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Your current IP address is 8.222.208.146. </p> <div class="paragraphbreak" style="margin-top:0.5em"></div><div style="font-size: 16px;"> <p>Even while globally blocked, you will <i>usually</i> still be able to edit pages on <a href="https://meta.wikimedia.org/wiki/" class="extiw" title="m:">Meta-Wiki</a>. </p> </div> <div class="paragraphbreak" style="margin-top:0.5em"></div><div style="font-size: 16px;"> <p>If you believe you were blocked by mistake, you can find additional information and instructions in the <a href="https://meta.wikimedia.org/wiki/Special:MyLanguage/Stewards/Wizard" class="extiw" title="m:Special:MyLanguage/Stewards/Wizard">Stewards Block Wizard</a>. <a href="/w/index.php?title=MediaWiki:Wikimedia-globalblocking-blockedtext-mistake-email-steward&action=edit&redlink=1" class="new" title="MediaWiki:Wikimedia-globalblocking-blockedtext-mistake-email-steward (page does not exist)">MediaWiki:Wikimedia-globalblocking-blockedtext-mistake-email-steward</a> </p> </div> <p>Other useful links: <a href="https://meta.wikimedia.org/wiki/Global_blocks" class="extiw" title="m:Global blocks">Global blocks</a> · <a href="/wiki/Help:I_have_been_blocked" title="Help:I have been blocked">Help:I have been blocked</a> </p> </div></li></ul><hr /> <div id="viewsourcetext">You can view and copy the source of this page:</div><textarea readonly="" accesskey="," id="wpTextbox1" cols="80" rows="25" style="" class="mw-editfont-monospace" lang="en" dir="ltr" name="wpTextbox1">{{Short description|Relationship between stroke volume and end diastolic volume}} {{distinguish|text=the [[Starling equation]] that describes transcapillary exchange}} [[File:Starling RAP combined.svg|thumb|440px|[[Cardiac function curve]]. In diagrams illustrating the '''Frank–Starling law of the heart''', the y-axis often describes the [[stroke volume]], [[stroke work]], or [[cardiac output]]. The x-axis often describes [[end-diastolic volume]], [[right atrial pressure]], or [[pulmonary capillary wedge pressure]]. The three curves illustrate that shifts along the same line indicate a change in [[Preload (cardiology)|preload]], while shifts from one line to another indicate a change in [[afterload]] or [[Myocardial contractility|contractility]]. A blood volume increase would cause a shift along the line to the right, which increases left ventricular end diastolic volume (x axis), and therefore also increases stroke volume (y axis). ]] The '''Frank–Starling law''' of the [[human heart|heart]] (also known as '''Starling's law''' and the '''Frank–Starling mechanism''') represents the relationship between [[stroke volume]] and [[End-diastolic volume|end diastolic volume]].<ref name=":0">Widmaier, E. P., Hershel, R., & Strang, K. T. (2016).''Vander's Human Physiology: The Mechanisms of Body Function''(14th ed.). New York, NY: McGraw-Hill Education. {{ISBN|978-1-259-29409-9}}</ref> The law states that the stroke volume of the heart increases in response to an increase in the volume of blood in the [[Ventricle (heart)|ventricles]], before contraction (the [[end diastolic volume]]), when all other factors remain constant.<ref name=":0" /> As a larger volume of blood flows into the ventricle, the blood stretches cardiac muscle, leading to an increase in the force of contraction. The Frank-Starling mechanism allows the cardiac output to be synchronized with the venous return, arterial blood supply and humoral length,<ref name="isbn0-7817-7311-3">{{cite book|title=Physiology|publisher=Lippincott Williams & Wilkins|year=2007|isbn=978-0-7817-7311-9|location=Hagerstwon, MD|pages=[https://archive.org/details/physiology00cost_0/page/81 81]|author=Costanzo, Linda S.|url-access=registration|url=https://archive.org/details/physiology00cost_0/page/81}}</ref> without depending upon external regulation to make alterations. The physiological importance of the mechanism lies mainly in maintaining left and right ventricular output equality.<ref name=":0" /><ref name=":1">{{cite journal | year = 1992 | title = Functional significance of the Frank-Starling mechanism under physiological and pathophysiological conditions | url = https://academic.oup.com/eurheartj/article-abstract/13/suppl_E/7/486060/Functional-significance-of-the-Frank-Starling?redirectedFrom=PDF | journal = European Heart Journal | volume = 13| pages = 7–14| doi = 10.1093/eurheartj/13.suppl_E.7 | author = Jacob R., Dierberger B., Kissling G. | pmid = 1478214 }}</ref> ==Physiology== The Frank-Starling mechanism occurs as the result of the '''length-tension relationship''' observed in striated muscle, including for example [[skeletal muscle]]s, arthropod muscle<ref>{{Cite journal|last1=West|first1=J. M.|last2=Humphris|first2=D. C.|last3=Stephenson|first3=D. G.|date=1992|title=Differences in maximal activation properties of skinned short- and long-sarcomere muscle fibres from the claw of the freshwater crustacean Cherax destructor|journal=Journal of Muscle Research and Cell Motility|volume=13|issue=6|pages=668–684|issn=0142-4319|pmid=1491074|doi=10.1007/BF01738256|s2cid=21089844}}</ref> and [[Cardiac muscle|cardiac (heart) muscle]].<ref name=":2">{{cite journal | author = Katz Arnold M | year = 2002 | title = "Ernest Henry Starling, His Predecessors, and the "Law of the Heart | url = http://circ.ahajournals.org/content/106/23/2986.1 | journal = Circulation | volume = 106 | issue = 23 | pages = 2986–2992 | doi = 10.1161/01.CIR.0000040594.96123.55 | pmid = 12460884 | doi-access = free | access-date = 2017-05-03 | archive-date = 2018-07-13 | archive-url = https://web.archive.org/web/20180713202316/http://circ.ahajournals.org/content/106/23/2986.1 | url-status = dead }}</ref><ref name=":6" /><ref name=":5" /> As striated muscle is stretched, active tension is created by altering the overlap of thick and thin filaments. The greatest isometric active tension is developed when a muscle is at its optimal length. In most relaxed skeletal muscle fibers, passive elastic properties maintain the muscle fibers length near optimal, as determined usually by the fixed distance between the attachment points of tendons to the bones (or the exoskeleton of arthropods) at either end of the muscle. In contrast, the relaxed [[sarcomere]] length of cardiac muscle cells, in a resting ventricle, is lower than the optimal length for contraction.<ref name=":0" /> There is no bone to fix sarcomere length in the heart (of any animal) so sarcomere length is very variable and depends directly upon blood filling and thereby expanding the heart chambers. In the human heart, maximal force is generated with an initial sarcomere length of 2.2 micrometers, a length which is rarely exceeded in a normal heart. Initial lengths larger or smaller than this optimal value will decrease the force the muscle can achieve. For longer sarcomere lengths, this is the result of there being less overlap of the thin and thick filaments;<ref>{{Cite journal|last1=Huxley|first1=H.|last2=Hanson|first2=J.|date=1954-05-22|title=Changes in the cross-striations of muscle during contraction and stretch and their structural interpretation|journal=Nature|volume=173|issue=4412|pages=973–976|issn=0028-0836|pmid=13165698|doi=10.1038/173973a0|bibcode=1954Natur.173..973H|s2cid=4180166}}</ref><ref>{{Cite journal|last1=Huxley|first1=A. F.|last2=Niedergerke|first2=R.|date=1954-05-22|title=Structural changes in muscle during contraction; interference microscopy of living muscle fibres|journal=Nature|volume=173|issue=4412|pages=971–973|issn=0028-0836|pmid=13165697|doi=10.1038/173971a0|bibcode=1954Natur.173..971H|s2cid=4275495}}</ref><ref name=":7" /> for shorter sarcomere lengths, the cause is the decreased sensitivity for calcium by the [[myofilament]]s.<ref name=":4">{{Cite journal|last1=Allen|first1=D.G.|last2=Kentish|first2=J.C.|date=1985|title=The cellular basis of the length-tension relation in cardiac muscle.|pmid=3900426|journal=Journal of Molecular and Cellular Cardiology|volume=17|issue=9|pages=821–840|doi=10.1016/S0022-2828(85)80097-3}}</ref><ref name=":5">{{Cite journal|last1=Stephenson|first1=D.G.|last2=Williams|first2=D.A.|date=1982|title=Effects of sarcomere length on the force-pCa relation in fast- and slow-twitch skinned muscle fibres from the rat|pmid=7182478|pmc=1197268|journal=Journal of Physiology|volume=333|pages=637–653|doi=10.1113/jphysiol.1982.sp014473}}</ref> An increase in filling of the ventricle increases the load experienced by each cardiac muscle cells, stretching their sarcomeres toward their optimal length.<ref name=":0" /> The stretching sarcomeres augments cardiac [[muscle contraction]] by increasing the [[calcium]] sensitivity of the [[myofibril]]s,<ref name="Cardiovascular Physiology Concepts">Klabunde, Richard E. "Cardiovascular Physiology Concepts". Lippincott Williams & Wilkins, 2011, p. 74.</ref> causing a greater number of [[actin]]-[[myosin]] cross-bridges to form within the muscle. Specifically, the sensitivity of [[troponin]] for binding Ca<sup>2+</sup> increases and there is an increased release of Ca<sup>2+</sup> from the [[sarcoplasmic reticulum]]. In addition, stretch of cardiac myocytes increases the releasability of Ca<sup>2+</sup> from the internal store, the [[sarcoplasmic reticulum]], as shown by an increase in Ca<sup>2+</sup> spark rate upon axial stretch of single cardiac myocytes.<ref>{{Cite journal |last1=Iribe |first1=G |last2=Ward |first2=CW |last3=Camelliti |first3=P |last4=Bollensdorff |first4=C |last5=Mason |first5=F |last6=Burton |first6=RAB |last7=Garny |first7=A |last8=Morphew |first8=MK |last9=Hoenger |first9=A |last10=Lederer |first10=WJ |last11=Kohl |first11=P |date=2009-03-27 |title=Axial stretch of rat single ventricular cardiomyocytes causes an acute and transient increase in Ca2+ spark rate |journal=Circulation Research |volume=104 |issue=6 |pages=787–795 |doi=10.1161/CIRCRESAHA.108.193334 |issn=1524-4571 |pmc=3522525 |pmid=19197074}}</ref> Finally, there is thought to be a decrease in the spacing between thick and thin filaments, when a cardiac muscle is stretched, allowing an increased number of [[Sliding filament theory|cross-bridges]] to form.<ref name=":0" /> The force that any single cardiac muscle cell generates is related to the sarcomere length at the time of muscle cell activation by calcium. The stretch on the individual cell, caused by ventricular filling, determines the sarcomere length of the fibres. Therefore the force (pressure) generated by the cardiac muscle fibres is related to the [[end-diastolic volume]] of the left and right ventricles as determined by complexities of the force-sarcomere length relationship.<ref name=":4" /><ref name=":5" /><ref name=":6">{{Cite journal|last1=Stephenson|first1=D.G.|last2=Stewart|first2=A.W.|last3=Wilson|first3=G.J.|date=1989|title=Dissociation of force from myofibrillar MgATPase and stiffness at short sarcomere lengths in rat and toad skeletal muscle|journal=Journal of Physiology|volume=410|pages=351–366|pmc=1190483|doi=10.1113/jphysiol.1989.sp017537|pmid=2529371}}</ref> Due to the intrinsic property of [[myocardium]] that is responsible for the Frank-Starling mechanism, the heart can automatically accommodate an increase in venous return, at any heart rate.<ref name=":0" /><ref name=":7">{{Cite journal|last1=Moss|first1=Richard L.|last2=Fitzsimons|first2=Daniel P.|date=2002-01-11|title=Frank-Starling Relationship|journal=Circulation Research|language=en|volume=90|issue=1|pages=11–13|issn=0009-7330|pmid=11786511|doi=10.1161/res.90.1.11|doi-access=free}}</ref> The mechanism is of functional importance because it serves to adapt left ventricular output to right ventricular output.<ref name=":1" /> If this mechanism did not exist and the right and left cardiac outputs were not equivalent, blood would accumulate in the pulmonary circulation (were the right ventricle producing more output than the left) or the systemic circulation (were the left ventricle producing more output than the right).<ref name=":0" /><ref name=":3">{{Cite book|title=Guyton and Hall textbook of medical physiology|last=Hall|first=John|publisher=Saunders/Elsevier|year=2016|isbn=978-1-4160-4574-8|edition=13th|location=Philadelphia, Pa.|pages=169–178 (Ch. 14)}}</ref> ==Clinical examples== === Premature ventricular contraction === [[Premature ventricular contraction]] causes early emptying of the [[left ventricle]] (LV) into the [[aorta]]. Since the next ventricular contraction occurs at its regular time, the filling time for the LV increases, causing an increased LV end-diastolic volume. Due to the Frank–Starling mechanism, the next ventricular contraction is more forceful, leading to the ejection of the larger than normal volume of blood, and bringing the LV end-systolic volume back to baseline.<ref name=":3" /> === Diastolic dysfunction – heart failure === [[Diastolic heart failure|Diastolic dysfunction]] is associated with a reduced compliance, or increased stiffness, of the ventricle wall. This reduced compliance results in an inadequate filling of the ventricle and a decrease in the end-diastolic volume. The decreased end-diastolic volume then leads to a reduction in stroke volume because of the Frank-Starling mechanism.<ref name=":0" /> ==History== The Frank–Starling law is named after the two physiologists, [[Otto Frank (physiologist)|Otto Frank]] and [[Ernest Starling|Ernest Henry Starling]]. However, neither Frank nor Starling was the first to describe the relationship between the end-diastolic volume and the regulation of cardiac output.<ref name=":2" /> The first formulation of the law was theorized by the Italian physiologist [[Dario Maestrini]], who on December 13, 1914, started the first of 19 experiments that led him to formulate the ''"legge del cuore" ''.<ref>{{cite book|title=Trattato di Fisiologia|last=Spadolini|first=Igino|year=1946|editor=UTET|volume=2|location=Torino|pages=}}</ref><ref>{{cite book|title=Fisiologia|last=Berne|first=Robert M.|year=2004|editor=Ambrosiana|location=Milano |author-link1=er al.}}</ref><ref name="urlwww.ancecardio.it">{{cite web|url=http://www.ancecardio.it/it/doc/1161358734110_4013.pdf|title=www.ancecardio.it|pages=29–31|language=it|archive-date=2013-11-09|access-date=6 August 2010|url-status=dead|archive-url=https://web.archive.org/web/20131109113213/http://www.ancecardio.it/it/doc/1161358734110_4013.pdf}}</ref><ref>{{cite journal|last=MAESTRINI|first=D.|date=February 1951|title=[Genesis of the so-called insufficient contractions of the heart in decompensation.]|journal=Policlinico Prat|volume=58|pages=257–68|pmid=14833944|number=9}}</ref><ref>{{cite journal|last=MAESTRINI|first=D.|date=July 1951|title=[The importance of the altered organic exchange (fatigue), of the structure and colloidal state of the fiber, for the genesis of the so-called small, insufficient contractions of the heart in failure.]|journal=Policlinico Prat|volume=58|pages=933–45|pmid=14864102|number=30}}</ref><ref>{{cite journal|last=MAESTRINI|first=D.|date=November 1951|title=[The law of the heart in biology and clinical medicine.]|journal=Minerva Med|volume=42|pages=857–64|pmid=14919226|number=80}}</ref><ref>{{cite journal|last=MAESTRINI|first=D.|date=June 1952|title=[A new theory of cardiac decompensation.]|journal=Policlinico Prat|volume=59|pages=797–814|pmid=14957592|number=24}}</ref><ref>{{cite journal|last=MAESTRINI|first=D.|title=[Not Available.]|journal=Gazz Sanit|volume=18|pages=162–4|pmid=18859625|number=5|year=1947}}</ref><ref>{{cite journal|last=PENNACCHIO|first=L.|date=September 1952|title=[Comment on a new theory of cardiac insufficiency.]|journal=Policlinico Prat|volume=59|pages=1223–4|pmid=13026471|author2=D. MAESTRINI|number=37}}</ref><ref name="MAESTRINI-1958Jan">{{cite journal|last=MAESTRINI|first=D.|date=January 1958|title=[The law of the heart from its discovery to the present time.]|journal=Minerva Med|volume=49|pages=Varia, 28–36|pmid=13516733|number=3–4}}</ref><ref name="MAESTRINI-1958Dec">{{cite journal|last=MAESTRINI|first=D.|date=December 1958|title=[Variations of cardiac volume dynamics in clinical practice, examined in the light of the law of the heart.]|journal=Minerva Cardioangiol|volume=6|pages=657–67|pmid=13643787|number=12}}</ref><ref name="MAESTRINI-1959Feb">{{cite journal|last=MAESTRINI|first=D.|date=February 1959|title=[S. Baglioni and the law of the heart.]|journal=Policlinico Prat|volume=66|pages=224–30|pmid=13645276|number=7}}</ref><ref name="MAESTRINI-1959Oct">{{cite journal|last=MAESTRINI|first=D.|date=October 1959|title=[On cardiac dynamics in the phase preceding right hypertrophy and on its electrocardiographic aspect in man.]|journal=Policlinico Prat|volume=66|pages=1409–13|pmid=13853750}}</ref>{{citation overkill|date=March 2021}} Otto Frank's contributions are derived from his 1895 experiments on frog hearts. In order to relate the work of the heart to skeletal muscle mechanics, Frank observed changes in diastolic pressure with varying volumes of the frog ventricle. His data was analyzed on a pressure-volume diagram, which resulted in his description of peak isovolumic pressure and its effects on ventricular volume.<ref name=":2" /> Starling experimented on intact mammalian hearts, such as from dogs, to understand why variations in arterial pressure, heart rate, and temperature do not affect the relatively constant cardiac output.<ref name=":2" /> More than 30 years before the development of the [[Sliding filament theory|sliding filament model]] of muscle contraction and the understanding of the relationship between active tension and sarcomere length, Starling hypothesized in 1914, "the mechanical energy set free in the passage from the resting to the active state is a function of the length of the fiber." Starling used a volume-pressure diagram to construct a length-tension diagram from his data. Starling's data and associated diagrams, provided evidence that the length of the muscle fibers, and resulting tension, altered the systolic pressure.<ref>{{Cite book|url=https://books.google.com/books?id=54mxMgO5H_YC&q=%C2%A0the+mechanical+energy+set+free+in+the+passage+from+the+resting+to+the+active+state+is+a+function+of+the+length+of+the+fiber&pg=PA547|title=Medical Physiology, 2e Updated Edition E-Book: with STUDENT CONSULT Online Access|last1=Boron|first1=Walter F.|last2=Boulpaep|first2=Emile L.|date=2012-01-13|publisher=Elsevier Health Sciences|isbn=978-1455711819|language=en}}</ref> ==See also== * [[Starling equation]] * [[Total peripheral resistance]] ==References== {{reflist}} {{Cardiovascular physiology}} {{DEFAULTSORT:Frank-Starling law of the heart}} [[Category:Cardiovascular physiology]] [[Category:Mathematics in medicine]] </textarea><div class="templatesUsed"><div class="mw-templatesUsedExplanation"><p><span id="templatesused">Pages transcluded onto the current version 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