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321:. These activate the muscle fibres within the spindle. Gamma motor neurons supply only muscle fibres within the spindle, whereas beta motor neurons supply muscle fibres both within and outside of the spindle. Activation of the neurons causes a contraction and stiffening of the end parts of the muscle spindle muscle fibers.
498:) often develops, whereby the stretch reflex in flexor muscles of the arms and extensor muscles of the legs is overly sensitive. This results in abnormal postures, stiffness and contractures. Hypertonia may be the result of over-sensitivity of alpha motor neurons and interneurons to the Ia and II afferent signals.
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Dynamic gamma activity is adjusted proactively during movement preparation in order to facilitate execution of the planned action. For example, if the intended movement direction is associated with stretch of the spindle-bearing muscle, Ia afferent and stretch reflex sensitivity from this muscle is
51:
Mammalian muscle spindle showing typical position in a muscle (left), neuronal connections in spinal cord (middle) and expanded schematic (right). The spindle is a stretch receptor with its own motor supply consisting of several intrafusal muscle fibres. The sensory endings of a primary (group Ia)
388:
of the receptor-bearing muscle. The reflexly evoked activity in the alpha motor neurons is then transmitted via their efferent axons to the extrafusal fibers of the muscle, which generate force and thereby resist the stretch. The Ia afferent signal is also transmitted polysynaptically through
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Static gamma activity is a "temporal template" of the expected shortening and lengthening of the receptor-bearing muscle. Dynamic gamma activity turns on and off abruptly, sensitizing spindle afferents to the onset of muscle lengthening and departures from the intended movement
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Gamma motor neurons are activated according to the novelty or difficulty of a task. Whereas static gamma motor neurons are continuously active during routine movements such as locomotion, dynamic gamma motorneurons tend to be activated more during difficult tasks, increasing Ia
429:
How does the central nervous system control gamma fusimotor neurons? It has been difficult to record from gamma motor neurons during normal movement because they have very small axons. Several theories have been proposed, based on recordings from spindle afferents.
52:
afferent and a secondary (group II) afferent coil around the non-contractile central portions of the intrafusal fibres. Gamma motor neurons activate the intrafusal muscle fibres, changing the resting firing rate and stretch-sensitivity of the afferents.
324:
Fusimotor neurons are classified as static or dynamic according to the type of muscle fibers they innervate and their effects on the responses of the Ia and II sensory neurons innervating the central, non-contractile part of the muscle spindle.
659:
Purves, Dale; Augustine, George J; Fitzpatrick, David; Hall, William C; Lamantia, Anthony Samuel; Mooney, Richard D; Platt, Michael L; White, Leonard E, eds. (2018). "Chapter 9 - The
Somatosensory System: Touch and Proprioception".
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The function of the gamma motor neurons is not to supplement the force of muscle contraction provided by the extrafusal fibers, but to modify the sensitivity of the muscle spindle sensory afferents to stretch. Upon release of
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by the active gamma motor neuron, the end portions of the intrafusal muscle fibers contract, thus elongating the non-contractile central portions (see "fusimotor action" schematic below). This opens stretch-sensitive
380:. Likewise, secondary type II sensory fibers respond to muscle length changes (but with a smaller velocity-sensitive component) and transmit this signal to the spinal cord. The Ia afferent signals are transmitted
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at either or both of the ends of the intrafusal muscle fibers and regulate the sensitivity of the sensory afferents, which are located in the non-contractile central (equatorial) region.
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Here it is posited that gamma motor neurons are activated in parallel with alpha motor neurons to maintain the firing of spindle afferents when the extrafusal muscles shorten.
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When a muscle is stretched, primary type Ia sensory fibers of the muscle spindle respond to both changes in muscle length and velocity and transmit this activity to the
634:
Pearson, Keir G; Gordon, James E (2013). "35 - Spinal
Reflexes". In Kandel, Eric R; Schwartz, James H; Jessell, Thomas M; Siegelbaum, Steven A; Hudspeth, AJ (eds.).
302:(medium diameter) end adjacent to the central regions of the static bag and chain fibres. These fibres send information by stretch-sensitive mechanically-gated
131:
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Heckmann CJ, Gorassini MA, Bennett DJ (February 2005). "Persistent inward currents in motoneuron dendrites: implications for motor output".
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reduced. Gamma fusimotor control therefore allows for the independent preparatory tuning of muscle stiffness according to task goals.
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fibers. They increase the firing rate of Ia and II afferents at a given muscle length (see schematic of fusimotor action below).
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Mancall, Elliott L; Brock, David G, eds. (2011). "Chapter 2 - Overview of the
Microstructure of the Nervous System".
17:
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Prochazka, A. (1996). "Proprioceptive feedback and movement regulation". In Rowell, L.; Sheperd, J.T. (eds.).
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393:(Ia inhibitory interneurons), which inhibit alpha motorneurons of antagonist muscles, causing them to relax.
298:(large diameter) spiral around all intrafusal muscle fibres, ending near the middle of each fibre. Secondary
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intrafusal muscle fibers. They increase the stretch-sensitivity of the Ia afferents by stiffening the bag
863:"Static and dynamic gamma-motor output to ankle flexor muscles during locomotion in the decerebrate cat"
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178:. The responses of muscle spindles to changes in length also play an important role in regulating the
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that primarily detect changes in the length of the muscle. They convey length information to the
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840:. Handbook of physiology. New York: American Physiological Society. pp. 89–127.
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firing, thus increasing the stretch-sensitivity of the muscle spindle afferents.
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912:"Goal-dependent tuning of muscle spindle receptors during movement preparation"
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Hulliger M (1984). "The mammalian muscle spindle and its central control".
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For the class of neurons characterized by a large spindle-shaped body, see
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Gray's
Clinical Neuroanatomy: The Anatomic Basis for Clinical Neuroscience
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https://www.ualberta.ca/~aprochaz/research_interactive_receptor_model.html
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The motor part of the spindle is provided by motor neurons: up to a dozen
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Innervated muscle structure involved in reflex actions and proprioception
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which spiral around muscle fibres within the spindle, and secondary
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It is also believed that muscle spindles play a critical role in
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Reviews of
Physiology, Biochemistry and Pharmacology, Volume 86
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638:(5th ed.). United States: McGraw-Hill. pp. 794–795.
243:. The regular muscle fibers outside of the spindle are called
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Taylor A, Durbaba R, Ellaway PH, Rawlinson S (March 2006).
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Light microscope photograph of a muscle spindle. HE stain.
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The muscle spindle has both sensory and motor components.
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Exercise: Regulation and
Integration of Multiple Systems
789:"Human muscle spindle response in a motor learning task"
664:(6th ed.). Sinauer Associates. pp. 201–202.
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of the endings, thereby increasing the probability of
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174:. This information can be processed by the brain as
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329:The static axons innervate the chain or static bag
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687:"Functional properties of human muscle spindles"
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411:of the sensory endings, leading to an influx of
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685:Macefield, VG; Knellwolf, TP (1 August 2018).
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585:Arthur Prochazka's Lab, University of Alberta
355:also terminate in muscle spindles; they make
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267:, of which there are three types: dynamic
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1020:at the U.S. National Library of Medicine
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787:Vallbo AB, al-Falahe NA (February 1990).
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490:or spinal cord injury in humans, spastic
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451:Fusimotor template of intended movement.
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197:Sensory information conveyed by primary
910:Papaioannou, S.; Dimitriou, M. (2021).
275:fibers), static nuclear bag fibers (bag
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376:in the form of changes in the rate of
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610:. Elsevier Saunders. pp. 29–30.
263:Muscle spindles are composed of 5-14
227:Muscle spindles are found within the
215:and to a lesser extent by one or two
1636:Connective tissue in skeletal muscle
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247:. Muscle spindles have a capsule of
211:within the spindle by up to a dozen
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336:The dynamic axons innervate the bag
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459:Goal-directed preparatory control.
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730:. Vol. 101. pp. 1–110.
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182:, for example, by activating
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636:Principles of Neural Science
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397:Sensitivity modification
245:extrafusal muscle fibers
241:intrafusal muscle fibers
1865:Fukutin-related protein
1184:Intrafusal muscle fiber
1823:Sarcoplasmic reticulum
1652:Neuromuscular junction
1560:elastic filament/titin
1282:Vascular smooth muscle
929:10.1126/sciadv.abe0401
300:type II sensory fibers
296:type Ia sensory fibers
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235:. Muscle spindles are
203:type II sensory fibers
199:type Ia sensory fibers
180:contraction of muscles
168:central nervous system
139:Anatomical terminology
1555:thick filament/myosin
704:10.1152/jn.00071.2018
482:Clinical significance
349:Efferent nerve fibers
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172:afferent nerve fibers
162:within the body of a
96:fusus neuromuscularis
446:stretch-sensitivity.
281:nuclear chain fibers
1550:thin filament/actin
1536:(a, i, and h bands;
1189:Nuclear chain fiber
1103:Merkel nerve ending
418:. This raises the
386:alpha motor neurons
353:gamma motor neurons
315:gamma motor neurons
253:Golgi tendon organs
213:gamma motor neurons
1083:Lamellar corpuscle
736:10.1007/bfb0027694
581:Animated version:
344:intrafusal fibers.
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269:nuclear bag fibers
217:beta motor neurons
120:H3.11.06.0.00018
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1113:Bulbous corpuscle
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1059:Sensory receptors
985:10.1002/mus.20261
745:978-3-540-13679-8
645:978-0-07-139011-8
617:978-1-4160-4705-6
502:Additional images
496:spastic paralysis
420:resting potential
378:action potentials
319:fusimotor neurons
249:connective tissue
160:stretch receptors
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1362:
1359:
1357:
1354:
1352:
1349:
1348:
1347:
1344:
1343:
1341:
1338:
1334:
1331:
1329:extracellular
1325:
1322:
1320:
1315:
1311:
1308:
1306:
1300:
1297:
1295:
1289:
1283:
1280:
1278:
1275:
1274:
1272:
1270:
1264:
1260:
1259:Muscle tissue
1253:
1248:
1246:
1241:
1239:
1234:
1233:
1230:
1218:
1215:
1213:
1210:
1209:
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1195:
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1190:
1187:
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1175:
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1128:
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1078:
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1064:
1060:
1053:
1048:
1046:
1041:
1039:
1034:
1033:
1030:
1023:
1019:
1016:
1015:
1002:
998:
994:
990:
986:
982:
977:
972:
969:(2): 135–56.
968:
964:
957:
949:
945:
940:
935:
930:
925:
921:
917:
913:
906:
898:
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868:
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857:
849:
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824:
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815:
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747:
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671:9781605353807
667:
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584:
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448:
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440:
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431:
427:
425:
421:
417:
414:
410:
405:
404:acetylcholine
394:
392:
387:
383:
379:
375:
360:
358:
354:
350:
335:
328:
327:
326:
322:
320:
316:
311:
309:
305:
301:
297:
288:
284:
282:
279:fibers), and
270:
266:
265:muscle fibers
256:
254:
250:
246:
242:
238:
234:
230:
218:
214:
210:
209:muscle fibres
206:
204:
200:
196:
195:
194:
191:
189:
185:
184:motor neurons
181:
177:
173:
169:
165:
161:
157:
146:
140:
136:
133:
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128:
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118:
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100:
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83:
80:
77:
75:
71:
68:
65:
61:
56:
48:
43:
38:
33:
19:
1729:Muscle fiber
1661:
1499:
1430:Dystrobrevin
1383:Dystroglycan
1217:Baroreceptor
1178:
1089:Light touch
966:
963:Muscle Nerve
962:
956:
919:
915:
905:
870:
866:
856:
837:
831:
796:
792:
782:
727:
721:
694:
690:
680:
662:Neuroscience
661:
654:
635:
607:
601:
577:
485:
473:sensorimotor
470:
458:
450:
442:
435:
428:
409:ion channels
400:
391:interneurons
371:
347:
323:
318:
312:
304:ion-channels
293:
262:
226:
192:
155:
154:
95:
1780:Muscle cell
1755:Myofilament
1574:Tropomyosin
1545:Myofilament
1346:Sarcoglycan
1174:Golgi organ
1148:Temperature
1138:Nociceptors
551:Alpha fiber
527:Gamma fiber
476:development
467:Development
454:trajectory.
374:spinal cord
259:Composition
85:Identifiers
1850:Telethonin
1811:Sarcolemma
1806:Sarcoplasm
1739:extrafusal
1734:intrafusal
1696:Myocardium
1657:Motor unit
1631:Endomysium
1626:Perimysium
1514:Caveolin 3
1447:Syntrophin
1425:Dystrophin
1277:Calmodulin
1212:Hair cells
1079:Vibration
867:J. Physiol
799:: 553–68.
793:J. Physiol
593:References
492:hypertonia
1855:Dysferlin
1838:ungrouped
1760:Sarcomere
1746:Myofibril
1706:Nebulette
1616:Epimysium
1532:Sarcomere
1484:Dysbindin
1479:Syncoilin
1396:Sarcospan
1327:Membrane/
1314:Costamere
1099:Pressure
971:CiteSeerX
772:ignored (
762:cite book
223:Structure
1884:Category
1845:Myotilin
1816:T-tubule
1776:Myoblast
1621:Fascicle
1581:Troponin
1500:related:
1303:Skeletal
1292:Striated
1109:Stretch
1001:17828664
993:15736297
948:33627426
916:Sci. Adv
897:16423858
713:29668385
539:1A fiber
384:to many
363:Function
357:synapses
294:Primary
237:fusiform
186:via the
1860:Fukutin
1686:Cardiac
1645:General
1565:nebulin
939:7904268
888:1805796
823:2140862
814:1190101
754:6240757
306:of the
108:D009470
63:Part of
58:Details
1836:Other/
1801:Desmin
1688:muscle
1305:muscle
1294:muscle
1269:muscle
1267:Smooth
1024:(MeSH)
999:
991:
973:
946:
936:
895:
885:
844:
821:
811:
752:
742:
711:
668:
642:
614:
488:stroke
486:After
413:sodium
74:System
67:Muscle
1794:Other
1769:Cells
1722:Fiber
1205:Other
1067:Touch
997:S2CID
569:Notes
308:axons
231:of a
229:belly
143:[
132:83607
91:Latin
1715:Both
1509:NOS1
1376:SGCZ
1371:SGCG
1366:SGCE
1361:SGCD
1356:SGCB
1351:SGCA
1319:DAPC
1125:Pain
989:PMID
944:PMID
893:PMID
842:ISBN
819:PMID
774:help
750:PMID
740:ISBN
709:PMID
666:ISBN
640:ISBN
612:ISBN
416:ions
271:(bag
170:via
158:are
103:MeSH
1337:DAP
981:doi
934:PMC
924:doi
883:PMC
875:doi
871:571
809:PMC
801:doi
797:421
732:doi
699:doi
695:120
457:4)
449:3)
441:2)
434:1)
351:of
127:FMA
1886::
1472:G2
1467:G1
1462:B2
1457:B1
995:.
987:.
979:.
967:31
965:.
942:.
932:.
918:.
914:.
891:.
881:.
869:.
865:.
817:.
807:.
795:.
791:.
766::
764:}}
760:{{
748:.
738:.
707:.
693:.
689:.
626:^
478:.
310:.
283:.
115:TH
1778:/
1753:/
1596:I
1591:C
1586:T
1534:/
1452:A
1440:B
1435:A
1339::
1316:/
1251:e
1244:t
1237:v
1051:e
1044:t
1037:v
1003:.
983::
950:.
926::
920:7
899:.
877::
850:.
825:.
803::
776:)
756:.
734::
715:.
701::
674:.
648:.
620:.
494:(
342:1
338:1
331:2
277:2
273:1
147:]
34:.
20:)
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