Fluctuating asymmetry in certain morphological traits in laboratory populations of Drosophila ananassae.

Fluctuating asymmetry (FA, subtle random deviations from perfect bilateral symmetry) is often used as a measure of developmental instability (DI), which results from perturbations in developmental pathways caused by genetic or environmental stressors. During the present study, we estimated FA in 5 morphological traits, viz. wing length (WL), wing to thorax ratio (W:T), sternopleural bristle number (SBN), sex-comb tooth number (SCTN), and ovariole number (ON) in 18 laboratory populations of Drosophila ananassae. FA levels of measured traits differed significantly among populations except for SBN (in males and females) and W:T ratio (in females). Positional fluctuating asymmetry (PFA), a sensitive measure of DI, also varied significantly among the populations for SBN in females and SCTN in males. Interestingly, both males and females were similar for nonsexual traits. However, when FA across all traits (sexual and nonsexual) was combined into a single composite index (CFA), significant differences were found for both populations and sexes. Males showed higher CFA values than females, suggesting that males are more prone to developmental perturbations. The magnitude of FA differed significantly among traits, being lowest for nonsexual traits (SBN, WL, W:T ratio) and highest for sexual traits (SCTN and ON). The trait size of sexual traits (SCTN and ON) was positively correlated with their asymmetry. The possible reasons for variation in FA both among traits and among populations, and the usefulness of FA as an indicator of developmental stress and phenotypic quality in D. ananassae are discussed.     

Does thermal-related plasticity in size and fat reserves influence supercooling abilities and cold-tolerance in Aphidius colemani (Hymenoptera: Aphidiinae) mummies?

(1) The parasitoid Aphidius colemani was reared at 15 or 25 °C to induce variation in size and fat reserves; SCP and cold-tolerance were compared. Insects from both temperatures were also exposed to constant or fluctuating cold-exposure.

(2) The lower SCP in mummies reared at 25 °C may be partially explained by their smaller size, a negative relationship being observed between SCP and size.

(3) A bimodality was observed in SCP distributions, with two modes around −26 and −22 °C, likely because of presence/absence of gut content.

(4) The type of exposure had a striking impact, mortality being considerably lower under fluctuating regime.

(5) While energy storage is an important factor, vulnerability to chill-injury is supposed to be the primary factor regulating survival at low temperature.     

Fluctuating asymmetry in hybrids of sibling species, Drosophila ananassae and Drosophila pallidosa, is trait and sex specific

Due to inconsistent results of the empirical studies, the relationship between fluctuating asymmetry (FA, a measure of developmental stability) and interspecific hybridization has been the subject of intense debates. In the present study, we have assessed the impact of interspecific hybridization between 2 sibling species of Drosophila: Drosophila ananassae and Drosophila pallidosa on the levels of FA over 3 generations. Trait size of different morphological traits, namely, sternopleural bristle number, wing length (WL), wing to thorax (W/T) ratio, sex comb tooth number (SCTN), and ovariole number differed significantly among parental species and their hybrids of different generations in both the sexes. However, the levels of FA of different morphological traits were similar in parental species and their hybrids of different generations in males (except SCTN) and in females (except for WL and W/T ratio). These results are interpreted in terms of developmental stability as a function of a balance between the level of heterozygosity and the disruption of coadapted gene complexes.     

Fluctuating asymmetry of sexual and nonsexual traits in stalk‐eyed flies: a poor indicator of developmental stress and genetic quality

It has been proposed that females use fluctuating asymmetry (FA) in sexual ornaments to assess male quality. FA of sexual traits is predicted to show greater sensitivity to stress than FA of nonsexual traits, and to be heritable. We used a half‐sib mating design and manipulation of larval food environment to test these predictions on stalk‐eyed flies, Cyrtodiopsis dalmanni, in which females prefer males with larger eyespans. We measured size and FA of eyestalks and of two nonsexually selected characters, wing length and width. We found no evidence of an increase in FA under larval food stress in any of the individual traits, although trait size decreased under stress. We combined FA across traits into a single composite index, and found that males reared in the most benign larval environment had significantly higher composite FA than males reared on other media. There was no such effect in females. Heritability of FA was not significantly different from zero in any of the traits, in any of the environments, although trait sizes showed high heritability. We conclude that FA in sexual and nonsexual traits is a poor indicator of developmental stress and genetic quality.     

Effect of mutations on developmental stability and canalization in morphological traits in Drosophila ananassae

The present study was designed to determine the effects of visible mutations of large effect on developmental stability and canalization in different morphological traits, namely, sternopleural bristle number, wing length, wing to thorax ratio, ovariole number, and sex comb tooth number (SCTN) in Drosophila ananassae. We have compared the mean trait size, fluctuating asymmetry (FA) (as an index of developmental stability), and morphological variation (as an index of canalization) of different mutant strains (yellow body color, y; claret eye color, ca; plexus wing, px; spread wing, spr; ebony body and sepia eye color, e se; yellow body and claret eye color, y ca; and cardinal eye color, curled wing, and ebony body color, cd cu e) with wild-type strain. The mean trait size of all morphological traits differs significantly among the wild-type and mutant strains. The wild-type and mutant strains vary significantly for the morphological variation and also for the levels of the FA in different morphological traits. However, we have found no increase in either the variance or in the degree of FA with the increase of the mutations (except in SCTN in y mutant). The plausible reasons for the variation in wild-type and mutant strains with particular reference to developmental stability and canalization have been discussed.     

Thermal Requirements and Parasitism Capacity of Trissolcus brochymenae (Ashmead) (Hymenoptera: Scelionidae) Under Constant and Fluctuating Temperatures, and Assessment of Development in Field Conditions

In order to understand the relative importance of Trissolcus brochymenae , a common egg parasitoid of the predatory stinkbug Podisus nigrispinus in areas where this predator is released, the thermal requirements and developmental periods (egg to adult) at constant (17, 20, 25, 28 and 32°C) and fluctuating (10-20, 13-23, 15-25, 17-27, 20-30 and 25-35°C) temperatures, and the effect of constant and fluctuating temperatures systems on the longevity and parasitism capacity of T. brochymenae were investigated. Based on thermal requirements of the parasitoid and field day/night temperature variation in the regions where P. nigrispinus was released, we calculated monthly and yearly number of generations of the parasitoid. Parameters of biological and thermal requirements of T. brochymenae were not affected when the insects were reared under constant or fluctuating temperatures. It was shown that males have a shorter developmental period and lower thermal requirements than females. The lowest parasitism capacity and shortest longevity of T. brochymenae were observed under fluctuating temperature regimes of 10-20oC and 25-35°C. A higher parasitism capacity was obtained under a 20-30°C regime (96.0 ±33.6 offspring per female), while longevity was longer at a constant temperature of 27°C (17.6 ±0.6 days). T. brochymenae showed a continuous development during all seasons through the year under field conditions, accomplishing 2.7, 2.4, 2.9 generations per month in the CAF, Pains, and Refloralje areas. An average of 26.4 and 25.8, 25.8 and 25.3, and 24.7 and 23.6 generations per year under constant and fluctuating temperatures systems was estimated for the CAF, Pains, and Refloralje areas, respectively. No significant difference between thermal systems in each area was observed; however, a significantly smaller number of generations per year were estimated to occur in the Refloralje area. Implications of these results for the forecasting of parasitoid population development and its impact on the host stinkbug predator are discussed.     

Is the number of possible QTL for asymmetry phenotypes dependent on thermal stress?

In bilateral organisms, fluctuating asymmetry (FA) was often used as an index of developmental instability but FA was not always found to be higher in stressful environments. An intercontinental set of recombinant inbred lines (RIL) was used to search for genetic variation in fluctuating asymmetry (FA) of both wing length (WL) and wing width (WW) in Drosophila melanogaster when reared at both benign (25 °C) and stressful (30 °C) temperatures. FA levels did not differ between benign and stressful temperatures. At benign temperature, no QTL was significant for FA. However, at stressful temperature, composite interval mapping revealed some QTL for FA in both WL and WW. QTL-based scans under stressful thermal environments may be an informative approach for either FA or developmental instability analyses, even when FA levels are similar between stressful and benign environments.     

Geography of fluctuating asymmetry in the greenfinch, Carduelis chloris

Levels of fluctuating asymmetry (FA) in 12 bilateral skeletal traits were estimated from 12 populations of greenfinches (Carduelis chloris) collected along a north‐south gradient across Europe. Average FA of measured traits was positively correlated with latitude indicating that the younger and genetically less diverse northern European populations are developmentally less stable than the older and genetically more diverse southern populations. Levels of FA differed significantly between different traits being lowest for functionally important traits (limb and wing bones) and highest for functionally less important traits such as foramina (apertures through bones)– a pattern that was highly concordant across different populations. Males tended to exhibit higher levels of FA than females, a finding consistent with the suggestions that males are more prone to developmental perturbations than females. Age differences in levels of FA were relatively clear, but inconsistent across traits with different degree of functionality. Individual heterozygosity – as enumerated from variation in allozyme loci – was unrelated to individual FA. No evidence for existence of individual asymmetry parameter (IAP) was found although traits related to locomotion indicated some degree of integration, which was expressed by correlations in the signed asymmetry. Nevertheless, an individual's overall asymmetry was poorly predicted by asymmetry of individual characters. Evidence for existence of population asymmetry parameter (PAP) was clear since all traits exhibited a similar degree of association with latitude. That the latitudinal cline of increasing FA towards north coincided with decreasing levels of genetic variability across the cline could be indicative of break down of developmental stability in the recently established and genetically impoverished populations. To what extent a reduced heterozygosity, the break up of co‐adapted gene complexes and/or environmental differences contributed to this process cannot be distinguished from our data.     

Effect of nifedipine on core cooling in rats during tail cold water immersion

Male rats (450 g, n=11/group) were heated at an ambient temperature of 42°C until a rectal temperature of 42.8°C was attained. Rats, then received either saline (30°C)+tail ice water immersion (F+I) or saline (30°C)+tail ice water immersion+Nifedipine, a peripheral vasodilator, (F+I+N) to determine cooling rate effectiveness and survivability. The time to reach a rectal temperature of 42.8°C averaged 172 min in both groups resulting in similar heating rates (0.029°C/min). The cooling rates in group F+I and F+I+N were not significantly different from each other. We conclude that since Nifedipine did not improve cooling rates when combined with fluid+tail ice water immersion, its use as a cooling adjunct does not seem warranted.     

Thermoregulatory use of heat increment of feeding in the tawny owl (Strix aluco)

The heat increment of feeding (HIF) was investigated in the tawny owl (Strix aluco) in central Norway (63°N, 10°E), close to the northern limit of its distribution. HIF was measured as the increase in heat production (measured as oxygen consumption) after force-feeding the owls with laboratory mice at thermoneutral conditions (20 °C) and during cold-exposure (5 °C and −5 °C). The basal metabolic rate of the owls (mean mass 419 g) was 4.39 kJ h⁻¹ and the lower critical temperature was approximately 16 °C. During cold conditions, HIF substituted for thermogenesis, and at an ambient temperature of −5 °C the substitution was complete. Calculations indicate that the substitution by HIF may save the owls as much as 60% of their daily thermoregulatory costs. This corresponds to about 10% of their total daily energy budget.     

Condition dependence of developmental stability in the sexually dimorphic fly Telostylinus angusticollis (Diptera: Neriidae)

Developmental stability is widely regarded as a condition‐dependent trait, but its relation to genotype and environment, and extent of developmental integration, remain contentious. In Telostylinus angusticollis, the dorsocentral bristles exhibit striking variation in developmental stability, manifested as fluctuating asymmetry (FA) in bristle position (‘positional FA’) and failure to develop some bristles (‘bristle loss’), in natural and laboratory populations. To determine whether this variation reflects condition, I tested for effects of genotype and environment (larval diet quality), and examined covariation with condition‐dependent traits. Positional FA was not affected by genotype or environment. However, positional FA covaried negatively with secondary sexual trait expression in males, and with sexual dimorphism in body shape, but covaried positively with body size in females. Bristle loss reflected both genotype and larval diet. Flies reared on poor‐quality diet exhibited a similar rate of bristle loss as wild flies. Both positional FA and bristle loss were greater in males. These results suggest that the relation between developmental stability and condition is complex and sex dependent.     

Stability of Development and Variability in Morphological Traits in a Natural Population of Drosophila melanogaster: Seasonal Dynamics in 1999

Seasonal dynamics of developmental stability and variability of morphological traits was examined in a natural population of Drosophila melanogaster in order to compare these two parameters as indicators of temperature stress. Morphometric (thorax length and wing length) and meristic (number of sternopelural and orbital bristles) were studied. Variability was measured as the coefficient of variation. Stability of development was estimated as fluctuating asymmetry (FA) of bilateral traits. Thorax length and wing length did not exhibit consistent seasonal trends whereas wing loading significantly decreased. Significant seasonal changes in FA were not detected in any trait examined. Two traits showed reduced variation in autumn. The use of FA as an indicator of ecological stresses in insects is discussed on the basis of these results and the literature evidence.     

Geographical variation in asymmetry in Gasterosteus aculeatus

We tested whether fluctuating asymmetry (FA) in undisturbed populations is associated with several natural environmental factors and whether FA is negatively correlated with fitness in the wild. We compared the FA of multiple bony structural defences among 87 endemic populations of threespine stickleback ( Gasterosteus aculeatus L.) inhabiting pristine freshwater habitats on the islands of Haida Gwaii, British Columbia, Canada. Multi-trait FA for adults and juvenile fish varied extensively among populations, but only in adults did it correlate with geography and two habitat characteristics (pH and water colour). Mean FA among individual traits varied concordantly among populations but was not correlated within individuals. While asymmetrical fish showed slightly higher levels of parasitism as predicted, selection differentials based on age class comparisons suggested that asymmetrical fish had the same or marginally higher survival than symmetrical fish. Selection differentials of FA varied significantly among traits and may reflect variability in their functional importance and in the strength of selection on their developmental stability. The data imply that FA/fitness associations are heterogeneous and character-specific.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 9–22.     

Temperature shock during development fails to increase the fluctuating asymmetry of a sexual trait in stalk-eyed flies.

The fluctuating asymmetry (FA) of bilateral traits is claimed to be a general indicator of environmental stress. Exaggerated sexual ornaments are thought to show elevated levels of FA and a greater response to stress than other traits. Previous work with stalk-eyed flies (Cyrtodiopsis dalmanni) has shown that the FA of the sexual trait (male eye stalks), wing length and wing width were unaffected by a continually applied food stress. Here we tested whether a transient stress (24-h heat shock at 31 degrees C during development) affected the FA of these traits. A second experiment tested the combined stresses of transient heat shock at 31 degrees C with continuous exposure to desiccation. In each experiment, temperature shock reduced the trait size, confirming that the treatments were stressful. However, stress had no effect on the FA of individual traits or the FA summed across all traits. Exposure to the combined stresses significantly elevated mortality and reduced trait size compared to the single-stress regime. However, FA did not differ significantly between flies from the two experiments. We found no evidence that FA in sexual and non-sexual traits reflects transient stress during the development of C. dalmanni.     

Experimentally Manipulated Growth Rate in Threespine Sticklebacks: Assessing Trade Offs with Developmental Stability

The evolution of adaptive growth rate and its influence on how other life history traits evolve is a neglected topic in biology. Growth rate influences life history because size strongly influences age-specific survival and fecundity, and because growth rate defines the relationship between age and size. Improved predictions about the evolution of life history traits may be possible with a greater understanding of the factors that influence the evolution of growth rate. We experimentally tested the hypothesis that a trade off may exist between growth rate and developmental stability in freshwater threespine sticklebacks, Gasterosteus aculeatus. We compared the degree of developmental instability (measured as fluctuating asymmetry = FA) in four lateral plate and two fin traits of fish reared under a high vs. low growth regime in response to food ration and temperature. We found evidence that symmetry was reduced (FA increased) in fast growing compared to slow growing fish, suggesting that a trade off between developmental stability and growth is possible. FA plausibly reflected developmental instability because of significant associations between rank FA levels across traits in individuals. These results are preliminary because of the possible confounding effects of temperature and food ration on asymmetry, and because we do not know if this trade off has fitness or other life history consequences. Our results also do not support the hypothesis of honest signaling sometimes invoked in studies of sexual selection because greater symmetry was found under poorer rather than better resource levels.     

Germination, Radial Growth, and Sporulation of Beauveria bassiana and Metarhizium anisopliae Isolates and Their Virulence to Chilo partellus (Lepidoptera: Pyralidae) at Different Temperatures

The effect of temperature regimes on conidia germination, radial growth and virulence of Beauveriabassiana and Metarhizium anisopliae against the spotted stalk borer Chilo partellus was investigated with Ethiopian isolates. Conidia germination, radial growth and sporulation of all isolates were retarded at 15 and 35°C. A suitable temperature range for the isolates was between 20 to 30°C. Conidia germination was less tolerant of low temperature (15°C) than radial growth. Radial growth and sporulation reacted differently to temperature. At both 25 and 30°C, all isolates induced 100% mortality to C. partellus larvae in six days. The LT₅₀ decreased for the isolates with increasing temperature.     

Stability in development and variability in morphological signs in a natural population of Drosophila melanogaster: seasonal dynamics in 1999

Seasonal dynamics of developmental stability and variability of morphological traits was examined in a natural population of Drosophila melanogaster in order to compare these two parameters as indicators of temperature stress. Morphometric (thorax length and wing length) and meristic (number of sternopelural and orbital bristles) were studied. Variability was measured as the coefficient of variation. Stability of development was estimated as fluctuating asymmetry (FA) of bilateral traits. Thorax length and wing length did not exhibit consistent seasonal trends whereas wing loading significantly decreased. Significant seasonal changes in FA were not detected in any trait examined. Two traits showed reduced variation in autumn. The use of FA as an indicator of ecological stresses in insects is discussed on the basis of these results and the literature evidence.     

Assemblage and population-level consequences of forest fragmentation on bilateral asymmetry in tropical montane birds

Habitat fragmentation has the potential to influence the development and thus the phenotype of organisms. The asymmetry of bilateral traits may be indicative of the extent to which developmental stability is compromised by the stressful conditions underlying fragmentation. Using an assemblage- and population-level approach, we explored asymmetry differences in tarsus and outermost tail feathers of birds inhabiting fragmented landscapes in the tropical Andes of Colombia. More than 2500 individuals of 185 species were mist-netted at nine forest sites representing continuous forest (> 1000 ha), medium- (70–110 ha), and small-sized (8–20 ha) fragments. Feathers showed true fluctuating asymmetry (FA), whereas tarsus presented a mixture of FA and directional asymmetry. Overall, asymmetry was lowest in continuous forest, and highest in small and medium fragments. These patterns remained unchanged when directionality and differences in species composition, abundance, and foraging tactics were considered. The population-level analyses showed a general trend of increased asymmetry variation in fragments, yet the responses were not always in the same direction. Increased asymmetry may represent an outcome of processes that contribute to the persistence of species in changing environments, and to the generation of phenotypic innovation, which suggests individual adjustments of development to deal with stress. This calls into question the deliberated application of FA as a biomonitoring tool for conservation.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 119–133.     

Morphological variability and developmental instability in subpopulations of the Eurasian badger (Meles meles) in Denmark

Aim Local populations from different geographical regions may differ in the selection regimes to which they are exposed. Differences in environmental factors and population density may affect the relative importance of different selective forces (e.g. natural vs. sexual selection). We suggest a direction of investigation concerned with the developmental instability of morphological traits. The goal is to disclose putative small‐scale geographical differences in the evolutionary forces, which may be hard to detect. Location Craniometrical investigations were carried out on ninety‐eight skulls and teeth of the Eurasian badger (Meles meles) collected during the period 1995–97 from three different populations in Denmark. One of these thrives at low population density, whereas the two others are characterized by high local density. Methods The skulls were investigated for developmental instability (DI) using fluctuating asymmetry (FA) as its estimator. FA was measured on canines, molars, premolar teeth and other skull and mandible traits. For the statistical analyses, we applied nonparametric permutation tests. Results Evidence was found suggesting differentiation among populations in mean degree of FA, and the FA values measured on canines were higher in the high‐density populations. FA of the canines was significantly higher in males than females, in contrast to FA of the other traits. Evidence of a negative relationship between canine size and their FA was found, whereas no significant correlations were found between the molar and premolar teeth measures and their FA. Main conclusions Our results suggest that canines could be under directional selection stemming from intrasexual competition, which may be stronger in high‐density zones. The other teeth investigated seem to be under a stabilizing regime hence their FA is mainly affected by environmental stresses. The negative relationship between canine size and FA found in males suggests the capacity of badgers to respond in an evolutionary way to environmental changes, despite the low genetic variability previously found at the molecular level.