Insect photoperiodism: measuring the night.

In studies of photoperiodic induction of over-wintering diapause, independent variation of the light (L) and dark (D) components of the daily (LD) cycle shows, with few exceptions, that the duration of the night (D) is more important than that of the day (L). Extensions of D to give cycle lengths up to 3 days or more in so-called Nanda-Hamner (NH) experiments suggest that night length is measured repeatedly in the extended night, with peaks of high diapause incidence occurring at intervals close to 24 h. This indicates a circadian involvement in night length measurement. The circadian oscillation revealed in NH experiments is shown to take its principal time cue from the beginning of the night - at a phase close to Circadian time, CT 12 - in series of such experiments with increasing light (L) components, in a manner comparable to other circadian oscillations such as that controlling the adult eclosion rhythm. It is considered that the photoperiodic circadian oscillation is causally involved in the discrimination between short (summer) and long (autumnal) nights, although further 'downstream' actions of the circadian system on the outcome of time measurement are also likely. Therefore Bünning's original hypothesis - or development of it - is considered to offer the most likely explanation for the photoperiodic mechanism.

Deciphering time measurement: the role of circadian 'clock' genes and formal experimentation in insect photoperiodism.

This review examines possible role(s) of circadian 'clock' genes in insect photoperiodism against a background of many decades of formal experimentation and model building. Since ovarian diapause in the genetic model organism Drosophila melanogaster has proved to be weak and variable, recent attention has been directed to species with more robust photoperiodic responses. However, no obvious consensus on the problem of time measurement in insect photoperiodism has yet to emerge and a variety of mechanisms are indicated. In some species, expression patterns of clock genes and formal experiments based on the canonical properties of the circadian system have suggested that a damped oscillator version of Pittendrigh's external coincidence model is appropriate to explain the measurement of seasonal changes in night length. In other species extreme dampening of constituent oscillators may give rise to apparently hourglass-like photoperiodic responses, and in still others there is evidence for dual oscillator (dawn and dusk) photoperiodic mechanisms of the internal coincidence type. Although the exact role of circadian rhythmicity and of clock genes in photoperiodism is yet to be settled, Bünning's general hypothesis (Bünning, 1936) remains the most persuasive unifying principle. Observed differences between photoperiodic clocks may be reflections of underlying differences in the clock genes in their circadian feedback loops.

Controversial aspects of photoperiodism in insects and mites.

This review examines several controversial aspects of photoperiodism in insects and mites including the role of the circadian system in night length measurement, the nature of apparent hourglass-like responses, and whether or not the circadian component in photoperiodism is the same as that in overt behavioural rhythms. These aspects of the phenomenon are discussed in terms of the entrainment of circadian oscillations by cycles of light and temperature. There is considerable variety of photoperiodic response within the insects (and other arthropods) to show, inter alia, circannual rhythms, internal and external coincidence night length timers, and in some species, non-circadian hourglass-like devices. Many apparent hourglass-like responses, however, could be circadian 'clocks' of the external coincidence type involving oscillations that dampen below threshold in extended periods of darkness. The review also concludes that there is little evidence in favour of the "Hourglass clock-oscillator counter" model proposed for the mite Tetranychus urticae by Vaz Nunes and Veerman (1982a). The responses of this species to complex light and temperature cycles may also be interpreted in terms of a damped oscillator version of external coincidence.

Erwin Bünning and Tony Lees, two giants of chronobiology, and the problem of time measurement in insect photoperiodism.

This paper examines the views of Erwin Bünning and Tony Lees on the mechanism of photoperiodic time measurement, the former advocating a circadian basis for the phenomenon and the latter a non-circadian hourglass-like timer. This difference in opinion led to a protracted split among workers on photoperiodism, some supporting an oscillatory clock and others an "hourglass", and gave rise to the often stated opinion that the two forms of time measurement were mutually exclusive. This paper, however, suggests that both oscillatory and hourglass-like properties are to be seen in insect photoperiodism. Furthermore, the differences between the two apparently conflicting models may be resolved if, following Bünning, "hourglasses" are regarded as damping circadian oscillators, with the more self-sustained (clearly oscillatory) and more highly damped (hourglass-like) responses being parts of a continuous series. Since circadian rhythmicity is an all-pervading and fundamental aspect of insect biology, currently opening up to genetic and molecular analysis, recognition of the basic similarity of a wide range of insect photoperiodic timers may help to unravel the biochemical nature of the mechanism(s) involved.

Selection for high diapause incidence in blow flies (Calliphora vicina) maintained under long days increases the maternal critical daylength: some consequences for the photoperiodic clock.

Using a population of Calliphora vicina from southern Scotland (55 degrees N), showing a critical day length for maternal induction of diapause of about 14.5 h per day, strains of flies were selected for a high incidence of larval diapause under long day length (LD 16:8 h). Diapause incidence was raised from under 10% to almost 100% within five or six generations. The selected flies showed an increase in their critical day length to over 16 h per day, and a high incidence of larval diapause under very long photophases. Selected flies, however, showed mean circadian periods for locomotor activity little different from the original stock, or from flies selected for high diapause under LD 12:12 h, and a Nanda-Hamner profile lacking peaks and troughs of diapause incidence at about 24 h intervals. These results are interpreted to show that the photoperiodic regulation of diapause and the control of overt behavioural rhythmicity are 'separate' physiological systems.

Quality of life among urban Canadian HIV/AIDS clinic outpatients.

Perceived health-related quality of life (QOL) of 113 adult people living with HIV/AIDS (PHA) attending an outpatient hospital clinic was assessed upon initial registration using the Medical Outcomes Study SF-36. QOL ratings on both physical functioning and psychological well-being aspects of the SF-36 were lower than for the general population, especially among PHA in the sample who were symptomatic or for whom AIDS had developed. Although not compromised in terms of physical functioning, asymptomatic PHA had well-being QOL scores comparable to other chronic medical conditions. In general, demographic, social, and many of the health and immune status variables examined were not significantly related to QOL. Number of symptoms was negatively associated with both physical status and emotional well-being dimensions of QOL. PHA in the asymptomatic stage of HIV infection reported better QOL than symptomatic/AIDS patients on few QOL dimensions. Mean QOL ratings did not significantly differ between AIDS patients and those with symptomatic (non-AIDS) infection. These findings suggest that although becoming symptomatic has a significant impact on a majority of dimensions of QOL, being diagnosed with HIV affects well-being dimensions the most. An emphasis on support for both adjusting to diagnosis and symptom management is as important as viral suppression in maximizing QOL for PHA.

Fate of fumonisins during the production of fried tortilla chips.

The fate of fumonisin B(1) (FB(1)), a mycotoxin found in corn, during the commercial manufacture of fried tortilla chips was studied. FB(1) and hydrolyzed FB(1) (HFB(1)) concentrations in four lots of corn and in the masa, other intermediates, liquid and waste byproducts, and fried chips were determined by HPLC. FB(1) concentrations in the masa and chips were reduced significantly, up to 80% in the fried chips, compared to that in the raw corn. HFB(1) was also found in the masa and chips, but at low concentrations compared to FB(1). LC-MS analyses corroborated HPLC findings and further showed the presence of partially hydrolyzed FB(1) (PHFB(1)), which, like HFB(1), was formed during the nixtamalization (cooking/steeping the corn in alkaline water to make masa) step and found predominantly in the cooking/steeping liquid and solid waste. No significant amounts of N-(carboxymethyl)-FB(1) or N-(1-deoxy-D-fructos-1-yl)-FB(1), indicative of fumonisin-sugar adduct formation, were found. Thus, FB(1) is removed from corn and diverted into liquid and waste byproducts during the commercial production of fried tortilla chips. Nixtamalization and rinsing are the critical steps, whereas grinding, sheeting, baking, and frying the masa had little effect.

Help-seeking patterns in HIV/AIDS outpatients.

This study investigates factors associated with requests for professional services, made by newly registered HIV/AIDS Clinic outpatients, referred by general practice physicians to a large urban hospital serving a broad metropolitan/rural area in Canada. Professional service requests were divided into two primary help-seeking categories: practical support (assistance with financial aid, housing, medication costs, etc.) and emotional support (i.e., psychological counselling). Slightly more than half of the sample (51%) of people living with HIV/AIDS (PHA) requested professional services. Service requesters are indistinguishable from non-requesters on a large number of variables, including age, gender, family awareness of HIV/AIDS diagnosis, time elapsed since diagnosis, employment status, spousal status, overall support network size, physical health indicators (Karnofsky Performance Status, CD4 count, symptoms, opportunistic infections) and receipt of community support services elsewhere. Similarly, requesters and non-requesters report comparable levels of overall perceived social support and quality of life. However, requesters of practical support services report significantly fewer friends, lower emotional-informational social support and poorer quality of life due to body pain than non-requesters. Requesters of emotional support services report experiencing significantly lower positive social interaction compared to non-requesters. Implications for the provision of practical and emotional support services for PHA are discussed.

Control of fumonisin: effects of processing.

Of about 10 billion bushels of corn that are grown each year in the United States, less than 2% is processed directly into food products, and about 18% is processed into intermediates such as high-fructose corn syrup, ethanol, and cornstarch. The vast majority of the annual crop is used domestically for animal feed (60%), and about 16% is exported. Thus, any program for controlling residues of fumonisin (FB) in food must recognize that most of the crop is grown for something other than food. Studies on the effects of wet milling on FB residues found these residues nondetectable in cornstarch, the starting material for high-fructose corn syrup and most other wet-milled food ingredients. Similar effects are noted for the dry-milling process. FB residues were nondetectable or quite low in dry flaking grits and corn flour, higher in corn germ, and highest in corn bran. Extrusion of dry-milled products reduces FB concentrations by 30-90% for mixing-type extruders and 20-50% for nonmixing extruders. Cooking and canning generally have little effect on FB content. In the masa process measurable FB is reduced following the cooking, soaking, and washing steps, with little conversion of FB to the hydrolyzed form. Sheeting, baking, and frying at commercial times and temperatures generally have no effect. In summary, all available studies on the effects of processing corn into food and food ingredients consistently demonstrate substantial reductions in measurable FB. No studies have shown a concentration in FB residues in food products or ingredients.

Perceived support in newly registered HIV/AIDS clinic outpatients.

This study compares levels of perceived social support in newly registered outpatients attending an HIV/AIDS clinic to those reported in a reference sample of other ambulatory medical populations. Perceived availability of four types of functional support is assessed: tangible, affectionate, positive social interaction and emotional-informational. Other elements of patients' support networks that might be associated with perceived support are also examined. In general, functional support levels in the outpatient PHA population were found to be similar to those of the MOS-SSS reference sample. Patient factors including age, gender, family awareness of HIV/AIDS diagnosis, time elapsed since diagnosis, employment status, physical health indicators, antiretroviral treatment history and support network size showed no relationship to functional support. Network composition factors significantly associated with higher social support ratings included the presence of a partner (all dimensions of functional support) and a greater proportion of close friends as opposed to close family members (emotional-informational support). Requesters of clinic support services tended to exhibit lower emotional-informational support scores compared to non-requesters. Implications for the provision of support services to people living with HIV/AIDS are discussed.

Larval diapause duration and fat metabolism in three geographical strains of the blow fly, Calliphora vicina.

Diapausing larvae of the blow fly, Calliphora vicina, from three geographical strains exposed, as adults, to short days, were maintained under identical conditions (darkness, 11-12 degrees C) and examined for changes in wet weight, dry weight, water and fat content during diapause development to the emergence of post-diapause adults. Larvae produced by flies originating from northern Finland (Nallikari, 65 degrees N) showed a longer, more intense, diapause than those from localities further south (Edinburgh, Scotland, 55 degrees N and Barga, Italy, 44 degrees N), but all three strains showed similar rates of loss of the parameters measured. This was also the case for post-diapause adults, flies of the Barga strain with its relatively short diapause emerging with greater residual fat reserves than flies from the Edinburgh or Nallikari strains with their more protracted diapause. It was concluded that the rates of water and fat loss were functions of the conditions used for diapause larval maintenance (probably temperature) rather than the maternally programmed degree of diapause incidence, or of its 'depth' or 'intensity'.

Effects of temperature and temperature-steps on circadian locomotor rhythmicity in the blow fly Calliphora vicina.

The free-running period (in darkness) of the locomotor activity rhythm in adult blow flies (Calliphora vicina) was temperature-compensated between 15 and 25 degrees C, showing Q(10) values between 0.98 and 1.04. Single steps-up (20 to 25 degrees C) or steps-down (20 to 15 degrees C) in temperature caused stable phase shifts of the activity rhythm, giving rise to temperature-step phase response curves (PRCs) with both advances and delays. Phase advances, however, were dominant for steps-up, and phase delays for steps-down; the two PRCs were almost "mirror images" of each other. Following protocols introduced by Zimmerman et al. [(1968) Temperature compensation of the circadian oscillation in Drosophila pseudoobscura and its entrainment by temperature cycles, Journal of Insect Physiology, 14, 669-684] for the rhythm of pupal eclosion in Drosophila pseudoobscura, the steps-up and steps-down PRCs for C. vicina were used to compute a theoretical PRC for a 6 h low temperature pulse, and from this a theoretical steady-state phase relationship of the locomotor activity rhythm to a train of such pulses making up a temperature cycle (18 h at 20 degrees and 6 h at 15 degrees C).

Geographical and diapause-related cold tolerance in the blow fly, Calliphora vicina.

Three geographical strains of the blow fly, Calliphora vicina, were tested for cold tolerance at 0 degrees, -4 degrees and -8 degrees C. Survival to eclosion after 1 to 18 days of cold exposure was greater for diapause-destined larvae than for nondiapause-destined larvae of the two northern strains (Nallikari, Finland 65 degrees N and Edinburgh, Scotland 55 degrees N) but not for the southernmost strain (Barga, Italy 44 degrees N) where no clear differences were apparent. Diapause-destined larvae of the Edinburgh strain were more cold tolerant than those from Nallikari, at both -4 degrees and -8 degrees C, a difference possibly attributable to the long-lasting snow cover in the more northern locality, which might insulate the overwintering soil microclimate. At 0 degrees C, however, Nallikari larvae were more cold tolerant than Edinburgh or Barga. This was also the case for nondiapause-destined larvae, indicating that cold tolerance may occur, in part, independently of the diapause programme. In all three strains diapausing larvae were more cold tolerant than same-age (nondiapausing) pupae. For Nallikari, but not Barga, wandering larvae from short-day exposed flies, therefore initially programmed for diapause, but diverted from the diapause pathway by larval breeding at 19 degrees C, were significantly more cold tolerant than nondiapause larvae from long-day parents, indicating some maternal regulation of larval cold tolerance. There was, however, no evidence for an additional cold hardiness in larvae acclimatised to cold by a gradual reduction of temperature.

Inheritance of the photoperiodic response controlling larval diapause in the blow fly, Calliphora vicina.

Larvae of the blow fly Calliphora vicina R-D. (Diptera: Calliphoridae) display a diapause in response to the exposure of their parents to short photoperiods. Due to geographic variation in photoperiodic response, flies from a southern, English population show a long-day response to the fixed photoperiod of L:D 15.5:8.5 whilst flies from a northern population from Finland show a short-day response to the same photoperiod. Crosses between these strains have shown previously that diapause incidence is a maternal characteristic; here we demonstrate that the hybrid female offspring of such crosses are not intermediate between the two parental strains but show a photoperiodic response biased towards their maternal line. Thus not only are males unable to influence directly the diapause incidence among their offspring but the indirect effects of inheritance down the male line are weaker than down the female. Diapause duration, in contrast, is influenced by each parent in a similar manner. Diapause lasts longer in larvae with a greater admixture of northern genes regardless of whether they were maternal or paternal.

Insect circadian rhythms and photoperiodism.

Two clock-controlled processes, overt circadian rhythmicity and the photoperiodic induction of diapause, are described in the blow fly, Calliphora vicina and the fruit fly, Drosophila melanogaster. Circadian locomotor rhythms of the adult flies reflect endogenous, self-sustained oscillations with a temperature compensated period. The free-running rhythms become synchronised (entrained) to daily light:dark cycles, but become arrhythmic in constant light above a certain intensity. Some flies show fragmented rhythms (internal desynchronisation) suggesting that overt rhythmicity is the product of a multioscillator (multicellular) system. Photoperiodic induction of larval diapause in C. vicina and of ovarian diapause in D. melanogaster is also based on the circadian system but seems to involve a separate mechanism at both the molecular and neuronal levels. For both processes in both species, the compound eyes and ocelli are neither essential nor necessary for photic entrainment, and the circadian clock mechanism is not within the optic lobes. The central brain is the most likely site for both rhythm generation and extra-optic photoreception. In D. melanogaster, a group of lateral brain neurons has been identified as important circadian pacemaker cells, which are possibly also photo-sensitive. Similar lateral brain neurons, staining for arrestin, a protein in the phototransduction 'cascade' and a selective marker for photoreceptors in both vertebrates and invertebrates, have been identified in C. vicina. Much less is known about the cellular substrate of the photoperiodic mechanism, but this may involve the pars intercerebralis region of the mid-brain.

S-antigen antibody partially blocks entrainment and the effects of constant light on the circadian rhythm of locomotor activity in the adult blow fly, Calliphora vicina.

Injection of S-antigen (arrestin) antibody into the brain of the blow fly, Calliphora vicina, appeared to reduce sensitivity to the photic effects of a light:dark cycle (LD; entrainment) or continuous "bright" light (LL; arrhythmicity). In LD, a proportion of the injected flies evaded entrainment or showed delayed entrainment. In bright LL, flies continued with a free-running rhythm, which remained unchanged (as in continuous darkness) or lengthened (as in "dim" LL). These results focus attention on four groups of arrestin-positive neurons in the fly's brain as potential components of the photoreceptive system.

Evaluation of the carcinogenic potential of pesticides. 4. Chloroalkylthiodicarboximide compounds with fungicidal activity.

The Health Effects Division of the Office of Pesticide Programs (OPP) assessed the carcinogenic potential of three structurally related chloroalkylthiodicarboximide fungicides using a consensus peer review process and EPA's 1986 guidelines for cancer risk assessment. All of the fungicides were categorized as Group B2 (probable human) carcinogens based upon findings of an increased incidence of malignant tumors, or combined malignant and benign tumors, in multiple experiments involving different strains of mice and rats. The primary sites of tumor formation with the chloroalkylthiodicarboximide fungicides in male and/or female mice (CD-1 and B6C3F1) were the gastrointestinal tract (captan, folpet, and captafol), the lymph system (folpet and captafol), and the vascular system (captafol). The main sites of tumor formation in rats of one or both sexes (CR CD, Wistar, or F344 strains) were the kidney (Captan and captafol), uterus (captan), mammary gland and liver (captafol). In addition, positive trends for thyroid, testicular, mammary gland, and lymph node tumors were observed with folpet in the same strains of rats. All three of the compounds exhibited positive mutagenic activity in a variety of in vitro short-term tests for gene mutation, DNA repair, and chromosomal aberrations in prokaryotic and eukaryotic cells, but were not genotoxic in available studies performed under in vivo conditions. The assessment of human cancer risk for captan, folpet, and captafol was made using low-dose extrapolation models.