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Smooth muscle tissue
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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.
It occurs in the walls of hollow organs, including the stomach, intestines, urinary bladder and uterus, and is found 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. Furthermore, it is present in the eyes, where it functions to change the size of the iris and alter the shape of the lens; and in the skin where it causes hair to stand erect in response to cold temperature or fear.[1]
Contents
[hide]
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 alternate 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 Page 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, ISBN 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): 725–744. 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 (Hoboken). 67 (10): 630–4. 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. (page 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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Shamrock
From Wikipedia, the free encyclopedia
Jump to: navigation, search
This article is about the plant that is a symbol of Ireland. Many places, organisations, businesses, products, creative works etc. use the shamrock in their names. For other uses, see Shamrock (disambiguation).
Not to be confused with four-leaf clover.
A shamrock
A shamrock is a young sprig of clover, used as a symbol of Ireland. Saint Patrick, Ireland's patron saint, is said to have used it as a metaphor for the Christian Holy Trinity.[1] The name shamrock comes from Irish seamróg [ˈʃamˠɾˠoːɡ], which is the diminutive of the Irish word for clover (seamair) and means simply "little clover" or "young clover".[2]
Shamrock usually refers to either the species Trifolium dubium (lesser clover, Irish: seamair bhuí)[3] or Trifolium repens (white clover, Irish: seamair bhán). However, other three-leaved plants—such as Medicago lupulina, Trifolium pratense, and Oxalis acetosella—are sometimes called shamrocks or clovers. The shamrock was traditionally used for its medicinal properties and was a popular motif in Victorian times.
Contents
[hide]
1 Botanical species
2 Early references
3 Link to St. Patrick
4 Symbol of Ireland
5 Use overseas
6 See also
7 References
7.1 Bibliography
8 External links
Botanical species[edit]
Trifolium dubium
Trifolium repens
There is still not a consensus over the precise botanical species of clover that is the "true" shamrock. John Gerard in his herbal of 1597 defined the shamrock as Trifolium pratense or Trifolium pratense flore albo, meaning Red or White Clover. He described the plant in English as "Three leaved grasse" or "Medow Trefoile", "which are called in Irish Shamrockes".[4] The Irish botanist Caleb Threlkeld, writing in 1726 in his work entitled Synopsis Stirpium Hibernicarum or A Treatise on Native Irish Plants followed Gerard in identifying the shamrock as Trifolium pratense, calling it White Field Clover.[5]
The botanist Carl von Linné in his 1737 work Flora Lapponica identifies the shamrock as Trifolium pratense, mentioning it by name as Chambroch, with the following curious remark: "Hiberni suo Chambroch, quod est Trifolium pratense purpureum, aluntur, celeres & promtissimi roburis" (The Irish call it shamrock, which is purple field clover, and which they eat to make them speedy and of nimble strength).[6][7] Linnaeus based his information that the Irish ate shamrock on the comments of English Elizabethan authors such as Edmund Spenser who remarked that the shamrock used to be eaten by the Irish, especially in times of hardship and famine. It has since been argued however, that the Elizabethans were confused by the similarity between the Irish (Gaelic) name for young clover seamróg, and the name for wood sorrel seamsóg.[8]
The situation regarding the identity of the shamrock was further confused by a London botanist James Ebenezer Bicheno, who proclaimed in a dissertation in 1830 that the real shamrock was Oxalis acetosella or Wood Sorrel.[9] Bichino falsely claimed that clover was not a native Irish plant and had only been introduced into Ireland in the middle of the 17th century, and based his argument on the same comments by Elizabethan authors that shamrock had been eaten. Bicheno argued that this fitted the wood sorrel better than clover, as wood sorrel was often eaten as a green and used to flavour food. Bicheno's argument has not been generally accepted however, as the weight of evidence favours a species of clover.
A more scientific approach was taken by English botanists James Britten and Robert Holland, who stated in their Dictionary of English Plant Names published in 1878, that their investigations had revealed that Trifolium dubium was the species sold most frequently in Covent Garden as shamrock on St. Patrick's Day, and that it was worn in at least 13 counties in Ireland.[10]
Finally, detailed investigations to settle the matter were carried out in two separate botanical surveys in Ireland, one in 1893[11][12] and the other in 1988.[13] The 1893 survey was carried out by Nathaniel Colgan, an amateur naturalist working as a clerk in Dublin; while the 1988 survey was carried out by E. Charles Nelson, Director of the Irish National Botanic Gardens. Both surveys involved asking people from all across Ireland to send in examples of shamrock, which were then potted up and allowed to flower, so that their botanical species could be identified. The results of both surveys were very similar, showing that the conception of the shamrock in Ireland had changed little in almost a hundred years. The results of the surveys are shown in the table below.
Medicago lupulina flowers in Ireland from May to October[14] and so is not in flower on St. Patrick's Day
Shamrock Survey
Botanical Name
Common Name
Percentage
1893 1988
Trifolium dubium Lesser Clover 51% 46%
Trifolium repens White Clover 34% 35%
Trifolium pratense Red Clover 6% 4%
Medicago lupulina Black Medick 6% 7%
Oxalis acetosella Wood Sorrel _ 3%
Various Trifolium spp., Oxalis spp. 3% 5%
The results show that there is no one "true" species of shamrock, but that Trifolium dubium (Lesser clover) is considered to be the shamrock by roughly half of Irish people, and Trifolium repens (White clover) by another third, with the remaining fifth split between Trifolium pratense, Medicago lupulina, Oxalis acetosella and various other species of Trifolium and Oxalis. None of the species in the survey are unique to Ireland, and all are common European species, so there is no botanical basis for the widespread belief that the shamrock is a unique species of plant that only grows in Ireland.
Early references[edit]
The word shamrock derives from seamair óg or young clover, and references to semair or clover appear in early Irish literature, generally as a description of a flowering clovered plain. For example, in the series of medieval metrical poems about various Irish places called the Metrical Dindshenchus, a poem about Tailtiu or Teltown in Co. Meath describes it as a plain blossoming with flowering clover (mag scothach scothshemrach).[15] Similarly, another story tells of how St. Brigid decided to stay in Co. Kildare when she saw the delightful plain covered in clover blossom (scoth-shemrach).[16] However, the literature in Irish makes no distinction between clover and shamrock, and it is only in English that shamrock emerges as a distinct word.
Three "wild" Irish kerns by Albrecht Dürer 1521
The first mention of shamrock in the English language occurs in 1571 in the work of the English Elizabethan scholar Edmund Campion. In his work Boke of the Histories of Irelande, Campion describes the habits of the "wild Irish" and states that the Irish ate shamrock: "Shamrotes, watercresses, rootes, and other herbes they feed upon".[17] The statement that the Irish ate shamrock was widely repeated in later works and seems to be a confusion with the Irish word seamsóg or wood sorrel (Oxalis).[8] There is no evidence from any Irish source that the Irish ate clover, but there is evidence that the Irish ate wood sorrel. For example, in the medieval Irish work Buile Shuibhne (The Frenzy of Sweeney), the king Sweeney, who has gone mad and is living in the woods as a hermit, lists wood sorrel among the plants he feeds upon.[18]
The English Elizabethan poet Edmund Spenser, writing soon after in 1596, described his observations of war-torn Munster after the Desmond Rebellion in his work A View of the Present State of Ireland. Here shamrock is described as a food eaten as a last resort by starving people desperate for any nourishment during a post-war famine:
Anatomies of death, they spake like ghosts, crying out of theire graves; they did eat of the carrions .... and if they found a plott of water cresses or shamrockes theyr they flocked as to a feast for the time, yett not able long to contynewe therewithall.[19]
The idea that the Irish ate shamrock is repeated in the writing of Fynes Moryson, one-time secretary to the Lord Deputy of Ireland. In his 1617 work An itinerary thorow Twelve Dominions, Moryson describes the "wild Irish", and in this case their supposed habit of eating shamrock is a result of their marginal hand-to-mouth existence as bandits. Moryson claims that the Irish "willingly eat the herbe Schamrock being of a sharpe taste which as they run and are chased to and fro they snatch like beasts out of the ditches." The reference to a sharp taste is suggestive of the bitter taste of wood sorrel.[20]
What is clear is that by the end of the sixteenth century the shamrock had become known to English writers as a plant particularly associated with the Irish, but only with a confused notion that the shamrock was a plant eaten by them. To a herbalist like Gerard it is clear that the shamrock is clover, but other English writers do not appear to know the botanical identity of the shamrock. This is not surprising, as they probably received their information at second or third hand. It is notable that there is no mention anywhere in these writings of St. Patrick or the legend of his using the shamrock to explain the Holy Trinity. However, there are two possible references to the custom of "drowning the shamrock" in "usquebagh" or whiskey. In 1607, the playwright Edward Sharpham in his play The Fleire included a reference to "Maister Oscabath the Irishman ... and Maister Shamrough his lackey".[21] Later, a 1630 work entitled Sir Gregory Nonsence by the poet John Taylor contains the lines: "Whilste all the Hibernian Kernes in multitudes, /Did feast with shamerags steeved in Usquebagh."[22]
Link to St. Patrick[edit]
St. Patrick depicted with shamrock in detail of stained glass window in St. Benin's Church, Wicklow, Ireland
Traditionally, shamrock is said to have been used by Saint Patrick to illustrate the Christian doctrine of the Holy Trinity when Christianising Ireland in the 5th century. The first evidence of a link between St Patrick and the shamrock appears in 1675 on the St Patrick's Coppers or Halpennies. These appear to show a figure of St Patrick preaching to a crowd while holding a shamrock, presumably to explain the doctrine of the Holy Trinity.[23] In pagan Ireland, three was a significant number and the Irish had many triple deities, which could have aided St Patrick in his evangelisation efforts.[24][25] Patricia Monaghan states that "There is no evidence that the clover or wood sorrel (both of which are called shamrocks) were sacred to the Celts". However, Jack Santino speculates that "The shamrock was probably associated with the earth and assumed by the druids to be symbolic of the regenerative powers of nature ... Nevertheless, the shamrock, whatever its history as a folk symbol, today has its meaning in a Christian context. Pictures of Saint Patrick depict him driving the snakes out of Ireland with a cross in one hand and a sprig of shamrocks in the other."[26] Roger Homan writes, "We can perhaps see St Patrick drawing upon the visual concept of the triskele when he uses the shamrock to explain the Trinity".[27]
The first written mention of the link does not appear until 1681, in the account of Thomas Dineley, an English traveller to Ireland. Dineley writes:
The 17th day of March yeerly is St Patricks, an immoveable feast, when ye Irish of all stations and condicions were crosses in their hatts, some of pinns, some of green ribbon, and the vulgar superstitiously wear shamroges, 3 leav'd grass, which they likewise eat (they say) to cause a sweet breath.[28]
There is nothing in Dineley's account of the legend of St. Patrick using the shamrock to teach the mystery of the Holy Trinity, and this story does not appear in writing anywhere until a 1726 work by the botanist Caleb Threlkeld.[5] Threlkeld identifies the shamrock as White Field Clover (Trifolium pratense album ) and comments rather acerbically on the custom of wearing the shamrock on St. Patrick's Day:
This plant is worn by the people in their hats upon the 17. Day of March yearly, (which is called St. Patrick's Day.) It being a current tradition, that by this Three Leafed Grass, he emblematically set forth to them the Mystery of the Holy Trinity. However that be, when they wet their Seamar-oge, they often commit excess in liquor, which is not a right keeping of a day to the Lord; error generally leading to debauchery.
The Rev Threlkeld's remarks on liquor undoubtedly refer to the custom of toasting St. Patrick's memory with "St. Patrick's Pot", or "drowning the shamrock" as it is otherwise known. After mass on St. Patrick's Day the traditional custom of the menfolk was to lift the usual fasting restrictions of Lent and repair to the nearest tavern to mark the occasion with as many St. Patrick's Pots as they deemed necessary. The drowning of the shamrock was accompanied by a certain amount of ritual as one account explains:[29][30]
Shamrock on an Irish Defence Forces UN beret
"The drowning of the shamrock" by no means implies it was necessary to get drunk in doing so. At the end of the day the shamrock which has been worn in the coat or the hat is removed and put into the final glass of grog or tumbler of punch; and when the health has been drunk or the toast honoured, the shamrock should be picked out from the bottom of the glass and thrown over the left shoulder.
The shamrock is still chiefly associated with Saint Patrick's Day, which has become the Irish national holiday, and is observed with parades and celebrations worldwide. The custom of wearing shamrock on the day is still observed and depictions of shamrocks are habitually seen during the celebrations.
Symbol of Ireland[edit]
Drawing of the medal awarded to the First Magherafelt Volunteers for skill with broadsword showing shamrocks.
As St. Patrick is Ireland's patron saint, shamrock has been used as a symbol of Ireland since the 18th century, in a similar way to how a rose is used for England, thistle for Scotland and daffodil for Wales. The shamrock first began to change from a symbol purely associated with St. Patrick to an Irish national symbol when it was taken up as an emblem by rival militias, during the turbulent politics of the late eighteenth century. On one side were the Volunteers (also known as the Irish Volunteers), who were local militias in late 18th century Ireland, raised to defend Ireland from the threat of French and Spanish invasion when regular British soldiers were withdrawn from Ireland to fight during the American Revolutionary War.[31] On the other side were revolutionary nationalist groups, such as the United Irishmen.
Among the Volunteers, examples of the use of the shamrock include its appearance on the guidon of the Royal Glin Hussars formed in July 1779 by the Knight of Glin, and its appearance on the flags of the Limerick Volunteers, the Castle Ray Fencibles and the Braid Volunteers.[32][33] The United Irishmen adopted green as their revolutionary colour and wore green uniforms or ribbons in their hats, and the green concerned was often associated with the shamrock. The song The Wearing of the Green commemorated their exploits and various versions exist which mention the shamrock. The Erin go bragh flag was used as their standard and was often depicted accompanied by shamrocks, and in 1799 a revolutionary journal entitled The Shamroc briefly appeared in which the aims of the rebellion were supported.[34]
British two shilling coin 1949
Since the 1800 Acts of Union between Britain and Ireland the shamrock was incorporated into the Royal Coat of Arms of the United Kingdom, depicted growing from a single stem alongside the rose of England, and the thistle of Scotland to symbolise the unity of the three kingdoms. Since then, the shamrock has regularly appeared alongside the rose, thistle and (sometimes) leek for Wales in British coins such as the two shilling and crown, and in stamps. The rose, thistle and shamrock motif also appears regularly on British public buildings such as Buckingham Palace.
Throughout the nineteenth century the popularity of the shamrock as a symbol of Ireland grew, and it was depicted in many illustrations on items such as book covers and St. Patrick's Day postcards. It was also mentioned in many songs and ballads of the time. For example, a popular ballad called The Shamrock Shore lamented the state of Ireland in the nineteenth century.[35] Another typical example of such a ballad appears in the works of Thomas Moore whose Oh the Shamrock embodies the Victorian spirit of sentimentality. It was immensely popular and contributed to raising the profile of the shamrock as an image of Ireland:[36]
Oh The Shamrock
-
Through Erin's Isle,
To sport awhile,
As Love and Valor wander'd
With Wit, the sprite,
Whose quiver bright
A thousand arrows squander'd.
Where'er they pass,
A triple grass
Shoots up, with dew-drops streaming,
As softly green
As emeralds seen
Through purest crystal gleaming.
Oh the Shamrock, the green immortal Shamrock!
Chosen leaf
Of Bard and Chief,
Old Erin's native Shamrock!
Rose thistle and shamrock motif on gate pillar at Buckingham Palace
Irish American Music sheet
St Patrick's Day postcard 1912
Throughout the nineteenth and twentieth centuries, the shamrock continued to appear in a variety of settings.[37] For example, the shamrock appeared on many buildings in Ireland as a decorative motif, such as on the facade of the Kildare Street Club building in Dublin, St. Patrick's Cathedral, Armagh, and the Harp and Lion Bar in Listowel, Co. Kerry. It also appears on street furniture, such as old lamp standards like those in Mountjoy Square in Dublin, and on monuments like the Parnell Monument, and the O'Connell Monument, both in O'Connell Street, Dublin. Shamrocks also appeared on decorative items such as glass, china, jewellery, poplin and Irish lace. Belleek Pottery in Co. Fermanagh, for example regularly features shamrock motifs.
Lamppost in Mountjoy Square, Dublin, early 20th Century
Design on Harp and Lion Bar Listowel, Co. Kerry
Work by Belleek Pottery, which often features shamrock motifs
2d Map of Ireland: the first Irish postage stamp featured the shamrock
The shamrock is used in the emblems of many state organisations, both in the Republic of Ireland and Northern Ireland. Some of these are all-Ireland bodies, (such as Tourism Ireland)[38] as well as organisations specific to the Republic of Ireland (such as IDA Ireland)[39] and Northern Ireland (such as Police Service of Northern Ireland). The Irish Postal Service An Post, regularly features the shamrock on its series of stamps. The airline Aer Lingus uses the emblem in its logos, and its air traffic control call sign is "SHAMROCK".
An Aer Lingus aircraft with a shamrock on its tail fin
The shamrock has been registered as a trademark by the Government of Ireland.[40] In the early 1980s, Ireland defended its right to use the shamrock as its national symbol in a German trademark case, which included high-level representation from taoiseach Charles Haughey. Having originally lost, Ireland won on appeal to the German Supreme Court in 1985.[41]
It has become a tradition for the Irish Taoiseach to present a bowl of shamrocks in a special Waterford Crystal bowl featuring a shamrock design to the President of the United States in the White House every St. Patrick's Day.[42]
Shamrock is also used in emblems of UK organisations with an association with Ireland, such as the Irish Guards. Soldiers of the Royal Irish Regiment of the British Army use the shamrock as their emblem, and wear a sprig of shamrock on Saint Patrick's Day. Shamrock are exported to wherever the regiment is stationed throughout the world. Queen Victoria decreed over a hundred years ago that soldiers from Ireland should wear a sprig of shamrock in recognition of fellow Irish soldiers who had fought bravely in the Boer War, a tradition continued by British army soldiers from both the north and the south of Ireland following partition in 1921. The coat of arms on the flag of the Royal Ulster Constabulary George Cross Foundation was cradled in a wreath of shamrock.[43]

The shamrock also appears in the emblems of a wide range of voluntary and non-state organisations in Ireland, such as the Irish Farmers Association,[44] the Boy Scouts of Ireland association, Scouting Ireland[45] Irish Girl Guides,[46] and the Irish Kidney Donors Association.,[47] In addition many sporting organisations representing Ireland use the shamrock in their logos and emblems. Examples include the Irish Football Association (Northern Ireland), Irish Rugby Football Union, Swim Ireland, Cricket Ireland, and the Olympic Council of Ireland. A sprig of shamrock represents the Lough Derg Yacht Club Tipperary, (est. 1836). The shamrock is the official emblem of Irish football club Shamrock Rovers.
This flowering Shamrock was purchased at a grocery store. It is a South American species of wood sorrel Oxalis regnellii.
Use overseas[edit]
Shamrock commonly appears as part of the emblem of many organisations in countries overseas with communities of Irish descent. Outside Ireland, various organisations, businesses and places also use the symbol to advertise a connection with the island.
The shamrock features in the emblem of the Ancient Order of Hibernians, the largest and oldest Irish Catholic organisation. Founded in New York City in 1836 by Irish immigrants, it claims a membership of 80,000 in the United States, Canada and Ireland.[48]
The Emerald Society, an organisation of American police officers or fire fighters of Irish heritage, includes a shamrock on its badge. Emerald Societies are found in most major US cities such as New York City, Milwaukee, WI, Jersey City, NJ, District of Columbia, Boston, Chicago, San Francisco, Los Angeles and Saint Paul, Minnesota.
The shamrock is featured in the "compartment" of the Royal Arms of Canada, as part of a wreath of shamrocks, roses, thistles, and lilies (representing the Irish, English, Scottish, and French settlers of Canada).
The flag of the city of Montreal, Quebec, Canada has a shamrock in the lower right quadrant. The shamrock represents the Irish population, one of the four major ethnic groups that made up the population of the city in the 19th century when the arms were designed, the other three being the French (represented by a fleur-de-lis in the upper-left), the English (represented by a rose in the upper-right), and the Scots (represented by a thistle in the lower-left).
The shamrock is featured on the passport stamp of Montserrat, many of whose citizens are of Irish descent.
The shamrock signified the Second Corps of the Army of the Potomac in the American Civil War, which contained the Irish Brigade. It can still be seen on the regimental coat of arms of "The Fighting Sixty-Ninth"
Flag of St. Patrick's Battalion of the Mexican army reconstructed from description of Jon Riley. The flag has since become an emblem of Irish nationalism
The Erin Go Bragh flag, used originally by the Saint Patrick's Battalion of the Mexican Army, uses an angelic Cláirseach, a medieval Irish harp, cradled in a wreath of clover. A flag strongly symbolic of Irish nationalism, it is often seen on Saint Patrick's Day, usually displayed during the parades.
The crest of Glasgow Celtic Football Club originally included a shamrock which was changed in 1938 to a four leaved clover for reasons that remain unclear.[49] The Club was founded in 1887 in Glasgow among the poor Irish immigrants of the city.
London Irish rugby football club has a shamrock on its crest. The club was founded in 1898 for the young Irishmen of London.
The Shamrocks Motorcycle Club is a USA-based traditional motorcycle club (composed of law enforcement personnel) which uses the Shamrock as its name and symbol.[50]
The basketball team, Boston Celtics, in the USA incorporate the shamrock in their logo. Former NBA player Shaquille O'Neal nicknamed himself the "Big Shamrock" after joining the team.
In Australia, the Melbourne Celtic Club features a shamrock on its emblem. The club was founded in 1887 for the Irish and other Celtic groups in the city.[51]
During the Russian Civil War a British officer Col. P.J. Woods, of Belfast, established a Karelian Regiment which had a shamrock on an orange field as its regimental badge.
A shamrock (Trifylli) is the official emblem of Greek multi-sport club Panathinaikos A.O., Greek football club Acharnaikos F.C. and Cypriot sports club AC Omonia. A red shamrock is also the emblem of Platanias F.C., a Cretan football team of Chania.
The Danish football club Viborg FF uses a shamrock in its badge and it has become a symbol of the town of Viborg.
The German football club SpVgg Greuther Fürth also has a shamrock in its badge as it is a symbol of the city of Fürth.
A shamrock is used in the Coat of Arms of the Italian Air Force.
According to the Anti-Defamation League, the Aryan Brotherhood symbol combines a shamrock with a swastika.[52]
The Flag of Montreal. The shamrock is located in the lower right corner
AOH logo
Emblem of the Roman Catholic Archdiocese of Newark, New Jersey
Panathinaikos F.C. shirt
Aryan Brotherhood shamrock with a Hakenkreuz
See also[edit]
Guernsey Lily
Ragwort (Isle of Man)
St. Patrick's Blue
Trefoil
References[edit]
Jump up ^ Treeck, Carl Van; Croft, Aloysius (1936). Symbols in the Church. Bruce Publishing Co. St. Patrick is said to have used the shamrock in explaining to the pagan Irish the idea of the Holy Trinity. |access-date= requires |url= (help)
Jump up ^ Nelson (1991), p. 14
Jump up ^ "Lesser Hop Trefoil, Trifolium dubium - Flowers - NatureGate". www.luontoportti.com.
Jump up ^ "The Herball or Generall Historie of Plantes (1597)". caliban.mpiz-koeln.mpg.de.
^ Jump up to: a b "NS07 Threlkeld Shamrock | a whole new world". dublincitypubliclibraries.com. Retrieved 4 July 2014.
Jump up ^ Linnæi, Caroli (1737). Flora Lapponica, exhibens plantas per Lapponiam crescentes, secundum systema sexuale, collectas in itinere impensis Soc. reg. scient. Upsaliensis, anno 1732 instituto. London: B White et Filiorum. pp. 229–230.
Jump up ^ Nelson (1991), p. 34
^ Jump up to: a b Kelly, Fergus, Early Irish Farming, (2000), Dublin, p 311
Jump up ^ "Journal of the Royal Institution of Great Britain". archive.org.
Jump up ^ "A dictionary of English plant-names". archive.org.
Jump up ^ Colgan, Nathaniel (1892). "The Shamrock: an attempt to fix its species". The Irish Naturalist. 1 (5): 95–97.
Jump up ^ Colgan, Nathaniel (1893). "The Shamrock: a further attempt to fix its species". The Irish Naturalist. 2 (8): 207–211.
Jump up ^ Nelson (1991), pp. 86–90, 139–144, 153
Jump up ^ design, Coded web. "Wildflower Trefoil, Lesser Irish Wild Flora Wildflowers of Ireland". www.wildflowersofireland.net.
Jump up ^ "The Metrical Dindshenchas". www.ucc.ie.
Jump up ^ Stokes, Whitley, Lives of the Saints from the Book of Lismore, (1890), p177
Jump up ^ Ware, Sir James (4 September 2017). "Ancient Irish Histories: The Works of Spencer, Campion, Hanmer, and Marleburrough". Reprinted at the Hibernia Press – via Google Books.
Jump up ^ "Buile Suibhne". www.ucc.ie.
Jump up ^ "A View of the Present State of Ireland". www.luminarium.org.
Jump up ^ Moryson, Fynes, An Itinerary etc. Vol IV, p200 Archived 28 March 2013 at the Wayback Machine.
Jump up ^ Nelson (1991), p. 22
Jump up ^ Taylor, John (4 September 2017). "Sir Gregory Nonsence His Newes from No Place". O[kes] – via Google Books.
Jump up ^ "Newbie's St. Patrick Coppers - Introduction". coins.nd.edu. Retrieved 4 July 2014.
Jump up ^ Monaghan, Patricia (1 January 2009). The Encyclopedia of Celtic Mythology and Folklore. Infobase Publishing. ISBN 9781438110370. There is no evidence that the clover or wood sorrel (both of which are called shamrocks) were sacred to the Celts in any way. However, the Celts had a philosophical and cosmological vision of triplicity, with many of their divinities appearing in three. Thus when St. Patrick, attempting to convert the Druids on Beltane, held up a shamrock and discoursed on the Christian Trinity, the three-in-one god, he was doing more than finding a homely symbol for a complex religious concept. He was indicating knowledge of the significance of three in the Celtic realm, a knowledge that probably made his mission far easier and more successful than if he had been unaware of that number's meaning.
Jump up ^ Hegarty, Neil (24 April 2012). Story of Ireland. Ebury Publishing. ISBN 9781448140398. In some ways, though, the Christian mission resonated: pre-Christian devotion was characterized by, for example, the worship of gods in groups of three, by sayings collected in threes (triads), and so on - from all of which the concept of the Holy Trinity was not so very far removed. Against this backdrop the myth of Patrick and his three-leafed shamrock fits quite neatly.
Jump up ^ Santino, Jack (1995). All Around the Year: Holidays and Celebrations in American Life. University of Illinois Press. p. 80. ISBN 9780252065163.
Jump up ^ Homan, Roger (2006). The Art of the Sublime: Principles of Christian Art and Architecture. Ashgate Publishing. p. 37.
Jump up ^ Journal of the Kilkenny Archaeological Society, 1 (1856), p183
Jump up ^ Journal of the Kildare Archaeological Society, 1908, p 443
Jump up ^ Danaher, Kevin, The Year in Ireland: Irish Calendar Customs, (1972), Dublin, pp 64–5
Jump up ^ Blackstock, Allan (2001). Issue 2 of Belfast Society publications, ed. Double traitors?: the Belfast Volunteers and Yeomen, 1778–1828. Ulster Historical Foundation. p. 2. ISBN 978-0-9539604-1-5. Retrieved 3 October 2009.
Jump up ^ Nelson (1991), p. 55
Jump up ^ Kieran Kennedy (10 May 2007). "Limerick Volunteers 1776–1793" (PDF). Retrieved 4 July 2014.
Jump up ^ [1][dead link]
Jump up ^ "Karan Casey - Shamrock Shore Lyrics". 11 April 2013. Archived from the original on 11 April 2013.
Jump up ^ Moore, Thomas (4 September 2017). "Melodies, national airs, miscellaneous poems and the Odes of Anacreon". Phillips, Sampson and Company – via Google Books.
Jump up ^ Nelson (1991), pp. 90–120
Jump up ^ "Corporate Logo". tourismirelandbrand.com. Archived from the original on 12 April 2013. Retrieved 4 July 2014.
Jump up ^ "Invest in Ireland, IDA Ireland, Foreign Direct Investment into Ireland, Business in Ireland". idaireland.com. Retrieved 4 July 2014.
Jump up ^ Use of the harp and the shamrock were registered by the Irish government as international trademarks. See "Record of the meeting of the Joint Oireachtas Committee on Enterprise and Small Business, 26 March 2003". Archived from the original on 17 November 2015. Retrieved 2015-11-13. . Retrieved 20 July 2008.
Jump up ^ Gartland, Fiona (31 December 2011). "How Ireland lost the battle for the shamrock in Germany". The Irish Times. Retrieved 13 November 2015.
Jump up ^ "The White House Gets a Bit of Luck from Waterford and Irish Prime Minister | House of Waterford". waterfordvisitorcentre.com. Retrieved 4 July 2014.
Jump up ^ "About Us - College of Arms". www.college-of-arms.gov.uk.
Jump up ^ "Irish Farmers' Association". ifa.ie. Retrieved 4 July 2014.
Jump up ^ "History of Scouting in Ireland Join the Adventure! | Scouting Ireland | www.scouts.ie". scouts.ie. Retrieved 4 July 2014.
Jump up ^ "Association Logo and Badge - Catholic Guides of Ireland". girlguidesireland.ie. Retrieved 4 July 2014.
Jump up ^ "IKA". ika.ie. Retrieved 4 July 2014.
Jump up ^ "Ancient Order of Hibernians — The Oldest and Largest Irish-Catholic Organization in the United States. Established 1836". aoh.com. Retrieved 4 July 2014.
Jump up ^ "Celtic badge - The Celtic Wiki". thecelticwiki.com. Retrieved 4 July 2014.
Jump up ^ [2] Archived 16 April 2015 at the Wayback Machine.
Jump up ^ "The Celtic Club - About". celticclub.com.au. Retrieved 13 November 2015.
Jump up ^ "Aryan Brotherhood". Anti-Defamation League.
Bibliography[edit]
Nelson, E. Charles (1991). Shamrock: Botany and History of an Irish Myth: a Biography of the Shamrock in History, Literature, Music and Art. Boethius Press. ISBN 0-86314-199-4.
External links[edit]
The truth behind the shamrock on the BBC News website, dated 17 March 2004. Retrieved 2008-07-20
Landscaping: Shamrocks and 4-Leaf Clovers on the About.com website. Retrieved 2008-07-20
www.decodeunicode.org/en/u+2618 Shamrock as a symbol in Unicode
Retrieved from "https://en.wikipedia.org/w/index.php?title=Shamrock&oldid=809501830"
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