Cephalization is an evolutionary trend in which, over many generations, the mouth, sense organs, and nerve ganglia become concentrated at the front end of an animal, producing a head region. This is associated with movement and bilateral symmetry, such that the animal has a definite head end. This led to the formation of a highly sophisticated brain in three groups of animals, namely the arthropods, cephalopod molluscs, and vertebrates.
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]
Cephalization is a characteristic feature of the Bilateria, a large group containing the majority of animal phyla. [2] 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. [3]
The Acoela are basal bilaterians, part of the Xenacoelomorpha. They are small and simple animals, and have very 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]
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]
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 |
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| 680 mya |
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]
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]
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 (sight by camera-type eyes, olfaction by nostrils, taste by taste buds, balance & hearing by otic vesicle); a large, multi-lobed brain (three-part at least); and later, in jawed ones, teeth and, in tetrapods, tongue (teeth like keratinous structures and tongue arose independently in jawless lampreys). Cephalochordates like Branchiostoma (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]
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.
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.
Bilateria is a large clade/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.
Tullimonstrum, colloquially known as the Tully monster or sometimes Tully's monster, is an extinct genus of soft-bodied bilaterian animal that lived in shallow tropical coastal waters of muddy estuaries during the Pennsylvanian geological period, about 300 million years ago. A single species, T. gregarium, is known. Examples of Tullimonstrum have been found only in the Essex biota, a smaller section of the Mazon Creek fossil beds of Illinois, United States. Its classification has been the subject of controversy, and interpretations of the fossil have likened it to molluscs, arthropods, conodonts, worms, tunicates, and vertebrates. This creature had a mostly cigar shaped body, with a triangular tail fin, two long stalked eyes, and a proboscis tipped with a mouth-like appendage. Based on the fossils, it seems this creature was a nektonic carnivore that hunted in the ocean’s water column. When Tullimonstrum was alive, Illinois was a mixture of ecosystems like muddy estuaries, marine environments, and rivers and lakes. Fossils of other organisms like crustacean Belotelson, the cnidarian Essexella, and the elasmobranch fish Bandringa have been found alongside Tullimonstrum.
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.
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.
A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may not have a head, but many bilaterally symmetric forms do, regardless of size.
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.
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.
Acoela, or the acoels, is an order of small and simple invertebrates in the subphylum Acoelomorpha of phylum Xenacoelomorpha, a deep branching bilaterian group of animals, which resemble flatworms. Historically they were treated as an order of turbellarian flatworms.
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. As of 2022, 2.16 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 around 7.77 million animal species. Animals range in length from 8.5 micrometres (0.00033 in) to 33.6 metres (110 ft). They have complex interactions with each other and their environments, forming intricate food webs. The scientific study of animals is known as zoology.
Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have evolved a shell or a hard exoskeleton. As on land and in the air, marine invertebrates have a large variety of body plans, and have been categorised into over 30 phyla. They make up most of the macroscopic life in the oceans.
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.
The urbilaterian is the hypothetical last common ancestor of the bilaterian clade, i.e., all animals having a bilateral symmetry.
Deuterostomes are bilaterian animals of the superphylum Deuterostomia, typically characterized by their anus forming before the mouth during embryonic development. The three major clades of extant deuterostomes include chordates, echinoderms and hemichordates.
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.
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.
Nephrozoa is a major clade of bilaterians, divided into the protostomes and the deuterostomes, containing almost all animal phyla and over a million extant species. Its sister clade is the Xenacoelomorpha. The Ambulacraria are occasionally thought to be sister to the Xenacoelomorpha, forming the Xenambulacraria as basal Deuterostomia, or basal Bilateria invalidating Nephrozoa and Deuterostomia in multiple studies. The coelom, the digestive tract and excretory organs (nephridia), and nerve cords developed in the Nephrozoa. It has been argued that, because protonephridia are only found in protostomes, they cannot be considered a synapomorphy of this group. This would make Nephrozoa an improper name, leaving Eubilateria as this clade's name.
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.
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.
Ocelli located at the base of the many tentacles represent one input to the B system, whereas the neurons of the O system are directly photosensitive. Many hydromedusae have ocelli of different levels of complexity (Singla, 1974). In addition, other marginal sensory structures associated with the outer nerve ring include statocysts (Singla, 1975), and mechanoreceptors, such as the tactile combs of Aglantha, which are located at the tentacle bases, and can activate the escape swimming circuitry (Arkett & Mackie, 1988; Mackie, 2004b).
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