Dauer larva

Dauer (German for "endurance", "duration" or "permanent") describes an alternative developmental stage of nematode worms, particularly Caenorhabditis elegans, whereby the larva goes into a type of stasis and can survive harsh conditions. Since the decision to enter the dauer stage is dependent on environmental cues, it represents a classic and well studied example of polyphenism The Dauer state is given other names in the various types of nematodes such as ‘diapause’, ‘hypobiosis’, or ‘spring rise’, but since the C. elegans 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 parasitic nematode larvae.

Under environmental conditions that are favorable for reproduction, C. elegans larvae develop through four stages or molts 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.

Dauer larvae are extensively studied by biologists because of their ability to survive harsh environments and live for extended periods of time. For example, C. elegans 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 daf-2 and daf-23. Dauer formation in C. elegans requires a nuclear receptor DAF-12 and a forkhead transcription factor daf-16. In favorable environments, DAF-12 is activated by a steroid hormone, called Dafachronic Acid, produced by the , DAF-9. DAF-9 and DAF-12 have been implicated by Cynthia Kenyon and colleagues as being required for extended longevity seen in animals that lack germlines. Kenyon showed that, although the daf-16 gene is required for life extension in C. elegans, 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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