Physiology

Reflexes: principles and

properties

levels: those involving the somatic nervous system, integrated at

the level of the spinal cord but of which the subject is generally
aware (walking, maintenance of posture); and those concerning
the autonomic nervous system (digestion of food, cardiac and
metabolic responses to a stressor), of which the subject is not
normally aware. These latter reflexes are integrated at the level
of the midbrain and medulla.

James Waterhouse
Iain Campbell

Somatic reflexes
The stretch reflex is the basis of posture and is responsible for
maintaining a set length for the muscles that support the weight
of the body, so preventing us from falling over as a result of
the influence of gravity. This reflex can be observed when tapping the tendon of a muscle; the rapid stretch produced by the
tap results in a reflex contraction of the muscle, seen most commonly with the patellar tendon of the quadriceps femoris or the
triceps tendon above the elbow. The sensor organ in the skeletal
muscles is the muscle spindle, which detects stretching of the
muscle; the effector organ is the voluntary striated muscle supplied by -motor neurones from the ventral root of the spinal
cord. These two components combine in a negative feedback
loop which maintains muscle length and posture.
The muscle spindle is a complex structure consisting of up to
ten muscle fibres enclosed in a connective tissue capsule. These
are termed intrafusal fibres, to distinguish them from the extrafusal fibres that make up the bulk of the muscle, and which
produce the contractile force when the muscle shortens. They
are connected to the tendons at each end of the muscle or to the
sides of the extrafusal fibres.
There are two types of intrafusal fibre nuclear bag and
nuclear chain fibres (Figure 1). A muscle spindle consists of about
two nuclear bag fibres and four or more nuclear chain fibres.
Nuclear bag fibres have contractile ends and a non-contractile
central dilated area containing several nuclei. The nuclear chain

Abstract
The body responds to changing circumstances and environmental threats
both consciously and subconsciously. The cognitive response to a physi
cal threat normally involves movement mediated by skeletal muscle.
There are a number of control mechanisms hardwired into the nervous
system which enable muscle systems to respond in an integrated fash
ion without involving a conscious decision, although the subject is usu
ally conscious of what has happened. These include the stretch reflex,
the withdrawal reflex and the crossed extensor reflexes. Muscle spindles,
Golgi tendon organs and cutaneous nociceptors provide the sensory
input to these reflexes, and muscle spindles also play a role in the con
trol of voluntary movement. The autonomic nervous system controls the
internal environment in response to environmental change. It consists
of the parasympathetic division, which controls basal and vegetative
mechanisms, and the sympathetic nervous system, which controls vis
ceral adaptive responses to any sort of environmental change or threat.

Keywords fight or flight; neurotransmitters; sympathetic and para

sympathetic nervous system; synapses; withdrawal and crossed extensor
reflexes

The concept of homeostasis maintaining the constancy of

the internal environment is fundamental to physiology; for
this constancy to be achieved, a sensor (to measure the variable being controlled), an effector (to change this variable) and
some linking pathway are required. Homeostasis results when
these components act together, forming negative feedback loops.
Similar components are needed when adaptation to change is
required escaping from a noxious stimulus or predator for
example although they will be wired together differently to
implement change rather than to maintain a steady state. The
activities of the various excitable tissues sense organs, neurones and muscles are integrated so that various controlled
processes can ensue. Many of these processes do not require
cognition (thought) but are automatic. They occur also on two

Diagrammatic representation of the main components

of mammalian muscle spindle

-efferent
To extrafusal fibres
Plate endings

James Waterhouse, DPhil, DSc, is Professor of Biological Rhythms at the

Research Institute for Sport and Exercise Sciences, Liverpool John
Moores University.

-efferent

II secondary afferent

Annulospiral
ending

Trail ending

Iain Campbell, MD, FRCA, is Consultant Anaesthetist at the University

Hospitals of South Manchester NHS Trust and Visiting Professor
of Human Physiology at Liverpool John Moores University. He
qualified from Guys Hospital Medical School, London, and trained in
anaesthesia in Zimbabwe, Southend, Montreal and Leeds.

ANAESTHESIA AND INTENSIVE CARE MEDICINE 9:5

Ia primary afferent
from nuclear bag fibre 2

Nuclear bag
fibre
Nuclear chain
fibre
Flower-spray ending

Each spindle has a capsule and usually contains two nuclear bag fibres and
four or more nuclear chain fibres. Reproduced from Ganong WF. Review of
Medical Physiology, 19th edn. Connecticut: Appleton and Lange, 1999. By
permission of the McGraw-Hill companies.

Figure 1

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fibres are smaller and their nuclei are laid out along the length
of the fibre. As implied above, the muscle spindles are stretch
receptors. They give rise to two types of large-diameter, afferent
(sensory) fibre: group Ia afferents arise from annulospiral endings around the central section of both types of intrafusal fibre;
group II afferents arise from flower-spray endings around the
ends of the nuclear chain fibres. Both types of afferent fibre pass
into the dorsal root of the spinal cord, the cell bodies lying in the
dorsal root ganglion, and form monosynaptic connections with
the -motor neurones supplying the extrafusal fibres of the same
muscle. The fact that both the afferent and efferent fibres are of
large diameter, coupled with the presence of only one synapse
in the pathway, means that the stimulus response delay time is
minimal, about 20 ms in the case of a simple knee jerk.
The spindles are arranged in parallel with the extrafusal fibres
of the muscle and monitor its length. If the muscle is stretched,
the spindles produce more afferent impulses and these cause the
muscle to contract (as with a knee jerk), so opposing the ori
ginal stretch. By contrast, if the muscle length decreases then the
spindle is unloaded and its rate of firing is reduced; the resultant
reduced rate of firing of the -motor neurones reduces muscle
contraction and allows it to lengthen.
Even though this negative feedback mechanism would, in
principle, act to preserve an individuals posture, in practice it
is too slow. One of the problems of the system as described so
far (and this applies to feedback loops in general) is that it meas
ures the length of the muscle and so has static sensitivity only.
As we begin to lose our balance the initial error in length of the
muscle is small, and so the response the feedback loop produces
is small too small to be effective. This problem is overcome
by the dynamic sensitivity of the spindles. Dynamic sensitivity
responds to change rather than state, and the dynamic compon
ent of the reflex comes from the Ia afferents arising from around
the nuclear bags of the nuclear bag fibres. These fibres are sensitive to the rate of lengthening of the muscle, although the precise
mechanism is not yet clear. Sensory nerves originating from the
nuclear chain fibres show only static responses. The result of this
dynamic input is that, as soon as the muscle starts to lengthen,
there is a burst of activity from the afferent (nuclear bag) nerves
that leads to a sufficiently large muscle contraction.
In practice the basic stretch reflex is complicated in three
main ways.
Reciprocal innervation through a pathway involving two
synapses, increased input from the Ia fibres causes inhibition of
the antagonistic muscle. There is an obvious benefit in having reflex changes to antagonistic muscles behaving inversely to each
other.
Efferent fibres the above mechanism is appropriate for
maintaining posture, where the length of the muscles needs to
be maintained at a set value, but a problem arises if movement,
which requires a controlled contraction or relaxation, is required.
Beta-efferent and -efferent fibres pass to the ends of both nuclear
bag and nuclear chain fibres and form plate endings and trail
endings (Figure 1). These efferent fibres are of two types and
can stimulate either the dynamic or static response of the muscle
spindle. Changes in sensitivity of the spindle are brought about
by contraction of the intrafusal fibres that stretch the area under
the sensory nerve endings. If the firing of these efferent fibres
increases, the increased sensory input from the spindles causes a

ANAESTHESIA AND INTENSIVE CARE MEDICINE 9:5

reflex contraction of the extrafusal fibres by the normal pathway.

Likewise, a decreased rate of firing of these efferents will lead to
muscle relaxation. In practice, when a voluntary movement is
initiated by the motor cortex, for example, there is a co-activation of the -motor neurones (which provide the contractile force
via the extrafusal fibres) and - and -fibres (which change the
set-point of the system the length at which the muscle is controlled by the stretch reflex). This co-activation of -, - and fibres enables a controlled contraction or relaxation to take place.
As a further example of the value of both intrafusal and extrafusal
fibres, there is a high ratio of intrafusal fibres to extrafusal fibres
in finely controlled muscles, such as those controlling the eye;
this enables the length of these muscles and changes in length
to be controlled with great accuracy. As a result, both eyes can
point in exactly the same direction, as required for stereoscopic
vision. The activity of the -motor neurones and their efferent
fibres is affected by several other sources, including noxious
stimuli to the skin and muscle effort elsewhere in the body. For
example, clasping the hands tightly and trying to pull them apart
will increase the tendon reflex due to the irradiation of nervous
activity down the spine.
Inverse stretch reflex the Golgi tendon organs are, as their
name suggests, found in the tendons; they have a network of
knobbly endings that ramify through the tendon bundles and
send afferent impulses to the spinal cord via group Ib afferents.
These afferent fibres make connections with the inhibitory inter
neurones that terminate on -motor neurones supplying the
extrafusal fibres of the same muscle. They also connect with excitatory fibres innervating antagonists to the muscle. Their position means that they are in series with the muscle and act as sensors of the force exerted by the contracting muscle rather than its
length. Although their threshold is low, so that they are activated
during normal muscle contractions, their effect is evident only
when the force exerted by the muscle becomes large. In such
cases, they inhibit muscle contraction and the muscle suddenly
relaxes the inverse stretch reflex. The value of this reflex is to
prevent the development of excessive forces during contraction.
Although the above changes are often described as simple
reflexes, this refers to the lack of requirement of any processing
by the central nervous system (CNS). Even so, information does
pass up the spinal cord into the CNS, and this input to the cerebellum is an important part of controlling fine movement. The
reflexes interact in several ways and integrate activities between
antagonistic muscles. Figure 2 summarizes the pathways and
neuronal arrangements of these reflexes.

The withdrawal and crossed extensor reflex

The withdrawal reflex is pre-potent, that is, its operation overrides that of the stretch and inverse stretch reflexes. It is polysynaptic, involving activation of some muscles and inhibition of
others. It is initiated by nociceptive stimuli, generally impinging
upon the skin, that are potentially harmful to the animal. There is
a flexion of the associated limb of the leg when a person stands
on something sharp, or of the arm when the hand touches something that is hot, for example and a relaxation (inhibition) of the
antagonist muscles. The stronger the stimulus, the more marked
the response and the wider the area of the body that is affected.
To take the example of standing on a sharp object: if the stimulus
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some of the neurones involved send excitatory impulses back to

earlier parts of the pathway, so forming reverberating circuits.
These circuits, as well as the multiple pathways, prolong the time
during which activation of the -motor neurones is present, a
phenomenon known as after-discharge.
Figure 3 summarizes the above reflexes in terms of the receptors and effectors involved and the nature of the connecting
neural pathway.

Summary of stretch and inverse stretch reflexes

Length of muscle
-efferents
-efferents

Spindles

Autonomic reflexes
Ia
(1)

II
(1)
+

II
(poly)

Ia
(2)

The autonomic nervous system (ANS) controls visceral function

and so regulates our internal environment, in particular by controlling smooth (involuntary) muscle in the blood vessels and
gastrointestinal tract. This is in contrast to the somatic nervous
system discussed above, which controls (voluntary) skeletal
muscle and thus the way we interact, usually consciously, with
our external environment.
The ANS is composed of two divisions the parasympathetic
(PNS) and the sympathetic (SNS) nervous systems. There is also
an enteric nervous system which controls many of the functions of
the gastrointestinal tract, and which is strongly influenced by the
input from both divisions of the ANS. The central components of
the ANS are in the midbrain and medulla: they include the limbic
system (emotion), the hypothalamus, parts of the brainstem and
the spinal cord. Myelinated axons arise in the CNS and synapse
in peripheral ganglia with cells whose axons innervate the organs
they control. These post-ganglionic neurones are un-myelinated.
The ANS also has afferent (sensory) neurones. They transmit not
only sensory information from visceral structures but also chemical and mechanical data from chemoreceptors (carotid body) and
pressure receptors (carotid sinus, aortic arch).

-motor neurones
Agonist

-motor neurones
Antagonist

Ib
(poly)
Golgi tendon organ
+
Tension in muscle
+, excite; , inhibit
Width of arrow compares importance
Number in brackets shows number of sensors; poly, many
Roman numeral indicates sensory group

Figure 2

Wiring diagrams of the three spinal reflexes

is relatively mild, then the leg is withdrawn; as the stimulus

gets stronger, flexion is faster and lasts longer (involving afterdischarge and reverberating circuits, see below). Even stronger
stimuli cause the whole weight of the body to be shifted onto the
other foot, thus supporting the body; this requires the extensors
of this limb to contract the crossed extensor reflex and there
might also be reflex changes in the upper limbs, with extension
of the ipsilateral upper limb (as if to push the person away from
the stimulus) and flexion of the contralateral limb. This spread of
activity up or down the spinal cord (as in the example above, or
factors affecting the activity of the -efferent fibres) is described
as the irradiation of the stimulus. Information reaches the CNS
and consciousness, the individual registering the effects of pain.
Not surprisingly, the reflex involves the simultaneous activity
of many neurones and neuronal circuits, some excitatory, others
inhibitory. Details of which circuits are involved, and whether
there is some degree of activation of abductor or adductor
muscles in addition to those involved in flexion, depends upon
the stimulus site. Moreover, whereas the stretch reflex generally
lasts for a short period of time (before another postural adjustment is required), the flexor response will last for longer. This
longer period of activation is achieved by the presence of many
polysynaptic pathways, the result of which is that the stimulus
affects the -motor neurones and muscles on multiple occasions,
having traversed many pathways of different length. Moreover,

ANAESTHESIA AND INTENSIVE CARE MEDICINE 9:5

Receptor
a. Spindle

b. Tendon
organ

Spinal cord

+
+

Effector and effect

+
+

c. Pain
receptor

Interneuron
+ pool with
reverberating
circuits

Agonist (homonymous)
muscle
Antagonist muscle
Agonist muscle
Antagonist muscle
Flexor muscle

Withdrawal
reflex Antagonist (extensor)
muscle
+
Crossed
extensor
reflex

Contra-lateral extensor
muscle
Contra-lateral flexor
muscle

a. Stretch reflex
b. Inverse stretch reflex
c. Withdrawal and crossed extensor reflexes

Figure 3

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Autonomic reflexes enable our physiological systems to cope

with a spectrum of challenges, and this is generally done subconsciously. The reflexes control basic functions, such as the
response to eating a meal or standing up. They respond also
to physiological and environmental demands, such as exercise,
injury, heat, cold, altitude, gravity (space travel) and extremes
of pressure (when deep-sea diving). The numerous individual
reflexes the ANS is involved in are too many and too complex to
cover in detail here. The reader is referred to the relevant chapters
elsewhere in this journal, in particular to those on cardiovascular,
respiratory and gastrointestinal physiology (including vomiting
and defecation), as well as micturition and those on integrated
and environmental physiology (exercise, injury, extremes of
barometric pressure and thermoregulation). This chapter will
cover only the general principles of autonomic control.

and the parasympathetic, the cervix. The sympathetic nerves

supplying the head leave the spinal cord with the upper thoracic nerves but synapse with their post-ganglionic cells in the
superior and middle cervical ganglia and in the stellate ganglion.
The detailed anatomy of the ANS is described elsewhere (Anaesthesia and intensive care medicine 2008 9:2: 3941).
Parasympathetic activity varies over a wide spectrum; in contrast, sympathetic activity tends to be an all-or-none phenomenon. A sympathetic discharge results in stimulation of all the
functions controlled by the SNS, including cardiac output, blood
pressure, blood sugar and sweating. As the SNS is usually stimulated in response to a stress, there is a need for an immediate
reaction in a widespread range of systems.

Neurotransmitters
The neurotransmitters differ between the two divisions of the
ANS. The ganglionic transmitter in both instances is acetyl choline, but the main transmitter released by post-ganglionic sympathetic nerves is noradrenaline and the transmitter released by the
post-ganglionic/post-synaptic terminals of the parasympathetic
nerves is acetyl choline. There are exceptions, however; acetyl
choline is released by some fibres innervating blood vessels in
skeletal muscle in some species and the sympathetic fibres innervating sweat glands are also cholinergic.
Cholinergic and noradrenergic receptors are subdivided according to their reaction to certain drugs. Cholinergic fibres are either
muscarinic or niocotinic. The receptors on post-ganglionic neurons are nicotinic, whereas the receptors on target organs are
usually muscarinic. Likewise, there are two types of noradrenergic
receptor, alpha () and beta (); -receptors respond to isoprenaline more than do -receptors. Both types of receptor have sub
divisions 1-receptors are found in post-synaptic nerve endings,
whereas 2-receptors are found predominantly on the pre-synaptic
terminals of the adrenergic fibres and inhibit further discharge of
noradrenaline. Both types of -receptor and 1-receptors respond
about equally to adrenaline and noradrenaline, whereas 2receptors respond more to adrenaline than noradrenaline.

Anatomy
The PNS consists of four cranial nerves (III occulomotor,
VII facial, IX glossopharyngeal and X vagus) and the sacral
nerves, S2S4, which supply the pelvic viscera bladder, lower
colon, rectum and uterine cervix; these are termed the cranial
and sacral outflows, respectively. Generally speaking, the
effects of parasympathetic activity are relatively discrete, limited
to the organ system(s) supplied by any particular nerve, although
a number of systems are supplied by the vagus; they include
the heart, bronchial tree and gastrointestinal tract. The effects of
vagal stimulation are therefore relatively widespread.
The parasympathetic pre-ganglionic fibres are small myelinated B axons; they synapse with post-ganglionic cells with (unmyelinated) C axons. The ganglia are situated on or near the
structures they innervate (Figure 4). The cell bodies of vagal
ganglia lie as microscopic collections of cells within the walls of
the viscera themselves; the post-ganglionic fibres therefore have
very short axons.
The cell bodies of the sympathetic nerves are situated in
the spinal cord, in the lateral horn of the grey matter, over the
whole length of the cord, although they exit the spinal cord only
between T1 and L3. They, too, are made up of relatively slow,
myelinated B axons, which leave the spinal cord as part of the
spinal nerves. They branch off from the spinal nerves and pass in
white (myelinated) rami communicantes (Figure 4) to the sympathetic ganglionic chain situated on the sides of the thoracic
vertebral bodies. Some synapse with post-ganglionic cells whose
un-myelinated C fibres pass via the grey rami communicantes
to their target organs. In line with their diffuse effects, one preganglionic fibre may synapse with several post-ganglionic fibres.
Post-ganglionic fibres usually travel to their target organs in close
proximity to the blood vessels supplying the same organ.
Other pre-ganglionic fibres pass straight through the thoracic
ganglia to synapse with ganglia in the periphery. The splanchnic
nerves contain such fibres; they are part of the thoracic sym
pathetic outflow and synapse in ganglia lying on and around the
major abdominal blood vessels the coeliac ganglion as well as
the superior and inferior mesenteric ganglia. Sympathetic nerves
from T10T12 supply branches to the ovarian and the superior and
inferior hypogastric plexuses, where they mingle with fibres from
the sacral parasympathetic outflow and innervate the uterus. The
sympathetic nerves innervate predominantly the uterine body

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Autonomic afferents
Sensory fibres from internal structures such as the viscera, as well
as muscle and bone, travel with both sympathetic and parasympathetic nerves to reach the CNS, their cell bodies lying in either
the dorsal roots or the homologous cranial nerve nuclei. The
nerve fibres are largely small un-myelinated C fibres. There are
no proprioceptors in viscera and few touch or temperature sense
organs. There are pain receptors, although fewer than in somatic
structures. Pain fibres from the lungs and bronchi and from the
pelvis travel largely in parasympathetic nerves; pain fibres from
the abdominal viscera and the heart follow sympathetic nervous
pathways. Pain felt in viscera is often referred to the corresponding somatic segments innervated by somatic nerves at the same
spinal level from which the visceral pain arises. The commonest
examples are cardiac pain going down the arm or into the neck,
and shoulder tip pain arising from the diaphragm.
Visceral pain is poorly localized; it frequently induces nausea
and sets up a reflex sympathetic discharge involving a tachycardia, increased blood pressure and sweating, although visceral
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Functional anatomy of the autonomic nervous system

III Oculomotor
Pupil
VII Facial
Midbrain and medulla

IX Glossopharyngeal
X Vagus

Ganglia

Lacrimal glands
Salivary glands
Mucus membranes
Nose, mouth

Carotid plexus

C
e
r
v
i
c
a
l

Thoracic
1

Heart

Larynx

Cardiac plexus
Trachea

4
Bronchi
Oesophagus

Thoracic

Coeliac ganglion

Stomach
Mesenteric blood vessels
Liver
Adrenal glands
Small intestine
Large intestine
Lumbar

Superior mesenteric ganglion

T
h
o
r
a
c
i
c
g
a
n
g
l
i
a

5
6
7
8
9
10
11
12
Lumbar

Kidney

1
Bladder
2
Sacrum

Sacral outflow

Inferior mesenteric ganglion

Sexual organs

2
3

Parasympathetic division

Sympathetic division

Figure 4

stimulation can produce vasovagal episodes associated with profound bradycardia, loss of cardiac output and loss of consciousness. Stimulation of parasympathetic receptors, traction on pelvic
organs, cervical dilatation, elevation of a fractured zygoma or even
prolonged suctioning of the pharynx can all result in bradycardia,
or even, although usually transient, cardiac standstill.

(VII) synapse in both the pterygopalatine and submandibular

ganglia to supply secretomotor fibres to the submandibular and
sublingual glands as well as sending a branch to the lacrimal
gland. They respond to food intake and to corneal/conjunctival stimulation. The PNS fibres in the glossopharyngeal nerves
synapse in the otic ganglion and supply secretory fibres to the
parotid gland. The glossopharyngeal nerve also carries afferent
nerves from the carotid body and the carotid sinus. The vagus
nerve supplies fibres to the heart (sino-atrial and atrioventricular
nodes) and carries afferents from the baroreceptors in the aortic
arch, as well as a motor nerve to the vocal cords and sensory
fibres to the trachea and bronchi. Further, the vagus provides the
motor supply to the whole of the digestive tract down to the midpoint of the colon, including the liver, gall bladder and pancreas.

General principles relating to autonomic reflexes

As outlined above, much of the anatomy and most of the effects
of the PNS are relatively discrete: the oculomotor nerve (III)
synapses in the ciliary ganglion and innervates the sphincter
muscle of the iris and the ciliary muscle, stimulation constricting the pupil; pre-ganglionic fibres carried by the facial nerve

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The vagus and glossopharyngeal nerves coordinate the swallowing reflex (Figure 4).
The PNS is a vegetative anabolic system that promotes basal
processes. It controls the digestive and absorptive processes
following the ingestion of food by increasing gastrointestinal
motility and stimulating gastrointestinal secretions, including
those of the gall-bladder and pancreas. It also facilitates childbirth and evacuation of the bladder and rectum. It slows the
heart and produces bronchoconstriction and stimulates bronchial
secretions, thereby increasing respiratory resistance.
Sympathetic stimulation produces the classic fight-or-flight
reaction. It responds to pain, blood loss and stress physical
and emotional by constricting peripheral blood vessels, directing blood centrally and to the muscles (away from non-essential
tissues such as the gut and the skin), arousing the central nervous
system and enabling the organism to cope with whatever threat
has arisen. It stimulates adrenaline secretion from the adrenal

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medulla, which essentially functions as a collection of postganglionic adrenergic neurones secreting adrenaline. It produces
pupillary dilatation, inhibits gastrointestinal motility and mobilizes energy substrate (stimulating glycogenolysis, lipolysis and
proteolysis), so releasing glucose, lactate, fatty acids, glycerol and
amino acids into the circulation to provide the energy to cope with
the threat. It regulates blood pressure in all circumstances and
controls thermoregulatory mechanisms in response to heat and
cold. The SNS also reacts to internal threats such as illness, with
sweating, tachycardia and fever being typical responses to such
disturbances as infection, myocardial infarction, pain and injury.
As stated earlier, the precise details of the autonomic reflexes
controlling the function of the various organ systems, such as
respiratory, cardiovascular and gastrointestinal systems, are
covered in the relevant chapters elsewhere in this journal. The
same applies to the integrated responses to, for example, injury,
exercise and high altitude.

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