Components of daily and seasonal timing systems in insects are reviewed. Photoperiod indicates seasonal position reliably, but signals can be much modified by habitat, latitude and season. Several receptor features and pigment systems are known, with different daily, seasonal and general functions, including differences between circadian and seasonal reception. Clocks can serve several different purposes, functioning as daily oscillators, interval timers or through successive requirements. The molecular functioning of circadian clocks is best known, but even so there is considerable complexity and diversity and much remains to be discovered. We know relatively little about the internal states that provide information for timed responses (such as the photoperiodic "counter"), about the central controlling mechanism, or about the effectors that transmit output signals. Nevertheless, temporal responses serve a very great range of purposes in insects, and the reported complexity in all of the components of timing systems reflects complex ecological needs across daily and seasonal intervals. The variety of components and the complexity of interactions reported (even within species), as well as the diversity of such elements as photosensitive pigments, molecular clock function and potential neurotransmitters, suggests that - unlike some earlier expectations - there is no single master clock for all timing functions in insects.
Insect photoperiodism and rhythmicity have been studied by both observational or direct approaches (examination of system elements or devices, and qualities such as survival), and by inferential or indirect approaches (such as interpretation of various responses to photoperiod, modelling, and estimating fitness). Many students work with only one approach, but the power of different approaches is not equal, and knowledge at one level may not give answers at another. These difficulties tend to limit our understanding of the linkages among components.
This overview suggests several lessons for the study of photoperiodism and rhythmicity. There are multiple elements, complex integration and a diversity of clocks, showing that different processes serve different purposes. The diversity of findings also results from the fact that different investigative approaches, which depend on the question being asked and on the perspective of the investigator, can influence the outcome of the investigation. Given these complexities, I believe that the key to interpreting photoperiodic and circadian responses is their ecological value. Notwithstanding the interest of timing mechanisms or their parts and of specific responses, daily rhythms and seasonal timing are best understood through the essential context provided by the ecological demands on the actual organisms under study.
Insect dormancy responses, in the broad sense of modifications of development, are examined from a general perspective. The range of responses is extraordinarily wide because environments are diverse, different taxa have different evolutionary histories, adaptations are needed for both seasonal timing and resistance to adversity, and not only development but also many other aspects of the life-cycle must be coordinated. Developmental options are illustrated by examining the wide range of ways in which development can be modified, the fact that each individual response consists of several components, and the different potential durations of the life-cycle. The concepts of alternative life-cycle pathways (chosen according to current and likely future environmental conditions) and of active and passive default responses are treated. Also introduced are aspects of variation and trade-offs.
Some general conclusions that help in understanding dormancy responses emerge from such an examination. Many options are available (cf. Table 1). The nature of the habitat, especially its predictability, determines the potential effectiveness of many of the developmental options. Any particular set of responses reflects evolutionary history and hence depends on past as well as current environments. It is not necessarily obvious what kinds of selection, especially requirements for timing versus resistance to adversity, explain a particular life cycle. Life-cycle pathways have multiple components, so that components cannot be analyzed in isolation. A given feature, such as delayed development, can have multiple roles. Default responses can be either active (development continues unless signalled otherwise) or passive (development stops unless signalled otherwise), making necessary a broad approach to understanding the action of environmental cues. Even relatively minor effects that fine-tune dormancy responses enhance survival, but may be difficult to detect or measure. Trade-offs are not inevitable, not only when certain resources are surplus, but also because resources in very short supply (constraints) cannot be traded off. Life-cycle components are widely, but not universally, coordinated. These conclusions confirm that the range of dormancy responses is wider, more complex and more integrated than has often been recognized.