Environmental and Genetic Effects on the Asymmetry Phenotype: Diazinon Resistance in the Australian Sheep Blowfly, Lucilia Cuprina

The asymmetry phenotype of diazinon-resistant flies lacking a fitness/asymmetry Modifier (+/+; R/-) was dominant and independent of developmental temperature, larval density and diazinon concentration. Asymmetry score, pooled over three bristle characters, was ~50% greater for these phenotypes than for those of modified genotypes (M/-; -/-) and unmodified susceptibles (+/+; S/S) reared under standard laboratory conditions. Modified and susceptible phenotypes showed increased asymmetry score for temperatures and larval densities above and below standard rearing conditions; a positive correlation was observed between diazinon concentration and asymmetry score. Single and multiple environmental stresses resulted in similar scores that approached, but never exceeded, those of unmodified resistant phenotypes. Irrespective of the developmental conditions anti-symmetry and fluctuating asymmetry were typically observed for each bristle character of unmodified resistant and the modified and susceptible phenotypes, respectively. Thus while similar asymmetry scores could arise from genetic or environmental effects, asymmetry pattern was genetically based. Population cage analyses at different temperatures and larval densities showed a negative association between mean asymmetry and relative fitness.     

Modification of developmental instability and fitness: Malathion-resistance in the Australian sheep blowfly,Lucilia cuprina

The evolution of resistance to malathion byLucilia cuprina initially results in an increase in fluctuating asymmetry. Resistant flies are at a selective disadvantage, relative to susceptibles, in the absence of the insecticide. A fitness/asymmetry modifier of diazinon-resistant phenotypes ameliorates these effects resulting in malathion-resistant phenotypes of relative fitness and asymmetry similar to susceptibles. For the nine genotypic combinations of the modifier and malathion-resistance alleles, developmental time increases linearly with increasing asymmetry. Percentage egg hatch decreases linearly with increasing asymmetry. The initially disruptive effect of the malathion-resistant allele was partially dominant, the effect of the modifier dominant. The results are discussed in terms of developmental perturbation, asymmetry estimation and relative fitness to consider whether it is adequate to use changes in fluctuating asymmetry alone as measures of developmental instability. It is suggested that in some circumstances antisymmetry may indicate developmental instability and that the diazinon/malathion-resistance systems inL. cuprina may allow the relative importance of genetical and/or environmental developmental perturbations to be ascertained.     

Fluctuating asymmetry as an indicator of fitness: can we bridge the gap between studies?

There is growing evidence from both experimental and non-experimental studies that fluctuating asymmetry does not consistently index stress or fitness. The widely held--yet poorly substantiated--belief that fluctuating asymmetry can act as a universal measure of developmental stability and predictor of stress-mediated changes in fitness, therefore staggers. Yet attempts to understand why the reported relationships between fluctuating asymmetry, stress and fitness are so heterogeneous--i.e. whether the associations are truly weak or non-existent or whether they become confounded during different stages of the analytical pathways remain surprisingly scarce. Hence, we attempt to disentangle these causes, by reviewing the various statistical and conceptual factors that are suspected to confound potential relationships between fluctuating asymmetry, stress and fitness. Two main categories of factors are discerned: those associated with the estimation of developmental stability through fluctuating asymmetry and those associated with the effects of genotype and environment on developmental stability. Next, we describe a series of statistical tools that have recently been developed to help reduce this noise. We argue that the current lack of a theoretical framework that predicts if and when relationships with developmental stability can be expected, urges for further theoretical and empirical research, such as on the genetic architecture of developmental stability in stressed populations. If the underlying developmental mechanisms are better understood, statistical patterns of asymmetry variation may become a biologically meaningful tool.     

Post-embryonic development of Ophyra aenescens (Wiedemann, 1830) (Diptera: Muscidae) with different diets, under laboratory conditions

The performance of various diets (bovine meat, fish-sardine, shrimp, dog faeces, and banana) in Ophyra aenescens development was evaluated. The biology was studied in an incubator (BOD) at 27 +/- 1 degrees C and 80 +/- 10% of RH. The developmental time from larvae to adult, the developmental time and viability of larvae and pupae, the weight of pupae as well as the sex ratio of the emerging adults were also determined. Beef and shrimp were the more efficient diets for rearing O. aenescens.     

A true summer diapause induced by high temperatures in the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae)

Summer diapause in the cotton bollworm, Helicoverpa armigera (Hübner), which prolongs the pupal stage, particularly in males, is induced by high temperatures. In the laboratory, summer-diapausing pupae of H. armigera were induced at high temperatures (33-39 degrees C) with a photoperiod of LD8:16; winter-diapausing and non-diapausing pupae, cultured at 20 degrees C with a photoperiod of LD8:16 and at 27 degrees C, LD16:8, respectively, acted as a control. Retention time of eye spots, weight, and lipid and glycogen levels were compared. At high temperatures, both body weight and energy storage capacity were much higher in summer-diapausing pupae than in non-diapausing pupae reared at 33-39 degrees C. At temperatures (>33 degrees C) high enough to maintain summer diapause, the eye spots of summer-diapausing pupae did not move during the 30-day experiment. However, eye spots of summer-diapausing pupae placed at 30 degrees C began to move about 10 days after they were transferred, significantly later than in non-diapausing pupae reared at 33-39 degrees C or non-diapausing pupae reared at 27 degrees C, which initiated eye spot movement 2 days after pupation. The differences in retention time of eye spots between summer- and winter-diapausing pupae shows that winter diapause is more intense than summer diapause in this insect. The weight loss, and lipid and glycogen metabolism curves indicate that the summer-diapausing pupae's metabolism is very low. We conclude that summer diapause in the cotton bollworm is a true diapause and that the summer diapause enables the cotton bollworm to withstand the high temperatures of summer.     

Developmental stability,sexual selection and speciation

Fluctuating asymmetry occurs when an individual is unable to undergo identical development of an otherwise bilaterally symmetric trait on both sides of its body. Since both sides of a bilaterally symmetric trait are the result of the actions of a single genome, fluctuating asymmetry represents an epigenetic measure of the sensitivity of development to stress. Different morphological traits may show a direct relationship between their functional importance and their degree of developmental canalization. This may explain why some characters show high degrees of fluctuating asymmetry, and why these characters more often become exaggerated secondary sexual ornaments. The degree of fluctuating asymmetry is generally larger in small marginal populations living in novel environments, and this will particularly lead to relatively large degrees of asymmetry in the least developmentally canalized traits. More stringent selection against heterozygotes in marginal populations may further break down developmental stability and linkage groups which would lead to increased genetic variance. Females may prefer to mate with males having large, but relatively symmetric morphological characters, because it is more difficult to make large traits (a good genes argument), a large trait is more easily perceived (a sensory bias preference), and because symmetry signals ability to cope with stress (a good genes argument). The low degree of developmental stability and the large amount of genetic variance in secondary sexual characters in small, marginal populations could set the scene for rapid development of divergence and speciation in marginal populations.     

低温对小菜蛾实验种群的影响

研究了低温(<8℃)对小菜蛾的发育、存活和繁殖的影响结果表明,卵和蛹在4℃和6℃下死亡率随处理时间的延长而增加,在分别处理55d和70d后,卵和蛹全部死亡;经4℃和6℃处理的蛹,在16℃下羽化成虫的平均产卵量随处理时间的延长而减少,处理45d时,产卵量均为0小菜蛾幼期各虫态在0℃以下,死亡率随低温强度加大和处理时问的延长而增高就耐寒力而言,3龄幼虫和蛹最强,其次是2龄和4龄幼虫,卵和1龄幼虫的耐寒力最弱不同低温和时间处理小菜蛾幼期虫态对其后继虫态的发育历期有较大影响,总体说来,经过处理的小菜蛾幼期虫态,其后继虫态的发育历期普遍延长,一般处理某一虫态对其相邻虫态发育历期的影响最大小菜蛾蛹经低温处理后其羽化成虫的产卵量随着蛹期所经历低温强度的增强和时间延长而减少。     

Influence of incubation temperature on hatching success,energy expenditure for embryonic development,and size and morphology of hatchlings in the oriental garden lizard,Calotes versicolor (Agamidae)

We incubated eggs of Calotes versicolor at four constant temperatures ranging from 24 degrees C to 33 degrees C to assess the effects of incubation temperature on hatching success, embryonic use of energy, and hatchling phenotypes that are likely to affect fitness. All viable eggs increased in mass throughout incubation due to absorption of water, and mass gain during incubation was dependent on initial egg mass and incubation temperature. The average duration of incubation at 24 degrees C, 27 degrees C, 30 degrees C, and 33 degrees C was 82.1 days, 60.5 days, 51.4 days, and 50.3 days, respectively. Incubation temperature affected hatching success, energy expenditure for embryonic development, and several hatchling traits examined, but it did not affect the sex ratio of hatchlings. Hatching success was lowest (3.4%) at 33 degrees C, but a higher incidence of deformed embryos was recorded from eggs incubated at this temperature compared to eggs incubated at lower temperatures. Most of the deformed embryos died at the last stage of incubation. Energy expenditure for embryonic development was, however, higher in eggs incubated at 33 degrees C than those similarly incubated at lower temperatures. A prolonged exposure of eggs of C. versicolor at 33 degrees C appears to have an adverse and presumably lethal effect on embryonic development. Hatching success at 24 degrees C was also low (43.3%), but hatchlings incubated at 24 degrees C did not differ in any of the examined traits from those incubated at two intermediate temperatures (27 degrees C and 30 degrees C). Hatchlings incubated at 33 degrees C were smaller (snout-vent length, SVL) than those incubated at lower incubation temperatures and had larger mass residuals (from the regression on SVL) as well as shorter head length, hindlimb length, tympanum diameter, and eye diameter relative to SVL. Hatchlings from 33 degrees C had significantly lower scores on the first axis of a principal component analysis representing mainly SVL-free head size (length and width) and fore- and hindlimb lengths, but they had significantly higher scores on the second axis mainly representing SVL-free wet body mass. Variation in the level of fluctuating asymmetry in eye diameter associated with incubation temperatures was quite high, and it was clearly consistent with the prediction that environmental stress associated with the highest incubation temperatures might produce the highest level of asymmetry. Newly emerged hatchlings exhibited sexual dimorphism in head width, with male hatchlings having larger head width than females.     

ANALYSIS OF GENOTYPIC DIFFERENCES IN DEVELOPMENTAL STABILITY IN ANNONA CHERIMOLA

The genetic basis of developmental stability, measured as asymmetry (fluctuating asymmetry in leaves), was analyzed in leaves and flowers of cherimoya (Annona cherimola Mill) and atemoya (A. cherimola × A. squamosa). The individuals analyzed belonged to a controlled collection of cultivars (clones) that had previously been characterized by means of isozymes. We used a nested design to analyze the differences in asymmetry at several sampling levels: individual leaves and flowers, individual trees, and genotypes. The clonal repeatability of developmental stability was not significantly different from zero, thus suggesting the absence of heritability of the asymmetry for leaves and flowers under these environmental conditions. No relationship between asymmetry and individual heterozygosity was found, but leaf fluctuating asymmetry was significantly related to particular isozymic genes. Petal and leaf size showed a phenotypically plastic response to the exposure zone of the tree (mainly due to light). Leaf fluctuating asymmetry also showed such a plastic response. No significant correlation was found between asymmetry and any pomological characters (some of these being fitness related). Finally, the hybrid species (atemoya) did not show larger developmental instability than did the parental species (cherimoya). All these data show that cherimoya asymmetry reveals the random nature of developmental noise, with developmental stability for leaves being possibly related to specific chromosome regions, but with weak evidence for genotypic differences in developmental stability.     

Thermal requirements of Galleria mellonella L. (Lepidoptera: Pyralidae) immature stages

The rearing of Galleria mellonella L. in laboratory is important for multiplication of entomopathogenic nematodes, mandatory for biological control studies. The objective of this study was to evaluate the effect of three thermal profiles on development stages of this insect, allowing synchronization of cycle production. Two distinct rearing phases were done: firstly, using nucleous of incubation for development of eggs and, secondly, using circular-aluminum manifolds for development of larvae and pupae. The time necessary for development of the immature stages decreased with higher temperatures. Incubation periods lasted 13.4 days at 22 degrees C, 8.3 at 27 degrees C and 6.8 days at 32 degrees C, while periods for larvae development lasted 40.4, 27.2, and 23.4 days, respectively, for the same temperatures. Development to pupal stage was observed 18.2, 15.0, and 12.2 days, respectively, for the same temperatures. Larval survival was higher at 32 degrees C, however embryonic stages and pupae survival were higher at 27 degrees C. and 22 degrees C, respectively. The threshold temperature was 11.209167 degrees C for the embryonic development stage, 7.695869 degrees C for larval stage, and 1.943050 degrees C for pupal stage of G. mellonella. Thermal constants were 138.380533 DG (degree day) for egg, 554.968830 DG for larvae, and 369.054080 DG for pupae.     

Is There a Genetic Basis for Fluctuating Asymmetry and Does it Predict Fitness in the Plant Lotus corniculatus Grown in Different Environmental Conditions?

Fluctuating asymmetry (FA) is considered to be a good measure of developmental stability. We measured the asymmetry of leaves and flowers of 16 different genotypes of Lotus corniculatus grown in four different experimental environments to estimate the plasticity or developmental stability of asymmetry itself. We found that an index of FA (absolute difference between size of left and right sides, corrected for trait size) differed significantly across environments, with the treatment CO2+/N+ inducing the greatest FA for both flowers and leaves. Genotypes did not differ in FAs. Individual plants showed significantly different FAs only for flowers. At the individual level, we found no significant relationship between flower FA and fitness. Previous work indicates that change in asymmetry in a poor or perturbing environment versus a good environment could reflect the intrinsic quality of a particular genotype. However, in our experiment, genotype effect was significant only for change in asymmetry of leaves, and this last trait was not significantly correlated with our fitness estimate for each genotype in either the most or the least perturbing environment.     

Human tolerance to heat strain during exercise: influence of hydration.

This study determined whether 1) exhaustion from heat strain occurs at the same body temperatures during exercise in the heat when subjects are euhydrated as when they are hypohydrated, 2) aerobic fitness influences the body temperature at which exhaustion from heat strain occurs, and 3) curves could be developed to estimate exhaustion rates at a given level of physiological strain. Seventeen heat-acclimated men [maximal oxygen uptake (VO2max) from 45 to 65 ml.kg-1.min-1] attempted two heat stress tests (HSTs): one when euhydrated and one when hypohydrated by 8% of total body water. The HSTs consisted of 180 min of rest and treadmill walking (45% VO2max) in a hot-dry (ambient temperature 49 degrees C, relative humidity 20%) environment. The required evaporative cooling (Ereq) exceeded the maximal evaporative cooling capacity of the environment (Emax); thus thermal equilibrium could not be achieved and 27 of 34 HSTs ended by exhaustion from heat strain. Our findings concerning exhaustion from heat strain are 1) hypohydration reduced the core temperature that could be tolerated; 2) aerobic fitness, per se, did not influence the magnitude of heat strain that could be tolerated; 3) curves can be developed to estimate exhaustion rates for a given level of physiological strain; and 4) exhaustion was rarely associated with a core temperature up to 38 degrees C, and it always occurred before a temperature of 40 degrees C was achieved. These findings are applicable to heat-acclimated individuals performing moderate-intensity exercise under conditions where Ereq approximates or exceeds Emax and who have high skin temperatures.     

The influence of hybridization between African and European honeybees, Apis mellifera, on asymmetries in wing size and shape

Abstract We examined the possible role of hybridization in the invasion process of the African honeybee by testing two hypotheses regarding fluctuating asymmetry (FA), a measure of developmental stability, in wing characteristics: (1) FA should be higher in hybrid versus parental genotypes of African and European races; (2) FA should be lower in African bees compared to hybrid and European workers. Parental and reciprocal hybrid worker genotypes were cross fostered in common-hive rearing environments. We did not find greater FA for wing size and shape in the hybrids compared to both parental types. However, we did find significantly lower FA of shape in the African workers compared to the European and hybrid workers, suggesting that European bees and their hybrids may have compromised fitness relative to African bees. We also found that the two hybrid genotypes significantly differed in overall wing size and shape. If these differences affect wing aerodynamics, then the paternity of hybrids may influence worker performance and could potentially contribute to the loss of European matrilines. Hybridization had few consistent effects on directional asymmetry for wing size and shape. Genotypic factors played a far greater role in determining the effect of hybridization on wing morphology than did differences in rearing environment. Thus, African bees may have lower FA for wing shape (and by inference greater developmental stability) relative to European and hybrid workers, which may contribute to the ability of African bees to displace European honeybee races in invaded regions.     

The effect of lead on fitness components and developmental stability in Drosophila subobscura

We analyzed the developmental time, egg-to-adult viability, and developmental stability (fluctuating wing size asymmetry) in Drosophila subobscura, maintained for six generations on different concentrations of lead. Development time is significantly affected by generation and lead concentration, but interaction of these factors is not a significant source of variability for this fitness component. Generation and the interaction generation x concentration of lead significantly affect egg-to-adult viability. Levene's test of heterogeneity of variance showed that variability of FA is not significant in any of the samples. Within both lead concentrations females showed significantly higher FA indices for the wing width than males. Within sexes, a significantly higher FA was found only in females for wing width FA between the control and the lower concentration of lead. The results show that if strong relationship between FA and the studied fitness components exists, it results in a stronger selection of unstable genotypes under lead as a stress factor and, consequently, FA needs to be used with caution as a biomarker in natural populations under environmental stress.     

Developmental and reproductive biology of Scirtothrips perseae (Thysanoptera: Thripidae): a new avocado pest in California

The developmental and reproductive biology of a new avocado pest, Scirtothrips perseae Nakahara, was determined in the laboratory at five constant temperatures, 15, 20, 25, 27.5 and 30 degrees C. At 20 degrees C, S. perseae exhibited greatest larval to adult survivorship (41%), and mated females produced a greater proportion of female offspring at this temperature when compared to 15, 25, 27.5 and 30 degrees C. Average lifetime fecundity and preoviposition period was greatest at 15 degrees C at 39.6 eggs per female and 17.6 days, respectively. Jackknifed estimates of net reproduction (Ro), capacity for increase (rc), intrinsic rate of increase (rm), and finite rate of increase (lambda) were all significantly greater at 20 degrees C than corresponding values at 15, 25 and 27.5 degrees C. Population doubling time (Td) was significantly lower at 20 degrees C, indicating S. perseae populations can double 33-71% faster at this temperature in comparison to 15, 25 and 27.5 degrees C. Mean adult longevity decreased with increasing temperature, from a maximum of 52.4 days at 15 degrees C to a minimum of 2.4 days at 30 degrees C. Developmental rates increased linearly with increasing temperatures for eggs and rates were non-linear for development of first and second instar larvae, propupae, pupae, and for egg to adult development. Linear regression and fitting of the modified Logan model to developmental rate data for egg to adult development estimated that 344.8 day degrees were required above a minimum threshold of 6.9 degrees C to complete development. An upper developmental threshold was estimated at 37.6 degrees C with an optimal temperature of 30.5 degrees C for egg to adult development. Unmated females produced only male offspring confirming arrhenotoky in S. perseae.     

Modelling developmental instability in relation to individual fitness: a fully Bayesian latent variable model approach

Since the influential paper by Palmer and Strobeck in 1986, the statistical analysis of fluctuating asymmetry and developmental stability has received much attention. Most studies deal with one of the following four difficulties: (i) correcting for bias in asymmetry estimates due to measurement error; (ii) quantifying sampling error in the estimation of individual developmental stability using individual asymmetry; (iii) the detection of directional asymmetry and antisymmetry; and (iv) combining data from several traits. Yet, few studies have focused on statistical properties of estimating a relationship between individual developmental stability and other factors (e.g. fitness). In this paper I introduce a fully Bayesian model in which the unobservable individual developmental stability is treated as a latent variable. The latter is then related to individual fitness. I show by means of the analysis of simulated data that this approach has several advantages over traditional techniques. First, the method provides unbiased (but slightly less accurate) estimates of slopes between developmental stability and fitness taking all sources of error into account. Secondly, it allows proper investigation of non‐linear associations. Finally, the model allows unbiased estimation of unobserved fitness of individuals that have been measured on left and right side.     

Resistance to thermal stress in preadult Drosophila buzzatii: variation among populations and changes in relative resistance across life stages

Resistance to a short term exposure to a high temperature stress was examined in eggs, larvae and pupae of Drosophila buzzfltii from seven localities. Across development, pupae were most resistant, followed by eggs, and then first and third-instar larvae. Variation among populations for resistance to heat stress was significant in all life stages. However, there was much less variation among populations where measured as eggs and pupae than for both first and third instar larvae. Older larvae showed large changes both in viability and developmental time, while exposure of young larvae to heat stress led to a decline in viability without delayed development. Populations that had the shortest developmental time at 25^oC were relatively the most resistant to heat stress as larvae. High relative resistance at one preadult life stage was not necessarily associated with relatively high resistance at another, or with previous measurements of resistance for adults from these populations. Comparison of populations that were more similar in their pattern of change in resistance across development suggested a relationship with the climate of origin. The possibility that developmental variation in the expression of heat shock proteins may cause variation in resistance to thermal stress for different life stages is discussed.     

The effects of temperature, diet, and other factors on development, survivorship, and oviposition of Aethina tumida (Coleoptera: Nitidulidae)

Developmental rate and survivorship of small hive beetle, Aethina tumida Murray (Coleoptera: Nitidulidae), life stages were measured across different temperatures (21, 25, 28, 32 and 35 degrees C) and diets, which included natural and artificial pollen, honey, and bee pupae. Temperature affected hatch success, time to hatching, and larval growth. Eggs hatched in 61 h at 21 degrees C but in < 22 h at 35 degrees C. Larvae achieved peak weight in < 8 d at 35 degrees C but needed 17 d at 21 degrees C. Diet had comparatively little effect on larval survivorship or maximum weight, although larvae fed only bee pupae had lower survivorship. Access to soil influenced pupation success. Duration of the life stage spent in the soil, during which pupation occurs, was also affected by temperature: adults emerged after 32.7 d at 21 degrees C but after only 14.8 d at 35 degrees C, albeit with high mortality. Minimum temperature for development was estimated at 13.5 degrees C for eggs, and 10.0 degrees C for larvae and pupae. Temperature influenced adult longevity and oviposition: on a honey and pollen diet average adult lifespan was 92.8 d at 24 degrees C but only 11.6 d at 35 degrees C. Beetles lived longer at 28 degrees C or lower but produced the most eggs per female, regardless of diet, at 32 degrees C. Beetle density influenced fecundity: beetles kept at three pairs per vial laid 6.7 times more eggs per female than those kept as single pairs. Overall, beetles fared best at 28-32 degrees C with mortality of all stages highest at 35 degrees C.     

Measuring fitness and intergenic interactions: The evolution of resistance to diazinon inLucilia cuprina

Resistance at the diazinon-resistance locus (Rop-1) is recessive with respect to fitness. Selection initially occurs at concentrations lower than those required to controlLucilia cuprina. The presence of theRop-1 allele initially disrupted development so that in the absence of diazinon, carriers were at a relative selective disadvantage. Continued use of the chemical, subsequent to resistance evolving, selected a modifier to ameliorate this effect. Modified resistant phenotypes show similar developmental stability and relative fitness to susceptible individuals. Frequency-dependent interactions are observed between resistant and susceptible phenotypes of theRop-1 locus. The interactions are determined by the concentration of diazinon and range from competitive to facilitative. The results are discussed in the context of the contribution insecticide resistance systems can make to the study of general evolutionary phenomena.     

Fluctuating asymmetry and fitness in Drosophila melanogaster

Models predict that developmental stability measured by fluctuating asymmetry should be positively correlated with fitness. Although such a correlation has often been suggested by indirect studies, there is still a lack of direct experimental evidence. In this note, I have measured the fluctuating asymmetry of sternopleural bristle counts in 32 lines of Drosophila melanogaster sharing the same genetic background but displaying all combinations of five visible mutations. Fluctuating asymmetry was heterogeneous among lines, suggesting a direct impact of the mutations on developmental stability. Two measures of fitness were made for each line: productivity (a combined measure of fecundity and egg‐to‐adult survivorship) and competitive male mating success. Fluctuating asymmetry was correlated with neither of these two components of fitness. This suggests that generalizations about fluctuating asymmetry must be taken with care.