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Amoeboid Movement

Amoeboid movement is a crawling-like movement accomplished by protrusion of cytoplasm and formation of pseudopodia in eukaryotic cells. Several hypotheses have been proposed to explain the mechanism, including that it involves contractility of the cytoplasm and formation of actin and myosin filaments, as well as sol-gel conversion of the cytoplasm and regulation of focal adhesions.

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64 views

Amoeboid Movement

Amoeboid movement is a crawling-like movement accomplished by protrusion of cytoplasm and formation of pseudopodia in eukaryotic cells. Several hypotheses have been proposed to explain the mechanism, including that it involves contractility of the cytoplasm and formation of actin and myosin filaments, as well as sol-gel conversion of the cytoplasm and regulation of focal adhesions.

Uploaded by

Sukanta Majumder
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© © All Rights Reserved
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Amoeboid movement

Amoeboid movement is the most common mode of locomotion in Eukaryotic cells. It is a


crawling-like type of movement accomplished by protrusion of cytoplasm of
the cell involving the formation of pseudopodia. The cytoplasm slides and forms a
pseudopodium in front to move the cell forward. This type of movement has been linked to
changes in action potential; the exact mechanism is still unknown. This type of movement is
observed in amoeboids, slime molds and some protozoans as well as some cells in humans
such as leukocytes. Sarcomas, or cancers arising from connective tissue cells, are
particularly adept at amoeboid movement, thus leading to their high rate of metastasis.

Hypothesis on Amoeboid movement


There is now almost general agreement that amoeboid movement is based on contractility
and that the cytoplasm undergoes a contraction in one region of the cell. There is still
some disagreement regarding the region that contracts. The first successful attempt at
the preservation of motility in a preparation of demembranated cytoplasm was the
isolation of "naked cytoplasm" in quartz capillaries.
Other model systems developed subsequently have helped to elucidate the mechanisms of
cytoplasmic streaming and contractility in amoeba. For example, Simard-Duquesne and
Couillard showed that glycerinated amoebae could be prepared that would undergo limited
contraction on the addition of ATP and magnesium ions. Amoeba cytoplasm was shown by
them to possess a calcium-activated ATPase activity resembling the ATPase of vertebrate
striated muscle myosin.
Bulk contractility and unorganized streaming activity have been observed in preparations
of pooled cytoplasm from Amoeba proteus by Thompson and Wolpert. They also were the
first to fractionate pooled cytoplasm and to demonstrate the presence of thin filaments in
a motile fraction.
Pollard and Ito improved the fractionation procedure and demonstrated the presence of
both thick and thin filaments in a fraction capable of movement and viscosity changes on
the addition of ATP. Motility was depressed by chilling during isolation and observed only
in the presence of ATP and on warming to room temperature.
However, the manner of movement in this pooled cytoplasm was different from that in
intact amoebae. Instead of streaming into pseudopodia, the pooled cytoplasm underwent a
massive contraction in which its material shifted in one direction. The thick and thin
filaments found in motile extracts of cytoplasm correspond to those observed in electron
micrographs of fixed intact amoebae by Nachmias. The 70-A filaments have been
identified as actin by heavy meromyosin binding in Amoeba Proteus and in Chaos
carolinensis. ATPase activity has been found in amoebae.
While several hypotheses have been proposed to explain the mechanism of amoeboid
movement, the exact mechanism is still unknown.
Sol-Gel Mechanism of Amoeboid movement
Amoeba contains a central elongated fluid portion (plasmasol), a rigid layer surrounding
this (plasmagel), a thin elastic surface layer (plasmalemma), and a hyaline layer between
the plasmagel and the plasmalemma which is fluid at the tip of active pseudopods and in
certain other regions.

The plasmasol is an emulsion. It consists of a fluid in which various vacuoles and granuoles
are suspended. The plasmagel is probably alveolar in structure. It contains the same kinds
of substances as the plasmasol, but some of the fluid appears to be gelated so as to form
alveoli. The plasmalemma probably consists of interwoven protein fibers and a lipoid which
fills the interstices.

The plasmasol is probably hypertonic; the plasmagel and the plasmalemma are probably
semipermeable. This and other factors result in an excess inflow of water, stretching the
plasmagel and the plasmalemma. When a pseudopod is formed, the inner portion of the
plasmagel liquefies locally. This produces a local decrease in elastic strength resulting in
the formation of a protuberance, a pseudopod. As this is formed there is contraction at
the posterior end, resulting in forward flow of the plasmasol and extension of the
pseudopod.

If the pseudopod is attached, the plasmalemma, being attached to the substratum and to
the adjoining plasmagel, slides over the plasmagel above and remains stationary below,
rolling movement results. If it is free, the plasmalemma is stretched out with movement in
it equal on all sides. If the free pseudopods become attached to the substratum at the tip
after they are thus formed, walking movement results.

During locomotion of either type, the plasmasol continuously gelates at the tip of the
extending pseudopods forming plasmagel, and the plasmagel continuously solates at the
posterior end forming plasmasol.

Response is due largely to changes in the elastic strength of the plasmagel in the
adhesiveness of the plasmalemma and in turgidity.

Molecular mechanism of cell motion


Based on some mathematical models recent studies hypothesize a novel biological model
for collective biomechanical and molecular mechanism of cell motion. It is proposed that
microdomains signal organizes cytoskeleton and its interaction with substratum. As
microdomains trigger and maintain active polymerization of actin filaments, their
propagation generates a highly interlinked network of curved or linear filaments. It is also
proposed that microdomain interaction marks the formation of new focal adhesion sites at
the cell periphery. Myosin interactions with the actin network then generate membrane
retraction, retrograde flow, and contractile forces for forward motion. Finally, continuous
application of stress on the old focal adhesion sites could result in the calcium-induced
calpain activation, and consequently the detachment of focal adhesions which completes
the cycle.
Another such proposed mechanism, the 'bleb-driven amoeboid locomotion' mechanism,
proposed that the cell cortex actomyosin contracts to increase hydrostatic pressure
inside the cell. The increase hydrostatic pressure causes the cell cortex to be broken in
the direction of the desired flow. And the cytoplasmic sol-gel state is regulated.
Locomotion of amoeba occurs due to the sol-gel conversion of the cytoplasm within its cell.
The ectoplasm being called the plasma gel and the endoplasm the plasma sol. 'Sol-gel
conversion is the contraction and relaxation events which are enforced by osmotic
pressure and other ionic charges. For example, when an amoeba moves, it extends the
gelatinous cytosol pseudopium, which then results in the more fluid cytosol flowing after
the gelatinous portion where it filled in the end of the pseudopium. Inside the amoeba,
there are proteins that can be activated to convert the gel sol state.

Cytoplasm consists largely of actin and actin is regulated by actin binding proteins. Actin
binding proteins are in turn regulated by calcium ions, hence, calcium ions are very
important in the sol-gel conversion process.

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