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1. We have used the glycogen-depletion technique, combined with myofibrillar ATPase (mATPase) staining for muscle fibre type, to study the fibre-type composition of four skeleto-fusimotor (beta) units in cat peroneus tertius, namely, one beta dynamic (beta d) unit and three beta static (beta s) units. 2. Depletion of glycogen was observed in serial cross-sections of thirty-four beta-unit extrafusal muscle fibres of various types traced from origin to insertion. No fibre was depleted of glycogen throughout its length; depletion was restricted to a number of zones, usually about five. Oxidative (type I) and oxidative-glycolytic (type IIA) fibres were depleted for a significantly greater proportion of their total length than glycolytic IIB fibres. 3. The fibre-type composition of the beta d unit was determined by tracing its fibres from end to end. The muscle unit consisted of one intrafusal bag1 fibre and ninety-three extrafusal muscle fibres comprising seventy-six type I fibres, eleven IIC fibres, and six fibres that changed from IIC to I during the course of their length (IIC/I fibres). The extrafusal fibre-type composition was thus 81.7% I plus 18.3% IIC and IIC/I. 4. The three beta s units (beta s1, beta s2, beta s3) were all fast-contracting and fatigued rapidly. Identification of their extrafusal fibre types, made in 1 mm2 areas sampled from different parts of each unit, gave mixed compositions as follows: beta s1, IIB + 6.7% IIA; beta s2, IIB + 5.8% IIA; beta s3, IIB + 29.9% IIA. The intrafusal component of each unit included either one or two long chain fibres. 5. In a discussion of the results, the fact that the continuous stimulation of extrafusal muscle fibres does not deplete them of glycogen throughout their length is examined in relation to the work of others who have assumed that it did. With regard to the finding of mixed extrafusal fibre types in the beta units, a distinction is drawn between minimal (around 5%) and moderate mixing. It is suggested that minimal mixing may occur in any motor unit as the outcome of endplate degeneration with foreign replacement, but that moderate mixing indicates an on-going process of conversion from one fibre type to another which in the adult may prove to occur only among beta units.

1. Intrafusal muscle fibres of cat tenuissimus spindles have been injected with the fluorescent dye Procion Yellow and identified histologically after recording their changes in membrane potential during 1/sec stimulation of single static or dynamic gamma axons. 2. Thirteen intrafusal muscle fibres innervated by static gamma axons were identified as eight bag2 and five chain fibres. The fact that none proved to be a bag1 fibre is not regarded as significant, for reasons given in the Discussion. 3. In one spindle Procion Yellow was injected into two intrafusal muscle fibres activated by the same static gamma axon; they were identified as a bag2 and a chain fibre. 4. Nine intrafusal muscle fibres innervated by dynamic gamma axons were identified as seven bag1 fibres, one bag2 fibre, and one long chain fibre. 5. In one spindle two bag fibres were injected, one activated by a dynamic gamma axon, the other by a static gamma axon; the former proved to be a bag1 fibre, the latter a bag2 fibre. 6. Stimulation of static gamma axons elicited junctional potentials in seven bag2 fibres and one damaged chain fibre, and action potentials in one bag2 and four chain fibres. In the whole sample of impaled intrafusal muscle fibres (identified and unidentified) activated by static axons, junctional potentials were recorded from twenty-three (62.2%), and action potentials from fourteen (37.8%). Stimulation of dynamic gamma axons always elicited junctional potentials. 7. In a number of instances it was possible to examine the ultrastructure of motor endings belonging to the stimulated gamma axon. The myoneural junctions of trail endings supplied by static gamma axons to bag2 and chain fibres were both smooth and folded; the deepest and most regular folding occurred on chain fibres. The terminals of p2 plates supplied to bag1 fibres by dynamic gamma axons had smooth myoneural junctions.

1. Eleven cat tenuissimus spindles have been analysed mainly by cutting serial, transverse, 1 micrometer thick sections following direct observation of the effects of dynamic motor (gamma or beta) stimulation. 2. Histological results from these spindles were also used to interpret the effects of static fusimotor stimulation of other spindles. 3. Dynamic motor stimulation usually produced contractions seen as convergent movements of sarcomeres in single bag fibres, identified as bag1 fibres for reasons given in the text. 4. In one spindle a single dynamic axon produced a translational movement in one pole of a bag1 fibre and a convergent movement in each pole of a bag2 fibre, together with movements in other unidentified (presumably chain) fibres. Subsequent analysis showed that besides innervating both bag fibres the axon also supplied two chain fibres. 5. Contrary to expectation, motor endings on the bag1 fibres seldom occurred at the sites of convergent movement. Only two cases of coincidence occurred among sixteen foci and twenty-one motor endings; otherwise focus and nearest ending were separated by distances of 0.85--2.5 mm. 6. Most of the convergent movements of sarcomeres observed in bag1 fibres occurred in a region of the pole that is ultrastructurally distinct from the region where most of the motor endings were located. The possible relevance of this to the production of contractions in the bag1 fibre is discussed. 7. Convergent movement foci in bag2 fibres produced by the stimulation of static axons occurred largely within the same regions of the pole as the motor endings were located, though, whereas foci were observed in both intra- and extracapsular regions, most of the endings were intracapsular.

The motor innervation of cat spindles was examined in hindlimb muscles using a variety of techniques employed in light and electron microscopy. Observations were made on teased, silver preparations of 267 spindles sampled from the peroneal, flexor hallucis longus, and soleus muscles, hereafter referred to as the PER/FHL/SOL series. The γ innervation. Trail endings are almost invariably present, and innervate both bag and chain muscle fibres. Trail fibres accounted for 64.6 to 74.8% of the total fusimotor supply to samples of spindle poles in the PER/FHL/SOL series, the mean number of fibres per pole varying from 2.7 to 5.0 in the different muscles, and the mean number of ramifications (areas of synaptic contact) per fibre being 3.7. By contrast, the p₂innervation of a spindle pole generally consists of a single fibre supplying only one plate. In the above samples p(2) fibres accounted for 4.1 to 28.0% of the total fusimotor supply, and the mean number of fibres per pole varied from 0.3 to 1.2 in the different muscles. Ninety per cent of p(2) plates innervate bag fibres. The α innervation. The structure of p₁plates as seen in both light and electron microscopy compares very closely with that of extrafusal plates. After nerve section p₁plates degenerate at the same time as extrafusal plates, being the first of the three types of fusimotor ending to disappear. The frequency of the p₁innervation is similar to that of the p₂innervation. In the same samples of PER/FHL/SOL spindle poles as above p₁ fibres accounted for 6.0 to 28.8% of the total fusimotor supply, the mean number of fibres per pole varying from 0.25 to 2.1 in the different muscles. The majority of p₁ fibres enter a pole to terminate in one plate only. Seventy-five per cent of the plates innervate bag fibres. The three types of fusimotor ending are thus not selectively distributed to the two types of intrafusal muscle fibre. All three types of fusimotor fibre may branch within the spindle so as to innervate both bag and chain fibres. Bag fibres receive both types of plate ending as well as trail endings. Most chain fibres receive trail endings only; the rest receive either a p₁or a p₂plate innervation in addition, 25% of the p₁and 10% of the p₂innervation being distributed to chain fibres. The significance of this nonselective innervation is interpreted as indicating that the type of contraction elicited by stimulating a fusimotor fibre depends upon the type of ending initiating it rather than upon the type of muscle fibre executing it. Reasons are given for concluding that the dynamic response is controlled via the p₁and p₂plates, and that the static response is controlled by the trail endings. The participation of the α fibres in mammalian fusimotor innervation, previously regarded as a vestigial feature, proved to be widespread in the muscles studied and more prevalent in fast muscles (FHL, peroneus digiti quinti) than slow (soleus). A low frequency of p₁innervation is offset by a high frequency of p₂(as in peroneus longus), and vice versa (as in FHL). It is unlikely that collaterals from slow α fibres innervating type B muscle fibres are wholly responsible for the high frequency of the p₁innervation in FHL, and it is suggested that collaterals may also be derived from fast α fibres innervating type C muscle fibres. The possibility of there being some motor fibres of α conduction velocity and with an exclusively fusimotor distribution is also taken into account.