Arterioles 5
increases it. According to Poiseuille’s equations, resistance constrictor substances, which act on the adjacent
to flow is proportional to the fourth power of the radius.
This means that a very small change in lumen radius will
have a significant impact of resistance and thus flow.
Control over arteriolar diameter is achieved by a num-
ber of tightly regulated mechanisms, which will be dis-
cussed in detail below, and has three important functions:
• Coordinated constriction or dilatation of a large pro-
portion of arterioles will alter the total peripheral
resistance and thus arterial blood pressure.
• Constriction or dilatation of arterioles in single
organs allows the distribution of the cardiac output
to different organs to be regulated. Dilatation of the
arterioles in a particular organ will decrease resis-
tance and increase flow to that organ.
• Constriction or dilatation of arterioles influences the
hydrostatic forces in the capillaries and as a result
has an effect on fluid filtration (explained below).
Constriction of arterioles increases resistance and
causes a greater pressure drop, reducing the pressure
in the distal vessels, including the capillaries.
Control of vascular smooth muscle
(VSM) tone
Tight control over the arteriolar vascular smooth muscle
tone is achieved via numerous mechanisms. Many of
the same factors also have an effect on the vascular
smooth muscle of the venous capacitance vessels. These
mechanisms can be broadly divided into local influ-
ences and systemic influences. The mechanisms of
VSM contraction and relaxation have been discussed
in Chapter 2 but in short, it is important to remember
that vasoconstriction is an active process brought about
by the contraction of vascular smooth muscle, which as
for cardiac muscle requires an increase in cytosolic
[Ca2þ]. Vasodilatation occurs when VSM relaxes and
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the pressure of blood within the vessel causes it to
distend.
Local influences
Myogenic tone
This describes the constrictor response of vascular
smooth muscle to stretching of the vessel wall as a result
of increased internal pressure. It plays an important role
in autoregulation and also contributes to basal tone.
The higher the internal hydrostatic pressure, the greater
the degree of myogenic constriction.
Endothelium-derived substances
Endothelial cells play an important role in the local reg-
ulation of blood flow. Although they do not have a con-
tractile function, they release a number of dilator and
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smooth muscle cells and influence arteriolar tone.
The importance of the balance between dilator and
constrictor influences on resting VSM can be
demonstrated by the administration of a nitric oxide
(NO) synthase inhibitor. This causes a 30% increase in
blood pressure by increasing total peripheral resistance.
This increase is a result of loss of basal release of NO.
Endothelium-derived dilator factors
Endothelium-derived dilating factors include nitric ox-
ide (NO), prostacyclin (PGI2) and endothelium derived
hyperpolarizing factor (EDHF). NO is the most impor-
tant dilator substance. In the endothelium, NO is pro-
duced by endothelial nitric oxide synthase (eNOS).
eNOS is activated by an increase in intracellular
[Ca2þ] which can occur in response to factors such as
bradykinin, acetylcholine and substance P. It is also ac-
tivated by the shear forces that are exerted on the endo-
thelial cells by the blood flowing through the vessel.
Following its production by eNOS, NO diffuses to the
adjacent vascular smooth muscle cells and activates
the enzyme guanylyl cyclase increasing levels of cGMP
and bringing about dilatation (see Fig. 2.56). In addi-
tion to its vasodilator properties, NO has other effects
including inhibition of platelet function and inhibition
of leukocyte adhesion to the endothelium. Endothelial
derived dilator substances, particularly NO, provide a
tonic vasodilator effect that counteracts the tonic myo-
genic vasoconstriction and it is the balance of these that
determines the resting tone of the vessel.
Endothelium-derived constrictor factors
Endothelium-derived constricting factors include end-
othelin 1, thromboxane A2 and prostaglandin H2.
Endothelin 1 is the most important constrictor released
by the endothelium. It is released in response to vasocon-
strictor substances including adrenaline/epinephrine, va-
sopressin and angiotensin II. It then diffuses to the
adjacent smooth muscle cells and causes constriction via
G-protein coupled mechanisms similar to those activated
when noradrenaline/norepinephrine binds to a adreno-
ceptors (see Fig. 2.54).
Metabolic factors
Locally produced products of metabolism diffuse into
the tunica media of the arterioles and cause relaxation
of VSM and thus vasodilatation. The amount of these
factors released correlates with the local metabolic rate
and as a result this process couples blood flow with the
57