Thermal preference in Drosophila

Environmental temperature strongly affects physiology of ectotherms. Small ectotherms, like Drosophila, cannot endogenously regulate body temperature so must rely on behavior to maintain body temperature within a physiologically permissive range. Here we review what is known about Drosophila thermal preference. Work on thermal behavior in this group is particularly exciting because it provides the opportunity to connect genes to neuromolecular mechanisms to behavior to fitness in the wild.     

Maximum voluntary temperature of insect larvae reveals differences in their thermal biology

We investigate the thermoregulatory behaviors of larvae of four species of Drosophila (D. melanogaster, D. subobscura, D. pseudoobscura, and D. mojavensis), a thermotolerant strain of Drosophila melanogaster (T strain) known to differ in thermal biology, and two mutant stocks of D. melanogaster that have (as adults) defective thermoregulatory behavior. We describe and evaluate new techniques to measure two indices of maximum voluntary temperature of insect larvae. Both measures were highly repeatable within lines (species, strains, or mutants). One measure (temperature at which larvae stood upright) differed among lines consistent with expectations based on adult thermal ecology, suggesting that this measure will be useful measures of thermoregulatory set-points of larvae. The second measure (temperature of emergence from media) is less discriminatory.     

Keeping warm in the cold: On the thermal benefits of aggregation behaviour in an intertidal ectotherm

Temperature is a primary determinant for species geographic ranges. In the context of global warming, most attention focuses upon the potential effects of heat stress on the future distribution of ectothermic species. Much less attention has, however, been given to cold thermal stress although it also sets species thermal window limits, hence distribution ranges. This study was conducted in winter on a South-Australian rocky shore in order to investigate the potential thermal benefits of the aggregation behavior observed in the dominant gastropod Nerita atramentosa. Thermal imaging was used to measure the body temperatures of 3681 aggregated individuals and 226 solitary individuals, and surrounding substratum temperature. N. atramentosa aggregates and solitary individuals were significantly warmer than their surrounding substratum. The temperature deviation between aggregates and substratum was, however, ca. 2 °C warmer than the one observed between solitary individuals and substratum. This result is critical since a body temperature increase of only a few degrees might enhance individual performance, hence organismal fitness, and could potentially drive changes in interspecific relationships. Besides, the potential higher thermal inertia of aggregates might increase the snail adaptive ability to abrupt environmental changes. We further investigate the potential thermal heterogeneity within an aggregate in order to identify any thermally advantageous position. Patch centers are significantly warmer than their edges, hence snails experience greater thermal advantages in the aggregate center. Finally, we examined the potential effect of aggregate size on snail temperature and thermal spatial heterogeneity. We identified an aggregate size threshold (216 individuals) beyond which all snails had equal thermal benefits, regardless of their spatial positions within an aggregate. While the determinism of this aggregate size threshold requires further investigations, the present work uniquely identified the thermal benefits of aggregation behavior for intertidal ectotherms under cold weather conditions. The implications of the present finding are discussed in the general framework of the ability of ectothermic populations to face environmental changes.     

Photoperiod modifies thermal reaction norms for growth and development in the red poplar leaf beetle Chrysomela populi (Coleoptera: Chrysomelidae)

Regression lines of development rate on temperature appeared significantly different between long (22 h) and short (12 h) day conditions and intersected each other at 23.8 °С. Thus, the rate of growth and development was higher at temperatures below the intersection point under short-day but above the intersection point it was higher under long day. Ecological relevance of this effect seems as follows: in autumn, as nights become longer and average daily temperature decreases, larvae have to speed up their development because it is only imago that overwinters. Conversely, midsummer offers long days and usually higher temperature, so again it is advantageous to develop as fast as possible in order to have at least one more generation per year. These results are compared with other studies showing interactions between photoperiod and temperature, and some possible general patterns are outlined. The lower thermal threshold for larval development depended on photoperiodic conditions; therefore rate isomorphy must be violated in this species. Development at higher temperatures generally resulted in smaller adults, as is usual with ectotherms according to the “temperature-size rule”, but body weight depended significantly on temperature only under short day. Our estimates of the lower temperature thresholds for growth and development in both cases did conform to the generalization made previously by Walters and Hassall (2006) in spite of another formula used by us. We briefly discuss this phenomenon and argue that relative position of these thresholds can be explained mathematically and per se may lack any biological sense.     

A place preference test in the fish Nile tilapia

The Nile tilapia fish (Oreochromis niloticus) has a high potential to be used as a model in neuroscience studies. In the present study, the preference of the Nile tilapia between a gravel-enriched (GEE), a shelter-enriched (SEE) or a non-enriched (NEE) environment was determined, for developing a place preference model. Nile tilapia had an initial preference for GEE, but after 1 day of observation, the fish stabilized their frequency of visits among compartments. Hence, any stimulus motivating tilapia increase in compartment visiting indicates a positively reinforcing effect. This feature is very useful for the development of new behavioural paradigms for fish in tests using environmental discrimination, such as the conditioning place preference test.     

Colonization and substratum preference of an introduced burrowing crustacean in a temperate estuary

Habitat use in marine invertebrates is often influenced by multiple abiotic and biotic factors. Substratum composition is one factor known to have a dramatic effect on habitat selection. The Australasian burrowing isopod (Sphaeroma quoianum, H. Milne Edwards 1840) is a common introduced species in many estuaries on the Pacific coast of North America. S. quoianum burrows into a variety of firm substrata including marsh banks (composed of peat, clay, and/or mud), wood, friable rock, and Styrofoam floats. In some areas, isopods achieve high densities and may accelerate the rate of shoreline erosion and damage marine structures; thus, understanding the substratum preference of this species may be important for conservation and management efforts. Field experiments were conducted in Coos Bay, Oregon to examine substratum preference, burrowing rates, and the life stage of colonizers. In three experimental trials (Fall 2005, Spring 2006, Fall 2006), replicates of four intertidal substrata (marsh banks, decayed wood, sandstone, Styrofoam) were deployed near intertidal populations of S. quoianum. The numbers of burrows created in each substratum were enumerated weekly or daily (depending on trial). After the trials were completed, the total numbers of isopods inhabiting each substratum were counted. In weeks, S. quoianum extensively burrowed the substrata but exhibited a distinct preference for decayed wood. Significantly more isopods were present in wood than the other substrata at the end of the experiments and rates of burrowing were greatest in wood, although significance varied across time in one trial. Nearly 90% of colonizing isopods were under 5 mm in length suggesting that juvenile isopods primarily colonize intertidal substrata. Differences between burrow densities measured in the field and the results from these preference trials may indicate other factors, such as relative availability of substrata, recruitment and dispersal limitations, and possible gregarious behavior also influence local isopod densities.     

Optimal foraging versus shared doom effects: interactive influence of mussel size and epibiosis on predator preference

In the western Baltic Sea, the highly competitive blue mussel Mytilus edulis tends to monopolize shallow water hard substrata. In many habitats, mussel dominance is mainly controlled by the generalist predator Carcinus maenas. These predator-prey interactions seem to be affected by mussel size (relative to crab size) and mussel epibionts.There is a clear relationship between prey size and predator size as suggested by the optimal foraging theory: Each crab size class preferentially preys on a certain mussel size class. Preferred prey size increases with crab size.Epibionts on Mytilus, however, influence this simple pattern of feeding preferences by crabs. When offered similarly sized mussels, crabs prefer Balanus-fouled mussels over clean mussels. There is, however, a hierarchy of factors: the influence of attractive epibiotic barnacles is weaker than the factor ‘mussel size’. Testing small mussels against large mussels, presence or absence of epibiotic barnacles does not significantly alter preferences caused by mussel size. Balanus enhanced crab predation on mussels in two ways: Additional food gain and, probably more important, improvement in handling of the prey. The latter effect is illustrated by the fact that artificial barnacle mimics increased crab predation on mussels to the same extent as do live barnacles.We conclude that crab predation preferences follows the optimal foraging model when prey belong to different size classes, whereas within size classes crab preferences is controlled by epibionts.     

Evolutionarily conserved thermal biology across continents: The North American lizard Plestiodon gilberti (Scincidae) compared to Asian Plestiodon

1.

The lizard genus Plestiodon (Scincidae; formerly Eumeces) is widespread in North America and Asia, but the thermal biology of only a few species has been studied.     

Myoblast fusion in Drosophila

The body wall musculature of a Drosophila larva is composed of an intricate pattern of 30 segmentally repeated muscle fibers in each abdominal hemisegment. Each muscle fiber has unique spatial and behavioral characteristics that include its location, orientation, epidermal attachment, size and pattern of innervation. Many, if not all, of these properties are dictated by founder cells, which determine the muscle pattern and seed the fusion process. Myofibers are then derived from fusion between a specific founder cell and several fusion competent myoblasts (FCMs) fusing with as few as 3-5 FCMs in the small muscles on the most ventral side of the embryo and as many as 30 FCMs in the larger muscles on the dorsal side of the embryo. The focus of the present review is the formation of the larval muscles in the developing embryo, summarizing the major issues and players in this process. We have attempted to emphasize experimentally-validated details of the mechanism of myoblast fusion and distinguish these from the theoretically possible details that have not yet been confirmed experimentally. We also direct the interested reader to other recent reviews that discuss myoblast fusion in Drosophila, each with their own perspective on the process [1], [2], [3] and [4]. With apologies, we use gene nomenclature as specified by Flybase (http://flybase.org) but provide Table 1 with alternative names and references.     

Thermal behavior of the Pacific sardine (Sardinops sagax) acclimated to different thermal cycles

The study of thermal behavior of fishes provides useful data to enable predictions of the effect of climatic change on populations so as to ensure good management of fisheries. The Pacific sardine has a complex population dynamics; three subpopulation have been proposed (cold, temperate and warm).We exposed Sardinops sagax caeruleus (temperate sub-population) to two different thermal cycles, which were chosen to be consistent with the temperatures reported in two distant places Cedros Island (CIc: 18–23 °C) and San Pedro (SPc: 13–18 °C).The thermal behavior of the SPc and CIc sardines was affected by acclimation treatment: the interval of thermal preference was 17.1–19.9 and 16.0–18.8 °C, while the lethal temperatures interval (LT50) was 7.7–25.6 and 6.9–24.3 °C, and the critical limits CTMax and CTMin were 7.1–32.2 and 5.5–30.4 °C, respectively.The results of thermal behavior showed that sardines of the temperate subpopulation are more tolerant to cold; this might suggest that they would be more able to survive in California and Oregon than in Baja California Sur and the Gulf of California.     

Seasonality provokes a shift of thermal preferences in a temperate lizard,but altitude does not

1. We compared the mean, limits and breadth of the preferred thermal range (PTR) of two Iberian populations of the lizard Psammodromus algirus separated by 700 m altitude in May and July.     

Effect of thermal environment on life histories of free living Drosophila melanogaster and D. subobscura

The variability of morphological and lifehistory traits in Drosophila melanogaster and D. subobscura, two sympatric Drosophila species of different climatic origin, were investigated with regard to seasonal and daily temperature and humidity fluctuations to assess thermal sensitivity and the responses of freeliving organisms to oscillating temperatures. Temperature and humidity were measured continuously at the site where Drosophila were observed throughout the day, and thus represent a realistic picture of the temperature environment of these animals. A phenotypic model (gaussian curve) of temperature performance was fitted to daily fecundities and provided estimates of the position of temperature optimum and of the maximum and the breadth of performance for both species. D. melanogaster had a higher optimum temperature than D. subobscura. A trade-off between the maximum and the breadth of performance was detected both within and between the species.     

Spatial and temporal regulation of the homeotic selector gene Antennapedia is required for the establishment of leg identity in Drosophila

Antennapedia is one of the homeotic selector genes required for specification of segment identity in Drosophila. Dominant mutations that ectopically express Antennapedia cause transformation of antenna to leg. Loss-of-function mutations cause partial transformation of leg to antenna. Here we examine the role of Antennapedia in the establishment of leg identity in light of recent advances in our understanding of antennal development. In Antennapedia mutant clones in the leg disc, Homothorax and Distal-less are coexpressed and act via spineless to transform proximal femur to antenna. Antennapedia is negatively regulated during leg development by Distal-less, spineless, and dachshund and this reduced Antennapedia expression is needed for the proper development of distal leg elements. These findings suggest that the temporal and spatial regulation of the homeotic selector gene Antennapedia in the leg disc is necessary for normal leg development in Drosophila.     

A critical role of teashirt for patterning the ventral epidermis is masked by ectopic expression of tiptop, a paralog of teashirt in Drosophila

The teashirt gene encodes a protein with three widely spaced zinc finger motifs that is crucial for specifying trunk identity in Drosophila embryos. Here, we describe a gene called tiptop, which encodes a protein highly similar to Teashirt. We have analyzed the expression patterns and functions of these two genes in the trunk of the embryo. Initially, teashirt and tiptop expressions are detected in distinct domains; teashirt in the trunk and tiptop in parts of the head and tail. In different mutant situations, we show that, in the trunk and head, they repress each other's expression. Unlike teashirt, we found that deletion of tiptop is homozygous viable and fertile. However, embryos lacking both gene activities display a more severe trunk phenotype than teashirt mutant embryos alone. Ectopic expression of either gene produces an almost identical phenotype, indicating that Teashirt and Tiptop have, on the whole, common activities. We conclude that Teashirt and Tiptop repress each other's expression and that Teashirt has a crucial role for trunk patterning that is in part masked by ectopic expression of Tiptop.     

Premature differentiation and aberrant movement of pituitary cells lacking both Hes1 and Prop1

In the pituitary, the transition from proliferating progenitor cell into differentiated hormone producing cell is carefully regulated in a time-dependent and spatially-restricted manner. We report that two targets of Notch signaling, Hes1 and Prop1, are needed to maintain progenitors within Rathke's pouch and for the restriction of differentiated cells to the ventral pituitary. We observed ACTH and αGSU producing cells that had prematurely differentiated within Rathke's pouch along with correlated ectopic expression of Mash1 only when both Prop1 and Hes1 were lost. We also discovered that downregulation of N-cadherin expression in cells as they transition from Rathke's pouch to the anterior lobe appears to be essential for their movement. In the Prop1 mutant, cells are trapped in Rathke's pouch and N-cadherin expression remains high. Also, Slug, a marker of epithelial-to-mesenchymal transition, is absent in the dorsal anterior lobe. When Hes1 is lost in the Prop1 mutant, N-cadherin is downregulated and cells are able to exit Rathke's pouch but have lost their migrational cues and form ectopic foci surrounding Rathke's pouch. Our data reveal important overlapping functions of Hes1 and Prop1 in cell differentiation and movement that are critical for pituitary organogenesis.     

Regulation of R7 and R8 differentiation by the spalt genes

Photoreceptor development begins in the larval eye imaginal disc, where eight distinct photoreceptor cells (R1-R8) are sequentially recruited into each of the developing ommatidial clusters. Final photoreceptor differentiation, including rhabdomere formation and rhodopsin expression, is completed during pupal life. During pupation, spalt was previously proposed to promote R7 and R8 terminal differentiation. Here we show that spalt is required for proper R7 differentiation during the third instar larval stage since the expression of several R7 larval markers (prospero, enhancer of split mdelta0.5, and runt) is lost in spalt mutant clones. In R8, spalt is not required for cell specification or differentiation in the larval disc but promotes terminal differentiation during pupation. We show that spalt is necessary for senseless expression in R8 and sufficient to induce ectopic senseless in R1-R6 during pupation. Moreover, misexpression of spalt or senseless is sufficient to induce ectopic rhodopsin 6 expression and partial suppression of rhodopsin 1. We demonstrate that spalt and senseless are part of a genetic network, which regulates rhodopsin 6 and rhodopsin 1. Taken together, our results suggest that while spalt is required for R7 differentiation during larval stages, spalt and senseless promote terminal R8 differentiation during pupal stages, including the regulation of rhodopsin expression.     

Drosophila homologue of the Rothmund-Thomson syndrome gene: essential function in DNA replication during development

Members of the RecQ family play critical roles in maintaining genome integrity. Mutations in human RecQL4 cause a rare genetic disorder, Rothmund-Thomson syndrome. Transgenic mice experiments showed that the RecQ4 null mutant causes embryonic lethality. Although biochemical evidence suggests that the Xenopus RecQ4 is required for the initiation of DNA replication in the oocyte extract, its biological functions during development remain to be elucidated. We present here our results in establishing the use of Drosophila as a model system to probe RecQ4 functions. Immunofluorescence experiments monitoring the cellular distribution of RecQ4 demonstrated that RecQ4 expression peaks during S phase, and RecQ4 is expressed only in tissues active in DNA replication, but not in quiescent cells. We have isolated Drosophila RecQ4 hypomorphic mutants, recq^EP and recq4²³, which specifically reduce chorion gene amplification of follicle cells by 4-5 fold, resulting in thin and fragile eggshells, and female sterility. Quantitative analysis on amplification defects over a 14-kb domain in chorion gene cluster suggests that RecQ4 may have a specific function at or near the origin of replication. A null allele recq4¹⁹ causes a failure in cell proliferation, decrease in DNA replication, chromosomal fragmentation, and lethality at the stage of first instar larvae. The mosaic analysis indicates that cell clones with homozygous recq4¹⁹ fail to proliferate. These results indicate that RecQ4 is essential for viability and fertility, and is required for most aspects of DNA replication during development.     

Suppressor of Hairless and Presenilin phenotypes imply involvement of canonical Notch-signalling in segmentation of the spider Cupiennius salei

Arthropods, vertebrates, and annelids all have a segmented body. Our recent discovery of involvement of Notch-signalling in spider segmentation revived the discussion on the origin of segmented body plans and suggests the sharing of a common genetic program in a common ancestor. Here, we analysed the spider homologues of the Suppressor of Hairless and Presenilin genes, which encode components of the canonical Notch-pathway, to further explore the role of Notch-signalling in spider segmentation. RNAi silencing of two spider Suppressor of Hairless homologues and the spider Presenilin homologue causes severe segmentation phenotypes. The most prominent defect is the consistent breakdown of segmentation after the formation of three (Suppressor of Hairless) or five (Presenilin) opisthosomal segments. These phenotypes indicate that Notch-signalling during spider segmentation likely involves the canonical pathway via Presenilin and Suppressor of Hairless. Furthermore, it implies that Notch-signalling influences both the formation and patterning of the spider segments: it is required for the specification of the posterior segments and for proper specification of the segment boundaries. We argue that alternative, partly redundant, pathways might act in the formation of the anterior segments that are not active in the posterior segments. This suggests that at least some differences exist in the specification of anterior and posterior segments of the spider, a finding that may be valid for most short germ arthropods. Our data provide additional evidence for the similarities of Notch-signalling in spider segmentation and vertebrate somitogenesis and strengthen our previous notion that the formation of the segments in arthropods and vertebrates might have shared a genetic program in a common ancestor.