REFLEXES INTRODUCTION The basic unit of integrated reflex activity is the reflex arc.

. This arc consists of a sense organ, an afferent neuron, one or more synapses in a central integrating station or sympathetic ganglion, an efferent neuron, and an effector. The afferent neurons enter via the dorsal roots or cranial nerves and have their cell bodies in the dorsal root ganglia or in the homologous ganglia on the cranial nerves. The efferent fibers leave via the ventral roots or corresponding motor cranial nerves. The principle that in the spinal cord the dorsal roots are sensory and the ventral roots are motor is known as the Bell- Magendie law. Activity in the reflex arc starts in a sensory receptor with a receptor potential whose magnitude is proportionate to the strength of the stimulus. This generates all-or-none action potentials in the afferent nerve, the number of action potentials being proportionate to the size of the generator potential. In the CNS, the responses are again graded in terms of EPSPs and IPSPs at the synaptic junctions.

 All-or-none responses are generated in the efferent nerve. When these reach the effector, they again set up a graded response. When the effector is smooth muscle, responses summate to produce action potentials in the smooth muscle, but when the effector is skeletal muscle, the graded response is always adequate to produce action potentials that bring about muscle contraction. The connection between the afferent and efferent neurons is usually in the CNS, and activity in the reflex arc is modified by the multiple inputs converging on the efferent neurons. The simplest reflex arc is one with a single synapse between the afferent and efferent neurons. Such arcs are monosynaptic, and reflexes occurring in them are monosynaptic reflexes. Reflex arcs in which one or more interneurons are interposed between the afferent and efferent neurons are polysynaptic, the number of synapses in the arcs varying from two to many hundreds. In both types, but especially in polysynaptic reflex arcs, activity is modified by spatial and temporal facilitation, occlusion, subliminal fringe effects, and other effects. MONOSYNAPTIC REFLEXES: THE STRETCH REFLEX Introduction

- When a skeletal muscle with an intact nerve supply is stretched, it contracts. This response is called the stretch reflex. - The stimulus that initiates the reflex is stretch of the muscle, and the response is contraction of the muscle being stretched. The sense organ is the muscle spindle. - The impulses originating in the spindle are conducted in the CNS by fast sensory fibers that pass directly to the motor neurons which supply the same muscle. - The neurotransmitter at the central synapse is glutamate. Clinical Examples Tapping the patellar tendon elicits the knee jerk, a stretch reflex of the quadriceps femoris muscle, because the tap on the tendon stretches the muscle. Tapping on the tendon of the triceps brachii, for example, causes an extensor response at the elbow as a result of reflex contraction of the triceps; tapping on the Achilles tendon causes an ankle jerk due to reflex contraction of the gastrocnemius; and tapping on the side of the face causes a stretch reflex in the masseter. Structure of Muscle Spindles Each muscle spindle consists of up to about 10 muscle fibers enclosed in a connective tissue capsule. They are called intrafusal fibers to

distinguish them from the extrafusal fibers, the regular contractile units of the muscle. The intrafusal fibers are in parallel with the rest of the muscle fibers because the ends of the capsule of the spindle are attached to the tendons at either end of the muscle or to the sides of the extrafusal fibers. There are two types of intrafusal fibers in mammalian muscle spindles. The first type contains many nuclei in a dilated central area and is therefore called a nuclear bag fiber. Typically there are two nuclear bag fibers per spindle, nuclear bag fiber 1 with a low level of myosin ATPase activity and nuclear bag fiber 2 with a high level of myosin ATPase activity. The second fiber type, the nuclear chain fiber, is thinner and shorter and lacks a definite bag. There are four or more of these fibers per spindle. Their ends connect to the sides of the nuclear bag fibers. The ends of the intrafusal fibers are contractile, whereas the central portions probably are not. There are two kinds of sensory endings in each spindle. The primary (annulospiral) endings are the terminations of rapidly conducting group Ia afferent fibers. One branch of the Ia fiber innervates nuclear bag fiber 1, whereas another branch innervates nuclear bag fiber 2 and nuclear chain fibers. These sensory fibers wrap around the center of the nuclear bag and nuclear chain fibers.

The secondary (flower-spray) endings are terminations of group II sensory fibers and are located nearer the ends of the intrafusal fibers but only on nuclear chain fibers. The spindles have a motor nerve supply of their own. These nerves are 3-6 um in diameter, constitute about 30% of the fibers in the ventral roots, and belong in Erlanger and Gasser's A group. Because of their characteristic size, they are called the efferents of Leksell or the small motor nerve system. They go exclusively to the spindles. In addition, larger motor neurons innervate both intrafusal and extrafusal fibers. The endings of the efferent fibers are of two histologic types. There are motor endplates (plate endings) on the nuclear bag fibers, and there are endings that form extensive networks (trail endings) primarily on the nuclear chain fibers. The spindles produce two kinds of sensory nerve patterns , dynamic and static (see below), and both and motor axons produce two functional types of responses. Stimulation of one type increases dynamic responses (dynamic fusimotor axons), and stimulation of the other increases static discharge at constant length (static fusiform axons).

Central Connections of Afferent Fibers