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Smooth muscle
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Smooth muscle tissue
Layers of Esophageal Wall:
Mucosa
Submucosa
Muscularis
Adventitia
Striated muscle
Striated and smooth
Smooth muscle
Lamina muscularis mucosae
Esophageal glands
Details
Identifiers
Latin
textus muscularis levis; textus muscularis nonstriatus
MeSH
D009130
TH
H2.00.05.1.00001
FMA
14070
Anatomical terminology
[edit on Wikidata]
Smooth muscle is an involuntary non-striated muscle. It is divided into two subgroups; the single-unit (unitary) and multiunit smooth muscle. Within single-unit cells, the whole bundle or sheet contracts as a syncytium.
Smooth muscle cells are found in the walls of hollow organs, including the stomach, intestines, urinary bladder and uterus, and in the walls of passageways, such as the arteries and veins of the circulatory system, and the tracts of the respiratory, urinary, and reproductive systems. These cells are also present in the eyes and are able to change the size of the iris and alter the shape of the lens. In the skin, smooth muscle cells cause hair to stand erect in response to cold temperature or fear.[1]
Contents
1 Structure
1.1 Molecular structure
1.1.1 Myosin
1.1.2 Actin
1.1.3 Other proteins of the contractile apparatus
1.1.4 Other tensile structures
1.1.5 Caveolae
2 Excitation-contraction coupling
2.1 Inducing stimuli and factors
2.1.1 External substances
2.2 Spread of impulse
2.3 Contraction
2.3.1 Phasic or tonic
2.3.2 Activation of myosin heads
2.3.3 Sustained maintenance
2.3.4 Other contractile mechanisms
2.4 Relaxation
3 Invertebrate smooth muscle
4 Specific effects
5 Growth and rearrangement
6 Related diseases
7 See also
8 References
9 External links
Structure[edit]
The dense bodies and intermediate filaments are networked through the sarcoplasm, which cause the muscle fiber to contract.
A series of axon-like swelling, called varicosities or "boutons", from autonomic neurons form motor units through the smooth muscle.
Most smooth muscle is of the single-unit variety, that is, either the whole muscle contracts or the whole muscle relaxes, but there is multiunit smooth muscle in the trachea, the large elastic arteries, and the iris of the eye. Single unit smooth muscle, however, is most common and lines blood vessels (except large elastic arteries), the urinary tract, and the digestive tract.
However, the terms single- and multi-unit smooth muscle represents an oversimplification. This is due to the fact that smooth muscles for the most part are controlled and influenced by a combination of different neural elements. In addition, it has been observed that most of the time there will be some cell to cell communication and activators/ inhibitors produced locally. This leads to a somewhat coordinated response even in multiunit smooth muscle.[2]
Smooth muscle is fundamentally different from skeletal muscle and cardiac muscle in terms of structure, function, regulation of contraction, and excitation-contraction coupling.
Smooth muscle cells known as myocytes, have a fusiform shape and, like striated muscle, can tense and relax. However, smooth muscle tissue tends to demonstrate greater elasticity and function within a larger length-tension curve than striated muscle. This ability to stretch and still maintain contractility is important in organs like the intestines and urinary bladder. In the relaxed state, each cell is spindle-shaped, 20–500 micrometers in length.
Molecular structure[edit]
A substantial portion of the volume of the cytoplasm of smooth muscle cells are taken up by the molecules myosin and actin,[3] which together have the capability to contract, and, through a chain of tensile structures, make the entire smooth muscle tissue contract with them.
Myosin[edit]
Myosin is primarily class II in smooth muscle.[4]
Myosin II contains two heavy chains which constitute the head and tail domains. Each of these heavy chains contains the N-terminal head domain, while the C-terminal tails take on a coiled-coil morphology, holding the two heavy chains together (imagine two snakes wrapped around each other, such as in a caduceus). Thus, myosin II has two heads. In smooth muscle, there is a single gene (MYH11[5]) that codes for the heavy chains myosin II, but there are splice variants of this gene that result in four distinct isoforms.[4] Also, smooth muscle may contain MHC that is not involved in contraction, and that can arise from multiple genes.[4]
Myosin II also contains 4 light chains, resulting in 2 per head, weighing 20 (MLC20) and 17 (MLC17) kDa.[4] These bind the heavy chains in the "neck" region between the head and tail.
The MLC20 is also known as the regulatory light chain and actively participates in muscle contraction.[4] Two MLC20 isoforms are found in smooth muscle, and they are encoded by different genes, but only one isoform participates in contraction.
The MLC17 is also known as the essential light chain.[4] Its exact function is unclear, but it's believed that it contributes to the structural stability of the myosin head along with MLC20.[4] Two variants of MLC17 (MLC17a/b) exist as a result of alternative splicing at the MLC17 gene.[4]
Different combinations of heavy and light chains allow for up to hundreds of different types of myosin structures, but it is unlikely that more than a few such combinations are actually used or permitted within a specific smooth muscle bed.[4] In the uterus, a shift in myosin expression has been hypothesized to avail for changes in the directions of uterine contractions that are seen during the menstrual cycle.[4]
Actin[edit]
The thin filaments that form part of the contractile machinery are predominantly composed of α- and γ-actin.[4] Smooth muscle α-actin (alpha actin) is the predominant isoform within smooth muscle. There are also lots of actin (mainly β-actin) that does not take part in contraction, but that polymerizes just below the plasma membrane in the presence of a contractile stimulant and may thereby assist in mechanical tension.[4] Alpha actin is also expressed as distinct genetic isoforms such as smooth muscle, cardiac muscle and skeletal muscle specific isoforms of alpha actin.[6]
The ratio of actin to myosin is between 2:1[4] and 10:1[4] in smooth muscle. Conversely, from a mass ratio standpoint (as opposed to a molar ratio), myosin is the dominant protein in striated skeletal muscle with the actin to myosin ratio falling in the 1:2 to 1:3 range. A typical value for healthy young adults is 1:2.2.;;[7][8][9][10]
Other proteins of the contractile apparatus[edit]
Smooth muscle does not contain the protein troponin; instead calmodulin (which takes on the regulatory role in smooth muscle), caldesmon and calponin are significant proteins expressed within smooth muscle.
Tropomyosin is present in smooth muscle, spanning seven actin monomers and is laid out end to end over the entire length of the thin filaments. In striated muscle, tropomyosin serves to block actin–myosin interactions until calcium is present, but in smooth muscle, its function is unknown.[4]
Calponin molecules may exist in equal number as actin, and has been proposed to be a load-bearing protein.[4]
Caldesmon has been suggested to be involved in tethering actin, myosin and tropomyosin, and thereby enhance the ability of smooth muscle to maintain tension.[4]
Also, all three of these proteins may have a role in inhibiting the ATPase activity of the myosin complex that otherwise provides energy to fuel muscle contraction.[4]
Other tensile structures[edit]
The myosin and actin are the contractile parts of continuous chains of tensile structures that stretch both across and between smooth muscle cells.
The actin filaments of contractile units are attached to dense bodies. Dense bodies are rich in α-actinin,[4] and also attach intermediate filaments (consisting largely of vimentin and desmin), and thereby appear to serve as anchors from which the thin filaments can exert force.[4] Dense bodies also are associated with β-actin, which is the type found in the cytoskeleton, suggesting that dense bodies may coordinate tensions from both the contractile machinery and the cytoskeleton.[4] Dense bodies appear darker under an electron microscope, and so they are sometimes described as electron dense.[11]
The intermediate filaments are connected to other intermediate filaments via dense bodies, which eventually are attached to adherens junctions (also called focal adhesions) in the cell membrane of the smooth muscle cell, called the sarcolemma. The adherens junctions consist of large number of proteins including α-actinin, vinculin and cytoskeletal actin.[4] The adherens junctions are scattered around dense bands that are circumfering the smooth muscle cell in a rib-like pattern.[3] The dense band (or dense plaques) areas alternate with regions of membrane containing numerous caveolae. When complexes of actin and myosin contract, force is transduced to the sarcolemma through intermediate filaments attaching to such dense bands.
During contraction, there is a spatial reorganization of the contractile machinery to optimize force development.[4] part of this reorganization consists of vimentin being phosphorylated at Ser56 by a p21 activated kinase, resulting in some disassembly of vimentin polymers.[4]
Also, the number of myosin filaments is dynamic between the relaxed and contracted state in some tissues as the ratio of actin to myosin changes, and the length and number of myosin filaments change.
Smooth muscle cells have been observed contracting in a spiral corkscrew fashion, and contractile proteins have been observed organizing into zones of actin and myosin along the axis of the cell.
Smooth muscle-containing tissue needs to be stretched often, so elasticity is an important attribute of smooth muscle. Smooth muscle cells may secrete a complex extracellular matrix containing collagen (predominantly types I and III), elastin, glycoproteins, and proteoglycans. Smooth muscle also has specific elastin and collagen receptors to interact with these proteins of the extracellular matrix. These fibers with their extracellular matrices contribute to the viscoelasticity of these tissues. For example, the great arteries are viscolelastic vessels that act like a Windkessel, propagating ventricular contraction and smoothing out the pulsatile flow, and the smooth muscle within the tunica media contributes to this property.
Caveolae[edit]
The sarcolemma also contains caveolae, which are microdomains of lipid rafts specialized to cell signaling events and ion channels. These invaginations in the sarcoplasma contain a host of receptors (prostacyclin, endothelin, serotonin, muscarinic receptors, adrenergic receptors), second messenger generators (adenylate cyclase, phospholipase C), G proteins (RhoA, G alpha), kinases (rho kinase-ROCK, protein kinase C, protein Kinase A), ion channels (L type calcium channels, ATP sensitive potassium channels, calcium sensitive potassium channels) in close proximity. The caveolae are often close to sarcoplasmic reticulum or mitochondria, and have been proposed to organize signaling molecules in the membrane.
Excitation-contraction coupling[edit]
A smooth muscle is excited by external stimuli, which causes contraction. Each step is further detailed below.
Inducing stimuli and factors[edit]
Smooth muscle may contract spontaneously (via ionic channel dynamics) or as in the gut special pacemakers cells interstitial cells of Cajal produce rhythmic contractions. Also, contraction, as well as relaxation, can be induced by a number of physiochemical agents (e.g., hormones, drugs, neurotransmitters – particularly from the autonomic nervous system).
Smooth muscle in various regions of the vascular tree, the airway and lungs, kidneys and vagina is different in their expression of ionic channels, hormone receptors, cell-signaling pathways, and other proteins that determine function.
External substances[edit]
For instance, blood vessels in skin, gastrointestinal system, kidney and brain respond to norepinephrine and epinephrine (from sympathetic stimulation or the adrenal medulla) by producing vasoconstriction (this response is mediated through alpha-1 adrenergic receptors). However, blood vessels within skeletal muscle and cardiac muscle respond to these catecholamines producing vasodilation because the smooth muscle possess beta-adrenergic receptors. So there is a difference in the distribution of the various adrenergic receptors that explains the difference in why blood vessels from different areas respond to the same agent norepinephrine/epinephrine differently as well as differences due to varying amounts of these catecholamines that are released and sensitivities of various receptors to concentrations.
Generally, arterial smooth muscle responds to carbon dioxide by producing vasodilation, and responds to oxygen by producing vasoconstriction. Pulmonary blood vessels within the lung are unique as they vasodilate to high oxygen tension and vasoconstrict when it falls. Bronchiole, smooth muscle that line the airways of the lung, respond to high carbon dioxide producing vasodilation and vasoconstrict when carbon dioxide is low. These responses to carbon dioxide and oxygen by pulmonary blood vessels and bronchiole airway smooth muscle aid in matching perfusion and ventilation within the lungs. Further different smooth muscle tissues display extremes of abundant to little sarcoplasmic reticulum so excitation-contraction coupling varies with its dependence on intracellular or extracellular calcium.[citation needed]
Recent research indicates that sphingosine-1-phosphate (S1P) signaling is an important regulator of vascular smooth muscle contraction. When transmural pressure increases, sphingosine kinase 1 phosphorylates sphingosine to S1P, which binds to the S1P2 receptor in plasma membrane of cells. This leads to a transient increase in intracellular calcium, and activates Rac and Rhoa signaling pathways. Collectively, these serve to increase MLCK activity and decrease MLCP activity, promoting muscle contraction. This allows arterioles to increase resistance in response to increased blood pressure and thus maintain constant blood flow. The Rhoa and Rac portion of the signaling pathway provides a calcium-independent way to regulate resistance artery tone.[12]
Spread of impulse[edit]
To maintain organ dimensions against force, cells are fastened to one another by adherens junctions. As a consequence, cells are mechanically coupled to one another such that contraction of one cell invokes some degree of contraction in an adjoining cell. Gap junctions couple adjacent cells chemically and electrically, facilitating the spread of chemicals (e.g., calcium) or action potentials between smooth muscle cells. Single unit smooth muscle displays numerous gap junctions and these tissues often organize into sheets or bundles which contract in bulk.
Contraction[edit]
Smooth muscle contraction is caused by the sliding of myosin and actin filaments (a sliding filament mechanism) over each other. The energy for this to happen is provided by the hydrolysis of ATP. Myosin functions as an ATPase utilizing ATP to produce a molecular conformational change of part of the myosin and produces movement. Movement of the filaments over each other happens when the globular heads protruding from myosin filaments attach and interact with actin filaments to form crossbridges. The myosin heads tilt and drag along the actin filament a small distance (10–12 nm). The heads then release the actin filament and then changes angle to relocate to another site on the actin filament a further distance (10–12 nm) away. They can then re-bind to the actin molecule and drag it along further. This process is called crossbridge cycling and is the same for all muscles (see muscle contraction). Unlike cardiac and skeletal muscle, smooth muscle does not contain the calcium-binding protein troponin. Contraction is initiated by a calcium-regulated phosphorylation of myosin, rather than a calcium-activated troponin system.
Crossbridge cycling causes contraction of myosin and actin complexes, in turn causing increased tension along the entire chains of tensile structures, ultimately resulting in contraction of the entire smooth muscle tissue.
Phasic or tonic[edit]
Smooth muscle may contract phasically with rapid contraction and relaxation, or tonically with slow and sustained contraction. The reproductive, digestive, respiratory, and urinary tracts, skin, eye, and vasculature all contain this tonic muscle type. This type of smooth muscle can maintain force for prolonged time with only little energy utilization. There are differences in the myosin heavy and light chains that also correlate with these differences in contractile patterns and kinetics of contraction between tonic and phasic smooth muscle.
Activation of myosin heads[edit]
Crossbridge cycling cannot occur until the myosin heads have been activated to allow crossbridges to form. When the light chains are phosphorylated, they become active and will allow contraction to occur. The enzyme that phosphorylates the light chains is called myosin light-chain kinase (MLCK), also called MLC20 kinase.[4] In order to control contraction, MLCK will work only when the muscle is stimulated to contract. Stimulation will increase the intracellular concentration of calcium ions. These bind to a molecule called calmodulin, and form a calcium-calmodulin complex. It is this complex that will bind to MLCK to activate it, allowing the chain of reactions for contraction to occur.
Activation consists of phosphorylation of a serine on position 19 (Ser19) on the MLC20 light chain, which causes a conformational change that increases the angle in the neck domain of the myosin heavy chain,[4] which corresponds to the part of the cross-bridge cycle where the myosin head is unattached to the actin filament and relocates to another site on it. After attachment of the myosin head to the actin filament, this serine phosphorylation also activates the ATPase activity of the myosin head region to provide the energy to fuel the subsequent contraction.[4] Phosphorylation of a threonine on position 18 (Thr18) on MLC20 is also possible and may further increase the ATPase activity of the myosin complex.[4]
Sustained maintenance[edit]
Phosphorylation of the MLC20 myosin light chains correlates well with the shortening velocity of smooth muscle. During this period there is a rapid burst of energy utilization as measured by oxygen consumption. Within a few minutes of initiation the calcium level markedly decrease, MLC20 myosin light chains phosphorylation decreases, and energy utilization decreases and the muscle can relax. Still, smooth muscle has the ability of sustained maintenance of force in this situation as well. This sustained phase has been attributed to certain myosin crossbridges, termed latch-bridges, that are cycling very slowly, notably slowing the progression to the cycle stage whereby dephosphorylated myosin detaches from the actin, thereby maintaining the force at low energy costs.[4] This phenomenon is of great value especially for tonically active smooth muscle.[4]
Isolated preparations of vascular and visceral smooth muscle contract with depolarizing high potassium balanced saline generating a certain amount of contractile force. The same preparation stimulated in normal balanced saline with an agonist such as endothelin or serotonin will generate more contractile force. This increase in force is termed calcium sensitization. The myosin light chain phosphatase is inhibited to increase the gain or sensitivity of myosin light chain kinase to calcium. There are number of cell signalling pathways believed to regulate this decrease in myosin light chain phosphatase: a RhoA-Rock kinase pathway, a Protein kinase C-Protein kinase C potentiation inhibitor protein 17 (CPI-17) pathway, telokin, and a Zip kinase pathway. Further Rock kinase and Zip kinase have been implicated to directly phosphorylate the 20kd myosin light chains.
Other contractile mechanisms[edit]
Other cell signaling pathways and protein kinases (Protein kinase C, Rho kinase, Zip kinase, Focal adhesion kinases) have been implicated as well and actin polymerization dynamics plays a role in force maintenance. While myosin light chain phosphorylation correlates well with shortening velocity, other cell signaling pathways have been implicated in the development of force and maintenance of force. Notably the phosphorylation of specific tyrosine residues on the focal adhesion adapter protein-paxillin by specific tyrosine kinases has been demonstrated to be essential to force development and maintenance. For example, cyclic nucleotides can relax arterial smooth muscle without reductions in crossbridge phosphorylation, a process termed force suppression. This process is mediated by the phosphorylation of the small heat shock protein, hsp20, and may prevent phosphorylated myosin heads from interacting with actin.
Relaxation[edit]
The phosphorylation of the light chains by MLCK is countered by a myosin light-chain phosphatase, which dephosphorylates the MLC20 myosin light chains and thereby inhibits contraction.[4] Other signaling pathways have also been implicated in the regulation actin and myosin dynamics. In general, the relaxation of smooth muscle is by cell-signaling pathways that increase the myosin phosphatase activity, decrease the intracellular calcium levels, hyperpolarize the smooth muscle, and/or regulate actin and myosin muscle can be mediated by the endothelium-derived relaxing factor-nitric oxide, endothelial derived hyperpolarizing factor (either an endogenous cannabinoid, cytochrome P450 metabolite, or hydrogen peroxide), or prostacyclin (PGI2). Nitric oxide and PGI2 stimulate soluble guanylate cyclase and membrane bound adenylate cyclase, respectively. The cyclic nucleotides (cGMP and cAMP) produced by these cyclases activate Protein Kinase G and Protein Kinase A and phosphorylate a number of proteins. The phosphorylation events lead to a decrease in intracellular calcium (inhibit L type Calcium channels, inhibits IP3 receptor channels, stimulates sarcoplasmic reticulum Calcium pump ATPase), a decrease in the 20kd myosin light chain phosphorylation by altering calcium sensitization and increasing myosin light chain phosphatase activity, a stimulation of calcium sensitive potassium channels which hyperpolarize the cell, and the phosphorylation of amino acid residue serine 16 on the small heat shock protein (hsp20)by Protein Kinases A and G. The phosphorylation of hsp20 appears to alter actin and focal adhesion dynamics and actin-myosin interaction, and recent evidence indicates that hsp20 binding to 14-3-3 protein is involved in this process. An alternative hypothesis is that phosphorylated Hsp20 may also alter the affinity of phosphorylated myosin with actin and inhibit contractility by interfering with crossbridge formation. The endothelium derived hyperpolarizing factor stimulates calcium sensitive potassium channels and/or ATP sensitive potassium channels and stimulate potassium efflux which hyperpolarizes the cell and produces relaxation.
Invertebrate smooth muscle[edit]
In invertebrate smooth muscle, contraction is initiated with the binding of calcium directly to myosin and then rapidly cycling cross-bridges, generating force. Similar to the mechanism of vertebrate smooth muscle, there is a low calcium and low energy utilization catch phase. This sustained phase or catch phase has been attributed to a catch protein that has similarities to myosin light-chain kinase and the elastic protein-titin called twitchin. Clams and other bivalve mollusks use this catch phase of smooth muscle to keep their shell closed for prolonged periods with little energy usage.
Specific effects[edit]
Although the structure and function is basically the same in smooth muscle cells in different organs, their specific effects or end-functions differ.
The contractile function of vascular smooth muscle regulates the lumenal diameter of the small arteries-arterioles called resistance vessels, thereby contributing significantly to setting the level of blood pressure and blood flow to vascular beds. Smooth muscle contracts slowly and may maintain the contraction (tonically) for prolonged periods in blood vessels, bronchioles, and some sphincters. Activating arteriole smooth muscle can decrease the lumenal diameter 1/3 of resting so it drastically alters blood flow and resistance. Activation of aortic smooth muscle doesn't significantly alter the lumenal diameter but serves to increase the viscoelasticity of the vascular wall.
In the digestive tract, smooth muscle contracts in a rhythmic peristaltic fashion, rhythmically forcing foodstuffs through the digestive tract as the result of phasic contraction.
A non-contractile function is seen in specialized smooth muscle within the afferent arteriole of the juxtaglomerular apparatus, which secretes renin in response to osmotic and pressure changes, and also it is believed to secrete ATP in tubuloglomerular regulation of glomerular filtration rate. Renin in turn activates the renin–angiotensin system to regulate blood pressure.
Growth and rearrangement[edit]
The mechanism in which external factors stimulate growth and rearrangement is not yet fully understood. A number of growth factors and neurohumoral agents influence smooth muscle growth and differentiation. The Notch receptor and cell-signaling pathway have been demonstrated to be essential to vasculogenesis and the formation of arteries and veins. The proliferation is implicated in the pathogenesis of atherosclerosis and is inhibited by nitric oxide.
The embryological origin of smooth muscle is usually of mesodermal origin, after the creation of muscle fibers in a process known as myogenesis. However, the smooth muscle within the Aorta and Pulmonary arteries (the Great Arteries of the heart) is derived from ectomesenchyme of neural crest origin, although coronary artery smooth muscle is of mesodermal origin.
Related diseases[edit]
"Smooth muscle condition" is a condition in which the body of a developing embryo does not create enough smooth muscle for the gastrointestinal system. This condition is fatal.
Anti-smooth muscle antibodies (ASMA) can be a symptom of an auto-immune disorder, such as hepatitis, cirrhosis, or lupus.
Vascular smooth muscle tumors are very rare. They can be malignant or benign, and morbidity can be significant with either type. Intravascular leiomyomatosis is a benign neoplasm that extends through the veins; angioleiomyoma is a benign neoplasm of the extremities; vascular leiomyosarcomas is a malignant neoplasm that can be found in the inferior vena cava, pulmonary arteries and veins, and other peripheral vessels. See Atherosclerosis.
See also[edit]
Atromentin has been shown to be a smooth muscle stimulant.[13]
Skeletal muscle
Cardiac muscle
References[edit]
Jump up ^ "10.8 Smooth Muscle – Anatomy and Physiology". opentextbc.ca. Archived from the original on 1 February 2018. Retrieved 28 April 2018.
Jump up ^ Berne & Levy. Physiology, 6th Edition
^ Jump up to: a b p. 174 in: The vascular smooth muscle cell: molecular and biological responses to the extracellular matrix. Authors: Stephen M. Schwartz, Robert P. Mecham. Editors: Stephen M. Schwartz, Robert P. Mecham. Contributors: Stephen M. Schwartz, Robert P. Mecham. Publisher: Academic Press, 1995. ISBN 0-12-632310-0, 978-0-12-632310-8
^ Jump up to: a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae Aguilar HN, Mitchell BF (2010). "Physiological pathways and molecular mechanisms regulating uterine contractility". Hum. Reprod. Update. 16 (6): 725–44. doi:10.1093/humupd/dmq016. PMID 20551073.
Jump up ^ Matsuoka R, Yoshida MC, Furutani Y, Imamura S, Kanda N, Yanagisawa M, Masaki T, Takao A (1993). "Human smooth muscle myosin heavy chain gene mapped to chromosomal region 16q12". Am. J. Med. Genet. 46 (1): 61–67. doi:10.1002/ajmg.1320460110. PMID 7684189.
Jump up ^ Perrin BJ, Ervasti JM (2010). "The actin gene family: function follows isoform". Cytoskeleton. 67 (10): 630–34. doi:10.1002/cm.20475. PMC 2949686. PMID 20737541.
Jump up ^ Aguilar_2010 (above reference) "In skeletal or striated muscle, there is 3-fold more myosin than actin."
Jump up ^ Trappe S, Gallagher P, et al. Single muscle fibre contractile properties in young and old men and women. J Physiol (2003), 552.1, pp. 47–58, Table 8
Jump up ^ Greger R, Windhorst U; Comprehensive Human Physiology, Vol. II. Berlin, Springer, 1996; Chapter 46, Table 46.1, Myosin 45%, Actin 22% of skeletal muscle myofibrillar proteins, p. 937
Jump up ^ Lawrie's Meat Science, Lawrie RA, Ledward, D; 2014; Chapter 4, Table 4.1, Chemical Composition of Typical Mammalian Adult Muscle, percent of skeletal muscle tissue wet weight; myosin 5.5%, actin 2.5%, p. 76
Jump up ^ Ultrastructure of Smooth Muscle, Volume 8 of Electron Microscopy in Biology and Medicine, Editor P. Motta, Springer Science & Business Media, 2012, ISBN 1461306833, 9781461306832. (p. 163 Archived 2017-05-10 at the Wayback Machine.)
Jump up ^ Scherer EQ, Lidington D, Oestreicher E, Arnold W, Pohl U, Bolz SS (2006). "Sphingosine-1-phosphate modulates spiral modiolar artery tone: A potential role in vascular-based inner ear pathologies?". Cardiovasc. Res. 70 (1): 79–87. doi:10.1016/j.cardiores.2006.01.011. PMID 16533504.
Jump up ^ Sullivan G, Guess WL (1969). "Atromentin: a smooth muscle stimulant in Clitocybe subilludens". Lloydia. 32 (1): 72–75. PMID 5815216.
External links[edit]
BBC – baby born with smooth muscle condition has 8 organs transplanted
Smooth muscle antibody
Stomach smooth muscle identified using antibody
UIUC Histology Subject 265
Histology at KUMC muscular-muscle08 "Smooth Muscle"
Histology image: 21701ooa – Histology Learning System at Boston University
Smooth muscle histology photomicrographs
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Padmé Amidala
From Wikipedia, the free encyclopedia
Jump to: navigation, search
"Padmé" redirects here. For the film, see Padmé (film).
Padmé Amidala
Star Wars character
Natalie Portman as Padmé Amidala in Star Wars: Episode III – Revenge of the Sith
First appearance
The Phantom Menace (1999)
Created by
George Lucas
Portrayed by
Natalie Portman
Voiced by
Catherine Taber (Star Wars: The Clone Wars film and TV series, Star Wars: The Clone Wars – Jedi Alliance, Star Wars: The Clone Wars – Republic Heroes, Disney Infinity 3.0 and Star Wars Forces of Destiny[1])
Grey DeLisle (Star Wars: Clone Wars, Star Wars Episode I: The Phantom Menace (video game), Star Wars Episode I: Jedi Power Battles, Star Wars: Galactic Battlegrounds, Star Wars: Super Bombad Racing and Star Wars: The Clone Wars)
Montana Norberg (Lego Star Wars: Droid Tales)
Information
Occupation
Queen of Naboo
Senator of Naboo
Co-founder of the Rebel Alliance
Affiliation
Galactic Senate of the Grand Republic
Alliance to Restore the Republic
Family
Jobal Naberrie (mother)
Ruwee Naberrie (father)
Sola Naberrie (sister)
Spouse(s)
Anakin Skywalker
Children
Luke Skywalker
Leia Organa
Relatives
Canon:
Han Solo (son-in-law)
Ben Solo (grandson)
Legends:
Mara Jade (daughter-in-law)
Ben Skywalker (grandson)
Jacen Solo (grandson)
Jaina Solo (granddaughter)
Anakin Solo (grandson)
Allana Solo (great-granddaughter)
Homeworld
Naboo
Padmé Amidala (née Naberrie) is a fictional character in the Star Wars franchise, appearing in the prequel trilogy portrayed by actress Natalie Portman. She served as the Princess of Theed and later Queen of Naboo. After her reign, she became a senator in the Galactic Senate, an anti-war movement spokesperson, and co-founder of the opposition-faction that later emerged as the Rebel Alliance.[2] She was secretly married to the Jedi Anakin Skywalker, and was the biological mother of Luke Skywalker and Leia Organa, which makes her the mother-in-law of Han Solo, and the grandmother of Kylo Ren.
Contents
[hide]
1 Concept and creation
1.1 Casting
1.2 Costumes
1.3 Character overview
1.3.1 Style
2 Appearances
2.1 Feature films
2.1.1 Original trilogy
2.1.1.1 Return of the Jedi
2.1.2 Prequel trilogy
2.1.2.1 Episode I: The Phantom Menace
2.1.2.2 Episode II: Attack of the Clones
2.1.2.3 Episode III: Revenge of the Sith
2.2 Animated series
2.2.1 The Clone Wars
2.2.2 Forces of Destiny (2017)
2.3 Legends
2.4 Novels
2.4.1 Clone Wars (2003)
2.4.2 Novels and comics
2.5 Video Games
3 Reception
4 Family tree
5 See also
6 References
7 Further reading
8 External links
Concept and creation[edit]
Padmé Amidala cosplay at Japan Expo in February 2012 in France
In initial drafts of the Star Wars story, Luke Skywalker's and Princess Leia's mother was not well developed. According to Dale Pollock, Luke Skywalker was originally Luke Starkiller and "Leia is the daughter of Owen Lars and his wife Beru and seems to be Luke's cousin – together they visit the grave of his mother, who perished with his father on a planet destroyed by the Death Star.[3] In an interview, Lucas answered a question about the development of characters like Obi-Wan Kenobi, Luke, and Leia; their mother was not a factor:
The first [version] talked about a princess and an old general. The second version involved a father, his son, and his daughter; the daughter was the heroine of the film. Now the daughter has become Luke, Mark Hamill's character. There was also the story of two brothers where I transformed one of them into a sister. The older brother was imprisoned, and the young sister had to rescue him and bring him back to their dad.[4]
Film historian Laurent Bouzereau reports that the second draft of the Return of the Jedi screenplay contained dialogue in which Obi-Wan Kenobi explains to Luke that he has a twin sister. She and their mother were "sent to the protection of friends in a distant system. The mother died shortly thereafter, and Luke's sister was adopted by Ben's friends, the governor of Alderaan and his wife."[5] Lucas is quoted in Star Wars: The Annotated Scripts (1997) as saying:
The part that I never really developed is the death of Luke and Leia's mother. I had a backstory for her in earlier drafts, but it basically didn't survive. When I got to Jedi, I wanted one of the kids to have some kind of memory of her because she will be a key figure in the new episodes I'm writing. But I really debated whether or not Leia should remember her.[6]
Revenge of the Sith does not explain how Leia remembers her "real mother". Film critic Peter Travers of Rolling Stone applauds Lucas's attempt to link the two trilogies in Revenge of the Sith's final scenes, but says, "It's too little and too late." He explains, "To hail Revenge of the Sith as a satisfying bridge to a classic is not just playing a game of the Emperor's New Clothes, it's an insult to what the original accomplished."[7]
When Lucas drafted the script for The Phantom Menace, he envisioned a "link between Padmé and Princess Leia, the daughter who follows so closely in her footsteps."[8] According to Natalie Portman, "It definitely did come into play how strong and smart a character Carrie Fisher portrayed, because I think that a lot of that is passed on from parent to child. I think George wrote Amidala as a strong, smart character, but it helped to know that I had this great woman before me who had portrayed her character as a fiery woman."[9] Paul McDonald notes that there are "inevitable comparisons" between the two characters: "both develop soft spots for rogue pilots, and both have a knack for slipping into and out of stilted British accents."[10] Despite being diplomats, each is also the best marksman of her respective trilogy, rarely missing.
Casting[edit]
George Lucas, Rick McCallum, and casting director Robin Gurland auditioned over 200 actresses for the part of Padmé Amidala.[11] They chose 16-year-old actress Natalie Portman to play the role. According to The Phantom Menace production notes, "The role required a young woman who could be believable as the ruler of that planet, but at the same time be vulnerable and open." Portman's performances in The Professional (1994) and Beautiful Girls (1996) impressed Lucas.[12] He stated, "I was looking for someone who was young, strong, along the lines of Leia. Natalie embodied all those traits and more."[13]
Portman was a unique choice because she was unfamiliar with Star Wars. "My cousins had always been obsessed with the films, yet I hadn't even seen them before I got the part," she says. "When it all happened for me, my cousins were exclaiming, 'Oh, my God, you're in Star Wars!'"[14] She told a CNN interviewer, "I really wasn't aware of how big a deal Star Wars was ... and when I saw the films, I really liked them, but I still didn't really understand how many ... were passionate fans of this film."[15] Portman was, however, enthusiastic over being cast as the queen of Naboo, a character she expected to become a role model: "It was wonderful playing a young queen with so much power. I think it will be good for young women to see a strong woman of action who is also smart and a leader."[14]
In The Phantom Menace, Portman had to portray a character younger than herself. In Attack of the Clones, however, her character had aged 10 years. Portman had aged only five years between the two films. She remarks, "[Lucas] wants to make sure I seem older than Anakin in Attack of the Clones, so it's believable that I can be bossing him around, and he's a little intimidated. She looks at him as a little boy – at least for the first half of the film."[16]
Portman signed a contract to play Padmé in the three prequel films. Reactions by critics to her performances were largely negative. James Berardinelli called her acting in The Phantom Menace "lackluster,"[17] while Annlee Ellingson of Box Office Magazine said "Portman's delivery is stiff and flat, perhaps hindered by the gorgeous but cumbersome costumes."[18] Mike Clark of USA Today complained about Portman and Hayden Christensen, who portrayed Anakin; he wrote, "Both speak in monotone for doubly deadly effect, though when not burdened by his co-star, Christensen often finds the emotion in his limited intonations."[19] A Revenge of the Sith reviewer for The Village Voice accused "computer-generated characters like wheezing cyborg baddie General Grievous and blippeting fireplug R2-D2" of "emot[ing] more convincingly than either Natalie Portman or Hayden Christensen."[20] Nonetheless, Mick LaSalle of the San Francisco Chronicle described Portman's performance in the third episode as "decorative and sympathetic."[21]
Critics have blamed Portman's performance on Lucas' direction and script. Roger Ebert, for example, charged that in Attack of the Clones "too much of ... the film is given over to a romance between Padmé and Anakin in which they're incapable of uttering anything other than the most basic and weary romantic clichés, while regarding each other as if love was something to be endured rather than cherished."[22] He offered a similar critique for Revenge of the Sith: "To say that George Lucas cannot write a love scene is an understatement; greeting cards have expressed more passion."[23] Todd McCarthy of Variety likewise lamented that "Lucas's shortcomings as a writer and director of intimate, one-on-one scenes" hampered Portman's performance.[24]
Costumes[edit]
Traditional Mongolian royal fashion (left) and the Senate Gown (right) inspired by such fashion
An extensive wardrobe was designed for Padmé Amidala by Lucasfilm concept artists and costume designers. Like Leia Organa, one of the inspirations for Padmé was the Flash Gordon character Dale Arden. The wardrobe in The Phantom Menace was designed by concept artist Iain McCaig and costume designer Trisha Biggar; concept artist Dermot Power joined McCaig and Biggar in the design process of Attack of the Clones.[25][26] Biggar worked as costume designer on the three films.[27] Many costumes were inspired by the historical royal fashions of different cultures. For example, in The Phantom Menace, the dress which Padmé wears when addressing the Senate is based on Mongolian imperial fashion worn by Empress Dondogdulam, the wife of Bogd Khan, and other monarchs into the early 20th century. Padmé's travel gown in Attack of the Clones is based on 17th-century Russian fashion photographed on Grand Duchess Xenia Alexandrovna at the Romanov Anniversary Ball in 1903.[28]
The costumes of the prequel trilogy are purposefully more elaborate than those of the original trilogy. Lucas asserts that galactic society in the prequels is much more sophisticated.[29] Commenting on the disparities between the two trilogies, Carrie Fisher mused, "Harrison Ford wears the same outfit for three flicks, and I was complaining that I wear, like, six outfits. And my mother – Natalie Portman – she wears three million. She walks through a doorway and there's another outfit. It's like the Liberace of sci-fi changing of clothes."[30] Trisha Biggar reveals that originally there were only three costumes planned for Amidala in The Phantom Menace, but "[Lucas] decided that every time we saw [her] she was going to have a different costume."[31] Lucas explains, "Someone of that stature would automatically be changing their costumes to fit the occasion."[29]
Grand Duchess Xenia Alexandrovna of Russia (left) and the Travel Gown (right) inspired by Russian fashion (similar to a kokoshnik)
Aesthetics aside, the wardrobe was designed to reflect key plot developments. In Attack of the Clones, Lucas wanted Padmé's wardrobe to mirror the romantic elements of the film. He suggested that her costumes be more "sultry in nature."[32] Trisha Biggar notes that Lucas wanted her to appear "sexy, gorgeous, and young in skimpy clothes."[33] Portman laughs, "I got over the hump of 18 so I'm allowed to show tummy now, I guess",[34] so during the battle of Geonosis the bottom part of her top is ripped off revealing her midriff. For Revenge of the Sith, Biggar says, "We knew that Padmé was going to be pregnant through the whole film, and nobody in the outside world could know that. Because she's pregnant, I wanted a soft quality to be apparent in the fabrics that were used."[35]
Some of the costumes created by Biggar's staff did not appear in the final version of the films. In Revenge of the Sith, for example, a multi-colored "Peacock Gown" and a "Green Cut Velvet Robe" worn by Padmé in scenes featuring the Delegation of 2000 were deleted during post-production. Biggar remarks that the Peacock Gown had been one of her favorite designs and that much time and money had been invested in these particular costumes.[36] Ultimately, the Peacock Gown would be used only for the film's theatrical poster. The velvet robe was ultimately re-used for a short scene filmed during pick-up photography, thus appearing in the film, and features on the DVD cover art.
Many of Padmé's costumes in The Phantom Menace were featured in the Japanese magazine High Fashion in 1999 and the Attack of the Clones costumes were in Vogue in 2002. The costumes went on display in the 2005 exhibit Dressing A Galaxy: The Costumes of Star Wars at the Fashion Institute of Design & Merchandising (FIDM) in Los Angeles.[37] Trisha Biggar won a Saturn Award for Best Costumes in 2000 for The Phantom Menace and in 2003 for Attack of the Clones. She was nominated in 2006 for Revenge of the Sith, but lost to Isis Mussenden, costume designer for The Chronicles of Narnia: The Lion, the Witch and the Wardrobe (2005).
Character overview[edit]
Padmé Amidala is depicted in Star Wars fiction as beautiful and graceful. In Cloak of Deception, she is described as having "a slight figure and a lovely, feminine face. She was remarkably solemn for one so young. It was clear that she took her responsibilities with the utmost seriousness."[38] Terry Brooks details the alien Nute Gunray's reaction to her appearance: "She was considered beautiful, Gunray had been told, but he had no sense of human beauty and by Neimoidian standards she was simply colorless and small-featured."[39] Brooks writes that she is "young, beautiful, and serene."[40]
The Star Wars Databank describes her as "one of Naboo's best and brightest"[41] and "interested in public service".[42] She demonstrates a devotion to the disadvantaged and deprived beings of the galaxy. Her childhood and adolescence is sacrificed to public service. In the Attack of the Clones novelization, Padmé's sister Sola Naberrie tells her, "You're so tied up in your responsibilities that you don't give any weight to your desires."[43]
Padmé relies on diplomacy to resolve disputes, often appearing as a pacifist. She is not, however, an advocate of appeasement, as she is willing to use "aggressive negotiations" to preserve democracy.[44] The Star Wars Databank notes, "Despite her initial objections to a Republic army, Padmé nonetheless fought alongside the newly created clone troopers against the Separatist droid forces."[42] Film critics Dominique Mainon and James Ursini classify her as a "modern Amazon," a reference to the warrior women of ancient Greek mythology.[45]
Her combat skills are explored further through the Star Wars universe. In Star Wars: Episode II – Attack of the Clones she fights squads of battle droids with hand-to-hand combat and a blaster. She is an expert markswoman; in the Star Wars: The Clone Wars episode "Assassin", she outguns bounty hunter Aurra Sing.
As a ruler and politician, Padmé is distrustful of bureaucracy, opposed to corruption, and a firm believer in democracy and the rule of law. She tells Anakin, "Popular rule is not democracy .... It gives the people what they want, not what they need."[46] According to Mainon and Ursini, "she tried to preach compromise and reason, [but] the disarray within the [Republic] ... led her to doubt the senate's effectiveness."[42] Her loyalty remains with the Republic until she suspects it no longer represents the democratic principles she espouses. In the novelization of Revenge of the Sith, Padmé advises Senator Bail Organa and Mon Mothma, the founders of what will become the Rebel Alliance, to "Be good little Senators. Mind your manners and keep your heads down. And keep doing ... all those things we can't talk about."[47]
Padmé is sometimes mysterious and deceptive. She is described in Brooks' The Phantom Menace novelization as a "chameleon of sorts, masking herself to the world at large and finding companionship almost exclusively with a cadre of handmaidens who were always with her."[48] Her decision to quietly marry Anakin and secret discussions with other senators about Palpatine add to the character's duplicity.[42] Paul F. McDonald of Space.com observes, "Amidala ... embod[ies] many of the dualities that inform Episode I— war and peace, queen and commoner, form and substance. Unlike other characters, whose personalities are divided and usually warring against one another, her dual nature works to her advantage." He explains, "Amidala can be cold and commanding when she needs to be, or warm and loving as Padmé."[10]
In February 2018, Portman reprised the role for a interviw-Rap sketch on Saturday Night Live.
Style[edit]
In The Phantom Menace, Padmé Amidala, in her capacity as queen, is addressed as "Your Majesty", "Your Royal Highness" and "Your Highness". Contrary to usage in real monarchies, where the style is fixed and tied to the person's rank, in Lucas's Star Wars universe they are apparently interchangeable.
After her tenure as monarch ended and she became a member of the Senate, Padmé Amidala was addressed as "Senator Amidala" with no formal style, although she continued to use her regnal name instead of her family name in accordance with Naboo custom.
Appearances[edit]
Feature films[edit]
Original trilogy[edit]
Return of the Jedi[edit]
Return of the Jedi, the final film in the original trilogy, alludes to the character. While in the Ewok village on the forest moon of Endor, Luke Skywalker asks if Leia Organa remembers their "real mother". Leia answers that "she died when I was very young" and also says that "she was very beautiful... kind... but sad", while Luke confesses to have no memories of their mother. An extended scene of Return of the Jedi expands on the fate of Luke and Leia's (then-unnamed) mother. It differs from the prequels in that Luke and Leia's mother sneaked away as Bail Organa's handmaiden after Anakin Skywalker turned to the dark side of the Force, and was with Leia until the Princess turned four, making the line Leia says to Luke about remembering their mother more believable.[49]
Prequel trilogy[edit]
Episode I: The Phantom Menace[edit]
Padmé Amidala makes her first film appearance in The Phantom Menace. She is introduced as the recently elected 14-year-old queen of Naboo, dedicated to ending the planet's occupation by the Trade Federation. She attempts to deal directly with Federation Viceroy Nute Gunray, who attempts to force her to sign a treaty which would legitimize the Trade Federation's occupation of Naboo, her planet. Padmé escapes from Naboo with the help of Jedi Master Qui-Gon Jinn and Padawan Obi-Wan Kenobi, but they are forced to land on the desert planet of Tatooine. Disguised as a handmaiden, Padmé meets nine-year-old slave Anakin Skywalker and his mother Shmi. Anakin gives her a hand-carved charm on a leather necklace. She witnesses Anakin win the Podrace at the Boonta Eve Classic that both aids her mission to Coruscant and secures his freedom.
Arriving on Coruscant, Padmé consults with Senator Palpatine, who encourages her to appeal to the Senate to resolve Naboo's dispute with the Trade Federation. Palpatine persuades her to make a motion in the Senate to have Supreme Chancellor Finis Valorum removed from office, which later enables Palpatine to be elected in his place. (She is unaware that he is in fact the Sith lord Darth Sidious, and is responsible for the crisis.) Padmé returns to Naboo to fight for her planet's freedom, enlisting the aid of Jar Jar Binks's Gungan warriors and having the handmaiden Sabé pose as her. As Sabé attempts a peace deal between Naboo and the Gungans, Padmé intervenes and reveals her true identity. The Gungans agree to help and offer a diversion to lure the droid armies away from the palace. Once in the palace, Padmé's forces storm the throne room and capture the Viceroy, ending the trade blockade of Naboo once and for all. A celebration is held to announce the unity between Naboo and the Gungans.
Episode II: Attack of the Clones[edit]
Padmé Amidala makes her second film appearance in Attack of the Clones, which is set a decade later. Now holding office as Senator following her term as queen, she represents Naboo in the Galactic Senate and leads a faction opposed to the Military Creation Act that would create an army of clones for the Republic, which has been threatened by a growing Separatist movement. As she arrives on Coruscant to cast her vote, assassins hired by the Trade Federation make an unsuccessful attempt on her life. Jedi Knight Obi-Wan Kenobi and Padawan Anakin Skywalker are assigned to protect Padmé. Chancellor Palpatine sends Padmé into hiding on Naboo, where she and Anakin struggle to maintain a platonic relationship despite their obvious mutual attraction. In a deleted scene, she introduces Anakin to her parents (Ruwee and Jobal Naberrie) and informs him of her charitable work with the Refugee Relief Movement, a galaxy-wide disaster relief and resettlement organization.
When Anakin has a vision of his mother in danger, Padmé accompanies him to Tatooine in a failed attempt to rescue Shmi from a band of Tusken Raiders. Anakin returns with Shmi's body and tearfully confesses to Padmé that he slaughtered the entire tribe. Padmé is troubled by what Anakin has done, but nevertheless comforts him. After they receive a message from Obi-Wan on the planet Geonosis, Padmé and Anakin rush to the Jedi Knight's aid, only to be captured themselves and condemned to death in a Geonosian coliseum by Separatist leader and Sith Lord Count Dooku. They declare their love to each other, and are saved at the last minute by Jedi Masters Mace Windu and Yoda leading an army of Jedi and clone troopers, thus marking the opening salvo of the Clone Wars. Afterwards, Padmé and Anakin are married in a secret ceremony on Naboo witnessed by the droids R2-D2 and C-3PO.
Episode III: Revenge of the Sith[edit]
Padmé Amidala makes her third film appearance in Revenge of the Sith, which is set three years later. After Anakin returns from rescuing Palpatine, she informs him that she is pregnant. Padmé detects changes in Anakin after he begins having dreams about her dying in childbirth. Eventually, these fears lead to Anakin turning to the dark side of the Force and becoming Palpatine's Sith apprentice, Darth Vader. Meanwhile, Padmé watches with increasing suspicion as Palpatine uses the Clone War in control over the Senate and judiciary. In another deleted scene, Padmé is seen as a dissenter in Palpatine's government and an early constituting member of the Alliance to Restore the Republic, joined by senators Bail Organa, Mon Mothma and Bana Breemu.
As Palpatine declares martial law by transforming the Republic into the Galactic Empire, Padmé remarks to Organa: "So this is how liberty dies – with thunderous applause". After Palpatine names himself emperor for life, Obi-Wan informs Padmé that her husband has turned to the dark side and killed everyone in the Jedi Temple. Unable to believe this, Padmé travels to the volcanic planet Mustafar (with Obi-Wan a stowaway aboard her ship). Padmé begs Vader to escape Palpatine's grasp and flee with her, but Vader insists that, together, they can overthrow Palpatine and rule the galaxy. Padmé recoils in horror, but still tries to persuade him to abandon the dark side. When Obi-Wan emerges from her ship, Vader accuses Padmé of betrayal and uses the Force to choke her into unconsciousness.
After Obi-Wan defeats Vader in a lightsaber duel, he brings Padmé to the secret asteroid base Polis Massa. Despite efforts of medical droids, Padmé dies after delivering twins, Luke and Leia. Just prior to her death, Padmé insists to Obi-Wan that she knows "there is still good" in Anakin. After Padmé's body is altered to appear still pregnant and given an elaborate funeral ceremony on Naboo, her twins are separated to be hidden from Darth Vader and Emperor Palpatine: Leia is adopted by Senator Organa and his wife Queen Organa on Alderaan to be raised as a princess, while Luke is brought to Tatooine to be raised by his step-grandfather's family, Owen and Beru Lars.
Animated series[edit]
The Clone Wars[edit]
Padmé Amidala makes her fourth film appearance on Star Wars: The Clone Wars. While Anakin and his Padawan Ahsoka Tano search for Jabba the Hutt's son Rotta, Padmé meets Jabba's uncle Ziro the Hutt at his palace on Coruscant to convince him to side with the Jedi. After Ziro forcibly removes Padmé, she escapes and eavesdrops on his communication with Count Dooku and the Separatists about an elaborate scheme to kill Rotta, frame the Jedi for his murder and force Jabba to attempt revenge, leaving Ziro as the Hutts' ruler. After being discovered, Dooku suggests Ziro collect the bounty placed on her head. When battle droids confiscate Padmé's comlink and blaster, she outwits and tricks one into activating her comlink as C-3PO is attempting to contact her before a droid smashes the device. C-3PO leads a squad of Coruscant Guard troopers to rescue her. Padmé then contacts Jabba just as the Hutt is about to execute Anakin and Ahsoka, and forces Ziro to confess his betrayal to Jabba. Padmé proceeds to negotiate an alliance between the Republic and the Hutts which would allow Republic warships to use unknown Hutt hyperspace lanes.
In the subsequent TV series, Padmé is mostly portrayed working in the Senate toward a peaceful resolution to the Clone Wars, although a few episodes have portrayed her fighting the Separatists alongside Anakin, Ahsoka and Jar Jar Binks. She has appeared in seven episodes in the first season and third season, four episodes in the second season, nine episodes in the fourth season, and one episode in the fifth season. A trilogy of episodes were set with her as the main focus in which she meets with her old friend Rush Clovis; these episode were later removed, however, and set to be bonus material.
Forces of Destiny (2017)[edit]
Padmé appears in the micro-series Star Wars: Forces of Destiny, voiced again by Catherine Taber.[50]
Legends[edit]
In April 2014, most of the licensed Star Wars novels and comics produced since the originating 1977 film Star Wars were rebranded by Lucasfilm as Star Wars Legends and declared non-canon to the franchise.[51][52][53]
Novels[edit]
The novelizations of the Star Wars prequel films introduced material about Padmé Amidala that was not included in the films. Terry Brooks' Phantom Menace (1999) includes a discussion between Qui-Gon Jinn and Obi-Wan Kenobi. Qui-Gon describes the then-queen of Naboo as "something of an unknown" before the Trade Federation blockade.[54] In the Attack of the Clones (2002) adaptation by R. A. Salvatore, there is a detailed conversation between Padmé and her sister Sola Naberrie shortly after Jamillia appoints her senator. Sola chides her for ignoring her personal life: "What about Padmé Amidala? Have you even thought about what might make your life better?"[55] Matthew Stover's Revenge of the Sith (2005) elaborates upon Padmé's role in the formation of the Rebel Alliance. Stover narrates Darth Vader's reaction to the death of his wife: Vader thinks to himself, "You killed her because, finally, when you could have saved her, when you could have gone away with her, when you could have been thinking about her, you were thinking about yourself ..."[56]
Clone Wars (2003)[edit]
Padmé Amidala appears in eight chapters of the Star Wars: Clone Wars micro-series that aired on Cartoon Network from 2003 to 2005. She is secluded on Coruscant and maintains a correspondence when Anakin Skywalker is fighting in the Clone Wars. In one chapter, Padmé travels with Yoda aboard her ship when he senses a disturbance in the Force coming from the ice planet Ilum. Despite Captain Typho's protest, she accompanies Yoda and helps rescue Jedi Master Luminara Unduli and Padawan Barriss Offee. In another chapter, she is thrilled by Anakin's graduation from Padawan to Jedi Knight. In the final chapter, she is briefly seen during General Grievous's assault on Coruscant.
Novels and comics[edit]
Padmé's background prior to her appearance in the prequel films is revealed in Star Wars novels and comics. In Terry Moore's comic "A Summer's Dream" printed in Star Wars Tales 5 (2000) and set a year before the events of The Phantom Menace, Padmé is the Princess of Theed, Naboo's capital city. A young man, Ian Lago, falls in love with Padmé, but she places her duty to the people over her personal happiness and rejects him. Lago is the son of an advisor to King Veruna, the reigning monarch of Naboo.[57][41]
In the novel, Cloak of Deception (2001) by James Luceno, King Veruna is forced to abdicate the throne following accusations of corruption. Padmé is elected Queen of Naboo and contacts Palpatine to inform him that Veruna has been mysteriously killed. She and Palpatine discuss the events that lead to the Trade Federation blockade of Naboo. She admits to him, "Naboo can scarcely afford to become embroiled in a dispute that pits the Republic against the Trade Federation."[58]
Star Wars literature focuses on Padmé's career as ruling monarch of Naboo. The young adult novel Star Wars Episode I Journal: Amidala (1999) by Jude Watson focuses on Padmé Amidala's early career and narrow escape from the Trade Federation.[59] The Queen's Amulet (1999) by Julianne Balmain narrates the close friendship between Padmé and her handmaiden Sabé immediately before the events of The Phantom Menace.[60] Erik Tiemens's comic "The Artist of Naboo" details the story of a young, unnamed artist on Naboo who becomes captivated by Padmé's beauty. The artist features her in a series of paintings and later risks his life to save her.[61]
Padmé's role in the Delegation of 2000 – the senatorial resistance movement to Palpatine's growing absolutism – is discussed in James Luceno's Labyrinth of Evil (2005). The Delegation of 2000 is primarily concerned with Palpatine's calls for public surveillance and restrictions on freedom of movement and action. Still, Padmé is confident Palpatine will relinquish his power when the crisis is over: "He's not stubborn," she tells Bail Organa. "You just don't know him as I do. He'll take our concerns to heart."[62]
Padmé appears in novels and comics set after the events of the original trilogy as holograms and flashbacks. In Troy Denning's The Joiner King (2005), the first book in the "Dark Nest Trilogy" set 35 years after the events of A New Hope, Luke Skywalker discovers a hologram recorded by R2-D2 of Anakin Skywalker informing Padmé of his vision of her death in childbirth. This is the first time Luke sees his mother.[63] Another hologram discovered in R2-D2 chronicles a conversation between Padmé and Obi-Wan. In this hologram, Luke and Leia hear their mother's name for the first time.[64] In the final novel of the trilogy, The Swarm War, Luke and Leia see their mother's death and their own births.[65]
Video Games[edit]
Padmé also appeared in console games based on the Star Wars franchise. Making prominent appearances in the ones based on the Clone Wars timeline and often appearing as a playable character.
Reception[edit]
In 2017, Rolling Stone magazine placed Padmé Amidala at the number 45 spot on their 50 Best Star Wars Characters of All Time list.[66]
Family tree[edit]
Main article: Skywalker family
[hide]
v
t
e
Skywalker family tree
Aika
Lars[a]
Cliegg
Lars[b]
Shmi
Skywalker[c]
Jobal
Naberrie[d]
Ruwee
Naberrie[d]
Beru
Whitesun[e]
Owen
Lars[f]
Anakin
Skywalker[g][h]
Padmé
Amidala[i]
Bail
Organa[j][k]
Breha
Organa[l]
Luke
Skywalker[m][n]
Leia
Organa[o]
Han
Solo[p]
Ben
Solo[p][q]
References:
Jump up ^ Hidalgo, Pablo (2016). Star Wars Character Encyclopedia: Updated and Expanded.
Jump up ^ "Databank: Cliegg Lars". StarWars.com. Retrieved December 3, 2015.
Jump up ^ "Databank: Shmi Skywalker Lars". StarWars.com. Retrieved December 3, 2015.
^ Jump up to: a b Attack of the Clones (2002) and Revenge of the Sith (2005)
Jump up ^ "Databank: Beru Lars". StarWars.com. Retrieved December 3, 2015.
Jump up ^ "Databank: Owen Lars". StarWars.com. Retrieved December 3, 2015.
Jump up ^ "Databank: Anakin Skywalker". StarWars.com. Retrieved December 4, 2015.
Jump up ^ "Databank: Darth Vader". StarWars.com. Retrieved December 4, 2015.
Jump up ^ "Databank: Padmé Amidala". StarWars.com. Retrieved December 4, 2015.
Jump up ^ Adopted father of Leia Organa, as established in Revenge of the Sith (2005).
Jump up ^ "Databank: Bail Organa". StarWars.com. Retrieved December 3, 2015.
Jump up ^ Adopted mother of Leia Organa, as established in Revenge of the Sith (2005).
Jump up ^ "Databank: Luke Skywalker". StarWars.com. Retrieved December 4, 2015.
Jump up ^ In the alternate Star Wars expanded universe (Legends), Luke is married to Mara Jade and has a son, Ben Skywalker.
Jump up ^ "Databank: Princess Leia Organa". StarWars.com. Retrieved December 4, 2015.
^ Jump up to: a b The Force Awakens. 2015.
Jump up ^ In the alternate Star Wars expanded universe (Legends) continuity, Han and Leia have three children: Jaina, Jacen and Anakin Solo.
See also[edit]
Star Wars portal
Women warriors in literature and culture
References[edit]
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^ Jump up to: a b McDonald, Paul F. (2000-06-12). "Amidala: The Goddess With Two Faces". Space.com. Archived from the original on 2007-04-20. Retrieved 2014-11-16.
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Jump up ^ "Return of the galaxy's new beauty New look, love interest for Portman's role in 'Star Wars'" at CNN; last accessed August 5, 2006.
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Jump up ^ Berardinelli, James (September 15, 2015). "Star Wars: The Phantom Menace". www.reelviews.net. Retrieved December 16, 2017.
Jump up ^ {{cite web| Annlee Ellingson, review of Star Wars Episode I: The Phantom Menace, Box Office Magazine, available here [1]Archived June 13, 2006, at the Wayback Machine.; last accessed August 5, 2006.
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Jump up ^ Ed Halter, "May the Force Be Over; The end of the beginning: Lucas's adolescent space opera concludes in a CGI Sith Storm," The Village Voice (New York), May 11, 2005, available here [3] Archived June 28, 2006, at the Wayback Machine.; last accessed August 5, 2006.
Jump up ^ LaSalle, Mick (May 18, 2005). "'Revenge of the Sith' review: Does Lucas know how good a movie he almost made?". San Francisco Chronicle. Retrieved December 16, 2017.
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Jump up ^ McCarthy, Todd (May 5, 2005). "Review: Star Wars Episode III: Revenge of the Sith". Variety. Retrieved December 16, 2017.
Jump up ^ Ian McCaig Biography, at Star Wars.com Archived August 7, 2006, at the Wayback Machine.; last accessed August 5, 2006.
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Jump up ^ "An Introduction to Episode III". StarWars.com. Lucasfilm. May 18, 2005. Archived from the original on December 14, 2005. Retrieved August 5, 2006.
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^ Jump up to: a b Padmé Amidala, Expanded Universe, at the Star Wars Databank; last accessed August 5, 2006.
^ Jump up to: a b c d Padmé Amidala at the Star Wars Databank; last accessed August 5, 2006.
Jump up ^ Salvatore, Attack of the Clones, p. 176.
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Jump up ^ Dominique Mainon and James Ursini, The Modern Amazons: Warrior Women On-Screen (book) (Pompton Plains, N.J.: Hal Leonard, 2006), p. 199, ISBN 0-87910-327-2
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Jump up ^ Terry Moore, "A Summer's Dream," in Star Wars Tales 5 (Dark Horse Comics, September 2000), ISBN 1-59307-286-4.
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Jump up ^ "50 Best 'Star Wars' Characters of All Time, 25. Qui-Gon Jinn". Rolling Stone.
Further reading[edit]
Biggar, Trisha. Dressing a Galaxy: The Costumes of Star Wars. New York: Harry N. Abrams, 2005. ISBN 0-8109-6567-4.
Wallace, Daniel. The New Essential Guide to Characters. New York: Del Rey, 2002. ISBN 0-345-44900-2.
Wallace, Daniel, and Kevin J. Anderson. The New Essential Chronology. New York: Del Rey, 2005. ISBN 0-345-49053-3.
External links[edit]
Wikimedia Commons has media related to Padmé Amidala.
Padmé Amidala in the StarWars.com Databank
Padmé Amidala on Wookieepedia, a Star Wars wiki
Dressing A Galaxy: The Costumes of Star Wars, an exhibit at the Fashion Institute of Design & Merchandising.
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