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Ganong's Review of Medical Physiology, 23rd Edition

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around the injury. The initial redness is due to capillary dilation,<br />

a direct response <strong>of</strong> the capillaries to pressure. The swelling<br />

(wheal) is local edema due to increased permeability <strong>of</strong> the capillaries<br />

and postcapillary venules, with consequent extravasation<br />

<strong>of</strong> fluid. The redness spreading out from the injury (flare)<br />

is due to arteriolar dilation. This three-part response—the red<br />

reaction, wheal, and flare—is called the triple response and is<br />

part <strong>of</strong> the normal reaction to injury (see Chapter 3). It persists<br />

after total sympathectomy. On the other hand, the flare is absent<br />

in locally anesthetized skin and in denervated skin after the sensory<br />

nerves have degenerated, but it is present immediately after<br />

nerve block or section above the site <strong>of</strong> the injury. This, plus<br />

other evidence, indicates that it is due to an axon reflex, a response<br />

in which impulses initiated in sensory nerves by the injury<br />

are relayed antidromically down other branches <strong>of</strong> the<br />

sensory nerve fibers (Figure 34–14). This is the one situation in<br />

the body in which there is substantial evidence for a physiologic<br />

effect due to antidromic conduction. The transmitter released at<br />

the central termination <strong>of</strong> the sensory C fiber neurons is substance<br />

P (see Chapter 7), and substance P and CGRP are present<br />

in all parts <strong>of</strong> the neurons. Both dilate arterioles and, in addition,<br />

substance P causes extravasation <strong>of</strong> fluid. Effective nonpeptide<br />

antagonists to substance P have now been developed,<br />

and they reduce the extravasation. Thus, it appears that these<br />

peptides produce the wheal.<br />

REACTIVE HYPEREMIA<br />

A response <strong>of</strong> the blood vessels that occurs in many organs but<br />

is visible in the skin is reactive hyperemia, an increase in the<br />

amount <strong>of</strong> blood in a region when its circulation is reestablished<br />

after a period <strong>of</strong> occlusion. When the blood supply to a<br />

limb is occluded, the cutaneous arterioles below the occlusion<br />

dilate. When the circulation is reestablished, blood flowing<br />

Spinal cord<br />

Orthodromic conduction<br />

Antidromic conduction<br />

Direction taken by impulses<br />

FIGURE 34–14 Axon reflex.<br />

Sensory neuron<br />

Endings<br />

in skin<br />

Endings<br />

near arteriole<br />

CHAPTER 34 Circulation Through Special Regions 581<br />

into the dilated vessels makes the skin become fiery red. O 2 in<br />

the atmosphere can diffuse a short distance through the skin,<br />

and reactive hyperemia is prevented if the circulation <strong>of</strong> the<br />

limb is occluded in an atmosphere <strong>of</strong> 100% O 2 . Therefore, the<br />

arteriolar dilation is apparently due to a local effect <strong>of</strong> hypoxia.<br />

GENERALIZED RESPONSES<br />

Noradrenergic nerve stimulation and circulating epinephrine<br />

and norepinephrine constrict cutaneous blood vessels. No<br />

known vasodilator nerve fibers extend to the cutaneous vessels,<br />

and thus vasodilation is brought about by a decrease in<br />

constrictor tone as well as the local production <strong>of</strong> vasodilator<br />

metabolites. Skin color and temperature also depend on the<br />

state <strong>of</strong> the capillaries and venules. A cold blue or gray skin is<br />

one in which the arterioles are constricted and the capillaries<br />

dilated; a warm red skin is one in which both are dilated.<br />

Because painful stimuli cause diffuse noradrenergic discharge,<br />

a painful injury causes generalized cutaneous vasoconstriction<br />

in addition to the local triple response. When the<br />

body temperature rises during exercise, the cutaneous blood<br />

vessels dilate in spite <strong>of</strong> continuing noradrenergic discharge in<br />

other parts <strong>of</strong> the body. Dilation <strong>of</strong> cutaneous vessels in<br />

response to a rise in hypothalamic temperature overcomes<br />

other reflex activity. Cold causes cutaneous vasoconstriction;<br />

however, with severe cold, superficial vasodilation may supervene.<br />

This vasodilation is the cause <strong>of</strong> the ruddy complexion<br />

seen on a cold day.<br />

Shock is more pr<strong>of</strong>ound in patients with elevated temperatures<br />

because <strong>of</strong> cutaneous vasodilation, and patients in shock<br />

should not be warmed to the point that their body temperature<br />

rises. This is sometimes a problem because well-meaning<br />

laymen have read in first-aid books that “injured patients<br />

should be kept warm,” and they pile blankets on accident victims<br />

who are in shock.<br />

PLACENTAL & FETAL<br />

CIRCULATION<br />

UTERINE CIRCULATION<br />

The blood flow <strong>of</strong> the uterus parallels the metabolic activity <strong>of</strong><br />

the myometrium and endometrium and undergoes cyclic<br />

fluctuations that correlate with the menstrual cycle in nonpregnant<br />

women. The function <strong>of</strong> the spiral and basilar arteries<br />

<strong>of</strong> the endometrium in menstruation is discussed in<br />

Chapter 25. During pregnancy, blood flow increases rapidly as<br />

the uterus increases in size (Figure 34–15). Vasodilator metabolites<br />

are undoubtedly produced in the uterus, as they are in<br />

other active tissues. In early pregnancy, the arteriovenous O 2<br />

difference across the uterus is small, and it has been suggested<br />

that estrogens act on the blood vessels to increase uterine<br />

blood flow in excess <strong>of</strong> tissue O 2 needs. However, even though<br />

uterine blood flow increases 20-fold during pregnancy, the

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