Diapause

Diapause, when referencing animal dormancy, is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions. Diapause is a mechanism used as a means to survive predictable, unfavorable environmental conditions, such as temperature extremes, drought, or reduced food availability. Diapause is most often observed in all the life stages of arthropods, especially insects. Embryonic diapause, a somewhat similar phenomenon, occurs in over 130 species of mammals, possibly even in humans, and in the embryos of many of the oviparous species of fish in the order Cyprinodontiformes. Diapause, when referencing animal dormancy, is the delay in development in response to regularly and recurring periods of adverse environmental conditions. It is considered to be a physiological state of dormancy with very specific initiating and inhibiting conditions. Diapause is a mechanism used as a means to survive predictable, unfavorable environmental conditions, such as temperature extremes, drought, or reduced food availability. Diapause is most often observed in all the life stages of arthropods, especially insects. Embryonic diapause, a somewhat similar phenomenon, occurs in over 130 species of mammals, possibly even in humans, and in the embryos of many of the oviparous species of fish in the order Cyprinodontiformes. Activity levels of diapausing stages can vary considerably among species. Diapause may occur in a completely immobile stage, such as the pupae and eggs, or it may occur in very active stages that undergo extensive migrations, such as the adult monarch butterfly, Danaus plexippus. In cases where the insect remains active, feeding is reduced and reproductive development is slowed or halted. Diapause in insects is a dynamic process consisting of several distinct phases. While diapause varies considerably from one taxon of insects to another, these phases can be characterized by particular sets of metabolic processes and responsiveness of the insect to certain environmental stimuli. Diapause can occur during any stage of development in arthropods, but each species exhibits diapause in specific phases of development. Reduced oxygen consumption is typical as is reduced movement and feeding. In Polistes exclamans only the queen is said to be able to undergo diapause. The sensitive stage is the period when stimulus must occur to trigger diapause in the organism. Examples of sensitive stage/diapause periods in various insects: The induction phase occurs at a genetically predetermined stage of life, and occurs well in advance of the environmental stress. This sensitive stage may occur within the lifetime of the diapausing individual, or in preceding generations, particularly in egg diapause. During this phase, insects are responsive to external cues called token stimuli, which trigger the switch from direct development pathways to diapause pathways. Token stimuli can consist of changes in photoperiod, thermoperiod, or allelochemicals from food plants. These stimuli are not in themselves favourable or unfavourable to development, but they herald an impending change in environmental conditions. The preparation phase usually follows the induction phase, though insects may go directly from induction to initiation without a preparation phase. During this phase, insects accumulate and store molecules such as lipids, proteins, and carbohydrates. These molecules are used to maintain the insect throughout diapause and to provide fuel for development following diapause termination. Composition of the cuticle may be altered by changing hydrocarbon composition and by adding lipids to reduce water loss, making the organism resistant to desiccation.Diapausing puparia of the flesh fly, Sarcophaga crassipalpis, increase the amount of cuticular hydrocarbons lining the puparium, effectively reducing the ability of water to cross the cuticle. Photoperiod is the most important stimulus initiating diapause. The initiation phase begins when morphological development ceases. In some cases, this change may be very distinct and can involve moulting into a specific diapause stage, or be accompanied by color change. Enzymatic changes may take place in preparation for cold hardening. For example, only diapausing adults of the fire bug, Pyrrhocoris apterus, have the enzymatic complement that allows them to accumulate polyhydric alcohols, molecules that help to lower their freezing points and thus avoid freezing. Insects may also undergo behavioural changes and begin to aggregate, migrate, or search for suitable overwintering sites. During the maintenance phase, insects experience lowered metabolism and developmental arrest is maintained. Sensitivity to certain stimuli which act to prevent termination of diapause, such as photoperiod and temperature, is increased. At this stage, insects are unresponsive to changes in the environment that will eventually trigger the end of diapause, but they grow more sensitive to these stimuli as time progresses. In insects that undergo obligate diapause, termination may occur spontaneously, without any external stimuli. In facultative diapausers, token stimuli must occur to terminate diapause. These stimuli may include chilling, freezing, or contact with water, depending on the environmental conditions being avoided. These stimuli are important in preventing the insect from terminating diapause too soon, for instance in response to warm weather in late fall. In the Edith's checkerspot butterfly, individuals must receive enough sunlight in order to terminate the diapause stage and become a fully grown butterfly. Termination may occur at the height of unfavourable conditions, such as in the middle of winter. Over time, depth of diapause slowly decreases until direct development can resume, if conditions are favourable.