Prey capture and accommodation in the sandlance, Limnichthyes fasciatus (Creediidae; Teleostei)

The eyes of the sandlance, Limnichthyes fasciatus (Creediidae, Teleostei) move independently and possess a refractive cornea, a convexiclivate fovea and a non-spherical lens giving rise to a wide separation of the nodal point from the axis of rotation of the eye much like that of a chameleon. To investigate this apparent convergence of the visual optics in these phylogenetically disparate species, we examine feeding behaviour and accommodation in the sandlance with special reference to the possibility that sandlances use accommodation as a depth cue to judge strike length. Frame-by-frame analysis of over 2000 strikes show a 100% success rate. Explosive strikes are completed in 50 ms over prey distances of four body lengths. Close-up video confirms that successful strikes can be initiated monocularly (both normally and after monocular occlusion) showing that binocular cues are not necessary to judge the length of a strike. Additional means of judging prey distance may also be derived from parallax information generated by rotation of the eye as suggested for chameleons. Using photorefraction on anaesthetised sandlances, accommodative changes were induced with acetylcholine and found to range between 120 D and 180 D at a speed of 600–720 D s⁻¹. The large range of accommodation (25% of the total power) is also thought to be mediated by corneal accommodation where the contraction of a unique cornealis muscle acts to change the corneal curvatures. Accepted: 8 December 1999     

The scaling of tongue projection in the veiled chameleon,Chamaeleo calyptratus

Within a year of hatching, chameleons can grow by up to two orders of magnitude in body mass. Rapid growth of the feeding mechanism means that bones, muscles, and movements change as chameleons grow while needing to maintain function. A previous morphological study showed that the musculoskeletal components of the feeding apparatus grow with negative allometry relative to snout–vent length (SVL) in chameleons. Here, we investigate the scaling of prey capture kinematics and muscle physiological cross-sectional area in the veiled chameleon, Chamaeleo calyptratus. The chameleons used in this study varied in size from approximately 3 to 18 cm SVL (1–200 g). Feeding sequences of 12 chameleons of different sizes were filmed and the timing of movements and the displacements and velocities of the jaws, tongue, and the hyolingual apparatus were quantified. Our results show that most muscle cross-sectional areas as well as tongue and hyoid mass scaled with isometry relative to mandible length, yet with negative allometry relative to SVL. Durations of movement also scaled with negative allometry relative to SVL and mandible length. Distances and angles generally scaled as predicted under geometric similarity (slopes of 1 and 0, respectively), while velocities generally scaled with slopes greater than 0 relative to SVL and mandible length. These data indicate that the velocity of jaw and tongue movements is generally greater in adults compared to juveniles. The discrepancy between the scaling of cross-sectional areas versus movements suggests changes in the energy storage and release mechanisms implicated in tongue projection.     

Binocular vision and accommodation in prey-catching chameleons

We have studied the role of accommodation and binocular convergence in the predatory behaviour of two chameleon species (Chamaeleocalyptratus, C. dilepis). Accommodation measurements support earlier observations that accommodation is the major distance cue. Specifically, accommodation speed (60 D s⁻¹), amplitude (45 D) and precision (no significant under-accommodation) were superior to those of other terrestrial vertebrates. Similar to other vertebrates, accommodation was accompanied by a prominent pupillary constriction (pupillary near response). Accommodation could be coupled or uncoupled in both eyes, depending on the experimental situation or the phase of the predatory behavioural sequence. Uncoupled accommodation occurred: 1. During scanning saccadic eye movements for prey detection. Only one eye accommodated appropriately, the other adopted a hyperopic resting refractive state. Attention switched from one eye to the other at approximately 1-s intervals. 2. During initial stages of distance estimation. Coupled accommodation only occurred shortly before the tongue shot. Coupling was demonstrated by either covering one eye with a lens or covering one eye with an infrared light transmitting cut-off filter which still permitted refraction to be measured. In both cases the amount of accommodation was identical in both eyes. Search-coil measurements showed that the angle of convergence of both eyes is too variable to permit triangulation or to provide the basic requirement for stereopsis (matching corresponding points). We conclude that coupling of accommodation serves to improve accommodation precision rather than to permit stereopsis. Accepted: 3 September 1997     

Morphology and histochemistry of the hyolingual apparatus in chameleons

We reexamined the morphological and functional properties of the hyoid, the tongue pad, and hyolingual musculature in chameleons. Dissections and histological sections indicated the presence of five distinctly individualized pairs of intrinsic tongue muscles. An analysis of the histochemical properties of the system revealed only two fiber types in the hyolingual muscles: fast glycolytic and fast oxidative glycolytic fibers. In accordance with this observation, motor-endplate staining showed that all endplates are of the en-plaque type. All muscles show relatively short fibers and large numbers of motor endplates, indicating a large potential for fine muscular control. The connective tissue sheet surrounding the entoglossal process contains elastin fibers at its periphery, allowing for elastic recoil of the hyolingual system after prey capture. The connective tissue sheets surrounding the m. accelerator and m. hyoglossus were examined under polarized light. The collagen fibers in the accelerator epimysium are configured in a crossed helical array that will facilitate limited muscle elongation. The microstructure of the tongue pad as revealed by SEM showed decreased adhesive properties, indicating a change in the prey prehension mechanics in chameleons compared to agamid or iguanid lizards. These findings provide the basis for further experimental analysis of the hyolingual system.     

Ultrastructure and function of the adhesion–capture apparatus of Stenus species (Coleoptera, Staphylinidae)

 The elongated labium of rove beetles of the genus Stenus forms an adhesion–capture apparatus that enables the animal to catch fast fleeing prey, for example, collemboles. Structural details of this labium have been reinvestigated by using transmission electron microscopy, and the functional model of the capture mechanism has been refined. The openings of glandular ductules have been found to be located at the outer margin of the sticky cushions formed by the paraglossae. These cushions can be expanded by hemolymph pressure and are compressed when the tip of the protrusible labium hits the prey. Endocuticular fibers stabilize the cushions internally and determine the shape of the cushions in both the expanded and the compressed state. Within the membranous connecting tube that connects the prementum with the head capsule, the existence of an extra inner membrane has been confirmed. It is formed by a portion of the epidermis that has become detached from the cuticle. The most important part of the functional model of the protrusion of the labium is that the membranous connecting tube turns itself inside out distally, but the extra inner membrane does so proximally. During protrusion of the labium by hemolymph pressure, the prementum is accelerated during the initial phase of the process, and the nerves, muscles, tracheae, and glandular ductules that are attached to it are passively drawn into the labium. The mechanoreceptive function of setae at the distal end of the prementum has been confirmed on the basis of their ultrastructural characteristics. Accepted: 4 September 1998     

Tests of a linear model of visual-vestibular interaction using the technique of parameter estimation

The goal of this study was to test whether a superposition model of smooth-pursuit and vestibulo-ocular reflex (VOR) eye movements could account for the stability of gaze that subjects show as they view a stationary target, during head rotations at frequencies that correspond to natural movements. Horizontal smooth-pursuit and the VOR were tested using sinusoidal stimuli with frequencies in the range 1.0–3.5 Hz. During head rotation, subjects viewed a stationary target either directly or through an optical device that required eye movements to be approximately twice the amplitude of head movements in order to maintain foveal vision of the target. The gain of compensatory eye movements during viewing through the optical device was generally greater than during direct viewing or during attempted fixation of the remembered target location in darkness. This suggests that visual factors influence the response, even at high frequencies of head rotation. During viewing through the optical device, the gain of compensatory eye movements declined as a function of the frequency of head rotation (P < 0.001) but, at any particular frequency, there was no correlation with peak head velocity (P > 0.23), peak head acceleration (P > 0.22) or retinal slip speed (P > 0.22). The optimal values of parameters of smooth-pursuit and VOR components of a simple superposition model were estimated in the frequency domain, using the measured responses during head rotation, as each subject viewed the stationary target through the optical device. We then compared the model's prediction of smooth-pursuit gain and phase, at each frequency, with values obtained experimentally. Each subject's pursuit showed lower gain and greater phase lag than the model predicted. Smooth-pursuit performance did not improve significantly if the moving target was a 10 deg × 10 deg Amsler grid, or if sinusoidal oscillation of the target was superimposed on ramp motion. Further, subjects were still able to modulate the gain of compensatory eye movements during pseudo-random head perturbations, making improved predictor performance during visual-vestibular interactions unlikely. We conclude that the increase in gain of eye movements that compensate for head rotations when subjects view, rather than imagine, a stationary target cannot be adequately explained by superposition of VOR and smooth-pursuit signals. Instead, vision may affect VOR performance by determining the context of the behavior. Received: 16 June 1997 / Accepted: 5 December 1997     

A reptilian smoking gun: first record of invasive Jackson’s chameleon (<Emphasis Type="Italic">Chamaeleo jacksonii</Emphasis>) predation on native Hawaiian species

Here we report the first conclusive evidence of an introduced reptile (Chamaeleo jacksonii) feeding on Hawaiian taxa, including 11 snails in four endemic genera from two families, including four individuals of an endangered species (Achatinella mustelina), and native insects in five genera. Native Hawaiian invertebrates were discovered in the dissected stomachs of wild caught Jackson’s chameleons collected from June to November 2009 on the island of Oahu. Although Jackson’s chameleons were introduced to the Hawaiian Islands in the early 1970s, ecological impacts have never been documented. Of particular concern is the fact that chameleons have previously only rarely been found in native Hawaiian habitat, although 12 were recently collected in a mid-elevation native forest, an area that is not likely to be suitable for their long-term persistence, but that is adjacent to higher elevation pristine forest where endemic prey are abundant and favorable climatic conditions exist for chameleon persistence. One concern is that Jacksons’s chameleons may be undergoing a range expansion into upper elevation pristine forests. If chameleons reach and establish populations in these areas, devastating impacts to the native ecosystem are possible. A thorough understanding of the impacts of chameleons on Hawaiian fauna will require additional evaluation and sampling, but dissemination of this discovery in a timely fashion is important as it provides new information regarding this threat. Monitoring and collection of chameleons is ongoing, particularly in native Hawaiian forest habitats at mid and upper elevations (600–1,300 m).     

Instability helps virtual flies to mate

In this paper we perform a bifurcation analysis for a discrete time dynamical system, describing the behavior of a virtual fly, developed by Böddeker and Egelhaaf (2003). Like real blowflies, the virtual counterparts exhibit a dichotomous behavior: they catch small targets but follow big objects at a constant distance. We consider this model for targets on linear and on circular trajectories. Then we transform the system into a ‘‘frozen’’ form, such that the position of the target is fixed. It turns out that the loss of stability of a fixed point in the frozen system due to a Neimark-Sacker bifurcation, explains the dichotomous behavior of the virtual fly.     

The application of co-ordination dynamics to the analysis of discrete movements using table-tennis as a paradigm skill

 The purpose of this experiment was to explore the application of co-ordination dynamics to the analysis of discrete rather than cyclical movements. Subjects, standing in a fixed position, were required to return table-tennis balls delivered to different spatial locations in the direction of a fixed target. This was achieved in condition 1 by systematically scaling, from left to right and vice versa, the `spatial location' of the ball–identified as a control parameter. In condition 2, the control condition, the spatial location was varied randomly over the same range. The changes between regimes of the stroke co-ordination pattern, defined at two different levels, (1) organisational – forehand or backhand drive, and (2) kinematic–the distance of the bat at ball–bat contact relative to the leading edge of the table, were identified as collective variables, the values of which changed spontaneously at the transition points exposed by the control parameter. The switch between regimes was shown to be dependent upon the direction of scaling, i.e. a hysteresis effect was identified in both conditions. These findings confirm that the conceptual and methodological frameworks of co-ordination dynamics can be applied, appropriately, to the analysis of discrete movements. Moreover, it would seem that control parameter values (spatial location of the ball) do not necessarily have to be scaled in a systematic way in order to produce the required effects. Received: 22 April 1999 / Accepted in revised form: 8 May 2000     

Generalist and specialist predators that mediate permanence in ecological communities

 General dynamic models of systems with two prey and one or two predators are considered. After rescaling the equations so that both prey have the same intrinsic rate of growth, it is shown that there exists a generalist predator that can mediate permanence if and only if there is a population density of a prey at which its per-capita growth rate is positive yet less than its competitor’s invasion rate. In particular, this result implies that if the outcome of competition between the prey is independent of initial conditions, then there exists a generalist predator that mediates permanence. On the other hand, if the outcome of competition is contingent upon initial conditions (i.e., the prey are bistable), then there may not exist a suitable generalist predator. For example, bistable prey modeled by the Ayala–Gilpin (θ-Logistic) equations can be stabilized if and only if θ<1 for one of the prey. It is also shown that two specialist predators always can mediate permanence between bistable prey by creating a repelling heteroclinic cycle consisting of fixed points and limit cycles. Received 10 August 1996; received in revised form 21 March 1997     

Object segmentation model: analytical results and biological implications

 A simple, biologically motivated neural network for segmentation of a moving object from a visual scene is presented. The model consists of two parts: an object selection model which employs a scaling approach for receptive field sizes, and a subsequent network implementing a spotlight by means of multiplicative synapses. The network selects one object out of several, segments the rough contour of the object, and encodes the winner object's position with high accuracy. Analytical equations for the performance level of the network, e.g., for the critical distance of two objects above which they are perceived as separate, are derived. The network preferentially chooses the object with the largest angular velocity and the largest angular width. An equation for the velocity and width preferences is presented. Additionally it is shown that for certain neurons of the model, flat receptive fields are more favourable than Gaussian ones. The network exhibits performances similar to those known from amphibians. Various electrophysiological and behavioral results – e.g., the distribution of the diameters of the receptive fields of tectal neurons, of the tongue-projecting salamander Hydromantes italicus and the range of optimal prey velocities for prey catching – can be understood on the basis of the model. Received: 7 December 2000 / Accepted: 13 February 2001     

Modulation of prey capture kinematics and the role of lingual sensory feedback in the lizard Pogona vitticeps

Most organisms feed on a variety of prey that may differ dramatically in their physical and behavioural characteristics (e.g. mobility, mass, texture, etc.). Thus the ability to modulate prey capture behaviour in accordance with the characteristics of the food appears crucial. In animals that use rapid tongue movements to capture prey (frogs and chameleons), the coordination of jaws and tongue is based on visual cues gathered prior to the prey capture event. However, most iguanian lizards have much slower tongue-based prey capture systems suggesting that sensory feedback from the tongue may play an important role in coordinating jaw and tongue movements. We investigated the modulation of prey capture kinematics in the agamid lizard Pogona vitticeps when feeding on a range of food items differing in their physical characteristics. As the lizard is a dietary generalist, we expected it to be able to modulate its prey capture kinematics as a function of the (mechanical) demands imposed by the prey. Additionally, we investigated the role of lingual sensory feedback by transecting the trigeminal sensory afferents. Our findings demonstrated that P. vitticeps modulates its prey capture kinematics according to specific prey properties (e.g. size). In addition, transection of the trigeminal sensory nerves had a strong effect on prey capture kinematics. However, significant prey type effects and prey type by transection effects suggest that other sources of sensory information are also used to modulate the prey capture kinematics in P. vitticeps.     

Sensory modulation and behavioral choice during feeding in the Australian frog, Cyclorana novaehollandiae

This study investigates how visual and tactile sensory information, as well as biomechanical effects due to differences in physical characteristics of the prey, influence feeding behavior in the frog Cyclorana novaehollandiae. Video motion analysis was used to quantify movement patterns produced when feeding on five prey types (termites, waxworms, crickets, mice and earthworms). Twelve kinematic variables differed significantly among prey types, and twelve variables were correlated with prey characteristics (including mass, length, height and velocity of movement). Results indicate that C.␣novaehollandiae uses a different strategy to capture each prey type. Visual assessment of prey characteristics appeared to be more important in modulating feeding behavior than tactile cues or biomechanical effects. We propose a hierarchical hypothesis of behavioral choice, in which decisions are based primarily on visual analysis of prey characteristics. In this model, the frogs first choose between jaw prehension and tongue prehension based on prey size. If they have chosen jaw prehension, they next choose between upward or downward head rotation based on length and height of the prey. If they have chosen tongue prehension, they next choose between behavior for fast and slow prey. Final decisions may be the result of behavioral fine tuning based on tactile feedback. Accepted: 5 August 1996     

Eye movements and target fixation during dragonfly prey-interception flights

The capture of flying insects by foraging dragonflies is a highly accurate, visually guided behavior. Rather than simply aiming at the prey’s position, the dragonfly aims at a point in front of the prey, so that the prey is intercepted with a relatively straight flight trajectory. To better understand the neural mechanisms underlying this behavior, we used high-speed video to quantify the head and body orientation of dragonflies (female Erythemis simplicicollis flying in an outdoor flight cage) relative to an artificial prey object before and during pursuit. The results of our frame-by-frame analysis showed that during prey pursuit, the dragonfly adjusts its head orientation to maintain the image of the prey centered on the “crosshairs” formed by the visual midline and the dorsal fovea, a high acuity streak that crosses midline at right angles about 60° above the horizon. The visual response latencies to drifting of the prey image are remarkably short, ca. 25 ms for the head and 30 ms for the wing responses. Our results imply that the control of the prey-interception flight must include a neural pathway that takes head position into account.     

Prey location,biomechanical constraints,and motor program choice during prey capture in the tomato frog, <Emphasis Type="Italic">Dyscophus guineti</Emphasis>

This study investigated how visual information about prey location and biomechanical constraints of the feeding apparatus influence the feeding behavior of the tomato frog, Dyscophus guineti. When feeding on prey at small azimuths (less than ± 40°), frogs aimed their heads toward the prey but did not aim their tongues relative to their heads. Frogs projected their tongues rapidly by transferring momentum from the lower jaw to the tongue. Storage and recovery of elastic energy by the mouth opening muscles amplified the velocities of mouth opening and tongue projection. This behavior can only occur when the lower jaw and tongue are aligned (i.e., within the range of motion of the neck). When feeding on prey at large azimuths (greater than ± 40°), frogs aimed both the head and tongue toward the prey and used a muscular hydrostatic mechanism to project the tongue. Hydrostatic elongation allows for frogs to capture prey at greater azimuthal locations. Because the tongue moves independently of the lower jaw, frogs can no longer take advantage of momentum transfer to amplify the speed of tongue projection. To feed on prey at different azimuthal locations, tomato frogs switch between alternative strategies to circumvent these biomechanical constraints.     

Ecological effects of longline fishing and climate change on the pelagic ecosystem off eastern Australia

Pelagic longline fisheries target (or catch incidently) large apex predators in the open ocean (e.g. tunas, billfish and sharks) and have the potential to disrupt the ecosystem functionality if these predators exert strong top–down control. In contrast, warming of oceans from climate change may increase bottom–up effects from increases in primary productivity. An ecosystem model of a large pelagic ecosystem off eastern Australia was constructed to explore the potential ecological effects of climate change and longlining by Australia’s Eastern Tuna and Billfish Fishery. The model reproduced historic biomass and fishery catch trends from 1952 to 2006 for seven functional groups. Simulated changes in fishing effort and fishing mortality rate on individual target species from 2008 to 2018 resulted in only modest (<20%) changes in the biomass of target species and their direct predators or competitors. A simulated increase in phytoplankton biomass due to climate change resulted in only small increases (<11%) in the biomass of all groups. However, climate-related changes to the biomass of micronekton fish (−20%) and cephalopods (+50%) resulted in trophic cascades. Our results suggest there may be ecological redundancy among high trophic level predators since they share a diverse suite of prey and collectively only represent <1% of the total system biomass. In contrast, micronekton fishes and cephalopods have high biomasses and high production and consumption rates and are important as both prey and predators. They appear to exert ‘wasp–waist’ control of the ecosystem rather than top–down or bottom–up processes reported to drive other pelagic systems.     

A linear canal-otolith interaction model to describe the human vestibulo-ocular reflex

A control systems model of the vestibulo-ocular reflex (VOR) originally derived for yaw rotation about an eccentric axis (Crane et al. 1997) was applied to data collected during ambulation and dynamic posturography. The model incorporates a linear summation of an otolith response due to head translation scaled by target distance, adding to a semi-circular canal response that depends only on angular head rotation. The results of the model were compared with human experimental data by supplying head angular velocity as determined by magnetic search coil recording as the input for the canal branch of the model and supplying linear acceleration as determined by flux gate magnetometer measurements of otolith position. The model was fit to data by determining otolith weighting that enabled the model to best fit the data. We fit to the model experimental data from normal subjects who were: standing quietly, walking, running, or making active sinusoidal head movements. We also fit data obtained during dynamic posturography tasks of: standing on a platform sliding in a horizontal plane at 0.2 Hz, standing directly on a platform tilting at 0.1 Hz, and standing on the tilting platform buffered by a 5-cm thick foam rubber cushion. Each task was done with the subject attending a target approximately 500, 100, or 50 cm distant, both in light and darkness. The model accurately predicted the observed VOR response during each test. Greater otolith weighting was required for near targets for nearly all activities, consistent with weights for the otolith component found in previous studies employing imposed rotations. The only exceptions were for vertical axis motion during standing, sliding, and tilting when the platform was buffered with foam rubber. In the horizontal axis, the model always fit near target data better with a higher otolith component. Otolith weights were similar with the target visible and in darkness. The model predicts eye movement during both passive whole-body rotation and free head movement in space implying that the VOR is controlled by a similar mechanism during both situations. Factors such as vision, proprioception, and efference copy that are available during head free motion but not during whole-body rotation are probably not important to gaze stabilization during ambulation and postural stabilizing movement. The linearity of the canal-otolith interaction was tested by re-analysis of the whole body rotation data on which the model is based (Crane et al. 1997). Normalized otolith-mediated gain enhancement was determined for each axis of rotation. This analysis uncovered minor non-linearities in the canal-otolith interaction at frequencies above 1.6 Hz and when the axis of rotation was posterior to the head. Received: 11 March 1998 / Received in revised form: 1 March 1999     

Sequence variation in the monoclonal-antibody-U36-defined CD44v6 epitope

 Monoclonal antibody (mAb) U36 was developed for the treatment of minimal residual disease of head and neck squamous cell carcinoma (HNSCC). The mAb-U36-defined antigen was characterized by cDNA cloning, and was shown to be identical to the keratinocyte-specific CD44 splice variant epican. The epitope recognized by mAb U36 was shown to be located in the v6 domain. Two amino acids within the epitope appeared to differ from the sequences that have been described in literature. The sequence of the epitope appeared to contain glutamic acid at position 367 and lysine at position 374, while valine and arginine respectively have been described before. Interestingly, another anti-CD44v6 antibody with possible clinical application, VFF18, recognizes an epitope in the same area. With respect to the applicability of these antibodies for tumor targeting, this variation might have an influence on antibody-antigen interaction and mAb accumulation in the tumor. Furthermore, this observation raised the question whether the different epitopes are related to the malignant behavior of tumor cells. In this paper we determine the relative affinity of mAb U36 for the variant epitope sequences by tumor cell binding assays using synthetic peptides for competition. The presence of glutamic acid instead of valine at position 367 caused strong competition. Further evaluation showed that the published valine variant does not exist in vivo, and is the result of a sequencing artefact. The effect of substitution of lysine for arginine at position 374 had no effect on the binding of mAb U36 to the cells. This amino acid variation was shown to be due to allelic polymorphism. There was no trend towards allelic imbalance in tumor cells as compared to normal cells. Received: 5 June 1997 / Accepted: 14 August 1997     

Extinction of top-predator in a three-level food-chain model

 In this paper we extend the Lyapunov functions, constructed by A. Ardito and P. Ricciardi for predator–prey system [1], to the three level food chain models. We first consider a general three-level food-chain model. A criterion for the extinction of top predator will be given. Then we restrict our attentions to the case in which the prey is of logistic growth and predators have Holling’s type II functional responses. Received: 10 October 1997