Frederick R. Prete

First Identification of a Putative Sex Pheromone in a Praying Mantid

Praying mantids are models for a wide variety of behavioral, physiological, and ecological studies, and sex pheromones have been assumed to be important components of their biology. However, no mantid pheromone has ever been identified. We collected volatiles emitted by females of the mantid, Sphodromantis lineola, via solid phase microextraction (SPME). Mass spectral analysis revealed the collected volatiles to be a mixture of pentadecanal and tetradecanal. We prepared a synthetic mixture of these compounds, and found that males were both attracted to this mixture and stimulated to exhibit typical precopulatory behavior. We then examined male antennae with scanning electron microscopy, and confirmed the presence of porous antennal sensilla typical of insect pheromone receptors, i.e., that male mantids are equipped with the appropriate morphological apparatus to receive volatile chemical signals. Pheromones, in conjunction with visual and tactile cues, are thus an important feature of the reproductive biology of this, and undoubtedly other species of mantids. In addition to adding a crucial aspect of behavioral biology to our knowledge of this group, identification and synthesis of mantid pheromones may be a first step in attracting and aggregating these generalist predators for use in pest control.

Responses of Movement-Sensitive Visual Interneurons to Prey-Like Stimuli in the Praying Mantis Sphodromantis lineola (Burmeister)

Previous behavioral work using both mechanical and computer-generated visual stimuli has demonstrated that mantids use a computational algorithm to recognize prey similar to that used by some amphibian predators: A stimulus elicits prey capture behavior if it falls within a perceptual envelope defined by five fundamental stimulus parameters: (1) overall size, (2) length of the leading edge, (3) contrast to the background, (4) location in the visual field, and (5) apparent speed. In this study, we recorded simultaneously from both cervical nerve cords of monocular Sphodromantis lineola while they viewed the same visual stimuli successfully used in the behavioral studies. Extracellular recordings showed three consistently proportioned amplitude classes of movement-elicited spikes in each cord and these were repeatedly and reliably identifiable across mantids. Overall, the movement-elicited activity in both cords was dominated by very large spikes suggesting the existence of several large, descending movement-sensitive interneurons projecting both ipsilaterally and contralaterally from the optic lobes. However, only the largest contralateral spikes occurred preferentially to prey-like stimuli, mirrored the behavioral response curves generated by S. lineola to the same visual stimuli, and displayed activity peaks that were correlated with the times at which the mantid emitted predatory strikes.

Visual stimulus characteristics that elicit tracking and striking in the praying mantises Parasphendale affinis, Popa spurca and Sphodromantis lineola.

SUMMARY We tested three species of praying mantis, Parasphendale affinis, Popa spurca and Sphodromantis lineola, with computer-generated stimuli that differed in size, contrast, configuration and movement pattern to determine the effects of these parameters on visual tracking and striking behavior. Overall, black disks moving erratically against a white background were strong releasers of both behaviors. When stimulus presentation order was randomized by size, P. affinis and P. spurca struck at progressively higher rates as the stimuli enlarged up to 44 deg; S. lineola struck most at intermediate sized (10–20 deg) disks. When disks were size-ordered from small to large, P. affinis and S. lineola struck at the smaller disks at higher rates; however, when the order was reversed, the early appearance of large disks suppressed subsequent responses to the smaller disks. Stimulus order did not differentially affect the responses of P. spurca. All species responded at higher rates to black disks moving against a white background versus the reverse. However, only P. spurca and S. lineola responded at higher rates to relatively darker grey disks, only P. affinis responded to mottled grey disks moving against an identically patterned background, and only P. spurca struck more frequently in response to rectangular stimuli oriented parallel (versus orthogonal) to their direction of movement. In conjunction with data on other species, these results support the hypothesis that praying mantises recognize prey based on assessment of several category-specific, spatiotemporal features, e.g. size, contrast, speed, movement pattern and leading edge length.

Visual stimuli that elicit visual tracking, approaching and striking behavior from an unusual praying mantis, Euchomenella macrops (Insecta: Mantodea).

In comparison to other similarly sized mantis species examined in previous studies, Euchomenella macrops has a significantly smaller head, shorter foreleg tibia, but longer prothorax which have been interpreted as specializations for the capture of smaller, slower prey. We tested this conjecture by assessing the rates at which computer generated stimuli elicit visual tracking, approaching, and striking behaviors by adult females. When presented with black disks moving erratically against a white background, strike rate rose progressively as disks enlarged up to 44 deg (visual angle) if the disks moved rapidly (e.g., 143 deg/s); at slower speeds (113, 127 deg/s), smaller disks (<27 deg) were preferred. When black moved linearly from the visual periphery to visual field center (at 73 or 143 deg/s) and then stopped, E. macrops struck consistently at disks as small as 5 deg after movement ceased. E. macrops also struck at higher rates in response to 23 deg erratically moving (subjective) red (versus subjective blue or green) disks that were luminance matched to a grey background although they tracked all colors at equally high rates. Unlike some other species, E. macrops did not strike at higher rates in response to elongated rectangular stimuli moving parallel (versus perpendicular) to their long axis, although the former elicited higher rates of approaching. An analysis of tracking behavior revealed that virtually all tracking movements were a result of head (versus) prothorax rotation.

Thoracic and prothoracic leg neuromuscular system of the praying mantid, Sphodromantis lineola (burmeister)

Historically, praying mantids have attracted attention because of their dramatic prey capture behavior, loosely termed the strike. However, little is known about the neuromuscular organization that underpins the behavior. Although once thought to be quite stereotyped, recent data indicate that the strike is quite plastic and can be aimed accurately within a relatively large three‐dimensional space. Hence, successful prey capture requires the integration of (1) visual information, indicating prey has been recognized; and (2) proprioceptive information, indicating head and prothorax (i.e., visual field) position and initial leg positions. This study was undertaken as part of a larger program examining how such sensory information is integrated with the appropriate motor systems. Our goals were (1) to describe the gross thoracic and foreleg neuromuscular system of Sphodromantis lineola and (2) to identify the soma locations of the motor neurons associated with the largest leg nerve, N4, which travels the length of each leg. We found that the thoracic and foreleg neuromusculature of S. lineola are similar but not identical to what is known about just three other species of mantid, and that motor neuron somata associated with N4 are arranged in stereotypical, bilaterally symmetrical groups as they are in other orthopteroids, suggesting that this is a general organizational feature of the insect CNS. J. Comp. Neurol. 409:325–338, 1999.

Circadian rhythms affect electroretinogram, compound eye color, striking behavior and locomotion of the praying mantis Hierodula patellifera

Many behaviors and physiological processes oscillate with circadian rhythms that are synchronized to environmental cues (e.g. light onset), but persist with periods of ~24 h in the absence of such cues. We used a multilevel experimental approach to assess whether circadian rhythms modulate several aspects of the visual physiology and behavior of the praying mantis Hierodula patellifera. We used electroretinograms (ERGs) to assess compound eye sensitivity, colorimetric photographic analyses to assess compound eye color changes (screening pigment migration), behavioral assays of responsiveness to computer-generated prey-like visual stimuli and analyses of locomotor activity patterns on a modified treadmill apparatus. Our results indicate that circadian clocks control and/or modulate each of the target behaviors. Strong rhythms, persisting under constant conditions, with periods of ~24 h were evident in photoreceptor sensitivity to light, appetitive responsiveness to prey-like stimuli and gross locomotor activity. In the first two cases, responsiveness was highest during the subjective night and lowest during the subjective day. Locomotor activity was strongly clustered around the transition time from day to night. In addition, pigment migration and locomotor behavior responded strongly to light:dark cycles and anticipated the light–dark transition, suggesting that the circadian clocks modulating both were entrained to environmental light cues. Together, these data indicate that circadian rhythms operate at the cellular, cellular systems and organismal level in H. patellifera. Our results represent an intriguing first step in uncovering the complexities of circadian rhythms in the Mantodea.