Locomotion in Euglena

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LOCOMOTION IN Euglena
1) INTRODUCTION:

Euglena is a single-celled organism. Currently, over


1,000 species of Euglena have been described.
Euglena is able to move through aquatic
environments by using a large flagellum for
locomotion. Flagellum is mainly of 2 types –
prokaryotic (which is found in- E. coli, Salmonella
and Helicobacter pylori etc.) and eukaryotic like the
one found in Euglena, Trypanosoma, Giardia etc.
Here the structure of eukaryotic flagellum and its Diagram of Euglena showing flagellum
role in locomotion in Euglena is described in details.

2) DEFINITION OF FLAGELLUM:
These are permanent thread-like projections of cytoplasm on the cell surface. They are similar in structure
to cilia but few and long.

3) STRUCTURE (E/M): Composed of two parts: a) Axoneme and b) Basal body / granule.

a) AXONEME: The elongated shaft and motile part of flagellum shows the following features:

1) Size: a few µm – 2mm in length and about 0.5 µm in diameter.

2) Structure:
i) Surrounded by an outer sheath, which is continuous with the plasmamembrane.
ii) Shows the fundamental 9+2 longitudinal microtubular (fiber) pettern.
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iii) The 9 are peripheral and paired – each with sub-fibers A & B.
iv) Subfiber A is smaller but complete (with 13 tubulin subunits) and lies close to the central axis.
v) Subfiber B is larger and incomplete and has only 11 subunits.
vi) Subfiber A has two rows (outer and inner) of
processes – the so called “Dynein arms” – that are
oriented in same direction in all fibers, clockwise
when viewed from base to tip.
vii) Arms contain dynein, an Mg2+ and Ca2+
activated enzyme (300 – 400 kD, mol. wt.).
viii) Nexin links: The doublets are linked by
interdoublet or nexin links – that comprise Nexin
Protein (150 – 160 kD, mol.wt.).
ix) 2 central fibers are singlets (unpaired) and
connected by a bridge.
x) Central sheath – that encloses the two central
fibers.
xi) Radial spokes – are the bridges or links between
subfiber – A and central sheath. The spokes
terminate in a dense knob or head.

3) All the components of the axoneme are


embedded within the flagellar matrix.

4) Axoneme ends at the base in Basal granule (Kinetosome / Blepharoplast).

b) BASAL BODY (GRANULE) OR KINETOSOME:


i) Cylindrical structure (0.2
µm x 0.5 µm) with both
ends open.
ii) Separated from axoneme
by basal plate, located at its
distal end.
iii) With 9 peripheral and no
central fibers (i.e; 9 + 0
pattern).
iv) Peripheral fibers are
triplet and arranged from
center to periphery and
designated as subfibers – A,
B, & C.
v) Subfibers A & B traverse
the basal plate and are
continuous with the
corresponding tubules in the
flagellum, while Subfiber –
C terminates near the plate.
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REMARK:
Kinetosomes are considered to be either derived from or developed into the centrioles (as they are
structurally quite homologous).

4) MOLECULAR MECHANISM OF FLAGELLAR MOVEMENT:

i) Each of the outer 9 doublet microtubules extends a pair of dynein arms (an "inner" and an "outer" arm) to
the adjacent microtubule.

ii) These dynein arms are responsible for flagellar beating, as the force produced by the arms causes the
microtubule doublets to slide against each other and the flagellum as a whole to bend.

iii) These dynein arms produce force through ATP hydrolysis in mitochondria that are contained in
blepharoplast.

iv) The flagellar axoneme also contains radial spokes, polypeptide complexes extending from each of the
outer 9 microtubule doublets towards the central pair, with the "head" of the spoke facing inwards.

v) The radial spoke is thought to be involved in the regulation of flagellar motion, although its exact
function and method of action are not yet understood.

5) THEORIES REGARDING FLAGELLAR MOVEMENT:


Mechanism of flagellar movement is not precisely clear. Following theories are usually put forwarded to
explain such type of movement.

i) BÜTSCHLI’S SCREW THEORY:


Bütschli advocated that Euglena undergoing a series of lateral movement exerts a pressure on water at right
angel to its body surface. The pressure resolves into 2 forces - one directed at parallel and the other at right
angle to the body axis. The former force drives the organism forward, while the later tends the
organism to rotate on its own axis like a screw (that follows spiral turning). This creates
Forward movement

propelling action pulling the animal forward.

ii) PADDLE STROKE THEORY OF


ULEHLA AND KRIJSMAN (1925):
They advocated the common movement of
flagellum is a sideways lash, directed
obliquely backward. The lash comprises an
effective down stroke, followed by a relaxed
recovery stroke to bring the flagellum
forward again. The beating of flagellum
drives water backward and propels the
animal forward. As the flagellum is
obliquely directed backward, the animal
rotates on its own axis during forward
movement.

(a) Down stroke (b) Recovery stroke


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iii) METZNER’S THEORY:


He advocated flagellum beats in a circle tracing a cone and generates
sufficient current to pull the animal forward.

iv) UNDULATING MOTION:


Wave-like undulation in flagellum leads movement of animal in
various directions:

a) Backward movement – When undulations pass from base to tip of


flagellum.

b) Forward movement - When undulation proceeds from tip to base of


flagellum.

c) Lateral movement – By base to tip undulation on a flagellum


positioned at about right angle to body axis.

d) Spiral undulations of flagellum cause the animal to rotate in Backward movement


opposite direction.

6) CONCLUSION:
The presence of mastigonemes on the flagella of certain organisms is known
to induce reversal of the direction of fluid motion. Although E. viridis
possesses a large number of mastigonemes on its flagellum, it moves in the
direction to be expected if the flagellum were smooth. The absence of
reversed mobility can be explained in several ways:
(a) The mastigonemes may be flexible and may lie almost parallel to
the flagellar membrane.
(6) The mastigonemes may be of insufficient length to provide a Lateral movement
force great enough to overcome that of the flagellum itself.
(c) The mastigonemes may lie in planes containing the axis of the
helix formed by the flagellum.
(d) The mastigonemes may be wrapped around the flagellum during
movement as suggested by Pitelka & Schooley (1955).

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