Chapter 14 · Nerves, reflexes and pathways · Topic 76

Reflexes

A&P ICell-to-cell communicationHomeostasisInteractive lesson

A reflex is a fast, automatic response that the same stimulus sets off the same way every time, and every reflex runs on the same five-part circuit, the reflex arc. This page walks through the reflex arc part by part, the stretch reflex behind the knee jerk (and why the "deep tendon reflex" is really a stretch reflex), how clinicians grade tendon reflexes, the tendon reflex that responds to muscle tension, the withdrawal and crossed-extensor reflexes that pull your foot off a tack and keep you standing, and the difference between somatic and visceral reflexes.

What makes a response a reflex

A doctor taps just below your kneecap and your lower leg kicks forward. You did not decide to kick, you could not easily stop it, and it happens the same way every time the tap lands in the same place. A moment later you notice the tap, but the kick has already happened.

That is a reflex (re- = back, flex- = bend: a signal "bent back" out of the CNS): a rapid, involuntary response to a particular stimulus, produced by a fixed circuit of neurons, so that the same stimulus produces the same response each time. Three features set a reflex apart from a voluntary movement:

Some reflexes are built in from birth, such as the knee jerk and a newborn's grasp. Others are learned by repetition, such as braking when a light turns red. This page is about the built-in ones.

The reflex arc

Every reflex runs on a reflex arc: the chain of structures a signal passes through from stimulus to response. It has five parts, always in this order (Figure 1):

  1. Sensory receptor. It detects the stimulus and produces a graded receptor potential. In the knee jerk, it is a sensory receptor inside the thigh muscle that responds to stretch.
  2. Sensory neuron. It carries action potentials into the CNS. For a spinal reflex, its cell body sits in a dorsal root ganglion and its axon enters through the dorsal root.
  3. Integration center. The place in the CNS where the incoming signal is passed on to outgoing neurons: one or more synapses in the gray matter of the spinal cord or brainstem. It may be a single synapse or a chain of interneurons.
  4. Motor neuron. It carries action potentials out of the CNS. For a spinal reflex to a skeletal muscle, this is the lower motor neuron, whose axon leaves through the ventral root.
  5. Effector. The muscle or gland that responds: in the knee jerk, the quadriceps femoris, which contracts and straightens the knee.
spinal cord (dorsal side up) 3 integration center: one synapse in the gray matter dorsal root ganglion 2 sensory neuron 4 motor neuron (out through the ventral root) 1 muscle spindle (sensory receptor, senses stretch) 5 quadriceps (effector) contracts; knee straightens tap stretches it
Figure 1. The five parts of a reflex arc, drawn for the knee jerk. Sensory (afferent) parts are in one color and motor (efferent) parts in another. Here the integration center is a single synapse.

A useful way to use the arc: if a reflex is missing, the fault can be in any of the five parts. A cut dorsal root, a damaged segment of the cord, a cut ventral root or a diseased muscle can each abolish the same knee jerk. The rest of the exam tells you which part failed.

One synapse or many

Somatic and visceral reflexes

Reflexes are sorted by their effector, which matches the two parts of the motor division you met in the organization of the nervous system:

The sensory half of the two kinds is alike. The motor half differs, as Figure 2 shows: a somatic reflex uses one motor neuron that runs all the way from the CNS to the muscle, while a visceral reflex uses a chain of two, which synapse in a ganglion outside the CNS.

Two pathways from a spinal cord cross section. Top, the autonomic efferent pathway: a myelinated axon runs from the lateral horn to a ganglion, where it synapses on a second neuron whose unmyelinated axon runs on to a curved strip of smooth muscle; an enlarged inset shows a swelling along that axon releasing transmitter onto the muscle surface. Bottom, the somatic efferent pathway: one myelinated axon runs from the ventral horn directly to a bundle of skeletal muscle fibers, and an inset shows it ending in a single synaptic terminal.
Figure 2. The motor half of a visceral reflex (top) and a somatic reflex (bottom). In the visceral reflex, a neuron in the spinal cord sends a myelinated axon to a ganglion, and a second neuron carries the signal on through an unmyelinated axon to smooth muscle. In the somatic reflex, one myelinated motor neuron runs from the cord straight to skeletal muscle. OpenStax Anatomy and Physiology 2e, Figure 15.6, openstax.org, CC BY 4.0.
Somatic reflexVisceral (autonomic) reflex
EffectorSkeletal muscleSmooth muscle, cardiac muscle or glands
Motor neurons from CNS to effectorOneTwo, synapsing in a ganglion outside the CNS
Motor axonsMyelinated all the way, fastFirst myelinated, second unmyelinated, slower
Effect on the targetAlways excites (contraction)Can excite or inhibit
Sensory sideSensory receptors mostly in skin, muscles and jointsSensory receptors mostly in internal organs and vessels
ExamplesKnee jerk; pulling your foot off a tackPupils narrowing in bright light; saliva flowing when you eat

The rest of this page is about somatic spinal reflexes, the ones tested in every physical exam. The visceral reflexes come back with the autonomic nervous system and with the organs they control.

The stretch reflex

The sensor: the muscle spindle

Scattered through almost every skeletal muscle are small sensory organs called muscle spindles, named for their shape: fat in the middle and tapering at both ends, like the spindle of a spinning wheel. Each is a capsule a few millimeters long holding a handful of thin, modified muscle fibers. Sensory nerve endings wrap around the middle of those fibers. Muscle spindles are proprioceptors: they lie parallel to the ordinary muscle fibers, so when the whole muscle is stretched, they are stretched too, and stretch opens mechanically gated channels in the wrapped endings. The more and the faster the muscle stretches, the faster the spindle's sensory neuron fires.

The circuit

The stretch reflex is the contraction of a muscle in response to its own stretch. Follow it for the knee jerk, which clinicians call the patellar reflex:

  1. The tap lands on the patellar ligament, just below the kneecap (clinicians usually call it the patellar tendon). The ligament is pushed in, which pulls on the quadriceps femoris and stretches it for a split second.
  2. Muscle spindles in the quadriceps stretch, and their sensory neurons fire a burst of action potentials.
  3. The sensory axons enter the spinal cord through the dorsal roots of about L2 to L4.
  4. In the ventral horn, they synapse directly on the lower motor neurons of the same muscle and release glutamate. One synapse: the stretch reflex is monosynaptic.
  5. Those motor neurons fire, and acetylcholine at their neuromuscular junctions makes the quadriceps contract.
  6. The knee straightens, and the lower leg kicks forward.

A stretch reflex resists a change in muscle length: stretch a muscle and it contracts back. That is why the same circuit helps keep you upright. When you start to sway forward, your calf muscles are stretched, their spindles fire, and the calf muscles contract and pull you back.

The antagonist relaxes: reciprocal inhibition

The quadriceps straightens the knee, and the hamstrings bend it. If the hamstrings contracted at the same moment, they would fight the kick. They do not, because branches of the same spindle sensory axons also excite inhibitory interneurons in the cord. These release glycine onto the motor neurons of the hamstrings, which produces IPSPs and stops them firing. Exciting one muscle while inhibiting its antagonist is called reciprocal inhibition. So a stretch reflex has a monosynaptic part (to the stretched muscle) and a polysynaptic part (to its antagonist).

Deep tendon reflexes and how they are graded

When a clinician taps a tendon with a reflex hammer, the test is called a deep tendon reflex. The name is misleading: the sensor is not in the tendon. The tap stretches the muscle through its tendon, and the muscle spindles respond. Every deep tendon reflex is a stretch reflex. Each one checks a particular set of spinal cord segments:

ReflexWhere you tapWhat happensMain segments
BicepsBiceps brachii tendon at the front of the elbowElbow bendsC5–C6
BrachioradialisBrachioradialis tendon above the wrist, thumb sideForearm bends and turnsC5–C6
TricepsTriceps brachii tendon above the back of the elbowElbow straightensC7
Patellar (knee jerk)Patellar ligament below the kneecapKnee straightensL3–L4
Ankle jerkCalcaneal (Achilles) tendon at the back of the ankleFoot points down (plantar flexion)S1

A rhyme many students use counts up from the ankle: "one, two, buckle my shoe" (S1–S2, ankle), "three, four, kick the door" (L3–L4, knee), "five, six, pick up sticks" (C5–C6, biceps), "seven, eight, lay them straight" (C7–C8, triceps). The table lists the main segment of each pair.

The grading scale

The briskness of each response is graded on a scale of 0 to 4+ and always compared with the same reflex on the other side:

GradeMeaning
0Absent, even with reinforcement
1+Present but weak (diminished); may need reinforcement to appear
2+Normal
3+Brisker than average; may or may not be abnormal
4+Very brisk, with clonus: a run of rhythmic jerks after a single stretch

Reinforcement means asking the person to clench their teeth or hook their fingers together and pull just as you tap. It makes a weak reflex easier to see, so a reflex is only called absent if it stays absent with reinforcement.

What the grades point to follows from the arc and from what you learned about motor pathways:

Worked example: how long does a knee jerk take?

Problem. In an adult, the sensory path from the quadriceps to the spinal cord segments of the knee jerk is about 0.6 m long, and the motor path back is also about 0.6 m. Spindle sensory axons conduct at about 70 m/s and the motor axons at about 60 m/s. The central synapse adds about 1 ms, and the neuromuscular junction plus the start of contraction add about 3 ms. Estimate the time from tap to muscle response.

  1. Time in the sensory axon. time = distance ÷ speed = 0.6 m ÷ 70 m/s = 0.0086 s = 8.6 ms.
  2. Time in the motor axon. 0.6 m ÷ 60 m/s = 0.010 s = 10 ms.
  3. Add the synapse. One central synapse: 1 ms.
  4. Add the muscle. Neuromuscular junction and start of contraction: 3 ms.
  5. Total. 8.6 + 10 + 1 + 3 = 22.6 ms.

Answer. About 23 ms, close to what is measured. Notice that almost all of it is spent in the axons. A disease that strips myelin from peripheral axons, and so slows conduction, lengthens the delay and weakens the reflex, while one extra synapse would add only about a millisecond.

The tendon reflex

Muscles have a second kind of proprioceptor. The Golgi tendon organ sits where muscle fibers join their tendon. It is a capsule of collagen strands woven through with sensory nerve endings. Because it lies in series with the muscle fibers, it is squeezed when the muscle pulls on its tendon, so it reports muscle tension (force), mostly from active contraction.

The tendon reflex runs from it: rising tension fires the tendon organ's sensory neuron, which excites inhibitory interneurons in the cord. These inhibit the motor neurons of the same muscle, and other interneurons excite the motor neurons of the antagonist. The contracting muscle's force is damped. Because interneurons are involved, the tendon reflex is polysynaptic.

Muscle spindleGolgi tendon organ
LocationAmong the muscle fibers, parallel to themAt the junction of muscle and tendon, in series with the fibers
DetectsMuscle length and how fast it changes (stretch)Muscle tension (force)
Reflex it startsStretch reflexTendon reflex
Synapses in the arcOne to the same muscle (monosynaptic)Two or more (polysynaptic)
Effect on the same muscleExcites: it contractsInhibits: its force drops
Effect on the antagonistInhibits itExcites it

The withdrawal and crossed-extensor reflexes

You step on a tack with your left foot. Before you feel any pain, your left knee and hip bend and lift the foot off the tack, and your right leg stiffens to take your whole weight. Two reflexes did this (Figure 3):

tack in the left sole: nociceptors fire sensory neurons enter the cord interneurons: several segments, some cross the midline left flexors contract left extensors relax right extensors contract right flexors relax same side: withdrawal reflex the left leg bends and lifts opposite side: crossed extensor the right leg straightens, bears weight
Figure 3. The withdrawal and crossed-extensor reflexes. One painful stimulus drives opposite patterns in the two legs, through interneurons in the cord. Solid arrows mean "causes".

These reflexes show what interneurons add:

Meanwhile, the same nociceptor signals also climb the spinothalamic tract to the thalamus and cortex. Only when they arrive do you feel the pain, which is why you often find your foot already lifted by the time it hurts.

Stretch reflexWithdrawal reflex
StimulusStretch of a muscleA harmful stimulus to the skin
Sensory receptorMuscle spindleNociceptors
SynapsesMonosynaptic to the stretched musclePolysynaptic
Segments involvedOne or twoMany
ResponseThe stretched muscle contractsFlexors of the limb contract, extensors relax
Opposite limbNot involvedExtends (crossed-extensor reflex)

Superficial reflexes and the Babinski sign

Not every reflex tested in a clinic is a stretch reflex. A superficial reflex is set off by stroking the skin. The one you met with motor pathways is the plantar reflex: stroking the outer sole curls the toes down in a healthy adult. Its arc runs through the lower lumbar and sacral segments, but, unlike a stretch reflex, its normal form depends on the corticospinal tract. When that tract is damaged, the response flips to the Babinski sign, with the big toe extending upward. Brisk deep tendon reflexes and a Babinski sign together are the reflex picture of upper motor neuron damage.

Reflexes and the brain

A spinal reflex does not need the brain, but in a healthy person the brain constantly adjusts it. Descending tracts from the brainstem and cortex excite or inhibit the motor neurons and interneurons of every reflex arc, and they set how sensitive the muscle spindles are. That is why:

That is what makes reflexes so useful at the bedside. Tapping one tendon tests one arc: its sensory neurons, its segments of the cord, its motor neurons and the descending control above it. The next topic puts reflex testing together with the rest of a clinician's checks of the nervous system.