Muscle

Muscle physiology
by Guesh M.
3/21/16
The Physiology of Muscles

Introduction
1. Introduction
1.1. muscles
a. Can be excited chemically, electrically and mechanically.
b. Contractile mechanisms (actin + myosin) that can be activated by AP.
1.2. Mass
a. 45-50% of the total body mass.
b. 40% skeletal muscles + 10% cardiac and smooth muscles (4550%).
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Introduction
1.3. O2 consumption
a. 25% total bodily O2 consumption at rest is consumed by the
muscles.
b. During strenuous exercise this amount can increase as much as
10-20 times.
2. Types/Classification
2.1. Anatomical
2.1.1. Striations:
Presence of alternating light and dark bands on the sarcolemma.
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Introduction
Classification
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Introduction
2.1.1.1. Skeletal muscle

i. Have well developed cross striations (interdigitating
thick and thin filaments).
ii. Voluntary muscle tissue.
iii. Cells are long and multinucleated.
iv. Contract only in response to stimuli (no syncytial
bridges between cells).
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Introduction
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Introduction
2.1.1.2. Cardiac muscle

i. Have cross striation (banding pattern of thick and thin
filaments).
ii. Involuntary muscle tissue.
iii. Cells are branched and mononucleated.
iv. Have intercalated disc with gap junctions.
2.1.2. Non-striations/Smooth Muscle
Alternating dark and light bands are absent.
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Introduction
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Introduction
2.1.2.1. Single unit smooth muscle (Visceral smooth muscle)

i. Are large sheets of mononucleated small cells.
ii. Have low resistance bridge of gap junctions.
iii. Show synchronous excitation and contractions.
(= functional syncytium)
iv. Have unstable resting membrane potential.
v. Found in gut, ureter, small blood vessels and uterus.
2.1.2.2. Multiunit smooth muscles

i. Found in iris, lungs, hair roots and large arteries.
ii. Have no gap junctions but each cell receive ANS nerve terminal.
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Introduction
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10

Introduction
2.2. Physiological
2.2.1. Voluntary Muscle
Skeletal muscle (CNS, somatic neurons).
2.2.2. Involuntary muscle

Cardiac muscle (Intrinsic + Extrinsic factors, ANS + Hormonal)
Smooth muscle (Intrinsic + Extrinsic factors, ANS + Hormonal)
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11

Skeletal Muscle
3. Skeletal Muscle
Interactions between the body and the external environment
(maintenance of posture and movement, speech, respiration).
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12

Skeletal Muscles
3.2. Structural arrangement and contractile unit

Muscle
epimysium
Fasciculus (20 muscle fibers)
perimysium
Muscle fiber (=10-100 m, L=30cm, multinucleated)
endomysium (Sarcolemma, sarcoplasm/myoplasm
Sarcoplasm reticulum
Myofibril (= 1-2m, longitudinal, Sarcomere,
75% muscle vol., Z-line, -actinin, desmin)
Myofilaments
Thick filaments
1500 molecules, myosin
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Thin filaments
3000 molecules, actin
13
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14
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15
Geometrical orientation of the contractile elements

Skeletal Muscles
16
Geometrical orientation of the contractile elements

Skeletal Muscles
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17

Skeletal Muscles
3.3. Functional unit (Sarcomere)

a. It is the distance between two z-lines.
b. Responsible for the striated appearance.
3.3.1. Dimensions
a. The resting length of a sarcomere is 2m-2.2m.
b. At this length it generates the greatest force of contraction.
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18

Skeletal Muscles
3.3.2. Molecular geometry

Differences in Refractive indices viewed under polarized light.
A-Band (A= Anisotropisch)*
a. The darker area in the center of the sarcomere.
b. It is due to the orderly arrangement of thick filaments.
c. Thin filaments may extend into the A-band.
H-Band (H = Hensens disc, ?*?)
a. It contains only myosin tails (no myosin heads/no crossbridges)
b. There are no thin filaments.
c. When the muscle is relaxed
* Germanic words The Physiology of Muscles
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19

Skeletal Muscles
M-line (M= Mittelmembran) )*

a. Site of the reversal polarity of the myosin molecules in each of
the thick filaments.
b. It vertically bisects the H-Band
c. It contains 2 important proteins:
Myomesin: a structural protein that links neighboring thick filaments
Creatinine Phosphokinase: an enzyme that maintains adequate
ATP conc. in working muscle fibers.
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20

Skeletal Muscles
I-Band (I= Isotropisch) )*

a. The lighter area on either side of the z-lines.
b. Each sarcomere contain half of the two I-bands.
c. Thin filaments
Z-Line/Disc (Z = Zwischenscheibe) )*
a. Dense line in the center of each light band.
b. Separates one sarcomere from the next.
c. It is the attachment site for the thin filaments.
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21

NB
Skeletal Muscles
1. During muscular contraction:

a. There is NO CHANGE in length of either the thick or the
thin
filaments.
b. H-zone disappears and Z-line gets considerably darker.
c. There is shortening of the sarcomere
( I-band and H zone, and A-band remains )
2. a. When a muscle increases its length in the N o of
sarcomeres
(NOT the length of each sarcomere)
b. The length of thick & thin filaments of sarcomeres is
identical
in a neonate & in the adult muscle.
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22

Skeletal Muscles
3.4. Molecular Basis of Contraction

3.4.1. Sliding-Filament Model (Hanson & Huxley, 1955)
This model theorizes that sliding of the filaments (thick &
thin) toward the center of muscle and sarcomere is responsible
for the shortening and force of contraction.
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23

Skeletal Muscles
3.4.2 Thick Filament

a. Is called myosin (an actin-binding protein).
b. Dimensions: = 11-18 nm, L = 1.6 m
c. Composition of Myosin:
d. 1 thick filament has 300 (500) myosin heads.
1 cross-bridge has 2 identical heads
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24

Skeletal Muscles
e. Orientation:
i. Head projects out on either side of the H-zone to swivel in
opposite directions to shorten the sarcomere.
ii. During rapid contraction each head form 5 cycles/sec, thus
sliding
myosin & actin filaments by about 5m/sec.
iii. In a resting muscle the cross-bridge is not attached to the thin
filaments & is oriented perpendicularly to the myosin
filaments.
f. The thick filaments are suspended or assumed their central
position
in the sarcomere
their attachments
to the z-disc by
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The because
Physiology of of
Muscles
25

Skeletal Muscles
Geometrical orientation of myosin and actin
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26

Skeletal Muscles
Geometrical orientation of myosin and actin
27

Skeletal Muscles
3.4.3. Thin Filaments

a. Are called actins.
b. Dimensions : = 5nm, L = 1 m
c. Components :
1. Globular proteins
i. Actin (300-400 molecules, and molecular weight of 45kd)
Globular -Actin (G-actin) or
Fibrillar- Actin (F-actin, a polymer of G-actins)
ii. Troponin (50 molecules)
2. Non-Globular proteins: tropomyosin (40-60 molecules, 70kD)
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28

Skeletal Muscles
3.4.4 Regulatory proteins

A. Tropomyosin
a. A rod-shaped molecule stretched along each strand of thin filament.
b. 1 tropomyosin molecule covers seven actin monomers in an actin filament.
c. It blocks the binding sites of myosin on actin.
B. Troponins
~ Small globular units located at intervals along the tropomyosin molecules.
a. Troponin T: it binds other troponin components to tropomyosin (37kD).
b. Troponin I: inhibits the interaction of myosin with actin (24kD)
c. Troponin C: it has the binding site for Ca2+ that initiates contraction (18kD)
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29
Actins, Troponins and Tropomyosin Filaments

Skeletal Muscles
30

Skeletal Muscles
3.5. Molecular mechanisms of regulating muscle contraction

3.5.1. Regulatory Role of Troponin-Tropomyosin System
a. Resting state: Troponin I and Tropomyosin covers the sites where
myosin heads bind to actin.
(At rest: interaction of thick and thin filaments is inhibited).
Troponin-Tropomyosin complex is called Relaxing-Protein
because it inhibits the interaction between myosin and
actin.
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Skeletal Muscles
b. Contractile state:
The invading action potential to T-Tubule Ca2+ released from
SR binds to troponin C binding of troponin I to actin is
weakened tropomyosin moves laterally uncovers binding
sites for myosin heads contraction (in the presence of ATP).
Seven myosin-binding sites are uncovered for each molecule of
troponin that binds a Ca2+.
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32

Skeletal Muscles
3.5.2. Regulatory action of calcium

2 tubular networks (Sarcotubular system) that are involved with Ca2+.
3.5.2.1. Transverse Tubule (T-tube)

a. It is an invagination of the surface of the muscle membrane
(sarcolemma).
b. It is found at the junction of A-band and I-band (at z-line in frog).
c. One end of the tube is open to the extracellular space, but its
other end is closed.
d. Function: Rapid transmission of the action potential from the
cell membrane to all the fibers on the muscle.
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33

Skeletal Muscles
3.5.2.2. Sarcoplasm Reticulum (SR)

a. It is an internal tubular structure that runs between the myofibrils.
b. It is closed at both ends.
c. It is not continuous with the sarcolemma.
d. Functions:
i. Stores Ca2+ in the terminal cisternae
(lateral sacs, 1calsequestrin = 43 Ca2+ )
ii. Uptake and release of Ca2+

Calsequestrin is sarcoplasmic protein that binds Calcium.
Triad: 2 lateral sacs + 1 transverse tubule
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Skeletal Muscles
3.5.3. Regulation of ATP

a. Actin + myosin + ATP + Ca2+ CONRACTION
b. Actin + myosin + ATP - Ca2+ RELAXATION
c. ATP is needed for relaxation
d. In the absence of Ca2+ ATP is not hydrolyzed.
e. 3 ATP molecules are needed:
i. For the formation of the actinmyosin complex
ii. For the initiation of relaxation.
iii. To pump out Ca2+ from the sarcoplasm to sequester it
into the SR.
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35

Skeletal Muscles: ECC
The Motor Unit

A motor neuron and the muscle fibers it innervates.
Elementary unit of motor control (Charles Sherrington, 1925).
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36

Motor Unit
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37

Skeletal Muscles: motor unit
Motor Unit and NMJ
38

Motor Units
39

3.6.1. Excitation-Contraction Coupling/

Electromechanical Coupling
Def. : is the process of linking Em/AP to muscle contraction.
Electrical events precedes mechanical events (2ms, 100ms)
Twitch
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40

Neuromuscular Junction
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41
Neuromuscular Junction

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42

Events at the
neuromuscular
junction that
lead to an
action potential
in the muscle
fiber plasma
membrane.
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43

3.6.2. Events at the neuromuscular junction:

3.6.2.1. Presynaptic end (-motor neuron)
AP in presynaptic -motor neuron terminals
Depolarization of plasma membrane of the presynaptic -motor neuron axon terminals
[ 30mV]
Opening of Ca2+ channels at the active zones Ca2+ permeability and

entry of Ca2+ into -motor neuron axon terminals
Release of Ach from the synaptic vesicles into the synaptic cleft
(200-300 vesicles/exocytosis)
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3.6.2.2. Postsynaptic end

Diffusion of Ach to Postjunctional membrane combination of Ach
with nAchR (107-108, 2 subunit) on the motor endplate
Conformational change opening of the gate permeability of the
postjunctional membrane to Na+ and K+
Transient change in the Em depolarization EPP
Depolarization of areas of muscle membrane adjacent to endplate and
initiation of AP
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Endplate Potential/EPP
i. A graded response (magnitude of depolarization No of open Ach channels)
ii. Transient (Ach is hydrolyzed to form choline and acetate).
iii. Amplitude: >50mV (= 70mV)
iv. No voltage-gated Na+ and K+ channels at the endplate region
( No AP, Na+, K+ channels are located on the muscle membrane
contiguous to the endplate).
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46

v. Ionic basis of EPP:

Ach activated channel.
Independent of membrane potential.
Em is between EK+ and ENa+ (-90mV . -20mV+60mV)
vi. EPP depolarizes the contiguous membrane regions by electronic
conduction to threshold and AP is generated.
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47

3.6.3. Propagation of AP into the T-tubule & release of Ca2+ from the
terminal cisternae
A. Transverse Tubule (T-tubule)
i. It is an invagination of the surface of the sarcolemma.
ii. It is found at the junction of A-I bands.
iii. One end of the tube is open to extracellular space, but its other
end is closed.
iv. Function: rapid transmission of AP from the cell membrane to
all the fibers on the muscle.
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B. Sarcoplasm Reticulum (SR)

i. It is an internal tubular structure that runs between the myofibrils.
ii. It is closed at both ends.
iii. It is not continuous with the sarcolemma.
iv. Functions:
Stores Ca2+ in the terminal cisternae (lateral sacs)
Uptake and release of Ca2+
NB
Calsequestrin is sarcoplasmic protein that binds Ca2+
(1 calsequestrin = 43 Ca2+ )
Triad: 2 lateral sacs + 1 transverse tubule
49

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50

3.6.3.1. Release of Ca2+ from the Terminal Cisternae

i. Voltage - gated Ca2+ channel ( Dihydropyridine receptors (DHPR) )
a. Found in the transverse tubule.
b. It is a pentamer (1,2, , , )
c. Voltage sensor which is responsible for triggering Ca2+ release
from the SR and for EC coupling.
d. Sensitive to a drug called Dihydroperidine which can block
this voltage-gated channels.
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Skeletal Muscles
ii. Ca2+- Release Channel (RyR)

a. Found in sarcoplasmic reticulum membrane.
b. It is a tetramer.
c. Sensitive for a drug called ryanodine.
Depolarization of the T-tubule
Opening of DHPR - RyR Ca2+ channels

(Connected by a foot process whose length is 20nm)
Release of calcium
Calcium flows out of the terminal cisternae into the sarcoplasm
[Ca2+] : 0.1 10mol/l

52

Skeletal Muscles
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53

3.6.4. The Activation of Muscle Proteins

3.6.4.1. Thick Filaments
a. Myosin (an actin binding protein)
b. 2 distinct structures:
Myosin head (cross bridge): actin-binding site
ATP binding site (ATPase):
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hydrolyzes ATP
54

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55

Resting state
i. Interaction of thick and thin filaments is inhibited.
ii. Troponin I & tropomyosin covers the sites where myosin heads bind to actin
Activated States:
Influx of Ca2+
Binds to Troponin C (4Ca2+)
Conformational change in troponin
Tropomyosin moves aside
Exposes the myosin-binding sites on actin
Myosin cross-bridge on the thick filament is exposed to actin filaments

56

3.6.5. Generation of Tension

Cross-bridge cycle: 4 steps
1. Binding of the cross-bridge to actin
A+M* . ADP. Pi
A.M* + ADP + Pi
Actin binding
2. Bending of the cross-bridge producing movement of thin filament.
A. M*. ADP. Pi
A.M*.+ ADP + Pi
Cross-bridge movement
3. Detachment of the cross-bridge from the thin filament
A.M+ ATP
A+ M.ATP
Cross-bridge dissociation from actin
4. The cross-bridge returns to its original upright position to repeat the cycle.
M.ATP
M*. ADP. Pi
ATP hydrolysis
57
NB. Tension is generated by repetitive cross-bridge cycling.

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58
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59

3.6.6. Relaxation of Muscle.

a. Removal of Ca2+ from the myoplasm (<0.1mol/l) into the SR.
b. For Ca2+ removal from the sarcoplasm the third ATP is consumed by
Ca2+-Mg2+ -ATPase/SERCA
c. After removal of Ca2+ :
i. Troponin returns to its original conformational state.
ii. Tropomyosin inhibition of Myosin-Actin interaction is
restored.
iii. Cross-bridge cycling stops and the muscle is returned
to its resting state.
d. Breakdown of Ach by AChE.
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60

Regulatory function of ATP

i. Actin + Myosin +ATP + Ca2+
Contraction
ii. Actin+ Myosin + ATP Ca2+
Relaxation
iii. In the absence of Ca2+ ATP is not hydrolyzed.

iv. 3 ATP molecules are needed:
a. For energizing the myosin cross-bridges
b. For dissociation of actinmyosin complex and initiation of relaxation
c. To pump out Ca2+ from the sacroplasm to sequester it into the SR
(Ca2+-Mg2+ - pump/SERCA pump)
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61

62

Changes
a. Banding
H-zone:Disappears
Z-line: Gets considerably darker
I-band: Narrower/smaller
A-band:
b. Contractile proteins:
c. Sarcomere:
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NO CHANGE in length of myosin or actin

Shortens
63

Changes in banding pattern
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64

Ion channels involved:

i.
Voltage-gated Ca2+ channels (active zone, -motor neuron)
ii.
Ligand-gated cationic channels (Ligand-gated Na+ channels, motor

endplate)
iii. Voltage-gated Na + channel (contiguous to the motor endplate).

iv. Voltage-gated Ca2+ channel (t-tubule)
v.
RyR Ca2+-release channel (SR)
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65

Summary
Discharge of motor neuron
Release of Ach at motor endplate
Binding of Ach to nAchR
gNa+ and gK + in endplate membrane
Generation of EPP
Generation of AP in muscle fibers

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66

Generation of AP in muscle fibers
Inward spread of depolarization along T-tubules
Release of Ca2+ from terminal cisterns of SR and diffusion to

thick and thin filaments
Binding of Ca2+ to troponin C, uncovering myosin-binding

sites on actin
Formation of cross-linkages between actin and myosin and

sliding of thin on thick filaments, producing movement
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67

Steps in relaxation
Ca2+ pumped back into SR
Release of Ca2+ from troponin
Cessation of interaction between actin and myosin
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68

Skeletal Muscles: Clinical correlates
Rigor Mortis
a. It is a state of muscle contracture, ie., contraction produced without
AP and not followed by relaxation.
b. It is a contracture which occurs in the muscles after death. It starts in
small muscles (2-3hrs) after death and involves all muscles in 12 hrs.
c. The rigidity is due to depletion of ATP from the muscle. Calcium
diffuses out of the SR & can not be recollected by the calcium pump.
d. Calcium initiates muscle contraction using the remaining ATP molecules,
relaxation does not occur because calcium is not recollected back into the
SR, and no ATP is available to disconnect the myosin heads from actin.
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Skeletal Muscles: Clinical correlates
e. It disappears when muscle fibers are autolysed by lysosomal enzymes

released after death.
f. It starts to disappear 14hrs after death and completed in 24hrs. High
environmental temperature accelerates the appearance and disappearance of
rigor mortis.
g. The extent of rigor mortis is used medically to determine the time of death.
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70

Cardiac Muscle
4. CARDIAC MUSCLE
4.1. Introduction
a. The physiological basis of heart pumping (contracting of the heart) is
to propel blood through the circulatory system.
b. It has SAME contractile machinery with some degree of modification.
(Actins, myosin, meromyosin, c-protein, nebulin, -actinin, tropomodulin)
c. REGULATION is done by intrinsic and extrinsic factors: neuronal

(ANS) + hormonal.
d. In the course of an average life span the heart will contract =
72 beats x 60 min x 24hrs x 365d x 70 years = 3x109
e. Abundance of connective tissue (prevent muscle rupture, prevent overstretching during periods of increased venous return).
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71

Cardiac Muscle
4.2. Cardiac Muscle Vs. Skeletal Muscle

4.2.1. Cardiac Muscle.
1. A cardiac myocyte has a single nucleus which is smaller ( = 15-20
m, L = 100m)
2. Has abundant amount of mitochondria (30%) + myoglobin which
makes it fatigue resistant (myoglobin facilitates the transport of
oxygen from the sarcolemma to the mitochondria)
3. A cardiac cells are joined end-to-end by intercalated discs:
i. Attach one cell to the next by means of desmosomes
ii. Connect the thin filament of the myofibrils of adjacent cells.
(Mechanical + electrical coupling)
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Cardiac Muscle
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Cardiac Muscle
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74

Cardiac Muscle
iii. Contain gap junctions which is synchronizing the contractions of

heart muscle cells.
4. The T-tubule is larger and it is found at z-line
5. The SR - makes contact with T-tubule and the cell membrane.
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75

Cardiac Muscle
4.3. Excitation-Contraction Coupling/Electrochemical Coupling

(Cardiac Vs Skeletal)
4.3.1. Cardiac Muscle.
1. Ca2+- release from the SR is triggered by Ca2+ (not by membrane
depolarization; extracellular Ca2+ SR (is responsible for
prolonged plateau phase)).
2. T-Tubule (DHPR) contains Ca2+ channel (through which Ca2+ enters
the cell during the AP).
3. SR-RyR containing Ca2+- release channel is opened by influx of Ca2+
from the T-Tubule.
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Cardiac Muscle
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77

Cardiac Muscle
4. Amount of Ca2+- release from the SR is under physiologic control.

a. Hormones (catecholamines) the amount of Ca2+ entering
the cell IC Ca2+
b. Pump (Na+-Ca2+ Exchanger, 1Ca2+, 3Na+) is increasing the
Ca2+ in the cell.
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Smooth Muscles
5. SMOOTH MUSCLE
5.1. Introduction
a. It is important in regulation of the airways, blood vessels, GIT, and
hollow organs (bladder, uterus...)
b. It is controlled by intrinsic factors (inherent rhythmicity): ANS +
Hormones.
c. It produces slower and longer-lasting contractions (slow & sustained)
(rates of cross-bridge cycling LATCH STATE maintain TONE
and little energy consumed).
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Smooth Muscles
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80

Smooth Muscles
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81

Smooth Muscles
5.2. Smooth Muscle Vs Striated Muscle.

5.2.1. Smooth Muscle.
1. It has NO STRIATIONS (sparse thick filaments and absence of
transverse registration).
(lacks visible cross-striations)
2. Has elongated spindle shaped cells with a single nucleus
(L = variable size, attached in series to bear equal stress).
3. Sarcomeres are absent.
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Smooth Muscles
4. The myofilaments are :

a. Thick filaments: myosin
b. Thin filaments: actin and tropomyosin (No troponin)
c. Thick and thin filaments are dispersed through out the cell.
They are not arranged in strictly ordered pattern.
(not arranged in regular arrays)
d. Thin filaments are attached to dense bodies
(functional equivalents of Z line)
i. Dense bodies: are found in cell membrane and cytoplasm
composed of actinin.
5. Tubules: T-tubules are absent and the SR is rudimentary (less
extensive than those observed in skeletal or cardiac muscle).
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Smooth Muscles
6.
a. smooth muscles contain few mitochondria and

b. depend, to a large extent, on glycolysis for their metabolic needs.
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84

Smooth Muscles
5.3. Excitation - Contraction Coupling (Smooth M.Vs Striated M.)

Cross-bridge cycling is regulated by Ca2+- induced phosphorylation of
myosin.
1. Myosin cross-bridge has 4 light chain
2. Myosin can not bind to actin unless one of these light chains
(LC20) is phosphorylated.
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Smooth Muscles
a. Phosphorylation of LC20 is by Myosin Light Chain Kinase (activated

by Calmodulin, 4Ca2+, Kinase - calmodulin - Ca2+- complex)
b. Ca2+ can enter the cell in a variety of ways:
i. Stimulation by a neurotransmitter (receptor-activated Ca2+ channel)
ii. Voltage-operated Ca2+channels
iii. Release from SR
(SR-IP3R) (IP3R = inositol triphosphate receptors)
3. The light chains are dephosphorylated by the enzyme Myosin Light
Chain Phosphorylase.
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Cardiac Muscle
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Smooth Muscles
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Smooth Muscles
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89

Smooth muscles: summary
Binding of Ach to mAchR
Increased Ca2+ influx into the cell
Activation of calmodulin-dependent myosin light chain kinase
Phosphorylation of myosin
Increased myosin ATPase activity and binding of myosin to actin
Contraction
Dephosphorylation of myosin by myosin light chain phosphatase
Relaxation, or sustained contraction due to the latch bridge and other

mechanisms
90

Smooth Muscles
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91

Muscle

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Muscle

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The Physiology of Muscles

3.4. Molecular Basis of Contraction

The Physiology of Muscles

The Physiology of Muscles

3.4.2 Thick Filament

The Physiology of Muscles

The Physiology of Muscles

The Physiology of Muscles

Geometrical orientation of myosin and actin

The Physiology of Muscles

The Physiology of Muscles

Geometrical orientation of myosin and actin

The Physiology of Muscles

3.4.3. Thin Filaments