Temperature during pupal development affects hoverfly developmental time, adult life span, and wing length.

Hoverflies (Diptera, Syrphidae) are cosmopolitan, generalist flower visitors and among the most important pollinators after bees and bumblebees. The dronefly Eristalis tenax can be found in temperate and continental climates across the globe, often synanthropically. Eristalis tenax pupae of different generations and different climate zones are thus exposed to vastly different temperatures. In many insects, the ambient temperature during the pupal stage affects development, adult size, and survival; however, the effect of developmental temperature on these traits in hoverflies is comparatively poorly understood. We here reared E. tenax pupae at different temperatures, from 10°C to 25°C, and quantified the effect on adult hoverflies. We found that pupal rearing at 17°C appeared to be optimal, with high eclosion rates, longer wings, and increased adult longevity. Rearing temperatures above or below this optimum led to decreased eclosion rates, wing size, and adult survival. Similar thermal dependence has been observed in other insects. We found that rearing temperature had no significant effect on locomotor activity, coloration or weight, despite evidence of strong sexual dimorphism for each of these traits. Our findings are important as hoverflies are key pollinators, and understanding the effects of developmental temperature could potentially be useful for horticulture.

Descending neurons of the hoverfly respond to pursuits of artificial targets.

Many animals use motion vision information to control dynamic behaviors. Predatory animals, for example, show an exquisite ability to detect rapidly moving prey, followed by pursuit and capture. Such target detection is not only used by predators but is also important in conspecific interactions, such as for male hoverflies defending their territories against conspecific intruders. Visual target detection is believed to be subserved by specialized target-tuned neurons found in a range of species, including vertebrates and arthropods. However, how these target-tuned neurons respond to actual pursuit trajectories is currently not well understood. To redress this, we recorded extracellularly from target-selective descending neurons (TSDNs) in male Eristalis tenax hoverflies. We show that they have dorso-frontal receptive fields with a preferred direction up and away from the visual midline. We reconstructed visual flow fields as experienced during pursuits of artificial targets (black beads). We recorded TSDN responses to six reconstructed pursuits and found that each neuron responded consistently at remarkably specific time points but that these time points differed between neurons. We found that the observed spike probability was correlated with the spike probability predicted from each neuron's receptive field and size tuning. Interestingly, however, the overall response rate was low, with individual neurons responding to only a small part of each reconstructed pursuit. In contrast, the TSDN population responded to substantially larger proportions of the pursuits but with lower probability. This large variation between neurons could be useful if different neurons control different parts of the behavioral output.

Dual Receptive Fields Underlying Target and Wide-Field Motion Sensitivity in Looming-Sensitive Descending Neurons.

Responding rapidly to visual stimuli is fundamental for many animals. For example, predatory birds and insects alike have amazing target detection abilities, with incredibly short neural and behavioral delays, enabling efficient prey capture. Similarly, looming objects need to be rapidly avoided to ensure immediate survival, as these could represent approaching predators. Male Eristalis tenax hoverflies are nonpredatory, highly territorial insects that perform high-speed pursuits of conspecifics and other territorial intruders. During the initial stages of the pursuit, the retinal projection of the target is very small, but this grows to a larger object before physical interaction. Supporting such behaviors, E. tenax and other insects have both target-tuned and loom-sensitive neurons in the optic lobes and the descending pathways. We here show that these visual stimuli are not necessarily encoded in parallel. Indeed, we describe a class of descending neurons that respond to small targets, to looming and to wide-field stimuli. We show that these descending neurons have two distinct receptive fields where the dorsal receptive field is sensitive to the motion of small targets and the ventral receptive field responds to larger objects or wide-field stimuli. Our data suggest that the two receptive fields have different presynaptic input, where the inputs are not linearly summed. This novel and unique arrangement could support different behaviors, including obstacle avoidance, flower landing, and target pursuit or capture.

Vision: Flies move their eyes.

As you are reading this text, your eyes scan across the page, even if you keep your head perfectly still. New research reveals that flies can perform analogous retinal movements, despite their eyes being rigidly fixed to their heads.

Hoverfly (Eristalis tenax) pursuit of artificial targets.

The ability to visualize small moving objects is vital for the survival of many animals, as these could represent predators or prey. For example, predatory insects, including dragonflies, robber flies and killer flies, perform elegant, high-speed pursuits of both biological and artificial targets. Many non-predatory insects, including male hoverflies and blowflies, also pursue targets during territorial or courtship interactions. To date, most hoverfly pursuits have been studied outdoors. To investigate hoverfly (Eristalis tenax) pursuits under more controlled settings, we constructed an indoor arena that was large enough to encourage naturalistic behavior. We presented artificial beads of different sizes, moving at different speeds, and filmed pursuits with two cameras, allowing subsequent 3D reconstruction of the hoverfly and bead position as a function of time. We show that male E. tenax hoverflies are unlikely to use strict heuristic rules based on angular size or speed to determine when to start pursuit, at least in our indoor setting. We found that hoverflies pursued faster beads when the trajectory involved flying downwards towards the bead. Furthermore, we show that target pursuit behavior can be broken down into two stages. In the first stage, the hoverfly attempts to rapidly decreases the distance to the target by intercepting it at high speed. During the second stage, the hoverfly's forward speed is correlated with the speed of the bead, so that the hoverfly remains close, but without catching it. This may be similar to dragonfly shadowing behavior, previously coined 'motion camouflage'.

The impulse response of optic flow-sensitive descending neurons to roll m-sequences.

When animals move through the world, their own movements generate widefield optic flow across their eyes. In insects, such widefield motion is encoded by optic lobe neurons. These lobula plate tangential cells (LPTCs) synapse with optic flow-sensitive descending neurons, which in turn project to areas that control neck, wing and leg movements. As the descending neurons play a role in sensorimotor transformation, it is important to understand their spatio-temporal response properties. Recent work shows that a relatively fast and efficient way to quantify such response properties is to use m-sequences or other white noise techniques. Therefore, here we used m-sequences to quantify the impulse responses of optic flow-sensitive descending neurons in male Eristalis tenax hoverflies. We focused on roll impulse responses as hoverflies perform exquisite head roll stabilizing reflexes, and the descending neurons respond particularly well to roll. We found that the roll impulse responses were fast, peaking after 16.5-18.0 ms. This is similar to the impulse response time to peak (18.3 ms) to widefield horizontal motion recorded in hoverfly LPTCs. We found that the roll impulse response amplitude scaled with the size of the stimulus impulse, and that its shape could be affected by the addition of constant velocity roll or lift. For example, the roll impulse response became faster and stronger with the addition of excitatory stimuli, and vice versa. We also found that the roll impulse response had a long return to baseline, which was significantly and substantially reduced by the addition of either roll or lift.

Efference copies: Context matters when ignoring self-induced motion.

Across the animal kingdom, efference copies of neuronal motor commands are used to ensure our senses ignore stimuli generated by our own actions. New work shows that the underlying motivation for an action affects whether visual neurons are responsive to self-generated stimuli.

Facilitation of neural responses to targets moving against optic flow.

For the human observer, it can be difficult to follow the motion of small objects, especially when they move against background clutter. In contrast, insects efficiently do this, as evidenced by their ability to capture prey, pursue conspecifics, or defend territories, even in highly textured surrounds. We here recorded from target selective descending neurons (TSDNs), which likely subserve these impressive behaviors. To simulate the type of optic flow that would be generated by the pursuer's own movements through the world, we used the motion of a perspective corrected sparse dot field. We show that hoverfly TSDN responses to target motion are suppressed when such optic flow moves syn-directional to the target. Indeed, neural responses are strongly suppressed when targets move over either translational sideslip or rotational yaw. More strikingly, we show that TSDNs are facilitated by optic flow moving counterdirectional to the target, if the target moves horizontally. Furthermore, we show that a small, frontal spatial window of optic flow is enough to fully facilitate or suppress TSDN responses to target motion. We argue that such TSDN response facilitation could be beneficial in modulating corrective turns during target pursuit.

Karin Nordström.

Interview with Karin Nordström, who studies hoverfly motion vision at Flinders University.

Persistent Firing and Adaptation in Optic-Flow-Sensitive Descending Neurons.

A general principle of sensory systems is that they adapt to prolonged stimulation by reducing their response over time. Indeed, in many visual systems, including higher-order motion sensitive neurons in the fly optic lobes and the mammalian visual cortex, a reduction in neural activity following prolonged stimulation occurs. In contrast to this phenomenon, the response of the motor system controlling flight maneuvers persists following the offset of visual motion. It has been suggested that this gap is caused by a lingering calcium signal in the output synapses of fly optic lobe neurons. However, whether this directly affects the responses of the post-synaptic descending neurons, leading to the observed behavioral output, is not known. We use extracellular electrophysiology to record from optic-flow-sensitive descending neurons in response to prolonged wide-field stimulation. We find that, as opposed to most sensory and visual neurons, and in particular to the motion vision sensitive neurons in the brains of both flies and mammals, the descending neurons show little adaption during stimulus motion. In addition, we find that the optic-flow-sensitive descending neurons display persistent firing, or an after-effect, following the cessation of visual stimulation, consistent with the lingering calcium signal hypothesis. However, if the difference in after-effect is compensated for, subsequent presentation of stimuli in a test-adapt-test paradigm reveals adaptation to visual motion. Our results thus show a combination of adaptation and persistent firing in the neurons that project to the thoracic ganglia and thereby control behavioral output.

Insect Vision: Novel Mechanism for Contrast Constancy in Dim Light.

A general problem of sensory systems is how to simultaneously encode prevailing input as well as deviations from this baseline. A new study shows how the fly visual system has solved this by using parallel processing.

Visual motion sensitivity in descending neurons in the hoverfly.

Many animals use motion vision information to control dynamic behaviors. For example, flying insects must decide whether to pursue a prey or not, to avoid a predator, to maintain their current flight trajectory, or to land. The neural mechanisms underlying the computation of visual motion have been particularly well investigated in the fly optic lobes. However, the descending neurons, which connect the optic lobes with the motor command centers of the ventral nerve cord, remain less studied. To address this deficiency, we describe motion vision sensitive descending neurons in the hoverfly Eristalis tenax. We describe how the neurons can be identified based on their receptive field properties, and how they respond to moving targets, looming stimuli and to widefield optic flow. We discuss their similarities with previously published visual neurons, in the optic lobes and ventral nerve cord, and suggest that they can be classified as target-selective, looming sensitive and optic flow sensitive, based on these similarities. Our results highlight the importance of using several visual stimuli as the neurons can rarely be identified based on only one response characteristic. In addition, they provide an understanding of the neurophysiology of visual neurons that are likely to affect behavior.

Image statistics of the environment surrounding freely behaving hoverflies.

Natural scenes are not as random as they might appear, but are constrained in both space and time. The 2-dimensional spatial constraints can be described by quantifying the image statistics of photographs. Human observers perceive images with naturalistic image statistics as more pleasant to view, and both fly and vertebrate peripheral and higher order visual neurons are tuned to naturalistic image statistics. However, for a given animal, what is natural differs depending on the behavior, and even if we have a broad understanding of image statistics, we know less about the scenes relevant for particular behaviors. To mitigate this, we here investigate the image statistics surrounding Episyrphus balteatus hoverflies, where the males hover in sun shafts created by surrounding trees, producing a rich and dense background texture and also intricate shadow patterns on the ground. We quantified the image statistics of photographs of the ground and the surrounding panorama, as the ventral and lateral visual field is particularly important for visual flight control, and found differences in spatial statistics in photos where the hoverflies were hovering compared to where they were flying. Our results can, in the future, be used to create more naturalistic stimuli for experimenter-controlled experiments in the laboratory.

Acute sleep loss induces signs of visual discomfort in young men.

Acute sleep loss influences visual processes in humans, such as recognizing facial emotions. However, to the best of our knowledge, no study till date has examined whether acute sleep loss alters visual comfort when looking at images. One image statistic that can be used to investigate the level of visual comfort experienced under visual encoding is the slope of the amplitude spectrum, also referred to as the slope constant. The slope constant describes the spatial distribution of pixel intensities and deviations from the natural slope constant can induce visual discomfort. In the present counterbalanced crossover design study, 11 young men with normal or corrected-to-normal vision participated in two experimental conditions: one night of sleep loss and one night of sleep. In the morning after each intervention, subjects performed a computerized psychophysics task. Specifically, they were required to adjust the slope constant of images depicting natural landscapes and close-ups with a randomly chosen initial slope constant until they perceived each image as most natural looking. Subjects also rated the pleasantness of each selected image. Our analysis showed that following sleep loss, higher slope constants were perceived as most natural looking when viewing images of natural landscapes. Images with a higher slope constant are generally perceived as blurrier. The selected images were also rated as less pleasant after sleep loss. No such differences between the experimental conditions were noted for images of close-ups. The results suggest that sleep loss induces signs of visual discomfort in young men. Possible implications of these findings are discussed.

A scoping review of empirical evidence on the impacts of the DRG introduction in Germany and Switzerland.

CONTEXT: Germany and Switzerland have introduced diagnosis-related groups (DRGs) for hospital reimbursement. This scoping review aims to evaluate if empirical evidence exists on the effect of the DRG introduction. METHODS: Medline via PubMed, Embase (Elsevier), CINAHL, PsychINFO, and Psyndex were systematically screened for studies from 2003 onwards using keywords-DRG, prospective payment system, and lump sum-in English, German, and French. Abstracts were screened for alignment with our inclusion criteria and classified as editorial/commentary, review, or empirical study. The full-text extraction included data on country, study design, collected data, study population, specialty, comparison group, and outcome measures. RESULTS: Our literature search yielded 1944 references, of which 1405 references were included in the abstract screening after removal of duplicates. 135 articles were relevant to DRG, including 94 editorials/comments/reviews and 41 empirical articles from 36 different samples. The most frequently used outcome parameters were length of stay (12), reimbursement/cost (9), and case numbers (9). CONCLUSIONS: Only a minority of identified articles (30.4%; 41 of 135) presented empirical data. This indicates that discussion on the topic is not totally evidence-based. The only common trend was a decrease in length of stay.

Integration of Small- and Wide-Field Visual Features in Target-Selective Descending Neurons of both Predatory and Nonpredatory Dipterans.

For many animals, target motion carries high ecological significance as this may be generated by a predator, prey, or potential mate. Indeed, animals whose survival depends on early target detection are often equipped with a sharply tuned visual system, yielding robust performance in challenging conditions. For example, many fast-flying insects use visual cues for identifying targets, such as prey (e.g., predatory dragonflies and robberflies) or conspecifics (e.g., nonpredatory hoverflies), and can often do so against self-generated background optic flow. Supporting these behaviors, the optic lobes of insects that pursue targets harbor neurons that respond robustly to the motion of small moving objects, even when displayed against syn-directional background clutter. However, in diptera, the encoding of target information by the descending neurons, which are more directly involved in generating the behavioral output, has received less attention. We characterized target-selective neurons by recording in the ventral nerve cord of male and female predatory Holcocephala fusca robberflies and of male nonpredatory Eristalis tenax hoverflies. We show that both species have dipteran target-selective descending neurons that only respond to target motion if the background is stationary or moving slowly, moves in the opposite direction, or has un-naturalistic spatial characteristics. The response to the target is suppressed when background and target move at similar velocities, which is strikingly different to the response of target neurons in the optic lobes. As the neurons we recorded from are premotor, our findings affect our interpretation of the neurophysiology underlying target-tracking behaviors.SIGNIFICANCE STATEMENT Many animals use sensory cues to detect moving targets that may represent predators, prey, or conspecifics. For example, birds of prey show superb sensitivity to the motion of small prey, and intercept these at high speeds. In a similar manner, predatory insects visually track moving prey, often against cluttered backgrounds. Accompanying this behavior, the brains of insects that pursue targets contain neurons that respond exclusively to target motion. We here show that dipteran insects also have target-selective descending neurons in the part of their nervous system that corresponds to the vertebrate spinal cord. Surprisingly, and in contrast to the neurons in the brain, these premotor neurons are inhibited by background patterns moving in the same direction as the target.

Rearing and Long-Term Maintenance of Eristalis tenax Hoverflies for Research Studies.

With an estimated 6000 species worldwide, hoverflies are ecologically important as alternative pollinators to domesticated honeybees. However, they are also a useful scientific model to study motion vision and flight dynamics in a controlled laboratory setting. As the larvae develop in organically polluted water, they are useful models for investigating investment in microbial immunity. While large scale commercial breeding for agriculture already occurs, there are no standardized protocols for maintaining captive populations for scientific studies. This is important as commercial captive breeding programs focusing on mass output during peak pollination periods may fail to provide a population that is consistent, stable and robust throughout the year, as is often needed for other research purposes. Therefore, a method to establish, maintain and refresh a captive research population is required. Here, we describe the utilization of an artificial hibernation cycle, in addition to the nutritional and housing requirements, for long term maintenance of Eristalis tenax. Using these methods, we have significantly increased the health and longevity of captive populations of E. tenax compared to previous reports. We furthermore discuss small scale rearing methods and options for optimizing yields and manipulating population demographics.

Visual approach computation in feeding hoverflies.

On warm sunny days, female hoverflies are often observed feeding from a wide range of wild and cultivated flowers. In doing so, hoverflies serve a vital role as alternative pollinators, and are suggested to be the most important pollinators after bees and bumblebees. Unless the flower hoverflies are feeding from is large, they do not readily share the space with other insects, but instead opt to leave if another insect approaches. We used high-speed videography followed by 3D reconstruction of flight trajectories to quantify how female Eristalis hoverflies respond to approaching bees, wasps and two different hoverfly species. We found that, in 94% of the interactions, the occupant female left the flower when approached by another insect. We found that compared with spontaneous take-offs, the occupant hoverfly's escape response was performed at ∼3 times higher speed (spontaneous take-off at 0.2±0.05 m s-1 compared with 0.55±0.08 m s-1 when approached by another Eristalis). The hoverflies tended to take off upward and forward, while taking the incomer's approach angle into account. Intriguingly, we found that, when approached by wasps, the occupant Eristalis took off at a higher speed and when the wasp was further away. This suggests that feeding hoverflies may be able to distinguish these predators, demanding impressive visual capabilities. Our results, including quantification of the visual information available before occupant take-off, provide important insight into how freely behaving hoverflies perform escape responses from competitors and predators (e.g. wasps) in the wild.

Caring for elder patients: Mutual vulnerabilities in professional ethics.

BACKGROUND:: Neglect and abuse of elders in care institutions is a recurring issue in the media. Elders in care institutions are vulnerable due to their physical, cognitive, and verbal limitations. Such vulnerabilities may make them more susceptible to mistreatment by caregivers on whom they are heavily dependent. OBJECTIVES:: The goal was to understand caregivers' concerns about ensuring correct and proper treatment, as well as their experiences with neglect and abuse of older patients. This article examines resources and challenges of professional ethics within the care setting. RESEARCH DESIGN:: A study was conducted to explore the quality of care provided to older patients in nursing homes, geriatrics institutions, and ambulant care in the northwest region of Switzerland. PARTICIPANTS AND RESEARCH CONTEXT:: A total of 23 semi-structured interviews were conducted with nursing staff of varying experience levels. ETHICAL CONSIDERATIONS:: Ethical approval was granted by the competent regional ethics commission, Ethikkomission Nordwest-und Zentralschweiz EKNZ [Ethics Commission Northwest and Central Switzerland] (2014-015). FINDINGS:: Three themes emerged from our data analysis: professional identity, professional context, and professional relationships. Our findings indicate mutual vulnerabilities within these three themes, characterizing the interactions between nursing staff and older patients. Study participants believe that incidences of error, neglect, and abuse are consequences of their own vulnerability since they are not able to meet the demands of an overstraining work situation. DISCUSSION:: Different aspects of this mutual vulnerability are described and critically discussed as challenges for professional ethics. CONCLUSION:: Early education, continuous training as well as better management and response from the institution are necessary to maintain professionalism while handling mutual vulnerabilities.

Image statistics and their processing in insect vision.

Natural scenes may appear random, but are not only constrained in space and time, but also show strong spatial and temporal correlations. Spatial constraints and correlations can be described by quantifying image statistics, which include intuitive measures such as contrast, color and luminance, but also parameters that need some type of transformation of the image. In this review we will discuss some common tools used to quantify spatial and temporal parameters of naturalistic visual input, and how these tools have been used to inform us about visual processing in insects. In particular, we will review findings that would not have been possible using conventional, experimenter defined stimuli.