Extraocular motor units: type classification and motoneuron stimulation frequency-muscle unit force relationships.

Extracellular and intracellular techniques were used to study single motor units of the abducens nucleus and lateral rectus muscle in the cat. Using a combination of two motor unit properties, the fusion frequency and an index of fatigability, the population of twitch motor units could be separated into 4 subgroups: fast fatigable (FF), fast fatigue resistant (FR), slow fatigable (SF) and slow fatigue resistant (S). Nontwitch motor units, a fifth subgroup (NT), formed 10% of the total studied population. The twitch tension and the maximum tetanic tension of the FF motor unit type were significantly stronger than all other motor unit types. The use of frequency varying stimulation patterns did not further differentiate the motor unit types. The relation between a series of single motoneuron stimulation frequencies and the resultant single muscle unit forces generated a slope defined as a motor unit's kt value. Motor units with low kt values had higher twitch tensions, higher maximum tetanic tensions, higher fusion frequencies and lower fatigue indices than motor units with high kt values. Motoneuron recruitment was tested by electrical stimulation of the medial rectus subdivision of the contralateral oculomotor nucleus. No correlations were seen between recruitment order and the mechanical parameters of the single abducens motor units.     

Recruitment order, contractile characteristics, and firing patterns of motor units in the temporalis muscle of monkeys.

Action potentials of single motor units were recorded in the temporalis muscle of rhesus monkeys trained to hold static forces on a bite bar. The average relationship between mandibular force and motor unit firing rate was determined for many motor units using a computer program, and from this relationship the threshold force at which a motor unit began to fire was determined. Several aspects of motor unit firing rate were examined in relation to recruitment threshold. Computer averaging was used to determine twitch tension and twitch contraction times for many motor units. Motor units recruited at low forces had longer contraction times and produced smaller twitch tensions than higher-threshold motor units. Motor units with small action potentials were nearly always recruited at lower force levels than those with larger action potentials. These results indicate that the motor units of the monkey temporalis muscle were recruited in an orderly fashion, which is in accord with the predictions of the “size principle”.     

MIF versus SIF Motoneurons,What Are Their Respective Contribution in the Oculomotor Medial Rectus Pool?

Multiply-innervated muscle fibers (MIFs) are peculiar to the extraocular muscles as they are non-twitch but produce a slow build up in tension on repetitive stimulation. The motoneurons innervating MIFs establish en grappe terminals along the entire length of the fiber, instead of the typical en plaque terminals that singly-innervated muscle fibers (SIFs) motoneurons establish around the muscle belly. MIF motoneurons have been proposed to participate only in gaze holding and slow eye movements. We aimed to discern the function of MIF motoneurons by recording medial rectus motoneurons of the oculomotor nucleus. Single-unit recordings in awake cats demonstrated that electrophysiologically-identified medial rectus MIF motoneurons participated in different types of eye movements, including fixations, rapid eye movements or saccades, convergences, and the slow and fast phases of the vestibulo-ocular nystagmus, the same as SIF motoneurons did. However, MIF medial rectus motoneurons presented lower firing frequencies, were recruited earlier and showed lower eye position (EP) and eye velocity (EV) sensitivities than SIF motoneurons. MIF medial rectus motoneurons were also smaller, had longer antidromic latencies and a lower synaptic coverage than SIF motoneurons. Peristimulus time histograms (PSTHs) revealed that electrical stimulation to the myotendinous junction, where palisade endings are located, did not recurrently affect the firing probability of medial rectus motoneurons. Therefore, we conclude there is no division of labor between MIF and SIF motoneurons based on the type of eye movement they subserve.SIGNIFICANCE STATEMENT In addition to the common singly-innervated muscle fiber (SIF), extraocular muscles also contain multiply-innervated muscle fibers (MIFs), which are non-twitch and slow in contraction. MIF motoneurons have been proposed to participate only in gaze holding and slow eye movements. In the present work, by single-unit extracellular recordings in awake cats, we demonstrate, however, that both SIF and MIF motoneurons, electrophysiologically-identified, participate in the different types of eye movements. However, MIF motoneurons showed lower firing rates (FRs), recruitment thresholds, and eye-related sensitivities, and could thus contribute to the fine adjustment of eye movements. Electrical stimulation of the myotendinous junction activates antidromically MIF motoneurons but neither MIF nor SIF motoneurons receive a synaptic reafferentation that modifies their discharge probability.     

Motoneurons of twitch and nontwitch extraocular muscle fibers in the abducens, trochlear, and oculomotor nuclei of monkeys

Eye muscle fibers can be divided into two categories: nontwitch, multiply innervated muscle fibers (MIFs), and twitch, singly innervated muscle fibers (SIFs). We investigated the location of motoneurons supplying SIFs and MIFs in the six extraocular muscles of monkeys. Injections of retrograde tracers into eye muscles were placed either centrally, within the central SIF endplate zone; in an intermediate zone, outside the SIF endplate zone, targeting MIF endplates along the length of muscle fiber; or distally, into the myotendinous junction containing palisade endings. Central injections labeled large motoneurons within the abducens, trochlear or oculomotor nucleus, and smaller motoneurons lying mainly around the periphery of the motor nuclei. Intermediate injections labeled some large motoneurons within the motor nuclei but also labeled many peripheral motoneurons. Distal injections labeled small and medium-large peripheral neurons strongly and almost exclusively. The peripheral neurons labeled from the lateral rectus muscle surround the medial half of the abducens nucleus: from superior oblique, they form a cap over the dorsal trochlear nucleus; from inferior oblique and superior rectus, they are scattered bilaterally around the midline, between the oculomotor nucleus; from both medial and inferior rectus, they lie mainly in the C-group, on the dorsomedial border of oculomotor nucleus. In the medial rectus distal injections, a "C-group extension" extended up to the Edinger-Westphal nucleus and labeled dendrites within the supraoculomotor area. We conclude that large motoneurons within the motor nuclei innervate twitch fibers, whereas smaller motoneurons around the periphery innervate nontwitch, MIF fibers. The peripheral subgroups also contain medium-large neurons which may be associated with the palisade endings of global MIFs. The role of MIFs in eye movements is unclear, but the concept of a final common pathway must now be reconsidered.     

Twitch and nontwitch motoneuron subgroups in the oculomotor nucleus of monkeys receive different afferent projections

Motoneurons in the primate oculomotor nucleus can be divided into two categories, those supplying twitch muscle fibers and those supplying nontwitch muscle fibers. Recent studies have shown that twitch motoneurons lie within the classical oculomotor nucleus (nIII), and nontwitch motoneurons lie around the borders. Nontwitch motoneurons of medial and inferior rectus are in the C group dorsomedial to nIII, whereas those of inferior oblique and superior rectus lie near the midline are in the S group. In this anatomical study, afferents to the twitch and nontwitch subgroups of nIII have been anterogradely labeled by injections of tritiated leucine into three areas and compared. 1) Abducens nucleus injections gave rise to silver grain deposits over all medial rectus subgroups, both twitch and nontwitch. 2) Laterally placed vestibular complex injections that included the central superior vestibular nucleus labeled projections only in twitch motoneuron subgroups. However, injections into the parvocellular medial vestibular nucleus (mvp), or Y group, resulted in labeled terminals over both twitch and nontwitch motoneurons. 3) Pretectal injections that included the nucleus of the optic tract (NOT), and the olivary pretectal nucleus (OLN), labeled terminals only over nontwitch motoneurons, in the contralateral C group and in the S group. Our study demonstrates that twitch and nontwitch motoneuron subgroups do not receive identical afferent inputs. They can be controlled either in parallel, or independently, suggesting that they have basically different functions. We propose that twitch motoneurons primarily drive eye movements and nontwitch motoneurons the tonic muscle activity, as in gaze holding and vergence, possibly involving a proprioceptive feedback system.     

Elasmobranch oculomotor organization: anatomical and theoretical aspects of the phylogenetic development of vestibulo-oculomotor connectivity

The oculomotor organization of two elasmobranch species, smooth dogfish (Mustelus canis) and little skate (Raja erinacea), was studied by investigating the extraocular muscle apparatus and the oculomotor motoneuron distribution. The macroscopic appearance of the eye muscles was similar to any lateral-eyed vertebrate species (e.g., goldfish, rabbit). The size of extraocular muscles was expressed by counting single muscle fibers and comparing cross-sectional areas of the extraocular muscles. There were significant differences in the number of fibers in the six extraocular muscles in dogfish, but not in skate. Fiber sizes varied considerably; thus, the number of fibers did not relate to cross-sectional areas. In the dogfish, no one pair of agonist-antagonist extraocular muscles was larger than the others, suggesting that there was no preference for eye movements in a particular plane of space. However, the lateral rectus was more than twice the size of most of the other muscles. In the skate, cross-sectional areas of the horizontal eye muscles were smaller than those of the vertical eye movers. This may indicate a reduced utilization of horizontal eye muscles, which may reflect the bottom-dwelling habitat and mode of locomotion of the skate. The distribution of the extraocular motoneurons was determined by injecting horseradish peroxidase (HRP) into single eye muscles. Medial rectus, superior rectus, and superior oblique motoneuron populations were located contralateral to their respective muscles. Lateral rectus, inferior rectus, and inferior oblique motoneurons were located ipsilateral to their muscles. This distribution is in contrast to almost all other vertebrates studied thus far, where medial rectus motoneurons are located ipsilateral to the muscle which they innervate. The oculomotor arrangement in elasmobranchs is likely to have consequences for the circuitry responsible for the production of conjugate compensatory eye movements in the horizontal plane. We hypothesize that, in contrast to other vertebrates, the basic elasmobranch vestibulo-ocular reflex pathway consists of three identically structured three-neuron-arcs connecting the three semicircular canals to their respective extraocular muscles. This innervation pattern may constitute a special feature of the elasmobranch brain or a phylogenetically older arrangement of eye movement pathways.     

Summation of extraocular motor unit tensions in the lateral rectus muscle of the cat

Contractile measures on 67 single muscle units in the cat lateral rectus muscle were made in response to motoneuron stimulation. Simultaneous activation of four to five additional units, using muscle nerve stimulation, allowed an examination of unit force summation. Linear force addition was found in 73% of the units, while 25% added only about half of their twitch force to the twitch force of the nerve-activated units. “Nonadditive” units had significantly weaker twitch tensions than the units which added linearly. Lengthening or shortening the whole muscle, from maximal isometric settings, reduced whole muscle twitch tension as well as muscle unit tension. Injury to the lateral rectus muscle did not significantly alter whole muscle tension. These findings suggest that the known serial and branching arrangement of these muscle fibers, as well as the complex interfiber matrix, may help explain the force reduction in some muscle units and the whole muscle's resistance to insult. © 1997 John Wiley & Sons, Inc. Muscle Nerve 20: 1229–1235, 1997     

Examination of feline extraocular motoneuron pools as a function of muscle fiber innervation type and muscle layer

This study explores two points related to the pattern of innervation of the extraocular muscles. First, species differences exist in the location of the motoneurons supplying multiply innervated fibers (MIFs) and singly innervated fibers (SIFs) in eye muscles. MIF motoneurons are located outside the extraocular nuclei in primates, but are intermixed with SIF motoneurons within rat extraocular nuclei. To test whether this difference is related to visual capacity and frontal placement of eyes, we injected retrograde tracers into the medial rectus muscle of the cat, a highly visual nonprimate with frontally placed eyes. Distal injections labeled smaller MIF motoneurons located ventrolaterally and rostrally within the oculomotor nucleus (III). More central injections also labeled a separate population of larger cells located dorsally in III. Thus, the cat shares with the nocturnal rat the feature of having MIF motoneurons located within the bounds of III. On the other hand, just as with monkeys, cats show segregation of the MIF and SIF medial rectus motoneuron pools, albeit in a different pattern. Second, extraocular muscles are divided into two layers; the inner, global layer inserts into the sclera, and the outer, orbital layer inserts into the connective tissue pulley. To test whether these layers are supplied by anatomically discrete motoneuron pools, we injected tracer into the orbital layer of the cat lateral rectus muscle. No evidence of either morphological or distributional differences was found, suggesting that the functional differences in these layers may be due mainly to their orbital anatomy, not their innervation. J. Comp. Neurol. 525:919–935, 2017. © 2016 Wiley Periodicals, Inc.     

Discharge variability and physiological properties of human masseter motor units

Intramuscular electrodes were used to study discharge variability in motor units of human masseter whose physiological properties were determined using spike-triggered averaging. Subjects voluntarily controlled the mean firing rate of a selected motor unit at 10 Hz for 15 min of continuous activation. Discharge variability was assessed at the beginning and end of this period. In 81% of units, the discharge variability at a mean interspike interval (ISI) of 100 ms increased after 15 min of continuous activity. There was a wide range of discharge variability within the population of masseter units studied, but no significant correlations were found between initial discharge variability and recruitment threshold, twitch tension or time-to-peak tension (TTP). There was, however, a significant correlation between motor unit fatigability and its initial discharge variability. This represents a link between the motoneuron and the functional properties of the muscle fibers in innervates.     

The regulation of synaptic strength within motor units of the frog cutaneous pectoris muscle

The physiological properties of frog neuromuscular junctions may vary widely in a single muscle. In order to understand the factors that contribute to this variation, we have studied populations of synapses belonging to individual motor units of the frog cutaneous pectoris muscle. Motor units in this muscle differ widely in twitch strength. A motor axon's synaptic contacts could be found throughout the muscle, at both singly and polyneuronally innervated endplates. Indeed, over 36% of the endplates contacted by each isolated motor axon were polyneuronally innervated. Comparisons of synapses on muscle fibers in large twitch motor units with those in small twitch motor units reveal that endplate potential amplitude, transmitter release, and muscle fiber diameter are positively correlated with the strength of the motor unit contraction. Large and small twitch motor units differ more in their transmitter release than in their nerve terminal length, indicating that larger twitch motor units have a higher release per unit length of terminal. Among motor units of roughly similar twitch tension, transmitter release at endplates receiving only one axonal input is remarkably constant, independent of postsynaptic muscle fiber input resistance, or, presumably, nerve terminal size. In cases where two different motor axons contribute to a single endplate, the synaptic strength of each input is again related to properties of the contributing motoneuron, although the individual synaptic inputs are markedly reduced in strength and size relative to singly innervated endplates. Additionally, the diameter of polyneuronally innervated muscle fibers appears related to properties of both innervating motoneurons. Thus, the pre- and postsynaptic characteristics of neuromuscular junctions may be determined both by the motoneuron and by peripheral interactions between motoneurons.     

Morphology and ultrastructure of medial rectus subgroup motoneurons in the macaque monkey

There are two muscle fiber types in extraocular muscles: those receiving a single motor endplate, termed singly innervated fibers (SIFs), and those receiving multiple small terminals along their length, termed multiply innervated fibers (MIFs). In monkeys, these two fiber types receive input from different motoneuron pools: SIF motoneurons found within the extraocular motor nuclei, and MIF motoneurons found along their periphery. For the monkey medial rectus muscle, MIF motoneurons are found in the C‐group, while SIF motoneurons lie in the A‐ and B‐groups. We analyzed the somatodendritic morphology and ultrastructure of these three subgroups of macaque medial rectus motoneurons to better understand the structural determinants controlling the two muscle fiber types. The dendrites of A‐ and B‐group motoneurons lay within the oculomotor nucleus, but those of the C‐group motoneurons were located outside the nucleus, and extended into the preganglionic Edinger–Westphal nucleus. A‐ and B‐group motoneurons were very similar ultrastructurally. In contrast, C‐group motoneurons displayed significantly fewer synaptic contacts on their somata and proximal dendrites, and those contacts were smaller in size and lacked dense‐cored vesicles. However, the synaptic structure of C‐group distal dendrites was quite similar to that observed for A‐ and B‐group motoneurons. Our anatomical findings suggest that C‐group MIF motoneurons have different physiological properties than A‐ and B‐group SIF motoneurons, paralleling their different muscle fiber targets. Moreover, primate C‐group motoneurons have evolved a special relationship with the preganglionic Edinger–Westphal nucleus, suggesting these motoneurons play an important role in near triad convergence to support increased near work requirements. J. Comp. Neurol. 522:626–641, 2014. © 2013 Wiley Periodicals, Inc.     

Lateral rectus emg and contractile responses elicited by cat abducens motoneurons

Stimulation of 41 single, abducens nucleus motoneurons in the cat evoked electromyographic (EMG) and contractile responses in the ipsilateral lateral rectus muscle. Separate, bipolar, fine wire EMG recording electrodes in the global and orbital muscle layers showed that 22 muscle units were confined to the global layer, 8 to the orbital layer, and 11 units were contained in both (“bilayer”) muscle layers. “Bilayer” units demonstrated significantly greater twitch (P ≤ 0.002) and maximum tetanic (P ≤ 0.001) tensions as well as faster fusion frequencies (P ≤ 0.022) than either global or orbital units. “Bilayer” units also showed the lowest average kt values (the slope of the linear relationship between motoneuron stimulation frequency and isometric tetanic tension). “Bilayer” units were predominantly fast fatigable (FF). Global units displayed all muscle unit types including all the nonwitch (NT) units. Orbital units were identified as slow fatigable (SF) and fast fatigue resistant (FR). © 1995 John Wiley & Sons, Inc.     

Extra- and intracellular HRP analysis of the organization of extraocular motoneurons and internuclear neurons in the guinea pig and rabbit

The distribution of extraocular motoneurons and abducens and oculomotor internuclear neurons was determined in guinea pigs by injecting horseradish peroxidase (HRP) into individual extraocular muscles, the abducens nucleus, the oculomotor nucleus, and the cerebellum. Motoneurons in the oculomotor nucleus innervated the ipsilateral inferior rectus, inferior oblique, medial rectus, and the contralateral superior rectus and levator palpebrae muscles. Most motoneurons of the trochlear nucleus projected to the contralateral superior oblique muscle although a small number innervated the ipsilateral superior oblique. The abducens and accessory abducens nuclei innervated the ipsilateral rectus and retractor bulbi muscles, respectively. The somata of abducens internuclear neurons formed a cap around the lateral and ventral aspects of the abducens nucleus. The axons of these internuclear neurons terminated in the medial rectus subdivision of the contralateral oculomotor nucleus. At least two classes of guinea pig oculomotor internuclear interneurons exist. One group, located primarily ventral to the oculomotor nucleus, innervated the abducens nucleus and surrounding regions. The second group, lying mainly in the dorsal midline area of the oculomotor nucleus, projected to the cerebellum. Intracellular staining with HRP demonstrated similar soma-dendritic organization for oculomotor and trochlear motoneurons of both guinea pigs and rabbits. Dendrites of oculomotor motoneurons radiated symmetrically from the soma to cover approximately one-third of the entire nucleus, and each motoneuron sent at least one dendrite into the central gray overlying the oculomotor nucleus. In both species, a small percentage of oculomotor motoneurons possessed axon collaterals that terminated both within and outside of the nucleus. The dendrites of trochlear motoneurons extended into the medial longitudinal fasciculus and the reticular formation lateral to the nucleus. Our data on the topography of motoneurons and internuclear neurons in the guinea pig and soma-dendritic organization of motoneurons in the guinea pig and rabbit show that these species share common organizational and morphological features. In addition, comparison of these data with those from other mammals reveals that dendritic complexity (number of dendrites per motoneuron) of extraocular motoneurons exhibits a systematic increase with animal size.     

Extraocular motoneuron pools develop along a dorsoventral axis in zebrafish,Danio rerio

Both spatial and temporal cues determine the fate of immature neurons. A major challenge at the interface of developmental and systems neuroscience is to relate this spatiotemporal trajectory of maturation to circuit‐level functional organization. This study examined the development of two extraocular motor nuclei (nIII and nIV), structures in which a motoneuron's identity, or choice of muscle partner, defines its behavioral role. We used retro‐orbital dye fills, in combination with fluorescent markers for motoneuron location and birthdate, to probe spatial and temporal organization of the oculomotor (nIII) and trochlear (nIV) nuclei in the larval zebrafish. We describe a dorsoventral organization of the four nIII motoneuron pools, in which inferior and medial rectus motoneurons occupy dorsal nIII, while inferior oblique and superior rectus motoneurons occupy distinct divisions of ventral nIII. Dorsal nIII motoneurons are, moreover, born before motoneurons of ventral nIII and nIV. The order of neurogenesis can therefore account for the dorsoventral organization of nIII and may play a primary role in determining motoneuron identity. We propose that the temporal development of extraocular motoneurons plays a key role in assembling a functional oculomotor circuit. J. Comp. Neurol. 525:65–78, 2017. © 2016 The Authors The Journal of Comparative Neurology Published by Wiley Periodicals, Inc.     

Muscle units divided among retractor bulbi muscle slips and between the lateral rectus and retractor bulbi muscles in cat

Single muscle units of the retractor bulbi and lateral rectus muscles were activated with brief (0.5-ms) current pulses delivered through an intracellular micropipet penetrating single motoneuron cell bodies or axons in the principal abducens nucleus of the cat. Muscle unit mechanical characteristics were measured. A total of 40 retractor bulbi muscle units was studied. Thirty muscle units were contained in 2, 3, or all 4 retractor bulbi muscle slips. Three muscle units involved one retractor bulbi muscle slip and seven units were contained in the lateral rectus muscle plus one or both of the lateral retractor bulbi muscle slips. The largest percentage of muscle units (32.5%) was contained in the two inferior retractor bulbi muscle slips. The 33 retractor bulbi muscle units, having no muscle fibers in the lateral rectus muscle, caused the contraction of 76 retractor bulbi muscle slips. Twitch tension was measured in 58 slips and maximum tetanic tension in 27 slips. The average twitch tension per measured slip was 68.0 mg and the average maximum tetanic tension was 459.8 mg. The average retractor bulbi muscle unit contraction time was 10 ms and the lateral rectus average contraction time was 7.9 ms. We propose that peripheral branching of abducens nerve axons can account for the division of muscle unit components among the retractor bulbi muscle slips and between the lateral rectus and retractor bulbi muscles.     

Morphological substrate for eyelid movements: innervation and structure of primate levator palpebrae superioris and orbicularis oculi muscles

The levator palpebrae superioris and orbicularis oculi are antagonistic muscles that function during movements of the eyelid. The levator also functions in conjunction with superior and inferior rectus muscles in coordinated eye/lid movements. The present study examined the innervation and morphology of these muscles in Cynomolgous monkeys (Macaca fascicularis) in order to provide a better understanding of the anatomical substrate for lid movements. Motoneurons innervating the levator and orbicularis muscles were identified and localized by retrograde transport of WGA/HRP and HRP. Retrogradely labelled levator motoneurons were distributed bilaterally throughout the caudal central division of the oculomotor nucleus. A few labelled cells were also present within the contralateral superior rectus division, possibly because of the spread of tracer at the injection site. The possibility that individual motoneurons collateralize to innervate the levator muscle bilaterally was tested by using double retrograde labelling techniques. Doubly labelled levator motoneurons could not be detected by using a combination of tracers (HRP and Fast Blue). Motoneurons innervating the upper lid portion of the orbicularis oculi muscle were distributed within the dorsal subdivision of the ipsilateral facial motor nucleus, with a few neurons in the corresponding locus of the contralateral facial nucleus. Species differences in levator motoneuron distribution, particularly distinctions in lateral-eyed versus frontal-eyed mammals, are discussed in relation to the neural control of lid movements. The levator palpebrae superioris contains three of the same ultrastructurally defined types of singly innervated muscle fiber found in the global layer of other extraocular muscles and an additional, unique slow-twitch fiber type. Moreover, the multiply innervated fiber types so characteristic of the other extraocular muscles are conspicuously absent from levator muscles. Unlike the rectus and oblique extraocular muscles, the levator lacks a layered distribution of fiber types. The morphological profiles of levator muscle fiber types are such that they generally do not respect traditional fiber classification schemes, but are consistent with a role for the levator in sustained elevation of the lid. The orbicularis oculi muscle, by contrast, exhibited three distinct fiber types that resembled categories of skeletal muscle twitch fibers. One slow-twitch and two fast-twitch fiber types were noted. On the basis of oxidative enzyme profiles and mitochondrial content, the majority of orbicularis oculi fibers would be fatigue-prone, an assessment consistent with their rapid onset/offset of acti     

Localization of motoneurons controlling the extraocular muscles of the rat

The localization of extraocular motoneurons in the rat was investigated by injecting horseradish peroxidase and [¹²⁵I]wheat germ agglutinin¹⁷ as retrogade tracer substances into individual eye muscles. The organization of subnuclei was found to be most similar to the rabbit. The subgroups representing the medial rectus and inferior rectus muscles are located in the rostral two thirds of the ipsilateral oculomotor nucleus (nIII) with some medial rectus motoneurons scattered laterally along the edge of the medial longitudinal fasciculus. The motor pool controlling the inferior oblique muscle is located in the middle third of the ipsilateral nIII. The motoneurons of the superior rectus muscles are in the caudal two-thirds of contralateral nIII while the levator palpebrae muscle has a bilateral innervation in the oculomotor nucleus. The motoneurons of the superior oblique are located in the contralateral trochlear nucleus although a few labeled neurons were scattered laterally in amongst the fibers of the medial longitudinal fasciculus. The cell bodies of lateral rectus motoneurons regional separation between the latter and internuclear neurons was found after injecting HRP into the oculomotor nucleus.     

Internal organization of medial rectus and inferior rectus muscle neurons in the C group of the oculomotor nucleus in monkey

Mammalian extraocular muscles contain singly innervated twitch muscle fibers (SIF) and multiply innervated nontwitch muscle fibers (MIF). In monkey, MIF motoneurons lie around the periphery of oculomotor nuclei and have premotor inputs different from those of the motoneurons inside the nuclei. The most prominent MIF motoneuron group is the C group, which innervates the medial rectus (MR) and inferior rectus (IR) muscle. To explore the organization of both cell groups within the C group, we performed small injections of choleratoxin subunit B into the myotendinous junction of MR or IR in monkeys. In three animals the IR and MR myotendinous junction of one eye was injected simultaneously with different tracers (choleratoxin subunit B and wheat germ agglutinin). This revealed that both muscles were supplied by two different, nonoverlapping populations in the C group. The IR neurons lie adjacent to the dorsomedial border of the oculomotor nucleus, whereas MR neurons are located farther medially. A striking feature was the differing pattern of dendrite distribution of both cell groups. Whereas the dendrites of IR neurons spread into the supraoculomotor area bilaterally, those of the MR neurons were restricted to the ipsilateral side and sent a focused bundle dorsally to the preganglionic neurons of the Edinger‐Westphal nucleus, which are involved in the “near response.” In conclusion, MR and IR are innervated by independent neuron populations from the C group. Their dendritic branching pattern within the supraoculomotor area indicates a participation in the near response providing vergence but also reflects their differing functional roles. J. Comp. Neurol. 523:1809–1823, 2015. © 2015 Wiley Periodicals, Inc.     

Calretinin inputs are confined to motoneurons for upward eye movements in monkey

Motoneurons of extraocular muscles are controlled by different premotor pathways, whose selective damage may cause directionally selective eye movement disorders. The fact that clinical disorders can affect only one direction, e.g., isolated up‐/downgaze palsy or up‐/downbeat nystagmus, indicates that up‐ and downgaze pathways are organized separately. Recent work in monkey revealed that a subpopulation of premotor neurons of the vertical eye movement system contains the calcium‐binding protein calretinin (CR). With combined tract‐tracing and immunofluorescence, the motoneurons of vertically pulling eye muscles in monkey were investigated for the presence of CR‐positive afferent terminals. In the oculomotor nucleus, CR was specifically found in punctate profiles contacting superior rectus and inferior oblique motoneurons, as well as levator palpebrae motoneurons, all of which participate in upward eye movements. Double‐immunofluorescence labeling revealed that CR‐positive terminals lacked the γ‐aminobutyric acid (GABA)‐synthesizing enzyme glutamate decarboxylase, which is present in inhibitory afferents to all motoneurons mediating vertical eye movements. Therefore, CR‐containing afferents are considered to be excitatory. In conclusion, a strong CR input is confined to motoneurons mediating upgaze, which derive from premotor pathways mediating saccades and smooth pursuit, but not from secondary vestibulo‐ocular neurons in the magnocellular part of the medial vestibular nucleus. The functional significance of CR in these connections is unclear, but it may serve as a useful marker to locate upgaze pathways in the human brain. J. Comp. Neurol. 521:3154–3166, 2013. © 2013 Wiley Periodicals, Inc.     

Short-term regulation of fatty acid synthesis in cultured glial and neuronal cells

The recruitment thresholds, twitch tension, twitch contraction time and fatiguability of human motor units have been studied in the first dorsal interosseous of the hand. Units recruited at contraction strengths <50 g had relatively low twitch tensions (0.18–1.9 g), long contraction times (59–146 msec) and were non-fatiguable. Units recruited at higher contraction strengths (>200 g) had higher twitch tensions (15–26 g), faster contraction times (33–57 msec) and were highly fatiguable. It is concluded that during graded voluntary muscle contractions motor units are recruited in order of increasing contraction strength and diminishing fatigue resistance.