Dauer larva
Dauer (
As E. Maupas first proposed in 1899-1900, all nematodes have five stages separated by four moults.[2] Under environmental conditions that are favorable for reproduction, C. elegans larvae develop through four stages or moults which are designated as L1, L2, L3 and L4. 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 temperature, food supply, and levels of a dauer-inducing pheromone, 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.[6]
Dauer larvae are extensively studied by biologists because of their ability to survive harsh environments and live for extended periods of time. For example,
A characteristic of the dauer stage is the pronounced
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
Parasitism in dauer larva
The Dauer Hypothesis
The dauer
Theory Development
The hypothesis was developed from the observation that roundworms, or nematodes, undergo the same four
A Theory for Parasitic Evolution
Broadly, the Dauer Hypothesis applies to all examples of parasitism in
- Non-Association 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, desiccation is a major selective force in only terrestrial environments, which the larva will combat by dauer dormancy.[18] Phylogenetic analysis of nematodes suggests that parasitic lineages are derived overwhelmingly from terrestrial ancestors, even with lineages that reside in water.[19][20] Both of these factors are supported by the dauer Hypothesis under the assumption that the dauer precedes the parasite, and is not influenced by earlier sources.
- Phoresy 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 Astigmata.[21] 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.[17] 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.[22]Therefore, an additional connection is formed between terrestrial species and eventual parasitism.
- Necromeny 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 proliferation.[16] 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.[23] This can create a habitat shift in group environments, and can further parasitic larvae development.
- Parasitsm: 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 digestive enzymes for this process, by secreting them into the environment as opposed to internal use.[24] 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 Heterorhabditis, 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.[13]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.
See also
References
- ^ Fuchs, Anton Gilbert (1937). Neue parasitische und halbparasitische Nematoden bei Borkenkäfern und einige andere Nematoden [New Parasitic and Half-parasitic Nematodes with Bark-Beetles and Some Other Nematodes] (in German). Fischer.
- ^ ISBN 978-0-85199-786-5.
- S2CID 4255657.
- PMID 17988074. Retrieved 2009-11-05.
- PMID 21959166.
- ^ RIDDLE, D.. 12 The Dauer Larva. Cold Spring Harbor Monograph Archive, North America, 17 January 1988. Available at: https://cshmonographs.org/index.php/monographs/article/view/5027/4126. Date accessed: 14 July 2016.
- PMID 21307052.
- PMID 8056303.
- S2CID 4332206.
- PMID 7638227.
- PMID 29686301.
- PMID 28274802.
- ^ ISBN 9781351088640. Retrieved 6 December 2023.
- S2CID 12869939. Retrieved 6 December 2023.
- ^ Kiontke, K. "Nematodes". Current Biology. Retrieved 6 December 2023.
- ^ PMID 24095839.
- ^ PMID 33005351.
- S2CID 209380774.
- PMID 24229807.
- doi:10.1139/z84-050.
- . Retrieved 6 December 2023.
- ^ Heip, C.H.R (1985). The ecology of marine nematodes. Oceanography and Marine Biology: An Annual Review. pp. 399–489. Retrieved 6 December 2023.
- PMID 31320757.
- PMID 24963797.