122:
221:, dating between 5.2 and 5.8 million years ago. The skeletal remains of this hominid exhibit a mosaic of morphological characteristics that would have been both adapted to an arboreal environment and walking upright terrestrially. The earliest evidence of a hominid exhibiting skeletal morphology capable of achieving orthograde posture dates to 9.5 million years ago, with the discovery of a Miocene ape, Dryopithecus in Can Llobateres, Spain.
323:. Evidence draws from three femur fragments, including the left shaft and head, and the head of the right femur. Linking the legs to the pelvis and lumbar vertebra, the femur quintessentially supports body weight as it is transferred from the pelvis to the knee and lower limbs. The femoral neck specifically, which connects the head of the femur to its primary shaft absorbs the force of impact when an upright biped assumes movement. In
25:
257:
316:, which produces the unique convex curvature seen in upright bipeds. The vertebral column of australopith fossils also share the curved morphology of modern humans. Lordosis in the lower lumbar spine centers the mass of the body on the lower joints such as the pelvis and femur such that the body is self-stabilizing and can remain upright.
306:
to the sacrum and pelvis. Primates with pronograde posture such as gorillas have four lumbar vertebrae that connect to twelve thoracic vertebrae. The difference in vertebra number results in a greater range of movement for humans with less thoracic vertebrae than gorillas with more lumbar vertebrae.
286:
The skull was an indicator of orthograde posture because of the location and orientation of the foramen magnum. The foramen magnum is the space in the skull that acts as the bridge to the central nervous system from the spinal cord to the brain. For animals with "pronograde posture, the foramen
242:
in that time period still used knuckle walking, a practice common in other apes. However habitual bipedalism in australopiths meant though they nested among the branches in trees at night, they moved with orthograde posture such that their hands could also be used for gathering, feeding, weight
264:
In order for animals to have the ability to walk upright, there are certain anatomical requirements. For mammals that exhibit orthograde posture, the scapula is more dorsally placed than in animals with a pronograde posture. The scapular index, the measure of width to length of the scapula, is
194:. This upright locomotion is called "orthograde posture". Orthograde posture in humans was made possible through millions of years of evolution. In order to walk upright with maximum efficiency, the skull, spine, pelvis, lower limbs, and feet all underwent evolutionary changes.
202:
The definition of orthograde posture can easily be derived from its roots “ortho-” meaning “upright” and “-grade” meaning “ascent.” This was true for the early hominidae, whose transition to upright walking took place approximately six to seven million years ago evident in
247:
and australopiths began to change, which is evident in morphological data accumulated from the remains of the different species. These major morphological changes differentiate them from pronograde hominin seen in the skull, vertebral column, pelvis, and femur fossils.
269:
is more flattened and the acromion process on the scapula is much larger. This is because there is more of a need for the deltoid muscle in orthograde posture, due to the availability of resource manipulation by the freeing up of hands.
287:
magnum is dorsally oriented, whereas in humans it is anteriorly located and forwardly inclined. In the Taung Child despite lacking the forward inclination seen in humans, the foramen magnum is also anteriorly oriented. Similarly in
243:
transfer, or balance during the day. From fossil evidence and hypotheses state that upright posture was a quintessential reaction to changes in environment and competition. Due to the more wooded barren savannahs of northern Africa,
311:
has six total lumbar vertebrae with also twelve thoracic vertebra
Another key characteristic that enforced upright posture in hominin was the shape of the lumbar vertebra. The “s” shape of the lumbar vertebra is called spinal
1038:
291:, the site of the space in the skull is even more human-like, inferiorly located such that the spinal cord would run perpendicularly to the ground. Relating this orientation to the
1048:
959:"Locomotion and posture from the common hominoid ancestor to fully modern hominins, with special reference to the last common panin/hominin ancestor"
164:, and they have a tendency to walk with their limbs swinging in parallel to one another. This differs from the manner of walking demonstrated by the
282:
in South Africa provided further evidence of bipedalism and orthograde posture. The skull belonged to a three-year-old child, later identified as
329:, the orientation of the head condyles of the broadened femur is wider and thicker in comparison to that of chimpanzees and other great apes.
691:
Falk D (2009). "The natural endocast of Taung (Australopithecus africanus): insights from the unpublished papers of
Raymond Arthur Dart".
42:
1028:
675:
108:
89:
1043:
190:
to use for balance). Disadvantages related to upright walking do exist for primates, since their primary mode of locomotion is
61:
238:
also practiced habitual bipedalism. These tree-dwellers were arboreal and inhabited the wooded areas of forest canopies. Some
46:
186:, when walking, walk upright, and their limbs swing in opposition to one another for balance (unlike monkeys, apes lack a
68:
917:
Pickford M, Senut B, Gommery D, Treil J (September 2002). "Bipedalism in
Orrorin tugenensis revealed by its femora".
75:
298:
More evidence in hominidae that enabled orthograde posture is present in the vertical column or lumbar vertebra of
35:
265:
decreased in animals exhibiting orthograde posture. This means that the scapula is broader than it is long. The
292:
57:
509:
Moyà-Solà S, Köhler M (January 1996). "A Dryopithecus skeleton and the origins of great-ape locomotion".
413:"Insights into the evolution of human bipedalism from experimental studies of humans and other primates"
295:
of hominin of the time, the position foramen magnum assisted in balance and supported upright posture.
215:
The first definitive evidence of habitual orthograde posture in human evolutionary lineage begins with
207:. These hominin were some of the first bipeds who propagated forward one leg at a time, step by step.
926:
622:
611:"Human bipedal instability in tree canopy environments is reduced by "light touch" fingertip support"
518:
465:
542:
303:
82:
609:
Johannsen L, Coward SR, Martin GR, Wing AM, Casteren AV, Sellers WI, et al. (April 2017).
1024:
988:
899:
848:
797:
756:
704:
671:
648:
591:
534:
491:
434:
157:
153:
1066:
978:
970:
934:
889:
879:
838:
828:
787:
746:
738:
696:
638:
630:
581:
573:
526:
481:
473:
424:
121:
1000:
930:
731:
Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
626:
566:
Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
522:
469:
983:
958:
894:
867:
843:
816:
751:
726:
643:
610:
586:
561:
486:
453:
191:
137:
938:
1060:
974:
727:"The cranial base of Australopithecus afarensis: new insights from the female skull"
387:
363:
546:
216:
279:
24:
821:
Proceedings of the
National Academy of Sciences of the United States of America
634:
477:
343:
338:
171:
454:"The evolution of the upright posture and gait--a review and a new synthesis"
302:. The human lumbar column consists of five vertebrae that connect the twelve
833:
256:
179:
992:
903:
884:
852:
801:
760:
742:
708:
652:
595:
577:
495:
438:
866:
Wagner H, Liebetrau A, Schinowski D, Wulf T, de
Lussanet MH (April 2012).
538:
313:
266:
239:
161:
868:"Spinal lordosis optimizes the requirements for a stable erect posture"
792:
775:
700:
324:
226:
175:
149:
429:
412:
530:
126:
774:
Williams SA, Middleton ER, Villamil CI, Shattuck MR (January 2016).
815:
Ward CV, Nalley TK, Spoor F, Tafforeau P, Alemseged Z (June 2017).
255:
183:
120:
187:
165:
152:
which is upright, with the independent motion of limbs. Both
18:
1021:
Windows On
Humanity: A Concise Introduction to Anthropology
560:Crompton RH, Sellers WI, Thorpe SK (October 2010).
49:. Unsourced material may be challenged and removed.
957:Crompton RH, Vereecke EE, Thorpe SK (April 2008).
319:The earliest habitual bipeds of the hominins were
1001:"What Does It Mean to Be Human? Walking Upright"
666:Bain GI, Itoi E, Di Giacomo G, Sugaya H (2015).
668:Normal and Pathological Anatomy of the Shoulder
720:
718:
8:
562:"Arboreality, terrestriality and bipedalism"
872:Theoretical Biology & Medical Modelling
1023:. New York, NY.: McGraw-Hill. p. 80.
982:
893:
883:
842:
832:
791:
780:American Journal of Physical Anthropology
750:
693:American Journal of Physical Anthropology
642:
585:
485:
428:
278:In 1924, the discovery of remains of the
109:Learn how and when to remove this message
776:"Vertebral numbers and human evolution"
355:
144:("right", "true", "straight") + Latin
148:(to walk) that describes a manner of
7:
47:adding citations to reliable sources
417:The Journal of Experimental Biology
14:
725:Kimbel WH, Rak Y (October 2010).
975:10.1111/j.1469-7580.2008.00870.x
23:
34:needs additional citations for
670:. Springer. pp. 403–413.
1:
939:10.1016/S1631-0683(02)00028-3
695:. 140 Suppl 49 (S49): 49–65.
252:Morphological characteristics
230:, australopithecines such as
1044:Dorland's Medical Dictionary
817:"Australopithecus afarensis"
224:Several million years after
284:Australopithecus africanus.
125:Orthograde locomotion of a
1083:
635:10.1038/s41598-017-01265-7
300:Australopithecus afarensis
289:Australopithecus afarensis
478:10.1007/s00114-009-0637-3
260:Annotated human shoulder.
211:Evolutionary significance
452:Niemitz C (March 2010).
1005:Smithsonian Institution
834:10.1073/pnas.1702229114
458:Die Naturwissenschaften
392:www.merriam-webster.com
368:www.merriam-webster.com
136:is a term derived from
919:Comptes Rendus Palevol
885:10.1186/1742-4682-9-13
743:10.1098/rstb.2010.0070
578:10.1098/rstb.2010.0035
411:Schmitt D (May 2003).
388:"Definition of -GRADE"
364:"Definition of ORTHO-"
261:
130:
274:Morphological changes
259:
124:
1051:on 20 February 2007.
786:(Suppl 61): S19-36.
321:Orrorin tungenenisis
218:Ardipithecus ramidus
58:"Orthograde posture"
43:improve this article
931:2002CRPal...1..191P
627:2017NatSR...7.1135J
523:1996Natur.379..156M
470:2010NW.....97..241N
1019:Kottak CP (2005).
963:Journal of Anatomy
793:10.1002/ajpa.22901
701:10.1002/ajpa.21184
615:Scientific Reports
326:Orrorin tugenensis
304:thoracic vertebrae
262:
227:Orrorin tugenensis
205:Orrorin tugenensis
131:
827:(23): 6000–6004.
737:(1556): 3365–76.
572:(1556): 3301–14.
430:10.1242/jeb.00279
423:(Pt 9): 1437–48.
158:Old World monkeys
119:
118:
111:
93:
16:Manner of walking
1074:
1052:
1047:. Archived from
1034:
1015:
1013:
1011:
996:
986:
943:
942:
914:
908:
907:
897:
887:
863:
857:
856:
846:
836:
812:
806:
805:
795:
771:
765:
764:
754:
722:
713:
712:
688:
682:
681:
663:
657:
656:
646:
606:
600:
599:
589:
557:
551:
550:
531:10.1038/379156a0
506:
500:
499:
489:
449:
443:
442:
432:
408:
402:
401:
399:
398:
384:
378:
377:
375:
374:
360:
114:
107:
103:
100:
94:
92:
51:
27:
19:
1082:
1081:
1077:
1076:
1075:
1073:
1072:
1071:
1057:
1056:
1055:
1037:
1031:
1018:
1009:
1007:
999:
956:
952:
950:Further reading
947:
946:
916:
915:
911:
865:
864:
860:
814:
813:
809:
773:
772:
768:
724:
723:
716:
690:
689:
685:
678:
665:
664:
660:
608:
607:
603:
559:
558:
554:
517:(6561): 156–9.
508:
507:
503:
451:
450:
446:
410:
409:
405:
396:
394:
386:
385:
381:
372:
370:
362:
361:
357:
352:
335:
293:encephalization
276:
254:
213:
200:
115:
104:
98:
95:
52:
50:
40:
28:
17:
12:
11:
5:
1080:
1078:
1070:
1069:
1059:
1058:
1054:
1053:
1035:
1030:978-0073258935
1029:
1016:
997:
953:
951:
948:
945:
944:
925:(4): 191–203.
909:
858:
807:
766:
714:
683:
677:978-3662457191
676:
658:
601:
552:
501:
444:
403:
379:
354:
353:
351:
348:
347:
346:
341:
334:
331:
275:
272:
253:
250:
212:
209:
199:
196:
192:quadrupedalism
160:are primarily
117:
116:
31:
29:
22:
15:
13:
10:
9:
6:
4:
3:
2:
1079:
1068:
1065:
1064:
1062:
1050:
1046:
1045:
1040:
1036:
1032:
1026:
1022:
1017:
1010:September 21,
1006:
1002:
998:
994:
990:
985:
980:
976:
972:
969:(4): 501–43.
968:
964:
960:
955:
954:
949:
940:
936:
932:
928:
924:
920:
913:
910:
905:
901:
896:
891:
886:
881:
877:
873:
869:
862:
859:
854:
850:
845:
840:
835:
830:
826:
822:
818:
811:
808:
803:
799:
794:
789:
785:
781:
777:
770:
767:
762:
758:
753:
748:
744:
740:
736:
732:
728:
721:
719:
715:
710:
706:
702:
698:
694:
687:
684:
679:
673:
669:
662:
659:
654:
650:
645:
640:
636:
632:
628:
624:
620:
616:
612:
605:
602:
597:
593:
588:
583:
579:
575:
571:
567:
563:
556:
553:
548:
544:
540:
536:
532:
528:
524:
520:
516:
512:
505:
502:
497:
493:
488:
483:
479:
475:
471:
467:
464:(3): 241–63.
463:
459:
455:
448:
445:
440:
436:
431:
426:
422:
418:
414:
407:
404:
393:
389:
383:
380:
369:
365:
359:
356:
349:
345:
342:
340:
337:
336:
332:
330:
328:
327:
322:
317:
315:
310:
309:Au. afarensis
305:
301:
296:
294:
290:
285:
281:
273:
271:
268:
258:
251:
249:
246:
245:O. tugenensis
241:
237:
236:Au. afarensis
233:
232:Au. africanus
229:
228:
222:
220:
219:
210:
208:
206:
197:
195:
193:
189:
185:
181:
177:
173:
169:
167:
163:
159:
155:
151:
147:
143:
139:
135:
128:
123:
113:
110:
102:
99:November 2015
91:
88:
84:
81:
77:
74:
70:
67:
63:
60: –
59:
55:
54:Find sources:
48:
44:
38:
37:
32:This article
30:
26:
21:
20:
1049:the original
1042:
1039:"orthograde"
1020:
1008:. Retrieved
1004:
966:
962:
922:
918:
912:
875:
871:
861:
824:
820:
810:
783:
779:
769:
734:
730:
692:
686:
667:
661:
618:
614:
604:
569:
565:
555:
514:
510:
504:
461:
457:
447:
420:
416:
406:
395:. Retrieved
391:
382:
371:. Retrieved
367:
358:
325:
320:
318:
308:
299:
297:
288:
283:
277:
263:
244:
235:
231:
225:
223:
217:
214:
204:
201:
170:
145:
141:
133:
132:
105:
96:
86:
79:
72:
65:
53:
41:Please help
36:verification
33:
621:(1): 1135.
280:Taung Child
172:Chimpanzees
397:2021-10-10
373:2021-10-10
350:References
344:Pronograde
339:Bipedalism
180:orangutans
134:Orthograde
69:newspapers
878:(1): 13.
1061:Category
993:18380868
904:22507595
853:28533391
802:26808105
761:20855310
709:19890860
653:28442732
596:20855304
496:20127307
439:12654883
333:See also
314:lordosis
267:rib cage
176:gorillas
162:arboreal
1067:Walking
984:2409101
927:Bibcode
895:3349546
844:5468642
752:2981961
644:5430707
623:Bibcode
587:2981953
547:4344346
539:8538764
519:Bibcode
487:2819487
466:Bibcode
240:hominin
150:walking
140:ὀρθός,
83:scholar
1027:
991:
981:
902:
892:
851:
841:
800:
759:
749:
707:
674:
651:
641:
594:
584:
545:
537:
511:Nature
494:
484:
437:
198:Origin
184:humans
142:orthos
129:lemur.
127:sifaka
85:
78:
71:
64:
56:
543:S2CID
146:gradi
138:Greek
90:JSTOR
76:books
1025:ISBN
1012:2015
989:PMID
900:PMID
849:PMID
798:PMID
757:PMID
705:PMID
672:ISBN
649:PMID
592:PMID
535:PMID
492:PMID
435:PMID
234:and
188:tail
182:and
166:apes
156:and
62:news
979:PMC
971:doi
967:212
935:doi
890:PMC
880:doi
839:PMC
829:doi
825:114
788:doi
784:159
747:PMC
739:doi
735:365
697:doi
639:PMC
631:doi
582:PMC
574:doi
570:365
527:doi
515:379
482:PMC
474:doi
425:doi
421:206
154:New
45:by
1063::
1041:.
1003:.
987:.
977:.
965:.
961:.
933:.
921:.
898:.
888:.
874:.
870:.
847:.
837:.
823:.
819:.
796:.
782:.
778:.
755:.
745:.
733:.
729:.
717:^
703:.
647:.
637:.
629:.
617:.
613:.
590:.
580:.
568:.
564:.
541:.
533:.
525:.
513:.
490:.
480:.
472:.
462:97
460:.
456:.
433:.
419:.
415:.
390:.
366:.
178:,
174:,
168:.
1033:.
1014:.
995:.
973::
941:.
937::
929::
923:1
906:.
882::
876:9
855:.
831::
804:.
790::
763:.
741::
711:.
699::
680:.
655:.
633::
625::
619:7
598:.
576::
549:.
529::
521::
498:.
476::
468::
441:.
427::
400:.
376:.
112:)
106:(
101:)
97:(
87:·
80:·
73:·
66:·
39:.
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
