Muscle and Muscle Tissue

Introduction

  • most distinguishing characteristic of muscle = its ability to turn chemical energy into mechanical energy → contraction

  • Function of Muscle Tissue: motion, heat production, posture and body support, breathing, speaking, protection of vital organs, regulating elimination of materials

Muscle Terminology/Organelles

  • sarcolemma: muscle fiber plasma membrane

  • sarcoplasm: muscle fiber cytoplasm → Contains many glycosomes for glycogen storage, as well as myoglobin for O2 storage

  • Modified organelles

    • Myofibrils

    • Sarcoplasmic reticulum

    • T tubules 

Types of Muscle Tissues

  • skeletal muscles

    • attach to and cover the bony skeleton

    • longest muscle cells

    • stripes = striations

    • voluntary

    • Function: Motility and Heat production

      • Contracts rapidly but tires easily

      • Adaptable → forces ranging from a fraction of an ounce to well over 70 lbs 

  • cardiac muscle

    • Only in the heart

    • Striated 

    • Not voluntary

    • Function: To pump blood throughout the body 

    • Under neural and hormonal controls

    • The heart and smooth muscle both have automaticity → Contains gap junctions

  • smooth muscle

    • Found in the walls of hollow visceral organs

    • Makes up the heart valves

    • Function: To force food and other substances through internal body channels

    • Not striated and involuntary 

  • functional characteristics of muscle tissue

    • Excitability (or irritability) and conductivity

    • Contractility

    • Extensibility

    • Elasticity 

Skeletal muscle

  • Each muscle is a discrete organ:Muscle tissue

    • Blood vessels

    • Nerve fibers

    • Connective tissue

    • Requires nerve stimulation 

  • Nerve and Blood Supply

    • Each muscle is served by one nerve, an artery, and one or more veins

    • Contracting fibers require continuous delivery of oxygen and nutrients

    • Wastes must be removed 

  • Skeletal Muscle: Attachments

    • Attached to bone in at least 2 places

      • Point of attachment to an immovable bone = the origin

      • Point of attachment to a movable bone= the insertion 

Mechanism of Skeletal Muscle Contraction

  • Sarcomeres:

    • Smallest contractile unit

    • Myofibril region between 2 Z discs

    • Composed of myofilaments:

      • Thick (myosin)

      • Thin (actin)

  • Ultrastructure of Myofilaments: Thick Filaments

    • Thick filaments: Composed of protein myosin that contains two heavy and four light polypeptide chains

      • Heavy chains intertwine to form myosin tail

      • Light chains form myosin globular head

      • During contraction, heads link thick and thin filaments together, forming cross bridges 

  • Ultrastructure of Myofilaments: Thin Filaments

    • Tropomyosin and troponin (Tn) are regulatory subunits

      • Troponin (Tn) complex of 3 proteins:

        • TnI: binds to actin

        • TnT: binds to tropomyosin

        • TnC: binds Ca+2 

    • Mostly made up of actin → helial polymer of globular subunits

    • 2 actin strands as an interwoven string

    • Active sites to which myosin heads attach during contraction 

  • Sliding Filament Theory

    • 2)Release of Pi upon binding cocks the myosin head, producing a power stroke that pulls the thin filament toward the center. 

    • After the power stroke, ADP is released and a new ATP binds.

      a) This makes myosin release actin.

      b) The new ATP is split.

    • The myosin head straightens out and rebinds to actin farther back.

    • Continues until the sarcomere has shortened 

  • Other Components of the Myofibril

    • Elastic filament: composed of protein titin → Holds thick filaments in place; helps recoil after stretch; resists excessive stretching

    • Dystrophin: Links thin filaments to proteins of sarcolemma

      • Nebulin, myomesin, C proteins bind filaments or sarcomeres together → Maintain alignment of sarcomere 

  • Sarcoplasmic Reticulum

    • Smooth endoplasmic surrounds each myofibril

    • Functions in the storage and regulation of intracellular Ca+2

    • T tubules (transverse) penetrate deep into the cell’s interior→ conduct impulses 

  • Skeletal Muscle Contraction

    • Skeletal muscle MUST

      • Be stimulated by a nerve ending

      • Propagate AP along its sarcolemma

      • ↑intracellular Ca2+

      • ATP

        • Linking the electrical signal to the contraction is excitation-contraction coupling 

  • Nerve Stimulus of Skeletal Muscle

    • Motor neurons of the somatic NS

    • Axons branch profusely

    • Each axonal branch forms a neuromuscular junction with a single muscle fiber 

  • Stimulating a Muscle Fiber

  • Action Potential in the Muscle

    • Ligand-gated sodium channels open → Na+ enters the cell

    • If the stimulus is strong enough, an action potential is initiated

  • Excitation-Contraction Coupling

    • Action potential:

      • Propagated along the sarcolemma

      • Travels down the T tubules

      • Triggers Ca2+ release from SR terminal cisternae

      • Ca2+ binds to troponin: Tropomyosin moves → Actin active binding sites exposed 

Contraction of Skeletal Muscle

  • The two types of muscle contractions are:

    • Isometric contraction – ↑ muscle tension (muscle does NOT shorten during contraction)

      • This muscle is attached to a weight that exceeds it’s maximum 

        tension producing capacity. 

    • Isotonic contraction – ↓ muscle length (muscle shortens during contraction) 

  • Motor Unit

    • An α motor neuron and all the muscle fibers it supplies

    • # of muscle fibers per motor unit can vary from four to several hundred 

    • ↓#muscle fibers = ↑ control = finer more precise movements

  • Muscle Twitch, Summation, and Tetanus

    • A muscle twitch =A single action potential

    • 3 phases= 

  • Graded Muscle Responses

    • Muscle twitches → NOT how muscles usually contract

    • Muscle contractions:

      • Smooth

      • Vary in strength depending on the demand placed on them (Graded) 

        • Responses are graded by:

          • Changing the frequency of stimulation

          • Changing the strength of the stimulus 

  • Muscle Response to Varying Frequency of Stimuli

    • ↑ frequency = ↑contractile force = twitch summation

    • Tetanus results:

      • Muscle relaxation disappears

      • Contractions fuse into a smooth, sustained contraction 

  • Muscle Response: Stimulation Strength

    • ↑strength of stimulus = ↑ strength of muscle contraction = Multiple Motor Unit Summation 

  • Stimulus Intensity and Muscle Tension

  • Size Principle

    • Recruitment is dictated by the size principle →

      • In any muscle, motor units with the smallest fibers are controlled by small ↑ excitable motor neurons → activated first 

    • The largest motor units have contractile forces ~50 times > small muscle fibers 

      → Innervated least excitable neurons (large) 

  • Treppe: The Staircase Effect

    • A long period of rest → initial contractions may be only ½ as strong as subsequent contractions

    • ↑ contraction strength in response to multiple stimuli of the same strength 

    • Contractions ↑ because:

      • ↑Ca2+ in the sarcoplasm

      • ↑heat generated = ↑efficiency enzymes 

Muscle Metabolism

  • Energy for Contraction

    • ATP is the only source used

    • ATP regenerated by 3 pathways:

      • ADP with creatine phosphate (CP) 

      • Anaerobic glycolysis 

      • Aerobic respiration 

  • Where Muscles Get Their Energy

    • At rest and for mild exercise: from the aerobic respiration of fatty acids

    • For moderate exercise: from glycogen stores

    • For heavy exercise: from blood glucose

      • As exercise intensity and duration increase, GLUT4 channels are inserted into the sarcolemma to allow more glucose into cells. 

  • Muscle Fatigue

    • Reversible reduction in the muscle’s ability to generate a force

    • Muscle fatigue occurs when ATP production fails to keep pace with ATP use 

    • There is a relative deficit of ATP, causing contractures (states of continuous contractions because cross bridges can’t detach)

      • Lactic acid accumulates in the muscle (cause?)

      • Ionic imbalances are present 

    • Intense exercise → rapid muscle fatigue

      • Na+-K+ pumps cannot restore ionic balances quickly enough

      • Prolonged ↓intensity exercise → slow-developing fatigue

        • SR is damaged → impairs Ca2+ regulation 

    • Adaptation to Endurance Exercise Training 

  • Skeletal Muscle Damage and Repair

    • Stem cells called satellite cells located near muscle fibers.

    • These can fuse to damaged muscle cells and repair them or fuse to each other to form new muscle fibers.

    • Myostatin is a paracrine regulator that inhibits satellite cells. 

Force of Muscle Contraction

  • Affected by:

    • # of muscle fibers contracting – the ↑ # motor fibers in a muscle, the stronger the contraction

    • The size of the muscle – the bulkier the muscle, the greater its strength 

  • Degree of muscle stretch –strongest when muscle fibers are 80-120% of their normal resting length

    • Length-tension relationship 

Smooth Muscle

    • Spindle-shaped fibers

    • Lack the coarse connective tissue sheaths 

  • Longitudinal layer contracts → organ dilates and contracts 

  • Circular layer contracts → organ elongates+ lumen narrows 

  • Innervation of Smooth Muscle

    • NO neuromuscular junctions

    • Innervating nerves have axonal terminal swellings called varicosities

    • Varicosities release neurotransmitters into wide synaptic clefts called diffuse junctions 

  • Microscopic Anatomy of Smooth Muscle

    • SR is less developed and lacks a specific pattern

    • NO T tubules

    • Plasma membranes have pouchlike infoldings called caveoli 

    • Ca2+ is sequestered by caveoli from outside the cell

    • NO striations + NO sarcomeres

    • Thin and thick filaments 

    • NO troponin complex

    • NO Z discs

    • Thick and thin filaments arranged diagonally → corkscrew contraction 

  • Smooth Muscle Contraction

    • They contract in unison

    • Gap junctions

    • Some smooth muscle cells: Act as pacemakers → Self-excitatory and depolarize without external stimuli 

  • Excitation-Contraction Coupling in Smooth Muscle

  • Special Features of Smooth Muscle Contraction

    • Takes 30 X longer to contract and relax

    • Smooth muscle tone: without fatiguing

    • Slow, prolonged contractile activity

    • ↓ energy requirements

    • Response to stretch 

  • Response to Stretch

    • Stress-relaxation response:

      • Responds to stretch only briefly, and then adapts to its new length

      • However, it retains its ability to contract

      • This enables organs such as the stomach and bladder to temporarily store contents 

  • Types of Smooth Muscle: Single Unit

    • Contract rhythmically as a unit

    • Are electrically coupled to one another via gap junctions

    • Often exhibit spontaneous action potentials

    • Are arranged in opposing sheets and exhibit stress-relaxation response 

  • Types of Smooth Muscle: Multiunit

    • Their characteristics include:

      • Rare gap junctions

      • Infrequent spontaneous depolarizations

      • Structurally independent muscle fibers 

      • A rich nerve supply, which, with a number of muscle fibers, forms motor units

      • Graded contractions in response to neural stimuli 

Comparison Between Cardiac, Smooth, and Skeletal Muscle