Physiology - Chapter 8: Muscle Physiology
Muscle fiber is excited
Ca+ pulls troponin tropomyosin complex from blocking binding site
The link between excitation and contraction
Calcium
What pulls the thin filaments inward?
Cycles of cross-bridge binding and bending, sliding filament mechanism
Is a motor protein, similar to kinesin
Myosin
Extrafusal vs intrafusal fibers
*Extrafusal* --> 99% of fibers -alpha motor neurons -contractile strength *Intrafusal* --> very few -gamma motor neurons -do NOT contribute to contractile strength
gradation accomplished in skeletal muscle
- # of motor units contracting - frequency of stimulation (twitch summation)
Three structures that play important roles in linking excitation and contraction
- Calcium - Sarcoplasmic reticulum - Transverse Tubules
Contractile activity at lengths lower than Lo
- actin-myosin overlap = no cross bridge - ends of myosin pushed up against discs - no calcium release (80% lower than Lo)
Where can single-unit smooth muscle be found?
- digestive - reproductive - urinary (tracts) - small blood vessels
How do muscles respond to demands and adapt?
- improve oxidative capacity (changing the # of mitochondria or capillaries) - muscle hypertrophy (increase actin/myosin production)
gradation accomplished in multiunit smooth muscle
- number of muscle fibers contracting - cytosolic calcium concentration in each fiber
polypeptide units that make up troponin
- one binds to tropomyosin - one binds to actin - one binds to Ca
smooth muscle contraction
- phasic - tonic
How are skeletal muscles stimulated to contract
- release of ACh at neuromuscular junctions between motor neuron terminal buttons and muscle fibers. - ACh binds with the motor end plate - permeability changes as a result - action p. conduction
how to override withdraw pain reflex (ex. vaccine shot)
- send epsps to motor neurons of triceps (to activate) 2 send ipsps to biceps (to inhibit)
categories of fibers
- slow oxidative (1) - fat oxidative (2a) - fast glycolytic (2x)
three kinds of motor activity
- somatic reflex (reflex arc: postural or protective) - voluntary movement (can become a reflex) - rhythmic (walking, chewing...etc)
How do muscles shorten?
- the individual sarcomeres shorten which pulls the z discs closer together - by moving specialized intracellular components, causing them to develop tension and shorten.
gradation accomplished in cardiac muscle
- varying length of fibers - cytosolic calcium concentration (through autonomic, hormonal, and local metabolite influences.
multiunit smooth muscle can be found in
- walls of large blood vessels - in small airways of lungs - ciliary muscle - muscle of iris - the base of hair follicles (goosebumps)
Functions of titin
1) It stabilizes the thick filaments and keeps them in place 2) It acts like a spring attached to myosin returning the muscle to its normal position after stretch 3) Participates in signal transduction (stimulus to activity)
Controlled contraction of muscle allows:
1) Purposeful movement of the whole body or parts of the body 2) Manipulation of external objects 3) Propulsion of contents through various hollow internal organs 4) Emptying contents of certain organs to the external environment
Adjustable factors to produce graded contractions?
1) number of muscle fibers contracting 2) tension developed by each contracting fiber
sites crucial for contractile processes in each cross bridge
1- actin binding site 2- myosin ATPase site
Influences each muscle fiber and the tension developed
1- frequency of stimulation 2- contraction onset 3- fatigue 4- fiber thickness
4 steps in the excitation, contraction, and relaxation processes that require ATP:
1- splitting of ATP into ADP and Pi by ATPase (provides E for powers stroke) 2- Bind ATP to myosin 3- Active transport of Ca2+ back into lateral sacs of the SR during relaxation 4- Activity of Na+-K+ pump during action potential production
More ATPase
2a and 2x fibers (more cycling=faster)
describe alpha muscle spindle reflex
1. change in muscle length 2. recognized by muscle spindles 3. alpha output to extrafusal fibers 4. contraction
types of muscle atrophy
1. disuse atrophy (cast/brace) 2. innervation atrophy (nerve supply to muscle is lost) 3. sarcopenia (age related; decrease in motor neurons, protein synthesis and hormone levels = 1/4 pound of muscle lost a year) 4. limited repair: muscle cannot divide mitotically to repair itself.
ways to increase size of muscle (hypertrophy) / increase the production of actin and myosin
1. influence of testosterone (increases synthesis) 2. interconversion of fast muscle types
proprioceptors send information about length and tension through
1. muscle spindles (length) 2. golgi tendon organs (tension)
gamma neuron output
1. output to edges of fibers 2. initiate contraction of intrafusal fibers
postural somatic reflexes input comes from...?
1. proprioceptors (muscles and joints) 2. touch receptors in the skin 3. vestibular apparatus 4. eyes (horizon)
why is there an increase in oxygen consumption after exercise?
1. reset medical disturbances 2. restore nutrients 3. repair O2 debt (resynthesize phosphate, pyruvate glucose...)
Power stroke
Actual motion (single power stroke) pulling the thin filaments toward the m-line, cross bridge bends pulling myofilament inward
T/F: only one myosin head can attach to the actin at a time
True
T/F: contractile activity outlasts electrical activity that initiates it
True, but why? - Ca+ takes more time to be reabsorbed as lateral sacs have to reabsorb all of it, action p. is over before binding of actin and myosin occurs
T/F neither thick nor thin filaments change length during shortening
True, contraction is accomplished by the movement of thin filaments from opposite sides, not shortening of the filaments themselves
more actin or myosin in a single muscle fiber?
actin (32 billion) > myosin (16 billion)
proteins of thin filaments
actin (spherical) tropomyosin, troponin
Myofibrils are composed of
actin and myosin myofilaments / highly organized cytoskeletal microfilaments
motor neurons in the spinal cord receive direct input from
afferent neurons, brain stem, primary motor cortex
influence motor output
afferent neurons, primary motor cortex, brainstem(multineural), cerebellum
detachment (3rd step of cross bridge activity)
after power stroke, cross bridge detaches and returns to its original conformation
maximum contraction
all muscle fibers activated
Why is contraction during muscle twitch not maximal?
Not all cross bridges find a binding site as cytosolic Ca declines.
phasic vs tonic
Phasic- change, short-lived things (hollow tubes) tonic - contracted all the time ex. arterioles
multineuronal motor system
Regulates body posture and involuntary movements of larger muscle groups
_________ allows for the troponin tropomyosin complex to slip back into its blocking position
Removing cytosolic Ca+ by SERCA pump
What splits ATP into ADP and Pi (inorganic phosphate)?
The enzyme ATPase on the myosin Inorganic phosphate is released during the power stroke
fulcrum
The fixed point around which a lever pivots
When do actin and myosin detach?
Until ATP attaches to myosin, ending the power stroke as it reduces binding affinity between myosin head and actin. ATP is then split by ATPase.
contraction does not take place in...
a relaxed muscle fiber
why can thin filaments be pulled along the thick filaments for further distance in smooth muscle?
because cross bridges are present along the entire smooth muscle filament length
Why is single-unit smooth muscle also called visceral smooth muscle?
because it can be found in the walls of hollow organs.
Why does tone exist in tonic smooth muscle?
because of its low resting potential
flexion and extension
bending and extension of a limb biceps and triceps
the formation and breaking apart of ATP is only needed for release not for...
binding
how are smooth muscle cells turned on?
by calcium dependent phosphorylation of myosin
chemical change in smooth muscle
calcium phosphorylation chain
force > load
can move
the two types of muscle that physically reposition troponin-tropomyosin complex to uncover actin cross bridge binding sites
cardiac and skeletal muscle (smooth muscle = chemically)
autonomic / involuntary muscle
cardiac and smooth muscle
Isometric contraction
constant length (ex. standing, holding beverage in between drinks)
isokinetic contraction
constant motion (ex. machines at a gym)
Isotonic contraction
constant tension (ex. lift object)
Basal nuclei receive input from
cortical motor and sensory areas
First source of supplying ATP for skeletal muscle contraction (that is, during the first few seconds, at the very beginning of contraction)
creatine phosphate
most rapid energy pathway
creatine phosphate (one equation)
creatine kinase equation
creatine phosphate + ADP (forward-reverse arrows) creatine + ATP
most energy is stored in muscle as
creatine phosphate pools
Availability of calcium in the cytosol allows for...?
cross bridges to participate in the cycling process for a longer time
load and velocity
directly related = eccentric contractions inversely related = concentric contractions
SERCA pump (Sarcoplasmic, endoplasmic reticulum, calcium ATPase).
energy consuming carrier in the SR, which actively transports Ca+ from cytosol to lateral sacs to relax the skeletal muscles
terminal cisternae
enlarged areas of the sarcoplasmic reticulum surrounding the transverse tubules; lateral sacs
the rise in cytosolic calcium is mostly from __________ in smooth muscle.
extacellular fluid (ECF)
The number of fibers contracting within a muscle depends on...?
extent of motor unit recruitment
motor neurons in the spinal cord serve as
final common pathway for efferent component for somatic spinal reflexes, voluntary motor activities, and rhythmic activities.
The brainstem serves as
final link in multineuronal motor system
muscle fiber more resistant to fatigue
has higher capacity for ATP production
asynchronous recruitment
helps avoid fatigue during sustained muscle contractions, refers to different motor units taking turns maintaining muscle tension, and makes an entire muscle seem to contract smoothly.
fatigue
inability to maintain a given level of muscle tension or intensity during activity
increase in tension
increase afferent firing
spastic paralysis
increased muscle tone (rigidity) and augmented limb reflexes (lowered inhibitory responses)
alpha motor neurons
innervate extrafusal muscle fibers
functional sycytium
interconnected muscle cells that function electrically and mechanically propagate action potentials in single unit smooth muscle, contract as a single unit
Where is tension produced?
internally within the sarcomeres
Muscle tension
is transferred to bone as titin
Three types of contraction
isotonic, isometric, isokinetic, concentric, eccentric
lowering object --> biceps ______
lengthen
muscle is attached to bone
limitation on muscle lengths
flaccid paralysis
low muscle tone; a state in which the muscles are limp and cannot contract, caused by nerve lesion
Tetanus
maximum number of cross bridge binding sites uncovered
more or less actin in smooth muscle
more (15-20 actin per myosin) skeletal m. = 2 to 1
the stronger the contraction
more motor units recruited
motor units _____ resistant to fatigue are recruited first
most
The brainstem receives input from:
motor cortex, cerebellum, basal nuclei, afferent neurons
Troponin when Ca+ is present
moves tropomyosin away from blocking the binding site
type of smooth muscle that initiates contraction
multiunit smooth muscle
compromises largest group of tissues in the body:
muscle
eccentric contraction
muscle lengthens (ex. lower object)
concentric contraction
muscle shortens
Tendons
muscle to bone attachment non-contractile elastic tissue
muscle fibers are composed of...?
myofibrils
can excite or inhibit to modify contraction
myogenic single-unit smooth muscle
oxidative fibers have more...?
myoglobin, mitochondria, capillaries
what protein does calmodulin activate?
myosin light chain kinase (MLC kinase, which phosphorylates myosin light chain)
approximately at Lo
naturally relaxed, vary 30% on average throughout the day
myogenic activity in single-unit smooth muscle
nerve independent in which muscle initiates contraction itself
neurogenic
nerve produced contractile activity (multiunit smooth muscle and skeletal muscle)
10% larger than Lo
no contraction
tetanus contractions
no relaxation; spasm and much more frequent
is threshold always reached in slow wave potentials?
no, depends on starting point of membrane potential at onset
I - band
non-projecting portion of thin filaments into A-band
areas without myosin in smooth muscle
none
troponin and tropomyosin in smooth muscle
none
Muscle twitch
one action potential in one muscle fiber
muscle tone
ongoing state of muscle tension even when at rest
where do slow wave potentials occur?
only in smooth muscle of the digestive tract
red fibers
oxidative fibers
used to form energy for light exercise such as walking or jogging
oxidative phosphorylation
pacemaker potential
pacemaker potentials: shifts in passive ionic fluxes accompany automatic changes in ion channel permeability resulting in the membrane potential gradually depolarizing on its own
series elastic component
passive elastic property of muscle derived from the tendons
All multiunit smooth muscle is
phasic
all musltiunit smooth muscle is _________
phasic
which type of smooth muscle contracts in bursts?
phasic smooth muscle
neuromuscular junction
point of contact between a motor neuron and a skeletal muscle cell releases ACh which diffuses and triggers action p.
examples of skeletal muscles being subject to involuntary regulation
posture, balance, walking
corticospinal tract
precise, coordinated limb movements
connective tissue role in muscle
prevents over-stretching
issues voluntary command to appropriate motor neurons
primary motor cortex descending system
send information about length and tension
proprioceptors
muscle atrophy decreases the number of ______________.
protein molecules (actin/myosin) NOT MUSCLE FIBERS
Muscles can only _____ never _____
pull, push
How is it impossible to achieve summation of action potentials?
refractory period occurs
calmodulin
regulatory protein that facilitates contraction in smooth muscles, calcium binding
functional unit of muscle
sarcomere (smallest component that can perform all functions of the organ)
person becomes aware of pain once impulse reaches __________.
sensory area of the cortex
Dihydropiridine Receptors in smooth muscles
serve as calcium channels
Foot proteins (ryanodine receptors)
serve as calcium release channels, in sarcoplasmic reticulum
how is tension transmitted from sarcomere to bone?
shortening of sarcomere ---> tightens series elastic component ---> force applied to bone moving it against a load
precise motor activity
smaller amount of motor fibers
clear cut length tension relationship is not present in
smooth muscle
has no T-tubules
smooth muscle
myosin molecules are arranged in a way so that cross bridges are present along the entire filament length in what muscle?
smooth muscle
only tropomyosin can be found in
smooth muscle
slowest speed of contraction
smooth muscle
to allow actin/myosin cross bridge in skeletal vs. smooth muscle
smooth muscle = chemical change skeletal muscle = physical change
what initiates slow wave potentials?
specialized clusters of nonmuscle pacemaker cells within the digestive tract and spread to adjacent smooth muscle via gap junctions
slow wave potentials
spontaneous; membrane potential undergoes gradually alternating depolarizing and hyperpolarizing swings
cells that repair minor injury in muscle
stallate cells (through giving rise of myoblasts)
maximum tension cannot be reached
start anywhere other than Lo
Rigor Mortis
stiffness of death; Ca+ released in excessive amounts as it leaks out of lateral sacs causing the troponin-tropomyosin complex to move aside and the actin and myosin contract. Because ATP is needed to detach them, and it cannot be produced after death, actin and myosin are attach forever.
Important for maintaining posture
submaximal isometric contractions
How does Ca+ get into the cytosol?
surface --> T-tubules --> SER --> cytosol
striations
t-tubules present (cardiac and skeletal)
Contraction ends when...?
the cross bridges become inactive and the reuptake of Ca+2 is finished
End plate potential
the depolarizations of skeletal muscle fibers caused by neurotransmitters binding to the postsynaptic membrane in the neuromuscular junction
while the rise in cytosolic calcium is what triggers contractions in skeletal and smooth muscle, the difference is in...?
the type of change induced (chemical/physical)
A- band
thick filaments + overlapping thin
H zone
thick filaments only, near M-line
Contractile activity at lengths greater than Lo
thin filaments pulled out of thick filaments
latent period
time between application of a stimulus and the beginning of a response in a muscle fiber
coiled portion at the end of the sacromere that connects the m-line to the z-disc
titin
involved in the complex pathway of muscle enlargement during weight lifting (signal transduction)
titin
Largest protein in the body?
titin (30,000 A.A)
examples of skeletal muscles that are not attached to 2 ends but produce movement
tongue external eye circular sphincters
can maintain low level tension even without action potentials, therefore are too myogenic
tonic single unit smooth muscle, why? - have sufficient cytosolic calcium
type of smooth muscle that maintains tone?
tonic smooth muscle
_________ covers actin sites that bind to cross bridges
tropomyosin
Troponin when not bound to Ca2+
troponin stabilizes tropomyosin in blocking position over actin's cross-bridge binding sites
How is tension generated by sarcomeres transmitted to bones when they do not come in contact?
via tendons
single unit
visceral smooth muscle ex: uterus gap junctions during birth
rhythmic activity examples
walking or chewing
example of tonic smooth muscle?
walls of arterioles
where is phasic smooth muscle most abundant?
walls of hollow digestive organs
Ca+ binds to troponin causing tropomyosin to change shape
what happens after calcium diffuses into cytosol?
Why may someone not be able to support a heavy object?
when all motor fibers participate, since it is a maximal contraction, it is impossible to alternate motor unit activity and prevent fatigue
how is a light chain phosphorylated in smooth muscle?
when inorganic phosphate attaches to it
when is energy released during creatine phosphate?
when the bond between creatine and phosphate is broken
when can myosin attach to actin in smooth muscle?
when the light chain is phosphorylated
levels of organization of muscle:
whole muscle muscle fiber (cell) myofibril (intracellular structure) cytoskeletal elements (thick and thin filaments) protein molecules (myosin and actin)
How do we get ATP?
- creatine phosphate (high energy phosphate from creatine phosphate to ADP) - oxidative phosphorylation (electron transport system +chemiosmosis) - glycolysis
gradation accomplished in single-unit smooth muscle
- cytosolic calcium concentration (through myogenic activity) - influences of autonomic nervous system, mechanical stretch, local metabolites, hormones
pain reflexes
1. withdraw (can override it with cortical influence) 2. cross extensor (require activation of both sides of the body)
skeletal muscle body weight
40% in males, 32% in females
Glycolysis
A metabolic process that breaks down carbohydrates and sugars through a series of reactions producing pyruvate which is then converted to lactate when there is a lack of oxygen
sarcoplasmic reticulum
A modified form of endoplasmic reticulum; stores calcium that is used to trigger contraction when muscle is stimulated
motor unit
A motor neuron and all of the muscle fibers it innervates
What leads to contraction of muscle fiber
Binding of myosin and actin at cross bridges
T/F: actin and myosin can only be found in muscles
F, they are only more abundant and organized in muscles
______ becomes smaller as thin filaments move inward, ______ narrows.
H-zone, I-band
what causes striations?
Highly organized internal arrangements, muscle fibers lying parallel to one another then bundled together by connective tissue.
Consists of thin filaments that do not overlap
I - band
Why does the width of the A-band remain unchanged as the thin filaments are pulled inward?
It is determined by the thick filaments, which do not change length during contraction
how is fatigue a protective mechanism?
It prevents reaching a point with no ATP (rigor)
genetic endowment of muscle fiber types
Largely determined by the type of activity for which the muscle is specialized
Multineural
Neural integration of muscle action Communicates with subcortical areas: - Basal ganglia - Thalamus - Hypothalamus - Reticular formation
serves as source of cytosolic calcium in all muscle types
SR
Dihydropiridine Receptors
Serve as voltage sensors in Transverse Tubules, activated when local depolarization of T tubules occurs. They trigger the opening of foot proteins
presence of gap junctions
Skeletal - no Cardiac - yes at intercalated discs Smooth - yes in unitary muscle (mostly single unit)
How does the muscle relax?
Through the reabsorption of Ca+
monitors motor commands from motor cortex and sensory feedback from muscles
cerebellum
how does smooth muscle permit the binding of cross-bridges
chemically brings about the phosphorylation of myosin cross bridges in thick filaments so that they can bind with actin
work
force x distance
dense bodies
found in smooth muscle, since they do not contain sarcomeres, they do not contain z-lines, so the dense bodies contain the same proteins the z-line contains.
what links fibers of single-unit smooth muscle?
gap junctions
What connects cardiac muscle cells?
gap junctions in intercalated discs
when oxidative phosphorylation cannot keep up
glycolysis
slower relaxation
slower removal of calcium
all fibers in a motor unit are
same type
The T tubules dip deep into
sarcolemma, between the A and I bands of myofibrils, allowing action potentials to reach deep into the muscle
reciprocal innervation
simultaneous stimulation of nerve supple and inhibition of nerves to antagonist muscle
What forms a functional syncytia?
single unit smooth muscle or visceral smooth muscle
single-unit vs multiunit smooth muscle
single unit: gap junctions so contractions occur as one unit myogenic (modulate by stretch, neural, and hormonal control) Ex. GI, uterus, small diameter blood vessels -> typically move contents within hollow organs multiunit: few to no gap junctions neurogenic-faster change in force via recruitment ex. hair and feather erectors, lens, iris **DO NOT have APs, cells are small enough Ca entry depolarization and graded potentials to spread
may be phasic or tonic
single-unit smooth muscle
How do each of the types of muscles maintain homeostasis
skeletal : generating heat, chewing, breathing، swallowing smooth: regulates the movement of blood, food, air, urine cardiac: pumps life sustaining blood
striated muscle
skeletal and cardiac muscle
troponin and tropomyosin
skeletal and cardiac muscle
ECF does not serve as source of cytosolic calcium in
skeletal muscle
Most abundant muscle?
skeletal muscle
no gap junctions
skeletal muscle
which type of muscle cannot be modified by hormones?
skeletal muscle
sarcoplasmic reticulum is best developed in _________ and most poorly developed in ____________.
skeletal muscle, smooth muscle
all types of muscle use the same mechanism of contraction which is
sliding filament mechanism
which type of fiber has the highest resistance to fatigue and why?
slow oxidative, has most capacity for ATP formation
muscle memory
the process by which the brain memorizes muscle coordination (motor skills) through repetition and stores the info. in the cerebellum
Sustained elevation in cytosolic calcium and more time to stretch tendons results in...?
twitch summation
What happens when muscle fiber is stimulated a second time before it has relaxed?
twitch summation: second contractile response or piggyback on top of the first twitch add together.
Myosin made up of
two identical heads
the sarcomere is the area between
two z-lines
The shortening of the muscle is in reality...
z lines being brought closer together as thin filaments attached to them slide inward
cytoskeletal disc that ,m,connects the thin filaments or two adjoining sarcomeres
z-line