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Nematode

The nematodes, roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (helminths) are the cause of soil-transmitted helminthiases.

Etymology
The word nematode comes from the Modern Latin compound of nema- 'thread' (from Greek nema, genitive nematos 'thread', from the stem nein 'to spin'; cf. needle) + -odes 'like, of the nature of' (cf. -oid). The addition firstly of '-oid' and then to '-ode' renders 'threadlike'. == Taxonomy and systematics==
Taxonomy and systematics
File:Eophasma jurasicum.JPG|Eophasma jurasicum, a fossilized nematode File:Celegans wt nhr80rnai.png|Caenorhabditis elegans File:Hookworms.JPG|Rhabditia File:Gravid adult female Nippostrongylus brasiliensis - image.pntd.v07.i08.g001.png|Nippostrongylus brasiliensis File:Anisakids.jpg|Unidentified Anisakidae (Ascaridina: Ascaridoidea) File:Threadworm.jpg|Oxyuridae Pinworm File:Microfilaria.jpg|Spiruridae Dirofilaria immitis History described nematodes including the parasitic Dracunculus medinensis, seen here under a person's skin. The name of the group Nematoda, informally called "nematodes", came from Nematoidea, originally defined by Karl Rudolphi in 1808, from Ancient Greek νῆμα (nêma, nêmatos, 'thread') and -ειδής (-eidēs, 'species') (cf. native German Fadenwurm In 1919, Nathan Cobb proposed that nematodes should be recognized alone as a phylum. He argued they should be called "nema" in English rather than "nematodes" and defined the taxon Nemates (later emended as Nemata, Latin plural of nema), listing Nematoidea sensu restricto as a synonym. In 1932, Potts elevated the class Nematoda to the level of phylum, leaving the name the same. Although Potts' and Cobb's classifications are equivalent, both names are used, and Nematode became a popular term in zoological science. However, some authors continued to accept phylum Aschelminthes in the second half of the 20th century. Phylogeny The phylogenetic relationships of the nematodes and their close relatives among the protostomes are still discussed. Initially, some morphology-based phylogenetic classifications accepted Aschelmithes/Nemathelminthes as a legitimate group. In the 1990s, nematodes were proposed to form the group Ecdysozoa together with moulting animals, such as arthropods. The identity of the closest living relatives of the Nematoda has always been considered to be well resolved. Morphological and molecular phylogenetics agree with placing the roundworms as a sister taxon to the parasitic Nematomorpha; together, they make up the Nematoida. Along with the Scalidophora (formerly Cephalorhyncha), the Nematoida might form the clade Cycloneuralia, but much disagreement occurs both between and among the available morphological and molecular data. The Cycloneuralia or the Introverta—depending on the validity of the former—are often ranked as a superphylum. Recent molecular analyses have placed Nematoida closer to Panarthropoda than to Scalidophora, forming clade Cryptovermes with the former. However, a few studies have supported alternative hypotheses, grouping nematodes with Tardigrada or with Loricifera. Advancements in molecular phylogeny have also clarified the internal branchings of Nematoda. Systematics Due to the lack of knowledge regarding many nematodes, their systematics is contentious. An early and influential classification was proposed by Chitwood and Chitwood—later revised by Chitwood—who divided the phylum into two classes—Aphasmidia and Phasmidia. These were later renamed Adenophorea (gland bearers) and Secernentea (secretors), respectively. The Secernentea share several characteristics, including the presence of phasmids, a pair of sensory organs located in the lateral posterior region, and this was used as the basis for this division. This scheme was adhered to in many later classifications, though the Adenophorea were not in a uniform group. Initial studies of incomplete DNA sequences suggested the existence of five clades: • DorylaimidaEnopliaSpirurinaTylenchinaRhabditina The Secernentea seem to be a natural group of close relatives, while the Adenophorea appear to be a paraphyletic assemblage of roundworms that retain a good number of ancestral traits. The old Enoplia do not seem to be monophyletic, either, but do contain two distinct lineages. The old group Chromadorea seems to be another paraphyletic assemblage, with the Monhysterida representing a very ancient minor group of nematodes. Among the Secernentea, the Diplogasteria may need to be united with the Rhabditia, while the Tylenchia might be paraphyletic with the Rhabditia. The understanding of roundworm systematics and phylogeny as of 2002 is summarised below: Phylum NematodaBasal order Monhysterida • Class Dorylaimida • Class Enoplea • Class Secernentea • Subclass Diplogasteria (disputed) • Subclass Rhabditia (paraphyletic?) • Subclass Spiruria • Subclass Tylenchia (disputed) • "Chromadorea" assemblage Later work has suggested the presence of 12 clades. In 2019, a study identified one conserved signature indel (CSI) found exclusively in members of the phylum Nematoda through comparative genetic analyses. The CSI consists of a single amino acid insertion within a conserved region of a Na(+)/H(+) exchange regulatory factor protein NRFL-1 and is a molecular marker that distinguishes the phylum from other species. • subclass Dorylaimia • orders Rhabditida, Trichinellida and Mermithida • suborder Rhabditina • infraorders Spiruromorpha and Oxyuridomorpha In 2022 a new classification of the entire phylum Nematoda was presented by M. Hodda. It was based on current molecular, developmental and morphological evidence. Under this classification, the classes and subclasses are: • Class Enoplea • Subclass Enoplia • Subclass Oncholaimia • Subclass Triplonchia • Class Dorylaimida • Subclass Dorylaimia • Subclass Bathyodontia • Subclass Trichocephalia • Class Chromadorea • Subclass Chromadoria • Subclass Plectia == Fossil record==
Fossil record
Nematode eggs from the clades Ascaridina, Spirurina, and Trichocephalida have been discovered in coprolites from the Oligocene-aged Tremembé Formation, which represented a palaeolake in present-day São Paulo with a diverse fossil assemblage of birds, fish, and arthropods that lent itself to fostering high nematode diversity. Nematodes have also been found in various lagerstätten, such as Burmese amber, the Moltrasio Formation, and the Rhynie chert, where the earliest known fossils are known from. == Anatomy==
Anatomy
''. The large circles filled with small green circles are the uterus and eggs. The long narrow feature is the digestive tract. The smaller red and orange circles are the ovaries and oviducts. The cluster of green and black blobs in the upper right and lower left are the nerve cords (ventral and dorsal). Surrounding the internal organs are the frilly green longitudinal muscles, the dark hypodermis, and the green outer cuticle. Nematodes are very small, slender worms. Most are free-living, often less than 2.5 mm long and some only about 1 mm. Many nematodes are microscopic. Some soil nematodes can reach up to 7 mm in length, and some marine species can reach up to 5 cm. Some are parasitic and can reach lengths of 50 cm or more. The body is often ornamented with ridges, rings, bristles, or other distinctive structures. The head is relatively distinct. Whereas the rest of the body is bilaterally symmetrical, the head is radially symmetrical, with sensory bristles and, in many cases, solid 'head-shields' radiating outwards around the mouth. The mouth has either three or six lips, which often bear a series of teeth on their inner edges. An adhesive 'caudal gland' is often found at the tip of the tail. Each nerve lies within a cord of connective tissue lying beneath the cuticle and between the muscle cells. The ventral nerve is the largest, and has a double structure forward of the excretory pore. The dorsal nerve is responsible for motor control, while the lateral nerves are sensory, and the ventral combines both functions. The body is covered in numerous sensory bristles and papillae that together provide a sense of touch. Behind the sensory bristles on the head lie two small pits, or 'amphids'. These are well supplied with nerve cells and are probably chemoreception organs. A few aquatic nematodes possess what appear to be pigmented eye-spots, but whether or not these are actually sensory in nature is unclear. == Reproduction==
Reproduction
, used for copulation, bar=100 μm Most nematode species are dioecious, with separate male and female individuals, though some, such as Caenorhabditis elegans, are androdioecious, consisting of hermaphrodites and rare males. Both sexes possess one or two tubular gonads. In males, the sperm are produced at the end of the gonad and migrate along its length as they mature. The testis opens into a relatively wide seminal vesicle and then during intercourse into a glandular and muscular ejaculatory duct associated with the vas deferens and cloaca. In females, the ovaries each open into an oviduct (in hermaphrodites, the eggs enter a spermatheca first) and then a glandular uterus. The uteri both open into a common vulva/vagina, usually located in the middle of the morphologically ventral surface. Eggs may be embryonated or unembryonated when passed by the female, meaning their fertilized eggs may not yet be developed. A few species are known to be ovoviviparous. The eggs are protected by an outer shell, secreted by the uterus. In free-living roundworms, the eggs hatch into larvae, which appear essentially identical to the adults, except for an underdeveloped reproductive system; in parasitic roundworms, the lifecycle is often much more complicated. Nematodes as a whole possess a wide range of modes of reproduction. Some nematodes, such as Heterorhabditis spp., undergo a process called endotokia matricida: intrauterine birth causing maternal death. Some nematodes are hermaphroditic, and keep their self-fertilized eggs inside the uterus until they hatch. The juvenile nematodes then ingest the parent nematode. This process is significantly promoted in environments with a low food supply. which is otherwise very rare among animals. The single genus Meloidogyne (root-knot nematodes) exhibits a range of reproductive modes, including sexual reproduction, facultative sexuality (in which most, but not all, generations reproduce asexually), and both meiotic and mitotic parthenogenesis. The genus Mesorhabditis exhibits an unusual form of parthenogenesis, in which sperm-producing males copulate with females, but the sperm do not fuse with the ovum. Contact with the sperm is essential for the ovum to begin dividing, but because no fusion of the cells occurs, the male contributes no genetic material to the offspring, which are essentially clones of the female. ==Aging==
Aging
The nematode Caenorhabditis elegans is often used as a model organism for studying aging at the molecular level. For example, in C. elegans aging negatively impacts DNA repair, and mutants of C. elegans that are long-lived were shown to have increased DNA repair capability. These findings suggest a genetically determined correlation between DNA repair capacity and lifespan. == Free-living species==
Free-living species
Different free-living species feed on materials as varied as bacteria, algae, fungi, small animals, fecal matter, dead organisms, and living tissues. Free-living marine nematodes are important and abundant members of the meiobenthos. They play an important role in the decomposition process, aid in recycling of nutrients in marine environments, and are sensitive to changes in the environment caused by pollution. One roundworm of note, C. elegans, lives in the soil and has found much use as a model organism. C. elegans has had its entire genome sequenced, the developmental fate of every cell determined, and every neuron mapped. == Parasitic species==
Parasitic species
parasitic (mostly) nematodes from stools of Old World monkeys Nematodes that commonly parasitise humans include ascarids (Ascaris), filarias, hookworms, pinworms (Enterobius), and whipworms (Trichuris trichiura). The species Trichinella spiralis, commonly known as the trichina worm, occurs in rats, pigs, bears, and humans, and is responsible for the disease trichinosis. Baylisascaris usually infests wild animals, but can be deadly to humans, as well. Dirofilaria immitis is known for causing heartworm disease by inhabiting the hearts, arteries, and lungs of dogs and some cats. Haemonchus contortus is one of the most abundant infectious agents in sheep around the world, causing great economic damage to sheep. In contrast, entomopathogenic nematodes parasitize insects and are mostly considered beneficial by humans, but some attack beneficial insects. One form of nematode is entirely dependent upon fig wasps, which are the sole source of fig fertilization. They prey upon the wasps, riding them from the ripe fig of the wasp's birth to the fig flower of its death, where they kill the wasp, and their offspring await the birth of the next generation of wasps as the fig ripens. of soybean cyst nematode (Heterodera glycines) and egg A parasitic tetradonematid nematode discovered in 2005, Myrmeconema neotropicum, induces fruit mimicry in the tropical ant Cephalotes atratus. Infected ants develop bright red gasters (abdomens), tend to be more sluggish, and walk with their gasters in a conspicuous elevated position. These changes likely cause frugivorous birds to confuse the infected ants for berries, and eat them. Parasite eggs passed in the bird's feces are subsequently collected by foraging C. atratus and are fed to their larvae, thus completing the lifecycle of M. neotropicum. Similarly, multiple varieties of nematodes have been found in the abdominal cavities of the primitively social sweat bee, Lasioglossum zephyrus. Inside the female body, the nematode hinders ovarian development and renders the bee less active, thus less effective in pollen collection. Agriculture and horticulture Depending on its species, a nematode may be beneficial or detrimental to plant health. From agricultural and horticulture perspectives, the two categories of nematodes are the predatory ones, which kill garden pests; and the pest nematodes, which attack plants, or act as vectors spreading plant viruses between crop plants. Predatory nematodes include Phasmarhabditis hermaphrodita which is a lethal parasite of gastropods such as slugs and snails. Some members of the genus Steinernema such as Steinernema carpocapsae and Steinernema riobrave are generalist parasites of webworms, cutworms, armyworms, girdlers, some weevils, wood-borers and corn earworm moths. These organisms are grown commercially as biological pest control agents which can be used as an alternative to pesticides; their use is considered very safe. Plant-parasitic nematodes include several groups causing severe crop losses, taking 10% of crops worldwide every year. The most common genera are Aphelenchoides (foliar nematodes), Ditylenchus, Globodera (potato cyst nematodes), Heterodera (soybean cyst nematodes), Longidorus, Meloidogyne (root-knot nematodes), Nacobbus, Pratylenchus (lesion nematodes), Trichodorus, and Xiphinema (dagger nematodes). Several phytoparasitic nematode species cause histological damages to roots, including the formation of visible galls (e.g. by root-knot nematodes), which are useful characters for their diagnostic in the field. Some nematode species transmit plant viruses through their feeding activity on roots. One of them is Xiphinema index, vector of grapevine fanleaf virus, an important disease of grapes, another one is Xiphinema diversicaudatum, vector of arabis mosaic virus. Other nematodes attack bark and forest trees. The most important representative of this group is Bursaphelenchus xylophilus, the pine wood nematode, present in Asia and America and recently discovered in Europe. This nematode is transmitted from tree to tree by sawyer beetles (Monochamus). Greenhouse growers use entomopathogenic nematodes as beneficial agents to control fungus gnats. The nematodes enter the larvae of the gnats by way of their anus, mouth, and spiracles (breathing pores) and then release bacteria which kills the gnat larvae. Commonly used nematode species to control pests on greenhouse crops include Steinernema feltiae for fungus gnats and western flower thrips, Steinernema carpocapsae used to control shore flies, Steinernema kraussei for control of black vine weevils, and Heterorhabditis bacteriophora to control beetle larvae. Rotations of plants with nematode-resistant species or varieties is one means of managing parasitic nematode infestations. For example, planting Tagetes marigolds as a cover crop just prior to planting a nematode-susceptible plant, has been shown to suppress nematodes. Another approach involves using natural antagonists, particularly bacteria and fungi, which have proven effective in suppressing plant-parasitic nematodes, such as the fungus Gliocladium roseum. Chitosan, a natural biocontrol, elicits plant defense responses to destroy parasitic cyst nematodes on roots of soybean, maize, sugar beet, potato, and tomato crops without harming beneficial nematodes in the soil. Soil steaming is an efficient method to kill nematodes before planting a crop, but indiscriminately eliminates both harmful and beneficial soil fauna. The golden nematode Globodera rostochiensis is a particularly harmful pest that has resulted in quarantines and crop failures worldwide. It can be controlled, however. CSIRO, the scientific research body of the Australian government, found a 13- to 14-fold reduction of nematode population densities in plots having Chinese mustard Brassica juncea green manure or seed meal in the soil. Disease in humans for intestinal nematode infections per 100,000 in 2002. effect of papain on Heligmosomoides bakeri A number of pathogenic intestinal nematodes cause diseases in humans, including ascariasis, trichuriasis, and hookworm disease. Anisakis species parasitise fish and marine mammals and when consumed by humans can cause anisakiasis, a gastric or gastroallergic disease. Gastrointestinal nematode infections in humans are common, with approximately 50% of the global population being affected. Developing countries are most heavily impacted, in part due to lack of access to medical care. Trichinosis starts in the intestines but larvae can migrate to muscle. Filarial nematodes cause filariases. Toxocariasis is a zoonotic infection caused by roundworms passed from dogs, and sometimes cats. It can give rise to different types of larva migrans, such as visceral larva migrans and ocular larva migrans. Studies have shown that parasitic nematodes infect American eels, causing damage to the eel's swim bladder, dairy animals like cattle and buffalo, and all species of sheep. == Soil ecosystems==
Soil ecosystems
About 90% of nematodes reside in the top 15 cm (6") of soil. Nematodes do not decompose organic matter, but, instead, are parasitic and free-living organisms that feed on living material. Nematodes can effectively regulate bacterial population and community composition—they may eat up to 5,000 bacteria per minute. Also, nematodes can play an important role in the nitrogen cycle by way of nitrogen mineralization. But plant parasitic nematodes cause billions of dollars in annual crop damage worldwide. One group of carnivorous fungi, the nematophagous fungi, are predators of soil nematodes. They can set enticements for the nematodes in the form of lassos or adhesive structures. They can also release powerful toxins when in contact with nematodes. == Survivability==
Survivability
The nematode Caenorhabditis elegans an important model organism, was used as part of an ongoing research project conducted on the 2003 Space Shuttle Columbia mission STS-107, and survived the re-entry breakup. It is believed to be the first known species to survive a virtually unprotected atmospheric descent to Earth's surface. The Antarctic nematode Panagrolaimus davidi was able to withstand intracellular freezing depending on how well it had been fed. In 2023 an individual of Panagrolaimus kolymaensis was revived after 46,000 years in Siberian permafrost. == See also==
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