The role of antennal hair plates in object-guided tactile orientation of the cockroach (Periplaneta americana)

The searching behavior of blinded cockroaches was examined under unrestrained conditions, in an arena, and on a treadmill. When cockroaches searching in a circular arena touched a stationary object (metal pole) with their antennae, they frequently approached the object more closely, and then climbed up it. Similar orientation behavior was observed in tethered animals in open loop conditions, walking on a Styrofoam ball. In these restrained cockroaches, a single antenna sufficed to distinguish the angular positions of an object, in the horizontal plane (0 degrees, 45 degrees, and 90 degrees). A group of mechanosensitive hairs on the basal segment of the antenna (scapal hair plate) appears to play a major role in antennal object detection in the horizontal plane, as gauged by shaving off these scapal hair plates. In unrestrained cockroaches, shaving the scapal hair plate increased the time needed to approach an object. Under tethered conditions, the ability to turn towards and to establish antennal contact with a test object was significantly impaired.     

Mechanosensory control of antennal movement by the scapal hair plate in the American cockroach

The voluntary movement of antennae of blinded cockroaches was examined in the tethered-walking condition. An object of metal plate was presented to a tip of a single antenna in order to induce tactile orientation behavior. Horizontal movements of the antenna before and during the object presentation were analyzed both before and after ablation of a mechanosensory organ, the scapal hair plate (S-HP), at the base of antenna. The resting antennal position shifted outwardly by about 20 degrees after the S-HP ablation. Spontaneous antennal movements in ablated animals became stiff and wider ranged. The tactile object was set at two different horizontal positions, 45 degrees and 90 degrees clockwise to the head, for the right side antenna. The number of contacts in a constant test period was significantly decreased in the tests at 45 degrees after ablation. Trajectories of antennal movements before and after contacts were categorized into four patterns. In the case that an antenna made contact with the object during its abduction (outward) movement, it then passed the object outwardly or withdrew inwardly. These were termed "outward-pass (O-P)" and "outward-withdrawal (O-W)" patterns, respectively. Similarly, contacts during the adduction (inward) movement were divided into "inward-pass (I-P)" or "inward-withdrawal (I-W)" pattern. Significant effects of the S-HP ablation appeared in the tests at 90 degrees : the I-P pattern mostly disappeared and was replaced by the I-W pattern. The results strongly suggest that the S-HP has crucial roles for controlling both spontaneous and stimulated movements of the cockroach antenna.     

Directionality of antennal sweeps elicited by water jet stimulation of the tailfan in the crayfish Procambarus clarkii

Summary Directionality and intensity dependence of antennal sweeps elicited by water jet stimulation of the tailfan in tethered, reversibly blinded adult and juvenile crayfish (Procambarus clarkii) were analyzed.Resting crayfish keep their antennae at about 50° symmetrically to the longitudinal body axis (Figs. 2 bottom, and 3).In adults, tailfan stimulation elicits synchronous backward sweeps of both antennae, which increase for more caudal stimulus directions (Figs. 2–4 and 5A). Directions differing by 30°–60° are significantly distinguished (Fig. 4). The mean sweep of the ipsilateral antenna significantly overrides that of the contralateral antenna for rostrolateral stimulation at 40–200 mm/s stimulus velocity and lateral to caudolateral stimulation at 40 mm/s and thus lateralization of the stimulus is revealed (Figs. 2 top, 4 and 5A). Mean antennal sweeps at a given stimulus direction and distance increase with increasing stimulus velocity (40–250 mm/s, Fig. 5A).In juveniles, the directional dependence of antennal sweeps is reduced compared to that of adults, while a similar intensity dependence is found (Fig. 5B).The pronounced directionality of the antennal response in adult crayfish vanishes and response latencies increase after reversibly covering the tailfan with a small bag or the telson with waterproof paste (Figs. 6 and 7). Thus, tailfan and especially telson mechanoreceptors play an important role in the localization of water movements elicited by predators or prey behind the crayfish.     

Tactile motor learning in the antennal system of the honeybee (Apis mellifera L.)

Honeybees fixed in small tubes scan an object within the range of the antennae by touching it briefly and frequently. In our experiments the animals were able to scan an object for several minutes with the antennae. After moving the object out of the range of the antennae, the animals showed antennal movements for several minutes that were correlated with the position of the removed object. These changes of antennal movements are called “behavioural plasticity” and are interpreted as a form of motor learning. Bees showed behavioural plasticity only for objects with relatively large surfaces. Plasticity was more pronounced in bees whose compound eyes were occluded. Behavioural plasticity was related to the duration of object presentation. Repeated presentations of the object increased the degree of plasticity. After presentation durations of 30 min the animals showed a significant increase of antennal positions related to the surface of the object and avoidance of areas corresponding to the edges. Behavioural plasticity was compared with reward-dependent learning by conditioning bees to objects. The results of motor learning and reward-dependent conditioning suggest that bees have tactile spatial memory. Accepted: 13 May 1997     

The antennal system and cockroach evasive behavior. II. Stimulus identification and localization are separable antennal functions

Cockroaches ( Periplaneta americana) orient their antennae toward moving objects based on visual cues. Presumably, this allows exploration of novel objects by the antennal flagellum. We used videographic and electrophysiological methods to determine if receptors on the flagellum are essential for triggering escape, or if they enable cockroaches to discriminate threatening from non-threatening objects that are encountered. When a flagellum was removed, and replaced with a plastic fiber, deflection of a "prosthetic flagellum" still activated the descending mechanosensory interneurons associated with escape and produced typical escape responses. However, escape was essentially eliminated by constraining the movement of the scape and pedicel at the antennal base. When cockroaches approached and briefly explored the surface of a spider or another cockroach with the flagellum, they produced escape significantly more often in response to subsequent controlled contact from a spider than from a cockroach. This discrimination did not depend on visual or wind-sensory input, but required flagellar palpation of the surface. The crucial sensory cues appear to involve texture rather than surface chemicals. These results indicate that cockroaches acquire basic information on stimulus identity during exploration of surfaces with flagellar receptors, but that basal receptors are triggers for escape behavior.     

Antennal movements induced by odour and central projection of the antennal neurones in the honey-bee

Movements of the antennae induced by odour were investigated. Odour presented to the antenna of one side induced both antennae to move to that side. The EMGs recorded from the flexor muscles of both scapes showed that the latency of the movement of the ipsilateral flagellum when induced by odour was about 71 msec shorter than that of the contralateral flagellum. Recording electrical activities from the antennal nerve showed that there are more than 14 neurones in the antenno-motor externus.The distribution of the antennal nerve in the brain was investigated histologically by the injection of fluorescent dye. Antennal sensory neurones terminated at the glomeruli in the antennal lobe, in the dorsal lobe, in the protocerebrum, and in the commissural part of the suboesophageal ganglion. Injection of the fluorescent dye into the antennal nerve after degeneration of the antennal sensory neurones showed that the antennal motoneurones run in the ventral side of the antennal and dorsal lobes, and terminate in the marginal region of the ipsilateral oesophageal connective.The difference in latency of odour-induced flagellar movements is discussed in relation to the histological results and the unitary responses in the brain reported previously.     

Electrophysiological identification of thermo- and hygro-sensitive receptor neurons on the antennae of the dragonfly Libellula depressa

Recent ultrastructural investigations on Odonata antennal flagellum describe two types of sensilla styloconica, T1 and T2. The styloconic sensilla are located in pits, at the bottom of deep cavities, and share common features typical of thermo-hygroreceptors. In order to ascertain if the Odonata antennae are involved in hygroreception and thermoreception, we carried out electrophysiological recordings (single cell recordings, SCR) from adult males and females of Libellula depressa L., 1758. After contact was established, the antenna was stimulated by rapid changes in temperature and humidity. The present research shows the occurrence of a dry (DC), a moist (MC) and a cold (CC) receptor neurons on the antennal flagellum of L. depressa. These data demonstrate for the first time the presence of functional thermo-hygroreceptors on the antennal flagellum of dragonflies. The present results extend our knowledge of the not visual sensory modalities of Odonata, a field of research unexplored so far.     

Bilateral and unilateral antennal lesions alter orientation abilities of the crayfish, Orconectes rusticus.

Numerous animals use chemical cues within their environments to execute various behaviors. One of these behaviors is orientation to an odor source. Crayfish, in particular, can orient to food sources under a number of different conditions. It has not been determined, however, what kind of search strategy these animals employ to successfully locate a food source. To determine the role of antennae and antennules in this behavior and to investigate different modes of orientation behavior, the orientation patterns of crayfish with complete and partial antennal lesions were examined. Detailed analysis of orientation paths confirmed that crayfish could not locate odor sources with either bilateral or unilateral lesions. This suggests that crayfish are using the spatial information obtained from these appendages to successfully orient. Animals using information from the bilaterally paired appendages in the control group exhibited increased walking speed, increased speed to source and decreased heading angles towards the source compared to these measurements taken from lesioned groups. There was no significant difference in any parameters between animals with unilateral or bilateral lesions. This strongly suggests that these animals are reliant on the spatial comparison of differences between bilaterally paired olfactory appendages for successful orientation.     

Antennal sense organs of the cockroach, Leucophaea maderae

Adult and nymphal antennae of the cockroach, Leucophaea maderae, contain nine or more different morphological types of sense organs. There is no outwardly apparent sexual dimorphism in adult antennae. Nymphs are dificient in gross numbers of sensilla. Sense organs are classified morphologically by their similarity to known types of sensila and are assigned functions on this basis and preliminary electrophysiological data: Sensilla chaetica (A), thick-walled mechanoreceptive hairs in groups on the antennal base; S. chaetica (B), thick-walled setae which are tactile and probably chemoreceptive, occurring in the antennal base and flagellum; S. trichodea (A), thin-walled chemoreceptive hairs of the flagellum; S. trichodea (B), minute hairs on the scape and pedicel; S. basiconica, thin-walled chemoreceptive pegs, and S. coeloconica (?pit-pegs”?) of the flagellum; S. campaniformia and scolopidia, mechanoreceptors in the base and flagellum; plus Johnston's organ and/or connective chrodotonal organs in the pedicel. Calculations based on absolute counts of sensilla and their known innervation yield an estimate of about 3.3 × 10⁴ sensilla and 10⁵ cells per antenna.     

Active tactile sensing for localization of objects by the cockroach antenna

Antennal movement during tactile orientation behavior was examined three-dimensionally in American cockroaches during tethered walking. When a wooden rod was presented to the tip of one antenna in an upright orientation at one of the three different horizontal positions (30°, 60°, or 90° from the center of the head), the animal touched it repeatedly with the antenna, and tried to approach it (positive thigmotaxis). Positional shifts were also observed for the contralateral unstimulated antenna. The ipsilateral antenna tended to touch the object during inward movement (adduction) at all three test angles. The cumulative turn angle made during a continuous test period of 24 s clearly depended on the object’s position; however, the contact frequencies were almost the same regardless of the position. The relationships between contact frequency and some locomotion parameters were also investigated on a shorter time scale of 3 s. The contact frequency positively correlated with the turn angle, with the accuracy of orientation at all three test angles, and with the translation velocity at test angles of 30° and 60°. It is concluded that the performance during tactile orientation can be represented effectively by the frequency with which the antennae touch the attractive objects.     

椰心叶甲啮小蜂复眼和触角在交配中的作用及其超微结构的扫描电镜观察

对寄生蜂交配行为的了解将有助于发展对其行为调控的技术,提高寄生蜂对害虫控制的效能。为探讨椰心叶甲啮小蜂Tetrastichus brontispae Ferrière的复眼和触角在交配中的作用,用水溶性黑色素和液体石蜡分别涂抹该蜂复眼和触角后观察其交配行为,并利用扫描电镜（SEM）观察其复眼和触角的超微结构,分析雌、雄蜂触角感器的分布与数量差异。结果表明：椰心叶甲啮小蜂雄蜂的复眼在交配过程中起重要作用,雌蜂复眼作用不显著。在椰心叶甲啮小蜂求偶识别和接受过程中,雄蜂触角柄节部位起主要作用,其次是棒节部位,再次是鞭节的索亚节部位,而雌蜂触角鞭节索亚节部位起主要作用,然后是棒节部位,最后是柄节部位。扫描电镜观察表明椰心叶甲啮小蜂触角上共有8种感器,其中毛形感器和板形感器是主要感器,雌、雄蜂触角有明显的性二型现象,表现为触角大小不同及触角感器类型、大小、数量、分布不同。     

Antennal malformations in light ocelli drones of Apis mellifera (Hymenoptera, Apidae)

Malformed antennae of Apis mellifera light ocelli drones were drawn, dissected and mounted permanently on slides containing Canada balsam, in order to count the olfactory discs present in each segment, in comparison with the number of those structures in normal antennae of their brothers. Some drones presented morphological abnormalities in a single segment of the right or left antenna, but others had two or more malformed segments in a same antenna. Drones with malformations in both antennae were also observed. The 4th and 5th flagellum segments were the most frequently affected. In a low number of cases the frequency of olfactory discs in malformed segments did not differ from that one recorded for normal segments. However, in most cases studied, the antennal malformations brought about a significant reduction in the number of olfactory discs from malformed segments.     

Structure and development of antennae in a moth,Manduca sexta

The antenna of the moth, Manduca sexta, comprises two small basal segments and a long (2 cm) flagellum, which is divided into nearly 80 annuli. The annuli bear cuticular scales and small sensory organs, sensilla. A trachea, a blood vessel, and two nerve trunks run through the lumen of the antenna and into the head. Sensilla are arranged in an orderly pattern that is repeated on each flagellar annulus. Each flagellum bears about 10⁵ sensilla, which contain about 2.5 × 10⁵ primary sensory neurons. Clumps of undifferentiated cells (imaginal disks), present in the larva, form pupal antennae during the larval-pupal molt. During the subsequent metamorphic development of the adult, cell divisions, changes in cell shape, and cellular differentiation transform pupal into adult antennae. Sensilla and scales arise and differentiate in the antenna during metamorphosis; regions in which sensilla and scales will arise can be recognized before overt differentiation occurs. All of the flagellar annuli develop synchronously. The dense innervation and neuronal simplicity of antennal flagella, as well as their synchronous development at a late and accessible stage in the animal's life cycle, suit them for studies of neuronal differentiation.     

Differentiation of insect sensory neurons in the absence of their normal synaptic targets.

Sensory neurons in the antenna of the moth, Manduca sexta, arise and differentiate during the 18 days of metamorphosis from pupa to adult, sending axons to the brain. To assess the trophic dependence of developing antennal neurons on their targets, we studied antennae from surgically debrained animals. If the brain is removed 1 to 45 hr after pupal ecdysis (before and during the period when antennal neurons arise by cell divisions), adult development can be triggered by injection of β-ecdysone; if the brain is removed 50 to 60 hr after pupal ecdysis (before antennal axons contact the brain), metamorphosis proceeds spontaneously. Neurons proliferate normally and differentiate extensively in the antennae of debrained animals. They acquire a characteristic size and shape, elaborate axons and dendrites, migrate to appropriate positions in the sensilla, accumulate components of a neurotransmitter system (acetylcholine, choline acetyltransferase, and acetylcholinesterase), and generate electrical responses to olfactory and mechanical stimuli. Antennal sensory neurons thus differ from a variety of vertebrate neurons, which fail to mature when deprived of their synaptic targets.     

伪鞘榕小蜂雌雄成虫触角感器的超微形态、分布及适生意义

伪鞘榕小蜂Sycoscapter trifemmensis是一种寄生于鸡嗉子榕间花期榕果的专性寄生蜂,雌雄两性繁殖策略分化明显,为更好理解和诠释雌蜂寄主定位和雄蜂配偶识别机制,有必要对两性的触角感器进行观察。运用环境扫描电镜观察,对比和探讨了伪鞘榕小蜂雌雄成虫的触角和触角感器类型、分布、数量及其生态适应性。结果表明:雌蜂触角鞭节由11鞭小节组成,总长817.82±33.23μm,分布有毛形感器、刺形感器(类型1)、锥形感器(类型1)、多孔板形感器(类型1)、栓锥形乳突状感器5类5种;雄蜂触角鞭节仅由6鞭小节组成,全长为雌蜂的1/3,且各节有明显的缩短和增粗特征,着生感器包括毛形感器、刺形感器(类型2和类型3)、锥形感器(类型1和类型2)、多孔板形感器(类型2)、腔锥形感器5类7种。雌蜂触角感器的数量与分布显著高于雄蜂,且同类型感器在雌蜂上具有明显的延伸、增粗、分支的特征,以板形感器和锥形感器最为突出。伪鞘榕小蜂雌雄成虫的触角及其感器有明显的性二型,特别是与化学信息识别相关的感器,反映了雌雄蜂在不同生态环境和繁殖压力下的形态分化、行为策略和生态适应。     

The antennal system and cockroach evasive behavior. I. Roles for visual and mechanosensory cues in the response

Cockroaches escape from predators by turning and then running. This behavior can be elicited when stimuli deflect one of the rostrally located and highly mobile antennae. We analyzed the behavior of cockroaches, under free-ranging conditions with videography or tethered in a motion tracking system, to determine (1) how antennal positional dynamics influence escape turning, and (2) if visual cues have any influence on antennal mediated escape. The spatial orientation of the long antennal flagellum at the time of tactile stimulation affected the direction of resultant escape turns. However, the sign of flagellar displacement caused by touch stimuli, whether it was deflected medially or laterally for example, did not affect the directionality of turns. Responsiveness to touch stimuli, and escape turn performance, were not altered by blocking vision. However, because cockroaches first orient an antenna toward stimuli entering the peripheral visual field, turn direction can be indirectly influenced by visual input. Finally, when vision was blocked, the run phase of escape responses displayed reduced average velocities and distances traveled. Our results suggest that tactile and visual influences are integrated with previously known wind-sensory mechanisms to achieve multisensory control of the full escape response.     

Proprioceptive input to a descending pathway conveying antennal postural information: Terminal organisation of antennal hair field afferents

Like several other arthropod species, stick insects use their antennae for tactile exploration of the near-range environment and for spatial localisation of touched objects. More specifically, Carausius morosus continuously moves its antennae during locomotion and reliably responds to antennal contact events with directed movements of a front leg. Here we investigate the afferent projection patterns of antennal hair fields (aHF), proprioceptors known to encode antennal posture and movement, and to be involved in antennal movement control. We show that afferents of all seven aHF of C. morosus have terminal arborisations in the dorsal lobe (DL) of the cerebral (=supraoesophageal) ganglion, and descending collaterals that terminate in a characteristic part of the gnathal (=suboesophageal) ganglion. Despite differences of functional roles among aHF, terminal arborisation patterns show no topological arrangement according to segment specificity or direction of movement. In the DL, antennal motoneuron neurites show arborizations in proximity to aHF afferent terminals. Despite the morphological similarity of single mechanoreceptors of aHF and adjacent tactile hairs on the pedicel and flagellum, we find a clear separation of proprioceptive and exteroceptive mechanosensory neuropils in the cerebral ganglion. Moreover, we also find this functional separation in the gnathal ganglion.     

Active tactile exploration for adaptive locomotion in the stick insect

Insects carry a pair of actively movable feelers that supply the animal with a range of multimodal information. The antennae of the stick insect Carausius morosus are straight and of nearly the same length as the legs, making them ideal probes for near-range exploration. Indeed, stick insects, like many other insects, use antennal contact information for the adaptive control of locomotion, for example, in climbing. Moreover, the active exploratory movement pattern of the antennae is context-dependent. The first objective of the present study is to reveal the significance of antennal contact information for the efficient initiation of climbing. This is done by means of kinematic analysis of freely walking animals as they undergo a tactually elicited transition from walking to climbing. The main findings are that fast, tactually elicited re-targeting movements may occur during an ongoing swing movement, and that the height of the last antennal contact prior to leg contact largely predicts the height of the first leg contact. The second objective is to understand the context-dependent adaptation of the antennal movement pattern in response to tactile contact. We show that the cycle frequency of both antennal joints increases after obstacle contact. Furthermore, inter-joint coupling switches distinctly upon tactile contact, revealing a simple mechanism for context-dependent adaptation.     

红火蚁触角及其上感受器的扫描电镜观察

应用扫描电镜对采自我国广东吴川和广西南宁的红火蚁Solenopsis invicta Buren工蚁、有翅雌蚁和雄蚁触角的形态和感受器类型进行了研究。结果表明,工蚁、雌蚁和雄蚁的触角存在着较大的差异。工蚁和雌蚁的触角为膝状,末端2或3节膨大为棒状,雌蚁的触角11节,比工蚁(10节)的多1节。雄蚁的触角为线状,12节,末端没有棒状结构。工蚁和雌蚁的触角上共有7种感受器,分别为毛形、曲毛形、锥形、腔锥形、坛形、刺形感受器和Böhm氏鬃毛,前6种感受器大多集中在触角末端的棒节,棒节以外的各鞭节上主要是毛形感受器,Böhm氏鬃毛则分布于柄节和梗节的基部,鞭节上无此感受器。雄蚁触角上也有7种感受器,但其分布与工蚁的有较大差异,Böhm氏鬃毛的分布与工蚁和雌蚁的相同,锥形感受器分布在末端2节,坛形感受器分布在末端3节,其余的4种感受器较为均匀地分布在各鞭节,而不是聚集在端节。另外,新发现雄蚁中存在一种与已知的毛型感受器不同的超短毛状感受器,定名为超短毛感受器。最后,对红火蚁的触角及感受器在蚁群社会性行为中的作用进行了讨论。     

Antennal and ovipositor sensilla of Pseudoligosita yasumatsui (Hymenoptera: Trichogrammatidae)

Pseudoligosita yasumatsui Viggiani and Subba Rao 1978 (Hymenoptera: Trichogrammatidae) is a common egg parasitoid of rice insect pests. The surface morphology of the antenna and ovipositor on P. yasumatsui was examined using scanning electron microscopy. The antenna of P. yasumatsui is geniculate in shape, hinged at the scape-pedicel joint, approximately 190 μm in length and consists of seven antennomeres. In total, the male and female antennae have ten different types of sensilla: trichoid sensilla type 1, 2, 3, 4, 5, 6, campaniform sensilla, basiconic sensilla, and placoid sensilla type 1 and 2. The flagellum of the female antenna is covered with cuticular pores, which are absent on the male antennal flagellum. The distal extremity of its ovipositor stylet has campaniform sensilla and styloconic sensilla. Trichoid sensilla found on its apical abdomen part may play a role in the host detection and egg placement. The types and distribution of antennal and ovipositor sensilla on the parasitoid were discussed.