Cephalization

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A lobster is heavily cephalized, with eyes, antennae, multiple mouthparts, and the brain (inside the armoured exoskeleton), all concentrated at the animal's head end. Lobster, National Lobster Hatchery 2.jpg
A lobster is heavily cephalized, with eyes, antennae, multiple mouthparts, and the brain (inside the armoured exoskeleton), all concentrated at the animal's head end.

Cephalization is an evolutionary trend in animals that, over many generations, the special sense organs and nerve ganglia become concentrated towards the front 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. Cephalization of the nervous system has led to the formation of a brain with varying degrees of functional centralization in three phyla of bilaterian animals, namely the arthropods, cephalopod molluscs, and vertebrates.

Contents

Animals without bilateral symmetry

Cnidaria, such as the radially symmetrical Hydrozoa, show some degree of cephalization. The Anthomedusae have a head end with their mouth, photoreceptive cells, and a concentration of neural cells. [1]

Bilateria

Idealised bilaterian body plan. With a cylindrical body (in the main clade, the nephrozoa) and a direction of travel, the animal has head and tail ends, favouring cephalization by natural selection. Sense organs, brain, and mouth form the basis of the head. Bilaterian body plan.svg
Idealised bilaterian body plan. With a cylindrical body (in the main clade, the nephrozoa) and a direction of travel, the animal has head and tail ends, favouring cephalization by natural selection. Sense organs, brain, and mouth form the basis of the head.

Cephalization is a characteristic feature of the bilaterians, a large group containing the majority of animal phyla. [3] These have the ability to move, using muscles, and a body plan with a front end that encounters stimuli first as the animal moves forwards, and accordingly has evolved to contain many of the body's sense organs, able to detect light, chemicals, and gravity. There is often also a collection of nerve cells able to process the information from these sense organs, forming a brain in several phyla and one or more ganglia in others. [2]

Acoela

The Acoela are basal bilaterians, part of the Xenacoelomorpha. They are small and simple animals with flat bodies. They have slightly more nerve cells at the head end than elsewhere, not forming a distinct and compact brain. This represents an early stage in cephalization. [4]

Flatworms

The yellow papillae flatworm, Thysanozoon nigropapillosum, is somewhat cephalized, with a distinct head end (at right) which has pseudotentacles and an eyespot. Yellow papillae flatworm (Thysanozoon nigropapillosum) (cropped).jpg
The yellow papillae flatworm, Thysanozoon nigropapillosum , is somewhat cephalized, with a distinct head end (at right) which has pseudotentacles and an eyespot.

The Platyhelminthes (flatworms) have a more complex nervous system than the Acoela, and are lightly cephalized, for instance having an eyespot above the brain, near the front end. [4]

Complex active bodies

The philosopher Michael Trestman noted that three bilaterian phyla, namely the arthropods, the molluscs in the shape of the cephalopods, and the chordates, were distinctive in having "complex active bodies", something that the acoels and flatworms did not have. Any such animal, whether predator or prey, has to be aware of its environment—to catch its prey, or to evade its predators. These groups are exactly those that are most highly cephalized. [5] [6] These groups, however, are not closely related: in fact, they represent widely separated branches of the Bilateria, as shown on the phylogenetic tree; their lineages split hundreds of millions of years ago. Other (less cephalized) phyla are not shown, for clarity. [7] [8] [9]

Planulozoa

Cnidaria Aurelia aurita NASA.png

Bilateria

Acoela etc Neochildia fusca.jpg

Nephrozoa
Deuterostomia
Echinoderms

Portugal 20140812-DSC01434 (21371237591).jpg

Chordates
Tunicates  (larva)

Larval tunicate drawing.png

Vertebrates

White shark (Duane Raver).png

 Large brains
Protostomia
Ecdysozoa
Nematodes  etc

Female Phasmarhabditis hermaphrodita (cropped).jpg

Arthropods

Long nosed weevil edit.jpg

Running legs
Spiralia
Flatworms  etc

Sorocelis reticulosa.jpg

Cephalopods

Octopusvulgaris.jpg

Active arms
610 mya
650 mya
 Cephalization 
680 mya


Arthropods

In arthropods, cephalization progressed with the gradual incorporation of trunk segments into the head region. This was advantageous because it allowed for the evolution of more effective mouth-parts for capturing and processing food. Insects are strongly cephalized, their brain made of three fused ganglia attached to the ventral nerve cord, which in turn has a pair of ganglia in each segment of the thorax and abdomen. The insect head is an elaborate structure made of several segments fused rigidly together, and equipped with both simple and compound eyes, and multiple appendages including sensory antennae and complex mouthparts (maxillae and mandibles). [4]

Cephalopods like this cuttlefish have advanced 'camera' eyes. Cuttlefish eye.jpg
Cephalopods like this cuttlefish have advanced 'camera' eyes.

Cephalopods

Cephalopod molluscs including octopus, squid, cuttlefish and nautilus are the most intelligent and highly cephalized invertebrates, with well-developed senses, including advanced 'camera' eyes and large brains. [10] [11]

Vertebrates

Cephalization in vertebrates, the group that includes mammals, birds, reptiles, amphibians and fishes, has been studied extensively. [4] The heads of vertebrates are complex structures, with distinct sense organs for sight, olfaction, and hearing, and a large, multi-lobed brain protected by a skull of bone or cartilage.[ citation needed ] Cephalochordates like the lancelet, a small fishlike animal with very little cephalization, are closely related to vertebrates but do not have these structures. In the 1980s, the new head hypothesis proposed that the vertebrate head is an evolutionary novelty resulting from the emergence of neural crest and cranial placodes (thickened areas of ectoderm), which result in the formation of all senses outside of the brain. [12] [13] However, in 2014, a transient larva tissue of the lancelet was found to be virtually indistinguishable from the neural crest-derived cartilage (later bone, in jawed ones) which forms the vertebrate skull, suggesting that persistence of this tissue and expansion into the entire head space could be a viable evolutionary route to formation of the vertebrate head. [14] Advanced vertebrates have increasingly elaborate brains. [4]

Idealised vertebrate body plan, showing brain and sense organs at the head end Vertebrate body plan.svg
Idealised vertebrate body plan, showing brain and sense organs at the head end

Anterior Hox genes

Bilaterians have many more Hox genes controlling the development, including of the front of the body than do the less cephalized Cnidaria (two Hox clusters) and the Acoelomorpha (three Hox clusters). In the vertebrates, duplication resulted in the four Hox clusters (HoxA to HoxD) of mammals and birds, while another duplication gave teleost fishes eight Hox clusters. Some of these genes, those responsible for the front (anterior) of the body, helped to create the heads of both arthropods and vertebrates. However, the Hox1-5 genes were already present in ancestral arthropods and vertebrates that did not have complex head structures. The Hox genes therefore most likely assisted in cephalization of these two bilaterian groups independently by convergent evolution, resulting in similar gene networks. [15]

See also

Related Research Articles

<span class="mw-page-title-main">Brain</span> Organ central to the nervous system

The brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for special senses such as vision, hearing and olfaction. Being the most specialized organ, it is responsible for receiving information from the sensory nervous system, processing those information and the coordination of motor control.

<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 primarily of the brain and spinal cord. 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">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 (efferent), while those nerves that transmit information from the body to the CNS are called sensory nerves (afferent). The PNS is divided into two separate subsystems, the somatic and autonomic, nervous systems. The autonomic nervous system is further subdivided into the sympathetic, parasympathetic and enteric 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. Nerves that exit from the brain are called cranial nerves while those exiting from the spinal cord are called spinal nerves.

<span class="mw-page-title-main">Nemertea</span> Phylum of invertebrates, ribbon worms

Nemertea is a phylum of animals also known as ribbon worms or proboscis worms, consisting of about 1300 known species. Most ribbon worms are very slim, usually only a few millimeters wide, although a few have relatively short but wide bodies. Many have patterns of yellow, orange, red and green coloration. The foregut, stomach and intestine run a little below the midline of the body, the anus is at the tip of the tail, and the mouth is under the front. A little above the gut is the rhynchocoel, a cavity which mostly runs above the midline and ends a little short of the rear of the body. All species have a proboscis which lies in the rhynchocoel when inactive but everts to emerge just above the mouth to capture the animal's prey with venom. A highly extensible muscle in the back of the rhynchocoel pulls the proboscis in when an attack ends. A few species with stubby bodies filter feed and have suckers at the front and back ends, with which they attach to a host.

<span class="mw-page-title-main">Evolutionary developmental biology</span> Comparison of organism developmental processes

Evolutionary developmental biology is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved.

<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 have pentaradial symmetry as adults, but are only bilaterally symmetrical as an embryo. Cephalization is a characteristic feature among most bilaterians, where the special sense organs and central nerve ganglia become concentrated at the front end.

<span class="mw-page-title-main">Eye</span> Organ that detects light and converts it into electro-chemical impulses in neurons

An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system.

<span class="mw-page-title-main">Panarthropoda</span> Animal taxon

Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada and Onychophora. Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphyletic group where the last common ancestor and basal members (stem-group) of each extant panarthropod phylum are thought to have risen. However the term "Lobopodia" is sometimes expanded to include tardigrades and onychophorans as well.

<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">Acoelomorpha</span> Phylum of marine, flatworm-like animals

Acoelomorpha is a subphylum of very simple and small soft-bodied animals with planula-like features which live in marine or brackish waters. They usually live between grains of sediment, swimming as plankton, or crawling on other organisms, such as algae and corals. With the exception of two acoel freshwater species, all known acoelomorphs are marine.

Hox genes, a subset of homeobox genes, are a group of related genes that specify regions of the body plan of an embryo along the head-tail axis of animals. Hox proteins encode and specify the characteristics of 'position', ensuring that the correct structures form in the correct places of the body. For example, Hox genes in insects specify which appendages form on a segment, and Hox genes in vertebrates specify the types and shape of vertebrae that will form. In segmented animals, Hox proteins thus confer segmental or positional identity, but do not form the actual segments themselves.

<span class="mw-page-title-main">Body plan</span> Set of morphological features common to members of a phylum of animals

A body plan, Bauplan, or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many.

<span class="mw-page-title-main">Ventral nerve cord</span> Structure of the invertebrate central nervous system

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">Animal</span> Kingdom of living things

Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a hollow sphere of cells, the blastula, during embryonic development. Animals form a clade, meaning that they arose from a single common ancestor. Over 1.5 million living animal species have been described, of which around 1.05 million are insects, over 85,000 are molluscs, and around 65,000 are vertebrates. It has been estimated there are as many as 7.77 million animal species on Earth. Animal body lengths range from 8.5 μm (0.00033 in) to 33.6 m (110 ft). They have complex ecologies and interactions with each other and their environments, forming intricate food webs. The scientific study of animals is known as zoology, and the study of animal behaviour is known as ethology.

<span class="mw-page-title-main">Marine invertebrates</span> Marine animals without a vertebral column

Marine invertebrates are invertebrate animals that live in marine habitats, and make up most of the macroscopic life in the oceans. It is a polyphyletic blanket term that contains all marine animals except the marine vertebrates, including the non-vertebrate members of the phylum Chordata such as lancelets, sea squirts and salps. As the name suggests, marine invertebrates lack any mineralized axial endoskeleton, i.e. the vertebral column, and some have evolved a rigid shell, test or exoskeleton for protection and/or locomotion, while others rely on internal fluid pressure to support their bodies. Marine invertebrates have a large variety of body plans, and have been categorized into over 30 phyla.

<i>engrailed</i> (gene) Protein family

engrailed is a homeodomain transcription factor involved in many aspects of multicellular development. First known for its role in arthropod embryological development, working in consort with the Hox genes, engrailed has been found to be important in other areas of development. It has been identified in many bilaterians, including the arthropods, vertebrates, echinoderms, molluscs, nematodes, brachiopods, and polychaetes. It acts as a "selector" gene, conferring a specific identity to defined areas of the body, and co-ordinating the expression of downstream genes.

<span class="mw-page-title-main">Protostome</span> Clade of animals whose mouth develops before the anus

Protostomia is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's members, although the reverse is typically true of its sister clade, Deuterostomia. Well-known examples of protostomes are arthropods, molluscs, annelids, flatworms and nematodes. They are also called schizocoelomates since schizocoely typically occurs in them.

<span class="mw-page-title-main">Cephalopod eye</span> Visual sensory organs of cephalopod molluscs

Cephalopods, as active marine predators, possess sensory organs specialized for use in aquatic conditions. They have a camera-type eye which consists of an iris, a circular lens, vitreous cavity, pigment cells, and photoreceptor cells that translate light from the light-sensitive retina into nerve signals which travel along the optic nerve to the brain. For the past 140 years, the camera-type cephalopod eye has been compared with the vertebrate eye as an example of convergent evolution, where both types of organisms have independently evolved the camera-eye trait and both share similar functionality. Contention exists on whether this is truly convergent evolution or parallel evolution. Unlike the vertebrate camera eye, the cephalopods' form as invaginations of the body surface, and consequently the cornea lies over the top of the eye as opposed to being a structural part of the eye. Unlike the vertebrate eye, a cephalopod eye is focused through movement, much like the lens of a camera or telescope, rather than changing shape as the lens in the human eye does. The eye is approximately spherical, as is the lens, which is fully internal.

The evolution of nervous systems dates back to the first development of nervous systems in animals. 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. 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.

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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