A comparison of the responses of tropical and temperate flies (Diptera: Sarcophagidae) to cold and heat stress

Summary Two flesh fly species from the tropical lowlands (Peckia abnormis and Sarcodexia sternodontis) were more susceptible to both cold-shock and heatshock injury than temperate flies (Sarcophaga crassipalpis and S. bullata) and a fly from a tropical high altitude (Blaesoxipha plinthopyga). A brief (2-h) exposure to 0°C elicits a protective response against subsequent cold injury at–10°C in the temperate flies and in B. plinthopyga but no such response was found in the flies from the tropical lowlands. However, both tropical and temperate flies could be protected against heat injury (45°C) by first exposing them to a mild heat shock (2 h at 40°C). The supercooling point is not a good indicator of cold tolerance: supercooling points of pupae were similar in all species, ranging from–18.9 to–23.0°C, and no differences were found between the tropical and temperate species. Among the temperate species, glycerol, the major cryoprotectant, can be elevated by short-term exposure to 0°C, but glycerol could not be detected in the tropical flies. Low-temperature (0°C) exposure also increased hemolymph osmolality of the temperate species, but no such increase was observed in the tropical lowland species. Adaptations to temperature stress thus differ in tropical and temperate flesh flies: while flies from both geographic areas share a mechanism for rapidly increasing heat tolerance, only the temperate flies appear capable of responding rapidly to cold stress. The presence of a heat shock response in species that lack the ability to rapidly respond to cold stress indicates that the biochemical and physiological bases for these two responses are likely to differ.     

Thermal response and reversibility of diapause in the eggs of Locusta migratoria

Abstract. Eggs of a local population of Locusta migratoria L. (Orthoptera: Acrididae) collected near Hirosaki (40.5^oN) entered diapause when incubated at temperatures between 20 and 35^oC. For diapause development the optimum temperature was 10^oC. The lower thermal threshold for post-diapause development was 14.7^oC. After chilling at 10^oC for 20 days, the rate of hatching varied with incubation temperature, being 0, 61% and 81% at 20, 25 and 30^oC, respectively. After chilling for 40 days or more, however, almost all eggs hatched at 20–30^oC. Diapause with a reduced intensity seemed to be eliminated easily by a high temperature of 25 or 30^oC.
When eggs chilled at 10^oC for 20 days were kept at 20^oC for 7 days or more before incubation at 25^oC, almost all eggs maintained diapause. Most eggs chilled at 10^oC for three 10 day periods separated by 3 days of warming at 25^oC failed to terminate diapause. Daily alternations of 10^oC (18 h) and 25^oC (6h) decreased the diapause-terminating effect of chilling. These facts suggest that diapause intensity can be restored if eggs chilled insufficiently are exposed to a moderately high temperature. This reversible change in diapause intensity would play an important role in maintaining diapause before winter.     

Effects of temperature on virogenesis of bluetongue virus serotype 11 in Culicoides variipennis sonorensis

Abstract. Culicoides variipennis sonorensis females were fed bluetongue virus serotype 11 mixed in sheep blood and were held at constant temperatures of 32, 27, 21 and 15^oC. Virogenesis, as measured by enzyme-linked immunosorbent assay (ELISA), proceeded significantly faster at higher temperatures. Based on ELISA absorbance ≥0.2, some flies first were categorized as infected after 1 day, 2 days and 4 days at 32, 27 and 21^oC, respectively. Peak levels of virus antigen were seen after 5–7, 7–13 and 18–22 days for flies held at 32, 27 and 21^oC, respectively. There was no significant virus replication in flies held at 15^oC for 22 days, but latent virus replicated and was detected easily (44% infection) 4–10 days after these flies were transferred to 27^oC. The implications for temperature effects on bluetongue epizootiology are discussed.     

Stress-induced accumulation of glycerol in the flesh fly, Sarcophaga bullata: evidence indicating anti-desiccant and cryoprotectant functions of this polyol and a role for the brain in coordinating the response

Nondiapausing larvae of the flesh fly, Sarcophaga bullata, responded to several forms of short-term environmental stress (low temperature, anoxia and desiccation) by accumulating glycerol. Elevation of this polyol, regardless of the type of stress that induced accumulation, conferred cold resistance: larvae with high glycerol levels were 3-4 times more tolerant of a 2h exposure to -10 degrees C than unstressed larvae. Protection against low temperature injury, as well as dehydration, was also attained by injection of exogenous glycerol into third instar larvae. This artificially induced cold hardiness was only temporary: when glycerol-injected larvae were exposed to -10 degrees C immediately after injection, survival was high, but none survived if they were injected and then held at 25 degrees C for 2 days before the -10 degrees C exposure. Larvae ligated behind the brain immediately after low temperature exposure failed to accumulate glycerol, but glycerol did accumulate in larvae ligated 6-24h after cold treatment, thus implying a critical role for the brain in initiating glycerol production. Interestingly, a much shorter exposure (2h) to low temperature was sufficient to reduce the maximum rate of water loss. Collectively, these observations suggest that multiple pathways may be exploited in response to stress: one pathway is most likely associated with rapid cold hardening (RCH) which generates immediate protection, and a second pathway remains activated for a longer period to enhance the initial protection afforded by glycerol.     

A mutant strain of Chymomyza costata (Diptera: Drosophilidae) insensitive to diapause-inducing action of photoperiod

ABSTRACT. From a Japanese population of Chymomyza costata which has been known to have a photoperiodic larval diapause, we selected a mutant strain which did not respond to photoperiod. However, about 70% of the individuals of this strain entered diapause at 11^oC irrespective of photoperiod, and about the same percentage of those of the photoperiod-sensitive strain also did so in continuous illumination at 11^oC. This indicates that low temperature induces diapause independently of photoperiod. On the other hand, a temperature drop from 18 or 25^oC to 15^oC and chilling at 4^oC did not induce diapause.     

The activation of resting tsetse flies (Glossina) in response to visual and olfactory stimuli in the field

ABSTRACT Studies were conducted in Zimbabwe of the responses of Glossina morsitans morsitans Westwood and Glossina pallidipes Austen resting in a refuge to various host stimuli. Tsetse took off in response to 100% ox odour, 0.08% carbon dioxide or a visual stimulus consisting of a 0.75 × 0.75 m black target placed c . 5 m from the refuge moving at 4^o s^-1, but the level of response was low with only 35%, 19% and 29% responding, respectively. Tsetse did not take off in response to any one of 25% ox odour, 0.8% carbon dioxide, acetone (3 μg 1^-1) or octenol (0.03 μg 1^-1). In the absence of any host stimuli, flies emerged from the refuge later on hotter days (35–37^oC) than on cooler days (32–34.5^oC). Male G.pallidipes emerging later in the afternoon contained significantly more haematin than those emerging relatively earlier. There were no significant differences between the responses of G.m. morsitans and G.pallidipes. It is suggested that the initial activation of resting flies is primarily mediated through endogenous, rather than host, stimuli.     

Induction of winged sexuparae in root-inhabiting colonies of the aphid Pemphigus betae

Abstract. Growth chamber experiments were conducted to determine what environmental factors induce the development of sexuparae in root-feeding colonies of the aphid Pemphigus betae Doane (Aphididae: Pemphiginae). Experimental colonies were derived from clones that had been reproducing for at least 15 months under constant laboratory conditions. At constant 20–23^oC, sexuparae were almost never produced regardless of the level of crowding or photoperiod. In contrast, every culture of several clones produced sexuparae when subjected to temperatures alternating between 20 and 15^oC during the light and dark phases respectively. This fluctuating temperature regime resulted in sexupara production regardless of photoperiod. All cultures also produced some sexuparae when cultures were kept in constant darkness at temperatures that were either alternated between 20 and 15^oC or kept constant at 15^oC. Constant 17^oC resulted in sexupara production in most cultures. At 17^oC, experimental manipulation of densities within cultures showed that crowding promotes sexupara production. Thus, within the range of conditions occurring at the time of sexupara induction in the field, photoperiod has no effect on sexupara production, temperature below a threshold between 17 and 20–23^oC triggers sexupara production, and crowding promotes sexupara production but only if temperature is low enough.     

Rapid cold hardening in the olive fruit fly Bactrocera oleae under laboratory and field conditions

A rapid cold hardening response was studied in females and males of the olive fruit fly Bactrocera (Dacus) oleae. When laboratory-reared females and males were transferred and maintained from the rearing temperature of 24 °C for 2 h to –6.5 °C approximately 5% survived. However, conditioning of both females and males for 2 h at various temperatures from 0 to 10 °C before their exposure for 2 h to –6.5 °C increased survival to 80 to 92%. A similar rapid cold hardening response in both females and males was also induced through gradual cooling of the flies at a rate of approximately 0.4 °C per min. The rapid increase in cold tolerance after prior conditioning of the flies to low temperatures, was rapidly lost when they returned to a higher temperature of 24 °C. In the field, in late February and early March, females and males were capable of a rapid cold hardening response. After exposure to the critical temperature they suffered a high mortality when tested in the afternoon and low mortality early in the morning on consecutive days, probably because of differences in the prevailing field temperatures a few hours before testing. This plasticity of cold tolerance gained through rapid cold hardening may allow the flies to survive during periods of the year with great fluctuation in circadian temperatures.     

Incubation temperature in the Australasian Gannet Moms senator

Gannets (Sulidae) and some other pelecaniforms incubate their eggs under the webs of their totipalmate feet. These species have a wide latitudinal distribution from tropics to subarctic, but little is known of the incubation temperatures attained. I measured egg temperatures of the Australasian Gannet Morns serrator at Cape Kidnappers, New Zealand, employing a data logger to obtain records every 15 min for day-long sample periods at undisturbed nests. Egg surface temperatures were relatively stable and little affected by ambient temperature. Mean surface temperature of natural eggs was 34.9^oC for samples taken during the first 4 days of incubation, but this then increased and stabilized at 36.5^oC. Internal temperature of pipped eggs was about 1^oC higher, attributable to embryonic heat production. Upper surface temperature of eggs kept in a fixed position was about 2^oC below deep adult body temperature (40.3^oC), suggesting heat is transferred directly from the body through the feet. Clutch size does not appear to be limited by an inability to warm two eggs. These results are in general agreement with measurements from other web incubators and are well within the range for species with conventional brood patch incubation.     

Environmental versus genetic influences on fluctuating asymmetry in the house fly, Musca domestica

The causes of fluctuating asymmetry (FA) are poorly understood, yet it has been widely used as a measure of fitness. Many studies have demonstrated that individuals with low FA are preferred as mates, and it has been argued that this is because FA provides an indicator of genetic quality. However, the relative importance of genes versus environment in determining the level of FA shown by an organism is currently the subject of much controversy. As yet there is no clear consensus as to whether FA generally has a heritable component and if so how large this might be, or indeed if it is sensible to generalize at all. In Musca domestica flies with low wing length FA have beeen found to enjoy higher mating success. In order to interpret this finding we assess whether wing length FA in this species is heritable, and also how it is influenced by environmental stresses induced by temperature and crowding. We also examine whether offspring viability is related to parental FA. We found that wing length FA in M. domestica had no detectable heritable components, and parental FA did not influence offspring viability. FA was influenced by rearing temperature, with flies exhibiting highest FA at the lowest rearing temperature (15^oC). Larval survival rate was greatest, and the resulting adults largest, at the intermediate rearing temperature (25^oC) compared to higher or lower temperatures, suggesting that 25^oC is close to the optimum for the development of M. domestica. Adult size appears to provide a better indicator of stress during development man does wing FA. These results are discussed in relation to the utility of FA as a tool for use in evolutionary studies.     

Thermoregulation in four Australian lizards of the genus Egernia (Sauria: Scincidae)

Mean thermal preferenda for the head and body were found to be similar in Egernia whitei, E. cunninghami, and E. bungana (range in T_H= 33.6^o -33.8^o C; range in T_B= 34.0^o-35.3^oC). However, those for E. majors were significantly lower (T_H= 31.1^o C, T_B=32.9^oC). Significant differences were found between mean head and body preferred temperatures in E. cunninghami and E. major. Higher temperatures were tolerated by the body than the head in all four species. Gaping was not observed to occur prior to cessation of respiration in any of the four species during radiant or solar heating. Based upon thermal preferenda and critical thermal maxima data, thermal safety margins for body temperature were calculated for E. whitei and E. cunninghami Postural orientation to the sun's rays increased and decreased heat gain. The seeking of shade was the most effective method employed by all four species for reducing head and body temperature.     

Effects of temperature on development of Otiorrhynchus sulcatus (Coleoptera: Curculionidae)

Development of the black vine weevil, Otiorrhynchus sulcatus , was investigated at constant temperatures from 9 to 27 ^oC. Pre-adult development was completed in 200 and 130 days at 15 and 24 ^oC, respectively. Eggs and larvae developed at 27 ^oC, but no pupation took place at this temperature. The developmental threshold temperatures for eggs, larvae + pre-pupae and pupae were 6, 2–6 and 12 ^oC, respectively. Reproductive maturation of females took place at temperatures from 12 to 27 ^oC, but only a few females became reproductive at 12 ^oC. Most females at 12 ^oC, like those at 9 ^oC, had a short maturation feed, but the ovaries did not develop. The pre-oviposition period was temperature dependent. The females showed a cyclic oviposition pattern with three peaks. The significance of soil temperatures for the phenology and distribution of O. sulcatus in Norway is discussed.     

Assessment of phosphine fumigation as a plant quarantine measure against Spodoptera littoralis

In laboratory tests first instar larvae of Spodoptera littoralis were killed by phosphine fumigations of 24 and 48 h at a concentration x time product (CTP) of 36 mg h/1. Eggs aged 1–2 days were, however, tolerant to phosphine at a CTP of 144 mg h/1 and this CTP caused severe damage to chrysanthemum cuttings (cv. Freedom). When eggs aged < 1 day and 1–2 days old were preconditioned at 20 ^oC for 72 h before phosphine fumigation, complete mortality occurred at CTPs at and above 36 mg h/1. Preconditioning for 48 h at 20 ^oC and for 48 and 72 h at 15 ^oC did not produce 100% mortality at a CTP of 288 mg h/1. It was suggested that eggs of S. littoralis pass through a phase of tolerance to phosphine which at 20 ^oC lasts about 4 days after oviposition.
There was no indication that phosphine was toxic to chrysanthemum cuttings (cv. Pollyanne) at concentrations of 0–75 mg/1 (CTPs of 18 and 36 mg h/1). Storage at 15 ^oC was not damaging to cuttings but at 20 ^oC some damage became apparent.     

The effect of temperature and inoculation access period on the transmission of barley yellow dwarf virus byRhopalosiphum padi (L.) and Sitobion avenae (F.)

Winged individuals of Rhopalosiphum padi and Sitobion avenae transmitted the PAV-like and MAV-like isolates of barley yellow dwarf virus respectively. Success of transmission after inoculation access periods of 2, 6, 12, 24, 48 and 72 h were examined and survival, reproduction and movement of the aphid vectors were recorded at these times. The experiment was done at four different temperatures: 6^oC, 12^oC, 18^oC and 23^oC. For both isolates the inoculation efficacy did not increase after a 24 h inoculation access period and there was no difference in inoculation efficiency at the three highest temperatures, that at 6^oC being significantly lower than at 12^oC to 23^oC. The results suggest that autumn temperature is a critical factor for BYDV epidemiology in Britain with a small increase in autumn temperature leading to greatly increased infection rates.     

Arabian sandflies (Diptera: Psychodidae) prefer the hottest nights?

Abstract. A vehicle-mounted net was used to collect hourly samples of sandflies on 15 nights during June in northern Oman. Every half hour, the temperature, relative humidity, wind velocity and light intensity were measured (there was no cloud or rainfall during this period).
The sandflies caught were mainly Phlebotomus alexandri and Sergentomyia clydei. Their circadian activity increased rapidly after sunset (18.50 hours). The high level of activity was fairly constant during 9 h of darkness until dawn, when it decreased rapidly. A few flies were still active at 07.00 hours, 1.5 h after sunrise.
A multiple regression showed that the main factor affecting sandfly activity was light intensity. When this factor was removed, by considering only the 135 catches collected during the 9 h of darkness, the second most important factor was low relative humidity, followed by low wind velocity. Temperature was not a significant factor in the analysis, because of its strong negative correlation with humidity. However, when the effect of humidity was removed from the regression, high temperature became significant, but less important than wind.
The regressions showed that, for flight activity, the optimum humidity was around 10%; the probable maximum wind velocity was 3.5 ms^-1 and 11^oC was the probable minimum temperature. Thus, when the 4 nights with highest catches (200–260 flies/night) were compared with the 4 nights with lowest catches (50–120 flies/night), the best nights had a low humidity (10–25%) and low wind speed (<0.3ms-¹) in combination with highest temperatures (31–43^oC).     

Hourglass and oscillator expressions of photoperiodic diapause response in the cabbage moth Mamestra brassicae

Abstract. Both oscillator and hourglass features are found in the photoperiodic response that controls the pupal winter diapause of Mamestra brassicae. The expression of oscillatory response to extended long-night cycles is temperature dependent, i.e. circadian resonance appears at 23 and 25^oC but not at 20 and 28^oC. At 20^oC, scanning of extended scotophases by a short light pulse does not reveal any clear circadian rhythmicity. However, a circadian feature of the photoperiodic response is indicated even at 20^oC by a bistability phenomenon, i.e. either one of the two dark periods in symmetrical skeleton photoperiods determines the diapause response depending on the phase angle with the preceding (entraining) light-dark cycles. At 20 and 25^oC, the incidence of diapause increases as a function of the number of light–dark cycles regardless of the cycle length (T) , if T is 24 h or 2 X 24h (with a 12 h light period). A non-diel cycle (r=36h) is less effective, suggesting that disturbance of the circadian organization partly impairs the diapause-inducing function. The inductive effect of a long night is largely affected by temperature, and becomes saturated with eight cycles at 20^oC and 14 cycles at 25^oC. Presumably, an hourglass mechanism measures the dark time, and a circadian component involved in some later sequence of the photoperiodic response may or may not be expressed depending on the mode of interaction between them.     

Development rates of some pre-adult stages in blowflies with reference to low temperatures

Abstract. Duration of some pre-adult stages of Calliphora vicina Robineau-Desvoidy, C.alpina Zett., C.vomitoria Linn., Phormia terraenovae R.-D. and Lucilia sericata Meig. were determined at 3.5–26^oC, in the context of use of these insects in forensic estimates of post-mortem intervals. Egg development in C. vicina occurs down to at least 3.5^oC, larval growth to 4^oC and puparial formation, development and perfect adult emergence to 5^oC. Growth curves by successive weighing of individual larvae showed specific differences in rates of development of the above species at constant temperatures. Alternating temperatures for 12 h periods (such as 10^oC and 20^oC) to simulate diurnal changes, usually, but not at all temperature pairs tested, caused acceleration of growth relative to the corresponding constant temperature (15^oC in the above case), in C.vomitoria, P.terraenovae and L.sericata , but usually caused retardation of larval growth in C.vicina. In this species, larvae as single individuals on beef liver in small tubes grew on average at the same rates as those (50–100 larvae) growing communally. All species showed variation in larval growth between individuals, more marked in C.vicina , and slower-growing larvae showed increased unexplained mortality. Because of this and to avoid complication by fast-growing larvae from later laid egg batches on a carcass, growth curves given are those of mean weights of the heaviest 25% (upper quartile) of groups of 40–90 feeding larvae. Variation in growth rates, and acceleration or retardation of growth in different species under alternating temperatures, thus affect suggested procedures in obtaining estimates of post-mortem intervals using these insects.     

Influence of thermoperiod and photoperiod on reproductive diapause in Bruchidius atrolineatus (Pic) (Coleoptera, Bruchidae)

Abstract Bruchidius atrolineatus (Pic) is a common pest infesting cowpea seeds ( Vigna unguiculata Walp) in Africa. In the Sahelian zone, the adults are in reproductive diapause during the dry season. Diapause induction depends on the climatic conditions during embryonic and post-embryonic development. A strain of B. atrolineatus originating from the Niamey (13^oN) was reared in different thermoperiodic and photoperiodic conditions. In the thermoperiodic regime 40:25^oC, in continuous darkness, induction of diapause was dependent on the duration of the thermophase. Photoperiod could also influence diapause induction but the response depended on the temperature. When the beetles were reared in conditions of LD 12:12 h, 40:25^oC, for seven generations, the proportion of diapausing adults decreased and their sensitivity to photoperiod increased. In LD 14:10 h, 40:25^CC, the proportion of diapausing adults remained high regardless of the photoperiod. In these conditions, the first emerging adults were sexually active and were used for selection of a strain with a low incidence of diapause. After twelve generations, the proportion of diapausing beetles was lower than 5%. The responses of this selected strain to photo- and thermoperiods were reduced. The sensitivity of B. atrolineatus to photoperiod and thermoperiod could be controlled by genetic systems as observed in other insect species.     

Calling behaviour in M.convecta females under different temperature and photoperiodic conditions

Abstract. . The effects of temperature and photoperiod on calling behaviour in females of the Australian common armyworm, Mythimna corwecta (Walker) (Lepidoptera: Noctuidae), were studied in the laboratory. Age at first calling was greater at 15 and 20^oC compared with 25^oC, but there were no significant differences between 25 and 30^oC. Females kept at 10^oC did not call, but if transferred at 10 days to 20^oC they called after a period similar to the pre-calling period at constant 20^oC. Onset time of calling was earlier at 15 and 20^oC compared with 25 and 30^oC, but there were no significant differences in calling duration. At 20^oC, age at first calling was increased by shorter day length (12h and 14h, compared with 16h), and there was an interaction between daylength and moth age affecting the duration of calling. Onset times advanced with longer daylength, but peak calling was later in relation to the lengtii of the scotophase. These results are discussed in connection with migration in M.convecta. Evidence for a circadian rhythm of calling was found in females entrained for 3 and 8 days after emergence under reverse-cycle conditions and then transferred to constant darkness. However, after 56 h and 80 h respectively of darkness, calling became de-synchronized. Subsequently, it appeared to re-synchronize to a different clock, which approximately correlated with the normal photoperiods the moths had experienced during larval development.     

Consistent differences among individual monarch butterflies (Danaus plexippus L.) in flight activity at low temperatures

Previous work suggested differences in allozyme frequencies between samples of monarch butterflies collected at different times of the day. This study examines the prediction that some individuals are active consistently earlier in the day than others. Four specific hypotheses were tested: (1) Individuals caught early on one day were more likely than those caught late to be caught early on subsequent days. This was true in some experiments. (2) Individuals caught early in the field were more likely than those caught late to fly early in outdoor flight cages. This was true when outdoor temperatures were low (between 10 and 20^oC) but not when they were above 20^oC. (3) Individuals caught early in the field were more likely to be able to fly at low body temperatures than those caught late. This was not true. (4) Individuals that flew early in outdoor flight cages were more likely to be able to fly at low body temperatures. This was not true. Overall we conclude, on the basis of field and flight cage experiments, that some individuals are active consistently earlier in the day than others. The lack of a relationship between field and temperature cabinet results indicates that this is not purely a result of differences in ability to fly at low body temperatures. The implications of these results are discussed.