Confronting physiology: how do infected flies die?

Fruit fly immunology is on the verge of an exciting new path. The fruit fly has served as a strong model for innate immune responses; the field is now expanding to use the fruit fly to study pathogenesis. We argue here that, to understand pathogenesis in the fly, we need to understand pathology - and to understand pathology, we need to confront physiology with molecular tools. When flies are infected with a pathogen, they get sick. We group the events following infection into three categories: innate immune responses (defence mechanisms by which the fly attempts to kill or neutralize the microbe, some of which can themselves cause harm to the fly); microbial virulence (mechanisms by which the microbe evades the immune response); and host pathology (physiologies adversely affected by either the immune response or microbial virulence). We divide this review into sections mirroring these categories. The molecular study of infection in the fruit fly has focused on the first category, has begun to explore the second, and has yet to tap the full potential of the fly regarding the third.     

Sexual selection: do flies lie with asymmetric legs?

A newly described species of empidid or 'dance fly' shows a bizarre polymorphism in their forelegs, which presumably serve as a mating lure. This trait may have evolved by frequency-dependent deceptive male signalling.     

The myosin filament superlattice in the flight muscles of flies: A-band lattice optimisation for stretch-activation?

Low-angle X-ray diffraction patterns from relaxed fruitfly (Drosophila) flight muscle recorded on the BioCat beamline at the Argonne Advanced Photon Source (APS) show many features similar to such patterns from the "classic" insect flight muscle in Lethocerus, the giant water bug, but there is a characteristically different pattern of sampling of the myosin filament layer-lines, which indicates the presence of a superlattice of myosin filaments in the Drosophila A-band. We show from analysis of the structure factor for this lattice that the sampling pattern is exactly as expected if adjacent four-stranded myosin filaments, of repeat 116 nm, are axially shifted in the hexagonal A-band lattice by one-third of the 14.5 nm axial spacing between crowns of myosin heads. In addition, electron micrographs of Drosophila and other flies (e.g. the house fly (Musca) and the flesh fly (Sarcophaga)) combined with image processing confirm that the same A-band superlattice occurs in all of these flies; it may be a general property of the Diptera. The different A-band organisation in flies compared with Lethocerus, which operates at a much lower wing beat frequency (approximately 30 Hz) and requires a warm-up period, may be a way of optimising the myosin and actin filament geometry needed both for stretch activation at the higher wing beat frequencies (50 Hz to 1000 Hz) of flies and their need for a rapid escape response.     

The erroneous courtship hypothesis: do insects really engage in aerial wars of attrition?

Males of various flying insects perform conspicuous aerial interactions around their mating stations. The broadly accepted interpretation of their aerial interaction is a war of attrition, where two contestants perform costly displays, and the one that reaches its cost threshold earlier gives up. The implicit but important requirement in this model is that some forces that match the intensity of display of the two contestants are necessary, and failure to enforce matching allows foul contestants that delay or stop their display to avoid paying contest costs. In addition, wars of attrition require flying insects to distinguish the sex of flying conspecifics because their aerial interactions begin when intruders fly into the territory. We investigated past research on the behaviour of odonates and butterflies aiming to clarify whether the two prerequisites of wars of attrition are fulfilled: (1) contestants can inflict substantial costs on nondisplaying opponents and (2) contestants can discriminate the sex of flying conspecifics. In odonates, we found an abundance of evidence suggesting that contests involve physical attack and that the ability of sexual discrimination is sufficient. Therefore, wars of attrition may occur in territorial odonates. In butterflies, however, we could not find any evidence that the two prerequisites are filled. The aerial interactions of butterflies are better interpreted as courtship between sexually active males (the erroneous courtship hypothesis). Based on these results, we discuss future directions of research on the aerial contests of flying insects.     

The energy cost of flight: do small bats fly more cheaply than birds?

Flapping flight is one of the most expensive activities in terms of metabolic cost and this cost has previously been considered equal for the two extant vertebrate groups which evolved flapping flight. Owing to the difficulty of obtaining accurate measurements without disturbing flight performance, current estimates of flight cost within the group of small birds and bats differ by more than a factor of five for given body masses. To minimize the potential problem that flight behaviour may be affected by the measurements, we developed an indirect method of measuring flight energy expenditure based on time budget analysis in which small nectar-feeding bats (Glossophaginae) could continue their natural rhythm of flying and resting entirely undisturbed. Estimates of metabolic flight power based on 172 24-h time and energy budget measurements were obtained for nine individual bats from six species (mass 7–28 g). Metabolic flight power (P_F) of small bats was found to increase with body mass following the relation P_F = 50.2 M^0.771 (r² = 0.96, n = 13, P_F in W, M in kg). This is about 20–25% below the majority of current predictions of metabolic flight cost for small birds. Thus, either the flight cost of small birds is significantly lower than has previously been thought or, contrary to current opinion, small bats require less energy to fly than birds. Accepted: 29 September 1997     

Free centrosomes: Where do they all come from?

Centrosomes act as major microtubule-organizing centers in most cell types. Their functions in interphase and mitosis are usually facilitated by their association with the nucleus. This may be particularly true in very large cells. Several papers report free centrosomes in syncytial Drosophila embryos. However, this phenotype often remains little explored. Yet, free centrosomes can occur by multiple mechanisms, including functional defects of the mitotic spindle, detachment of centrosomes from the nuclear envelope, centrosome inactivation upon DNA damage, and de novo centrosome genesis. Deciphering the cellular mechanism leading to free centrosomes upon a given perturbation such as a mutation or injection of a drug, can provide valuable clues regarding the nature of the molecular pathway affected. To this end, genetic and cytological tests, as well as time-lapse imaging are available. These studies can inform on the biology of centrosomes, cell cycle regulation and cytoskeletal dynamics. Here we briefly discuss what to make of free centrosomes in the fly embryo.     

Info-gap robust-satisficing model of foraging behavior: do foragers optimize or satisfice?

In this note we compare two mathematical models of foraging that reflect two competing theories of animal behavior: optimizing and robust satisficing. The optimal-foraging model is based on the marginal value theorem (MVT). The robust-satisficing model developed here is an application of info-gap decision theory. The info-gap robust-satisficing model relates to the same circumstances described by the MVT. We show how these two alternatives translate into specific predictions that at some points are quite disparate. We test these alternative predictions against available data collected in numerous field studies with a large number of species from diverse taxonomic groups. We show that a large majority of studies appear to support the robust-satisficing model and reject the optimal-foraging model.     

How do trabeculae affect the calculation of structural properties of bone?

One difficulty that arises in an analysis of the cross-sectional properties of bone is whether to include cancellous bone in the analysis. The purpose of this paper is to determine how different amounts of cancellous bone affect the measurement of structural properties of bone cross-sections. Thirty-two tibial and femoral cross-sections were chosen at random from a series of cross-sectioned nonhuman primate bones. Geometrical properties were calculated for the cross-sections, and torsional and bending stress analyses were performed. The results suggest that the effect of including cancellous bone in the analysis is closely related to the amount of bone, where it lies within the cross-section, and the type of analysis performed. Including cancellous bone in calculations of structural properties of bone cross-sections may cause the strength and stiffness of the bone to be exaggerated.     

Distribution of recreational boating across lakes: do landscape variables affect recreational use?

1. Because people impact lake ecosystems, it is important to consider factors influencing the human use of freshwater resources. We investigated the influence of the landscape position, as well as lake area, recreational facilities, and distance to highways and urban centres, on lake use by boaters in the Northern Highland Lake District of Wisconsin, U.S.A.
2. In aerial surveys of ninety-nine randomly selected lakes, we did not see boats on over half of the lakes. Of the lakes with boats, we found a strong correlation between the number of boats and lake area. Recreational boats tended to be found on large, accessible lakes with good boating facilities. Boats were not seen on small, stained lakes with few recreational facilities.
3. Regression models showed that lake size and landscape position explained 63% of the variability in the average number of boats per lake and landscape position explained 24% of the variability in boat density on all ninety-nine lakes. Social variables representing the quality of boating facilities and the perception of good fishing explained 70% of the variability in number of boats per lake and 54% of the variation in boat density on all lakes. A combined model using both physical and social variables increased the explanatory power for both number and density. Lake use by boaters was correlated with landscape position, the quality of fishing and the availability of recreational facilities. When the analysis was restricted to the forty-six lakes where boats were found, only the availability of recreational facilities proved a significant predictor of boat density.
4. Our results suggest that lake choice by recreational boaters may be best predicted by a combination of the location of a lake in a regional hydrologic landscape, and considerations of available facilities and perceptions regarding fishing.     

Environmental fluctuations: How do they affect the topography of the adaptive landscape?

A Phenotypic Adaptive Landscape is defined with fitness as the ordinate, and longevity of the juvenile phase and duration of disturbances to the adult phase as the horizontal axes. The effect of local environmental perturbations on the landscape’s shape is studied using a semistochastic population model. In this model the intrinsic population dynamics takes the form of a differential-delay nonlinear equation and the environmental disturbance appears as a multiplicativetelegraphic noise. We demonstrate that the landscape has no single characteristic scale. Rather, it shows adaptive peaks corresponding to an integer relation between the biological and the environmental periodicities. Since the system is constrained by a finite time and a finite physiological range, the landscape may have different topographies for different local environmental regimes. A very simple fully deterministic model is presented, predicting landscapes that are similar to those obtained by the semistochastic model. Application to life history strategies are discussed.     

The influence of snow on lynx and coyote movements: does morphology affect behavior?

Summary We studied sympatric lynx (Lynx canadensis) and coyotes (Canis latrans) to assess how morphological disadvantages to locomotion over snow affected movement patterns. Both species are of similar size and mass, but the feet of lynx are much larger, and coyotes were found to have 4.1–8.8 times the foot-load (ratio of body mass to foot area) of lynx. This resulted in greater mean sinking depths of coyote limbs, although the magnitude of the difference was less than that in foot-load. Coyotes exhibited stronger use of behavioral patterns that reduced negative effects of snow on movements. Coyotes were most abundant at low elevations where snow was shallow, whereas lynx were mostly at higher elevations. Coyotes also used areas at both elevations where snow was shallower than average, while lynx used areas where snow was deeper. further, both species used travel routes where snow was shallower than it was near the track. Coyotes traveled on harder snow and used trails more frequently, thereby tending to reduce sinking depths to those similar to lynx. The behavioral repertoire of coyotes reduced the morphological advantage of large feet possessed by lynx; however, overall sinking depths were still greater in coyotes. Snowshoe hares (Lepus americanus) were the main prey of both species, and their foot-load was less than that of either predator. Hare kills by coyotes occurred after fewer bounds than did those by lynx, and the large difference between foot-loads of both species of predators may have forced coyotes to ambush rather than chase hares, as did lynx.     

Butterfly contests and flight physiology: why do older males fight harder?

The males of many butterfly species compete for territoriesvia conspicuous aerial wars of attrition, in which the determinantsof persistence ability are largely unclear. Flight performancefeatures, such as stamina, acceleration, and maneuverability,are often assumed to be important in this context, yet thereis no direct evidence by which to evaluate these possibilities.Recent research has indicated that competitive ability increaseswith age in a notably territorial species, Hypolimnas bolina,which could arise from lifetime morphological or physiologicalchanges that directly affect flight performance. I evaluatedthis hypothesis by investigating how size-independent variancein body composition, energy stores, flight muscle ratio (FMR),and wing condition change with age in this species. Males infive age categories (spanning the functional life span of territorialindividuals) were sampled from encounter sites in tropicalAustralia. Analysis of body composition with respect to anestimate of eclosion mass (forewing length) indicated thattotal body mass, abdomen mass, and wing area decrease throughoutan individual's lifetime, but thorax mass remains unchanged.Wing loading (the ratio of wing area to body mass) is lowestin intermediately aged individuals, but FMR and energetic statusremain largely similar regardless of age. On average, therefore,the energetic cost of sustained flight should first decrease,then increase, with age in a male H. bolina (of standardizedbody size), while available energy reserves decline slightly. Acceleration and maneuverability should remain relatively constant.These results, coupled with the fact that body size is unrelatedto contest success in this territorial butterfly, fail to supportthe idea that age-related competitive ability is mediated simplyby energetics or flight performance.     

How do cytokinins affect the cell?

The lecture presents modern knowledge of the mechanisms of cytokinin perception and signal transduction to the genes of primary and secondary responses. It also demonstrates the relations between the rapid cytokinin-induced processes and cytokinin-induced physiological effects. The characteristics of the cytokinin regulatory system and its role in the control of plant growth and development are discussed.     

How do synonymous mutations affect fitness?

While it has often been assumed that, in humans, synonymous mutations would have no effect on fitness, let alone cause disease, this position has been questioned over the last decade. There is now considerable evidence that such mutations can, for example, disrupt splicing and interfere with miRNA binding. Two recent publications suggest involvement of additional mechanisms: modification of protein abundance most probably mediated by alteration in mRNA stability and modification of protein structure and activity, probably mediated by induction of translational pausing. These case histories put a further nail into the coffin of the assumption that synonymous mutations must be neutral.     

How do toxicants affect epidemiological dynamics?

Populations are formed of their constituent interacting individuals, each with their own respective within‐host biological processes. Infection not only spreads within the host organism but also spreads between individuals. Here we propose and study a multilevel model which links the within‐host statuses of immunity and parasite density to population epidemiology under sublethal and lethal toxicant exposure. We analyse this nested model in order to better understand how toxicants impact the spread of disease within populations. We demonstrate that outbreak of infection within a population is completely determined by the level of toxicant exposure, and that it is maximised by intermediate toxicant dosage. We classify the population epidemiology into five phases of increasing toxicant exposure and calculate the conditions under which disease will spread, showing that there exists a threshold toxicant level under which epidemics will not occur. In general, higher toxicant load results in either extinction of the population or outbreak of infection. The within‐host statuses of the individual host also determine the outcome of the epidemic at the population level. We discuss applications of our model in the context of environmental epidemiology, predicting that increased exposure to toxicants could result in greater risk of epidemics within ecological systems. We predict that reducing sublethal toxicant exposure below our predicted safe threshold could contribute to controlling population level disease and infection.     

Climb and flight speeds of shorebirds embarking on an intercontinental flight; do they achieve the predicted optimal behaviour?

Most Arctic-breeding waders wintering in West Africa cover the first 4000 km of their northward journey in spring by a single flight to western Europe. We examined the extent to which waders economize their flight behaviour during departure by comparing climb rates and forward flight speeds with predictions based on flight mechanic theory and the relevant morphological measurements made of birds collected on the site. With an optical range finder, we followed 98 wader flocks on their departure from Banc d'Arguin in Mauritania. We also measured wind speed and direction at different altitudes by tracking helium-filled balloons and thus were able to deduce airspeeds from groundspeeds of the departing flocks. Of the nine species examined, six showed the predicted negative relationship between climb rate and airspeed, although only one was statistically significant. By normalizing the data, we found a statistically significant negative correlation across all species. Although 17% of the observed climb rates were greater than the predicted theoretical maximum, the average observed climb rate was lower than the predicted optimum and the average observed airspeed was higher. The absolute deviations of climb rates from theory may have been because of the existence of pockets of rising and sinking air at the boundary of desert and ocean. That the absolute deviations in average climb rate and airspeed followed the predicted negative relationship is in accordance with the current theory of flight mechanics.     

Ecophysiology of cognition: How do environmentally induced changes in physiology affect cognitive performance?

Cognitive performance is based on brain functions, which have energetic demands and are modulated by physiological parameters such as metabolic hormones. As both environmental demands and environmental energy availability change seasonally, we propose that cognitive performance in free‐living animals might also change seasonally due to phenotypic plasticity. This is part of an emerging research field, the ‘ecophysiology of cognition’: environmentally induced changes in physiological traits, such as blood glucose and hormone levels, are predicted to influence cognitive performance in free‐living animals. Energy availability for the brain might change, and as such cognition, with changing energetic demands (e.g. reproduction) and changes of energy availability in the environment (e.g. winter, drought). Individuals spending more energy than they can currently obtain from their environment (allostatic overload type I) are expected to trade off energy investment between cognition and other life‐sustaining processes or even reproduction. Environmental changes reducing energy availability might thus impair cognition. However, selection pressures such as predation risk, mate choice or social demands may act on the trade‐off between energy saving and cognition. We assume that different environmental conditions can lead to three different trade‐off outcomes: cognitive impairment, resilience or enhancement. Currently we cannot understand these trade‐offs, because we lack information about changes in cognitive performance due to seasonal changes in energy availability and both the resulting changes in homeostasis (for example, blood glucose levels) and the associated changes in the mechanisms of allostasis (for example, hormone levels). Additionally, so far we know little about the fitness consequences of individual variation in cognitive performance. General cognitive abilities, such as attention and associative learning, might be more important in determining fitness than complex and specialized cognitive abilities, and easier to use for comparative study in a large number of species. We propose to study seasonal changes in cognitive performance depending on energy availability in populations facing different predation risks, and the resulting fitness consequences.     

Control of tsetse flies and trypanosomiasis: Myth or reality?

The African trypanosomiasis are among Africa's most devastating diseases. The human disease, sleeping sickness, and the animal disease, nagana, are caused by trypanosomes, protozoan parasites transmitted by tsetse flies, Glossina spp. Attempts have been made to control tsetse and trypanosomiasis for over 70 years, supported by ever-increasing amounts of foreign aid. Although progress has been made in the control of sleeping sickness, this disease still persists in many countries. Nogono excludes cattle from many of the potentially most productive areas of Africa and is a major constraint on economic development. In this paper, Robert Dransfield, Brian Williams and Robert Brightwell review the control of tsetse and trypanosomiasis in the light of recent progress in our understanding of tsetse population dynamics, with special reference to the experience gained in tsetse control on a Maasai ranch at Ngurumon in the Rift Valley of Kenya, and make suggestions for the management and funding of future control programmes in relation to rural development.     

Social behavior: how do fish find their shoal mate?

Fish form social aggregations called shoals which often consist of fish with similar morphologies. Experiments using zebrafish pigment variants demonstrate that fish can select shoal mates solely on the basis of their color patterns, and that early experience plays a key role in determining these shoaling preferences.