Egomotion estimation with optic flow and air velocity sensors

We develop a method that allows a flyer to estimate its own motion (egomotion), the wind velocity, ground slope, and flight height using only inputs from onboard optic flow and air velocity sensors. Our artificial algorithm demonstrates how it could be possible for flying insects to determine their absolute egomotion using their available sensors, namely their eyes and wind sensitive hairs and antennae. Although many behaviors can be performed by only knowing the direction of travel, behavioral experiments indicate that odor tracking insects are able to estimate the wind direction and control their absolute egomotion (i.e., groundspeed). The egomotion estimation method that we have developed, which we call the opto-aeronautic algorithm, is tested in a variety of wind and ground slope conditions using a video recorded flight of a moth tracking a pheromone plume. Over all test cases that we examined, the algorithm achieved a mean absolute error in height of 7% or less. Furthermore, our algorithm is suitable for the navigation of aerial vehicles in environments where signals from the Global Positioning System are unavailable.     

Nocturnal insects use optic flow for flight control

To avoid collisions when navigating through cluttered environments, flying insects must control their flight so that their sensory systems have time to detect obstacles and avoid them. To do this, day-active insects rely primarily on the pattern of apparent motion generated on the retina during flight (optic flow). However, many flying insects are active at night, when obtaining reliable visual information for flight control presents much more of a challenge. To assess whether nocturnal flying insects also rely on optic flow cues to control flight in dim light, we recorded flights of the nocturnal neotropical sweat bee, Megalopta genalis, flying along an experimental tunnel when: (i) the visual texture on each wall generated strong horizontal (front-to-back) optic flow cues, (ii) the texture on only one wall generated these cues, and (iii) horizontal optic flow cues were removed from both walls. We find that Megalopta increase their groundspeed when horizontal motion cues in the tunnel are reduced (conditions (ii) and (iii)). However, differences in the amount of horizontal optic flow on each wall of the tunnel (condition (ii)) do not affect the centred position of the bee within the flight tunnel. To better understand the behavioural response of Megalopta, we repeated the experiments on day-active bumble-bees (Bombus terrestris). Overall, our findings demonstrate that despite the limitations imposed by dim light, Megalopta-like their day-active relatives-rely heavily on vision to control flight, but that they use visual cues in a different manner from diurnal insects.     

Visual perception of surface curvature. The spin variation and its physiological implications

In the context of the models of structure from motion visual processing, we propose that the optic-flow field is a source of information for the perception of the curvature of a smooth surface in motion. In particular, it is shown how the spin variation (SV), a second spatial derivative of the retinal velocity field, is mathematically related to the curvature of the surface. Under the hypothesis that the visual system relies on SV to analyse the structure of a moving surface, a neural scheme for SV detection is proposed and psychophysical predictions are developed. Results obtained on artificial images show that the SV scheme presents a rather weak sensitivity to noise in conditions of low image velocity.     

Conserving insect assemblages in urban landscapes: accounting for species-specific responses and imperfect detection

Understanding how global environmental change impacts insect biodiversity is central to the core principals of conservation biology. To preserve the ecosystem services provided by insects in cities, it is crucial to understand how insect species are influenced by the degree of urbanization of the surrounding landscape. Using a hierarchical occupancy–detection model, we estimated the effect of urbanization on heteropteran bug species richness and occupancy, an approach that concurrently accounts for species-specific responses and imperfect detection. We found that species richness decreased along a gradient of increasing urbanization. This trend corresponded well with species-specific trends, as approximately two-thirds of all herbivores and predatory species experienced a strong mean negative response to urbanization. These results indicate that many species are potentially at risk of local extinction as cities grow and expand in the future. A second group of species, however, showed a weak mean negative response, indicating that they are ubiquitous urban species that thrive regardless of the surrounding degree of urban disturbance. Our research suggests that as cities develop, many of the species that are currently present will become less likely to occur, and therefore assemblages in the future are likely to become more simplified. In order to preserve or increase insect biological diversity in cities, it is critical to understand how individual species are influenced by urbanization. Our finding that insects display species-specific responses to urbanization has important repercussions for decision makers charged with preserving and improving urban biodiversity and the deliverance of ecosystem services in cities.     

Control of Reproductive Floral Organ Identity Specification in Arabidopsis by the C Function Regulator AGAMOUS

The floral organ identity factor AGAMOUS (AG) is a key regulator of Arabidopsis thaliana flower development, where it is involved in the formation of the reproductive floral organs as well as in the control of meristem determinacy. To obtain insights into how AG specifies organ fate, we determined the genes and processes acting downstream of this C function regulator during early flower development and distinguished between direct and indirect effects. To this end, we combined genome-wide localization studies, gene perturbation experiments, and computational analyses. Our results demonstrate that AG controls flower development to a large extent by controlling the expression of other genes with regulatory functions, which are involved in mediating a plethora of different developmental processes. One aspect of this function is the suppression of the leaf development program in emerging floral primordia. Using trichome initiation as an example, we demonstrate that AG inhibits an important aspect of leaf development through the direct control of key regulatory genes. A comparison of the gene expression programs controlled by AG and the B function regulators APETALA3 and PISTILLATA, respectively, showed that while they control many developmental processes in conjunction, they also have marked antagonistic, as well as independent activities.     

Mechanisms underlying insect freeze tolerance

Freeze tolerance – the ability to survive internal ice formation – has evolved repeatedly in insects, facilitating survival in environments with low temperatures and/or high risk of freezing. Surviving internal ice formation poses several challenges because freezing can cause cellular dehydration and mechanical damage, and restricts the opportunity to metabolise and respond to environmental challenges. While freeze‐tolerant insects accumulate many potentially protective molecules, there is no apparent ‘magic bullet’ – a molecule or class of molecules that appears to be necessary or sufficient to support this cold‐tolerance strategy. In addition, the mechanisms underlying freeze tolerance have been minimally explored. Herein, we frame freeze tolerance as the ability to survive a process: freeze‐tolerant insects must withstand the challenges associated with cooling (low temperatures), freezing (internal ice formation), and thawing. To do so, we hypothesise that freeze‐tolerant insects control the quality and quantity of ice, prevent or repair damage to cells and macromolecules, manage biochemical processes while frozen/thawing, and restore physiological processes post‐thaw. Many of the molecules that can facilitate freeze tolerance are also accumulated by other cold‐ and desiccation‐tolerant insects. We suggest that, when freezing offered a physiological advantage, freeze tolerance evolved in insects that were already adapted to low temperatures or desiccation, or in insects that could withstand small amounts of internal ice formation. Although freeze tolerance is a complex cold‐tolerance strategy that has evolved multiple times, we suggest that a process‐focused approach (in combination with appropriate techniques and model organisms) will facilitate hypothesis‐driven research to understand better how insects survive internal ice formation.     

Honeybees change their height to restore their optic flow

To further elucidate the mechanisms underlying insects’ height and speed control, we trained outdoor honeybees to fly along a high-roofed tunnel, part of which was equipped with a moving floor. Honeybees followed the stationary part of the floor at a given height. On encountering the moving part of the floor, which moved in the same direction as their flight, honeybees descended and flew at a lower height, thus gradually restoring their ventral optic flow (OF) to a similar value to that they had percieved when flying over the stationary part of the floor. This was therefore achieved not by increasing their airspeed, but by lowering their height of flight. These results can be accounted for by a control system called an optic flow regulator, as proposed in previous studies. This visuo-motor control scheme explains how honeybees can navigate safely along tunnels on the sole basis of OF measurements, without any need to measure either their speed or the clearance from the surrounding walls.     

Visual flight control in naturalistic and artificial environments

Although the visual flight control strategies of flying insects have evolved to cope with the complexity of the natural world, studies investigating this behaviour have typically been performed indoors using simplified two-dimensional artificial visual stimuli. How well do the results from these studies reflect the natural behaviour of flying insects considering the radical differences in contrast, spatial composition, colour and dimensionality between these visual environments? Here, we aim to answer this question by investigating the effect of three- and two-dimensional naturalistic and artificial scenes on bumblebee flight control in an outdoor setting and compare the results with those of similar experiments performed in an indoor setting. In particular, we focus on investigating the effect of axial (front-to-back) visual motion cues on ground speed and centring behaviour. Our results suggest that, in general, ground speed control and centring behaviour in bumblebees is not affected by whether the visual scene is two- or three dimensional, naturalistic or artificial, or whether the experiment is conducted indoors or outdoors. The only effect that we observe between naturalistic and artificial scenes on flight control is that when the visual scene is three-dimensional and the visual information on the floor is minimised, bumblebees fly further from the midline of the tunnel. The findings presented here have implications not only for understanding the mechanisms of visual flight control in bumblebees, but also for the results of past and future investigations into visually guided flight control in other insects.     

Flight of Heliothis virescens males in the field in response to sex pheromone

Abstract. The behaviour of Heliothis virescens males flying upwind in the field in a sex pheromone plume was videorecorded and analysed. Males flew faster and straighter, with less counterturning, and heading more directly into the wind when they were 9-11m away from the odour source than when they were 1–3 m away. Regardless of their distance from the source or the windspeed, they maintained an average groundspeed of c. 200 cm s^_1, except when they arrived within 1 m of the source, when their groundspeed slowed significantly. Two or more males flying in the plume at the same instant often exhibited either extremely straight and directly upwind tracks or else zigzagging tracks with significant counterturning (as did males flying through the field of view of the cameras at slighdy different times). The males' position, either in the centre of the plume's axis or along one side, might explain these differences in track straightness, which previous studies with H.virescens have shown to be caused by higher frequencies of contact with plume filaments. When a significant shift in wind direction occurred, males tended to make an initial movement in the direction of the shift, perhaps due to latencies of response in both the olfactory and visual systems associated with flying into clean air. The males' behaviour in the field overall was similar to that observed in the wind tunnel, except that their airspeeds and groundspeeds were significantly higher than those observed in the laboratory. The fact that they flew faster in the field can be explained both by the significandy higher windspeeds that males need to compensate for in the field to attain a preferred velocity of image motion, as well as by a higher height of flight over the ground in die field causing a slower apparent motion of images at a given groundspeed compared with the laboratory.     

The control of optic flow during learning flights

Hymenopteran insects perform systematic learning flights on departure from their nest, during which they acquire a visual representation of the nest environment. They back away from and pivot around the nest in a series of arcs while turning to view it in their fronto-lateral visual field. During the initial stages of the flights, turning rate and arc velocity relative to the nest are roughly constant at 100–200° s⁻¹ and are independent of distance, since the insects increase their flight speed as they back away from the pivoting centre. In this paper I analyse how solitary wasps control their flight by having them perform learning flights inside a rotating striped drum. The wasps' turning velocity is under visual control. When the insects fly inside a drum that rotates around the nest as a centre, their average turning rate is faster than normal when they fly an arc into the direction of drum rotation and slower when they fly in the opposite direction. The average slip speed they experience lies within 100–200° s⁻¹. The wasps also adjust their flight speed depending on the rotation of the drum. They modulate their distance from the pivoting centre accordingly and presumably also their height above ground, so that maximal ground slip is on average 200°␣s⁻¹. The insects move along arcs by short pulses of translation, followed by rapid body turns to correct for the change in retinal position of the nest entrance. Saccadic body turns follow pulses of translation with a delay of 80–120 ms. The optomotor response is active during these turns. The control of pivoting flight most likely involves three position servos, to control the retinal position of both the azimuth and the altitude of nest and the direction of flight relative to it, and two velocity servos, one constituting the optomotor reflex and the other one serving to clamp ground slip at about 200° s⁻¹. The control of ground slip is the prime source of the dynamic constancy of learning flights, which may help wasps to scale the pivoting parallax field they produce during these flights. Constant pivoting rate may in addition be important for the acquisition of a regular sequence of snapshots and in scanning for compass cues. Accepted : 31 July 1996     

Interspecific interactions in phytophagous insects revisited: a quantitative assessment of competition theory

The importance of interspecific competition is a highly controversial and unresolved issue for community ecology in general, and for phytophagous insects in particular. Recent advancements, however, in our understanding of indirect (plant- and enemy-mediated) interactions challenge the historical paradigms of competition. Thus, in the context of this rapidly developing field, we re-evaluate the evidence for interspecific competition in phytophagous insects using a meta-analysis of published studies. Our analysis is specifically designed to test the assumptions underlying traditional competition theory, namely that competitive interactions are symmetrical, necessitate spatial and temporal co-occurrence, and increase in intensity as the density, phylogenetic similarity, and niche overlap of competing species increase. Despite finding frequent evidence for competition, we found very little evidence that plant-feeding insects conform to theoretical predictions for interspecific competition. Interactions were highly asymmetrical, similar in magnitude within vs. between feeding guilds (chewers vs. sap-feeders), and were unaffected by the quantity of resources removed (% defoliation). There was mixed support for the effects of phylogeny, spatial/temporal separation, and the relative strength of intra- vs. interspecific competition. Clearly, a new paradigm that accounts for indirect interactions and facilitation is required to describe how interspecific competition contributes to the organization of phytophagous insect communities, and perhaps to other plant and animal communities as well.     

Advances in Transgenic Research for Insect Resistance in Sugarcane

The first phase of transgenic research in sugarcane concentrated on the development and evaluation of transgenic lines transformed for resistance to biotic stresses, particularly diseases and insect pests. Sugarcane is attacked by a range of insects including tissue borers, sucking pests and canegrubs. Losses due to these pests are estimated to be around 10%. Although chemical control and integrated pest management are regularly practiced for the control of insect pests, success is often limited due to practical difficulties. The genetic complexity of sugarcane coupled with the non-availability of resistance genes in the germplasm has made conventional breeding for insect resistance difficult. In this context, transgenic technology has become a handy tool for imparting insect resistance to an elite variety which is otherwise superior for most other agronomic traits. A number of transgenic sugarcane lines have been developed with genes expressing Cry proteins, proteinase inhibitors or lectins resistant to borers, sucking insects or grubs. While commercializing transgenic lines, issues such as higher and stable transgene expression, preparedness for resistance management and non-target effects need to be addressed. To manage the constant threat of resistance development in target insects, it is imperative to deploy field-level strategies taking clues from other crops coupled with the search for new potent replacement molecules for transformation.     

Central pattern generating networks in insect locomotion

Central pattern generators (CPGs) are neural circuits that based on their connectivity can generate rhythmic and patterned output in the absence of rhythmic external inputs. This property makes CPGs crucial elements in the generation of many kinds of rhythmic motor behaviors in insects, such as flying, walking, swimming, or crawling. Arguably representing the most diverse group of animals, insects utilize at least one of these types of locomotion during one stage of their ontogenesis. Insects have been extensively used to study the neural basis of rhythmic motor behaviors, and particularly the structure and operation of CPGs involved in locomotion. Here, we review insect locomotion with regard to flying, walking, and crawling, and we discuss the contribution of central pattern generation to these three forms of locomotion. In each case, we compare and contrast the topology and structure of the CPGs, and we point out how these factors are involved in the generation of the respective motor pattern. We focus on the importance of sensory information for establishing a functional motor output and we indicate behavior‐specific adaptations. Furthermore, we report on the mechanisms underlying coordination between different body parts. Last but not least, by reviewing the state‐of‐the‐art knowledge concerning the role of CPGs in insect locomotion, we endeavor to create a common ground, upon which future research in the field of motor control in insects can build.     

Field performance of a genetically engineered strain of pink bollworm

Pest insects harm crops, livestock and human health, either directly or by acting as vectors of disease. The Sterile Insect Technique (SIT)--mass-release of sterile insects to mate with, and thereby control, their wild counterparts--has been used successfully for decades to control several pest species, including pink bollworm, a lepidopteran pest of cotton. Although it has been suggested that genetic engineering of pest insects provides potential improvements, there is uncertainty regarding its impact on their field performance. Discrimination between released and wild moths caught in monitoring traps is essential for estimating wild population levels. To address concerns about the reliability of current marking methods, we developed a genetically engineered strain of pink bollworm with a heritable fluorescent marker, to improve discrimination of sterile from wild moths. Here, we report the results of field trials showing that this engineered strain performed well under field conditions. Our data show that attributes critical to SIT in the field--ability to find a mate and to initiate copulation, as well as dispersal and persistence in the release area--were comparable between the genetically engineered strain and a standard strain. To our knowledge, these represent the first open-field experiments with a genetically engineered insect. The results described here provide encouragement for the genetic control of insect pests.     

Orientation Cues for High-Flying Nocturnal Insect Migrants: Do Turbulence-Induced Temperature and Velocity Fluctuations Indicate the Mean Wind Flow?

Migratory insects flying at high altitude at night often show a degree of common alignment, sometimes with quite small angular dispersions around the mean. The observed orientation directions are often close to the downwind direction and this would seemingly be adaptive in that large insects could add their self-propelled speed to the wind speed, thus maximising their displacement in a given time. There are increasing indications that high-altitude orientation may be maintained by some intrinsic property of the wind rather than by visual perception of relative ground movement. Therefore, we first examined whether migrating insects could deduce the mean wind direction from the turbulent fluctuations in temperature. Within the atmospheric boundary-layer, temperature records show characteristic ramp-cliff structures, and insects flying downwind would move through these ramps whilst those flying crosswind would not. However, analysis of vertical-looking radar data on the common orientations of nocturnally migrating insects in the UK produced no evidence that the migrants actually use temperature ramps as orientation cues. This suggests that insects rely on turbulent velocity and acceleration cues, and refocuses attention on how these can be detected, especially as small-scale turbulence is usually held to be directionally invariant (isotropic). In the second part of the paper we present a theoretical analysis and simulations showing that velocity fluctuations and accelerations felt by an insect are predicted to be anisotropic even when the small-scale turbulence (measured at a fixed point or along the trajectory of a fluid-particle) is isotropic. Our results thus provide further evidence that insects do indeed use turbulent velocity and acceleration cues as indicators of the mean wind direction.     

Insect-plant interactions on a planet of weeds

Two conflicting views confront ecologists and evolutionary biologists on the degree of symmetry in interactions between plants and phytophagous insects. The symmetrical view holds that insects and plants have strong effects on one another's evolutionary and ecological dynamics. Thus, herbivores are regarded as a major influence on plant distribution and abundance in contemporary ecosystems, and coevolution is commonly invoked to explain adaptive radiation in plants and insects, host specialization in insects, as well as much of the morphological and chemical variety observed in plants. The asymmetrical view acknowledges that plants have major effects on insects, but claims that insects seldom impose significant effects on plants. Proponents of the asymmetric view tend to ignore or discount insect-plant interactions in communities and ecosystems altered by human impacts. If we recognize the scope and scale of human impacts, and ways in which these impacts change insect-plant interactions, then our views about symmetry or asymmetry in insect-plant interactions will change. To understand, predict, and manage insect herbivory we need to study it in all its manifestations. In particular, the study of interactions involving alien species is both an urgent priority for environmental management and potentially a source of ecological insights on the role of herbivores in plant population and community dynamics. A complete theory of insect/host plant interactions must explain and predict interactions both within and beyond the native range. Such a theory might guide efforts to deal with environmental problems stemming from rapid rates of extinction and homogenization of the world's biota.     

Almond orchards with living ground cover host more wild insect pollinators

Wild pollinators are becoming more valuable to global agriculture as the commercial honeybee industry is increasingly affected by disease and other stressors. Perennial tree crops are particularly reliant on insect pollination, and are often pollen limited. Research on how different tree crop production systems influence the richness and abundance of wild pollinators is, however, limited. We investigated, for the first time, the richness and abundance of potential wild pollinators in commercial temperate almond orchards in Australia, and compared them to potential pollinator communities in proximate native vegetation. We quantified ground cover variables at each site and assessed the value of ground cover on the richness and abundance of potential wild pollinators in commercial almond systems focussing on three common taxa: bees, wasps and flies. More insects were caught in orchards with living ground cover than in native vegetation or orchards without ground cover, although overall species richness was highest in native vegetation. Percent ground cover was positively associated with wasp richness and abundance, and native bee richness, but flies showed no association with ground cover. The strongest positive relationship was between native bee abundance and the richness of ground cover plants. Our results suggest that maintaining living ground cover within commercial almond orchards could provide habitat and resources for potential wild pollinators, particularly native bees. These insects have the potential to provide a valuable ecosystem service to pollinator-dependent crops such as almond.     

Predicting Visibility of Aircraft

Visual detection of aircraft by human observers is an important element of aviation safety. To assess and ensure safety, it would be useful to be able to be able to predict the visibility, to a human observer, of an aircraft of specified size, shape, distance, and coloration. Examples include assuring safe separation among aircraft and between aircraft and unmanned vehicles, design of airport control towers, and efforts to enhance or suppress the visibility of military and rescue vehicles. We have recently developed a simple metric of pattern visibility, the Spatial Standard Observer (SSO). In this report we examine whether the SSO can predict visibility of simulated aircraft images. We constructed a set of aircraft images from three-dimensional computer graphic models, and measured the luminance contrast threshold for each image from three human observers. The data were well predicted by the SSO. Finally, we show how to use the SSO to predict visibility range for aircraft of arbitrary size, shape, distance, and coloration.     

Heat tolerance variation reveals vulnerability of tropical herbivore–parasitoid interactions to climate change

Assessing the heat tolerance (CTmax) of organisms is central to understand the impact of climate change on biodiversity. While both environment and evolutionary history affect CTmax, it remains unclear how these factors and their interplay influence ecological interactions, communities and ecosystems under climate change. We collected and reared caterpillars and parasitoids from canopy and ground layers in different seasons in a tropical rainforest. We tested the CTmax and Thermal Safety Margins (TSM) of these food webs with implications for how species interactions could shift under climate change. We identified strong influence of phylogeny in herbivore–parasitoid community heat tolerance. The TSM of all insects were narrower in the canopy and parasitoids had lower heat tolerance compared to their hosts. Our CTmax-based simulation showed higher herbivore–parasitoid food web instability under climate change than previously assumed, highlighting the vulnerability of parasitoids and related herbivore control in tropical rainforests, particularly in the forest canopy.     

The ‘mother knows best’ principle: should soil insects be included in the preference–performance debate?

Abstract.  1. Few entomological studies include soil-dwelling insects in mainstream ecological theory, for example the preference–performance debate. The preference–performance hypothesis predicts that when insect herbivores have offspring with limited capacity to relocate in relation to a host plant, there is a strong selection pressure for the adult to oviposit on plants that will maximise offspring performance.
2. This paper discusses the proposition that insect herbivores that live above ground, but have soil-dwelling offspring, should be included in the preference–performance debate. Twelve relevant studies were reviewed to assess the potential for including soil insects in this framework, before presenting a preliminary case study using the clover root weevil ( Sitona lepidus ) and its host plant, white clover ( Trifolium repens ).
3. Maternal S. lepidus preferentially oviposited on T. repens plants that had rhizobial root nodules (which enhance offspring performance) rather than T. repens plants without nodules, despite plants having similar foliar nutritional quality. This suggests that adult behaviour above ground was influenced by below-ground host-plant quality.
4. A conceptual model is presented to describe how information about the suitability for offspring below ground could underpin oviposition behaviour of parental insects living above ground, via plant- and soil-mediated semiochemicals. These interactions between genetically related, but spatially separated, insect herbivores raise important evolutionary questions such as how induced plant responses above ground affect offspring living below ground and vice versa.