonic axis.

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