 | This article may be too technical for most readers to understand.(September 2024) |
The protocerebrum is the first segment of the panarthropod brain.
Recent studies suggest that it comprises two regions. [1] : 5
six3 is a transcription factor that marks the anteriormost part of the developing body in a whole host of Metazoa. [1]
In the panarthropod brain, the anteriormost (rostralmost) part of the germband expresses six3. This region is described as medial, and corresponds to the annelid prostomium. [1]
In arthropods, it contains the pars intercerebralis and pars lateralis. [1]
six3 is associated with the euarthropod labrum [2] and the onychophoran frontal appendages (antennae).
The other region expresses homologues of orthodenticle, Otx or otd. This region is more caudal and lateral, and bears the eyes. [1] [3]
Orthodenticle is associated with the protocerebral bridge, part of the central complex, traditionally a marker of the prosocerebrum. [4]
In the annelid brain, Otx expression characterises the peristomium, but also creeps forwards into the regions of the prostomium that bear the larval eyes. [1]
Inconsistent use of the terms archicerebrum and the prosocerebrum makes them confusing. [1] : 5
The regions were defined by Siewing (1963): the archicerebrum as containing the ocular lobes and the mushroom bodies (= corpora pedunculata), and the prosocerebrum as comprising the central complex. [5] [6]
The archicerebrum has traditionally been equated with the anteriormost, 'non-segmental' part of the protocerebrum, equivalent to the acron in older terminology. The prosocerebrum is then equivalent to the 'segmental' part of the protocerebrum, bordered by segment polarity genes such as engrailed, and (on one interpretation) bearing modified segmental appendages (= camera-type eyes).
But Urbach and Technau (2003) complicate the matter by seeing the prosocerebrum (central complex) + labrum as the anteriormost region
{{cite journal}}: CS1 maint: multiple names: authors list (link)An alternative usage of the terms seems to have been introduced by Steinmetz et al (2010), who seem to flip the terms?
protocerebrum, can be further subdivided into a more lateral region bearing, for example, the optic lobes (archicerebrum) and a median region that includes, for example, the pars intercerebralis (prosocerebrum). Most authors think that the archicerebrum represents the tip of the neuraxis, but this has been disputed."
Lobopodians are members of the informal group Lobopodia, or the formally erected phylum Lobopoda Cavalier-Smith (1998). They are panarthropods with stubby legs called lobopods, a term which may also be used as a common name of this group as well. While the definition of lobopodians may differ between literatures, it usually refers to a group of soft-bodied, marine worm-like fossil panarthropods such as Aysheaia and Hallucigenia. However, other genera like Kerygmachela and Pambdelurion are often referred to as “gilled lobopodians”.
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.
The mushroom bodies or corpora pedunculata are a pair of structures in the brain of arthropods, including insects and crustaceans, and some annelids. They are known to play a role in olfactory learning and memory. In most insects, the mushroom bodies and the lateral horn are the two higher brain regions that receive olfactory information from the antennal lobe via projection neurons. They were first identified and described by French biologist Félix Dujardin in 1850.
Dinocaridida is a proposed fossil taxon of basal arthropods, which flourished during the Cambrian period and survived up to Early Devonian. Characterized by a pair of frontal appendages and series of body flaps, the name of Dinocaridids refers to the suggested role of some of these members as the largest marine predators of their time. Dinocaridids are occasionally referred to as the 'AOPK group' by some literatures, as the group compose of Radiodonta, Opabiniidae, and the "gilled lobopodians" Pambdelurion and Kerygmachelidae. It is most likely paraphyletic, with Kerygmachelidae and Pambdelurion more basal than the clade compose of Opabiniidae, Radiodonta and other arthropods.
The supraesophageal ganglion is the first part of the arthropod, especially insect, 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, each a pair of ganglia that may be more or less pronounced, reduced, or fused depending on the genus:
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.
Kerygmachela kierkegaardi is a kerygmachelid gilled lobopodian from the Cambrian Stage 3 aged Sirius Passet Lagerstätte in northern Greenland. Its anatomy strongly suggests that it, along with its relative Pambdelurion whittingtoni, was a close relative of radiodont and euarthropods. The generic name "Kerygmachela" derives from the Greek words Kerygma (proclamation) and Chela (claw), in reference to the flamboyant frontal appendages. The specific name, "kierkegaardi" honors Danish philosopher Søren Kierkegaard.
The (pan)arthropod head problem is a long-standing zoological dispute concerning the segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other. While the dispute has historically centered on the exact make-up of the insect head, it has been widened to include other living arthropods, such as chelicerates, myriapods, and crustaceans, as well as fossil forms, such as the many arthropods known from exceptionally preserved Cambrian faunas. While the topic has classically been based on insect embryology, in recent years a great deal of developmental molecular data has become available. Dozens of more or less distinct solutions to the problem, dating back to at least 1897, have been published, including several in the 2000s.
Fuxianhuia is a genus of Lower Cambrian fossil arthropod known from the Chengjiang fauna in China. Its purportedly primitive features have led to its playing a pivotal role in discussions about the euarthropod stem group. Nevertheless, despite being known from many specimens, disputes about its morphology, in particular its head appendages, have made it one of the most controversial of the Chengjiang taxa, and it has been discussed extensively in the context of the arthropod head problem.
Homeobox protein SIX3 is a protein that in humans is encoded by the SIX3 gene.
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.
Kenyon cells are the intrinsic neurons of the mushroom body, a neuropil found in the brains of most arthropods and some annelids. They were first described by F. C. Kenyon in 1896. The number of Kenyon cells in an organism varies greatly between species. For example, in the fruit fly, Drosophila melanogaster, there are about 2,500 Kenyon cells per mushroom body, while in cockroaches there are about 230,000.
Megacheira is an extinct class of predatory arthropods defined by their possession of spined "great appendages". Their taxonomic position is controversial, with studies either considering them stem-group euarthropods, or stem-group chelicerates. The homology of the great appendages to the cephalic appendages of other arthropods is also controversial. Uncontested members of the group were present in marine environments worldwide from the lower to middle Cambrian.
Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. They may be referred to as radiodonts, radiodontans, radiodontids, anomalocarids, or anomalocaridids, although the last two originally refer to the family Anomalocarididae, which previously included all species of this order but is now restricted to only a few species. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and used for a variety of functions. Radiodonts included the earliest large predators known, but they also included sediment sifters and filter feeders. Some of the most famous species of radiodonts are the Cambrian taxa Anomalocaris canadensis, Hurdia victoria, Peytoia nathorsti, Titanokorys gainesi, Cambroraster falcatus and Amplectobelua symbrachiata. The later surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco and the Early Devonian member Schinderhannes bartelsi from Germany.
The cephalon is the head section of an arthropod. It is a tagma, i.e., a specialized grouping of arthropod segments. The word cephalon derives from the Greek κεφαλή (kephalē), meaning "head".
The labrum is a flap-like structure that lies immediately in front of the mouth in almost all extant Euarthropoda. The most conspicuous exceptions are the Pycnogonida, which probably are chelicerates. In entomology, the labrum amounts to the "upper lip" of an insect mouth, the corresponding "lower lip" being the labium.
Cardiodictyon is a genus of lobopodian known from 518 millions years old Chengjiang Lagerstätte. 525 millions years old partial fossil is also reported. It has ~25 pairs of legs, each associated with a pair of dorsal plates.
Lyrarapax is a radiodont genus of the family Amplectobeluidae that lived in the early Cambrian period 520 million years ago. Its neural tissue indicates that the radiodont frontal appendage is protocerebral, resolving parts of the arthropod head problem and showing that the frontal appendage is homologous to the antennae of Onychophorans and labrum of euarthropods. Its fossilized remains were found in Yunnan in southwestern China. A second species was described in 2016, differing principally in the morphology of its frontal appendages. It is a small animal, with L. unguispinus and L. trilobus measuring up to 8.3 cm (3.3 in) and 3.9 cm (1.5 in) in body length respectively.
Cheloniellon is a monotypic genus of cheloniellid arthropod, known only by one species, Cheloniellon calmani, discovered from the Lower Devonian Hunsrück Slate of Germany.
Deuteropoda is a proposed clade of arthropods whose members are distinguished from more basal stem-group arthropods like radiodonts by an anatomical reorganization of the head region, namely the appearance of a differentiated first appendage pair, a multisegmented head, a hypostome/labrum complex, and by bearing pairs of segmented biramous limbs.
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