David L. Denlinger

Ohio State University

Diapause is a form of developmental arrest in insects that is much like hibernation in higher animals. It enables insects and related arthropods to circumvent adverse seasons. Winter is most commonly avoided in temperate zones, but diapause is also used to avoid hot, dry summers and periods of food shortage in the tropics. Unlike quiescence, which represents a halt in development elicited immediately at any stage by an adverse condition, diapause is a developmental response that is expressed only during a specific developmental stage, which depends on the species of insect. For example, the commercial silkworm (Bombyx mori) always diapauses as an early embryo, the European corn borer (Ostrinia nubilalis) as a fifth instar, the cecropia moth (Hyalophora cecropia) as a pupa, and the Colorado potato beetle (Leptinotarsa decemlineata) as an adult. A few species are capable of entering diapause several times, but this usually occurs only in species living at high latitudes for which several years may be required for the completion of development. If the diapause occurs in response to environmental cues it is referred to as "facultative diapause," but if it occurs during each generation regardless of the environmental cues it receives, it is considered an "obligatory diapause." Facultative diapause is by far the more common, but several important species such as the gypsy moth (Lymantria dispar) have an obligatory diapause.

Embryonic diapauses are common in many of the Lepidoptera, in many Hemiptera, and in some Diptera such as mosquitoes. The arrest can occur at any stage of embryonic development, from shortly after fertilization (e.g., commercial silkworm) until after the first instar has already been fully formed (e.g., gypsy moth). Larval diapauses, especially common in the Lepidoptera, are most frequent in the final instar but they sometimes occur in earlier instars as well, e.g., the second instar of the spruce budworm (Choristoneura fumi-ferana). Pupal diapause is well known for the Lepidoptera and Diptera. Usually the arrest of pupal development occurs in the true pupal stage, but there are a few examples of diapause occurring in pharate adults (after completion of adult differentiation but before adult eclosion). Adult diapause is common in the Coleoptera, Hemiptera and Homoptera, Hymenoptera, Orthoptera, and Neuroptera, as well as some Diptera and Lepidoptera. Adult diapause, sometime referred to as a reproductive diapause, represents a halt in reproduction. Ovaries of females remain small, and the oocytes within the ovarioles contain little or no yolk. In males of some species, the testes remain small during diapause, but in others the testes are well developed and contain sperm. Male accessory glands, the organs that produce spermatophores and factors responsible for sperm activation, usually remain small and inactive during diapause. Mating behavior is strongly suppressed during diapause. In wasps, mating takes place in the autumn; males die soon thereafter and only the females overwinter in diapause. In many other insects, both sexes overwinter and mating takes place in the spring, after diapause has been terminated. Some species, such as lacewings and weevils, mate both before and after diapause.

In preparation for diapause, the insect usually sequesters additional energy reserves and moves to a site that is somewhat protected from the full onslaught of the inclement environmental conditions. Such sites may be underground, beneath debris on the soil, within galls and other plant tissues, or inside cocoons or other structures constructed by the insect. A migratory flight may be a preparatory step for diapause. This may include a short flight to a fence row or a local wooded area, but in the extreme it may be a long-distance flight, as made by the monarch butterfly (Danaus plexippus) when it leaves its summer habitat in Canada and the northern regions of the United States and flies to the highlands of Mexico or California to spend the winter in an adult diapause.

Upon entering diapause, development (or reproduction if it is an adult diapause) is halted and metabolic activity is suppressed. Usually, feeding ceases during diapause; thus, the insect is forced to survive on the energy reserves it has garnered prior to the onset of diapause. It is not unusual for an insect destined for diapause to sequester twice as much lipid reserves as its counterpart that is not programmed to enter diapause. The economic utilization of these reserves is enhanced by the suppression of metabolism, and for poikilotherms such as insects, the low temperatures prevailing during winter further serve to conserve energy reserves. Another challenge faced by diapausing insects is the lack of access to free water. Although some insects may drink during diapause, certain stages such as embryos and pupae do not have this option. This lack of water poses special constraints for an organism as small as an insect. Their large surface-to-volume ratios make insects particularly vulnerable to water loss across the surface of their integument. Two features appear to be common adaptations for maintaining water balance during the long months of diapause. The cuticles of many diapausing insects are coated with extra thick layers of wax that are effective in retarding water loss. In addition, a number of diapausing insects are capable of absorbing atmospheric water vapor directly through their cuticle using a mechanism that is not yet clearly understood.

Color changes are sometimes noted for diapausing individuals. For example, diapausing larvae of the southwestern corn borer, Diatraea grandiosella, are white, whereas their nondiapausing counterparts are brown. Reproductively active adults of a lacewing, Chrysopa carnea, are green but turn brown when they enter diapause in the autumn. In the spring, when the lacewings become reproductively active, they again turn green. Such changes presumably serve to camouflage the insect and help it blend with the dominant colors of the seasonal environment.

Flight muscles in many beetles and bugs degenerate when the adults enter diapause. Flight muscles are particularly expensive to maintain, thus their degeneration presumably saves energy that would otherwise be expended for maintenance of this tissue.

Several species that diapause as adults, especially beetles, bugs, and butterflies, are found in aggregations. For species that are distasteful, aggregations are likely to provide protection from predators. Such aggregations, however, may also provide another important function by providing a more stable microenvironment. In diapausing aggregations of a tropical fungus beetle, Stenotarsus rotundus, the beetle's metabolic rate is inversely related to group size and relative humidity. By forming an aggregation the beetles create a stable, high humidity in their environment, a feature that serves to reduce metabolic rate.

Being in diapause does not, by itself, ensure winter survival. The small size of insects implies that they quickly assume a body temperature close to that of the environment, and their body water is thus vulnerable to freezing. Diapausing insects that live in temperate and polar regions have a host of behavioral, physiological, and biochemical adaptations that enable them to survive at low temperature. A few insects such as the goldenrod gall fly, Eurosta solidaginis, are freeze tolerant, which implies that they can actually survive body freezing. But, the majority of insects cannot tolerate body freezing. Such freeze-intolerant or freeze-avoiding insects prevent body freezing by several mechanisms. For example, selection of a thermally buffered microhabitat is a first line of defense. Ice nucleators such as food particles or microbes are usually eliminated from the digestive tract to reduce sites for ice formation. Glycerol, sorbitol, or other polyols serve as classic antifreezes that are synthesized and released into the body to suppress the supercooling point. Several proteins, including thermal hysteresis proteins, ice nucleator proteins, and heat-shock proteins, also contribute to cold hardiness. In some insects, such as flesh flies (Sarcophaga), cold hardiness is directly linked to diapause, indicating that the same genetic program that dictates diapause also results in cold hardiness. In other insects, for example the European corn borer, the two programs are regulated independently: the European corn borer enters diapause without initially being cold hardy, but it becomes cold hardy later in the season in response to prevailing low temperatures.

Diapause thus represents a syndrome of developmental, physiological, biochemical, and behavioral attributes that together serve to enhance survival during seasons of environmental adversity.

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