Cat Motoneurons Research Articles

Morphological analysis of neck muscle nerves and neurons in cats

Previous studies have failed to show morphological differences between neck muscle alpha and gamma motor fibers or alpha and gamma motoneurons. The present study aimed to investigate the morphological features of neck muscle motor nerves and motoneurons in cats. To determine the morphological features of peripheral motor fibers, the value of the outer contours of each fiber was converted into a perfect circle after ganglionectomy to remove sensory fibers, and the fiber diameters were calculated based on their circumferences. The sizes of neck motor fibers in the peripheral nerves had an evident bimodal distribution into small and large fiber groups, as depicted in histograms. The sizes of small and large motor fibers ranged from 2 to 12 µm and from 12 to 40 µm, respectively. The small fiber group is likely to correspond to gamma motor fibers and the large fiber group to alpha motor fibers. The morphological features of neck muscle motoneurons sectioned in the horizontal plane were examined using the horseradish peroxidase (HRP) retrograde labeling technique. The diameters of the biventer cervicis and complexus motoneurons had bimodal distributions. The inflection point between the small and large diameter population was 28 µm for the biventer cervicis and 26 µm for the complexus. We also observed that larger neurons displayed more dendrites. In conclusion, we could identify morphological differences likely to correlate with alpha and gamma motoneurons in both neck muscle peripheral nerves and neck motoneurons.

Enlargement of the Nerve Fibers of Silenced Lumbosacral Motoneurons in Cats.

Whether neuromuscular activity influences the size of motor nerves is controversial. All neuromuscular activity in cat hindlimbs was eliminated by spinal cord isolation (SCI), namely, spinal cord transection above and below the medial gastrocnemius (MG) and soleus (SOL) motoneuron pools and L5-S3 dorsal root transection. MG, SOL and sural (SUR) nerves were removed for size measurements, eight months after SCI surgery and from age-matched control cats. Nerve fiber number, the linear relationship between axon size and myelin thickness, and the bimodal distributions of nerve fiber area and diameter were maintained in all three nerves after SCI. The distributions of myelinated sensory fibers were unchanged in SUR nerves in contrast to the myelinated motor fibers in the MG and SOL nerves that were significantly larger. These findings provide evidence that all lumbar motoneurons survive SCI and that their nerve fibers enlarge. Thus, motor nerve fiber size in addition to the properties of the motoneurons and their muscle fibers is dynamic, responding to neuromuscular activity.

Synaptic control of the shape of the motoneuron pool input-output function.

Although motoneurons have often been considered to be fairly linear transducers of synaptic input, recent evidence suggests that strong persistent inward currents (PICs) in motoneurons allow neuromodulatory and inhibitory synaptic inputs to induce large nonlinearities in the relation between the level of excitatory input and motor output. To try to estimate the possible extent of this nonlinearity, we developed a pool of model motoneurons designed to replicate the characteristics of motoneuron input-output properties measured in medial gastrocnemius motoneurons in the decerebrate cat with voltage-clamp and current-clamp techniques. We drove the model pool with a range of synaptic inputs consisting of various mixtures of excitation, inhibition, and neuromodulation. We then looked at the relation between excitatory drive and total pool output. Our results revealed that the PICs not only enhance gain but also induce a strong nonlinearity in the relation between the average firing rate of the motoneuron pool and the level of excitatory input. The relation between the total simulated force output and input was somewhat more linear because of higher force outputs in later-recruited units. We also found that the nonlinearity can be increased by increasing neuromodulatory input and/or balanced inhibitory input and minimized by a reciprocal, push-pull pattern of inhibition. We consider the possibility that a flexible input-output function may allow motor output to be tuned to match the widely varying demands of the normal motor repertoire.NEW & NOTEWORTHY Motoneuron activity is generally considered to reflect the level of excitatory drive. However, the activation of voltage-dependent intrinsic conductances can distort the relation between excitatory drive and the total output of a pool of motoneurons. Using a pool of realistic motoneuron models, we show that pool output can be a highly nonlinear function of synaptic input but linearity can be achieved through adjusting the time course of excitatory and inhibitory synaptic inputs.

Response: Commentary: Age-related neurodegenerative disease research needs aging models.

In response to my recent article (Johnson, 2015), Wallace and Howlett have provided a thoughtful commentary pointing out that aging might be defined not just by the passage of time, but by the accumulation of defects in multiple regions throughout the organism that can be represented by a frailty index (Wallace and Howlett, 2016). It occurs to me that this approach might help overcome two major barriers to aging research: (i) the time taken for animals to age, and (ii) the fact that animals tend to die as they age. In our research on aging (24 m-old) ad-libitum fed rats, we found approximately 50% of animals died before the age of 24 m. In contrast, caloric-restricted rats did not die before they reached 24 m, did not show typical age-related pathologies, and had motoneurones that responded differently to injury compared to age-matched ad-libitum-fed rats (Aperghis et al., 2003; Johnson and Duberley, 1998). For 10–15-years-old ad-libitum fed cats we also found increasing health problems with age, but no evidence that the spinal motoneurones in these aging cats were any more vulnerable to injury than those in 1–2-years-old cats (Johnson et al., 1991). It would be interesting to know if the creation in younger animals of age-related defects, such as those associated with metabolic syndrome or changes in inflammatory status, also causes nervous system pathologies typical of aging. Such models may well be quicker and cheaper than aging animals. They might also remove the criticism that results obtained from the study of aging animals are unrepresentative because they are based on the “survivors.” The use of a frailty index to determine age and perhaps create artificially aged models is an exciting concept, notwithstanding the possibility that the largely post-mitotic population of neurones in the nervous system may respond differently to cells in the rest of the organism.

Distribution and density of contacts from noradrenergic and serotonergic boutons on the dendrites of neck flexor motoneurons in the adult cat.

Serotonergic (5-HT) and noradrenergic (NA) input to spinal motoneurons is essential for generating plateau potentials and self-sustained discharges. Extensor motoneurons are densely innervated by 5-HT and NA synapses and have robust plateau potentials and self-sustained discharges. Conversely, plateau potentials and self-sustained discharges are very rare in flexor motoneurons. The most likely reasons for this difference are that flexor motoneurons have few 5-HT and NA synapses and/or they are distributed distant to the channels responsible for plateau potentials and self-sustained discharges. However, the distribution of 5-HT and NA synapses on flexor motoneurons is unknown. Here we describe the distribution and density of 5-HT and NA synapses on motoneurons that innervate the flexor neck muscle, rectus capitis anterior (RCA), in the adult cat. Using a combination of intracellular staining, fluorescent immunohistochemistry, and 3D reconstruction techniques, we found that 5-HT and NA synapses are widely distributed throughout the dendritic trees of RCA motoneurons, albeit with a strong bias to small-diameter dendrites and to medial dendrites in the case of NA contacts. The number of 5-HT and NA contacts per motoneuron ranged, respectively, from 381 to 1,430 and from 642 to 1,382, which is 2.3- and 1.4-fold less than neck extensor motoneurons (Montague et al., J Comp Neurol 2013;521:638-656). These results suggest that 5-HT and NA synapses on flexor motoneurons may provide a powerful means of amplifying synaptic currents without incurring plateau potentials or self-sustained discharges. This feature is well suited to meet the biomechanical demands imposed on flexor muscles during different motor tasks.

Postsynaptic inhibition of hypoglossal motoneurons produces atonia of the genioglossal muscle during rapid eye movement sleep.

Hypoglossal motoneurons were recorded intracellularly to determine whether postsynaptic inhibition or disfacilitation was responsible for atonia of the lingual muscles during rapid eye movement (REM) sleep. Intracellular records were obtained of the action potentials and subthreshold membrane potential activity of antidromically identified hypoglossal motoneurons in cats during wakefulness, nonrapid eye movement (NREM) sleep, and REM sleep. A cuff electrode was placed around the hypoglossal nerve to antidromically activate hypoglossal motoneurons. The state-dependent changes in membrane potential, spontaneous discharge, postsynaptic potentials, and rheobase of hypoglossal motoneurons were determined. During quiet wakefulness and NREM sleep, hypoglossal motoneurons exhibited spontaneous repetitive discharge. In the transition from NREM sleep to REM sleep, repetitive discharge ceased and the membrane potential began to hyperpolarize; maximal hyperpolarization (10.5 mV) persisted throughout REM sleep. During REM sleep there was a significant increase in rheobase, which was accompanied by barrages of large-amplitude inhibitory postsynaptic potentials (IPSPs), which were reversed following the intracellular injection of chloride ions. The latter result indicates that they were mediated by glycine; IPSPs were not present during wakefulness or NREM sleep. We conclude that hypoglossal motoneurons are postsynaptically inhibited during naturally occurring REM sleep; no evidence of disfacilitation was observed. The data also indicate that glycine receptor-mediated postsynaptic inhibition of hypoglossal motoneurons is crucial in promoting atonia of the lingual muscles during REM sleep.

Biomechanical benefits of the onion-skin motor unit control scheme

Muscle force is modulated by varying the number of active motor units and their firing rates. For the past five decades, the notion that the magnitude of the firing rates is directly related to motor unit size and recruitment threshold has been widely accepted. This construct, here named the After-hyperpolarization scheme evolved from observations in electrically stimulated cat motoneurons and from reported observations in voluntary contractions in humans. It supports the assumption that the firing rates of motor units match their mechanical properties to “optimize” force production, so that the firing rate range corresponds to that required for force-twitch fusion (tetanization) and effective graduation of muscle force. In contrast, we have shown that, at any time and force during isometric voluntary constant-force contractions in humans, the relationship between firing rate and recruitment threshold is inversely related. We refer to this construct as the Onion-Skin scheme because earlier-recruited motor units always have greater firing rates than latter-recruited ones. By applying a novel mathematical model that calculates the force produced by a muscle for the two schemes we found that the Onion-Skin scheme is more energy efficient, provides smoother muscle force at low to moderate force levels, and appears to be more conducive to evolutionary survival than the After-hyperpolarization scheme.

Delayed depolarization and firing behavior of human motoneurons during voluntary muscle contractions.

The firing behavior of motoneurons is governed by the interaction between the intrinsic properties of motoneurons and the synaptic inputs that they receive. In particular, long-lasting after-hyperpolarization (AHP) following each motoneuron spike and thus decreasing motoneuron excitability immediately after spike was found to be a key mechanism in controlling of interspike interval (ISI) duration (Eccles et al., 1958; Kernell, 1965). As a result, motoneurons commonly fire single discharges at rather low firing rates, especially during gentle voluntary muscle contractions and postural tasks. However, some motoneurons, under conditions of weak synaptic drive, can fire double discharges (doublets) with very short intradoublet ISIs (up to a few ms). Each doublet is commonly followed by a prolonged post-doublet (interdoublet) ISI resulting from the AHP summation (Calvin and Schwindt, 1972). Two types of doublets have been observed: single and repetitive ones. Single doublets have been reported many times in cat and human motoneurons (e.g., Eccles and Hoff, 1932; Hoff and Grant, 1944; Denslow, 1948; Calvin and Schwindt, 1972; Kudina, 1974; Kudina and Churikova, 1990; Kirkwood and Munson, 1996; Piotrkiewicz et al., 2008; Stephenson and Maluf, 2010). Repetitive doublets have also been demonstrated in some of the reports above, but as a rule by the way; and only a few studies have directly addressed this topic (Bawa and Calancie, 1983; Kudina and Alexeeva, 1992; Kudina and Andreeva, 2010). However, a number of questions are still unanswered, in particular the question of whether there are certain muscles and (or) certain motoneurons for which repetitive doublet firing is likely to appear. This question will be taken up below based on an analysis of doublet firing in different limb and trunk muscles in healthy humans. With regard to the origin of this surprising firing, currently, the short delayed depolarization (DD) appearing as a “hump” on the falling phase of an action potential, preceding the AHP (Granit et al., 1963; Kernell, 1964; Nelson and Burke, 1967; Kernell, 2006) and thus shortly increasing MN excitability is widely accepted as the mechanism responsible for single doublets in cat (Calvin and Schwindt, 1972; Kirkwood and Munson, 1996; see also Kernell, 2006), and human motoneurons (Kudina, 1974; Bawa and Calancie, 1983; Kudina and Alexeeva, 1992; Bawa and Lemon, 1993; Garland and Griffin, 1999; Piotrkiewicz et al., 2008; Stephenson and Maluf, 2010). As to the origin of repetitive doublets, the question still remains open. At first sight, one could assume that they have the same underlying mechanism like single doublets. However, as known from findings by Granit et al. (1963) and Calvin (1975), in cat motoneurons, the hump-DD changes systematically during rhythmic firing evoked by intracellular depolarization, becoming smaller with each successive spike. Therefore, as may be expected, in firing motoneurons the DD can reach the threshold only occasionally and generate single doublets only. Consequently, it might be suggested that the DD alone can hardly play a pivotal role in producing repetitive doublets. In our recent studies (Kudina and Andreeva, 2010, 2013), we hypothesized that the mechanism underlying repetitive doublet firing in human motoneurons during voluntary muscle contraction is complex, including the hump-DD as the primary determinant that possibly becomes persistent due to a plateau potential probably activated in parallel with a common synaptic input. The assumption about a plateau potential role in repetitive doublet mechanisms was inferred from the comparison between essential characteristics of human motor unit (MU) repetitive doublet firing and properties of cat motoneuron firing in the presence of a plateau potential (e.g., Conway et al., 1988). If this assumption is true, then the question arises what are the DD characteristics which become persistent during repetitive doublet firing. The question will be considered below.

INFLUENCE OF DIFFERENT MODELING PROTOCOLS FOR CALCIUM CURRENTS IN THE AHP FEATURES OF A TYPE-S MOTONEURON

INFLUENCE OF DIFFERENT MODELING PROTOCOLS FOR CALCIUM CURRENTS IN THE AHP FEATURES OF A TYPE-S MOTONEURON

Frequency-dependent amplification of stretch-evoked excitatory input in spinal motoneurons.

Voltage-dependent calcium and sodium channels mediating persistent inward currents (PICs) amplify the effects of synaptic inputs on the membrane potential and firing rate of motoneurons. CaPIC channels are thought to be relatively slow, whereas the NaPIC channels have fast kinetics. These different characteristics influence how synaptic inputs with different frequency content are amplified; the slow kinetics of Ca channels suggest that they can only contribute to amplification of low frequency inputs (<5 Hz). To characterize frequency-dependent amplification of excitatory postsynaptic potentials (EPSPs), we measured the averaged stretch-evoked EPSPs in cat medial gastrocnemius motoneurons in decerebrate cats at different subthreshold levels of membrane potential. EPSPs were produced by muscle spindle afferents activated by stretching the homonymous and synergist muscles at frequencies of 5-50 Hz. We adjusted the stretch amplitudes at different frequencies to produce approximately the same peak-to-peak EPSP amplitude and quantified the amount of amplification by expressing the EPSP integral at different levels of depolarization as a percentage of that measured with the membrane hyperpolarized. Amplification was observed at all stretch frequencies but generally decreased with increasing stretch frequency. However, in many cells the amount of amplification was greater at 10 Hz than at 5 Hz. Fast amplification was generally reduced or absent when the lidocaine derivative QX-314 was included in the electrode solution, supporting a strong contribution from Na channels. These results suggest that NaPICs can combine with CaPICs to enhance motoneuron responses to modulations of synaptic drive over a physiologically significant range of frequencies.

Revisiting the role of spike afterhyperpolarization and spike threshold in motoneuron current-frequency gain

The relationship between synaptic or injecting input level and firing rate is an important metric to characterize neuron input-output dynamics. In this study, we examine two long-held, but never validated, assumptions in the "algebraic summation of afterhyperpolarization" theory, which explains how firing rate varies with input (typically referred to as input current-frequency modulation or "F-I gain"). In the theory, the afterhyperpolarizations themselves, along with spike threshold, were assumed constant. That is, whereas they were central concepts in the theory, they were not included in any feedback loops, wherein they could both affect and be affected by firing rate. We performed intracellular recordings from spinal motoneurons in the adult cat to determine whether F-I gain correlates with the afterhyperpolarization and/or spike threshold. We observe that the afterhyperpolarization does indeed appear to be out of the F-I gain mechanism loop, and thus that original assumption holds. However, the presented experimental evidence indicates that the spike threshold appears to be in the loop. That is, spike threshold variation associated with input correlates with F-I gain. We present an extension to the original theory, which explains the F-I gain experimental correlations.

Motoneuron afterhyperpolarisation duration in amyotrophic lateral sclerosis

Motor unit (MU) potentials were registered from 20 ALS patients and 13 age-matched control individuals during isometric constant force contractions of brachial biceps (BB). The registered signals were decomposed into single MU potential trains. The estimates of duration of the afterhyperpolarisation (AHP) in MNs, derived from the interspike interval variability, was compared between ALS patients (124 MNs) and control subjects (111 MNs) and no significant differences were encountered. However, the relationship between TI and age for patients appeared to be qualitatively different from that of the control group. The dependence of patients' AHPs on relative force deficit (RFD), which quantified muscle involvement, was more specific. For RFDs below 30%, the AHP estimate was significantly lower than control values and then increased thereafter with increasing RFDs. Moreover, firing rates of patients with the smallest RFDs were significantly higher while firing rates of patients with the greatest RFDs were significantly lower than control values. The AHP shortening in the early stages of muscle impairment is consistent with the decrease in firing threshold of ‘fast' MNs found in spinal cord slices from neonatal SOD1 mice. The later elongation of the AHP may be caused by the higher vulnerability of ‘fast' MNs to degeneration and by the influence of reinnervation. Our results are comparable to what has been observed in acute experiments in animal models, providing a bridge between animal and clinical research that may be relevant for identification of mechanism(s) underlying neurodegeneration in ALS.

Time Structure of Impulsation of Spinal Motoneurons of Cats under Conditions of Realization of the Stretch Reflex

We analyzed the characteristics of impulse activity recorded intracellularly from motoneurons of the mm. gastrocnemius + soleus in decerebrated cats; this activity was induced by controlled stretching of the above muscle group. Long-lasting (30 min long or more) sufficiently qualitative records (membrane potential, MP, greater than −50 mA throughout the entire recording period) were obtained from 17 motoneurons. Muscle stretching (sinusoidal half-waves of different amplitudes and combinations of such waves) allowed us to obtain well-reproducible successions of impulses with no significant changes in the resting MP of the studied motoneurons. In 14 of such cells, we found significant correlation between normalized variations of the instantaneous frequency (in a sequence of preceding and subsequent action potentials, APs), indicative of a considerable probability of alternation of high and low values of this parameter. The same trend (toward alternation of high and low values) was observed in successive magnitudes of afterhyperpolarization (AH) that followed APs. Between the values of the extrema of AH trajectories and values of the instantaneous frequency of impulsation, we observed significant negative correlation. The mechanisms underlying the formation of AH in spinal motoneurons and also the effects of this hyperpolarization on the characteristics of impulse activity generated by these cells are discussed.

Reversal of the late phase of spike frequency adaptation in cat spinal motoneurons during fictive locomotion.

In spinal motoneurons, late spike frequency adaptation (SFA) is defined as the slowing of the firing rate over tens of seconds and can be seen during sustained or intermittent current injection. Although the function of late SFA is not known, it may result in a decrease in force production over time, or muscle fatigue. Because locomotion can persist for long periods of time without fatigue, late SFA was studied using intracellular recordings from adult cat motoneurons during fictive locomotion. Of eight lumbar motoneurons studied, all showed late adaptation during control conditions, but none demonstrated late adaptation during locomotor activity. The most consistent properties that correlated with the presence or absence of late SFA were those related to availability of fast, inactivating sodium channels, particularly action potential rate of rise. Evidence of the reversal of late SFA during locomotion was present for several minutes following locomotor trials, consistent with the suggestion that SFA is modulated through slow metabotropic pathways. The abolition of late adaptation in spinal motoneurons during fictive locomotion is an example of a state-dependent change in the "intrinsic" properties of mammalian motoneurons. This change contributes to increased excitability of motoneurons during locomotion and results in robust firing during sustained locomotion.

Characteristics and organization of discharge properties in rat hindlimb motoneurons.

The discharge properties of hindlimb motoneurons in ketamine-xylazine anesthetized rats were measured to assess contributions of persistent intrinsic currents to these characteristics and to determine their distribution in motoneuron pools. Most motoneurons (30/37) responded to ramp current injections with adapting patterns of discharge and the frequency-current (f-I) relations of nearly all motoneurons included a steep subprimary range of discharge. Despite the prevalence of adapting f-I relations, responses included indications that persistent inward currents (PICs) were activated, including increased membrane noise and prepotentials before discharge, as well as counterclockwise hysteresis and secondary ranges in f-I relations. Examination of spike thresholds and afterhyperpolarization (AHP) trajectories during repetitive discharge revealed systematic changes in threshold and trajectory within the subprimary, primary, and secondary f-I ranges. These changes in the primary and secondary ranges were qualitatively similar to those described previously for cat motoneurons. Within the subprimary range, AHP trajectories often included shallow approaches to threshold following recruitment and slope of the AHP ramp consistently increased until the subprimary range was reached. We suggest that PICs activated near recruitment contributed to these slope changes and formation of the subprimary range. Discharge characteristics were strongly correlated with motoneuron size, using input conductance as an indicator of size. Discharge adaptation, recruitment current, and frequency increased with input conductance, whereas both subprimary and primary f-I gains decreased. These results are discussed with respect to potential mechanisms and their functional implications.

Nerve Growth Factor Regulates the Firing Patterns and Synaptic Composition of Motoneurons

Target-derived neurotrophins exert powerful synaptotrophic actions in the adult brain and are involved in the regulation of different forms of synaptic plasticity. Target disconnection produces a profound synaptic stripping due to the lack of trophic support. Consequently, target reinnervation leads to synaptic remodeling and restoration of cellular functions. Extraocular motoneurons are unique in that they normally express the TrkA neurotrophin receptor in the adult, a feature not seen in other cranial or spinal motoneurons, except after lesions such as axotomy or in neurodegenerative diseases like amyotrophic lateral sclerosis. We investigated the effects of nerve growth factor (NGF) by retrogradely delivering this neurotrophin to abducens motoneurons of adult cats. Axotomy reduced the density of somatic boutons and the overall tonic and phasic firing modulation. Treatment with NGF restored synaptic inputs and firing modulation in axotomized motoneurons. When K252a, a selective inhibitor of tyrosine kinase activity, was applied to specifically test TrkA effects, the NGF-mediated restoration of synapses and firing-related parameters was abolished. Discharge variability and recruitment threshold were, however, increased by NGF compared with control or axotomized motoneurons. Interestingly, these parameters returned to normal following application of REX, an antibody raised against neurotrophin receptor p75 (p75(NTR)). In conclusion, NGF, acting retrogradely through TrkA receptors, supports afferent boutons and regulates the burst and tonic signals correlated with eye movements. On the other hand, p75(NTR) activation regulates recruitment threshold, which impacts on firing regularity. To our knowledge, this is the first report showing powerful synaptotrophic effects of NGF on motoneurons in vivo.

A diverse pattern of the spike threshold changes in feline gastrocnemius–soleus motoneurons during stretch reflex activation

The aim of the study was to define the spike threshold changes in motoneurons during naturally evoked activation. Spike activity was evoked in the gastrocnemius-soleus motoneurons of decerebrate cats by controlled stretches of the homonymous muscles. The spike thresholds were defined by using the first derivative of the membrane potential and statistical boundaries of its change. The evoked firing was analyzed in terms of the spike thresholds and the membrane potential trajectories between spikes, which were dependent on the intensity of the firing. The main inference of the study is the absence of a single-valued dependence between the spike rates and thresholds; the last ones can both increase and decrease with a rise in the firing rate; moreover, opposite directions in the threshold changes are often observed at various phases of the stretch-evoked firing. The dependency of the spike thresholds on firing rates was checked in 42 of the total set of 57 motoneurons. In a group of cells (n = 27) showing statistically significant correlation (P < 0.05), this parameter was negative in 18 and positive in 9 motoneurons. The obtained results indicate that the threshold-crossing models with a single-valued dependency of the spike thresholds on the firing rates are not completely suitable to analyze the spike generation processes in the motor unit records.

The Polysynaptic Pathways from the Low Threshold Cutaneous Afferents of the Trunk to Motoneurons Innervating Hindlimb Muscles in the Spinal Cat

We studied the reflex actions of the cutaneous afferents innervating the trunk to hindlimb motoneurons in the spinal cat using an intracellular recording technique. Stimulation of the trunk cutaneous afferents entering into the L2-L5 spinal segment produced different types of polysynaptic potentials in hindlimb motoneurons via polysynaptic neuronal pathways. The trunk cutaneous afferents predominantly caused excitatory PSPs in the flexor motoneurons and inhibitory PSPs in the extensor motoneurons. The size and latency of polysynaptic potentials were related to the proximity of the spinal segments of the nerves stimulated to the spinal segments of motoneurons. These findings suggest that the neuronal pathways from trunk cutaneous afferents to hindlimb motoneurons play an important role in coordinating between the trunk and hindlimbs.

Accommodation in motoneuron models with passive dendrite: Response to linearly rising currents

The biophysical properties of cat motoneuron (MN) models with passive dendrite and different sizes have been extensively tested. However, the behavior of these models in relation to accommodation has not been tested yet. The experimental data reported have shown that MNs accommodate to a very limited degree, exhibiting, in some extension, breakdown of accommodation phenomenon. One possible explanation for the latter behavior would be the presence of persistent sodium conductance. The accommodation can be evaluated by observing threshold latency (TL) curve and the accommodation coefficient ( I/ I 0) s , obtained from the response to linearly rising currents. The simulation results have showed that the models exhibit breakdown of accommodation phenomenon and much similar accommodation coefficients, despite their type: 1.463, 1.456 and 1.418 to S, FR and FF MN models, respectively. According to these data, all the MN models can be classified as slowly accommodating. Our data suggests that accommodation coefficients cannot be associated to the size or to the type of the MN, and persistent sodium conductance might not be necessary to explain breakdown of accommodation phenomenon in MN soma at all.

Cholinergic terminals in the ventral horn of adult rat and cat: Evidence that glutamate is a cotransmitter at putative interneuron synapses but not at central synapses of motoneurons

Until recently it was generally accepted that the only neurotransmitter to be released at central synapses of somatic motoneurons was acetylcholine. However, studies on young mice (P0-10) have provided pharmacological evidence indicating that glutamate may act as a cotransmitter with acetylcholine at synapses between motoneurons and Renshaw cells. We performed a series of anatomical experiments on axon collaterals obtained from intracellularly labeled motoneurons from an adult cat and labeled by retrograde transport in adult rats to determine if glutamate is co-localized with acetylcholine by these terminals. We could find no evidence for the presence of vesicular glutamate transporters in motoneuron axon terminals of either species. In addition, we were unable to establish any obvious relationship between motoneuron terminals and the R2 subunit of the AMPA receptor (GluR2). However we did observe a population of cholinergic terminals in lamina VII which did not originate from motoneurons but were immunoreactive for the vesicular glutamate transporter 2 and formed appositions to GluR2 subunits. These were smaller than motoneuron terminals and, unlike them, formed no relationship with Renshaw cells. The evidence suggests that glutamate does not act as a cotransmitter with acetylcholine at central synapses of motoneurons in the adult cat and rat. However, glutamate is present in a population of cholinergic terminals which probably originate from interneurons where its action is via an AMPA receptor.