Defence behaviours of the praying mantis Tenodera aridifolia in response to looming objects

Defence responses to approaching objects were observed in the mantis Tenodera aridifolia. The mantis showed three kinds of behaviour, fixation, evasion and cryptic reaction. The cryptic reaction consisted of rapid retraction of the forelegs under the prothorax or rapid extending of the forelegs in the forward direction. Obstructing the mantis’ sight decreased its response rates, suggesting that the visual stimuli generated by an approaching object elicited the cryptic reaction. The response rate of the cryptic reactions was highest for objects that approached on a direct collision course. Deviation in a horizontal direction from the direct collision course resulted in a reduced response. The response rate of the cryptic reaction increased as the approaching velocity of the object increased, and the rate decreased as the object ceased its approach at a greater distance from the mantis. These results suggest that the function of the observed cryptic reactions is defence against impending collisions. The possible role of the looming-sensitive neuron in the cryptic reaction is also discussed.     

Investigating saccade programming in the praying mantis Tenodera aridifolia using distracter interference paradigms

To investigate the saccadic system in the mantis, I applied distracter interference paradigms. These involved presenting the mantis with a fixation target and one or several distracters supposed to affect saccades towards the target. When a single target was presented, a medium-sized target located in its lower visual field elicited higher rates of saccade response. This preference for target size and position was also observed when a target and a distracter were presented simultaneously. That is, the mantis chose and fixated the target rather than a distracter that was much smaller or larger than the target, or was located above the target. Furthermore, the mantis' preference was not affected by increasing the number of distracters. However, the presence of the distracter decreased the occurrence rate of saccade and increased the response time to saccade. I conclude that distracter interference paradigms are an effective way of investigating the visual processing underlying saccade generation in the mantis. Possible mechanisms of saccade generation in the mantis are discussed.     

Coordinated movements of the head and body during orienting behaviour in the praying mantis Tenodera aridifolia

The visual orienting behaviour towards prey in the free-moving mantis was investigated with a high-speed camera. The orienting behaviour consisted of head, prothorax, and abdomen rotations. Coordinated movements of these body parts in the horizontal plane were analysed frame-by-frame. Rotations of these body parts were initiated with no or slight (≤40 ms) differences in timing. The initiation timing of prothorax-abdomen rotation was affected by its initial angle before the onset of orienting. There were positive correlations in amplitude among head-prothorax, prothorax-abdomen, and abdomen rotations. The ratio of these rotations to total gaze rotation was affected by the initial prothorax-abdomen angle before the onset of orienting. Our data suggest that coordinated movements of the head, prothorax, and abdomen during orienting are ballistic events and are pre-determined according to visual and proprioceptive information before the onset of orienting.     

Predatory behavior of the praying mantis,Tenodera aridifolia I. Effect of prey size on prey recognition

Predatory behavior and effect of prey size of the praying mantis,Tenodera aridifolia, was investigated using three-dimensional prey models. Parameters of prey such as width, length, shape (length/width), area (width×length) and volume were investigated for effects on Watch and Attack responses. Regression analyses suggest that volume is the most important variable in prey recognition. This behavioral trait is considered adaptive, since prey volume is the most reliable measurement for the energy content of the prey.     

Spectral sensitivity studies on the visual system of the praying mantis, Tenodera sinensis

In these studies a constant ERG response was used as a measure of visual sensitivity to different wavelengths of light. The dark-adapted compound eye of Tenodera sinensis is dominated by a single class of photoreceptors. with a major peak of sensitivity at about 510–520 nm, and with a minor peak of sensitivity in the near-ultraviolet region at about 370 nm. The dark-adapted dorsal ocellus does not contain a homogeneous population of sensory receptors. The sensitivity function of the dark-adapted ocellus to longer wavelength light (yellow and red) is determined by a single receptor with a major peak of sensitivity in the green at 510–520 nm with some sensitivity in the near-ultraviolet. Sensitivity at shorter wavelengths (near-ultraviolet and blue), however, involves the stimulation of both this and a near-ultraviolet-sensitive receptor with a maximum sensitivity at about 370 nm. Anatomically, the sensory cells of the dorsal ocellus of Tenodera were determined histologically to be grouped into two distinct regions, each group making its own separate contribution to the ocellar nerve. This may represent the separation of two different photoreceptor types in the ocellus of the mantis.     

Responses to non-locomotive prey models by the praying mantis,Tenodera angustipennis Saussure

Adult females of the praying mantisTenodera angustipennis were presented with computer-generated images, and the attractiveness of “non-locomotive” prey models was examined. Mantises fixated and struck the “body and leg” model (consisting of an immobile black square on a white background with 2 black lines oscillating randomly at its sides) more frequently than the “leg” model (only oscillating lines) or the “body” model (static square only). This indicates that the model consisting of a static object and moving lines effectively elicits mantis strike behavior, although it is “non-locomotive.”     

Saccadic tracking of a light grey target in the mantis,Tenodera aridifolia

I presented a horizontally moving square on a computer display to the mantis, Tenodera aridifolia, and examined the effects of target brightness and velocity, and background brightness on its tracking behavior. The mantis tracked a light grey square with more saccadic head movements than a black square, although these squares moved on a homogeneous background. The amplitude of saccades was larger when the light grey square moved at a lower velocity. The background brightness had little effect on the type (smooth or saccadic) of tracking behavior. These results suggest that the saccadic tracking of light grey objects on a homogeneous background may not be caused by low contrast, i.e., the difficulty in discriminating the object from the background. The possible biological significance of saccadic tracking on a homogenous background is discussed.     

The antennal sensilla of the praying mantis Tenodera aridifolia: A new flagellar partition based on the antennal macro-, micro- and ultrastructures

In insects, the antenna consists of a scapus, a pedicellus, and a flagellum comprising many segments (flagellomeres). These segments possess many morphological types of sensory organs (sensilla) to process multimodal sensory information. We observed the sensilla on flagellomeres in praying mantis (Tenodera aridifolia) with both scanning and transmission electron microscopes. We classified the sensilla into six types: chaetic, campaniform, coelocapitular, basiconic, trichoid and grooved peg sensilla, and inferred their presumptive functions on the basis of their external and internal structures. In addition, based on their distribution, we newly divided the flagellum into 6 distinct parts. This new division leads to a better understanding about the sexual dimorphism and the antennal development in the mantises. The sexual difference in distribution of the grooved peg sensilla suggests that this type of sensilla may play a role in sex-pheromone detection in mantis, which is a rare case of double-walled sensilla mediating this function.     

Predatory behavior of the praying mantis,Tenodera aridifolia II. Combined effect of prey size and predator size on the prey recognition

Predatory behavior of the praying mantis,Tenodera aridifolia, as a function of the combined effect of its size and the size of the prey was investigated by using prey models. Behavioral responses were almost identical through the nymphal development in the predator. As the mantis grew, it attacked larger prey models, suggesting that it recognizes the prey's size in accordance with its own body size. Regression analyses demonstrate that the ratio of the prey's volume to the cube and the square of the predator's length is a more important parameter for prey recognition than are the one-dimensional parameters of the prey's and the predator's sizes.     

Enzymic activities involved in the oothecal sclerotization of the praying mantid,Tenodera aridifolia sinensis saussure

The enzyme(s) responsible for the sclerotization of mantid ootheca is secreted by the left colleterial gland. From an extract of the glands of Tenodera aridifolia sinensis, two soluble enzyme fractions of different activities were obtained. One fraction acted on N-acetyldopamine (NADA), a precursor of a representative sclerotizing agent, and produced NADA-quinone. The other did not act on NADA itself but converted the quinone to a highly reactive intermediate, such as quinone methide, which was able to react nonenzymically with nucleophilic compounds. Other insoluble enzyme preparations obtained from the silk and pupal cuticle of the Japanese giant silk moth, Dictyoploca japonica, also had these two activities.     

Responses to computer-generated visual stimuli by the male praying mantis, Sphodromantis lineola (Burmeister)

We tested male mantis' responses to square, computer-generated visual stimuli of various sizes to determine the stimulus parameters that affect their striking behaviour. Males in a high-hunger group displayed a sharp preference for solid black 12×12-degree squares moving linearly against a white background, especially when the squares moved downward (versus horizontally). Males in a low-hunger group visually tracked but, with rare exception, did not strike at any linearly moving squares. In contrast, when solid black squares moved erratically around visual field centre, males struck regularly at them irrespective of hunger level, even when the squares were as large as 47×47 degrees. Males also recognized black-and-white Julesz-patterned square stimuli moving against a similarly patterned background, indicating that they recognized the synchronous movement of a group of stimulus elements as a single moving object. Finally, we compared allometric and life history data between male and females S. lineola to elucidate the possible reasons for the differences between male response patterns and previously published data on females. These comparisons suggest that males employ a different behavioural strategy than do females when faced with large, erratically moving visual stimuli.     

Responses of descending neurons to looming stimuli in the praying mantis <Emphasis Type="Italic">Tenodera aridifolia</Emphasis>

Responses to visual stimuli of some neurons that descend the nerve cord from the brain were recorded extracellularly in the mantis Tenodera aridifolia. Most of the recorded neurons showed their largest responses to looming stimuli that simulated a black circle approaching towards the mantis. The neurons showed a transient excitatory response to a gradually darkening or receding circle. The neurons showed sustained excitation to the linearly expanding stimuli, but the spike frequency decreased rapidly. The responses of the neurons were affected by both the diameter and the speed of looming stimuli. Faster or smaller looming stimuli elicited a higher peak frequency. These responses were observed in both recordings from the connective between suboesophageal and prothoracic ganglia and the connective between prothoracic and mesothoracic ganglia. There was a one-to-one correspondence of spike firing between these two recordings with a fixed delay. The neurons had the receptive field on ipsilateral side to its axon at the cervical connective. These results suggest that there is a looming-sensitive descending neuron, with an axon projecting over prothoracic ganglion, in the mantis nervous system.     

Saccadic head movements of the praying mantis, with particular reference to visual and proprioceptive information

ABSTRACT. Horizontal head movements of the praying mantis, Sphodromantis lineola Burm., were recorded continuously. They responded to the presence of a live blowfly prey in the antero-lateral visual field with a rapid saccadic head movement. The angular movement of a fixation saccade was correlated positively to the displacement of the prey from the prothoracic midline. Saccade magnitude and velocity are related. After the stimulus moved out of the visual field, the mantis made a second saccadic head movement, a return saccade towards the body midline. We observed return saccades in which the head overshot or undershot the body midline, as well as saccades which returned the head exactly to its initial position. In 92% of trials with intact mantids, the return movement succeeded eventually in rotating the head back to its initial position, whereas after removal of the neck hair plates this occurred in only 47% of trials. There is a consistent relation between saccade extent and velocity. Velocities of return saccades were slower than those of fixation saccades. It is suggested that sensory inputs from the neck hair plate proprioceptors modify both the magnitude and the angular velocity of fixation and return saccadic head movements.     

Mantisbot is a robotic model of visually guided motion in the praying mantis

Insects use highly distributed nervous systems to process exteroception from head sensors, compare that information with state-based goals, and direct posture or locomotion toward those goals. To study how descending commands from brain centers produce coordinated, goal-directed motion in distributed nervous systems, we have constructed a conductance-based neural system for our robot MantisBot, a 29 degree-of-freedom, 13.3:1 scale praying mantis robot. Using the literature on mantis prey tracking and insect locomotion, we designed a hierarchical, distributed neural controller that establishes the goal, coordinates different joints, and executes prey-tracking motion. In our controller, brain networks perceive the location of prey and predict its future location, store this location in memory, and formulate descending commands for ballistic saccades like those seen in the animal. The descending commands are simple, indicating only 1) whether the robot should walk or stand still, and 2) the intended direction of motion. Each joint's controller uses the descending commands differently to alter sensory-motor interactions, changing the sensory pathways that coordinate the joints' central pattern generators into one cohesive motion. Experiments with one leg of MantisBot show that visual input produces simple descending commands that alter walking kinematics, change the walking direction in a predictable manner, enact reflex reversals when necessary, and can control both static posture and locomotion with the same network.     

Purification and characterization of a serine protease with fibrinolytic activity from Tenodera sinensis (praying mantis)

Mantis egg fibrolase (MEF) was purified from the egg cases of Tenodera sinensis using ammonium sulfate fractionation, gel filtration on Bio-Gel P-60 and affinity chromatography on DEAE Affi-Gel blue gel. The protease was assessed homogeneous by SDS-polyacrylamide gel electrophoresis and has a molecular mass of 31500 Da. An isoelectric point of 6.1 was determined by isoelectric focusing. Amino acid sequencing of the N-terminal region established a primary structure composed of Ala-Asp-Val-Val-Gln-Gly-Asp-Ala-Pro-Ser. MEF readily digested the Aalpha- and Bbeta-chains of fibrinogen and more slowly the gamma-chain. The nonspecific action of the enzyme results in extensive hydrolysis of fibrinogen and fibrin releasing a variety of fibrinopeptide. The enzyme is inactivated by Cu2+ and Zn2+ and inhibited by PMSF and chymostatin, yet elastinal, aprotinin, TLCK, TPCK, EDTA, EGTA, cysteine, beta-mercaptoethanol, iodoacetate, E64, benzamidine and soybean trypsin inhibitor do not affect activity. Antiplasmin was not sensitive to MEF but antithrombin III inhibited the enzymatic activity of MEF. Among chromogenic protease substrates, the most sensitive to MEF hydrolysis was benzoyl-Phe-Val-Arg-p-nitroanilide with maximal activity at pH 7.0 and 30 degrees C. MEF preferentially cleaved the oxidized B-chain of insulin between Leu15 and Tyr16. D-Dimer concentrations increased on incubation of cross-linked fibrin with MEF, indicating the enzyme has a strong fibrinolytic activity.     

Visual stimuli that elicit appetitive behaviors in three morphologically distinct species of praying mantis

We assessed the differences in appetitive responses to visual stimuli by three species of praying mantis (Insecta: Mantodea), Tenodera aridifolia sinensis, Mantis religiosa, and Cilnia humeralis. Tethered, adult females watched computer generated stimuli (erratically moving disks or linearly moving rectangles) that varied along predetermined parameters. Three responses were scored: tracking, approaching, and striking. Threshold stimulus size (diameter) for tracking and striking at disks ranged from 3.5 deg (C. humeralis) to 7.8 deg (M. religiosa), and from 3.3 deg (C. humeralis) to 11.7 deg (M. religiosa), respectively. Unlike the other species which struck at disks as large as 44 deg, T. a. sinensis displayed a preference for 14 deg disks. Disks moving at 143 deg/s were preferred by all species. M. religiosa exhibited the most approaching behavior, and with T. a. sinensis distinguished between rectangular stimuli moving parallel versus perpendicular to their long axes. C. humeralis did not make this distinction. Stimulus sizes that elicited the target behaviors were not related to mantis size. However, differences in compound eye morphology may be related to species differences: C. humeralis’ eyes are farthest apart, and it has an apparently narrower binocular visual field which may affect retinal inputs to movement-sensitive visual interneurons.     

Head grooming behaviour in the praying mantis

Mantis head grooming is a rhythmic behaviour pattern involving co-ordinated movements of head and foreleg. The probability of grooming is influenced by releaser concentration, while the number of cycles/episode is influenced by concentration and amount of releaser. Within a grooming episode (usually 4 to 6 cycles) successive cycles are longer in duration; this is primarily caused by an increase in the actual head cleaning phase, although all three phases increase in duration. The phase cleaning the femur brush is most sensitive to feedback effects from releaser substances. It is concluded that head grooming is a programmed, i.e. centrally organized, movement pattern responsive to peripheral feedback. First approximations to the model-equations and model-hypotheses are presented.