278 BIOLOGY
You have learnt that all living cells have to be provided with nutrients, O
2
and other essential substances. Also, the waste or harmful substances
produced, have to be removed continuously for healthy functioning of
tissues. It is therefore, essential to have efficient mechanisms for the
movement of these substances to the cells and from the cells. Different
groups of animals have evolved different methods for this transport. Simple
organisms like sponges and coelenterates circulate water from their
surroundings through their body cavities to facilitate the cells to exchange
these substances. More complex organisms use special fluids within their
bodies to transport such materials. Blood is the most commonly used body
fluid by most of the higher organisms including humans for this purpose.
Another body fluid, lymph, also helps in the transport of certain substances.
In this chapter, you will learn about the composition and properties of
blood and lymph (tissue fluid) and the mechanism of circulation of blood
is also explained herein.
18.1 BLOOD
Blood is a special connective tissue consisting of a fluid matrix, plasma,
and formed elements.
18.1.1 Plasma
Plasma is a straw coloured, viscous fluid constituting nearly 55 per cent of
the blood. 90-92 per cent of plasma is water and proteins contribute 6-8
per cent of it. Fibrinogen, globulins and albumins are the major proteins.
B
ODY
F
LUIDS AND
C
IRCULATION
C
HAPTER
18
18.1 Blood
18.2 Lymph (Tissue
Fluid)
18.3 Circulatory
Pathways
18.4 Double
Circulation
18.5 Regulation of
Cardiac Activity
18.6 Disorders of
Circulatory
System
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Fibrinogens are needed for clotting or coagulation of blood. Globulins
primarly are involved in defense mechanisms of the body and the albumins
help in osmotic balance. Plasma also contains small amounts of minerals
like Na
+
, Ca
++
, Mg
++
, HCO
3
–
, Cl
–
, etc. Glucose, amino acids, lipids, etc., are
also present in the plasma as they are always in transit in the body. Factors
for coagulation or clotting of blood are also present in the plasma in an
inactive form. Plasma without the clotting factors is called serum.
18.1.2 Formed Elements
Erythrocytes, leucocytes and platelets are collectively called formed
elements (Figure 18.1) and they constitute nearly 45 per cent of the blood.
Erythrocytes or red blood cells (RBC) are the most abundant of all
the cells in blood. A healthy adult man has, on an average, 5 millions to
5.5 millions of RBCs mm
–3
of blood. RBCs are formed in the red bone
marrow in the adults. RBCs are devoid of nucleus in most of the mammals
and are biconcave in shape. They have a red coloured, iron containing
complex protein called haemoglobin, hence the colour and name of these
cells. A healthy individual has 12-16 gms of haemoglobin in every
100 ml of blood. These molecules play a significant role in transport of
respiratory gases. RBCs have an average life span of 120 days after which
they are destroyed in the spleen (graveyard of RBCs).
Leucocytes are also known as white blood cells (WBC) as they are
colourless due to the lack of haemoglobin. They are nucleated and are
relatively lesser in number which averages 6000-8000 mm
–3
of blood.
Leucocytes are generally short lived. We have two main categories of WBCs
– granulocytes and agranulocytes. Neutrophils, eosinophils and basophils
are different types of granulocytes, while lymphocytes and monocytes
are the agranulocytes. Neutrophils are the most abundant cells (60-65
per cent) of the total WBCs and basophils are the least (0.5-1 per cent)
among them. Neutrophils and monocytes (6-8 per cent) are phagocytic
cells which destroy foreign organisms entering the body. Basophils secrete
histamine, serotonin, heparin, etc., and are involved in inflammatory
reactions. Eosinophils (2-3 per cent) resist infections and are also
R B C
Platelets
Eosinophil
Basophil
Neutrophil
Monocyte
T lymphocyte
B lymphocyte
Figure 18.1 Diagrammatic representation of formed elements in blood
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280 BIOLOGY
associated with allergic reactions. Lymphocytes (20-25 per cent) are of
two major types – ‘B’ and ‘T’ forms. Both B and T lymphocytes are
responsible for immune responses of the body.
Platelets also called thrombocytes, are cell fragments produced from
megakaryocytes (special cells in the bone marrow). Blood normally
contains 1,500,00-3,500,00 platelets mm
–3
. Platelets can release a variety
of substances most of which are involved in the coagulation or clotting of
blood. A reduction in their number can lead to clotting disorders which
will lead to excessive loss of blood from the body.
18.1.3 Blood Groups
As you know, blood of human beings differ in certain aspects though it
appears to be similar. Various types of grouping of blood has been done.
Two such groupings – the ABO and Rh – are widely used all over the
world.
18.1.3.1 ABO grouping
ABO grouping is based on the presence or absence of two surface antigens
(chemicals that can induce immune response) on the RBCs namely A
and B. Similarly, the plasma of different individuals contain two natural
antibodies (proteins produced in response to antigens). The distribution
of antigens and antibodies in the four groups of blood, A, B, AB and O
are given in Table 18.1. You probably know that during blood transfusion,
any blood cannot be used; the blood of a donor has to be carefully matched
with the blood of a recipient before any blood transfusion to avoid severe
problems of clumping (destruction of RBC). The donor’s compatibility is
also shown in the Table 18.1.
Blood Group Antigens on Antibodies Donor’s Group
RBCs in Plasma
A A anti-B A, O
B B anti-A B, O
AB A, B nil AB, A, B, O
O nil anti-A, B O
TABLE 18.1 Blood Groups and Donor Compatibility
From the above mentioned table it is evident that group ‘O’ blood can
be donated to persons with any other blood group and hence ‘O’ group
individuals are called ‘universal donors’. Persons with ‘AB’ group can
accept blood from persons with AB as well as the other groups of blood.
Therefore, such persons are called ‘universal recipients’.
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18.1.3.2 Rh grouping
Another antigen, the Rh antigen similar to one present in Rhesus monkeys
(hence Rh), is also observed on the surface of RBCs of majority (nearly 80
per cent) of humans. Such individuals are called Rh positive (Rh+ve)
and those in whom this antigen is absent are called Rh negative (Rh-ve).
An Rh-ve person, if exposed to Rh+ve blood, will form specific antibodies
against the Rh antigens. Therefore, Rh group should also be matched
before transfusions. A special case of Rh incompatibility (mismatching)
has been observed between the Rh-ve blood of a pregnant mother with
Rh+ve blood of the foetus. Rh antigens of the foetus do not get exposed to
the Rh-ve blood of the mother in the first pregnancy as the two bloods are
well separated by the placenta. However, during the delivery of the first
child, there is a possibility of exposure of the maternal blood to small
amounts of the Rh+ve blood from the foetus. In such cases, the mother
starts preparing antibodies against Rh antigen in her blood. In case of
her subsequent pregnancies, the Rh antibodies from the mother (Rh-ve)
can leak into the blood of the foetus (Rh+ve) and destroy the foetal RBCs.
This could be fatal to the foetus or could cause severe anaemia and
jaundice to the baby. This condition is called erythroblastosis foetalis.
This can be avoided by administering anti-Rh antibodies to the mother
immediately after the delivery of the first child.
18.1.4 Coagulation of Blood
You know that when you cut your finger or hurt yourself, your wound
does not continue to bleed for a long time; usually the blood stops flowing
after sometime. Do you know why? Blood exhibits coagulation or clotting
in response to an injury or trauma. This is a mechanism to prevent
excessive loss of blood from the body. You would have observed a dark
reddish brown scum formed at the site of a cut or an injury over a period
of time. It is a clot or coagulam formed mainly of a network of threads
called fibrins in which dead and damaged formed elements of blood are
trapped. Fibrins are formed by the conversion of inactive fibrinogens in
the plasma by the enzyme thrombin. Thrombins, in turn are formed from
another inactive substance present in the plasma called prothrombin. An
enzyme complex, thrombokinase, is required for the above reaction. This
complex is formed by a series of linked enzymic reactions (cascade
process) involving a number of factors present in the plasma in an inactive
state. An injury or a trauma stimulates the platelets in the blood to release
certain factors which activate the mechanism of coagulation. Certain
factors released by the tissues at the site of injury also can initiate
coagulation. Calcium ions play a very important role in clotting.
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18.2 LYMPH (TISSUE FLUID)
As the blood passes through the capillaries in tissues, some water along
with many small water soluble substances move out into the spaces
between the cells of tissues leaving the larger proteins and most of the
formed elements in the blood vessels. This fluid released out is called the
interstitial fluid or tissue fluid. It has the same mineral distribution as
that in plasma. Exchange of nutrients, gases, etc., between the blood and
the cells always occur through this fluid. An elaborate network of vessels
called the lymphatic system collects this fluid and drains it back to the
major veins. The fluid present in the lymphatic system is called the lymph.
Lymph is a colourless fluid containing specialised lymphocytes which
are responsible for the immune responses of the body. Lymph is also an
important carrier for nutrients, hormones, etc. Fats are absorbed through
lymph in the lacteals present in the intestinal villi.
18.3 CIRCULATORY PATHWAYS
The circulatory patterns are of two types – open or closed. Open
circulatory system is present in arthropods and molluscs in which blood
pumped by the heart passes through large vessels into open spaces or
body cavities called sinuses. Annelids and chordates have a closed
circulatory system
in which the blood pumped by the heart is always
circulated through a closed network of blood vessels. This pattern is
considered to be more advantageous as the flow of fluid can be more
precisely regulated.
All vertebrates possess a muscular chambered heart. Fishes have a
2-chambered heart with an atrium and a ventricle. Amphibians and the
reptiles (except crocodiles) have a 3-chambered heart with two atria and a
single ventricle, whereas crocodiles, birds and mammals possess a
4-chambered heart with two atria and two ventricles. In fishes the heart
pumps out deoxygenated blood which is oxygenated by the gills and
supplied to the body parts from where deoxygenated blood is returned to
the heart (single circulation). In amphibians and reptiles, the left atrium
receives oxygenated blood from the gills/lungs/skin and the right atrium
gets the deoxygenated blood from other body parts. However, they get mixed
up in the single ventricle which pumps out mixed blood (incomplete double
circulation). In birds and mammals, oxygenated and deoxygenated blood
received by the left and right atria respectively passes on to the ventricles of
the same sides. The ventricles pump it out without any mixing up, i.e., two
separate circulatory pathways are present in these organisms, hence, these
animals have double circulation. Let us study the human circulatory
system.
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18.3.1 Human Circulatory System
Human circulatory system, also called the blood vascular system consists
of a muscular chambered heart, a network of closed branching blood
vessels and blood, the fluid which is circulated.
Heart, the mesodermally derived organ, is situated in the thoracic
cavity, in between the two lungs, slightly tilted to the left. It has the size of
a clenched fist. It is protected by a double walled membranous bag,
pericardium, enclosing the pericardial fluid. Our heart has four
chambers, two relatively small upper chambers called atria and two larger
lower chambers called ventricles. A thin, muscular wall called the inter-
atrial septum separates the right and the left atria, whereas a thick-walled,
the inter-ventricular septum, separates the left and the right ventricles
(Figure 18.2). The atrium and the ventricle of the same side are also
separated by a thick fibrous tissue called the atrio-ventricular septum.
However, each of these septa are provided with an opening through which
the two chambers of the same side are connected. The opening between
the right atrium and the right ventricle is guarded by a valve formed of
three muscular flaps or cusps, the tricuspid valve, whereas a bicuspid
or mitral valve guards the opening between the left atrium and the left
ventricle. The openings of the right and the left ventricles into the
Figure 18.2 Section of a human heart
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284 BIOLOGY
pulmonary artery and the aorta respectively are provided with the
semilunar valves. The valves in the heart allows the flow of blood only in
one direction, i.e., from the atria to the ventricles and from the ventricles
to the pulmonary artery or aorta. These valves prevent any backward
flow.
The entire heart is made of cardiac muscles. The walls of ventricles
are much thicker than that of the atria. A specialised cardiac musculature
called the nodal tissue is also distributed in the heart (Figure 18.2). A
patch of this tissue is present in the right upper corner of the right atrium
called the sino-atrial node (SAN). Another mass of this tissue is seen in
the lower left corner of the right atrium close to the atrio-ventricular septum
called the atrio-ventricular node (AVN). A bundle of nodal fibres, atrio-
ventricular bundle (AV bundle) continues from the AVN which passes
through the atrio-ventricular septa to emer
ge on the top of the inter-
ventricular septum and immediately divides into a right and left bundle.
These branches give rise to minute fibres throughout the ventricular
musculature of the respective sides and are called purkinje fibres. The
nodal musculature has the ability to generate action potentials without
any external stimuli, i.e., it is autoexcitable. However, the number of action
potentials that could be generated in a minute vary at different parts of
the nodal system. The SAN can generate the maximum number of action
potentials, i.e., 70-75 min
–1
,
and is responsible for initiating and
maintaining the rhythmic contractile activity of the heart. Therefore, it is
called the pacemaker. Our heart normally beats 70-75 times in a minute
(average 72 beats min
–1
).
18.3.2 Cardiac Cycle
How does the heart function? Let us take a look. To begin with, all the
four chambers of heart are in a relaxed state, i.e., they are in joint
diastole. As the tricuspid and bicuspid valves are open, blood from the
pulmonary veins and vena cava flows into the left and the right ventricle
respectively through the left and right atria. The semilunar valves are
closed at this stage. The SAN now generates an action potential which
stimulates both the atria to undergo a simultaneous contraction – the
atrial systole. This increases the flow of blood into the ventricles by about
30 per cent. The action potential is conducted to the ventricular side by
the AVN and AV bundle from where the bundle of His transmits it through
the entire ventricular musculature. This causes the ventricular muscles
to contract, (ventricular systole), the atria undergoes relaxation
(diastole), coinciding with the ventricular systole. Ventricular systole
increases the ventricular pressure causing the closure of tricuspid and
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bicuspid valves due to attempted backflow of blood into the atria. As
the ventricular pressure increases further, the semilunar valves guarding
the pulmonary artery (right side) and the aorta (left side) are forced open,
allowing the blood in the ventricles to flow through these vessels into
the circulatory pathways. The ventricles now relax (ventricular diastole)
and the ventricular pressure falls causing the closure of semilunar valves
which prevents the backflow of blood into the ventricles. As the
ventricular pressure declines further, the tricuspid and bicuspid valves
are pushed open by the pressure in the atria exerted by the blood which
was being emptied into them by the veins. The blood now once again
moves freely to the ventricles. The ventricles and atria are now again in
a relaxed (joint diastole) state, as earlier. Soon the SAN generates a new
action potential and the events described above are repeated in that
sequence and the process continues.
This sequential event in the heart which is cyclically repeated is called
the cardiac cycle and it consists of systole and diastole of both the atria
and ventricles. As mentioned earlier, the heart beats 72 times per minute,
i.e., that many cardiac cycles are performed per minute. From this it could
be deduced that the duration of a cardiac cycle is 0.8 seconds. During a
cardiac cycle, each ventricle pumps out approximately 70 mL of blood
which is called the stroke volume. The stroke volume multiplied by the
heart rate (no. of beats per min.) gives the cardiac output. Therefore, the
cardiac output can be defined as the volume of blood pumped out by each
ventricle per minute and averages 5000 mL or 5 litres in a healthy individual.
The body has the ability to alter the stroke volume as well as the heart rate
and thereby the cardiac output. For example, the cardiac output of an
athlete will be much higher than that of an ordinary man.
During each cardiac cycle two prominent sounds are produced which
can be easily heard through a stethoscope. The first heart sound (lub) is
associated with the closure of the tricuspid and bicuspid valves whereas
the second heart sound (dub) is associated with the closure of the
semilunar valves. These sounds are of clinical diagnostic significance.
18.3.3 Electrocardiograph (ECG)
You are probably familiar with this scene from a typical hospital television
show: A patient is hooked up to a monitoring machine that shows voltage
traces on a screen and makes the sound “... pip... pip... pip.....
peeeeeeeeeeeeeeeeeeeeee” as the patient goes into cardiac arrest. This type
of machine (electro-cardiograph) is used to obtain an electrocardiogram
(ECG). ECG is a graphical representation of the electrical activity of the
heart during a cardiac cycle. To obtain a standard ECG (as shown in the
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286 BIOLOGY
Figure 18.3), a patient is connected to the
machine with three electrical leads (one to each
wrist and to the left ankle) that continuously
monitor the heart activity. For a detailed
evaluation of the heart’s function, multiple
leads are attached to the chest region. Here,
we will talk only about a standard ECG.
Each peak in the ECG is identified with a
letter from P to T that corresponds to a specific
electrical activity of the heart.
The P-wave represents the electrical
excitation (or depolarisation) of the atria,
which leads to the contraction of both the atria.
The QRS complex represents the depolarisation of the ventricles,
which initiates the ventricular contraction. The contraction starts shortly
after Q and marks the beginning of the systole.
The T-wave represents the return of the ventricles from excited to normal
state (repolarisation). The end of the T-wave marks the end of systole.
Obviously, by counting the number of QRS complexes that occur in a
given time period, one can determine the heart beat rate of an individual.
Since the ECGs obtained from different individuals have roughly the same
shape for a given lead configuration, any deviation from this shape indicates
a possible abnormality or disease. Hence, it is of a great clinical significance.
18.4 DOUBLE CIRCULATION
The blood flows strictly by a fixed route through Blood Vessels—the
arteries and veins. Basically, each artery and vein consists of three layers:
an inner lining of squamous endothelium, the tunica intima, a middle
layer of smooth muscle and elastic fibres, the tunica media, and an
external layer of fibrous connective tissue with collagen fibres, the tunica
externa
. The tunica media is comparatively thin in the veins (Figure
18.4).
As mentioned earlier, the blood pumped by the right ventricle enters
the pulmonary artery, whereas the left ventricle pumps blood into the
aorta. The deoxygenated blood pumped into the pulmonary artery is
passed on to the lungs from where the oxygenated blood is carried by
the pulmonary veins into the left atrium. This pathway constitutes the
pulmonary circulation. The oxygenated blood entering the aorta is
carried by a network of arteries, arterioles and capillaries to the tissues
from where the deoxygenated blood is collected by a system of venules,
veins and vena cava and emptied into the right atrium. This is the
systemic circulation (Figure 18.4). The systemic circulation provides
nutrients, O
2
and other essential substances to the tissues and takes
CO
2
and other harmful substances away for elimination. A unique
vascular connection exists between the digestive tract and liver called
Figure 18.3 Diagrammatic presentation of a
standard ECG
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hepatic portal system. The hepatic portal vein carries blood from intestine
to the liver before it is delivered to the systemic circulation. A special
coronary system of blood vessels is present in our body exclusively for
the circulation of blood to and from the cardiac musculature.
18.5 REGULATION OF CARDIAC ACTIVITY
Normal activities of the heart are regulated intrinsically, i.e., auto regulated
by specialised muscles (nodal tissue), hence the heart is called myogenic.
A special neural centre in the medulla oblangata can moderate the cardiac
function through autonomic nervous system (ANS). Neural signals through
the sympathetic nerves (part of ANS) can increase the rate of heart beat,
the strength of ventricular contraction and thereby the cardiac output.
On the other hand, parasympathetic neural signals (another component
of ANS) decrease the rate of heart beat, speed of conduction of action
potential and thereby the cardiac output. Adrenal medullary hormones
can also increase the cardiac output.
18.6 DISORDERS OF CIRCULATORY
SYSTEM
High Blood Pressure (Hypertension): Hypertension is the term for blood
pressure that is higher than normal (120/80). In this measurement 120
mm Hg (millimetres of mercury pressure) is the systolic, or pumping,
pressure and 80 mm Hg is the diastolic, or resting, pressure. If repeated
checks of blood pressure of an individual is 140/90 (140 over 90) or
Figure 18.4 Schematic plan of blood circulation in human
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higher, it shows hypertension. High blood pressure leads to heart diseases
and also affects vital organs like brain and kidney.
Coronary Artery Disease (CAD): Coronary Artery Disease, often referred
to as atherosclerosis, affects the vessels that supply blood to the heart
muscle. It is caused by deposits of calcium, fat, cholesterol and fibrous
tissues, which makes the lumen of arteries narrower.
Angina: It is also called ‘angina pectoris’. A symptom of acute chest pain
appears when no enough oxygen is reaching the heart muscle. Angina
can occur in men and women of any age but it is more common among
the middle-aged and elderly. It occurs due to conditions that affect the
blood flow.
Heart Failure: Heart failure means the state of heart when it is not pumping
blood effectively enough to meet the needs of the body. It is sometimes
called congestive heart failure because congestion of the lungs is one of
the main symptoms of this disease. Heart failure is not the same as cardiac
arrest (when the heart stops beating) or a heart attack (when the heart
muscle is suddenly damaged by an inadequate blood supply).
SUMMARY
Vertebrates circulate blood, a fluid connective tissue, in their body, to transport essential
substances to the cells and to carry waste substances from there. Another fluid, lymph
(tissue fluid) is also used for the transport of certain substances.
Blood comprises of a fluid matrix, plasma and formed elements. Red blood cells (RBCs,
erythrocytes), white blood cells (WBCs, leucocytes) and platelets (thrombocytes) constitute
the formed elements. Blood of humans are grouped into A, B, AB and O systems based
on the presence or absence of two surface antigens, A, B on the RBCs. Another blood
grouping is also done based on the presence or absence of another antigen called Rhesus
factor (Rh) on the surface of RBCs. The spaces between cells in the tissues contain a fluid
derived from blood called tissue fluid. This fluid called lymph is almost similar to blood
except for the protein content and the formed elements.
All vertebrates and a few invertebrates have a closed circulatory system. Our circulatory
system consists of a muscular pumping organ, heart, a network of vessels and a fluid, blood.
Heart has two atria and two ventricles. Cardiac musculature is auto-excitable. Sino-atrial node
(SAN) generates the maximum number of action protentials per minute (70-75/min) and
therefore, it sets the pace of the activities of the heart. Hence it is called the Pacemaker. The
action potential causes the atria and then the ventricles to undergo contraction (systole) followed
by their relaxation (diastole). The systole forces the blood to move from the atria to the ventricles
and to the pulmonary artery and the aorta. The cardiac cycle is formed by sequential events in
the heart which is cyclically repeated and is called the cardiac cycle. A healthy person shows 72
such cycles per minute. About 70 mL of blood is pumped out by each ventricle during a
cardiac cycle and it is called the stroke or beat volume. Volume of blood pumped out by each
ventricle of heart per minute is called the cardiac output and it is equal to the product of stroke
volume and heart rate (approx 5 litres). The electrical activity of the heart can be recorded from
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the body surface by using electrocardiograph and the recording is called
electrocardiogram (ECG) which is of clinical importance.
We have a complete double circulation, i.e., two circulatory pathways, namely,
pulmonary and systemic are present. The pulmonary circulation starts by the
pumping of deoxygenated blood by the right ventricle which is carried to the lungs
where it is oxygenated and returned to the left atrium. The systemic circulation
starts with the pumping of oxygenated blood by the left ventricle to the aorta
which is carried to all the body tissues and the deoxygenated blood from there is
collected by the veins and returned to the right atrium. Though the heart is
autoexcitable, its functions can be moderated by neural and hormonal mechanisms.
EXERCISES
1. Name the components of the formed elements in the blood and mention one
major function of each of them.
2. What is the importance of plasma proteins?
3. Match Column I with Column II :
Column I Column II
(a) Eosinophils (i) Coagulation
(b) RBC (ii) Universal Recipient
(c) AB Group (iii) Resist Infections
(d) Platelets (iv) Contraction of Heart
(e) Systole (v) Gas transport
4. Why do we consider blood as a connective tissue?
5. What is the difference between lymph and blood?
6. What is meant by double circulation? What is its significance?
7. Write the differences between :
(a) Blood and Lymph
(b) Open and Closed system of circulation
(c) Systole and Diastole
(d) P-wave and T-wave
8. Describe the evolutionary change in the pattern of heart among the vertebrates.
9. Why do we call our heart myogenic?
10. Sino-atrial node is called the pacemaker of our heart. Why?
11. What is the significance of atrio-ventricular node and atrio-ventricular bundle
in the functioning of heart?
12. Define a cardiac cycle and the cardiac output.
13. Explain heart sounds.
14. Draw a standard ECG and explain the different segments in it.
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