The 3D ultrastructure of the chordotonal organs in the antenna of a microwasp remains complex although simplified.

Insect antennae are astonishingly versatile and have multiple sensory modalities. Audition, detection of airflow, and graviception are combined in the antennal chordotonal organs. The miniaturization of these complex multisensory organs has never been investigated. Here we present a comprehensive study of the structure and scaling of the antennal chordotonal organs of the extremely miniaturized parasitoid wasp Megaphragma viggianii based on 3D electron microscopy. Johnston's organ of M. viggianii consists of 19 amphinematic scolopidia (95 cells); the central organ consists of five scolopidia (20 cells). Plesiomorphic composition includes one accessory cell per scolopidium, but in M. viggianii this ratio is only 0.3. Scolopale rods in Johnston's organ have a unique structure. Allometric analyses demonstrate the effects of scaling on the antennal chordotonal organs in insects. Our results not only shed light on the universal principles of miniaturization of sense organs, but also provide context for future interpretation of the M. viggianii connectome.

Antennal and palpal sensilla of three predatory Lispe species (Diptera: Muscidae): an ultrastructural investigation.

Antennae and maxillary palps are the most important chemical reception organs of flies. So far, the morphology of antennae and maxillary palps of flies of most feeding habits have been well described, except for that of relatively rare aquatic predatory species. This study describes sensilla on antennae and maxillary palps of three aquatic predatory Lispe species: Lispe longicollis, L. orientalis and L. pygmaea. Types, distribution, and density of sensilla are characterised via light and scanning electron microscopy. One type of mechanoreceptors is found on antennal scape. Mechanoreceptors (two subtypes) and one single pedicellar button (in L. pygmaea) are located on antennal pedicel. Four types of sensilla are discovered on antennal postpedicel: trichoid sensilla, basiconic sensilla (three subtypes), coeloconic sensilla and clavate sensilla. A unique character of these Lispe species is that the coeloconic sensilla are distributed sparsely on antennal postpedicel. Mechanoreceptors and basiconic sensilla are observed on the surface of maxillary palps in all three species. We demonstrated clear sexual dimorphism of the maxillary palps in some of the Lispe species, unlike most other Muscidae species, are larger in males than females. This, along with their courtship dance behaviour, suggest their function as both chemical signal receiver and visual signal conveyer, which is among the few records of a chemical reception organ act as a signal conveyer in insects.

Dorsal setae in Raoiella (Acari: Tenuipalpidae): Their functional morphology and implication in fluid secretion.

The setae of mites are not regarded as secretory structures, yet in the flat mite genus Raoiella, each developmental stage presents droplets of fluid associated with the tips of their dorsal setae. To understand the origin of this fluid, the ultrastructure of the dorsal setae is investigated in females of Raoiella bauchani Beard & Ochoa and the invasive pest species Raoiella indica Hirst using scanning and transmission electron microscopy techniques. The dorsal setae are barbed along their entire length and have either a broadened plumose or a flat spatulate tip. Ultrastructurally, they present the typical features of mechanoreceptors, but have a "hollow" axis represented by a protoplasmatic core containing dendritic branches. This combination of ultrastructural characters indicates that the setae might be multimodal receptors: acting as both mechanoreceptors and contact chemoreceptors. The epidermal cells that underlie the setal sockets are columnar and have an ultrastructure that suggests they have a glandular function. Moreover, these cells present regular microvilli apically and form extracellular cuticular canals, containing epicuticular filaments, that are connected with the microvilli proximally and which open via pores onto the surface of the setal base distally. This arrangement indicates that the secretion from the microvilli passes into the canals and is then conducted to pores at the base of the seta, where it then accumulates and moves up the setal shaft, along the longitudinal grooves of the barbs. Based on similar arrangements in some insect taxa, the organization of the structures here observed in Raoiella suggests the passage of a non-polar, water insoluble, lipoid fluid through the cuticle, the function of which is still obscure.

Neurochemical and Ultrastructural Characterization of Unmyelinated Non-peptidergic C-Nociceptors and C-Low Threshold Mechanoreceptors Projecting to Lamina II of the Mouse Spinal Cord.

C-nociceptors (C-Ncs) and non-nociceptive C-low threshold mechanoreceptors (C-LTMRs) are two subpopulations of small unmyelinated non-peptidergic C-type neurons of the dorsal root ganglia (DRGs) with central projections displaying a specific pattern of termination in the spinal cord dorsal horn. Although these two subpopulations exist in several animals, remarkable neurochemical differences occur between mammals, particularly rat/humans from one side and mouse from the other. Mouse is widely investigated by transcriptomics. Therefore, we here studied the immunocytochemistry of murine C-type DRG neurons and their central terminals in spinal lamina II at light and electron microscopic levels. We used a panel of markers for peptidergic (CGRP), non-peptidergic (IB4), nociceptive (TRPV1), non-nociceptive (VGLUT3) C-type neurons and two strains of transgenic mice: the TAFA4Venus knock-in mouse to localize the TAFA4+ C-LTMRs, and a genetically engineered ginip mouse that allows an inducible and tissue-specific ablation of the DRG neurons expressing GINIP, a key modulator of GABABR-mediated analgesia. We confirmed that IB4 and TAFA4 did not coexist in small non-peptidergic C-type DRG neurons and separately tagged the C-Ncs and the C-LTMRs. We then showed that TRPV1 was expressed in only about 7% of the IB4+ non-peptidergic C-Ncs and their type Ia glomerular terminals within lamina II. Notably, the selective ablation of GINIP did not affect these neurons, whereas it reduced IB4 labeling in the medial part of lamina II and the density of C-LTMRs glomerular terminals to about one half throughout the entire lamina. We discuss the significance of these findings for interspecies differences and functional relevance.

Comparative study of the cephalic trichobothria in plant bugs (Hemiptera: Heteroptera: Miridae).

A pair of ventral cephalic trichobothria was observed for the first time and so far the only one in a representative of single species of Miridae (Fulvius carayoni) in 2013. The purpose of our research was to verify the hypothesis that this is not an exception, but a characteristic feature of all plant bugs. Twenty-three representatives of all seven subfamilies of Miridae were examined using a scanning electron microscope. The results presents detailed data on the distribution and ultramorphology of the cephalic trichobothria in plant bugs. A pair of ventral cephalic trichobothria was observed in all of the examined species. Each trichobothrium of this pair is located laterally to the first article of the rostrum, on the gula (between the buccula and the antennal tubercle). Moreover, a pair of dorsal cephalic trichobothria was observed for the first time. They were found in nine species, located above the antennal tubercle, towards the center of the frons.

The effect of axotomy on firing and ultrastructure of the crayfish mechanoreceptor neurons and satellite glial cells.

Neurotrauma is among main causes of human disability and death. We studied effects of axotomy on ultrastructure and neuronal activity of a simple model object - an isolated crayfish stretch receptor that consists of single mechanoreceptor neurons (MRN) enwrapped by multilayer glial envelope. After isolation, MRN regularly fired until spontaneous activity cessation. Axotomy did not change significantly MRN spike amplitude and firing rate. However, the duration of neuron activity from MRN isolation to its spontaneous cessation decreased in axotomized MRN relative to intact neuron. [Ca2+] in MRN axon and soma increased 3-10 min after axotomy. Ca2+ entry through ion channels in the axolemma accelerated axotomy-stimulated firing cessation. MRN incubation with Ca2+ionophore ionomycin accelerated MRN inactivation, whereas Ca2+-channel blocker Cd2+ prolonged firing. Activity duration of either intact, or axotomized MRN did not change in the presence of ryanodine or dantrolene, inhibitors of ryanodin-sensitive Ca2+ channels in endoplasmic reticulum. Thapsigargin, inhibitor of endoplasmic reticulum Ca2+-ATPase, or its activator ochratoxin were ineffective. Ultrastructural study showed that the defect in the axon transected by thin scissors is sealed by fused axolemma, glial and collagen layers. Only the 30-50 µm long segment completely lost microtubules and contained swelled mitochondria. The microtubular bundle remained undamaged at 300 µm away from the axotomy site. However, mitochondria within the 200-300 µm segment were strongly condensed and lost matrix and cristae. Glial and collagen layers exhibited greater damage. Swelling and edema of glial layers, collagen disorganization and rupture occurred within this segment. Thus, axotomy stronger damages glia/collagen envelope, axonal microtubules and mitochondria.

The morphology, size and density of the touch dome in human hairy skin by scanning electron microscopy.

The touch domes of mammalian hairy skin are mechanoreceptors characterized by the accumulation of Merkel cell-neurite complexes at the epidermal base. In this study, we examined the shape, size, and density of the touch dome of human skin of the forearm and the abdomen through scanning electron microscopy (SEM). Human skin samples were obtained from donated bodies, as well as a patient who underwent biopsy. Skin pieces were treated with a KOH-collagenase method for the separation of the epidermis from the dermis. The basal surface of the separated epidermis was then observed using SEM. The touch dome was clearly determined as a concave area bordered by a thick epidermal ridge, where neural components accumulated. The touch dome was rather independent from hair follicles, although they were sometimes located beside the touch dome. The average size and density of the touch dome were 0.06 mm2 and 3.82 cm2 in the forearm, and 0.10 mm2 and 1.30 cm2 in the abdomen, respectively. Our results suggested that the regional difference in size and density of the touch dome might be related to the sensation's sensitivity as touch spots in human hairy skin.

Ecological associations among epidermal microstructure and scale characteristics of Australian geckos (Squamata: Carphodactylidae and Diplodactylidae).

A first step in examining factors influencing trait evolution is demonstrating associations between traits and environmental factors. Scale microstructure is a well-studied feature of squamate reptiles (Squamata), including geckos, but few studies examine ecology the of microstructures, and those focus mainly on toe pads. In this study, the ecomorphology of cutaneous microstructures on the dorsum was described for eight Australian species of carphodactylid (Squamata: Carphodactylidae) and 19 diplodactylid (Squamata: Diplodactylidae) geckos. We examined scale dimensions, spinule and cutaneous sensilla (CS) morphology, using scanning electron microscopy, and described associations of these traits with microhabitat selection (arboreal, saxicoline or terrestrial) and relative humidity of each species' habitat (xeric, mesic or humid). We used a phylogenetic flexible discriminant analysis (pFDA) to describe relationships among all traits and then a modeling approach to examine each trait individually. Our analysis showed that terrestrial species tended to have long spinules and CS with more bristles, saxicoline species larger diameter CS and arboreal species tended to have large granule scales and small intergranule scales. There was high overlap in cutaneous microstructural morphology among species from xeric and mesic environments, whereas species from humid environments had large diameter CS and few bristles. Significant associations between epidermal morphology and environmental humidity and habitat suggest that epidermal microstructures have evolved in response to environmental variables. In summary, long spinules, which aid self-cleaning in terrestrial geckos, are consistent with greater exposure to dirt and debris in this habitat. Long spinules were not clearly correlated to environmental humidity. Finally, more complex CS (larger diameter with more bristles) may facilitate better perception of environmental variation in geckos living in drier habitats.

Early Triassic marine reptile representing the oldest record of unusually small eyes in reptiles indicating non-visual prey detection.

The end-Permian mass extinction (EPME) led to reorganization of marine predatory communities, through introduction of air-breathing top predators, such as marine reptiles. We report two new specimens of one such marine reptile, Eretmorhipis carrolldongi, from the Lower Triassic of Hubei, China, revealing superficial convergence with the modern duckbilled platypus (Ornithorhynchus anatinus), a monotreme mammal. Apparent similarities include exceptionally small eyes relative to the body, snout ending with crura with a large internasal space, housing a bone reminiscent of os paradoxum, a mysterious bone of platypus, and external grooves along the crura. The specimens also have a rigid body with triangular bony blades protruding from the back. The small eyes likely played reduced roles during foraging in this animal, as with extant amniotes (group containing mammals and reptiles) with similarly small eyes. Mechanoreceptors on the bill of the animal were probably used for prey detection instead. The specimens represent the oldest record of amniotes with extremely reduced visual capacity, utilizing non-visual cues for prey detection. The discovery reveals that the ecological diversity of marine predators was already high in the late Early Triassic, and challenges the traditional view that the ecological diversification of marine reptiles was delayed following the EPME.

Design principles of hair-like structures as biological machines.

Hair-like structures are prevalent throughout biology and frequently act to sense or alter interactions with an organism's environment. The overall shape of a hair is simple: a long, filamentous object that protrudes from the surface of an organism. This basic design, however, can confer a wide range of functions, owing largely to the flexibility and large surface area that it usually possesses. From this simple structural basis, small changes in geometry, such as diameter, curvature and inter-hair spacing, can have considerable effects on mechanical properties, allowing functions such as mechanosensing, attachment, movement and protection. Here, we explore how passive features of hair-like structures, both individually and within arrays, enable diverse functions across biology. Understanding the relationships between form and function can provide biologists with an appreciation for the constraints and possibilities on hair-like structures. Additionally, such structures have already been used in biomimetic engineering with applications in sensing, water capture and adhesion. By examining hairs as a functional mechanical unit, geometry and arrangement can be rationally designed to generate new engineering devices and ideas.

The mechanosensory-motor apparatus of antennae in the Oleander hawk moth (Daphnis nerii, Lepidoptera).

Insect antennae are sensory organs of great importance because they can sense diverse environmental stimuli. In addition to serving as primary olfactory organs of insects, antennae also sense a wide variety of mechanosensory stimuli, ranging from low-frequency airflow or gravity cues to high-frequency antennal vibrations due to sound, flight or touch. The basal segments of the antennae house multiple types of mechanosensory structures that prominently include the sensory hair plates, or Böhm's bristles, which measure the gross extent of antennal movement, and a ring of highly sensitive scolopidial neurons, collectively called the Johnston's organs, which record subtle flagellar vibrations. To fulfill their multifunctional mechanosensory role, the antennae of insects must actively move thereby enhancing their ability to sense various cues in the surrounding environment. This tight coupling between antennal mechanosensory function and antennal movements means that the underlying mechanosensory-motor apparatus constitutes a highly tuned feedback-controlled system. Our study aims to explore how the sensory and motor components of this system are configured to enable such functional versatility. We describe antennal mechanosensory neurons, their central projections in the brain relative to antennal motor neurons and the internal morphology of various antennal muscles that actuate the basal segments of the antenna. We studied these in the Oleander hawk moth (Daphnis nerii) using a combination of techniques such as neural dye fills, confocal microscopy, scanning electron microscopy and X-ray tomography. Our study thus provides a detailed anatomical picture of the antennal mechanosensory-motor apparatus, which in turn provides key insights into its multifunctional role.

The antennae of damselfly larvae.

The larval antennal sensilla of two Zygoptera species, Calopteryx haemorroidalis (Calopterygidae) and Ischnura elegans (Coenagrionidae) are investigated with SEM and TEM. These two species have different antennae (geniculate, setaceous) and live in different environments (lotic, lentic waters). Notwithstanding this, similarities in the kind and distribution of sensilla are outlined: in both species the majority of sensilla types is located on the apical portion of the antenna, namely a composed coeloconic sensillum (possible chemoreceptor), two other coeloconic sensilla (possible thermo-hygroreceptors) and an apical seta (direct contact mechanoreceptor). Other mechanoreceptors, such as filiform hairs sensitive to movements of the surrounding medium or bristles positioned to sense the movements of the flagellar segments, are present on the antenna. Similarities in the antennal sensilla types and distribution are observed also with other dragonfly species, such as Onychogomphus forcipatus and Libellula depressa. A peculiar structure with an internal organization similar to that of a gland is observed in the apical antenna of C. haemorroidalis and I. elegans and it is present also in O. forcipatus and L. depressa. The possible function of this structure is at the moment unknown but deserves further investigations owing to its widespread presence in Odonata larvae.

Morphology and hydro-sensory role of superficial neuromasts in schooling behaviour of yellow-eyed mullet (Aldrichetta forsteri).

The lateral line system is a mechanosensory organ found in all fish species and located on the skin or in subdermal canals. The basic functional units are superficial and canal neuromasts, which are involved in hydrodynamic sensing and cohesion in schooling fish. Yellow-eyed mullet (Aldrichetta forsteri) are an obligate schooling species found commonly in shallow coastal areas of New Zealand and Australia. Schooling is a fundamental part of their behavioural repertoire, yet little is known about the structure or functionality of the lateral line in this species. We used scanning electron microscopy to characterise the morphology of trunk superficial neuromasts. We then took a multi-sensory approach and conducted behavioural experiments comparing school structure in groups of fish with and without fully functioning lateral lines, under photopic and scotopic conditions. A highly developed hydro-sensing system exists on the trunk of yellow-eyed mullet consisting of superficial neuromasts containing hundreds of hair cells aligned, with respect to their most sensitive axis, in a rostrocaudal direction. Without functioning superficial neuromasts, schooling behaviour was disrupted under both photopic and scotopic conditions and the ability to detect stationary objects decreased. Results highlight the importance of this component of the lateral line system to schooling behaviour.

A population of G2-arrested cells are selected as sensory organ precursors for the interommatidial bristles of the Drosophila eye.

Cell cycle progression and differentiation are highly coordinated during the development of multicellular organisms. The mechanisms by which these processes are coordinated and how their coordination contributes to normal development are not fully understood. Here, we determine the developmental fate of a population of precursor cells in the developing Drosophila melanogaster retina that arrest in G2 phase of the cell cycle and investigate whether cell cycle phase-specific arrest influences the fate of these cells. We demonstrate that retinal precursor cells that arrest in G2 during larval development are selected as sensory organ precursors (SOPs) during pupal development and undergo two cell divisions to generate the four-cell interommatidial mechanosensory bristles. While G2 arrest is not required for bristle development, preventing G2 arrest results in incorrect bristle positioning in the adult eye. We conclude that G2-arrested cells provide a positional cue during development to ensure proper spacing of bristles in the eye. Our results suggest that the control of cell cycle progression refines cell fate decisions and that the relationship between these two processes is not necessarily deterministic.

Haltere morphology and campaniform sensilla arrangement across Diptera.

One of the primary specializations of true flies (order Diptera) is the modification of the hind wings into club-shaped halteres. Halteres are complex mechanosensory structures that provide sensory feedback essential for stable flight control via an array of campaniform sensilla at the haltere base. The morphology of these sensilla has previously been described in a small number of dipteran species, but little is known about how they vary across fly taxa. Using a synoptic set of specimens representing 42 families from all of the major infraorders of Diptera, we used scanning electron microscopy to map the gross and fine structures of halteres, including sensillum shape and arrangement. We found that several features of haltere morphology correspond with dipteran phylogeny: Schizophora generally have smaller halteres with stereotyped and highly organized sensilla compared to nematoceran flies. We also found a previously undocumented high variation of haltere sensillum shape in nematoceran dipterans, as well as the absence of a dorsal sensillum field in multiple families. Overall, variation in haltere sensillar morphology across the dipteran phylogeny provides insight into the evolution of a highly specialized proprioceptive organ and a basis for future studies on haltere sensory function.

[The quill mite Syringophilopsis fringilla (Fritsch) (Acari: Trombidiformes: Syringophilidae): The structure of receptor organs providing feeding of the parasite inside the feather quill].

The structure of sensory organs situated on palps and inside the cheliceral stylet of the quill mite Sringophilopsis fringilla (Fritsch, 1958) was examined in scanning and transmitting electron microscopes. Eight sensilla of 3 types were revealed on palptarsus, including two contact chemo-mechanosensory sensilla, a single distant chemosensory (probably olfactory) sensillum, and 5 tactile mechanosensitive sensilla. All other sensilla situated on basas parts of the palp and on the outer surface of gnathosoma are represented by tactile mechanoreceptors. A proprioceptor sensillum was revealed in the movable digit of the chelicera; modified cilia of dendrites of 5 sensory neurons run in the inner non-sclerotized core of the stylet, ending at different levels as electron-dense rods connected with the sclerotized wall of the stylet. The authors assume that the proprioceptor sensillum of the stylet detects the pressing force of the movable digit on the inner wall of the quill during piercing process, while papal sensilla determine the optimal place for piercing.

Novel structure in sciaenid fish skulls indicates continuous production of the cephalic neuromast cupula.

The presence of a conspicuous and frequent but never-described structure in the skull cavities of sciaenid fish was noted during population studies in an urbanized bay. The ultrastructure closely resembles the cupula of neuromasts, an organ associated with the perception of the environment in teleost fish. The bodies were recorded detached in both preserved and freshly sampled individuals and without associated cilia. Prominent characteristics are acellularity, the elliptic-conic shape composed of stack-like protein lamellas, and a mesh-like appearance in cross section. These acellular lamellar cephalic bodies (ALCBs) were more abundant in larger individuals and showed temporal peaks of abundance independently of the fish size. The conic and lamellar features suggest that the deposition of protein layers follows fish growth, and the bimodality of the size of these structures in individuals indicates temporal peaks of production. These results indicate that these ALCBs are a consequence of the accretion of the cupula of neuromasts at a faster rate than they degrade. Given the novelty of this structure and the increasing records of diseases of marine organisms worldwide, an important question is whether these bodies occur subsequently to some environmental change and whether their accumulation in the skull cavities has consequences to fish health.

Morphology, Ultrastructure and Possible Functions of Antennal Sensilla of Sitodiplosis mosellana Géhin (Diptera: Cecidomyiidae).

To better understand the olfactory receptive mechanisms involved in host selection and courtship behavior of Sitodiplosis mosellana (Diptera: Cecidomyiidae), one of the most important pests of wheat, scanning and transmission electron microscopy were used to examine the external morphology and ultrastructure of the antennal sensilla. The moniliform antennae exhibit obvious sexual dimorphism: antennae of the males are markedly longer than those of the females. Furthermore, each male flagellomere consists of two globular nodes, whereas each female flagellomere is cylindrical. Seven types of sensilla were identified in both sexes. Two types of s. chaetica have a lumen without dendrites and thick walls, suggesting that they are mechanoreceptors. S. trichodea and s. circumfila are typical chemoreceptors, possessing thin multiporous walls encircling a lumen with multiple dendrites. There are significantly more s. trichodea in female than in male, which may be related to host plant localization. In contrast, male s. circumfila are highly elongated compared to those of females, perhaps for pheromone detection. Peg-shaped s. coeloconica are innervated with unbranched dendrites extending from the base to the distal tip. Type 1 s. coeloconica, which have deep longitudinal grooves and finger-like projections on the surface, may serve as olfactory or humidity receptors, whereas type 2 s. coeloconica, smooth with a terminal pore, may be contact chemoreceptors. Also, this is the first report of Böhm' bristles at proximal scape on antennae of Cecidomyiid species potentially functioning as mechanoreceptors.

Atypical mechanoreceptors in larvae of biting midges Forcipomyia nigra (Diptera: Ceratopogonidae).

The dorsolateral setae of Forcipomyia nigra have been investigated. They are mechanoreceptors, sensilla trichoidea, innerved by a single neuron. The tubular body is located at the base of the hair shaft. Although the fine structure of the dorsolateral setae is similar to dorsal setae that perform a double function: secretion of a hygroscopic viscous substance through the pores and mechanoreception dorsolateral setae, they do not secrete any fluid. In both types of setae, trichogen cells (hair forming cells) produce the hair shafts and thereafter do not retract from the cavity of the setae. They contain a large polyploid nucleus and expanded bundles of microtubules. In dorsolateral mechanoreceptors, the microtubules form a network around the nucleus of the trichogen cell and are especially numerous in the cytoplasm invading the interior of the seta, which is evidenced by immunofluorescence light microscopy. No tormogen cell, responsible for the production of the setal socket, was found. Our observations indicate that the dorsolateral setae are solely mechanoreceptors but their trichogen cells reveal some glandular activity.

Electrophysiological evidence for the existence of a rare population of C-fiber low threshold mechanoreceptive (C-LTM) neurons in glabrous skin of the rat hindpaw.

The mammalian skin in innervated by distinct classes of low-threshold mechanoreceptive (LTM) primary afferent neurons that are classified as Aß-, Aδ- or C-LTMs according to their axonal conduction velocities (CVs). C-LTMs are thought to signal pleasant and erotic touch sensations in humans, and to exist only in the hairy skin of primates and other species. Using intracellular recordings from rat L4/L5 dorsal root ganglion (DRG) neurons that were classified in vivo as C-nociceptors or C-LTMs, according to their dorsal root CVs and their responses to mechanical and thermal stimuli, the present study provides the first electrophysiological evidence that C-LTMs exist in the glabrous skin of the rat's hindpaw. Indeed 6.4% (5/78) of the total sample of lumbar C-fiber DRG neurons with receptive fields in the glabrous skin of the rat hindpaw were C-LTMs. The electrophysiological properties of this rare subpopulation of C-fiber neurons (mean CV=0.48±0.06m/s) are distinct from those of C-fiber high threshold mechanoreceptors (HTMs). Indeed, their mean mechanical (1.7±1.1mN) and electrical (4.0±0.4V) thresholds was significantly different from that of C-HTMs. They also exhibited faster action potential and afterhyperpolarization kinetics than C-HTMs. The present study lends support to previous studies that have provided indirect evidence for the presence of C-LTMs in glabrous skin. If C-LTMs are present in human glabrous skin, they may, in this type of skin, represent a novel peripheral neuronal substrate for the pleasant/social touch sensation, and account for or contribute to touch hypersensitivity after injury.