338:. Phoresy describes a non-parasitic relationship between two organisms, where one organism uses the other as a mode of transportation. In phoresy, there is a phoront, which is then transported species, and the vector, the mobile species through which the phoront travels. It is crucial to the evolution of parasitism due to both its initiation of close contact between the phoront and vector, as well as being a constraint on parasite size. The stressors of phoresy and parasitism are closely related, such as desiccation and starvation. This reliance on the vector mirrors the reliance on a host, both of which act on the affected organism's fitness. The need for phoresy is also much lower in marine environments, as marine nematodes can utilize currents as methods of low-effort transport. Therefore, an additional connection is formed between terrestrial species and eventual parasitism.
353:. However, necromeny has been found to select traits that reinterpret the vector not simply as transport, but also as a habitat. It is important to note that necromeny does not necessarily eliminate the further need for phoresy. Because of this, it is thought that developing nematodes rely on both environmental signals, as well as communication with other larvae while making the choice between continuing development on their vector (necromeny) or attempting to find a new one (phoresy). For example, it has been found that dauers can communicate with other dauers via pheromones, in which adult nematodes signal larvae to continue their development. This can create a
110:, a population density cue, influence this dauer decision. Dauer larvae are thus considered an alternative L3 stage larva, and this stage is sometimes preceded by L2d. L2d animals are considered pre-dauer and are characterised by delayed development and dark intestines produced by storage of fat. L2d larvae can either continue normal development or enter dauer stage depending on whether the conditions that triggered their formation persist. Dauer is not, however, a permanent condition. In fact, if the food supply and the population density become optimal for growth the dauer larvae can exit this stage and become L4s and then adults.
266:. Models of parasitic evolution are difficult to confirm because they are difficult to test. Like other methods of studying evolution, researchers can make use of genomic data, specifically while comparing data from closely related, non-parasitic species. Parasitism is common, and it is even more common in nematodes, which have evolved into parasitism on up to eighteen separate occasions throughout their evolutionary history. This calls into question what exactly about the nematode leads to such an inclination toward parasitism.
377:, a genus of nematodes that harbors symbiotic bacteria that are highly pathogenic to hosts, but completely harmless to them. After the bacteria kill the host, they proliferate on the host's dead body. The Heterorhabditis then feeds on this new growth of bacteria for development. In both cases of feeding, the parasitic nematodes make direct use of the host's body, possible only through the evolutionary pathway aided by phoresy.
311:. Four steps of an evolutionary sequence pathway to animal parasitism have been proposed. The steps are as follows: 1.) Free-living ancestors that do not associate with a larger species, 2) phoretic relationships in which nematodes superficially attach to a larger animal for dispersal, 3) necromeny, in which nematodes may feed on their dead hosts without directly contributing to the death themselves, and 4) parasitism.
368:: Through the development of phoresy to necromeny, developing larvae can officially reach a state of parasitism in their adulthood. In parasitic nematodes, there are two main methods of feeding: direct feeding and indirect feeding. In direct feeding, nematodes switch from their ancestral food source, such as bacteria, to their host vector's tissue. They utilize
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is a major selective force in only terrestrial environments, which the larva will combat by dauer dormancy. Phylogenetic analysis of nematodes suggests that parasitic lineages are derived overwhelmingly from terrestrial ancestors, even with lineages that reside in water. Both of these factors are
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between different habitats through carrier animals. In both of these cases, the alternative stage is called the dauer. In parasitic species of nematodes, this alternative stage is called the “infective juvenile”, and facilitates transmission not between environments, but hosts. All three of these
62:
nematode has become the most studied nematode, the term ‘dauer stage’ or 'dauer larvae' is becoming universally recognised when referring to this state in other free-living nematodes. The dauer stage is also considered to be equivalent to the infective stage of
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for this process, by secreting them into the environment as opposed to internal use. However, in indirect feeding, nematodes weaponize bacteria to kill a host. For example, in George O. Poinar Jr's 1990 book on
Nematodes and Biological Control, he describes
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Dauer larvae generally remain motionless, but can react to touch or vibrations. They can stand on their tails, waving their bodies in the air, and attach themselves to any passing animals, particularly insects, enabling them to
102:. After L4, animals moult to the reproductive adult stage. However, when the environment is unfavorable, L1 and L2 animals have the option to divert their development from reproduction to dauer formation. Signals such as
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50:, whereby the larva goes into a type of stasis and can survive harsh conditions. Since the entrance of the dauer stage is dependent on environmental cues, it represents a classic and well studied example of
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dauer larvae can survive up to four months, much longer than their average lifespan of about three weeks during normal reproductive development. Two genes that are essential for dauer formation are
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Necromeny is most effectively thought of as a parasitic extension of phoresy, in which the phoront will feed on the vector if it dies in transit, as well as using the body as a place for
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Galles, Celina; Prez, Gastón M.; Penkov, Sider; Boland, Sebastian; Porta, Exequiel O. J.; Altabe, Silvia G.; Labadie, Guillermo R.; Schmidt, Ulrike; Knölker, Hans-Joachim (2018-04-23).
186:, the life extension effect can be uncoupled from dauer growth arrest. The lifespan increase was shown to be associated with an increase in stress resistance.
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Dauer larvae are extensively studied by biologists because of their ability to survive harsh environments and live for extended periods of time. For example,
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Nematodes can live both on land and in water, residing in both soil and underwater sediment. However, as found by
Rebecci et al. in their 2020 study,
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The next step in Crook's proposed plan is phoresy. Phoresy as a step for parasitism is not confined to nematode development and is seen similarly in
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optional stages share the common function of facilitating organism survival under states of high stress during larval stages and are similar in
649:"Natural variation in Pristionchus pacificus dauer formation reveals cross-preference rather than self-preference of nematode dauer pheromones"
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supported by the dauer
Hypothesis under the assumption that the dauer precedes the parasite, and is not influenced by earlier sources.
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282:, leading them to an optional alternative life stage during times of high stress. In some species this alternative stage leads to
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286:, pausing organism development until conditions are more favorable, and in others that alternative stage is used for group
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and that the parasitic “infective juvenile” life stage is derived from the ancestral, non-parasitic dauer larva.
698:"daf-2, daf-16 and daf-23: genetically interacting genes controlling Dauer formation in Caenorhabditis elegans"
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802:"Thermotolerance and extended life-span conferred by single-gene mutations and induced by thermal stress"
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The hypothesis was developed from the observation that roundworms, or nematodes, undergo the same four
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Synthesis and
Activity of Dafachronic Acid Ligands for the C. elegans DAF-12 Nuclear Hormone Receptor
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Neue parasitische und halbparasitische
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inhibit the dauer formation caused by PUFA deficiency or impaired cholesterol trafficking.
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Dafadine inhibits DAF-9 to promote dauer formation and longevity of
Caenorhabditis elegans
1200:"Caenorhabditis elegans dauers vary recovery in response to bacteria from natural habitat"
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New
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209:. For example, dauer larvae of rhabditids are often found in parallel rows under the
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1441:"An excreted small molecule promotes C. elegans reproductive development and aging"
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579:"Hormone Signaling and Phenotypic Plasticity in Nematode Development and Evolution"
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and colleagues as being required for extended longevity seen in animals that lack
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first proposed in 1899 and 1900, all nematodes have five stages separated by four
54:. The dauer state is given other names in the various types of nematodes such as ‘
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stages, some species only differing by having extra components to their
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Broadly, the Dauer hypothesis applies to all examples of parasitism in
295:. From this, the Dauer Hypothesis suggests that these three stages are
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lineages evolved into parasites through two major steps,
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A characteristic of the dauer stage is the pronounced
147:. In favorable environments, DAF-12 is activated by a
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58:’ or ‘hypobiosis’, but since the
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716:10.1093/genetics/137.1.107
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129:. Dauer formation in
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242:The dauer hypothesis
142:transcription factor
40:worms, particularly
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866:"The Dauer Cuticle"
820:1995PNAS...92.7540L
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320:desiccation
227:cholesterol
104:temperature
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399:References
297:homologous
293:morphology
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280:life cycle
252:parasitism
248:hypothesis
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1287:209380774
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366:Parasitsm
347:Necromeny
336:Astigmata
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256:nematode
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520:"Dauer"
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355:habitat
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