ANIMAL PHYSIOLOGY

SUBMITTED BY: QIRRAT QAISER

SUBMITTED TO: MAM IFRA
ROLL.NO: 12060814-010
SEMESTER: SUMMER -2014

MSc ZOOLOGY

RESPIRATORY CENTER AND CONTROL OF

RESPIRATION BY MEDULLARY AND PERIPHERAL
CHEMORECEPTORS
The nervous system normally adjusts the rate of alveolar ventilation almost exactly to the
demands of our body so that the oxygen pressure and the carbon dioxide pressure in the arterial
blood are hardly altered, even during heavy exercise and most other types of respiratory stress.
The following topic describes the function of respiratory center for regulation of respiration.

RESPIRATORY CENTER
The respiratory center is composed of several groups of neurons located bilaterally in the
medulla oblongata (lowermost part of brainstem) and pons of the brain stem.
It is divided into three major collections of neurons:

Inspiratory center (Dorsal respiratory group)

Location: Dorsal portion of medulla

Nucleus: nucleus tractus solitarius (NTS)

Function: causes inspiration while stimulated.

Expiratory center (Ventral respiratory group)

Location: Antero- lateral part of medulla, about 5 mm anterior and lateral to dorsal
respiratory group.

Function: It generally causes expiration but can cause either expiration or inspiration
depending upon which neuron in the group is stimulated. It sends inhibitory impulse to the
Apneustic center.

Pneumotaxic centre

Location: Pons(upper part )

Function: It controls both rate and pattern of breathing. Limit inspiration.

Apneustic center

Location: Pons(lower part)

Functions:
a. It discharges stimulatory impulse to the inspiratory centre causing inspiration.
b. It receives inhibitory impulse from pneumotaxic centre and from stretch receptor of lung.
c. It discharges inhibitory impulse to expiratory centre.

DORSAL RESPIRATORY GROUP OF NEURONS

The dorsal respiratory group of neurons plays the most fundamental role in the control of
respiration and it extends most of the length of the medulla. Most of its neurons are located

within the nucleus of tractus solitarius. The NTS is the sensory termination of both the vagal and
the glossopharyngeal nerves, which transmit sensory into the respiratory center from
1. Peripheral chemoreceptors
2. Baroreceptors
3. Several types of receptors in the lungs
RHYTHMICAL INSPIRATORY DISCHARGES FROM DORSAL RESPIRATORY
GROUP
The basic rhythm of respiration is generated mainly in the dorsal respiratory group. Even when
all the peripheral nerves entering the medulla have been sectioned and the brainstem transected
both above and below the medulla, this group of neurons still emits repetitive burst of inspiratory
neuronal action potential. In primitive animals, neural network have been found in which activity
of one set of neurons excites a second set, which in turn inhibits the first. Then, after a period of
time, the mechanism repeats itself, continuing throughout the life of animal. Therefore, most
respiratory physiologists believe that some similar networks of neurons is present in the human
being, located entirely within the medulla; it probably involves not only the dorsal respiratory
group but adjacent areas of the medulla as well, and it is responsible for the basic rhythm of
respiration.
INSPIRATORY RAMP SIGNAL:
The nervous signal that is transmitted to the inspiratory muscles, mainly the diaphragm is not as
instantaneous burst of action potential. Instead, it begins weakly and increases steadily on a ramp
manner for about 2 seconds in normal respiration. Then it ceases abruptly for approximately the
next 3 seconds, which turns off the excitation of the diaphragm and allows elastic recoil of the
lungs and the chest wall to cause expiration. Next, the inspiratory signals begin again for another
cycle; this cycle repeats again and again, with expiratory occurring in between. Thus, the
inspiratory signal is a ramp signal. The obvious advantage of the ramp is that it causes a steady
increase in the volume of the lungs during inspiration.
There are two qualities of the inspiratory ramp that are controlled as follows:

Control of the limiting point at which the ramp suddenly ceases. This is the usual method
for controlling the rate of respiration; the earlier ramp ceases, the shorter the duration of

the inspiration. This also shortens the duration of expiration.

Control of the rate of increase of ramp signal so that during heavy respiration, the ramp
increases rapidly and therefore fills the lungs rapidly.

A PNEUMOTAXIC CENTER LIMITS THE DURATION OF INSPIRATION AND

INCREASES THE RESPIRATORY RATE
A pneumotaxic center, located dorsally in the nucleus parabrachialis of the upper pons,
transmit signals to the inspiratory area. The primary effect of this center is to control the
switch off point of the inspiratory ramp, thus controlling the duration of filling phase of the
lung cycle. When the pneumotaxic signal is strong, inspiration lasts for as little as 0.5 second,
thus filling the lungs only slightly. When the pneumotaxic signal is weak, inspiration might
continue for 5 or more seconds, thus filling the lungs with great excess of air.
The function of the pneumotaxic center is primarily to limit inspiration. This has a secondary
effect of increasing the rate of breathing because limitation of inspiration also shortens
expiration, and the entire period of each respiration.
A strong pneumotaxic signal can increase the rate of breathing to 30 to 40 breaths per minute,
whereas the weak pneumotaxic signal may reduce the rate to 3 to 5 breaths per minute.

VENTRAL RESPIRATORY GROUP OF NEURONS

Located in each side of the medulla, about 5 millimeter anterior and lateral to the dorsal
respiratory group of neurons, is the ventral respiratory group of neurons, which is found in
the nucleus ambiguous rostrally and the nucleus retroambiguus caudally. This group of
neurons is different from dorsal group of neurons in these ways:

The neurons of ventral respiratory group remain almost totally inactive during normal
quiet respiration. So thats why normal quiet breathing is caused by inspiratory

signals from dorsal respiratory group

The ventral respiratory neurons do not appear to participate in the basic rhythmical
oscillations that control respiration.

Electrical stimulation of few of the neurons in the ventral group causes inspiration,
whereas stimulation of other causes expiration. Therefore these neurons contribute to
both inspiration and expiration. They are especially important in providing the
powerful expiratory signal to the abdominal muscles during very heavy expiration.

Mechanism of respiratory center

CHEMICAL CONTROL OF RESPIRATION

The ultimate goal of respiration is to maintain proper concentrations of oxygen, carbon dioxide and
hydrogen ions in the tissues. Excess carbon dioxide and excess hydrogen ions in the blood mainly act
directly on the respiratory center, causing greatly increased strength of both inspiratory and

expiratory motor signals to the respiratory muscles.

In contrast, oxygen does not have significant direct effect on the respiratory center of the
brain in controlling respiration.it acts almost entirely on peripheral chemoreceptors located in
the carotid and the aortic bodies, and these in turn transmit nervous signals to the respiratory
center.
Two types of specialized nerve endings are of importance. These are given below
1. Chemosensitive receptors in the Medulla Oblongata
2. Chemosensitive receptors in the carotid and aortic bodies; these are called peripheral
chemoreceptor
.

Medullary Chemosensitive Receptors

Central chemoreceptors of the central nervous system, located on the ventrolateral
medullary surface in the vicinity of the exit of the 9th and 10th cranial nerves, are
sensitive to the pH of their environment. This area is therefore quite distinct from the
respiratory center. These act to detect the changes in pH of nearby cerebral spinal fluid
(CSF) that are indicative of altered oxygen or carbon dioxide concentrations available to
brain tissues. An increase in carbon dioxide tension of the arteries, often resulting from
increased CO2 intake, indirectly causes the blood to become more acidic; the cerebral

spinal fluid pH is closely comparable to plasma, as carbon dioxide easily diffuses across
the blood/brain barrier.

When a pledged soaked in cerebrospinal fluid saturated with CO 2 or soaked in an acidic

solution is applied to this area, an immediate response showing hypernea (increased
respiration) results. Application of drugs like acetylcholine and nicotine to this area also
results in hyperpnea. On the other hand, an application of procaine, a local anesthetic
agent, or cold which depress nerve activity of this area results in a stop of breathing or
apnea. These methods have been used to localize the Chemosensitive area of the brain
stem.

The primary stimulus for chemosensitive receptor is a fall in the pH of extracellular fluid
which lies close to the surface of the brain. The extracellular fluid in this region resembles
the cerebrospinal fluid with which it freely communicates in respect to HCO3- and CO2. The
PCO2 of the arterial blood serves to regulate the pH of the extracellular fluid in this area. A
rise in the arterial blood PCO2 leads to a greater CO2 diffusion into the cerebrospinal fluid
and the extracellular fluid thus reducing their pH leading to an increased ventilation of the
lungs or hyperpnea. The fall in arterial blood PCO2 has the reverse effect, i.e. there is a rise
in the pH causing a decrease in pulmonary ventilation.
Experiments on human beings have shown that breathing 4% CO2 increases pulmonary
ventilation to twice the normal value. Breathing 10% of CO2 increases pulmonary ventilation
to 10 times the normal value. The maximum effect on pulmonary ventilation is produced by
breathing about 20% of CO2. Any further increase in CO2 results in lowering of pulmonary
ventilation. At 10% CO2, dyspnea, headache, faintness, and restlessness are produced. At
15% CO2 there result rigidity of muscles, tremors and convulsions and the subject may lose
consciousness. Breathing 20% CO2 results in surgical anesthesia.

Peripheral Chemoreceptors

In addition to control of respiratory activity by respiratory center itself, still other mechanism
is available for controlling respiration. This is the peripheral chemoreceptors system. Special
nervous chemical receptors called chemoreceptors are located in several areas outside the
brain. They are especially important for detecting changes in oxygen in the blood, although
they also respond to a lesser extent to changes in carbon dioxide and hydrogen ion
concentrations. The chemoreceptors transmit nervous signals to the respiratory center in the
brain to help regulate respiration activity.
These lie in the carotid and aortic bodies. The carotid body is a small pinkish nodule located
in the bifurcation of the common carotid artery. It is the most vascular tissues in the body
receiving every minute 20 ml of blood per gram of tissue. The blood on flowing through it
comes in contact with nerve endings sensitive to chemical changes in the blood. The afferent
nerve fibers travel in the carotid branch of the glossopharyngeal nerve which also carries
afferents from the stretch receptors present in the carotid sinus.
The aortic bodies are groups of cells resembling cells of the carotid bodies; these are situated
in the aortic arch. The afferents from these bodies travel via the aortic nerves.

The most effective stimulus to peripheral chemoreceptor is a fall in arterial PO2

(hypoxemia). A rise in PCO2 and fall in pH of the arterial blood can also stimulate these
chemoreceptors but to a much lesser extent than hypoxemia. The stimulation of peripheral
chemoreceptors results in a reflex hyperpnea. If the peripheral chemoreceptors are enervated
or their afferent never fibers blocked by a local anesthetic agent, then the reflex hyperpnea is

not obtained. The carotid body has 7 times more chemoreceptor activity than the aortic
bodies.

Decreased arterial oxygen stimulates the chemoreceptors:

When the oxygen concentration in the arterial blood falls below normal, the chemoreceptors
become strongly stimulated. The impulse rate is particularly sensitive to changes in arterial
oxygen pressure in the range of 60 to 30 mm Hg, a range in which hemoglobin saturation
with oxygen decreases rapidly.

Increased carbon dioxide and hydrogen ion concentration stimulates the

chemoreceptors:
An increase in either carbon dioxide concentration or hydrogen ion concentration also excites
the chemoreceptors and in this way, indirectly increases respiratory activity. However, the
direct effects of both of these factors in the respiratory centers itself are much more powerful
than their effects mediated through the receptors. The stimulation by way of the peripheral
chemoreceptors occurs as much as five times as rapidly as central stimulation. So the
peripheral chemoreceptors might be especially important in increasing the rapidity of
response to carbon dioxide at the onset of exercise.
In addition to the above mentioned chemical changes in the arterial blood, i.e. fall in PO2 and
pH; and rise in PCO2, some other changes in blood also stimulate chemoreceptor activity.
These are the following.

Cyanide: This acts by inhibiting cytochrome oxidase which produces hypoxia within

chemoreceptor tissue by a non-utilization of O2.

Nicotine: It stimulates sympathetic ganglion cells.
Acetylcholine, serotonin and sulfides
Raised temperature of blood.
Decreased blood flow through carotid and aortic bodies.

Effects of stimulation of Peripheral Chemoreceptors

An increase in pulmonary ventilation.

Vasoconstriction in limb vessels.

Rise in blood pressure.
Increased pulmonary vascular resistance.
Increased secretion of hormones of adrenal medulla.

FLOW CHART OF CONTROL OF RSPIRATION BY MEDULLARY AND

PERIPHERAL CHEMORECEPTORS