The effects of stimulus location on the gating of touch by heat- and cold-induced pain

The influence of heat- and cold-induced pain on tactile sensitivity, a "touch gate", was measured under conditions in which the location of the noxious stimuli was varied with respect to the tactile stimulus applied to the thenar eminence of humans. Vibrotactile thresholds were measured in the absence of pain and during administration of a painful stimulus, with the stimulus frequencies selected to activate independently the four psychophysical channels hypothesized to exist in human glabrous skin. Heat-induced pain produced by spatially co-localizing the noxious stimuli with the tactile stimuli was found, on average, to elevate threshold amplitude by 2.2 times (6.7 dB). Co-localized, cold-induced pain raised the average thresholds by about 1.5 times (3.6 dB). Heat-induced pain presented contralaterally produced no change in vibrotactile sensitivity indicating that the effect is probably not due to attentional mechanisms. Ipsilateral heat-induced pain caused an elevation in tactile thresholds even when the noxious and non-noxious stimuli were not co-localized, and the effect may seem to require that the painful stimulus be within the somatosensory region defined possibly in terms of dermatomal organization. Thus the effect is probably related to somatotopic organization and is not peripherally mediated. A brief discussion as to the possible locus of the touch gate within the nervous system is also given.     

The effects of stimulus location on the gating of touch by heat- and cold-induced pain

The influence of heat- and cold-induced pain on tactile sensitivity, a "touch gate", was measured under conditions in which the location of the noxious stimuli was varied with respect to the tactile stimulus applied to the thenar eminence of humans. Vibrotactile thresholds were measured in the absence of pain and during administration of a painful stimulus, with the stimulus frequencies selected to activate independently the four psychophysical channels hypothesized to exist in human glabrous skin. Heat-induced pain produced by spatially co-localizing the noxious stimuli with the tactile stimuli was found, on average, to elevate threshold amplitude by 2.2 times (6.7 dB). Co-localized, cold-induced pain raised the average thresholds by about 1.5 times (3.6 dB). Heat-induced pain presented contralaterally produced no change in vibrotactile sensitivity indicating that the effect is probably not due to attentional mechanisms. Ipsilateral heat-induced pain caused an elevation in tactile thresholds even when the noxious and non-noxious stimuli were not co-localized, and the effect may seem to require that the painful stimulus be within the somatosensory region defined possibly in terms of dermatomal organization. Thus the effect is probably related to somatotopic organization and is not peripherally mediated. A brief discussion as to the possible locus of the touch gate within the nervous system is also given.     

Modulation of skin sensitivity by dynamic and isometric exercise in man

The effect of dynamic cycle ergometer exercise and isometric leg exercise on skin sensitivity was studied in man. Exercise was performed at different loads. Cutaneous sensitivity to innocuous and noxious thermal stimuli was tested using a contact thermostimulator and sensitivity to tactile stimuli was tested using electrical stimuli. During isometric exercise a segmental (the exercising limb), but not a multisegmental, phasic decrease of cutaneous thermal sensitivity to innocuous stimuli was found. At the isometric forces used the effect on tactile and heat pain sensitivity was not significant. During dynamic exercise a multisegmental, load-dependent decrease of sensitivity in all tested sensory modalities was found and this attenuation disappeared gradually after the end of exercise. In contrast to isometric exercise, the decrease of sensitivity produced by dynamic exercise was most evident in tactile sensitivity. The size of the stimulus area (7.9 vs 11.8 cm2) did not have a significant effect on the magnitude of the exercise-induced decrease of cutaneous thermal sensitivity to innocuous stimuli. It was concluded that underlying the modulation of skin sensitivity by dynamic and isometric exercise were mechanisms that were different, at least to a small extent. Isometric exercise produced a segmental modulation of skin sensitivity due to central neuronal mechanisms, independent of exercise-induced stress. Exercise-induced stress could have caused the modulation of skin sensitivity by dynamic exercise.     

Representation of moving stimuli by somatosensory neurons.

The findings obtained in neurophysiological and psychophysical investigations using tactile stimuli that move at constant velocity across the skin are reviewed. For certain neurons in the postcentral gyrus of the cerebral cortex (S-I) of macaque monkeys, direction of stimulus motion is a "trigger feature" i.e., moving tactile stimuli evoke vigorous discharge activity in these neurons only if the stimuli are moved in a particular direction across the receptive field. This directional selectivity is maximal when stimulus velocity is between 5 and 50 cm/sec, and falls off rapidly at lower or higher velocities. The capacity for human subjects to correctly identify the direction of stimulus motion on the skin exhibits a similar dependence on stimulus velocity. The similar effects of velocity on neural and psychophysical measures of directional sensitivity support the idea that direction of stimulus motion on the skin can only be recognized if the moving stimulus optimally activates the group of S-I neurons for which that directions of simulus motion is the trigger feature.     

Conditions for noise reduction and stable encoding of spatial structure by cortical neural networks

Cortical circuits have been proposed to encode information by forming stable spatially structured attractors. Experimentally in the primary somatosensory cortex of the monkey, temporally invariant stimuli lead to spatially structured activity patterns. The purpose of this work is to study a recurrent cortical neural network model with lateral inhibition and examine what effect additive random noise has on the networks' ability to form stable spatially structured representations of the stimulus pattern. We show numerically that this network performs edge enhancement and forms statistically stationary, spatially structured responses when the lateral inhibition is of moderate strength. We then derive analytical conditions on the connectivity matrix that ensure stochasticly stable encoding of the stimulus spatial structure by the network. For stimuli whose strength falls in the near linear region of the sigmoid, we are able to give explicit conditions on the eigenvalues of the connection matrix. Finally, we prove that a network with a connection matrix, where the total excitation and inhibition impinging upon a neural unit are nearly balanced, will yield stable spatial attractor responses. Received: 16 October 1998 / Accepted in revised form: 25 November 1999     

Neural mechanisms of tactile motion integration in somatosensory cortex

How are local motion signals integrated to form a global motion percept? We investigate the neural mechanisms of tactile motion integration by presenting tactile gratings and plaids to the fingertips of monkeys, using the tactile analogue of a visual monitor and recording the responses evoked in somatosensory cortical neurons. The perceived directions of the gratings and plaids are measured in parallel psychophysical experiments. We identify a population of somatosensory neurons that exhibit integration properties comparable to those induced by analogous visual stimuli in area MT and find that these neural responses account for the perceived direction of the stimuli across all stimulus conditions tested. The preferred direction of the neurons and the perceived direction of the stimuli can be predicted from the weighted average of the directions of the individual stimulus features, highlighting that the somatosensory system implements a vector average mechanism to compute tactile motion direction that bears striking similarities to its visual counterpart.     

The effects of cross-modal manipulations of attention on the detection of vibrotactile stimuli in humans

Although it is well known that attention to a visual or auditory stimulus can enhance its perception, less is known concerning the effects of attention on the perception of natural tactile stimuli. The present study was conducted to examine the magnitude of the effect of cross-modal manipulations of attention in human subjects on the detection of weak, low-frequency vibrotactile stimuli delivered to the glabrous skin of the finger pad of the right index finger via an Optacon. Three suprathreshold vibrotactile arrays (40 Hz), varying in the number of activated pegs and hence the area of skin stimulated, were used. Subjects were trained to detect the occurrence of vibrotactile or visual stimuli and to respond by pressing a foot pedal as quickly as possible thereafter. Two instructional lights were used to cue the subjects as to which stimulus modality they should attend, in three experimental conditions. In the first cue condition, the forthcoming stimulus modality was indicated by the illumination of its associated light. In the second cue condition, both instructional lights were illuminated, and the subjects were asked to divide their attention equally between the two modalities. In the third cue condition, the stimulus modality was falsely indicated by the illumination of the cue not associated with the stimulus to be presented. Reaction times (RTs) were calculated for each trial. For each modality, tactile and visual, the RTs varied significantly with the cue condition, with the mean RT changing in a graded manner across the experimental conditions (being shortest for the correctly cued condition, intermediate for the neutrally cued condition, and longest for the incorrectly cued condition.(ABSTRACT TRUNCATED AT 250 WORDS)     

Localization of Chemical Stimulation: Capsaicin on Hairy Skin

The ability to localize a chemical stimulus applied to the skin of the forearm was compared to the ability to localize a punctate tactile stimulus. The chemical stimulus was a single, 6-μ1 drop of a 1.0% solution of capsaicin in an ethanol vehicle; the tactile stimulus was a polyester monofilament that exerted 7.5 g of force. Subjects attempted to localize the stimuli at 30-sec intervals for a period of 13.5 min, and rated the perceived intensity and quality of the chemogenic sensations. To avoid generating potentially confounding tactile sensations, localization attempts were made by pointing to the area of sensation with a focused light beam. The results showed that overall, chemical localization was inferior to tactile localization: The absolute error of localization averaged 2.5 cm for capsaicin compared to 1.4 cm for the monofilament. The experiment also revealed that chemical localization (1) varied significantly across arms, (2) exhibited a relatively strong bias toward the elbow, and (3) appeared to be unaffected by the perceived intensity of the sensation. The dominant sensation quality reported was itch. The results are discussed in the context of cutaneous localization in general and localization in the nociceptive system in particular.     

Social facilitation of insect reproduction with motor-driven tactile stimuli

Tactile stimuli provide animals with important information about the environment, including physical features such as obstacles, and biologically relevant cues related to food, mates, hosts and predators. The antennae, the principal sensory organs of insects, house an array of sensory receptors for olfaction, gustation, audition, nociception, balance, stability, graviception, static electric fields, and thermo-, hygro- and mechanoreception. The antennae, being the anteriormost sensory appendages, play a prominent role in social interactions with conspecifics that involve primarily chemosensory and tactile stimuli. In the German cockroach (Blattella germanica) antennal contact during social interactions modulates brain-regulated juvenile hormone production, ultimately accelerating the reproductive rate in females. The primary sensory modality mediating this social facilitation of reproduction is antennal mechanoreception. We investigated the key elements, or stimulus features, of antennal contact that socially facilitate reproduction in B. germanica females. Using motor-driven antenna mimics, we assessed the physiological responses of females to artificial tactile stimulation. Our results indicate that tactile stimulation with artificial materials, some deviating significantly from the native antennal morphology, can facilitate female reproduction. However, none of the artificial stimuli matched the effects of social interactions with a conspecific female.     

Spatially structured oscillations in a two-dimensional excitatory neuronal network with synaptic depression

We study the spatiotemporal dynamics of a two-dimensional excitatory neuronal network with synaptic depression. Coupling between populations of neurons is taken to be nonlocal, while depression is taken to be local and presynaptic. We show that the network supports a wide range of spatially structured oscillations, which are suggestive of phenomena seen in cortical slice experiments and in vivo. The particular form of the oscillations depends on initial conditions and the level of background noise. Given an initial, spatially localized stimulus, activity evolves to a spatially localized oscillating core that periodically emits target waves. Low levels of noise can spontaneously generate several pockets of oscillatory activity that interact via their target patterns. Periodic activity in space can also organize into spiral waves, provided that there is some source of rotational symmetry breaking due to external stimuli or noise. In the high gain limit, no oscillatory behavior exists, but a transient stimulus can lead to a single, outward propagating target wave.     

Mesenteric and tactile Pacinian corpuscles are anatomically and physiologically comparable

Pacinian corpuscles (PCs) in cat mesentery have been studied extensively to help determine the structural and functional bases of tactile mechanotransduction. Although we, like many other investigators, have found that the mesenteric receptors are anatomically very similar to those found in mammalian skin, few physiological characteristics of the mesenteric PCs and those of the skin have been compared. Action-potential rate-amplitude and frequency characteristics (10 Hz–1 KHz), as well as interval (IH) and peri-stimulus-time (PSTH) histograms in response to sinusoidal displacements were obtained from nerve fibers innervating mesenteric PCs and from PC fibers innervating cat glabrous skin. The intensity characteristics obtained on both preparations showed similar response profiles, including equal slopes for low stimulus intensities (approximately 10, with impulse ratios/20 dB displacement) and one and two impulse/cycle entrainment. The frequency characteristics of both groups were U-shaped with similar low-frequency slopes (?12.5 dB/octave) and bandwidths (Q_3dB = 1.4). The best frequency for both the tactile PCs' and mesenteric PCs was 250 Hz, which is in the expected range. The IHs showed entrainment and the PSTHs showed neither transient responses nor adaptation to steady-state sinusoidal stimuli. The functional similarity between mesenteric PCs' nerve responses and those of tactile PC afferents, as well as the receptors’ anatomical similarity, lead us to suggest that the mesenteric PC can act as a model for those in the skin. Furthermore, since the frequency characteristics of the two PC types are similar, it is concluded that the skin, while attenuating stimulus intensity, does not impart temporal filtering of vibratory stimuli.     

The capacity of human subjects to process directional information provided at two skin sites

The ability of human subjects to discriminate direction of tactile stimulus motion on the dorsum of the hand was determined (1) in the absence and (2) in the presence of a moving stimulus delivered to a second skin site on the ipsilateral or contralateral forelimb. When the two skin sites were simultaneously contacted by stimuli moving in the same direction, directional sensitivity was typically below that predicted for a hypothetical subject who could independently process the information provided at each of the two skin sites. Even when the stimulus delivered to a second site was deliberately ignored, it could still alter a subject's perception of stimulus direction on the dorsal hand. Moreover, its influence was greatest whenever it moved in a direction opposite to that of the attended stimulus. Whenever the two moving stimuli were delivered nonsimultaneously to two skin sites, directional sensitivity rarely matched the levels predicted for a hypothetical subject who could independently process the information provided at each site. This, in part, resulted from the subjects' utilization of "long-range" cues provided by the temporal order of stimulation. Subjects frequently failed to distinguish these cues from the sensation of stimulus direction provided at each skin site.     

Mesenteric and tactile Pacinian corpuscles are anatomically and physiologically comparable

Pacinian corpuscles (PCs) in cat mesentery have been studied extensively to help determine the structural and functional bases of tactile mechanotransduction. Although we, like many other investigators, have found that the mesenteric receptors are anatomically very similar to those found in mammalian skin, few physiological characteristics of the mesenteric PCs and those of the skin have been compared. Action-potential rate-amplitude and frequency characteristics (10 Hz-1 KHz), as well as interval (IH) and peri-stimulus-time (PSTH) histograms in response to sinusoidal displacements were obtained from nerve fibers innervating mesenteric PCs and from PC fibers innervating cat glabrous skin. The intensity characteristics obtained on both preparations showed similar response profiles, including equal slopes for low stimulus intensities (approximately 10, with impulse ratios/20 dB displacement) and one and two impulse/cycle entrainment. The frequency characteristics of both groups were U-shaped with similar low-frequency slopes (-12.5 dB/octave) and bandwidths (Q(3dB) = 1.4). The best frequency for both the tactile PCs' and mesenteric PCs was 250 Hz, which is in the expected range. The IHs showed entrainment and the PSTHs showed neither transient responses nor adaptation to steady-state sinusoidal stimuli. The functional similarity between mesenteric PCs' nerve responses and those of tactile PC afferents, as well as the receptors' anatomical similarity, lead us to suggest that the mesenteric PC can act as a model for those in the skin. Furthermore, since the frequency characteristics of the two PC types are similar, it is concluded that the skin, while attenuating stimulus intensity, does not impart temporal filtering of vibratory stimuli.     

Dependence of Subjective Traverse Length on Velocity of Moving Tactile Stimuli

Two series of experiments were performed to assess the effects of stimulus velocity on human subjects' perception of the distance traversed by a moving tactile stimulus. In all experiments, constant-velocity stimuli were applied to the dorsal surface of the left forearm; velocities ranging between 1.0 and 256 cm/sec were used. In some experiments the stimuli moved from distal to proximal over the skin, and in others they moved from proximal to distal. The length of skin contacted by the moving stimulus was defined by a plate having an aperture of 4.0 × 0.5 cm.

In the first series of experiments, subjects were required to compare the distance traversed by a test stimulus delivered 2 sec after a standard stimulus, and also to report the on-locus and the off-locus of the brushing stimulus. In the second series of experiments, the subjects rated the perceived distance on the skin using a free-magnitude-estimation procedure. The data from both series of experiments defined the same relationship between stimulus velocity and perceived stimulus distance. More specifically, although the length of skin contacted by the stimulus was the same at all velocities, subjects' estimates of stimulus distance decreased with increasing stimulus velocity. In addition, the function relating estimates of stimulus distance to velocity was flat for velocities between 5 and 20 cm/sec, but possessed an appreciable negative slope at lower and higher velocities.

It is interesting that the plateau of the relationship between perceived stimulus distance and velocity occurred within the range of velocities that human subjects employ to scan textured surfaces; it also corresponded precisely with the range of stimulus velocities at which the directional sensitivity of somatosensory cortical neurons and human subjects is optimal.     

High-content behavioral analysis of Caenorhabditis elegans in precise spatiotemporal chemical environments

To quantitatively understand chemosensory behaviors, it is desirable to present many animals with repeatable, well-defined chemical stimuli. To that end, we describe a microfluidic system to analyze Caenorhabditis elegans behavior in defined temporal and spatial stimulus patterns. A 2 cm × 2 cm structured arena allowed C. elegans to perform crawling locomotion in a controlled liquid environment. We characterized behavioral responses to attractive odors with three stimulus patterns: temporal pulses, spatial stripes and a linear concentration gradient, all delivered in the fluid phase to eliminate variability associated with air-fluid transitions. Different stimulus configurations preferentially revealed turning dynamics in a biased random walk, directed orientation into an odor stripe and speed regulation by odor. We identified both expected and unexpected responses in wild-type worms and sensory mutants by quantifying dozens of behavioral parameters. The devices are inexpensive, easy to fabricate, reusable and suitable for delivering any liquid-borne stimulus.     

Behavioral response to antennal tactile stimulation in the field cricket Gryllus bimaculatus

We examined behavioral responses of the field cricket Gryllus bimaculatus to tactile stimuli to the antennae. Three stimulants of similar shape and size but different textures were used: a tibia from the hunting spider Heteropoda venatoria (potential predator), a tibia from the orb-web spider Argiope bruennichi (less likely predator), and a glass rod. Each stimulus session comprised a first gentle contact and a second strong contact. The evoked behavioral responses were classified into four categories: aversion, aggression, antennal search, and no response. Regardless of the stimulants, the crickets exhibited antennal search and aversion most frequently in response to the first and second stimuli, respectively. The frequency of aversion was significantly higher to the tibia of H. venatoria than to other stimulants. The most striking observation was that aggressive responses were exclusive to the H. venatoria tibia. To specify the hair type that induced aggression, we manipulated two types of common hairs (bristle and fine) on the tibia of the predatory spider. When bristle hairs were removed from the H. venatoria tibia, aggression was significantly reduced. These results suggest that antennae can discriminate the tactile texture of external objects and elicit adaptive behavioral responses.     

四乙酰基植物鞘氨醇生物合成的研究进展

四乙酰基植物鞘氨醇(tetraacetyl phytosphingosine, TAPS)是一种性能卓越的天然护肤品原料,经去乙酰化后生成的植物鞘氨醇可作为前体合成保湿护肤品神经酰胺,因此广泛应用于护肤化妆品行业。非常规酵母威克汉姆西弗酵母(Wickerhamomyces ciferrii)是已知的唯一可天然分泌四乙酰基植物鞘氨醇的微生物,目前已成为四乙酰基植物鞘氨醇工业生产的宿主。本文介绍了四乙酰基植物鞘氨醇的发现、功能及其生物合成途径,综述了近年来利用单倍体筛选、诱变育种和代谢工程改造威克汉姆西弗酵母高产四乙酰基植物鞘氨醇的研究进展,并展望了实现四乙酰基植物鞘氨醇工业生产的未来发展方向。     

Bumblebees require visual pollen stimuli to initiate and multimodal stimuli to complete a full behavioral sequence in close‐range flower orientation

Flower visits are complex encounters, in which animals are attracted by floral signals, guided toward the site of the first physical contact with a flower, land, and finally take up floral rewards. At close range, signals of stamens and pollen play an important role to facilitate flower handling in bees, yet the pollen stimuli eliciting behavioral responses are poorly known. In this study, we test the response of flower‐naive bumblebees (Bombus terrestris) toward single and multimodal pollen stimuli as compared to natural dandelion pollen. As artificial pollen stimuli, we used the yellow flavonoid pigment quercetin, the scent compound eugenol, the amino acid proline, the monosaccharide glucose, and the texture of pollen‐grain‐sized glass pellets as a tactile stimulus. Three test stimuli, dandelion pollen, one out of various uni‐ and multimodal stimulus combinations, and a solvent control were presented simultaneously to individual bumblebees, whose response was recorded. The results indicate that bumblebees respond in an irreversible sequence of behavioral reactions. Bumblebees approached the visual stimulus quercetin as often as natural dandelion pollen. An additional olfactory stimulus resulted in slightly more frequent landings. The multimodal stimulus combinations including visual, olfactory, gustatory, and tactile stimuli elicited approaches, antennal contacts, and landings as often as natural pollen. Subsequent reactions like proboscis extension, mandible biting, and buzzing were more often but not regularly observed at dandelion pollen. Our study shows that visual signals of pollen are sufficient to trigger initial responses of bumblebees, whereas multimodal pollen stimuli elicit full behavioral response as compared to natural pollen. Our results suggest a major role of pollen cues for the attraction of bees toward flowers and also explain, why many floral guides mimic the visual signals of pollen and anthers, that is, the yellow and UV‐absorbing color, to direct bumblebees toward the site where they access the floral rewards.     

Cross-modal interactions between olfaction and touch

We report two experiments designed to investigate the nature of any cross-modal interactions between olfactory and tactile information processing. In Experiment 1, we assessed the influence of olfactory cues on the tactile perception of fabric softness using computer-controlled stimulus presentation. The results showed that participants rated fabric swatches as feeling significantly softer when presented with a lemon odor than when presented with an animal-like odor, demonstrating that olfactory cues can modulate tactile perception. In Experiment 2, we assessed whether this modulatory effect varied as a function of the particular odors being used and/or of the spatial coincidence between the olfactory and tactile stimuli. The results replicated those reported in Experiment 1 thus further supporting the claim that people's rating of tactile stimuli can be modulated by the presence of an odor. Taken together, the results of the two experiments reported here support the existence of a cross-modal interaction between olfaction and touch.     

Figuring space by time.

Sensory information is encoded both in space and in time. Spatial encoding is based on the identity of activated receptors, while temporal encoding is based on the timing of activation. In order to generate accurate internal representations of the external world, the brain must decode both types of encoded information, even when processing stationary stimuli. We review here evidence in support of a parallel processing scheme for spatially and temporally encoded information in the tactile system and discuss the advantages and limitations of sensory-derived temporal coding in both the tactile and visual systems. Based on a large body of data, we propose a dynamic theory for vision, which avoids the impediments of previous dynamic theories.