Synchronization of wing beat cycle of the desert locust, Schistocerca gregaria, by periodic light flashes.

Studies on the generation of rhythmic motor patterns have shown that peripheral sensory input may contribute substantially to the rhythm generating network. A prominent example is the wing beat frequency of desert locusts, which can be entrained to rhythmic mechanosensory input, but also to the frequency of periodic light flashes. To further analyze the entrainment by light, tethered flying locusts were presented with periodic light flashes, while the position of the forewing was filmed. We show that entrainment of wing beat occurs both in the UV and green range of light. Animals maintained a characteristic phase relationship to the light stimulus with the most elevated wing position occurring at the end of the dark phase. Speed and time course of entrainment varied greatly and ranged from the duration of a single wing beat cycle to several seconds. To identify the visual system mediating entrainment, synchronization to UV light was tested after cutting the optic stalks to the optic lobes/compound eyes or the ocellar nerves. The results show that light entrainment of the locust flight pattern is largely and perhaps exclusively mediated via the fast ocellar pathway and may have a role to stabilize flight with respect to the horizon.

Applicability of white-noise techniques to analyzing motion responses.

Motion processing in visual neurons is often understood in terms of how they integrate light stimuli in space and time. These integrative properties, known as the spatiotemporal receptive fields (STRFs), are sometimes obtained using white-noise techniques where a continuous random contrast sequence is delivered to each spatial location within the cell's field of view. In contrast, motion stimuli such as moving bars are usually presented intermittently. Here we compare the STRF prediction of a neuron's response to a moving bar with the measured response in second-order interneurons (L-neurons) of dragonfly ocelli (simple eyes). These low-latency neurons transmit sudden changes in intensity and motion information to mediate flight and gaze stabilization reflexes. A white-noise analysis is made of the responses of L-neurons to random bar stimuli delivered either every frame (densely) or intermittently (sparsely) with a temporal sequence matched to the bar motion stimulus. Linear STRFs estimated using the sparse stimulus were significantly better at predicting the responses to moving bars than the STRFs estimated using a traditional dense white-noise stimulus, even when second-order nonlinear terms were added. Our results strongly suggest that visual adaptation significantly modifies the linear STRF properties of L-neurons in dragonfly ocelli during dense white-noise stimulation. We discuss the ability to predict the responses of visual neurons to arbitrary stimuli based on white-noise analysis. We also discuss the likely functional advantages that adaptive receptive field structures provide for stabilizing attitude during hover and forward flight in dragonflies.

Curiosity and context revisited: crassulacean acid metabolism in the Anthropocene.

Having gained some understanding of the consequences of the CO(2)-concentrating mechanisms in crassulacean acid metabolism (CAM) that internalize the photosynthetic environment of the Cretaceous on a daily basis, it may be time to consider potential long-term effects of the planetary CO(2)-concentrating mechanism on growth and ecology of these plants in the Anthropocene. This paper emphasizes our limited understanding of the carbohydrate economy of CAM in relation to growth processes and briefly reviews recent studies of the diel cycles of growth in these plants. An inadvertent long-term, regional-scale experiment from the past is revisited in which an Opuntia monoculture grew to occupy >25 million hectares of farmland in central eastern Australia, producing a total biomass of about 1.5 billion tonnes in about 80 years. Although at the time it does not seem to have been recognized that this invasion involved CAM, a botanist from the University of Melbourne, Jean White-Haney emerges as a heroic pioneer in the control of the invader by poison and pioneered its biological control. The Opuntia population was expanding at 10-100 ha h(-1) when it was brought to a halt within a decade by the voracious appetite of Cactoblastis cactorum larvae. It is now known that the female parent moth of this predator detects CAM in O. stricta prior to oviposition by deploying the most sensitive CO(2) detector system yet found in the Lepidoptera. The O. stricta invasion is a dramatic demonstration of the capacity of CAM plants to attain and sustain high biomass; to sequester and retain atmospheric CO(2). In conclusion, experiments are reviewed that show stimulation of CO(2) assimilation, growth, and biomass of CAM plants by elevated atmospheric [CO(2)], and the proposition that these plants may have a role in atmospheric CO(2) sequestration is re-examined. This role may be compromised by predators such as Cactoblastis. However the moth CO(2) sensors are adapted to pre-industrial atmospheric [CO(2)] and FACE (free-air CO(2) enrichment) experiments show this exquisite system of biological control is also compromised by rising global [CO(2)] in the Anthropocene.

Directional selectivity in the simple eye of an insect.

Among other sensory modalities, flight stabilization in insects is performed with the aid of visual feedback from three simple eyes (ocelli). It is thought that each ocellus acts as a single wide-field sensor that detects changes in light intensity. We challenge this notion by providing evidence that, when light-adapted, the large retinal L-neurons in the median ocellus of the dragonfly respond in a directional way to upward moving bars and gratings. This ability is pronounced under UV illumination but weak or nonexistent in green light and is optimal at angular velocities of approximately 750 degrees s(-1). Using a reverse-correlation technique, we analyze the functional organization of the receptive fields of the L-neurons. Our results reveal that L-neurons alter the structure of their linear spatiotemporal receptive fields with changes in the illuminating wavelength, becoming more inseparable and directional in UV light than in green. For moving bars and gratings, the strength of directionality predicted from the receptive fields is consistent with the measured values. Our results strongly suggest that, during the day, the retinal circuitry of the dragonfly median ocellus performs an early linear stage of motion processing. The likely advantage of this computation is to enhance pitch control.

Form vision in the insect dorsal ocelli: an anatomical and optical analysis of the Locust Ocelli.

The dorsal ocelli are commonly considered to be incapable of form vision, primarily due to underfocused dioptrics. We investigate the extent to which this is true of the ocelli of the locust Locusta migratoria. Locust ocelli contain thick lenses with a pronounced concavity on the inner surface, and a deep clear zone separating retina and lens. In agreement with previous research, locust ocellar lenses were found to be decidedly underfocused with respect to the retina. Nevertheless, the image formed at the level of the retina contains substantial information that may be extractable by individual photoreceptors. Contrary to the classical view it is concluded that some capacity for resolution is present in the locust ocelli.

Form vision in the insect dorsal ocelli: an anatomical and optical analysis of the dragonfly median ocellus.

Previous work has suggested that dragonfly ocelli are specifically adapted to resolve horizontally extended features of the world, such as the horizon. We investigate the optical and anatomical properties of the median ocellus of Hemicordulia tau and Aeshna mixta to determine the extent to which the findings support this conclusion. Dragonfly median ocelli are shown to possess a number of remarkable properties: astigmatism arising from the elliptical shape of the lens is cancelled by the bilobed shape of the inner lens surface, interference microscopy reveals complex gradients of refractive index within the lens, the morphology of the retina results in zones of high acuity, and the eye has an exceedingly high sensitivity for a diurnal terrestrial invertebrate. It is concluded that dragonfly ocelli employ a number of simple, yet elegant, anatomical and optical strategies to ensure high sensitivity, fast transduction speed, wide fields of views and a modicum of spatial resolving power.

The mapping of visual space by dragonfly lateral ocelli.

We study the extent to which the lateral ocelli of dragonflies are able to resolve and map spatial information, following the recent finding that the median ocellus is adapted for spatial resolution around the horizon. Physiological optics are investigated by the hanging-drop technique and related to morphology as determined by sectioning and three-dimensional reconstruction. L-neuron morphology and physiology are investigated by intracellular electrophysiology, white noise analysis and iontophoretic dye injection. The lateral ocellar lens consists of a strongly curved outer surface, and two distinct inner surfaces that separate the retina into dorsal and ventral components. The focal plane lies within the dorsal retina but proximal to the ventral retina. Three identified L-neurons innervate the dorsal retina and extend the one-dimensional mapping arrangement of median ocellar L-neurons, with fields of view that are directed at the horizon. One further L-neuron innervates the ventral retina and is adapted for wide-field intensity summation. In both median and lateral ocelli, a distinct subclass of descending L-neuron carries multi-sensory information via graded and regenerative potentials. Dragonfly ocelli are adapted for high sensitivity as well as a modicum of resolution, especially in elevation, suggesting a role for attitude stabilisation by localization of the horizon.

The mapping of visual space by identified large second-order neurons in the dragonfly median ocellus.

In adult dragonflies, the compound eyes are augmented by three simple eyes known as the dorsal ocelli. The outputs of ocellar photoreceptors converge on relatively few second-order neurons with large axonal diameters (L-neurons). We determine L-neuron morphology by iontophoretic dye injection combined with three-dimensional reconstructions. Using intracellular recording and white noise analysis, we also determine the physiological receptive fields of the L-neurons, in order to identify the extent to which they preserve spatial information. We find a total of 11 median ocellar L-neurons, consisting of five symmetrical pairs and one unpaired neuron. L-neurons are distinguishable by the extent and location of their terminations within the ocellar plexus and brain. In the horizontal dimension, L-neurons project to different regions of the ocellar plexus, in close correlation with their receptive fields. In the vertical dimension, dendritic arborizations overlap widely, paralleled by receptive fields that are narrow and do not differ between different neurons. These results provide the first evidence for the preservation of spatial information by the second-order neurons of any dorsal ocellus. The system essentially forms a one-dimensional image of the equator over a wide azimuthal area, possibly forming an internal representation of the horizon. Potential behavioural roles for the system are discussed.

A spatiotemporal white noise analysis of photoreceptor responses to UV and green light in the dragonfly median ocellus.

Adult dragonflies augment their compound eyes with three simple eyes known as the dorsal ocelli. While the ocellar system is known to mediate stabilizing head reflexes during flight, the ability of the ocellar retina to dynamically resolve the environment is unknown. For the first time, we directly measured the angular sensitivities of the photoreceptors of the dragonfly median (middle) ocellus. We performed a second-order Wiener Kernel analysis of intracellular recordings of light-adapted photoreceptors. These were stimulated with one-dimensional horizontal or vertical patterns of concurrent UV and green light with different contrast levels and at different ambient temperatures. The photoreceptors were found to have anisotropic receptive fields with vertical and horizontal acceptance angles of 15 degrees and 28 degrees, respectively. The first-order (linear) temporal kernels contained significant undershoots whose amplitudes are invariant under changes in the contrast of the stimulus but significantly reduced at higher temperatures. The second-order kernels showed evidence of two distinct nonlinear components: a fast acting self-facilitation, which is dominant in the UV, followed by delayed self- and cross-inhibition of UV and green light responses. No facilitatory interactions between the UV and green light were found, indicating that facilitation of the green and UV responses occurs in isolated compartments. Inhibition between UV and green stimuli was present, indicating that inhibition occurs at a common point in the UV and green response pathways. We present a nonlinear cascade model (NLN) with initial stages consisting of separate UV and green pathways. Each pathway contains a fast facilitating nonlinearity coupled to a linear response. The linear response is described by an extended log-normal model, accounting for the phasic component. The final nonlinearity is composed of self-inhibition in the UV and green pathways and inhibition between these pathways. The model can largely predict the response of the photoreceptors to UV and green light.

Volatile organic compounds as signals in a plant-herbivore system: electrophysiological responses in olfactory sensilla of the moth Cactoblastis cactorum.

The morphological sensillum types on the antennae of male and female Cactoblastis cactorum were visualized by scanning electron microscopy. Electrophysiological recordings were performed for the first time on single olfactory sensilla of C. cactorum. The male sensilla trichodea house a receptor cell responding to the putative pheromone component (9Z,12E)-tetradecadienyl acetate. The sensilla trichodea of the females were much shorter than those of the males and contained specialized receptor cells responding to certain terpenoids, the most frequent being the nerolidol-sensitive cell. The sensilla auricillica and sensilla basiconica of both sexes contained cells responding less specifically to terpenoid compounds as well as to green leaf volatiles. Cells of the sensilla coeloconica responded to aliphatic aldehydes and acids. Eight volatile organic compounds emitted by Opuntia stricta, a host plant of C. cactorum, were identified using gas chromatography-mass spectrometry, beta-caryophyllene being the major compound. Five compounds identified by gas chromatography in the headspace of O. stricta elicited responses in olfactory receptor cells of C. cactorum, nonanal being the most active compound and therefore a candidate attractant of C. cactorum.

The renal type IIa Na/Pi cotransporter: structure-function relationships.

The type IIa Na/Pi cotransporter mediates proximal tubular brush-border membrane secondary active phosphate (Pi) flux. It is rate limiting in tubular Pi reabsorption and, thus, a final target in many physiological and pathophysiological situations of altered renal Pi handling. In the present short review, we will briefly summarize our current knowledge about the transport mechanism (cycle) as well as particular regions of the transporter protein ("molecular domains") that potentially determine transport characteristics.

Effects of changes in atmospheric carbon dioxide on the location of hosts by the moth, Cactoblastis cactorum.

Sensory organs that detect CO2 are common in herbivorous moths and butterflies, but their function has been unclear until now. As the CO2 gradients in the vicinity of a host plant depend on its physiological condition, CO2 could provide a sensory cue for the suitability of the plant as a larval food source. This study investigated whether changing the atmospheric CO2 concentration affected oviposition by Cactoblastis cactorum on its host, the cactus Opuntia stricta. On host plants exposed to rapid fluctuations in CO2 concentration, the frequency of oviposition was reduced by a factor of 3.2 compared to the control. As the fluctuations mask the much smaller CO2 signals generated by the plants, this suggests that those signals constitute an important component of the host identification process. On host plants exposed to a constant background of doubled CO2, oviposition was also reduced, by a factor of 1.8. An increased background reduces host signal detectability, partially as a consequence of a general principle of sensory physiology (Weber-Fechner's law), and partially due to other factors specific to CO2-receptor neurons.