Repeatability (R) in landmark coordinates and centroid size for constant condition and short-term heat stress experiments.

Repeatability (R) in landmark coordinates and centroid size for constant condition and short-term heat stress experiments.

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Hsp70 genes may influence the expression of wing abnormalities in Drosophila melanogaster but their effects on variability in quantitative characters and developmental instability are unclear. In this study, we focused on one of the six Hsp70 genes, Hsp70Ba, and investigated its effects on within- and among-individual variability in orbital bristle...

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... We can speculate that the differences in maneuverability during oviposition or mating may act as selective forces between genders. Wing morphology has been used as an indicator of changes in environmental conditions by measuring changes in developmental instability on the shape using fluctuating asymmetry (Woods et al., 1999;Hoffmann and Shirriffs, 2002;2005b;Takahashi et al., 2011). ...
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Butterflies display remarkable variation in wing shape and size, which is associated with tremendous ecological diversity. In butterflies, wing color patterns have been extensively studied, but wing shape and size diversity is less understood. In the present study, we investigated the altitude-specific wing asymmetry (right and left wings), sexual asymmetry, and wing covariation (proximal-distal and anterior-posterior wing organization) of Tirumala septentrionis (Butler, 1874). Using the geometric morphometric (GMM) method, we discovered that broad and expanded wings were found in lowland areas, whereas long and more rounded slender wings were found in highland areas. Furthermore, the ANOVA analysis, found significant (p < 0.05) altitude-specific shape and size differences in forewings and hindwings of both sexes. Thus, compared to size variations, the shape can be utilized as a valuable tool to distinguish sex (male and female), wing asymmetry (right and left) as well as altitudes (high and lowlands) specific variations of T. septentrionis. Furthermore, we noticed that lowland butterfly population had more asymmetric wings than the highland in the wing asymmetry analysis. We suspected that the unique wing morphology observed in the highland T. septentrionis population favored long-distance flight (indication of a migratory population). In contrast, the lowland population wing morphology indicated a newly colonized butterfly population. In addition to GMM analysis, we documented the FE-SEM wing scale morphology of T. septentrionis collected from two different altitudes. Interestingly, we observed that there is no difference in scale morphology related to altitude variations. Finally, we concluded that altitude-specific variations were observed mainly on wing shape and size, rather than the wing scales. These morphological variations were primarily associated with an adaptive response like migration, host searching, flight energy management and so on.
... They can be easily observed and measured, and, to the best of our knowledge, the FA index has not been determined in D. melanogaster larvae using these structures to date. Various studies have shown differences in the morphology of wings reflected in the FA are sensitive indicators of environmental stress [36][37][38]. Moreover, Debat et al. [36] indicated that wing shape can vary in more flexible ways and be more informative than simple traits such as the size of other body parts when investigating deteriorated larval survival, development, and adult fitness of insects exposed to environmental stressors (temperature, insecticides, and other chemical pollutants, dietary additives such as antimicrobial agents). ...
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The fruit fly Drosophila melanogaster is a model species used for a wide range of studies. Contamination of Drosophila cultures with bacterial infection is common and is readily eradicated by antibiotics. Neomycin antibiotics can cause stress to D. melanogaster's larvae and imagoes, which may affect the interpretation of the results of research using culture from neomycin-based medium. In the present study, fluctuating asymmetry (FA), one of the important bioindicators of stress, was measured. Larvae and imagoes of a wild-type D. melanogaster strain were exposed to various concentrations of neomycin. The size of anal papillae and selected wing veins were measured using scanning electron and light microscopy, respectively. Next, the FA was checked. The values obtained for larval anal papillae appeared to be concentration-dependant; the FA indices increased with the concentration of neomycin. The wing FA presented a large but variable correlation, depending on the measured vein. However, the mean length of veins was the highest for the control group, with neomycin-exposed groups showing lower values. The research showed that neomycin may cause sublethal stress in D. melanogaster, which manifests in increased FA indices. This suggests that neomycin can cause physiological and developmental stress in insects, which should be taken into account when interpreting the results of studies using these model organisms.
... The study showed that deletions do not affect the asymmetry of the Drosophila wing, but the development of canalization in a period of time changes. Consequently, genetic deletions have an impact on the canalization of the development of traits, but not on the stability of the development [34,35]. Genetic assimilation is the subsequent genetic fixing of the new trait in the population. ...
... Here, we highlight the role of the main classes of factors known to contribute to developmental buffering during cell and animal development. For specific examples and further theoretical and experimental advances see [176][177][178][179][180][181] and further factors can be uncovered by genome-wide screen using chromosomal deficiencies and classical screens in multicellular organisms [182]. As some factors with a demonstrated role in filtering noise and environmental/mutational variation may also be important for protecting genome stability, our classification as 'defences' and 'buffers' is only for convenience. ...
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Symmetric growth and the origins of fluctuating asymmetry are unresolved phenomena of biology. Small, and sometimes noticeable, deviations from perfect bilateral symmetry reflect the vulnerability of development to perturbations. The degree of asymmetry is related to the magnitude of the perturbations and the ability of an individual to cope with them. As the left and right sides of an individual were presumed to be genetically identical, deviations of symmetry were traditionally attributed to non-genetic effects such as environmental and developmental noise. In this review, we draw attention to other possible sources of variability, especially to somatic mutations and transposons. Mutations are a major source of phenotypic variability and recent genomic data have highlighted somatic mutations as ubiquitous, even in phenotypically normal individuals. We discuss the importance of factors that are responsible for buffering and stabilizing the genome and for maintaining size robustness and quality through elimination of less-fit or damaged cells. However, the important question that arises from these studies is whether this self-correcting capacity and intrinsic organ size controls are sufficient to explain how symmetric structures can reach an identical size and shape. Indeed, recent discoveries in the fruit fly have uncovered a conserved hormone of the insulin/IGF/relaxin family, Dilp8, that is responsible for stabilizing body size and symmetry in the face of growth perturbations. Dilp8 alarm signals periphery growth status to the brain, where it acts on its receptor Lgr3. Loss of Dilp8-Lgr3 signaling renders flies incapable of detecting growth perturbations and thus maintaining a stable size and symmetry. These findings help to understand how size and symmetry of somatic tissues remain undeterred in noisy environments, after injury or illnesses, and in the presence of accumulated somatic mutations.
... The shape of such norm of reaction curves can vary among genoytpes (Fig. 1B). However, they might also result from other factors such as destabilizing effects of environmental stress or genetic perturbations [44] (Fig. 1C). Scharloo's classic experiments with the Drosophila cubitus interruptus and hairless mutants showed that selection can alter the phenotypic responses to temperature variation [45,46]. ...
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Canalization, or robustness to genetic or environmental perturbations, is fundamental to complex organisms. While there is strong evidence for canalization as an evolved property that varies among genotypes, the developmental and genetic mechanisms that produce this phenomenon are very poorly understood. For evolutionary biology, understanding how canalization arises is important because, by modulating the phenotypic variation that arises in response to genetic differences, canalization is a determinant of evolvability. For genetics of disease in humans and for economically important traits in agriculture, this question is important because canalization is a potentially significant cause of missing heritability that confounds genomic prediction of phenotypes. We review the major lines of thought on the developmental-genetic basis for canalization. These fall into two groups. One proposes specific evolved molecular mechanisms while the other deals with robustness or canalization as a more general feature of development. These explanations for canalization are not mutually exclusive and they overlap in several ways. General explanations for canalization are more likely to involve emergent features of development than specific molecular mechanisms. Disentangling these explanations is also complicated by differences in perspectives between genetics and developmental biology. Understanding canalization at a mechanistic level will depend on conceptual and methodological approaches that integrate quantitative genetics and developmental biology.
... In Drosophila, other molecular chaperones-Hsp22, Hsp67, and Hsp70-were also observed to affect either within-individual variation (measured by asymmetry of bilateral traits) or among-individual variation in morphology [67]. In eukaryotes, Hsp90 impairment has been found to reveal CGV in organisms ranging from yeast to flies to vertebrates to plants [68,69]. ...
... Therefore, deformation in geometric shape may not fully reflect biological shape deformation in such cases. The interpretation of geometric shape deformation in a biologically relevant way is a common challenge in studies of genetic and developmental processes leading to wing shape in D. melanogaster (Palsson & Gibson 2004;Takahashi et al. 2011a). ...
... Geometric morphometrics technology has gained considerable momentum and is making rapid progress. In entomology, 2-D data have been the primary sources for analyses, particularly on flat organs such as the wings and the posterior lobe of epandrium (a part of male genitalia) in Drosophila (Zeng et al. 2000;Takahashi et al. 2011a), and even on cubic organs such as mandibles of stag beetles (Tatsuta et al. 2004). Obviously, in the latter case, the comparison of shape features should ideally be analyzed in 3-D. ...
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The recent expansion of a variety of morphometric tools has brought about a revolution in the comparison of morphology in the context of the size and shape in various fields including entomology. First, an overview of the theoretical issues of geometric morphometrics is presented with a caution about the usage of traditional morphometric measurements. Second, focus is then placed on two broad approaches as tools for geometric morphometrics; that is, the landmark-based and the outline-based approaches. A brief outline of the two methodologies is provided with some important cautions. The increasing trend of entomological studies in using the procedures of geometric morphometrics is then summarized. Finally, information is provided on useful toolkits such as computer software as well as codes and packages of the R statistical software that could be used in geometric morphometrics.
... Some studies investigated how variation patterns are affected by developing under suboptimal conditions, and used canalization and DS as proxies for detecting disturbance (e.g., Parsons, 1990Parsons, , 1992, providing evidence that development may easily be hampered by both environmental and genetic factors, which can be identified as increased levels of fluctuating asymmetry (FA), variance, deviance from allometric trajectories, or all the above (Lazić, Carretero, Crnobrnja-Isailović, & Kaliontzopoulou, 2015). Other studies focused on determining the mechanisms responsible for DS and canalization by identifying the genes involved in the development of the examined morphological trait(s) (e.g., Debat & Peronnet 2013;Debat, Debelle, & Dworkin, 2009;Debat et al., 2011), or, by investigating the effects of heatshock proteins (e.g., Debat, Milton, Rutherford, Klingenberg, & Hoffmann, 2006;;Milton, Huynh, Batterham, Rutherford, & Hoffmann, 2003;Takahashi, Daborn, Hoffmann, & Takano-Shimizu, 2011) or mutations (e.g., DeLaurier et al., 2014, highlighting the important, although variable, contribution of genetic factors in shaping DS and canalization. ...
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Canalization and developmental stability (DS) are important organismal properties involved in determining the level of phenotypic variation. Ontogenetic patterns of phenotypic variance components can shed light on the mechanistic basis of developmental buffering (DB). Here, we analyze how individual FA and among-individual variation in head shape change in ontogenetic series of three lizard species raised in laboratory. The degree of asymmetry increased slightly with size, suggesting that developmental mechanisms hypothesized to correct for deviations either do not exist, or that their efficiency is truncated with increasing size. Alternatively, they may need the disturbance as a trigger. The relationship between asymmetry and age was complex, with asymmetry being stable across the age range in two species but increased with age in the third. Lack of congruence in ontogenetic patterns of asymmetry might be due to intrinsic differences in buffering mechanisms or a result of species-specific growth patterns. Head shape was shown to be equally canalized across both size and age range in all species, probably as a result of a balance between the buffering mechanisms and mechanisms generating variance. The patterns of symmetric and asymmetric head shape variation were highly correlated across species meaning that DS and canalization may rely on similar mechanisms.
... hsp22, hsp67 и hsp70) are also apparently involved in the control of normal morphogenesis and developmental stability. [27][28][29] Therefore, the accumulated data implicated hsps genes not only as components of powerful antistress system enabling organism to survive various challenges but also as agents necessary for normal development providing genome stability under various conditions. It is not quite clear whether HS system as a whole and individual Hsps may interact with components of piRNA-system and modify MEs activity under normal conditions and after stress. ...
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Different types of stress including heat shock may induce genomic instability, due to the derepression and amplification of mobile elements (MEs). It remains unclear, however, whether piRNA-machinery regulating ME expression functions normally under stressful conditions. The aim of this study was to explore the features of piRNA expression after heat shock (HS) exposure in Drosophila melanogaster. We also evaluated functioning of piRNA-machinery in the absence of major stress protein Hsp70 in this species. We analyzed the deep sequence data of piRNA expression after HS treatment and demonstrated that it modulates the expression of certain double-stranded germinal piRNA-clusters. Notable, we demonstrated significant changes in piRNA levels targeting a group of MEs after HS only in the strain containing normal set of hsp70 genes. Surprisingly, we failed to detect any correlation between the levels of piRNAs and the transcription of complementary MEs in the studied strains. We propose that modulation of certain piRNA-clusters expression upon HS exposure in D. melanogaster occurs due to HS-induced altering of chromatin state at certain chromosome regions.
... There appear to be remarkably few exceptions to this pattern. A few studies of shape in Drosophila found no significant directional asymmetry of wing shape [115][116][117][118][119] or mixed results [120][121][122][123][124][125][126], although a series of other studies did find it [15,16,54,56,62,77,85,123]. Similarly, one study on human skulls [127] found no directional asymmetry of shape, whereas several others reported directional asymmetry of the skull [47,68,84,86,90,94,109] and soft tissues of the face and ears [38,66,98,104,105,108]. ...
... Some empirical studies, using geometric morphometric methods, focused on molecular chaperones such as Hsp90 and other heat shock proteins [124,126,177,337] and genes involved in various ways in the development of the structures under study [62,332,338] and found variable effects of those genes on shape asymmetry. Additional studies have found pronounced effects of mutations in various genes on the fluctuating asymmetry in the size of structures and, even though they have not specifically investigated shape, sometimes found evident asymmetries of shape [333,339,340]. ...
... Genetic studies of fluctuating asymmetry are complicated by the fact that the measurable asymmetries themselves are of non-genetic nature: what is of interest, therefore, is the genetic basis of developmental instability, the tendency of organisms with particular genotypes and in the given environment to produce left-right asymmetry. The most direct way to implement experiments that reflect this situation is to use model organisms where it is easy to obtain many individuals with identical genotypes [52,56,77,[124][125][126], but traditional approaches from quantitative genetics can also be used, such as parent-offspring regression [128], analyses of variation among inbred lines [85], or pedigree-based methods [85]. ...
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Approximately two decades after the first pioneering analyses, the study of shape asymmetry with the methods of geometric morphometrics has matured and is a burgeoning field. New technology for data collection and new methods and software for analysis are widely available and have led to numerous applications in plants and animals, including humans. This review summarizes the concepts and morphometric methods for studying asymmetry of shape and size. After a summary of mathematical and biological concepts of symmetry and asymmetry, a section follows that explains the methods of geometric morphometrics and how they can be used to analyze asymmetry of biological structures. Geometric morphometric analyses not only tell how much asymmetry there is, but also provide information about the patterns of covariation in the structure under study. Such patterns of covariation in fluctuating asymmetry can provide valuable insight about the developmental basis of morphological integration, and have become important tools for evolutionary developmental biology. The genetic basis of fluctuating asymmetry has been studied from empirical and theoretical viewpoints, but serious challenges remain in this area. There are many promising areas for further research that are only little explored at present.