239:
legs in stance. Tetrapod coordination (when 4 legs are in stance) is where diagonally opposite pairs of legs swing together. Wave (sometimes called a metachronal wave) describes walking where only 1 leg enters swing at a time. This movement propagates from back to front on side of the body and then the opposite. Stick
Insects, a larger hexapod, only shows a tripod gait during the larval stage. As adults at low speeds, they are most likely to walk in a metachronal wave, where only 1 leg swings at a time. At higher speeds, they walk in a tetrapod coordination with 2 legs paired in swing or a metachronal wave, only moving one leg at a time.
231:
bounding gait). Lateral sequence gaits during walking and running are most common in mammals, but arboreal mammals such as monkeys, some opossums, and kinkajous use diagonal sequence walks for enhanced stability. Diagonal sequence walks and runs (aka trots) are most frequently used by sprawling tetrapods such as salamanders and lizards, due to the lateral oscillations of their bodies during movement. Bipeds are a unique case, and most bipeds will display only three gaits—walking, running, and hopping—during natural locomotion. Other gaits, such as human skipping, are not used without deliberate effort.
583:
1212:
38:
122:
81:
classified according to footfall patterns, but recent studies often prefer definitions based on mechanics. The term typically does not refer to limb-based propulsion through fluid mediums such as water or air, but rather to propulsion across a solid substrate by generating reactive forces against it (which can apply to walking while underwater as well as on land).
238:
where 3 legs swing together while 3 legs remain on the ground in stance. However, variability in gait is continuous. Flies do not show distinct transitions between gaits but are more likely to walk in a tripod configuration at higher speeds. At lower speeds, they are more likely to walk with 4 or 5
303:
mammals moving from a walk to a run to a gallop as speed increases. Each of these gaits has an optimum speed, at which the minimum calories per metre are consumed, and costs increase at slower or faster speeds. Gait transitions occur near the speed where the cost of a fast walk becomes higher than
80:
that prevent use of certain gaits, or simply due to evolved innate preferences as a result of habitat differences. While various gaits are given specific names, the complexity of biological systems and interacting with the environment make these distinctions "fuzzy" at best. Gaits are typically
230:
Animals typically use different gaits in a speed-dependent manner. Almost all animals are capable of symmetrical gaits, while asymmetrical gaits are largely confined to mammals, who are capable of enough spinal flexion to increase stride length (though small crocodilians are capable of using a
292:
104:
pioneered the contemporary scientific analysis and the classification of gaits. The movement of each limb was partitioned into a stance phase, where the foot was in contact with the ground, and a swing phase, where the foot was lifted and moved forwards. Each limb must complete a
173:, lizards and salamanders must expand and contract their body wall in order to force air in and out of their lungs, but these are the same muscles used to laterally undulate the body during locomotion. Thus, they cannot move and breathe at the same time, a situation called
109:, otherwise one limb's relationship to the others can change with time, and a steady pattern cannot occur. Thus, any gait can completely be described in terms of the beginning and end of stance phase of three limbs relative to a cycle of a reference limb, usually the left
215:
202:
156:
relationship between the limb pairs. If the same-side forelimbs and hindlimbs initiate stance phase at the same time, the phase is 0 (or 100%). If the same-side forelimb contacts the ground half of the cycle later than the hindlimb, the phase is 50%.
84:
Due to the rapidity of animal movement, simple direct observation is rarely sufficient to give any insight into the pattern of limb movement. In spite of early attempts to classify gaits based on footprints or the sound of footfalls, it was not until
144:-hindlimb phase relationship. Duty factor is simply the percent of the total cycle which a given foot is on the ground. This value will usually be the same for forelimbs and hindlimbs unless the animal is moving with a specially trained gait or is
255:. In this scheme, movements are divided into walking and running. Walking gaits are all characterized by a "vaulting" movement of the body over the legs, frequently described as an inverted pendulum (displaying fluctuations in kinetic and
133:. In a symmetrical gait, the left and right limbs of a pair alternate, while in an asymmetrical gait, the limbs move together. Asymmetrical gaits are sometimes termed "leaping gaits", due to the presence of a suspended phase.
429:
Tasch, U.; Moubarak, P.; Tang, W.; Zhu, L.; Lovering, R. M.; Roche, J.; Bloch, R. J. (2008). "An
Instrument That Simultaneously Measures Spatiotemporal Gait Parameters and Ground Reaction Forces of Locomoting Rats".
221:
220:
217:
216:
222:
206:
204:
331:
model of running, "walks" and "runs" are seen in animals with 2, 4, 6, or more legs. The term "gait" has even been applied to flying and swimming organisms that produce distinct patterns of wake
557:
Ayali A, Borgmann A, Buschges A, Cousin-Fuchs E, Daun-Gruhn S, Holmes P (2015). "The comparative investigation of the stick insect and cockroach models in study of animal locomotion".
219:
205:
394:
Hildebrand, Milton (1 December 1989). "The
Quadrupedal Gaits of Vertebrates: The timing of leg movements relates to balance, body shape, agility, speed, and energy expenditure".
1113:
802:
Cavagna, G. A.; Heglund, N. C.; Taylor, R. C. (1977). "Mechanical work in terrestrial locomotion: two basic mechanisms for minimizing energy expenditure".
203:
1469:
1070:
218:
1474:
954:
447:
189:
to act as a piston, inflating and deflating the lungs as the animal's spine flexes and extends, increasing ventilation and allowing greater
860:
604:
1106:
626:
304:
the cost of a slow run. Unrestrained animals will typically move at the optimum speed for their gait to minimize energy cost. The
152:. Duty factors over 50% are considered a "walk", while those less than 50% are considered a run. Forelimb-hindlimb phase is the
1500:
234:
Hexapod gaits have also been well characterized, particularly for drosophila and stick insects (Phasmatodea). Drosophila use a
463:
Strauss R, Heisenberg M (August 1990). "Coordination of legs during straight walking and turning in
Drosophila melanogaster".
1099:
432:
Volume 2: Automotive
Systems; Bioengineering and Biomedical Technology; Computational Mechanics; Controls; Dynamical Systems
129:
Gaits are generally classed as "symmetrical" and "asymmetrical" based on limb movement. These terms have nothing to do with
1211:
1233:
986:
597:
591:
1325:
1161:
1080:
608:
853:
166:
251:
and force-plate records has given rise to an alternative classification scheme, based on the mechanics of the
641:
Hildebrand, M. (1989). "Vertebrate locomotion an introduction how does an animal's body move itself along?".
174:
125:
Gait graphs in the style of
Hildebrand. Dark areas indicate times of contact, bottom axis is % of cycle
1464:
1408:
1403:
1388:
1020:
964:
949:
869:
90:
263:. In running, the kinetic and potential energy fluctuate in-phase, and the energy change is passed on to
1479:
944:
31:
101:
165:
Gait choice can have effects beyond immediate changes in limb movement and speed, notably in terms of
1418:
1330:
737:
679:
773:
Blickhan, R.; Full, R. J. (1993). "Similarity in multilegged locomotion: Bouncing like a monopode".
1413:
1340:
939:
893:
846:
328:
280:
137:
130:
76:, and energetic efficiency. Different animal species may use different gaits due to differences in
93:
began taking rapid series of photographs that proper scientific examination of gaits could begin.
1428:
1393:
1146:
827:
790:
761:
695:
658:
488:
411:
344:
317:
308:
is used to compare the energetics of different gaits, as well as the gaits of different animals.
170:
86:
1444:
1423:
1383:
1263:
1181:
819:
753:
716:
707:
Carrier, D. (1987). "Lung ventilation during walking and running in four species of lizards".
539:
480:
443:
374:
324:
305:
291:
65:
1454:
1228:
811:
782:
745:
687:
650:
562:
529:
519:
472:
435:
403:
354:
260:
256:
252:
106:
68:
over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed,
49:
1398:
1302:
1285:
1156:
1075:
913:
178:
741:
683:
1459:
1350:
1273:
1220:
1191:
1151:
1126:
1031:
534:
507:
182:
53:
670:
Hoyt, D. F.; Taylor, R. C. (1981). "Gait and the energetics of locomotion in horses".
1494:
1280:
1196:
1186:
991:
349:
831:
815:
794:
765:
699:
492:
1345:
1057:
903:
654:
364:
149:
145:
61:
1357:
1307:
1248:
1243:
1238:
1047:
1026:
1011:
1001:
996:
959:
897:
300:
235:
566:
37:
1258:
1201:
1171:
1166:
1052:
1016:
369:
359:
320:
248:
185:. In contrast, the spinal flexion of a galloping mammal causes the abdominal
17:
1362:
1335:
838:
749:
190:
121:
73:
543:
439:
757:
728:
Bramble, D. M.; Carrier, D. R (1983). "Running and breathing in mammals".
720:
484:
1367:
1290:
1268:
1176:
1006:
823:
332:
276:
247:
While gaits can be classified by footfall, new work involving whole-body
141:
110:
1091:
524:
1295:
978:
923:
918:
908:
786:
662:
476:
415:
186:
77:
69:
57:
1253:
691:
272:
264:
407:
290:
213:
200:
36:
1317:
1130:
268:
153:
30:
This article is about gaits of all animals. For other uses, see
1095:
842:
1138:
1122:
576:
506:
DeAngelis BD, Zavatone-Veth JA, Clark DA (June 2019).
1437:
1376:
1316:
1219:
1137:
1040:
977:
932:
886:
876:
316:In spite of the differences in leg number shown in
259:which are out of phase), a mechanism described by
279:acting as springs (thus it is described by the
226:Alternating tripod gait of walking desert ants.
1107:
854:
299:Speed generally governs gait selection, with
210:A hamster walking on a transparent treadmill.
8:
27:Pattern of movement of the limbs of animals
1114:
1100:
1092:
883:
861:
847:
839:
1470:Tradeoffs for locomotion in air and water
627:Learn how and when to remove this message
533:
523:
590:This article includes a list of general
120:
386:
1071:Animal locomotion on the water surface
181:, can circumvent this restriction via
140:for gait are the duty factor and the
7:
775:Journal of Comparative Physiology A
465:Journal of Comparative Physiology A
596:it lacks sufficient corresponding
25:
559:Current Opinion in Insect Science
1210:
581:
243:Energy-based gait classification
107:cycle in the same length of time
816:10.1152/ajpregu.1977.233.5.R243
1:
161:Physiological effects of gait
655:10.1093/bioscience/39.11.764
987:Comparative foot morphology
197:Differences between species
1517:
1326:Flying and gliding animals
1162:Fin and flipper locomotion
567:10.1016/j.cois.2015.07.004
29:
1208:
1066:
870:Animal locomotion on land
750:10.1126/science.6849136
611:more precise citations.
177:, though some, such as
1501:Terrestrial locomotion
1465:Terrestrial locomotion
1409:Evolution of cetaceans
1404:Origin of avian flight
1389:Evolution of tetrapods
950:Rectilinear locomotion
440:10.1115/ESDA2008-59085
296:
227:
211:
169:. Because they lack a
126:
42:
1480:Undulatory locomotion
1429:Homologous structures
945:Undulatory locomotion
327:model of walking and
294:
225:
209:
124:
40:
32:Gait (disambiguation)
1424:Analogous structures
1419:Convergent evolution
709:Experimental Biology
175:Carrier's constraint
1475:Rotating locomotion
1414:Comparative anatomy
940:Concertina movement
894:Arboreal locomotion
742:1983Sci...219..251B
684:1981Natur.292..239H
525:10.7554/eLife.46409
323:, according to the
131:left-right symmetry
91:Étienne-Jules Marey
1394:Evolution of birds
1147:Aquatic locomotion
787:10.1007/bf00197760
477:10.1007/BF00192575
434:. pp. 45–49.
345:Bipedal gait cycle
312:Non-tetrapod gaits
297:
228:
212:
127:
87:Eadweard Muybridge
48:is the pattern of
43:
1488:
1487:
1445:Animal locomotion
1384:Evolution of fish
1264:facultative biped
1089:
1088:
973:
972:
736:(4582): 251–256.
678:(5820): 239–240.
637:
636:
629:
449:978-0-7918-4836-4
375:Parkinsonian gait
325:inverted pendulum
306:cost of transport
281:spring-mass model
223:
207:
102:Milton Hildebrand
16:(Redirected from
1508:
1455:Robot locomotion
1229:Limb development
1214:
1187:Lobe-finned fish
1116:
1109:
1102:
1093:
884:
863:
856:
849:
840:
835:
810:(5): R243–R261.
798:
769:
724:
703:
692:10.1038/292239a0
666:
632:
625:
621:
618:
612:
607:this article by
598:inline citations
585:
584:
577:
571:
570:
554:
548:
547:
537:
527:
503:
497:
496:
460:
454:
453:
426:
420:
419:
391:
355:Gait abnormality
261:Giovanni Cavagna
257:potential energy
224:
208:
41:Elephant walking
21:
1516:
1515:
1511:
1510:
1509:
1507:
1506:
1505:
1491:
1490:
1489:
1484:
1433:
1399:Origin of birds
1372:
1312:
1234:Limb morphology
1215:
1206:
1192:Ray-finned fish
1157:Fish locomotion
1133:
1120:
1090:
1085:
1076:Fish locomotion
1062:
1036:
969:
928:
914:Knuckle-walking
872:
867:
801:
772:
727:
706:
669:
649:(11): 764–765.
640:
633:
622:
616:
613:
603:Please help to
602:
586:
582:
575:
574:
556:
555:
551:
505:
504:
500:
462:
461:
457:
450:
428:
427:
423:
408:10.2307/1311182
393:
392:
388:
383:
341:
314:
295:Bison galloping
289:
245:
214:
201:
199:
191:oxygen exchange
179:monitor lizards
163:
119:
99:
35:
28:
23:
22:
15:
12:
11:
5:
1514:
1512:
1504:
1503:
1493:
1492:
1486:
1485:
1483:
1482:
1477:
1472:
1467:
1462:
1457:
1452:
1447:
1441:
1439:
1435:
1434:
1432:
1431:
1426:
1421:
1416:
1411:
1406:
1401:
1396:
1391:
1386:
1380:
1378:
1374:
1373:
1371:
1370:
1365:
1363:Pterosaur wing
1360:
1355:
1354:
1353:
1348:
1343:
1333:
1328:
1322:
1320:
1314:
1313:
1311:
1310:
1305:
1300:
1299:
1298:
1288:
1283:
1278:
1277:
1276:
1271:
1266:
1261:
1256:
1251:
1246:
1241:
1231:
1225:
1223:
1217:
1216:
1209:
1207:
1205:
1204:
1199:
1194:
1189:
1184:
1179:
1174:
1169:
1164:
1159:
1154:
1152:Cephalopod fin
1149:
1143:
1141:
1135:
1134:
1121:
1119:
1118:
1111:
1104:
1096:
1087:
1086:
1084:
1083:
1081:Volant animals
1078:
1073:
1067:
1064:
1063:
1061:
1060:
1055:
1050:
1044:
1042:
1038:
1037:
1035:
1034:
1029:
1024:
1014:
1009:
1004:
999:
994:
989:
983:
981:
975:
974:
971:
970:
968:
967:
962:
957:
952:
947:
942:
936:
934:
930:
929:
927:
926:
921:
916:
911:
906:
901:
890:
888:
881:
874:
873:
868:
866:
865:
858:
851:
843:
837:
836:
804:Am. J. Physiol
799:
781:(5): 509–517.
770:
725:
704:
667:
635:
634:
589:
587:
580:
573:
572:
549:
498:
455:
448:
421:
385:
384:
382:
379:
378:
377:
372:
367:
362:
357:
352:
347:
340:
337:
313:
310:
288:
285:
244:
241:
198:
195:
183:buccal pumping
162:
159:
118:
115:
98:
95:
72:, the need to
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
1513:
1502:
1499:
1498:
1496:
1481:
1478:
1476:
1473:
1471:
1468:
1466:
1463:
1461:
1458:
1456:
1453:
1451:
1448:
1446:
1443:
1442:
1440:
1436:
1430:
1427:
1425:
1422:
1420:
1417:
1415:
1412:
1410:
1407:
1405:
1402:
1400:
1397:
1395:
1392:
1390:
1387:
1385:
1382:
1381:
1379:
1375:
1369:
1366:
1364:
1361:
1359:
1356:
1352:
1349:
1347:
1344:
1342:
1339:
1338:
1337:
1334:
1332:
1329:
1327:
1324:
1323:
1321:
1319:
1315:
1309:
1306:
1304:
1301:
1297:
1294:
1293:
1292:
1289:
1287:
1284:
1282:
1279:
1275:
1272:
1270:
1267:
1265:
1262:
1260:
1257:
1255:
1252:
1250:
1247:
1245:
1242:
1240:
1237:
1236:
1235:
1232:
1230:
1227:
1226:
1224:
1222:
1218:
1213:
1203:
1200:
1198:
1197:Pectoral fins
1195:
1193:
1190:
1188:
1185:
1183:
1180:
1178:
1175:
1173:
1170:
1168:
1165:
1163:
1160:
1158:
1155:
1153:
1150:
1148:
1145:
1144:
1142:
1140:
1136:
1132:
1128:
1124:
1117:
1112:
1110:
1105:
1103:
1098:
1097:
1094:
1082:
1079:
1077:
1074:
1072:
1069:
1068:
1065:
1059:
1056:
1054:
1051:
1049:
1046:
1045:
1043:
1039:
1033:
1030:
1028:
1025:
1022:
1018:
1015:
1013:
1010:
1008:
1005:
1003:
1000:
998:
995:
993:
992:Arthropod leg
990:
988:
985:
984:
982:
980:
976:
966:
963:
961:
958:
956:
953:
951:
948:
946:
943:
941:
938:
937:
935:
931:
925:
922:
920:
917:
915:
912:
910:
907:
905:
902:
899:
895:
892:
891:
889:
885:
882:
879:
875:
871:
864:
859:
857:
852:
850:
845:
844:
841:
833:
829:
825:
821:
817:
813:
809:
805:
800:
796:
792:
788:
784:
780:
776:
771:
767:
763:
759:
755:
751:
747:
743:
739:
735:
731:
726:
722:
718:
714:
710:
705:
701:
697:
693:
689:
685:
681:
677:
673:
668:
664:
660:
656:
652:
648:
644:
639:
638:
631:
628:
620:
610:
606:
600:
599:
593:
588:
579:
578:
568:
564:
560:
553:
550:
545:
541:
536:
531:
526:
521:
517:
513:
509:
502:
499:
494:
490:
486:
482:
478:
474:
471:(3): 403–12.
470:
466:
459:
456:
451:
445:
441:
437:
433:
425:
422:
417:
413:
409:
405:
401:
397:
390:
387:
380:
376:
373:
371:
368:
366:
363:
361:
358:
356:
353:
351:
350:Gait analysis
348:
346:
343:
342:
338:
336:
334:
330:
326:
322:
319:
311:
309:
307:
302:
293:
286:
284:
282:
278:
274:
270:
266:
262:
258:
254:
250:
242:
240:
237:
232:
196:
194:
192:
188:
184:
180:
176:
172:
168:
160:
158:
155:
151:
147:
143:
139:
134:
132:
123:
116:
114:
112:
108:
103:
96:
94:
92:
88:
82:
79:
75:
71:
67:
63:
59:
55:
51:
47:
39:
33:
19:
18:Leaping gaits
1449:
904:Hand-walking
877:
807:
803:
778:
774:
733:
729:
715:(1): 33–42.
712:
708:
675:
671:
646:
642:
623:
614:
595:
561:(12): 1–10.
558:
552:
515:
511:
508:"Drosophila"
501:
468:
464:
458:
431:
424:
399:
395:
389:
365:Gait (human)
315:
298:
246:
233:
229:
164:
150:decelerating
146:accelerating
135:
128:
100:
83:
60:, including
45:
44:
1358:Insect wing
1308:Webbed foot
1249:unguligrade
1244:plantigrade
1239:digitigrade
1048:Canine gait
1021:Facultative
1007:Unguligrade
1002:Plantigrade
997:Digitigrade
965:Other modes
960:Sidewinding
898:Brachiation
617:August 2009
609:introducing
402:(11): 766.
329:spring-mass
321:vertebrates
318:terrestrial
301:quadrupedal
236:tripod gait
167:ventilation
1286:Cephalopod
1202:Pelvic fin
1172:Dorsal fin
1167:Caudal fin
1058:Human gait
1053:Horse gait
643:BioScience
592:references
396:BioScience
381:References
370:Horse gait
360:Gait (dog)
287:Energetics
249:kinematics
66:locomotion
1377:Evolution
1336:Bird wing
1281:Arthropod
1274:quadruped
1032:Quadruped
277:ligaments
171:diaphragm
138:variables
117:Variables
64:, during
1495:Category
1368:Wingspan
1351:feathers
1346:skeleton
1331:Bat wing
1291:Tetrapod
1177:Fish fin
1041:Specific
832:15842774
795:19751464
766:23551439
700:26841475
544:31250807
493:12965869
339:See also
333:vortices
253:movement
154:temporal
142:forelimb
136:The key
111:hindlimb
97:Overview
74:maneuver
50:movement
1438:Related
1296:dactyly
1182:Flipper
979:Anatomy
955:Rolling
933:Legless
924:Walking
919:Running
909:Jumping
758:6849136
738:Bibcode
730:Science
721:3666097
680:Bibcode
663:1311182
605:improve
535:6598772
485:2121965
416:1311182
273:tendons
265:muscles
187:viscera
78:anatomy
70:terrain
58:animals
52:of the
1460:Samara
1269:triped
1254:uniped
1027:Triped
1012:Uniped
887:Legged
830:
824:411381
822:
793:
764:
756:
719:
698:
672:Nature
661:
594:, but
542:
532:
491:
483:
446:
414:
62:humans
1318:Wings
1303:Digit
1259:biped
1221:Limbs
1131:wings
1127:limbs
1017:Biped
880:class
828:S2CID
791:S2CID
762:S2CID
696:S2CID
659:JSTOR
512:eLife
489:S2CID
412:JSTOR
269:bones
54:limbs
1450:Gait
1341:keel
1139:Fins
1129:and
1123:Fins
878:Gait
820:PMID
754:PMID
717:PMID
540:PMID
481:PMID
444:ISBN
275:and
89:and
46:Gait
812:doi
808:233
783:doi
779:173
746:doi
734:219
688:doi
676:292
651:doi
563:doi
530:PMC
520:doi
473:doi
469:167
436:doi
404:doi
283:).
148:or
56:of
1497::
1125:,
826:.
818:.
806:.
789:.
777:.
760:.
752:.
744:.
732:.
713:47
711:.
694:.
686:.
674:.
657:.
647:39
645:.
538:.
528:.
518:.
514:.
510:.
487:.
479:.
467:.
442:.
410:.
400:39
398:.
335:.
271:,
267:,
193:.
113:.
1115:e
1108:t
1101:v
1023:)
1019:(
900:)
896:(
862:e
855:t
848:v
834:.
814::
797:.
785::
768:.
748::
740::
723:.
702:.
690::
682::
665:.
653::
630:)
624:(
619:)
615:(
601:.
569:.
565::
546:.
522::
516:8
495:.
475::
452:.
438::
418:.
406::
34:.
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.