Dr. S.

Sharma
Phylum Protozoa: Methods of Locomotion, Amoeboid,
Metabolic, swimming and Gliding movement
METHODS OF LOCOMOTION IN PROTOZOANS
Basically there are four known methods by which the protozoans move
1. Amoeboid movement
2. Swimming movement
3. Gliding movement
4. Metabolic movement

AMOEBOID MOVEMENT
This type of locomotion is also called as pseudopodial locomotion. Here
locomotion is brought about by the pseudopodia. It is the characteristic of
rhizopod protozoans like Amoeba proteus and Entamoeba histolytica. Also such
movement is exhibited by amoeboid cells, macrophages and phagocytic
leucocytes like monocytes and neutrophils of metazoans. Various theories
have been proposed to explain the amoeboid locomotion.
Name of theory Proposed by

Contraction theory Schultuz

Walking theory Dellinger

Rolling move ment theory Jenning

Surface tension theory Berthold

Adhesion theory Jenning

Fountain zone theory Allen

Folding and unfolding theory Goldacre and Lorch

Hydraulic pressure theory Renoldi and Jaun

Sol-gel theory Hyman

Dr. S.Sharma

Sol Gel theory

Sol Gel theory convincingly explains the mechanism involved in the
formation of pseudopodia. This theory, also known as Change in viscosity
theory was advocated by Hyman. Later Pantin and Mast explained this
theory. According to this theory, the cytoplasm of amoeba can be
distinguished into outer ectoplasm/Plasmagel and the inner
endoplasm/Plasmosol.
The plasmagel which forms the outer layer of the cytoplasm is thick, less
in quantity, non-granular, transparent and contractile. The plasmosol
which forms the inner layer of the cytoplasm is more in quantity, less
viscous, fluid like, more granular and opaque. Due to change in the
viscosity, the plasmagel and plasmosol inter -convert and consequently the
pseudopodia form and disappear causing the movement of Amoeba.
This inter-convertibility of plasmagel and plasmosol is physicochemical
change. Gel becomes sol by taking water and sol becomes gel by losing
water.
Amoeboid locomotion can be ex plained in the following steps:
Step 1: Initially Amoeba attaches itself to the solid substratum by the
plasma lemma at the temporary anterior end.
Step 2: Then the hyaline layer of the ectoplasm at the anterior end forms a
thickened hyaline cap. It is the first stage in the formation of the
pseudopodium.

Step 3: Behind the hyaline cap, a point of weakness in the elasticity of

plasmagel is formed. Hence the inner plasmosol flows forward, forming a
pseudopodium.
Dr. S.Sharma
Step 4: The plasmosol that flows outward behind the hyaline cap changes
its colloidal state from sol to gel and joins the ectoplasm.
Step 5: The outer region of the plasmosol, which is flowing forward
undergoes gelation and produces a rigid plasmagel tube. The gelation of
plasmosol extends the plasmagel tube forward.
Step 6: Two ends appear in Amoeba at this stage. The anterior end is
smooth with the rounded surface which the retractile end also called as
Uroid has a wrinkled surface.
Step 7: Around the region of the hyaline cap, an annular region of sol to
gel transformation is formed. It is called the zone of gelation. At the uroid
end a region where gel transforms into sol is called as zone of solation.
Step 8: Plasmagel at the uroid end changes into sol and flows forward
continuously through the gelatinized tube. As the plasmosol flows
forward, the pseudopodium elongates further and the body of amoeba
moves in that direction. The ectoplasm does not move but grows at the
leading tip and is broken down at the uroid end.
Step 9: The gelation at the advancing end and the solation at the trailing
end occur simultaneously and at the same rate thus making the forward
movement of amoeba continuous.
Step 10: The contraction of the plasmagel at the trailing end causes
hydraulic pressure on the sol and makes the plasmosol flow forward
continuously in the plasmagel tube.
Step 11: As the pseudopodium advances continuously in the direction of the
movement the body of amoeba also moves.

Basis of action of protein molecules: Sol gel theory

Amoeboid locomotion is brought about by the protein molecules (actin and
myosin) present in the cytoplasm. Goldacre and Lorsch explained the
phenomenon of gelation and solation based on the folding and unfolding
of these protein molecules. According to the m, the cytoplasm gelates
when the protein molecules unfold by losing water and the cytoplasm
solates when the protein molecules folds by absorbing water.
Hence, the proteins in the plasmosol are in folded state and the proteins
in the plasmagel are in the unfolded state. This folding and unfolding of
the protein molecules lead to the formation of the pseudopodia and thus
the amoeboid movement. The energy required for this process is made
available from the ATP.
Dr. S.Sharma

According to the foundation zone theory put forth by Allen, the plasmosol
flows forward due to the pulling force caused by the sliding action of the
actin molecules over the myosin molecules at the advancing end.
This interconvertibility of sol and gel is mainly due to the assembly and
disassembly o f actin filaments. Assembly results in gel formation and the
disassembly leads to the sol formation.
METABOLIC MOVEMENT
In protozoans a pellicle is present in the ectoplasm which is composed of
proteinaceous strips supported by dorsal and ventral microtubu les. In
many protozoans these protein strips can slide past one another, causing
wriggling motion. This wriggling motion is called as metaboly or metabolic
movement. This movement is mainly caused by the change in the shape of
the body.
This metabolic movement is observed in most of the sporozoans at certain
stages of life cycle. These kinds of movement are also referred to as
Gregarine movements as this movement is the characteristic of most of
the gregarines.

SWIMMING MOVEMENT
Swimming locomotion in protozoans is caused by the flagella and cilia.
Flagella bring about the movement of some parasites in the body fluids of
the hosts. As the movement in this case is caused by the beating flagella
and cilia are also known as undulipodia.
Dr. S.Sharma
Depending on the structure involved swimming movement can be of two
types namely,
* Flagellar movement
* Ciliary movement

Type 1 swimming movement: Flagellar movement

A flagellum pushes the fluid medium at right angles to the surface of its
attachment, by its bending movement. The bending movement of
flagellum is made by the sliding of microtubules past each other with the
help of dynein arms. The dynein arms show a com plex cycle of movement
with the energy provided by ATP. These dynein arms attaché to the outer
microtubule of an adjacent doublet and pull the neighboring doublet. As
the result the doublets slide past each other in opposite direction. The
arms release and attach a little farther on the adjacent doublet and again
pull the neighboring doublet.
The doublets of the flagellum are physically held in place by the radial
spokes and thus the doublets cannot slide past much and their sliding is
limited by the radial spokes. Instead the doublets can curve causing a
bend in the flagellum and this bending has an important role in the
flagellar movement. Flagellum shows the following movements,
Undulation moveme nt: Undulation from the base to the tip causes pushing
force and pushes the organism backwards. Similarly undulation from the
tip to the base causes pulling force and causes the organism to pull
forward. Also when the flagellum ends to one side and shows wave like
movement from base to tip the organism moves in lat erally in opposite
direction. Finally when the undulation is spiral, it causes rotation of the
organism in the opposite direction and this is called as gyration.
Dr. S.Sharma
Sidewise lash moveme nt: The flagellar movement of many organisms is a
paddle-like beat or sidewise lash consisting of strokes namely effective
stroke and recovery stroke.

Effective stroke-During effective stroke the flagellum becomes rigid and

starts bending against the water. This beating in water at right angles to
the longitudinal axis of the body causes the organism to move forward.
Recovery stroke- During recovery stroke, the flagellum becomes
comparatively soft and will be less resistant to the water. This helps the
flagellum move backwards and then to the original position.
Simple conical gyration moveme nt: In this kind of movement the flagellum
turns like a screw. This propelling action pulls the organism forward
through the water with a spiral rotation around the axis of movement and
gyration on its own.

Type 2 swimming movement: Ciliary movement

Just like the flagellum, the cilium also shows back and forth movements
during the locomotion. These back and forth movements of the cilia are
also called as effective and recovery strokes respectively. Cilium moves
just like a pendulum or a paddle. The cilium moves the water parallel to
the surface of its attachment like that of paddle stroke movement. The
movement of water is perpendicular to the longitudinal axis of cilium.
Effective stroke: During effective stroke, the cilium bends and beats against
water thus bringing the body forward and sending the water backwards.
Recovery stroke: During recovery stroke, the cilium comes back to original
position by its backward movement without any r esistance.
Dr. S.Sharma

Cilia shows two types of coordinated rhythms,

* Synchronous rhythm, where in the cilia beast simultaneously in a
transverse row.
* Metachronous rhythm, where in cilia beat one after another in a
longitudinal row. The metachronal waves pass from anterior to posterior
end.
The beating of the cilia can be reversed to move backwards when a
Paramoecium encounters any undesirable object in its path. The ciliary
movement is coordinated by infraciliary system though neuromotor center
called as motorium present near the cytopharynx in the ciliates like
Paramoecium. The infraciliary system together with motorium form
neuromotor system which helps in coordination of the beating of the cilia.
Ciliary movement is the fastest locomotion in prot ozoans.

GLIDING MOVEMENT
The zigzag movement in the protozoans brought about by the contraction
and relaxation of myonemes present below the pellicle in the ectoplasm is
called as the gliding movement. The movement by gliding is comparatively
small. Myonemes are the contractile fibrils which are similar to the
myofibrils. This kind of gliding movement is shown by flagellates,
Sporozoans, Cnidospora and some ciliates.
 Dr. S.Sharma

1. What are the basic methods by which the protozoans move?

2. Who proposed the Sol-Gel threory?

3. Enumerate the steps involved in the movement of Amoeba as per Sol -Gel theory.

4. Write about the gliding moveme nt of protozoans.

5. Discuss the swimming moveme nt of protozoans.

6. Explain the effective and recovery strokes in Ciliary movement.

Phylum Protozoa: Locomotary Organs (Pseudopodia,

Myonemes, Flagella and Cilia)
Protozoan movement in water
Protozoans in water are subjected to forces of water resistance like
pressure drag and viscous drag. Pressure drag is due to the difference of
pressure between two ends of the body. Viscous drag is due to the water
molecules attached to the surface of the body.
For protozoans which are small in size, viscous drag is of much
importance. These organisms are not streamlined to minimize the
pressure drag.

Locomotory Organs in Protozoa

Locomotion is the movement of the animals from place to place. It is
performed in search of food, mate, and shelter or to escape from
predators etc. it is influenced by external and internal stimuli.
Protozoans are very primitive, sin gle celled animals which show great
adaptability in their locomotion. They exhibit slowest locomotion like
amoeboid locomotion and also the fastest locomotion like ciliary
locomotion.
In protozoans, locomotion is brought about by

 Cellular extensions like Pse udopodia (Eg: Amoeba)

 Pellicular contractile structures like Myonemes (Eg: Euglena and Sporozoans)

 Locomotory orga nelles like Flagella (Eg: Paramecium) and Cilia (Eg: Euglena)
Dr. S.Sharma
PSEUDOPODIA (CELLULAR EXTENSION)
They are also known as false feet. These ar e the temporary outgrowths of
the cell. They are formed on the surface of the body by the movement of
the cytoplasm.

Depending on number of pseudopodia formed on the surface:

Polypodia- Several pseudopodia formed on the surface of the body.
Eg: Amoeba proteus
Monopodia- Only single pseudopodia is formed on the surface of the body.
Eg: Entamoeba histolytica

Depending on the structure of the pseudopodia:

Lobopodia: These are lobe like and blunt structures with broad and
rounded ends. These structures co mposed of endoplasm and ectoplasm.
Lobopodia move by pressure flow mechanism.
Eg: Amoeba proteus, Entamoeba histolytica

Filopodia: These are slender filamentous pseudopodia tapering from base

to tip. Sometimes these may be branched out but they are not f used to
form a network. They are composed of only ectoplasm.
Eg: Euglypha, Lecithium
Dr. S.Sharma

Reticulopodia: They are also known as rhizopodia or myxopodia. They are

filamentous, profusely interconnected and branched. They form a network.
The primary function of these pseudopodia in ingestion of food and the
secondary function is locomotion. They exhibit two way flow of the
cytoplasm. They are commonly found in foraminifers.
Eg: Elphidium, Globigerina

Axopodia: These are fine needle like, straight pseudopodia ra diating from
the surface of the body. Each Axopodia contain a central axial rod which is
covered by granular and adhesive cytoplasm. The main function of these
axopodia is food collection. Axopodia also exhibit two -way flow of
cytoplasm. Axopodia are mainl y found in Heliozoans and radiolarians.
Dr. S.Sharma
Eg: Actinosphaerium, Actinophrys, Collozoum

MYONEMES (PELLICULAR CONTRACTILE EXTENSIONS)

Many protozoans have contractile structures in the pellicle or ectoplasm
called as myonemes. These may be in the form of,
* Ridges or grooves (Eg: Euglena)
* Contractile myofibrils (Eg: Larger ciliates)
* Microtubules (Eg: Trypanosoma)
FLAGELLUM (LOCOMOTORY ORGANELLES)
Flagella are the locomotory organelles of flagellate mastigophoran
protozoans. They are mostly thread like pr ojection on the cell surface. A
typical flagellum consists of an elongated, stiff axial fiber called as axial
filament or axoneme enclosed by an outer sheath. The axoneme arises
from basal granule called as blepharoplast or kinetosome which is further
derived from Centrioles. Blepharoplast lies below the cell surface in the
ectoplasm. The region around blepharoplast is called microtubular
organizing center that controls the assembly of microtubules.
Dr. S.Sharma

When the axial filament is viewed under an electron micr oscope 9 + 2

arrangement can be observed. The 2 central longitudinal fibers are
enclosed by membranous inner sheath. The 2 central longitudinal fibers
are surrounded by 9 longitudinal peripheral doublets (each with
microtubules A and B) which form a cylind er between the inner and the
outer sheath. Each peripheral paired fiber is connected to the internal
membranous sheath by radial spokes.
Each peripheral doublet also has pairs of arms directed towards
neighboring doublet. These arms are made of the protein called as dynein.
The arms create the sliding force. The peripheral doublets are surrounded
by an outer membranous sheath called as protoplasmic sheath, which is
an extension of the plasma membrane. Some flagella also bear lateral
appendages called as flimmers or mastigonemes along the length of the
axoneme above the level of the pellicle.

Types of Flagella
Number and arrangement of flagella vary in Mastigophora from one to
eight or more. Free living species usually have one to eight flagella
Dr. S.Sharma
whereas the parasitic forms may have one to many flagella. Flagella are
classified based on the arrangement of lateral appendages and the nature
of the axial filament.
Stichonematic: Only one row of lateral appendages occurs on the axoneme
up to tip.
Eg: Euglena, Astasia
Pantonematic: Two or more rows of lateral appendages occur on the
axoneme
Eg: Peranema, Monas
Acronematic: Lateral appendages are absent and axoneme ends as a
terminal ‘naked’ axial filament
Eg: Chlamydomonas, Polytoma

Pantacronematic: Flagellum is provided with two or more rows of lateral

appendages and the axoneme ends in a terminal naked axial filament.
Eg: Urceolus
Anematic: In some cases the flagella is simple without any lateral
appendages and a terminal naked filament.
Eg: Chilomonas, Cryptomonas
Dr. S.Sharma
CILIA (LOCOMOTORY ORGANELLES)
Cilia are short hair like structures present all over the surface of the
body. They may be also confined to specific regions of the ciliate
protozoan. Cilia help in locomotion as well as in food collection .
Cilia greatly resemble the flagella in the basic structure. The major
difference between the flagella and the cilia is that cilia are smaller
compared to the flagella. Cilia arise from the kinetosome. Cilia consist of
an axial filament called as axoneme surrounded by the protoplasmic outer
sheath.
Electron microscopic studies of axoneme reveal 9 + 2 organization of the
peripheral doublet fibrils and central singlet fibrils. The details of the 9 +
2 organization and the presence of the dynein arms are simi lar to that of
the flagellum. All these fibrils are embedded in a matrix. The central
fibrils are enclosed within a delicate sheath.
The infraciliary system is located just beneath the pellicle. It consists of
kinetosomes at the bases of cilia, kinetodesmo s or kinetodesmal fibrils
that are connected to the kinetosomes and running along the right side of
each row of kinetosomes as cord of fibers known as kinetodesmata. A
longitudinal row of kinetosomes, kinetodesmal fibrils and their
kinetodesmata form a unit called kinety. All the kineties together form an
infraciliary system that lies in the ectoplasm. The infraciliary system is
connected to the motorium, a neuromotor center neat the cytopharynx
and forms the neuromotor system. This neuromotor system contro ls and
coordinates the movement of cilia.

Types of cilia
* In some primitive forms like holotrichs (Eg: Paramecium) cilia are present
all over the body
* In some forms like peritrichs (Eg: Vorticella) cilia are present only in the
peristomial region
* In Suctorians (Eg: Acineta) cilia are present in only in the young ones
which are later replaced by sucking tentacles in the adults

Differences between Flagella and cilia

Flagella Cilia

They may be one to four in number. Mor than four Generally cilia are more in number comp
flagella are present in mastigophoran parasites flagella. Cilia vary from 3,000 to 14,000 in num
 Dr. S.Sharma
Flagella Cilia

A flagellum is about 150 microns in length A cilium is about 5 to 10 microns in length

Flagella are commonly found at one end of the cell Cilia occur either all over the body surfac
specific regions of the cell

Flagella produce undular movement Cilia produce pendular movement

Flagella help in locomotion only Cilia help both in feeding and in locomotion

Flagella do not form compound orga nelles Cilia may form undulation me mbranes and
compound ciliary organelles

Compound ciliary organelles

* Cilia in compound ciliary organelles do not fuse, but their basal granules
are sufficiently close to introduce a sort of coupling.
* A group of cilia that forms a bundle is called as cirrus. An undulating
membrane is a row of adhering cilia forming long sheet.
* The smaller rows of adhering cilia form the membranelles

1. Write about the structure of flagellum.

2. Explain the structure and function of cilia.

3. Diffrentiate between flagella and cilia.

4. What are the different organelles which help locomotion in Protozoans?

5. Describe the process of moveme nt of protozoan in water.

6. Mention the types of pse udopodia with examples.

 Dr. S.Sharma

Cilia and Flagella- Definition, Structure, Functions and

Diagram
Table of Contents
 Cilia and Flagella Definition
 Structure of cilia and flagella
 Working of Cilia and Flagella
 Functions of Cilia
 Functions of Flagella

Cilia and Flagella Definition

 Cilia and Flagella are complex filamentous cytoplasmic structures
protruding through a cell wall.
 They are minute, especially differentiated appendices of the cell.
 Flagella (singular = flagellum) are long, hair-like structures that extend
from the plasma membrane and are used to move an entire cell.
 Cilia (singular = cilium) are short, hair-like structures that are used to move
entire cells (such as paramecia) or substances along the outer surface of
the cell (for example, the cilia of cells lining the Fallopian tubes that move
the ovum toward the uterus, or cilia lining the cells of the respiratory tract
that trap particulate matter and move it toward the nostrils).
 The terms cilium (meaning an eyelash) and flagellum (meaning a whip) are
often used arbitrarily.
 Generally, cilia are shorter than flagella (<10 μm compared to >40μm).
 Cilia are present on the surface of the cell in much greater numbers
(ciliated cells often have hundreds of cilia but flagellated cells usually have
a single flagellum).
 The real difference, however, lies in the nature of their movement. Cilia row
like oars. The movement is biphasic, consisting of an effective stroke in
which the cilium is held rigid and bends only at its base and a recovery
stroke in which the bend formed at the base passes out to the tip.
 Flagella wriggle like eels. They generate waves that pass along their length,
usually from base to tip at constant amplitude.
 Thus the movement of water by a flagellum is parallel to its axis while a
cilium moves water perpendicular to its axis and, hence, perpendicular to
the surface of the cell.
Dr. S.Sharma

Figure: Diagram of Cilia and Flagella

Structure of cilia and flagella
Despite their different pattern of beating, cilia and flagella are
indistinguishable structurally.
All cilia and flagella are built on a common fundamental plan:
1. A bundle of microtubules called the axoneme (1 to 2 nm in length and 0.2
μm in diameter) is surrounded by a membrane that is part of the plasma
membrane.
2. The axoneme is connected with the basal body which is an intracellular
granule lying in the cell cortex and which originates from the centrioles.
3. Each axoneme is filled with ciliary matrix, in which are embedded two
central singlet microtubules, each with the 13 protofilaments and nine
outer pairs of microtubules, called doublets. This recurring motif is known
as the 9 + 2 array.
4. Each doublet contains one complete microtubule, called the A sub fiber,
containing all the 13 protofilaments. Attached to each A sub fiber is a B
sub fiber with 10 protofilaments.
5. Subfibre A has two dynein arms which are oriented in a clockwise
direction. Doublets are linked together by nexin links.
6. Dynein is an ATPase that converts the energy released by ATP hydrolysis
into the mechanical work of ciliary and flagellar beating.
7. Each sub fiber A is also connected to the central microtubules by radial
spokes terminating in fork-like structures, called spoke knobs or heads.
This regular arrangement of microtubules and associated proteins with the
nine-way pattern is also seen in centrioles. But unlike centrioles, cilia and
Dr. S.Sharma
flagella have a central pair of microtubules, so that the overall structure is
called the 9 + 2 axoneme.
Note: Eukaryotic flagella diverge from prokaryotes in composition. Flagella in
eukaryotes contain far more proteins and bear some similarity to motile cilia,
with the same general motion and control patterns.
Working of Cilia and Flagella
In eukaryotes
Using ATP produced by mitochondria near the base of the cilium or flagellum
as fuel, the dynein arms push on the adjacent outer doublets, forcing a sliding
movement to occur between adjacent outer doublets. Because the arms are
activated in a strict sequence both around and along the axoneme and
because the amount of sliding is restricted by the radial spokes and inter-
doublet links, sliding is converted into bending.
In prokaryotes
Bacterial flagella use a fundamentally different mechanism. Like the propeller
of a boat, the motion of the bacterial flagellum is entirely driven by the rotary
motor at its base. The bacterial flagellum itself is a specialized piece of
extracellular cell wall, made of one protein (flagellin) that has no similarity to
tubulin or dynein. Cilia and flagella are full of cytosol all the way to their tips
and use the ATP in that cytosol to generate force all the way along their
length.
Functions of Cilia
 Cilia are used for locomotion in isolated cells, such as certain protozoans
(e.g., Paramecium).
 Motile cilia use their rhythmic undulation to sweep away substances, as in
clearing dirt, dust, micro-organisms and mucus, to prevent disease.
 Cilia play roles in the cell cycle as well as animal development, such as in
the heart.
 Cilia selectively allow certain proteins to function properly.
 Cilia also play a role in cellular communication and molecular trafficking.
 Non-motile cilia serve as sensory apparatus for cells, detecting signals.
They play crucial roles in sensory neurons. Non-motile cilia can be found in
the kidneys to sense urine flow, as well as in the eyes of the
photoreceptors of the retina.
 They also provide habitats or recruitment areas for symbiotic microbiomes
in animals.
 Cilia have also been discovered to participate in vesicular secretion of
ectosomes.
 Dr. S.Sharma

Functions of Flagella
 Flagella are generally used for locomotion of cells, such as the
spermatozoon and Euglena (protozoan).
 Flagella have an active role in aiding cell feeding and eukaryotic
reproduction.
 In prokaryotes such as bacteria, flagella serve as propulsion mechanisms;
they’re the chief way for bacteria to swim through fluids.
 It also provides a mechanism for pathogenic bacteria to aid in colonizing
hosts and therefore transmitting diseases.
 Flagella also function as bridges or scaffolds for adhesion to host tissue.

Differences Between Cilia and Flagella

Flagella are the complex filamentous cytoplasmic structure protruding
through cell wall. These are unbranched, long, thread like structures, mostly
composed of the protein flagellin, intricately embedded in the cell
envelope.
Cilia are slender, microscopic, hair-like structures or organelles that extend
from the surface of nearly all mammalian cells (multiple or single).

S.N. Characteristics Cilia Flagella

Cilia are short, hair like appendages Flagella are long, threadlike
1 Definition extending from the surface of a living appendages on the surface of a living
cell. cell.

2 Number Numerous Less in Number

3 Length Short and hair like organelle (5-10µ) Long wipe like organelle (150µ)

Presence at one end or two ends or

4 Occurrence Occurs throughout the cell surface.
all over the surface.

5 Cross section Nexin arm present. Nexin arm absent

 Dr. S.Sharma
6 Density Many (hundreds) per cell Few (less than 10) per cell

Cilia beat in a coordinated rhythm

7 Beating either simultaneously (synchronous) They beat independent of each other.
or one after the other (metachronic).

Rotational, like a motor, very fast Wave-like, undulating, sinusoidal,

8 Motion
moving slow movement compared to cilia

9 Found in Eukaryotic cells Eukaryotic and prokaryotic cells

Cilia use ‘kinesin’ which has an Flagella are powered by the proton-
10 Energy Production ATPase activity that produces energy motive force by the plasma
to perform the movement. membrane.

Helps in locomotion, feeding

11 Functions Help mainly in locomotion only.
circulation, aeration, etc.

12 Examples Cilia present in Paramecium Flagella present in Salmonella

Ciliary Locomotion in
Paramoecium | Protozoa
Introduction to Paramoecium:
Paramoecium is the fastest moving organism of all protozoans
found in fresh water ponds. It possesses cilia or compound ciliary
structures as locomotors and food-capturing organelles. This
organism is found in abundance in stagnant ponds and organic
infusions.

Orientation of Cilia in Paramoecium:

The body of Paramoecium is covered with a distinct tough and
flexible pellicle. The pellicle has inflated donut shaped alveoli which
produce a polygonal space. One or two cilia arise through this space.
In Paramoecium, the ciliature is divided into body ciliature, which
Dr. S.Sharma
occurs over the general body surface, and the oral ciliature, which is
associated with the mouth region. The latter includes larger cilia
(Fig. 1.15).

Structure of a Cilium:
The cilia are tiny hair-like appendages about 0-25 µm in diameter.
Cilia are shorter than the flagella and are generally more numerous
than the flagella. The ultrastructure of a cilium is identical to that of
a flagellum. Thus, each cilium is made up of an axial filament — the
axoneme, surrounded by a protoplasmic sheath. The cilium arises
from a basal body or kinetosome located in the alveolar layer.

The kinetosomes that form a particular longitudinal row, are

connected by means of fine striated fibres, called a kinetodesma.
Actually, a single kinetodesma in Paramoecium is a cable of still
smaller kinetodesmal fibrils, each of which originates from a
kinetosome (Fig. 2.7). The cilia, kinetosome and kinetodesma
together make up a kinety.

Role of Cilia in Locomotion:

Paramoecium exhibits the following types of movement by means of
cilia which are adapted for acting in fluid media.

i. Swimming:
Dr. S.Sharma
During swimming, each cilium moves in a whip-like motion. It first
gives a forward active stroke in which the cilium is fully extended
and beating against the surrounding liquid. It is followed by a
recovery stroke, in which the cilium returns to its original position
with an unrolling movement that minimizes the viscous drag (Fig.
2.9a).

The direction of the effective stroke is oblique to the long axis of the
body of Paramoecium. This causes the animal to swim in a spiral
course and at the same time to rotate on its longitudinal axis (Fig.
2.8).

In Paramoecium, cilia do not beat simultaneously rather groups of

cilia bend in a coordinated unidirectional waves. The movement of
adjacent cilia occurs as a result of interference effect of the
surrounding water layer. Thus, the hydrodynamic forces impose a
co-ordination on the cilia. This coordinated sequential activation of
cilia over the surface of the cell body is seen as a wave, called
metachronal waves (Fig. 2.9b).
Dr. S.Sharma

The ciliary beat can be reversed and the animal can move backward.
The backward movement is associated with the so-called avoiding
reaction. When a Paramoecium comes in contact with some
undesirable substances or objects, the ciliary beat is reversed.

The ciliate moves a short distance backward, turns slightly

clockwise or counterclockwise, and moves forward again (Fig. 2.10).
This avoiding reaction is repeated, till the existence of unfavorable
condition. In Paramoecium, energy necessary for ciliary beating is
controlled by changing the levels of Ca++ and K+ ions of cellular fluid.

ii. Creeping:
During creeping, Paramoecium uses its cilia of the oral surface as
miniature legs and simply glides over the obstacles. As the pellicle is
Dr. S.Sharma
thin and elastic, the ciliate can easily bend and squeeze through
gaps narrower than its own body diameter.

Flagellar Locomotion in Euglena |

Invertebrates
Introduction to Flagella in Euglena:
A common plan of organization in the non-muscular contractile
system of animals is found both in flagella and cilia. These struc-
tures with certain associated fibrillar systems, provide organelles of
movement not only for different protozoa, but also in many
metazoan animals where that function as an important effector
structure.

The effect of flagella upon the movement of a protozoa is best

exemplified by Euglena — an organism, 55-100 µm in length, found
swimming freely on the surface of fresh water bodies like pond,
canal, lake etc.

Structure of Flagellum in Euglena:

In general, flagellum is a long whip like organ which protrudes to
the exterior from the cell body and permits mechanical work
without any marked change in the form of the effector cell. In
Euglena, there are two flagella. One of them is equal in length to
body while other is short. They emerge out through the gullet — a
narrow depression at the exterior end of the spindle-shaped body.

The gullet leads to a flask-shaped non-contractile reservoir (Fig.

2.1). The flagellum bifurcates into two at the middle of the reservoir.
These two flagella originate from two compact basal granules or
blepharoplasts, situated in the cytoplasm just beneath the base of
the reservoir.

In most species of Euglena, the two flagella originate separately

from two blepharoplasts and the shorter one soon after its
emergence unites with the longer one (Fig. 2.1).
Dr. S.Sharma

Ultrastructure of Flagellar in Euglena:

Electron microscopy has shown that the long flagellum in
Euglena has two parts:
It is the inner core, composed of microtubules and other proteins.
Microtubules are normally long, hollow tubes formed of two types
of proteins viz., a tubulin and p tubulin.

In the axoneme, the microtubules are modified and arranged in a

ring of nine special doublets of microtubules surrounding a central
pair of single microtubule (Fig. 2.2). This “9 + 2” array is the
characteristic of axoneme of almost all forms of cilia and flagella.

This microtubules extend continuously throughout the length of

axoneme. In the centre, the pair of single microtubules are complete
microtubules, while in the outer ring, each doublet is composed of
one complete and one partial microtubules known as the A and B
Dr. S.Sharma
tubules respectively. Each doublet in the outer ring is provided with
sets of arms that join neighbouring doublets.

Each arm is composed of a protein called dynein. Pairs of inner and

outer arms are spaced all along each A tubule at regular 24 nm
intervals. The outer doublets are connected circumferentially by
another protein called nexin links at intervals of about 96 nm. A
series of radial spoke with a periodicity of 88 to 96 nm extends from
the A sub-tubule to the central pair of microtubules (Fig. 2.2).

All flagella arise from a basal body. When the basal bodies are
distributed to daughter cells during mitosis, they typically arrange
themselves at each pole of the mitotic spindle and are then
designated as centrioles. A region around the basal bodies and
centrioles, called the microtubule organizing centre, controls the
above „mentioned organized assembly of microtubules.

The ultrastructure of the basal bodies is like that of an axoneme

except that the central singlet are absent and the nine fibrils in the
outer circle are triplets, two of these being continuous with the dou-
Dr. S.Sharma
blets of the flagellum. Dynein arms however, are absent in the
triplets.

Role of Flagella in Locomotion:

In Euglena, the movement of flagella commonly involves the
generation of waves that are transmitted along it, either in a single
plane or in a corkscrew pattern. The waves arise at the base of the
flagellum, from the wall of the reservoir, apparently by two roots.
The waves then pass to tip of the main flagellum, which beats at a
rate of about 12 strokes per second and also shows a movement of
rotation.

This rotation causes the tip of the organism to rotate (Fig. 2.3),
while at the same time pushing it to one side (Fig. 2.4). Because of
this, Euglena rotates as it swims (at a rate of about 1 turn per
second) and it also follows a corkscrew course (Fig. 2.4).

The movement of its body is thus comparable with that of propeller,

for it sets up forces on the water that bring about forward
displacement. When an undulation moves along the flagellum, it
also generates lateral forces. These forces are usually symmetrical,
the left-directed forces cancel the right directed forces, and only the
longitudinal force remains to move the cell forward (Fig. 2.5a &
2.5b).
Dr. S.Sharma

The relationship of flagellar ultrastructure to movement has

received much attention in recent years and the sliding tubule
model is now widely accepted. According to this theory, the
movement of a flagellum is produced by the bending of the core or
axoneme. The bending force is produced due to active sliding of
adjacent outer doublets against each other.
Dr. S.Sharma
In the presence of ATP, the dynein arm on one doublet attaches to
the adjacent doublet and flexes, causing the doublets to slide past
each other by one increment. Successive attachments and flexes
cause the doublets to slide smoothly past one another over a
distance sufficient to bend the flagellum.

If a flagellum is severed from a cell by a laser beam, the isolated

structure continues to propagate bending movements in a normal
way, indicating that the motile machinery is contained in the
axoneme itself and its movement do not depend on a motor at its
base.

Sometimes, Euglena shows a very peculiar motion in which waves

of contraction pass along the body from anterior to posterior end
and the animal creeps forward. This contraction is brought about by
the stretching of protoplasm on the pellicle or by localised fibrils
called myonemes in the cytoplasm.

This type of locomotion is known as Euglenoid movement (Fig.

2.6A). An Euglena can also move by rowing. During rowing, the
flagellum is held rigid and is slightly arched in the direction of the
stroke. In recovering the position, it bends as it is drawn back so as
to face minimum resistance (Fig. 2.6B).
Dr. S.Sharma