Low Threshold Points Research Articles

When do pupils talk about their problems? Explaining pupils’ intentions to seek help from school social work services

School social work aims to be a low threshold point of contact and advice for pupils. In practice, however, pupils often avoid seeking help when needed. Although the factors that influence help-seeking are well established in the literature, there has been very little research on this for school social work. Adapting the behavioral model of health services use, the present study addresses this gap by examining individual and contextual factors promoting or inhibiting pupils’ intentions to seek help from social work services directly integrated into their schools. A total of 4,420 pupils (grades 5 to 9) from 32 schools in Switzerland completed a questionnaire in which they were asked how likely they would be to seek help when they face a family problem, peer problem, or difficulties in everyday school life they cannot solve on their own. Results from the multi-level analysis reveal that, overall, pupils’ intentions to seek help are rather low. Predisposing factors and enabling factors influence willingness to seek help, with trust emerging as the most important facilitator. It is therefore important that all pupils can establish a trusting relationship with their school social worker and gain positive experiences with him or her.

The treatment of migraine patients with triptans – is there a need for further Rx-to-OTC switches?

Abstract Objective The disease migraine affects a large share of the German population and is linked to a high economic potential in terms of direct and indirect sickness costs. Triptans are the recommended treatment in the case of an acute migraine attack. Both, the disease and its appropriate treatment are of interest in the context of self-medication and Rx-to-OTC switch efforts. Therefore, a survey was carried out with the intention to collect data among migraine patients regarding the use of analgesics in general and of triptans specifically. This data can be the basis for further health economic considerations. Methods By an online survey among 206 migraine patients, different data regarding patients suffering from migraine, the frequency and sort of symptoms and the use of analgesics in general as well as triptans specifically was collected and analysed. A special focus was on symptoms affecting patients’ productivity, their use of triptans and their satisfaction with the current supply situation with triptans. Results The survey revealed among other findings that most patients suffer from symptoms which limit their productivity or their ability to work and make an immediate treatment necessary. Most patients know well about their disease and feel able to treat themselves in the context of self-medication once they are diagnosed by a physician. In this context a demand for further triptans available without a prescription could be identified. This is specifically, because patients respond differently to various triptans. Therefore, a larger variety of prescription free triptans would increase the number of patients with access to OTC triptans significantly. Conclusion Most survey respondents know well about their disease and the appropriate treatment and feel able to treat themselves within the scope of self-medication. Nevertheless, they mostly respond to one triptan only. In the case of an acute migraine attack an immediate treatment is required, ideal is an intake of triptans as soon as first symptoms occur. In this context pharmacies play an important role as fast and low-threshold access point to medications. Because only a limited number of patients responds to the already prescription free available triptans, there is a demand for further Rx-to-OTC switches of triptans among migraine patients which should be considered in further switch efforts in Germany.

Stabilization of weld pool through jet flow argon gas backing in C-Mn steel keyhole TIG welding

Welding tests were carried out to join the 16Mn steel with three different shielding modes, including no shielding, fully covered Argon gas, and backing jet flow Argon gas. It was found that: (1) it was hard to achieve a stable K-TIG welding process if the backside weld pool was exposed in the air; (2) stable keyhole welding process could be created in the current range of 410A–450A if the backside weld pool was covered with static argon gas; and (3) stable keyhole welding current range was increased to 420A–560A if the argon gas jets directly flowed towards the backside weld pool. The jet flow gas provides additional pressure supplementing gravity to balance the arc gas pressure. Fully penetrated weld could be obtained even though the welding current was beyond the upper threshold point. The welds obtained in higher currents had comparable quality to those obtained at the low threshold point. The results give new solution to stabilize the fully penetrated weld pool.

Auditory sensitivity to local stimulation of the head surface in a beluga whale (Delphinapterus leucas).

Using the auditory evoked response technique, sensitivity to local acoustic stimulation of the ventro-lateral head surface was investigated in a beluga whale (Delphinapterus leucas). The stimuli were tone pip trains of carrier frequencies ranging from 16 to 128 kHz with a pip rate of 1 kHz. For higher frequencies (90-128 kHz), the low-threshold point was located next to the medial side of the middle portion of the lower jaw. For middle (32-64 kHz) and lower (16-22.5 kHz) frequencies, the low-threshold point was located at the lateral side of the middle portion of the lower jaw. For lower frequencies, there was an additional low-threshold point next to the bulla-meatus complex. Based on these data, several frequency-specific paths of sound conduction to the auditory bulla are suggested: (i) through an area on the lateral surface of the lower jaw and further through the intra-jaw fat-body channel (for a wide frequency range); (ii) through an area on the ventro-lateral head surface and further through the medial opening of the lower jaw and intra-jaw fat-body channel (for a high-frequency range); and (iii) through an area on the lateral (near meatus) head surface and further through the lateral fat-body channel (for a low-frequency range).

An engineered mammalian band-pass network

Gene expression circuitries, which enable cells to detect precise levels within a morphogen concentration gradient, have a pivotal impact on biological processes such as embryonic pattern formation, paracrine and autocrine signalling, and cellular migration. We present the rational synthesis of a synthetic genetic circuit exhibiting band-pass detection characteristics. The components, involving multiply linked mammalian trans-activator and -repressor control systems, were selected and fine-tuned to enable the detection of ‘low-threshold’ morphogen (tetracycline) concentrations, in which target gene expression was triggered, and a ‘high-threshold’ concentration, in which expression was muted. In silico predictions and supporting experimental findings indicated that the key criterion for functional band-pass detection was the matching of componentry that enabled sufficient separation of the low and high threshold points. Using the circuitry together with a fluorescence-encoded target gene, mammalian cells were genetically engineered to be capable of forming a band-like pattern of differentiation in response to a tetracycline chemical gradient. Synthetic gene networks designed to emulate naturally occurring gene behaviours provide not only insight into biological processes, but may also foster progress in future tissue engineering, gene therapy and biosensing applications.

Position of spinothalamic tract axons in upper cervical spinal cord of monkeys.

Percutaneous upper cervical cordotomy continues to be performed on patients suffering from several types of severe chronic pain. It is believed that the operation is effective because it cuts the spinothalamic tract (STT), a primary pathway carrying nociceptive information from the spinal cord to the brain in humans. In recent years, there has been controversy regarding the location of STT axons within the spinal cord. The aim of this study was to determine the locations of STT axons within the spinal cord white matter of C2 segment in monkeys using methods of antidromic activation. Twenty lumbar STT cells were isolated. Eleven were classified as wide dynamic range neurons, six as high-threshold cells, and three as low-threshold cells. Eleven STT neurons were recorded in the deep dorsal horn and nine in superficial dorsal horn. The axons of the examined neurons were located at antidromic low-threshold points (<30 microA) within the contralateral lateral funiculus of C2. All low-threshold points were located ventral to the denticulate ligament, within the lateral half of the ventral lateral funiculus (VLF). None were found in the dorsal half of the lateral funiculus. The present findings support our previous suggestion that STT axons migrate ventrally as they ascend the length of the spinal cord. Also, the present findings indicate that surgical cordotomies that interrupt the VLF in C2 likely disrupt the entire lumbar STT.

Locations of spinothalamic tract axons in cervical and thoracic spinal cord white matter in monkeys.

The spinothalamic tract (STT) is the primary pathway carrying nociceptive information from the spinal cord to the brain in humans. The aim of this study was to understand better the organization of STT axons within the spinal cord white matter of monkeys. The location of STT axons was determined using method of antidromic activation. Twenty-six lumbar STT cells were isolated. Nineteen were classified as wide dynamic range neurons and seven as high-threshold cells. Fifteen STT neurons were recorded in the deep dorsal horn (DDH) and 11 in superficial dorsal horn (SDH). The axons of 26 STT neurons were located at 73 low-threshold points (<30 microA) within the lateral funiculus from T(9) to C(6). STT neurons in the SDH were activated from 33 low-threshold points, neurons in the DDH from 40 low-threshold points. In lower thoracic segments, SDH neurons were antidromically activated from low-threshold points at the dorsal-ventral level of the denticulate ligament. Neurons in the DDH were activated from points located slightly ventral, within the ventral lateral funiculus. At higher segmental levels, axons from SDH neurons continued in a position dorsal to those of neurons in the DDH. However, axons from neurons in both areas of the gray matter were activated from points located in more ventral positions within the lateral funiculus. Unlike the suggestions in several previous reports, the present findings indicate that STT axons originating in the lumbar cord shift into increasingly ventral positions as they ascend the length of the spinal cord.

Spinohypothalamic tract neurons in the cervical enlargement of rats: locations of antidromically identified ascending axons and their collateral branches in the contralateral brain.

Antidromic activation was used to determine the locations of ascending spinohypothalamic tract (SHT) axons and their collateral projections within C1, medulla, pons, midbrain, and caudal thalamus. Sixty-four neurons in the cervical enlargement were antidromically activated initially by stimulation within the contralateral hypothalamus. All but one of the examined SHT neurons responded either preferentially or specifically to noxious mechanical stimuli. A total of 239 low-threshold points was classified as originating from 64 ascending (or parent) SHT axons. Within C1, 38 ascending SHT axons were antidromically activated. These were located primarily in the dorsal half of the lateral funiculus. Within the medulla, the 29 examined ascending SHT axons were located ventrolaterally, within or adjacent to the lateral reticular nucleus or nucleus ambiguus. Within the pons, the 25 examined ascending SHT axons were located primarily surrounding the facial nucleus and the superior olivary complex. Within the caudal midbrain, the 23 examined SHT ascending axons coursed dorsally in a position adjacent to the lateral lemniscus. Within the anterior midbrain, SHT axons traveled rostrally near the brachium of the inferior colliculus. Within the posterior thalamus, all 17 examined SHT axons coursed rostrally through the posterior nucleus of thalamus. A total of 114 low-threshold points was classified as collateral branch points. Sixteen collateral branches were seen in C1; these were located primarily int he deep dorsal horn. Forty-five collateral branches were located in the medulla. These were primarily in or near the medullary reticular nucleus, nucleus ambiguus, lateral reticular nucleus, parvocellular reticular nucleus, gigantocellular reticular nucleus, cuneate nucleus, and the nucleus of the solitary tract. Twentysix collateral branches from SHT axons were located in the pons. These were in the pontine reticular nucleus caudalis, gigantocellular reticular nucleus, parvocellular reticular nucleus, and superior olivary complex. Twenty-three collateral branches were located in the midbrain. These were in or near the mesencephalic reticular nucleus, brachium of the inferior colliculus, cuneiform nucleus, superior colliculus, central gray, and substantia nigra. Int he caudal thalamus, two branches were in the posterior thalamic nucleus and two were in the medial geniculate. These results indicate that SHT axons ascend toward the hypothalamus in a clearly circumscribed projection in the lateral brain stem and posterior thalamus. In addition, large numbers of collaterals from SHT axons appears to project to a variety of targets in C1, the medulla, pons, midbrain, and caudal thalamus. Through its widespread collateral projections, the SHT appears to be capable of providing nociceptive input to many areas that are involved in the production of multifaceted responses to noxious stimuli.

Dorsal horn projection targets of ON and OFF cells in the rostral ventromedial medulla.

1. The rostral ventromedial medulla (RVM) participates in the modulation of nociceptive transmission by spinal cord neurons. Previous anatomic studies have demonstrated that RVM neurons project to laminae I, II, and V of the dorsal horn; laminae VII and VIII of the intermediate and ventral horns; the intermediolateral column; and lamina X. The RVM contains at least three physiologically defined classes of neurons, two of which, the ON and the OFF cells, have been implicated in nociceptive modulation. Because these cells classes are intermingled in the RVM, it has not been possible to determine the spinal laminar projection targets of ON and OFF cells by anatomic methods. Therefore in the current study we employed antidromic microstimulation methods to determine the laminar projections of two of the three classes of RVM neurons, the ON and the OFF cells. 2. In lightly anesthetized (with methohexital sodium) rats, single-unit extracellular recordings were made from 48 RVM neurons that were physiologically characterized as ON (30) or OFF (18) cells. The recording locations of 45 of these neurons were recovered. Thirty-seven were found in the nucleus raphe magnus and eight were located near its dorsal and lateral borders. 3. Thirty-two physiologically identified RVM neurons (18 ON and 14 OFF cells) were antidromically activated from the cervical spinal cord using a monopolar stimulating electrode. The stimulating electrode was moved systematically in the white matter until antidromic activation could be produced with currents of < or = 20 microA (6.1 +/- 0.7 microA, mean +/- SE). The points from which minimum currents were required to antidromically activate the neurons were located mainly in the ipsilateral dorsolateral funiculus (DLF) (27 of 32). In a few cases, lowest antidromic threshold currents were found near the border between the DLF and ventrolateral funiculus (VLF) or, rarely, in the VLF itself. In these cases, the cell recordings were found to be near the dorsal boundary of the RVM. 4. While one electrode was used to stimulate the parent axon in the lateral funiculus, a second was used to explore the gray matter for the presence of collateral branches. The identification of a branch was initially determined by an increase in antidromic latency. At the same rostrocaudal plane of the spinal cord, stimulation of the DLF induced an antidromic spike that invaded the neuron earlier than the antidromic spike elicited by stimulation in the gray matter. Collateral branches were confirmed by establishing that the location of the minimum threshold point for antidromic activation of the neurons from the second electrode was in the gray matter, that the minimum current required to antidromically activate the neuron from that point was too low to activate the parent axon in the DLF, and that a collision occurred between the spikes induced by the two stimulating electrodes. 5. In 17 cases, physiologically identified RVM neurons (10 ON and 7 OFF cells) were antidromically activated from the gray matter of the cervical spinal cord using a current of 8.4 +/- 2.1 (SE) microA. Minimum threshold points for antidromic activation were found in laminae I-II (3 ON and 4 OFF cells), lamina V (5 ON and 6 OFF cells), and regions ventral to the lateral reticulated area (3 ON and 2 OFF cells) of the gray matter. As indicated by these numbers, some neurons were antidromically activated from more than one gray matter region. In general, all OFF cells and 9 of 10 ON cells were antidromically activated from low threshold points in either laminae I-II or lamina V. 6. In six cases, neurons were activated from separate points located in two or three different laminae of the gray matter. Three OFF cells were activated from laminae I-II and V, one OFF cell and one ON cell were activated from lamina V and from more ventral points, and one ON cell was activated from laminae I-II and from points ventral to lamina V.

Spinothalamic and spinohypothalamic tract neurons in the cervical enlargement of rats. III. Locations of antidromically identified axons in the cervical cord white matter

1. Fifty-five neurons in the cervical enlargement (C6-C8) of urethan-anesthetized rats were antidromically activated from the contralateral posterior diencephalon. In all cases, antidromic thresholds were < or = 30 microA. The locations of the axons of these neurons within the white matter of segments C2-C6 were determined by tracking systematically using a second antidromic stimulating electrode. 2. The recording locations of 51 neurons were marked and recovered. Twenty neurons were recorded in the superficial dorsal horn (SDH) and 31 were in the deep dorsal horn (DDH). Eighty-three lowest threshold points for antidromic activation within the white matter of segments C2-C6 were determined for these 51 neurons. The mean antidromic threshold at these points was 9.5 +/- 0.5 (SE) microA. For 26 neurons, the lowest threshold point for antidromic activation was determined at one segmental level. We also attempted to determine whether individual axons maintained their position as they ascended through the cervical cord white matter. In 25 cases, lowest threshold points were determined at two or more segmental levels. 3. In segments C5-C6, 88% (7/8) of the lowest threshold points of the examined axons were located in the contralateral ventral funiculus, indicating that the majority of examined axons crossed the midline within one or two segments. 4. In segments C3-C4, 32% (14/44) of all examined axons were found in the dorsal lateral funiculus (DLF) and 66% (29/44) were within the ventral quadrant [ventral lateral funiculus (VLF) and ventral funiculus (VF)]. Sixty-nine percent (11/16) of the axons of neurons recorded in the SDH were located in the contralateral DLF and 31% (5/16) were located in the ventral quadrant (VQ). In contrast, only 11% (3/28) of the axons of neurons recorded in the DDH were located in the contralateral DLF and 86% (24/28) were located in the VQ. Therefore, in segments C3-C4, the locations of axons differed significantly. Those from neurons recorded in the SDH were located primarily in the DLF and those from neurons recorded in the DDH were located principally in the VQ. 5. In segment C2, 74% (23/31) of all examined axons were found in the DLF, 23% (7/31) were in the VQ, and 3% (1/31) were in the dorsal horn. Thus, the percentage of all examined axons in the DLF in C2 was approximately 2.5 times greater than it was in C3-C4.(ABSTRACT TRUNCATED AT 400 WORDS)

Magnetic coil stimulation of straight and bent amphibian and mammalian peripheral nerve in vitro: locus of excitation.

1. According to classical cable theory, a magnetic coil (MC) should excite a linear nerve fibre in a homogeneous medium at the negative-going first spatial derivative of the induced electric field. This prediction was tested by MC stimulation of mammalian phrenic and amphibian sciatic nerve and branches in vitro, immersed in Ringer solution within a trough, and identifying the sites of excitation by recording responses of similar latency to local electrical stimulation. Subsequently, the identified sites of excitation were compared with measurements of the induced electric field and its calculated first spatial derivative. A special hardware device was used to selectively reverse MC current direction and to generate predominantly monophasic- or polyphasic-induced pulse profiles whose initial phases were identical in polarity, shape and amplitude. When using the amphibian nerve preparation, a complication was excitation at low threshold points related to cut branches. 2. Reversal of monophasic current resulted in latency shifts corresponding approximately to the distance between induced cathode and anode. The location of each site of excitation was at, or very near, the negative-going first spatial derivative peaks of the induced electric field measured parallel to the straight nerve. Significantly, excitation of the nerve did not occur at the peak of the induced electric field above the centre of the 'figure of eight' MC junction. 3. A polyphasic pulse excited the nerve at both sites, by the negative-going first phase at one location, and approximately 150 microseconds later, by the reversed negative-going second phase at the other location. Polyphasic and monophasic pulses elicited responses with similar latency when the induced current flowed towards the recording electrode. 4. Straddling a nerve with non-coding solid lucite cylinders created a localized spatial narrowing and increase in the induced electric field, resulting in a lowered threshold of excitation. The corresponding closer spacing between first spatial derivative peaks was exhibited by a significant reduction in latency shift when MC current direction was reversed. 5. When a nerve is bent and the induced current is directed along the nerve towards the bend, the threshold of excitation is reduced there. Increasing the angle of the bend from 0 deg to more than 90 deg graded the decrease in threshold. 6. In a straight nerve the threshold was lowest when current was directed towards the cut end.(ABSTRACT TRUNCATED AT 400 WORDS)

Reticulospinal and reticuloreticular pathways for activating the lumbar back muscles in the rat.

These experiments tested hypotheses about the logic of reticulospinal and reticuloreticular controls over deep back muscles by examining descending efferent and contralateral projections of the sites within the medullary reticular formation (MRF) that evoke EMG responses in lumbar axial muscles upon electrical stimulation. In the first series of experiments, retrograde tracers were deposited at gigantocellular reticular nucleus (Gi) sites that excited the back muscles and in the contralateral lumbar spinal cord. The medullary reticular formation contralateral to the Gi stimulation/deposition site was examined for the presence of single- and double-labeled cells from these injections. Tracer depositions into Gi produced labeled cells in the contralateral Gi and Parvocellular reticular nucleus (PCRt) whereas the lumbar injections retrogradely labeled cells only in the ventral MRF, indicating that separate populations of medullary reticular cells project to the opposite MRF and the lumbar cord. In the second series of experiments the precise relationships between the location of neurons retrogradely labeled from lumbar spinal cord depositions of the retrograde trace, Fluoro-Gold (FG) and effective stimulation tracks through the MRF were examined. The results indicate that the Gi sites that are most effective for activation of the back muscles are dorsal to the location of retrogradely labeled lumbar reticulospinal cells. To verify that cell bodies and not fibers of passage were stimulated, crystals of the excitatory amino acid agonist, N-methyl-D-aspartate (NMDA) were deposited at effective stimulation sites in the Gi. NMDA decreased the ability of electrical stimulation to activate back muscles at 5 min postdeposition, indicating a local interaction of NMDA with cell bodies at the stimulation site. In the third series of experiments, electrical thresholds for EMG activation along a track through the MRF were compared to cells retrogradely labeled from FG deposited into the cervical spinal cord. In some experiments, Fast Blue was also deposited into the contralateral lumbar cord. Neurons at low threshold points on the electrode track were labeled following cervical depositions, indicating a direct projection to the cervical spinal cord. The lumbar depositions, again, labeled cells in MRF areas that were ventral to the locations of effective stimulation sites, primarily on the opposite side of the medulla. In addition, the lumbar depositions back-filled cells in the same cervical segments to which the Gi neurons project. These results suggest that one efferent projection from effective stimulation sites for back muscle activation is onto propriospinal neurons in the cervical cord, which in turn project to lumbar cord levels.(ABSTRACT TRUNCATED AT 400 WORDS)

Integration in descending motor pathways controlling the forelimb in the cat

Collateralization and termination of single C3-C4 propriospinal neurones (PNs) have been studied in the C6-Th1 segments of the cat using two methods: threshold mapping for antidromic activation of C3-C4 PNs and intra-axonal injection of horseradish peroxidase. Low threshold points for antidromic activation of C3-C4 PNs were found in the region of different motor nuclei in lamina IX both at one level and at different segmental levels, in all parts of lamina VII, in the lateral part of lamina VI and in the dorsal and ventral parts of lamina VIII. Collaterals were found from C6 to Th1. A marked decrease of conduction velocity of the stem axon occurred in the caudal region of termination, while it was almost constant in the rostral region of termination. HRP was injected iontophoretically in C6-Th1 into stem axons of neurones, which were activated antidromically from the ventral part of the lateral funiculus in C5/C6, from the lateral reticular nucleus (LRN) and monosynaptically from the corticospinal fibres (stimulated in the contralateral pyramid) which were transected in C5/C6. Reconstruction of successfully stained stem axons, revealed collaterals with terminals on presumed motoneurones in different parts of lamina IX and on interneurones in laminae IV-VIII. These findings confirm previous results which showed monosynaptic projections from C3-C4 PNs to forelimb motoneurones and Ia inhibitory interneurones. With respect to termination in the region of the motoneurones in lamina IX and in the region of Ia inhibitory interneurones in lamina VII, three patterns were found: 1) termination mainly in lamina IX (n = 1) 2) termination in laminae IX and VII (n = 15) and 3) termination mainly in lamina VII (n = 2). However, in some cases the same stem axon gave off collaterals which terminated either on motoneurones in lamina IX or on presumed Ia inhibitory interneurones in lamina VII. Furthermore, when the stem axons had collaterals which terminated in different motor nuclei only some of these collaterals had additional terminations on presumed Ia inhibitory interneurones. This result suggest that C3-C4 PNs do not follow a strict Ia pattern of reciprocal innervation. It is tentatively proposed that the difference of innervation may be related to the type of multi-joint movement, such as target-reaching with the forelimb, which has been shown to be controlled by the C3-C4 PNs. Termination in laminae VI, VIII and different parts of lamina VII indicates that C3-C4 PNs also project to other types of neurones than motoneurones and Ia inhibitory interneurones.(ABSTRACT TRUNCATED AT 400 WORDS)

Projections of the group y of the vestibular nuclei and the dentate and fastigial nuclei of the cerebellum to the interstitial nucleus of Cajal

Experiments were performed to study the projection of the group y of the vestibular nuclei and the dentate and fastigial nuclei of the cerebellum to the interstitial nucleus of Cajal (INC) in cats by using retrograde axonal transport of horseradish peroxidase (HRP) and electrophysiological methods; and to study the vestibular responses of such projection neurons. Following injections of HRP into the unilateral INC, with partial involvement of the surrounding reticular formation, including the nucleus of Darkschewitsch (ND), many retrogradely labeled neurons were found in the dorsal part of the group y nucleus contralateral to the injection site. Labeled cells were also seen in the contralateral dentate nucleus, frequently in its caudal-ventral part, and in the contralateral fastigial nucleus at all rostrocaudal levels, but most frequently in its caudal part. In electrophysiological experiments performed on cats anesthetized with α-chloralose or N 2O and paralyzed with gallamine, group y, dentate and fastigial nuclei neurons were antidromically activated by weak stimuli that were confined to the contralateral INC. Depth-threshold curves for antidromic activation of such neurons revealed that the lowest threshold points were within the INC, but not in the ND. The INC-projecting neurons in the group y and dentate nuclei did not respond to electrical stimulation of the ipsilateral or contralateral vestibular nerve, indicating that they do not receive direct labyrinthine inputs. On the contrary, many fastigial neurons projecting to the INC responded to labyrinthine stimulation, suggesting that they may be involved in the vestibular reflexes. These results suggest a difference in properties of INC-projecting neurons in these nuclei.

Augmenting responses evoked in area 17 of the cat by intracortical axon collaterals of cortico-geniculate cells.

Evoked potentials were recorded in the visual cortex of the cat after electrical stimulation of the lateral geniculate nucleus (l.g.n.). The primary response, mediated by geniculo-cortical fibres, was depressed at stimulation frequencies above 7 Hz and replaced by a late potential, the incremental response, which gradually increased in amplitude with successive stimuli. The incremental response was a negative-positive potential in the depth of the cortex with the negative component having maximal amplitude in layer 4. The response reversed polarity in layer 1 to become a positive-negative potential at the surface. The latency of the negative component of the incremental response was about 3.5-4 ms in layer 4, compared to about 1.5 and 2.5 ms for the mono- and disynaptic components of the primary response. The incremental response could only be evoked from the l.g.n. and the optic radiation, not from the optic tract, superior colliculus or other surrounding structures. Within the l.g.n., the effect was only evoked from stimulation sites in approximate retinotopic register with the recording site in the cortex. Low threshold points were found in the A laminae, completely overlapping with the low threshold points for the primary response. Thresholds increased steeply when the stimulation electrode was lowered into the C laminae. The incremental response could still be evoked ten days after the destruction of all cells in the l.g.n. complex by kainic acid. It is concluded that the described incremental response is identical to the augmenting response of Dempsey & Morison (1943) and is mediated by intracortical axon collaterals of antidromically activated cortico-geniculate neurones.

Responses of vestibular neurons to stimulation of the interstitial nucleus of Cajal in the cat.

(1) Spikes of neurons in the medial and descending vestibular nuclei were recorded extracellularly and their responses to stimulation of the interstitial nucleus of Cajal (INC) were studied in cerebellectomized cats under chloralose anesthesia. Stimuli applied in the ipsilateral INC excited 37% of neurons that did not exhibit spontaneous activity. About 84% of spontaneously discharging neurons were influenced by the INC; typical responses were excitation (35%), inhibition (22%) and excitation followed by inhibition (27%). Of the neurons that were excited, 24% fired monosynaptically. Such monosynaptic activation was evoked by stimulating the INC and midbrain medial longitudinal fasciculus (MLF), but was not evoked by stimulating the lateral midbrain reticular formation. Polysynaptic excitation or inhibition was evoked more widely, but the lowest threshold points were within the INC. Stimulation of the contralateral INC also evoked polysynaptic excitation or inhibition. However, the frequency of occurrence of the evoked responses was significantly smaller compared to the ipsilateral responses. (2) Intracellular recordings revealed that some medial and lateral vestibular neurons received monosynaptic excitatory postsynaptic potentials (EPSPs), others received polysynaptic EPSPs or inhibitory postsynaptic potentials (IPSPs) from the ipsilateral INC. The minimum latency for the IPSPs suggests that the pathway is at least disynaptic. No significant collision was observed between monosynaptic EPSPs evoked by the ipsilateral INC and contralateral vestibular nuclei. Acute lesions that damaged the pontine MLF and part of the reticular formation did not abolish monosynaptic responses of vestibular neurons by the INC. Depth threshold curves for mono- or polysynaptic responses drawn before and after the lesions were virtually similar. Antidromic thresholds of interstitio-vestibular fibers evoked from the pontine MLF showed that a great majority of these fibers run outside the MLF at the pontine level. These results control for vestibular axon reflexes, since vestibulo-interstitial fibers ascend within the MLF (cf. Gacek 1971). (3) Responses to stimulation of the INC were not different among different types of canal responding neurons; vertical and horizontal canal responding neurons received similar effects. However, canal responding neurons that received excitation from the contralateral vestibular nerve were activated more frequently by the INC than those that received inhibition (χ2 test, p<0.01). Qualitatively similar results were obtained from vestibular neurons that had different projection sites; vestibulospinal, contralateral INC-projecting and contralateral vestibular nuclei-projecting neurons received similar effects. (4) Vestibulo-collic reflexes, studied with EMG, were modified by preceding INC stimulation. Intracellular recordings from some neck motoneurons showed that disynaptic EPSPs evoked by stimulation of the contralateral vestibular nerve were modified by preceding INC stimulation applied ipsilateral to the stimulated vestibular nerve. INC stimulation alone did not evoke any response in these motoneurons, suggesting that the interaction of the labyrinthine and interstitial effects occurred at least in part at the vestibular nuclei. (5) Some medial and descending vestibular neurons showed multiple branching, projecting to the contralateral INC, C1 or contralateral vestibular nuclei. About 34% of neurons that projected to the contralateral INC were also antidromically activated from the C1; some of them received vertical canal inputs.

Responses of cat mesencephalic reticulospinal neurons to stimulation of superior colliculus, pericruciate cortex, and neck muscle afferents.

Neurons were recorded extracellularly in the mesencephalic reticular formation outside the interstitial nucleus of Cajal in cerebellectomized cats anesthetized with alpha chloralose. Reticulospinal neurones were identified by antidromic stimulation of the upper cervical segments. Stimulation in the deep layers of the ipsilateral superior colliculus evoked firing in 36% of reticulospinal neurons. For many neurons thresholds for activation were high in the intermediate tectal layers and declined as the electrodes entered the underlying tegmentum. However, low threshold points were found above the deep fiber layer within the superior colliculus for some cells. Stimulation of the contralateral superior colliculus excited 10% of neurons and thresholds for activation were high above the deep fiber layer for all neurons. Stimulation of the ipsilateral and contralateral pericruciate cortex excited 39 and 21% of neurons, respectively. The lowest threshold area was found in the frontal eye fields. Sixteen percent of neurons received excitation from neck muscle afferents (C2 biventer-cervicis) bilaterally. Comparison of responses between mesencephalic reticulospinal neurons and interstitiospinal neurons (Fukushima et al. 1981) showed that responses of the two groups of neurons were similar when the pericruciate cortex and neck muscle afferents were stimulated. However, a difference was observed in tectal responses. since low threshold points were rarely observed above the deep fiber layer for interstitiospinal neurons.

Effects of stimulation of frontal cortex, superior colliculus, and neck muscle afferents on interstitiospinal neurons in the cat.

Interstitiospinal neurons were activated by antidromic stimulation of the spinal cord ventromedial funiculus at C1 and C4 in cerebellectomized cats under chloralose anesthesia. Neurons responding only to C1 were classified as N cells and those responding both to C1 and C4 were classified as D cells, as in previous experiments (Fukushima et al. 1980a). Vestibular branching interstitiospinal and reticulospinal neurons were also identified as in the previous experiments. Stimulation of the ipsilateral pericruciate cortex evoked firing in 31% of N cells, 41% of D cells and 35% of the vestibular branching neurons, while stimulation of the contralateral cortex excited 6% of N cells, 29% of D cells and 14% of vestibular branching neurons. Response latencies ranged from 2 to 15 ms after the effective pulse. By measuring the thresholds of activation of these neurons while changing the depth of the stimulating electrodes, and by mapping the cortical areas, it was shown that the lowest threshold areas were in the frontal eye fields and the anterior sigmoid gyrus near the presylvian sulcus (Area 6). Stimulation of the latter area often evoked neck or shoulder muscles contraction. Stimulation in the deep layers of the ipsilateral superior colliculus evoked firing in about 20% of interstitiospinal neurons and about 42% of vestibular branching neurons, with typical latencies 2-3 ms after the effective pulse, while stimulation of the contralateral superior colliculus was rarely effective. N cells and D cells responded similarly. Thresholds for activation were high in the intermediate tectal layers and declined as the electrodes entered the underlying tegmentum. This suggests that the superior colliculus is not the main source of synaptic inputs to these neurons. Low threshold points were found above the deep fiber layer when stimulating electrodes were inserted into the pretectum. Stimulation of the C2 biventer cervicis nerve excited about 8% of N cells, 18% of D cells, and 15% of vestibular branching neurons bilaterally with typical latencies around 10 ms. Similar results were obtained when C2 splenius nerves were stimulated. The fibers responsible for such excitation are probably group II, since stimuli stronger than 1.8 times threshold of the lowest threshold fibers were needed to evoke excitation. Response decrement was often observed when stimuli were repeated at 1/s, while no such decrement was observed at the rate of 1/3 s. When the convergence of cortical and labyrinthine excitatory inputs was studied, 36% of interstitiospinal neurons received single inputs either from the pericruciate cortex or from the labyrinth, 22% of neurons received convergent excitation from both and the remaining 42% did not respond to either stimulus. Although vestibular branching neurons rarely received labyrinthine inputs, they frequently showed convergence of excitation to stimulation of the frontal cortex, superior colliculus and vestibular nuclei.

Organization of the face representation in macaque motor cortex.

We stimulated with microelectrodes the face representation in precentral motor cortex in macaque monkeys. Responses were very discrete; at threshold current levels the usual response was a small focus of movement in part of a muscle. Facial muscles cluster together in the posterior and anterior portions of the precentral gyrus with tongue movements represented in the intervening region and along the lateral extent. Within each cluster there are multiple representations of individual muscle movements. In long penetrations down the anterior wall of the central sulcus we were able to advance the electrode tangentially through cortex. In these penetrations we encountered a series of discrete zones each of which was related to the movement of a particular muscle or part of a muscle in the face. The lowest threshold points were found in the center of each zone, and as the microelectrode progressed toward the edge, thresholds rose until there was a shift to a new muscle movement. Successive stimulation points separated by as little as 50 micrometer could yield different responses. These zones could be either roughly cylindrical or take the form of narrow curving bands running mediolaterally across cortex. There is a tendency for adjacent muscles to occur together, and the representation may be roughly topographical within the limits set by the morphological structure of the muscles themselves. The most commonly evoked muscle response was in zygomaticus, which retracts the corners of the mouth in expressions of fear and anger.

Low-threshold mechanisms and cortical activation pathways.

In chronic experiments with alert cats, stimulation with electrical current of moderate strength at various points on the lateral surface of the cerebral hemispheres leads to activation of various brain regions. In addition to high-threshold cortical points, low-threshold points have been discovered which are located in the sensorimotor region and in the Ep field of the auditory zone. The latter possess the same low thresholds for evoking an activation response as do points in the mesencephalic RF and the thalamic CM, VPL, and GM. Connections have been discovered (in the morphological part of the study) between the auditory Ep field and the intralaminar nuclei of the thalamus and the brain-stem part of the RF; the major projections run from the dorsal part of the Ep field into the lateral zone of the tegmentum. It is proposed that the role of the cortical low-threshold foci could involve the triggering of the nonspecific activation apparatus in accord with the biological significance of the signals being analyzed.