In muscle spindles of the cat, independent control of dynamic and
static components of the response of the
primary sensory ending to stretch is provided by separate motor inputs
to the various kinds of intrafusal
muscle fibre: dynamic axons (γ or β) to the bag1
fibres and static axons to the bag2 (typically γ only)
and
chain (γ or β) fibres. Nonlinear summation of separately evoked
effects during combined stimulation of
dynamic and static motor axons appears to be due to mutual resetting by
antidromic invasion of separate
encoding sites, leading to partial occlusion of the momentarily lesser
response by the greater. The encoding
sites are thought to be located within the primary ending's preterminal
branches which from first-order level
are normally segregated to the bag1 fibre and to the bag2
and chain fibres. Here we describe the analysis of a
special case that arose in a histophysiological study which had shown that
the degree of occlusion was
related to the minimum number of nodes between the putative encoding sites.
Three-dimensional
reconstruction of the primary ending revealed that the terminals of one
chain fibre were derived entirely
from the first-order branch that supplied the bag1 fibre, including
one terminal that was shared directly with
the bag1 (sensory cross-terminal). The other first-order branch
supplied the bag2 and remaining chain fibres
as normal. The degree of occlusion seen during simultaneous stimulation
of a dynamic β axon and a static γ
axon indicated that the encoding sites were separated by both first-order
branches. Schematic reconstruction
of the motor innervation revealed that the static γ axon was most
unlikely to have supplied the chain fibre
which shared sensory terminals with the bag1, but that these
fibres also shared a motor input with
histological characteristics of β type. Ramp-frequency stimulation
of the dynamic β axon at constant length
evoked a driving effect which persisted after fatiguing the extrafusal
component and was therefore explicable
on the basis of the observed pattern of motor innervation, though the identity
of the axon could not be
conclusively proved. Individually, instances of shared sensory terminals
and motor input of bag1 and chain
fibres are rare in the cat; their combination in a single spindle with
correlated physiology is described here
for the first time. The observation is considered in relation to the importance
of dynamic and static
segregation in motor control, since it may imply that there is a lower
limit to the degree of segregation that
the developmental programme can provide.