PHYSIOLOGY

The physiology of
micturition
Christopher Fry

The bladder has a dual function.

• For most of the time it is a compliant muscular organ that stores
up to 1 litre of urine. It is therefore important that the intravesical
pressure is less than that of the upper tract to allow filling.
• Its second function is to facilitate controlled voiding and for
this purpose a sufficient intravesical pressure must be generated
to overcome the resistance of the outflow tract.

Lower urinary tract anatomy

Figure 1 shows the components of the lower urinary tract.
Figure 2 shows the three layers of the bladder dome:
• a transitional epithelium facing the lumen with an underly-
ing submucosal layer containing sensory nerves and blood
vessels
• a detrusor smooth muscle layer
• an adventitial outer layer.
Towards the base of the bladder the ureters open into the lumen
and the triangular region formed with the bladder neck as the
apex, is the trigone (see Figure 1). Towards the bladder neck and
urethra the muscle bundles become smaller with more connective
tissue. The bladder neck forms an internal sphincter mechanism,
because it is closed during filling. However, it is not under vol-
untary control, and there is no readily identifiable anatomical
structure. In men the prostate gland merges with the bladder neck
and contributes to this sphincter mechanism, mainly to prevent
retrograde ejaculation.
The proximal part of the urethra contains a distinct distal
sphincter mechanism, a layer of skeletal muscle (the intrinsic
rhabdosphincter) and a pubo-urethral sling of muscle. This is
common to men and women. However, in men the overall length
is much longer than in women (about 4 cm vs. 20 cm) as it passes
through the penis. This will inevitably increase the resistance
to urine flow in men, so that when the sphincter mechanisms
become dysfunctional the chance of urine leakage is much more
likely in women. The intrinsic rhabdosphincter is probably the
most important component; it forms a horseshoe-like structure
round the urethra, which kinks the tract when it is contracted

Christopher Fry is Professor of Cell Physiology at the Institute of Urology,

University College London. He achieved a PhD in physiology at the
University of Leicester and spent time at the University of Bern and St
Thomas’ Hospital, London. His research interests are in the cellular
processes that regulate excitation–contraction coupling in smooth and
cardiac muscle.

WOMEN’S HEALTH MEDICINE 2:6 53 © 2005 The Medicine Publishing Company Ltd
 PHYSIOLOGY

Lower urinary tract in the female Innervation of the lower urinary tract

Detrusor
dome Efferent Afferent
innervation innervation

Parasympathetic
motor nuclei S2–S4

Sympathetics
T10–L2

Ureteric orifice
Trigone

Bladder neck Onuf’s

nucleus
Afferent
Pelvic nerve
Urethral smooth Somatic nerves
muscle efferent Bladder
nerves

Periurethral musculature

Intrinsic rhabdosphincter Pelvic

1 Pudendal
and reduces flow. The lower urinary tract lies within a number of
supporting structures.1
Nervous control of the bladder (Figure 3)
Efferent innervation: the lower urinary tract receives a dual auto-
nomic innervation; a parasympathetic supply from S2–S4 and a
sympathetic supply from T10–L2.
There is also a dual somatic supply which runs in the nervi
erigentes. Typical motor neurons in levels S2–S4 provide axons
in the pudendal nerve to innervate the pelvic floor musculature
and part of the urethra. A second component provides fibres to
the intrinsic rhabdosphincter.
Afferent innervation from the lower urinary tract is less well
understood. Afferent fibres originate in the suburothelial layer, the
Bladder lumen
Urothelium
Submucosa
Detrusor
muscle
Serosa
2 Three-layered structure of the bladder wall.
plexus
Distal
urethral
nerve sphincter
3
urothelium and detrusor layers. They convey a feeling of bladder
fullness and are involved in micturition reflexes. The transduction
mechanism between bladder filling and afferent excitation was
assumed to be mediated by stretch receptors in the suburothelial
layer, a hypothesis marred only by the inability to identify such
receptors convincingly. More recently, it has been proposed that
the urothelium releases chemical mediators such as ATP when
stretched, and it is these that initiate afferent activity.
Afferent fibres also originate from the trigone and urethral
wall and run in the sympathetic hypogastric nerves and pudendal
nerves, respectively. They respond to greater degrees of bladder
fullness and convey varying levels of urge to void.
Coordination of micturition (Figure 4)2
Micturition requires that the lower urinary tract undergoes two
coordinated processes:
• reduction of bladder outflow resistance
• increase of bladder vesical pressure.
Normal micturition is impossible if these two processes do not
occur together. Although afferent and efferent fibres form spinal
connections, a simple spinal reflex is insufficient to explain this
coordinated process and control from higher nervous centres is
required.
• Most afferent fibres make synaptic connections with ascend-
ing fibres that travel in lateral tracts. It is thought that these tracts
project to the periaqueductal grey, which controls a region of the
rostral pons. This ‘micturition centre’ coordinates different pelvic
functions including micturition, defecation and coitus.

WOMEN’S HEALTH MEDICINE 2:6 54 © 2005 The Medicine Publishing Company Ltd
 PHYSIOLOGY
The coordinated process of bladder filling and voiding
Pressures recorded in the urethra and bladder during filling and
voiding show the coordinated process of outlet relaxation and
detrusor contraction. During filling with 500 ml of liquid, bladder
pressure remains fairly constant despite the large increase in
volume and there is a small, but definite, rise of urethral pressure.
Voiding is characterized by an initial fall of urethral pressure
followed almost immediately by a rise of detrusor pressure.
Detrusor
50 cm H2O
pressure
Urethral
50 cm H2O
pressure
20 ml/second Flow rate
Volume
500 ml
infused
• Other fibres travel to areas in the cerebral cortex (the paracentral
lobule) that control the pelvic floor musculature and to the medial
pre-optic area. Fibres that project to the frontal cortex allow social
and conscious control over micturition.
The ‘micturition centre’ is thought to act as a final common
pathway and controls the coordinated voiding process involving
bladder outlet relaxation and detrusor contraction.
• Descending fibres reduce motor neuron activity to the intrinsic
rhabdosphincter by acting on inhibitory sacral interneurons using
glycine and γ-aminobutyric acid (GABA) as transmitters.
• Other fibres to the urethral smooth muscle may be acti-
vated to relax this tissue, possibly by the release of a relaxing
neurotransmitter (e.g. nitric oxide).
• Descending fibres also probably activate parasympathetic fibres
that cause the detrusor smooth muscle to contract.
The role of sympathetic fibres is less clear. Sympathetic innerva-
tion is most dense to the trigone and bladder neck and a decrease
of activity would relax both these components.
WOMEN’S HEALTH MEDICINE 2:6
Bladder filling
The tissues that comprise the bladder wall are fairly compliant.
They show a remarkable ability to realign themselves when sub-
jected to strain, thus dissipating stress within the tissue (stress
relaxation). Whether there is active relaxation of detrusor smooth
muscle is more contentious, but there is no conclusive evidence.
However, different gating mechanisms may attenuate parasympa-
thetic efferent activity during filling, including:
• sympathetic inhibition on the pelvic plexuses
• gating of afferent information, such that low-threshold activ-
ity does not break through to activate a micturition reflex.
The rise of urethral pressure may be a spinal reflex from local
afferents that either excite parasympathetic fibres and/or inhibit
relaxatory nitrergic efferents.
Smooth muscle function in the lower urinary tract3
The end target of efferent nerves is largely the muscular layers in the
wall of the urinary tract. Spontaneous activity in bladder smooth
muscle is discussed on pages 20–23. In the normally functioning
bladder wall, detrusor is under the exclusive control of cholinergic
fibres. Parasympathetic fibres release acetylcholine that binds to
muscarinic M3 receptors, which elicits a transient rise of intra-
cellular calcium via the intermediate generation of the second
messenger inositol trisphosphate. The importance of muscarinic
receptors in mediating detrusor contraction is the basis of most
of the therapeutic control of bladder overactivity.
In the trigone and bladder neck, sympathetic activation via
α-adrenoreceptor activation plays an important role, alongside
muscarinic activation, in regulating contractile activity. This tissue
tends to show more spontaneous activity, suggesting that there is
more tone in this region of the bladder wall, which may help to
preserve a functional proximal sphincter. 
REFERENCES
1 Mundy A R. Structure and function of the lower urinary tract.
In: Mundy A R, Fitzpatrick J M, Neal D E, George N J R, eds. The
scientific basis of urology. Oxford: Isis Medical Media, 1999:
217–42.
2 Morrison J et al. Neural control. In: Khoury S, Wein A, eds.
Incontinence, 3rd International Consultation on Incontinence,
vol 1. Plymouth: Health Publications Ltd, 2005: 363–422.
3 Fry C H et al. Cell biology. In: Khoury S, Wein A, eds.
Incontinence, 3rd International Consultation on Incontinence,
vol 1. Plymouth: Health Publications Ltd, 2005: 313–62.
55 © 2005 The Medicine Publishing Company Ltd