MUSCULAR SYSTEM
Introduction
Locomotion is important to all organisms for various purposes like to find
food, to mate, to escape from predators, for survival purpose, etc. and
altogether, they influence in locomotive organ evolution process in animals.
Locomotive organs are different in nature from one species to other. Muscles
play important role in movement of the animals. In higher animals along with
muscles, bones also associate for the movement. Animal movements can
differentiate as walk, run, swim, fly, crawl and jump. Bacterial movement can
classify as flagellar, spirochaetal and gliding movement. Single cell organisms
also show crawling like movement often called amoeboidal movement as well
as they show cillary movement, flagellar movement. Nature of movement is
different in plants in which roots, branches, leaves move in response to
environmental abiotic factors but overall the plant is non-movable. Here we
discuss the different types of locomotion or movement and their mechanisms
briefly.
Definition
Locomotion can defined as an ability to move from one place to another.
The organs which helps for the locomotion is called as locomotive organs. Ex:
limbs, flagella, cilia, etc.
Locomotion in single cell organisms
Prokaryotic and eukaryotic single cell organism do movement for their
survival, growth and reproduction.
Bacterial movements
Bacteria are single cell prokaryotic organisms. They show three different
types of movements. They are
1. Flagellar movement
2. Spirochaetal movement and
3. Gliding movement.
Mechanism of these movements are partially understood.
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Flagellar movement
Bacteria can move in liquids with help of flagellum. Depends on number
of flagella, arrangement over cell surface, bacteria were named differently like
monotrichous, lophotrichous, amphitrichous and peritrichous. According to the
arrangement they are classified into different classes and those bacteria are
motile.
Figure 1: Structure of Flagellum
Flagellum (Figure 1) is a lash-like cell surface appendages and it consists
of protein flagellin. It generally divided into 3 portions and they are basal body,
hook and filament. Basal body areimplanted in cell wall. Basal body of the
flagellum has M (motor) ring, S(stator) ring and C ring. M and s ring together
bind with peptidoglycan layer of the bacteria whereas c ring present in the
cytoplasmic region. Basal body is connected with the filament by hook.
Flagellum can rotate either clockwise or anticlockwise. Further basal body has
around 40 proteins including ‘mot’ proteins and ‘fli’ proteins which are
essential in the active movement through proton pumping mechanism. Ex. for
flageller bacteria Vibrio, Spirillum, Salmonella,Klebsiella, etc.
Spirochetal movement:
Spirochetes are Class V bacteria. The movement of Spirochetes is called
as spirochetal movement. Spirochetes are long right handed helically shaped
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bacteria. Axial fibrils and periplasmic flagellum,present in the periplasmic
region of the bacteria,overlapped together and run one sub-polar end to other of
the Spirochetes (Figure 2). The mechanism of Spirochetal movement is not
clear, however the proposed anticlockwise rotation of the flagellum can lead the
bacterial movement. Classical example for this type movement is
Treponemapallidum.
Figure 2: Structure of Spirochetes
Gliding movement:
Group of gliding bacteria show non flagellum based gliding movement.
Fimbriae like appendages (pili), present at polar region, involve in the gliding
movement with the help of motor system A and S along chemosensory system
‘Frz’. Here coordinated movement of two different motor system, known as S
and A system, response for the motility of bacteria (Figure 3).But their mode of
action is unclear. Ex. Myxococcus, cyanobacterium, Oscillatoria.
Figure 3: Gliding movement of Treponema Pallidum
But other hypotheses like generation of contractile waves or surface
tension or pushing by secreted slime was also proposed as possible mechanisms
of gliding.
Eukaryotic single cell movement
Both single cell life forms as well as multicellular life forms are exist in
eukaryotes. According to the life form they adopt their own mode of locomotion.
Protozoan are single cell eukaryotes. They are following different type of
locomotion according their life form. Amoeba is a single cell protozoa. It can
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migrate with the help of pseudopodia and this movement is called as amoeboid
movement. Most of the time amoeba move to get the food. Various hypotheses
regarding the amoeba movement: front contraction hypothesis, tail contraction
hypothesis, ectoplasmic contraction theory, cytoplasmic streaming, sol-gel
theory, etc., Here we discuss the simple ameboidal locomotive sol-gel theory.
The simple mechanism of amoeba movement (Figure 4) as follows
1. Swelling of the plasmagel results pseudopodia in response of
semipermeable membrane turgidity.
2. Movement of plasmasol towards the pseudopodia with simultaneous
contraction of plasmagel
3. Gelation of plasmosol results enlargement of pseudopodia
4. Adhesion to the food particle.
Figure 4: Sol-Gel Ameoba Movement
Most of non-flagellated cells often follow the amoeboid movement. Ex.
Macrophages
Euglenoid movement
Euglena is another single cell eukaryotic organism. Euglena contains
flagella as well as pellicular structure. Euglena can move by waving the flagella.
Pellicular structure of the euglena also help them to move. Adjacent pellicular
strips move very quickly thus by changing own shape euglena can move
without the help of flagella.
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Paramesium (ciliary) movement
Paramesium is one more single cell eukaryotic organism which show cilia
movement. Dense hair like cilia is present on entire outer surface of the euglena.
In a coordinated manner, cilia move one after one all over the body result the
paramecium movement.
Movements in animal cells
Animals, insects, birds, reptiles and fishes are multicellular eukaryotes.
They migrate in three different modes. They are amoeboid movement, ciliary
movement and muscular movement.
Amoeboid movement: In animal cell migration is possible by cytoplasm
streaming which followed by the formation of pseudopodia. In animal cells,
microfilament involvement is observed in amoeboid movement.
Ciliary movements: Cilia is the short form of the flagella present in
ciliated epithelium. Coordinated movements of cilia helping those cells to
migrate. The function of cilia almost similar to flagellum.
Muscular movement: In higher organisms, muscles based movement is
very common. Coordinated function of muscles, skeletal and neural systems
results the muscle movement.
Locomotion in lower animals
In lower animals skeletal system is underdeveloped or absent. In such
condition they have to use muscles or specialized locomotive organs for the
locomotion. Ex. Earthworms, snails, snake, fishes, etc.
Locomotion in Earthworm:
Long tube like structured earthworm moves underground by means of
waves of muscular contraction. During muscular contraction body segments are
alternatively shorten and lengthen. A special type S-shaped setae ring present in
all segments of the earthwork except first, last and clitellum which help to
anchor the surrounding soil. Mucus lubrication secretion make easier of their
movement in soil.
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Locomotion in Snake:
Snakes also use muscles for their locomotion. Four types of locomotion
were identified in snakes. They are lateral undulation, rectilinear movement,
concertina movement and sidewinding. In common lateral undulation mode,
wave like muscle reflexion from head to tail results the locomotion.
Locomotion in Sea-anemone:
Bilateral sea-anemone moves by radial operation of radial pedal. Sea-
anemone creeps in the direction of plane of symmetry.
Locomotion in fishes
Fishes swim by using their specialized locomotive organ fins. While
moving the fins, the generated thrust help to move to the fish. With the help of
multiple fins fishes can control the direction of movement.
Locomotion in cockroach
Cockroaches have wings as well as legs. Primarily they use the legs for
the locomotion and as well as they can use the wings too for the locomotion.
Locomotion in Higher animals:
Large number of animals use limbs for their movement in which muscles
are attached endoskeleton or exoskeleton. In vertebrates endoskeleton muscle
system is present in which muscles are attached external portion of the bones
(Figure 5 (a)). But in arthropods exoskeleton muscle system is present in which
muscles are attached inside of the skeleton system (Figure 5 (b))
Figure 5: Exoskeleton and endoskeleton system
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In birds, limbs are developed into wings as well as legs. Such differentiation is
absent in vertebrates (Figure 6).
Figure 6: Modified limb systems
Most of the insects use limb system for their movement but they follow
different mode of locomotion according to their systems.
Movements in plants:
Plants also show movements like phototrophism, gravitrophism. In
response to light, plant grow towards the light. This phenomenon is called as
phototropism. Whereas signalling molecules have their own roles in this
process. Auxin is one them. It triggers shoot growth. Stems show positive
phototropism and roots show negative phototropism.
The growth in response to gravity is called as geotrophism. Roots show
positive geotrophism and stems show negative trophism. Here again auxin plays
an important role in geotrophism.
Plant movement against contact stimuli is called as thigmotropism.
Climbing plants such as vines are the examples for thigmotropism. When the
plant touches some surface, the cell which contact the surface secrete the auxin
to the untouched neighbour cell and induce them to grow long. This results
bending around the object.
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Muscle is one type of tissue among four different tissues. Muscle is a
specialised type of tissue and it originating from mesoderm. Muscles alone
contribute around 40 to 50 percent body weight of the human. Excitability,
contractility, elasticity and extensibility are the special characteristic nature of
the muscle cells.
Based on their location, muscles are classified into three categories:
1. Skeletal muscles,
2. Visceral muscles and
3. Cardiac muscles.
Skeletal Muscles: Muscles which are associated with skeletal
components are known as skeletal muscles. Microscopically their appearance is
look like striped. Hence they are also called as striated muscles. These muscle
activities are regulated voluntarily by the nervous system. The principle action
of skeletal muscle is locomotion and it can modulate body shape.
Visceral muscles: Visceral muscle present in inner walls of hollow
visceral organs of the body. Example includes, alimentary canal, reproductive
tract, etc. Unlike skeletal muscle, they are smooth in appearance. They are
involuntary in nature so nervous system cannot control voluntarily. Visceral
muscle involves in food transport in digestive tract and gamete transport
through the genital tract.
Cardiac muscles: Heart consists of cardiac muscles in which cells
assemble in a branching pattern. Like skeletal muscles, they are also striated.
Activity wise, they are involuntary in nature thus nervous system cannot control
the cardiac muscles directly. The different between these muscle forms is given
in Table 1.
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Table 1: Difference Between skeletal, visceral and cardiac muscle.
Skeletal Muscle Visceral Muscle Cardiac Muscle
Location Around bone all Inner walls of Heart
over the body alimentary canal,
reproductive tract
Morphological Striped Smooth Striped with cross
appearance linking
Nerve control Involuntary Voluntary Involuntary
Function Movement Transport Pumping
Contraction Short twitch and Concentric, Long tetranic
types long tetranic eccentric,
isometric and
isotonic
Elasticity Elastic in nature Elastic in nature Elastic in nature
Diameter 10 to 80 μM 1 to 5μM ~ 10μM
Structure of skeletal muscle:
Muscle cells are also termed as muscle fibres. (Figure 7) They are long,
cylindrical and multinucleated cells. Muscle fibres were organised in three
levels. They are epimysium, endomysium and perimysium. Muscles are covered
by thick and tough connective tissues called as epimysium. Epimysium
separates one muscle from another. Collagen fibres of the eipmysium are wavy
in appearance and it has connection with the perimysium. Each perimysium
covers 100 to 150 muscle fibres and forms fascicles. Interstitial space between
muscle fibres is around 1 μm which allow development of the tunnel in the
perimysium thus arteries, veins and nerves can pass through it. In perimysium,
collagen fibres are arranged as wavy forms with cross links which helps to
improve the strength and stability of the muscle fibres. Endomysium consists of
loose connective tissues and they additionally add up the strength of the muscle
fibres. Endomysium also connects with the perimysium for the stability.
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Figure 7: Organization of muscle fibre
Muscle fibres are lined by the plasma membrane (sarcolemma) enclosing
sarcoplasm. Muscle fibres contains multiple nucleus (syncytium) and the
sarcoplasm contains relatively higher amount of glycogen granules, myoglobin.
Moreover high amount of calcium stored in endoplasmic reticulum. Parallel
arrangement of myofilaments in sarcoplasm is one of the characteristic nature of
the muscle fibre. Two types of myofilaments, known as myofibril present in the
muscle fibres (Figure 8). Based on the thickness, myofibril termed as thin and thick
myofilament which is made up of actin and myosin filaments respectively. Actin
filament otherwise termed as ‘I’ band or isoelectric band whereas myosin filament
termed as ‘A’ band or anisotrophic band. Actin and myosin fibrils are arranged
alternatively and run across the muscle fibres longitudinally. Z – line and M – line,
which are fibres with elastic in nature present in between I band and A band
respectively. The portion in between to A band is termed as sarcomere which is the
functional unit of the muscles which is responsible for contraction.
Figure 8: A thin myofilament
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Actin: Actin is one of the major muscle proteins, which is a polymeric in
nature. G-actin is the monomer unit of the actin filament and it is globular in
nature. Its molecular weight is around 43 kda. Number of actin monomer units
polymerize and form multimeric fibrous F-actin. Two F-actins coil around each
other to form α-helice (Figure. 16.3). Thickness of F-actin filament is around 6-
7 nm and it consists of 14 G-actin molecules per turn. Association with
tropomyosin and troponin, F-actin forms I band of the myofibril. Troponin
masks the active binding sites for myosin on actin filaments.
Figure 9: Actin Fibre
Myosin: Myosin is another major muscle protein. It is a hexameric
protein. It consists of two heavy chains and four light chains. Molecular weight
of the heavy chains and light chains are around 200 kda and 15 to 27 kda
respectively. Heavy chain forms dimeric filaments by coiling α-helically each
other except N terminal (Figure 10). Unwound N terminal of myosin is globular
in nature and it serves the binding sites for other light chains as well as F-actin.
N terminal of the myosin has ATPase activity. Carboxy terminal of myosin
filament meet together at H zone whereas N terminals meet together at the
margins of A band.
Figure 10: Myosin fibre
Tropomyosin: Tropomyosin is a rod like fibrous molecule. Its molecular
weight is around 66 kda. It made up of two different α and β peptides and both
are α-helical in nature. Double stranded tropomyosin run parallel with thin
filaments in grooves of the F-actin strands.
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Troponin: Troponin is a globular protein and exit as three interconnected
protein system. They are (a) TpT (tropomyosin binding protein), (b) TpC
(calcium binding protein) and (c) TpI (actin and TpC binding protein). Troponin
lie on thin filaments with an interval of 38.5 nm, but the rest attached with F-
actin and tropomyosin.
Polymerization of desmin results intermediate filaments which is
predominantly present in Z line. Α-α-actinin is a homodimeric protein and often
anchors the ends of f-actin molecule to the Z line.
Neuromuscular junction in skeletal muscle:
Neuromuscular junction is a connective region of efferent nerve fibre of
nerve system and muscle fibre of muscle system through synapses. The entire
junction is covered by a cell, which known as Schwann cell, for insulation purpose.
An invaginated membrane appearance present in this junction is called as synaptic
gutter where axon neuronal end. Sub-neuronal clefts surround the axon terminal
and it increases the surface area for synaptic transmitters. (Figure. 11)
Figure 11: Neuromuscular Junction
Structure of Visceral muscle
Visceral muscle or smooth muscles are fusiform in shape (Figure 12).
They mostly present free in nature i.e., single fibre form. It has greater elasticity
than the muscle fibre. Contractility and extencity are also higher than muscle
fibre. Myosin, actin, troponin, tropomyosin, calponin, caldesporin are some of
the protein present in the smooth muscles.
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Figure 12: Structure of smooth muscle
Neuromuscular junction in smooth muscle:
Branches of autonomous nerve fibres innervate the smooth muscle in a
diffuse manner. The junctions are not connected directly with muscle, but they
secrete the neurotransmitters at external space of muscle fibre. Secreted
neurotransmitters diffuse through the muscular fibre membrane to transmit the
signal.
Structure of Cardiac muscle:
Cardiac muscles having many features as like skeletal muscle. Cardiac
muscle cells are separated by a dark area which called as intercalated discs
(Figure 13). They are striated and interconnected. Gap junctions and
desmosomes also present in the cardiac muscle. Interconnected cardiac cells
called as syncytium. Heart consists of two syncytium, known as atrial
syncytium and ventricular syncytium.
Figure 13: Structure of cardiac muscle
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In human well developed limb system exists for the locomotion. Together
muscle-skeletal-nerve system coordination results the locomotion. Here we
discuss the different types of muscle contraction and their mode of action.
Types of muscle contraction in human:
Three types of muscles present in human which we discussed in previous
lecture. Skeletal muscles are mainly involving in the locomotion by contraction.
Contraction of the skeletal muscle underlie in two different class. They are
Twits contraction and tetranic contraction. Voluntary muscles contractions are
differentiated as concentric, eccentric, isometric and isotonic contraction.
Cardiac muscle contraction can be include in tetranic contraction but molecular
level difference can be seen (Figure 14).
Twitch contraction: It is a very short timed muscle contraction. In
response to stimuli, muscle contract and relax but before reaching the peak
muscle start to relax.
Tetranic contraction: In comparison to twitch contraction, tetranic
contraction is a longer contraction. In response to stimuli, muscles contract,
stabilize and then relax in a sequence manner. Duration of the stable condition
vary depend on the strength of the stimuli.
Concentric contraction: In order to overcome the resistance with
generated energy, muscle contracts and shortens. This contraction can be seen
in bicep contraction, angle change in joints.
Eccentric contraction: Due to the lack of energy generation to overcome
the external response, muscle contract and lengthen.The response is slightly
delayed one when compare to the concentric contraction.
Isometric contraction: No variation can find in muscle length in
response to the external force. So there is no movement.
Isotonic contraction: The excessive force on the muscle initially
contracts and later increases the muscle length. The muscle length increase in
response to the external force.
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General mechanism
In response to signals action potential generated from the brain reach to
the muscle cells through the neurons. At the end of the axon terminal, it secretes
a small amount of neurotransmitter, acetylcholine. Acetylcholine diffuses from
the membrane receptors which present on the muscle membrane to transmit the
signal. Acetylcholine binding leads to the opening of sodium channels results
influx of sodium inside the cell.
Sodium influx generate an action potential in the muscle fibre and it
travels along the muscle cell membrane. This action potential depolarizes the
muscle membrane which results the release of calcium ions from sarcoplasmic
reticulum to the sarcoplasma. Calcium mediated attraction in between the actin
and myosin lead to the sliding alongside each other. This sliding movement
results the muscle contraction. After a fraction of second, calcium pump
actively transfer the calcium from sarcoplasm to sarcoplasmic reticulum. This
removal of calcium from the sarcoplasm results the muscle relaxation.
Figure 14: Skeletal muscle contraction
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Molecular mechanism:
Muscle contraction mechanism is described by sliding filament theory.
This theory described by Andrew F Huxle, Rolf Niedergerke, Hugh Huxley and
Jean Hanson in 1954. Muscle contraction is a cyclic repetitive process. In
which, actin filament slide over myosin and generate tension in the muscle.
Action potential from CNS reaches neuromuscular junction and release the
acetylcholine near to muscle fibre. Acetylcholine diffuse the synapse where it
binds to the nicotinic acetylcholine receptors. After binding, receptors get
activated on neuromuscular junction and it lead to the opening of the
sodium/potassium channel. It results in sodium influx and potassium outflow
from the cell. Sodium/potassium movement from the muscle cell generate an
action potential which leads to the depolarization of the inner muscle fibre.
Depolarization in the inner muscle fibre activates L-type voltage dependent
calcium channels (Ex. Dihydropyridinereceptors) in the sarcoplasmic reticulum.
This channels are closely resembles the calcium release channels such as
ryanodine receptors. Activation of voltage gated calcium channels presence in
the sarcoplasmic reticulum start to release the calcium.
The released calcium binds to the troponin C which present in the myosin
fibril and modulates the tropomyosin allosterically. In normal condition
tropomyosin do not allow the myosin to bind on thin fibres. But in presence of
calcium, troponin (troponin T), it undergoes the allosteric modification and
troponin loose the affinity over thin filament. It results in tropomyosin
migration from the thin filament. Immediately, myosin binds strongly with the
freed area of thin filament. This process is an energy coupled process, so
myosin utilizes the ATP energy by cleaving into ADP and inorganic phosphate.
Actin also involve in this process, here it helps to release the inorganic
phosphate from the myosin. The overall results in sarcomere shortening. In this
state, the distance in between the Z band shortens. The movement of myosin
was observed around 10 to 12 nm per power stroke. The availability of ATP
increases the power stroke cycles (energy generation by lysing the ATP).
Meanwhile, sarcoplasmic reticulum actively collect the calcium present
in the sarcoplasm. This results in the decrease in sarcoplasmic calcium level.
So, the calcium start to migrate from the thin filaments which results in
tropomysin remodelling to the native form. The native form of tropomyosin
replaces the myosin and bind with the thin filament (Figure 15).
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Figure 15: Molecular mechanism of Skeletal muscle contraction
In smooth muscles, troponin is replaced by p-light chain protein. In
sarcoplasm, calmodulin and myosin-light chain kinase are present. In excited or
action potential generated condition, sarcoplasmic calcium level increases
calmodulin which immediately bind with four calcium level. Ca++-calmodulin
complex immediately binds with the enzyme p-light chain kinase. P-light chain
kinase-calmodulin complex actively phosphorylate the p-light chain.
Phosphorylated p-light chain readily dismantle from the actin filament.
Immediately, myosin bind with actin after ATP lysis and results contraction of
actin filament. When ATP binds with myosin, it again disassemble from the
actin filament and p-light chain bind back to the actin to lead relaxed condition.
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