Smooth muscle
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Smooth Muscle
Physiology • Muscles
SMOOTH MUSCLE
Distribution, structure, types, electrical activity, contraction, plasticity, neuromuscular junction and control of smooth muscle.
1. DISTRIBUTION OF SMOOTH MUSCLE
- Smooth muscles are nonstriated (plain) and involuntary muscles.
- Smooth muscles are present in almost all organs in the form of:
- Sheets
- Bundles
- Sheaths around other tissues
- Smooth muscles form the contractile tissues of various organs.
2. STRUCTURES HAVING SMOOTH MUSCLE
- Wall of organs such as:
- Esophagus
- Stomach
- Intestine in gastrointestinal tract
- Ducts of digestive glands.
- Trachea, bronchial tube and alveolar ducts of respiratory tract.
- Ureter, urinary bladder and urethra in excretory system.
- Wall of blood vessels in circulatory system.
- Arrector pili of skin.
- Mammary glands, uterus, genital ducts, prostate gland and scrotum in reproductive system.
- Iris and ciliary body of the eye.
3. FUNCTIONS OF SMOOTH MUSCLE
Smooth muscles are concerned with very important functions in different parts of the body.
1. In Cardiovascular System
- Smooth muscle fibers around the blood vessels regulate:
- Blood pressure
- Blood flow through different organs and regions of the body
2. In Respiratory System
- Contraction and relaxation of smooth muscle fibers of air passages:
- Alter the diameter of air passage.
- Regulate the inflow and outflow of air.
3. In Digestive System
- Smooth muscle fibers in the digestive tract help in:
- Movement of food substances.
- Mixing of food substance with digestive juices.
- Absorption of digested material.
- Elimination of unwanted substances.
- Sphincters along the digestive tract regulate flow of food materials.
4. In Renal System
- Smooth muscle fibers in renal blood vessels regulate:
- Renal blood flow
- Glomerular filtration
- Smooth muscles in ureters propel urine from kidneys to urinary bladder through ureters.
- Smooth muscles present in urinary bladder help in voiding urine to the exterior.
5. In Reproductive System
In Males
- Smooth muscle fibers facilitate the movement of:
- Sperms
- Secretions from accessory glands
In Females
- Smooth muscles:
- Accelerate the movement of sperms through genital canal after sexual act.
- Move ovum into uterus through fallopian tube.
- Help in expulsion of menstrual fluid.
- Help in delivery of the fetus.
4. STRUCTURE OF SMOOTH MUSCLE
- Smooth muscle fibers are fusiform or elongated cells.
- Nucleus is single, elongated and centrally placed.
- Normally, two or more nuclei are present in the nucleus.
- Smooth muscle fibers are generally very small:
- 2–5 microns in diameter
- 50–200 microns in length
- Smooth muscle fibers are covered by connective tissue.
- T tubules are absent.
Myofibrils and Sarcomere
- Well-defined myofibrils and sarcomere are absent in smooth muscles.
- Therefore, alternating dark and light bands are absent.
- Absence of dark and light bands gives the nonstriated appearance to smooth muscle.
Myofilaments and Contractile Proteins
- Contractile proteins in smooth muscle fiber are:
- Actin
- Myosin
- Tropomyosin
- Troponin or troponin-like substance is absent.
- Thick and thin filaments are present in smooth muscle fibers.
- Filaments are not arranged in an orderly fashion as in skeletal muscle.
- Thick filaments formed by myosin molecules:
- Have many numbers of cross bridges than in skeletal muscle.
- Thin filaments are formed by:
- Actin
- Tropomyosin molecules
Dense Bodies
- Dense bodies are special structures of smooth muscle fibers to which:
- Actin
- Tropomyosin
Sarcotubular System
- Sarcotubular system in smooth muscle fibers is present in the form of a network.
- T-tubules are absent.
- L-tubules are poorly developed.
5. TYPES OF SMOOTH MUSCLE FIBERS
1. Single-unit or Visceral Smooth Muscle Fibers
2. Multiunit Smooth Muscle Fibers
6. SINGLE-UNIT OR VISCERAL SMOOTH MUSCLE FIBERS
- Single-unit smooth muscle fibers are fibers with interconnecting gap junctions.
- Gap junctions allow rapid spread of action potential throughout the tissue.
- All muscle fibers show synchronous contraction as a single unit.
- Single-unit smooth muscle fibers are also called visceral smooth muscle fibers.
Features of Single-Unit Smooth Muscle Fibers
- Muscle fibers are arranged in:
- Sheets
- Bundles
- Cell membrane of adjacent fibers fuses at many points to form gap junctions.
- Through gap junctions, impulses move freely from one cell to another.
- Thus, a functional syncytium is developed.
- Syncytium contracts as a single unit.
- In this way, visceral smooth muscle resembles cardiac muscle more than skeletal muscle.
- Single-unit smooth muscle fibers are present in walls of organs such as:
- Gastrointestinal organs
- Uterus
- Ureters
- Respiratory tract
- Etc.
7. MULTIUNIT SMOOTH MUSCLE FIBERS
- Multiunit smooth muscle fibers are muscle fibers without interconnecting gap junctions.
- Multiunit smooth muscle fibers resemble skeletal muscle fibers in many ways.
Features of Multiunit Smooth Muscle Fibers
- Muscle fibers are individual fibers.
- Each muscle fiber is innervated by a single nerve ending.
- Each muscle fiber has an outer membrane made up of myofilament, which helps to insulate and separate the muscle fibers from one another.
- Control of muscle fibers is mainly by nerve signals.
- Smooth muscle fibers do not exhibit spontaneous contractions.
Multiunit Smooth Muscles are Present in
- Ciliary muscles of the eye
- Iris of the eye
- Nictitating membrane (in cat)
- Arrector pili
- Smooth muscles of blood vessels
- Urinary bladder
8. DIFFERENCES BETWEEN SINGLE-UNIT AND MULTIUNIT SMOOTH MUSCLE
| Feature | Single-unit Smooth Muscle | Multiunit Smooth Muscle |
|---|---|---|
| Presence | More common | Less common |
| Appearance | Arranged like sheets or bundles of tissue | Discrete individual muscle fibers |
| Situation | In small blood vessels and walls of hollow organs, such as gastrointestinal tract, respiratory tract, urinary system, etc. | Ciliary muscles of the eye, iris of the eye, nictitating membrane (in cat), arrector pili and larger blood vessels |
| Interconnection | Has gap junctions which allow rapid passage of action potential | No gap junctions; each muscle fiber is innervated by single nerve ending |
| Pacemaker cells | Self-excitable pacemaker cells are present; spontaneous rhythmical contractions occur | No pacemaker cells and no spontaneous contractions |
| Control of action | Myogenic | Neurogenic |
| Resting membrane potential | Unstable resting membrane potential with slow spike potentials due to rhythmic modulations in sodium-potassium pump | Stable resting membrane potential |
| Action potential | Can be generated spontaneously; can be elicited by electrical or hormonal stimulation; spreads rapidly throughout the sheet of cells and makes the cells act as a single unit; occurs with a plateau due to long depolarization and slow repolarization | Cannot be generated spontaneously; can be elicited by neural and hormonal stimulation; selective activation of each muscle fiber that can contract independently of each other; no plateau |
| Activity | All the fibers in an organ behave as a single unit, i.e. all the muscle fibers of an organ contract or relax together and permit contraction or relaxation of the whole organ | Each fiber behaves independently, i.e. each fiber contracts and relaxes on its own |
9. ELECTRICAL ACTIVITY IN SINGLE-UNIT SMOOTH MUSCLE
- Usually, 30 to 40 smooth muscle fibers are simultaneously depolarized, which leads to development of self-propagating action potential.
- This is possible because of:
- Gap junctions
- Syncytial arrangements of single-unit smooth muscles
Resting Membrane Potential
- Resting membrane potential in single-unit smooth muscle fiber is very much unstable.
- It ranges between −50 mV and −75 mV.
- Sometimes, it reaches a low level of −25 mV.
Cause for Unstable Resting Membrane Potential
- Unstable resting membrane potential is caused by appearance of some wave-like fluctuations called slow waves.
- Slow waves occur in a rhythmic fashion at a frequency of 4 to 10 per minute with an amplitude of 10 to 15 mV.
- Slow-wave rhythm may be due to rhythmic modulations in the activity of sodium-potassium pump.
- Slow wave is not an action potential and cannot cause contraction of the muscle.
- It initiates the action potential.
10. ACTION POTENTIAL IN SINGLE-UNIT SMOOTH MUSCLE
Three types of action potential occur in single-unit smooth muscle:
- Spike potential
- Spike potential initiated by slow-wave rhythm
- Action potential with plateau
1. Spike Potential
- Spike potential in single-unit smooth muscle is different from that in skeletal muscles.
- Average duration of spike potential varies between 30 and 50 milliseconds.
- Its amplitude is very low and it does not reach the isoelectric base.
- It is due to nervous and other stimuli.
- It leads to contraction of the muscle.
2. Spike Potential Initiated by Slow-Wave Rhythm
- Sometimes, slow-wave rhythm of resting membrane potential initiates spike potentials, which lead to contraction of the muscle.
- Spike potentials occur rhythmically at a rate of about one or two spikes at the peak of each slow wave.
- The slow waves cause rhythmic contractions of smooth muscles.
- This type of potential appears mostly in muscles which are self-excitatory and contract themselves without any external stimuli.
- Therefore, spike potentials initiated by slow-wave rhythm are also called pacemaker waves.
- Smooth muscles showing rhythmic contractions are present in some visceral organs such as intestine.
3. Action Potential With Plateau
- This type of action potential starts with rapid depolarization as in skeletal muscle.
- But repolarization does not occur immediately.
- Muscle remains depolarized for long periods of about 100 to 1,000 milliseconds.
- This forms the plateau (stable period) in action potential.
- This type of action potential is responsible for sustained contraction of smooth muscle fibers.
- After long depolarized state, slow repolarization occurs.
Tonic Contraction of Smooth Muscle Without Action Potential
- Smooth muscles of some visceral organs maintain a state of partial contraction called tonus or tone.
- Tonic contraction of the muscle occurs without any action potential or any stimulus.
- Sometimes, tonic contraction occurs due to action of some hormones.
11. IONIC BASIS OF ACTION POTENTIAL
In Skeletal Muscle
- Depolarization occurs due to opening of sodium channels.
- Sodium ions enter from extracellular fluid into the muscle fiber.
In Single-Unit Smooth Muscle
- Depolarization is due to entry of calcium ions rather than sodium ions.
- Calcium channels open and close slowly.
- This is responsible for prolonged potential with plateau in smooth muscles.
- Calcium ions play an important role during contraction of smooth muscle.
12. ELECTRICAL ACTIVITY IN MULTIUNIT SMOOTH MUSCLE
- Electrical activity in multiunit smooth muscle is different from that in single-unit smooth muscle.
- Electrical changes leading to contraction of multiunit smooth muscle are triggered by nervous stimuli.
- Nerve endings secrete neurotransmitters such as:
- Acetylcholine
- Noradrenaline
- Neurotransmitters depolarize the membrane of smooth muscle fiber slightly, leading to contraction.
- Such depolarization does not develop action potential.
- This type of depolarization is called local depolarization of junctional potential.
- Local depolarization travels throughout the entire smooth muscle fiber and causes contraction.
- Local depolarization is developed because the multiunit smooth muscle fibers are too small to develop action potential.
13. CONTRACTILE PROCESS IN SMOOTH MUSCLE
- Compared to skeletal muscles, contraction and relaxation processes in smooth muscles are slow.
- Process of excitation and contraction is very slow in smooth muscles because of poor development of T-tubules (sarcoplasmic reticulum).
- Therefore, calcium ions needed for excitation-contraction coupling must be obtained from the extracellular fluid.
- This makes the process of excitation-contraction coupling slow.
Calcium-Calmodulin Complex
- Stimulation of ATPase activity of myosin in smooth muscle is different from that in skeletal muscle.
- In smooth muscle, myosin has to be phosphorylated for activation of myosin ATPase.
Phosphorylation of Myosin
- Calcium, which enters the sarcoplasm from extracellular fluid, combines with a protein calmodulin and forms calcium-calmodulin complex.
- This complex activates an enzyme called calmodulin-dependent myosin light chain kinase.
- This enzyme in turn causes phosphorylation of myosin followed by activation of myosin ATPase.
- Now, the sliding of actin filaments starts.
- Phosphorylated myosin gets attached to the actin molecule for longer period.
- This is called latch-bridge mechanism.
- It is responsible for sustained contraction of the muscle with expenditure of little energy.
- Relaxation of the muscle occurs due to dissociation of calcium-calmodulin complex.
14. LENGTH-TENSION RELATIONSHIP: PLASTICITY
- Smooth muscle fibers have the property of plasticity.
- Plasticity → adaptability of smooth muscle fibers to a wide range of lengths.
- If the smooth muscle fiber is stretched, it adapts to this new length and contracts when stimulated.
- This adaptability exists to a wide range of lengths.
Significance of Plasticity
- Tension produced by muscle fiber is not directly proportional to resting length of the muscle fiber.
- In other words, Starling’s law is not applicable to smooth muscle.
- In skeletal and cardiac muscles, Starling’s law is applicable and the force of contraction is directly proportional to the initial length of the muscle fibers.
15. NEUROMUSCULAR JUNCTION IN SMOOTH MUSCLE
- Well-defined neuromuscular junctions are absent in smooth muscle.
- Nerve fibers (axons) do not end in the form of terminal branches.
- Instead, these nerve fibers end on smooth muscle fibers in three different ways.
Three Types of Nerve Endings on Smooth Muscle Fibers
-
In some smooth muscles, nerve fibers diffuse on the sheet of smooth muscle fibers without making any direct contact with the muscle.
- The diffused nerve fibers form diffused junctions which contain neurotransmitters.
- In some smooth muscle fibers, axon terminal ends in the form of many varicosities which contain the neurotransmitter.
-
In some of the multiunit smooth muscle fibers, a synaptic cleft is present between the varicosities and the membrane of smooth muscle fibers, which resembles the contact junction in skeletal muscle.
- This gap is a junction and it functions as a neuromuscular junction of skeletal muscle.
16. CONTROL OF SMOOTH MUSCLE ACTIVITIES
- Activities of smooth muscle fibers are controlled by both:
- Nervous factors
- Humoral factors
Nervous Factors
- Single-unit smooth muscle and multiunit smooth muscle are innervated by nerves of both divisions of the autonomic nervous system.
- All these nerves initiate contraction of multiunit smooth muscles only.
- Nerves supplying single-unit smooth muscles regulate the rate and force of contraction.
Humoral Factors
- Activity of smooth muscle is also controlled by humoral factors which include:
- Hormones
- Neurotransmitters
- Other humoral factors
- Action of hormones and neurotransmitters depends upon the type of receptors present in the membrane of smooth muscle fibers in particular area.
- There are two types of receptors:
- Excitatory receptors
- Inhibitory receptors
- If excitatory receptors are present, hormones or neurotransmitters contract the muscle by producing depolarization.
- If inhibitory receptors are present, hormones or neurotransmitters relax the muscle by producing hyperpolarization.
Humoral Factors Which Cause Contraction of Smooth Muscles
- Acetylcholine
- Antidiuretic hormone (ADH)
- Adrenaline
- Angiotensin I, II and IV
- Endothelin
- Histamine
- Oxytocin
- Serotonin
Humoral Factors Which Cause Relaxation of Smooth Muscles
- Lack of oxygen
- Excess carbon dioxide
- Increase in hydrogen ion concentration
- Adenosine
- Lactic acid
- Excess potassium ion
- Decrease in calcium
- Nitric oxide (NO), the endothelium-derived relaxing factor (EDRF)