Evolution Of Bipedalism Research Articles

Positional behavior and substrate use in wild adult bearded capuchin monkeys (Sapajus libidinosus).

Natural selection for positional behavior (posture and locomotion) has at least partially driven the evolution of anatomical form and function in the order Primates. Examination of bipedal behaviors associated with daily activity patterns, foraging, and terrestrial habitat use in nonhuman primates, particularly those that adopt bipedal postures and use bipedal locomotion, allows us to refine hypotheses concerning the evolution of bipedalism in humans. This study describes the positional behavior of wild bearded capuchins (Sapajus libidinosus), a species that is known for its use of terrestrial substrates and its habitual use of stones as tools. Here, we test the association of terrestrial substrate use with bipedal posture and locomotion, and the influence of sex (which co-varies with body mass in adults of this species) on positional behavior and substrate use. Behavior and location of 16 wild adult bearded capuchins from two groups were sampled systematically at 15 s intervals for 2 min periods for 1 year (10,244 samples). Despite their different body masses, adult males (average 3.5 kg) and females (average 2.1 kg) in this study did not differ substantially in their positional behaviors, postures, or use of substrates for particular activities. The monkeys used terrestrial substrates in 27% of samples. Bipedal postures and behaviors, while not a prominent feature of their behavior, occurred in different forms on the two substrates. The monkeys crouched bipedally in trees, but did not use other bipedal postures in trees. While on terrestrial substrates, they also crouched bipedally but occasionally stood upright and moved bipedally with orthograde posture. Bearded capuchin monkeys' behavior supports the suggestion from anatomical analysis that S. libidinosus is morphologically better adapted than its congeners to adopt orthograde postures.

A quantification of calcaneal lateral plantar process position with implications for bipedal locomotion in Australopithecus

The evolution of bipedalism in the hominin lineage has shaped the posterior human calcaneus into a large, robust structure considered to be adaptive for dissipating peak compressive forces and energy during heel-strike. A unique anatomy thought to contribute to the human calcaneus and its function is the lateral plantar process (LPP). While it has long been known that humans possess a plantarly positioned LPP and apes possess a more dorsally positioned homologous structure, the relative position of the LPP and intraspecific variation of this structure have never been quantified. Here, we present a method for quantifying relative LPP position and find that, while variable, humans have a significantly more plantar position of the LPP than that found in the apes. Among extinct hominins, while the position of the LPP in Australopithecus afarensis falls within the human distribution, the LPP is more dorsally positioned in Australopithecus sediba and barely within the modern human range of variation. Results from a resampling procedure suggest that these differences can reflect either individual variation of a foot structure/function largely shared among Australopithecus species, or functionally distinct morphologies that reflect locomotor diversity in Plio-Pleistocene hominins. An implication of the latter possibility is that calcaneal changes adaptive for heel-striking bipedalism may have evolved independently in two different hominin lineages.

There Is No "Obstetrical Dilemma": Towards a Braver Medicine with Fewer Childbirth Interventions.

Humans give birth to big-brained babies through a bony birth canal that metamorphosed during the evolution of bipedalism; they have a tighter fit at birth between baby and bony birth canal than do our closest relatives the chimpanzees; and they are incapable of grasping onto caregivers as early as infant chimpanzees develop the skill. Since the mid-20th century, these observations and more have been linked together into the "obstetrical dilemma" (OD): human babies are helpless because they are born early to escape before they outgrow the mother's pelvis, the expansion of which is prevented by natural selection for bipedalism. The OD continues to be a popular idea, often expressed as incontrovertible fact, but it no longer deserves its popularity. There are better explanations for gestation length, childbirth difficulty, and the developmental biology of newborns than mainly or only because of natural selection's constraints on women's hips. And humans are not born early either, as is widely assumed. This all-too-powerful human evolutionary narrative deserves our skeptical consideration. Bias from OD thinking is likely amplifying the perceived risk of cephalopelvic and fetopelvic disproportion during labor-contributing, even if slightly, to medicine's underestimation of women's bodies and over-implementation of childbirth interventions.

Paleoanthropology's uses of the bipedal criterion.

Bipedalism is one of the criteria that paleoanthropologists use in order to interpret the fossil record and to determine if a specimen belongs to the human lineage. In the context of such interpretations, bipedalism is considered to be a unique characteristic of this lineage that also marks its origin. This conception has largely remained unchallenged over the last decades, in spite of fossil discoveries that led to the emergence of bipedalism in the human lineage being shifted back by several millions of years. In this paper, I analyze the uses of this criterion in paleoanthropology and demonstrate that interpretative biases (such as underdetermined inferences and circular reasoning) are at play in interpretations of hominin remains. By discussing Darwin's hypotheses about the evolution of bipedalism, I identify major theoretical issues that need to be addressed in the current debates on hominin evolution. First, the assumption that "man alone has become a biped" (Darwin in The descent of man, and selection in relation to sex, John Murray, London, 1871) is analyzed in the light of recent empirical data. Three major issues are discussed: the definition of "man", i.e. "human", the uniqueness of human bipedalism, and the equivocal meaning of being a "biped". Then, I highlight some of Darwin's remarks that may be helpful for current debates in paleoanthropology, regarding natural selection in locomotor evolution, as well as taxonomic and phylogenetic significance of functional features. Finally, I analyze two examples of how fossil discoverers referred to Darwin in the recent years and discuss his role as an intellectual support.

Sexual anatomy differs between orthograde and pronograde primates and other mammals: Implications for human evolution

Detailed descriptions of the perineal anatomy of most primates are lacking so that studies of modern human perineal anatomy in the context of general primate evolutionary morphology still require a broader framework. Our dissections of human, non‐human primate, and non‐primate mammal perinea (human (n=43), dog (n=16), cat (n=3), goat (n=9), pig (n=9), horse (n=3), cotton‐top tamarin (n=2), ring‐tailed lemur (n=1), potto (n=1), and Verreaux's sifaka (n=1)) suggest that modern human perineal anatomy likely follows a g eneral primate pattern. Primates with a habitually orthograde locomotor position copulate ventro‐ventrally and initial observations suggest that they have different perineal anatomy than primates and non‐primate mammals that employ habitually pronograde locomotor position and copulate dorso‐ventrally. Specifically, females of orthograde primate species have a relatively shallow vulva with easy access to a ventrally facing, prominent clitoris – easily stimulated during sexual courtship. Conversely, females of pronograde primate and non‐primate mammal species possess deep vulvae with subcutaneous constrictor muscles. These restrict access to a dorsally facing, recessed clitoris and vaginal orifice during courtship and the clitoris is only available during copulation. We suggest that adaptation of the pelvis for orthograde locomotor posture during the evolution of bipedalism followed a general pattern of primate perineal anatomy. This pattern of female perineal morphology enables easier access to the clitoris so that pre‐copulatory stimulation is possible. This female perineal anatomy possibly set the stage for loss of estrus in women that facilitates the modern male‐female pair bond social structure.Support or Funding InformationSupport received from Midwestern University intramural funds

Temporal and phylogenetic evolution of the sauropod dinosaur body plan.

The colossal size and body plan of sauropod dinosaurs are unparalleled in terrestrial vertebrates. However, to date, there have been only limited attempts to examine temporal and phylogenetic patterns in the sauropod bauplan. Here, we combine three-dimensional computational models with phylogenetic reconstructions to quantify the evolution of whole-body shape and body segment properties across the sauropod radiation. Limitations associated with the absence of soft tissue preservation in fossils result in large error bars about mean absolute body shape predictions. However, applying any consistent skeleton : body volume ratio to all taxa does yield changes in body shape that appear concurrent with major macroevolutionary events in sauropod history. A caudad shift in centre-of-mass (CoM) in Middle Triassic Saurischia, associated with the evolution of bipedalism in various dinosaur lineages, was reversed in Late Triassic sauropodomorphs. A craniad CoM shift coincided with the evolution of quadrupedalism in the Late Triassic, followed by a more striking craniad shift in Late Jurassic–Cretaceous titanosauriforms, which included the largest sauropods. These craniad CoM shifts are strongly correlated with neck enlargement, a key innovation in sauropod evolution and pivotal to their gigantism. By creating a much larger feeding envelope, neck elongation is thought to have increased feeding efficiency and opened up trophic niches that were inaccessible to other herbivores. However, we find that relative neck size and CoM position are not strongly correlated with inferred feeding habits. Instead the craniad CoM positions of titanosauriforms appear closely linked with locomotion and environmental distributions, potentially contributing to the continued success of this group until the end-Cretaceous, with all other sauropods having gone extinct by the early Late Cretaceous.

Relationship between human evolution and neurally mediated syncope disclosed by the polymorphic sites of the adrenergic receptor gene α2B-AR.

The objective of this study was to clarify the effects of disease on neurally mediated syncope (NMS) during an acute stress reaction. We analyzed the mechanism of the molecular interaction and the polymorphisms of the alpha-2 adrenoreceptor (α2B-AR) gene as the potential psychiatric cause of incentive stress. We focused on the following three genotypes of the repeat polymorphism site at Glu 301–303 in the α2B-AR gene: Glu12/12, Glu12/9, and Glu9/9. On the basis of our clinical research, NMS is likely to occur in people with the Glu12/9 heterotype. To verify this, we assessed this relationship with the interaction of Gi protein and adenylate cyclase by in silico analysis of the Glu12/9 heterotype. By measuring the difference in the dissociation time of the Gi-α subunit twice, we found that the Glu12/9 heterotype suppressed the action of adenylate cyclase longer than the Glu homotypes. As this difference in the Glu repeat number effect is thought to be one of the causes of NMS, we investigated the evolutionary significance of the Glu repeat number. Glu8 was originally repeated in simians, while the Glu12 repeats occurred over time during the evolution of bipedalism in humans. Taken with the Glu12 numbers, NMS would likely become a defensive measure to prevent significant blood flow to the human brain.

New Perspectives on the Evolution of Bipedalism

New Perspectives on the Evolution of Bipedalism

Body mass estimation in non‐avian bipeds using a theoretical conversion to quadruped stylopodial proportions

Summary Body mass is strongly related to both physiological and ecological properties of living organisms. As a result, generating robust, broadly applicable models for estimating body mass in the fossil record provides the opportunity to reconstruct palaeobiology and investigate evolutionary ecology on a large temporal scale. A recent study provided strong evidence that the minimum circumference of stylopodial elements (humerus and femur) is conservatively associated with body mass in living quadrupeds. Unfortunately, this model is not directly applicable to extinct bipeds, such as non‐avian dinosaurs. This study presents a new equation that mathematically corrects the quadruped equation for use in bipeds. It is derived from the systemic difference in the circumference‐to‐area scaling relationship of two circles (hypothetical quadruped) and one circle (hypothetical biped), which represent the cross‐section of the main weight‐bearing limb bones. When applied to a newly constructed data set of femoral circumferences and body masses in living birds, the new equation reveals errors that are significantly lower than other published equations, but significantly higher than the error inherent in the avian data set. Such errors, however, are expected given the unique overall femoral circumference–body mass scaling relationship found in birds. Body mass estimates for a sample of bipedal dinosaurs using the new model are consistent with recent estimates based on volumetric life reconstructions, but, in contrast, this equation is simpler to use, with the concomitant potential to provide a wider set of body mass estimates for extinct bipeds. Although it is evident that no one estimation model is flawless, the combined use of the corrected quadrupedal equations and the previously published quadrupedal equation offer a consistent approach with which to estimate body masses in both quadrupeds and bipeds. These models have implications for conducting large‐scale macroevolutionary analyses of body size throughout the evolutionary history of terrestrial vertebrates, and, in particular, across major changes in body plan, such as the evolution of bipedality in archosaurs and quadrupedality in dinosaurs.

Three‐dimensional shape variation of talar surface morphology in hominoid primates

The hominoid foot is of particular interest to biological anthropologists, as changes in its anatomy through time reflect the adoption of terrestrial locomotion, particularly in species of Australopithecus and Homo. Understanding the osteological morphology associated with changes in whole foot function and the development of the plantar medial longitudinal foot arch are key to understanding the transition through habitual bipedalism in australopithecines to obligate bipedalism and long-distance running in Homo. The talus is ideal for studying relationships between morphology and function in this context, as it is a major contributor to the adduction-abduction, plantar-dorsal flexion and inversion-eversion of the foot, and transmits all forces encountered from the foot to the leg. The talar surface is predominantly covered by articular facets, which have different quantifiable morphological characters, including surface area, surface curvature and orientation. The talus also presents challenges to the investigator, as its globular shape is very difficult to quantify accurately and reproducibly. Here we apply a three-dimensional approach using type 3 landmarks (slid semilandmarks) that are geometrically homologous to determine overall talar shape variations in a range of living and fossil hominoid taxa. Additionally, we use novel approaches to quantify the relative orientations and curvatures of talar articular facets by determining the principal vectors of facet orientation and fitting spheres to articular facets. The resulting metrics are analysed using phylogenetic regressions and principal components analyses. Our results suggest that articular surface curvatures reflect locomotor specialisations with, in particular, orangutans having more highly curved facets in all but the calcaneal facet. Similarly, our approach to quantifying articular facet orientation appears to be effective in discriminating between extant hominoid species, and may therefore provide a sound basis for the study of fossil taxa and evolution of bipedalism in Australopithecus and Homo.

Delegation to automaticity: the driving force for cognitive evolution?

The ability to delegate control over repetitive tasks from higher to lower neural centers may be a fundamental innovation in human cognition. Plausibly, the massive neurocomputational challenges associated with the mastery of balance during the evolution of bipedality in proto-humans provided a strong selective advantage to individuals with brains capable of efficiently transferring tasks in this way. Thus, the shift from quadrupedal to bipedal locomotion may have driven the rapid evolution of distinctive features of human neuronal functioning. We review recent studies of functional neuroanatomy that bear upon this hypothesis, and identify ways to test our ideas.

Hand before foot? Cortical somatotopy suggests manual dexterity is primitive and evolved independently of bipedalism

People have long speculated whether the evolution of bipedalism in early hominins triggered tool use (by freeing their hands) or whether the necessity of making and using tools encouraged the shift to upright gait. Either way, it is commonly thought that one led to the other. In this study, we sought to shed new light on the origins of manual dexterity and bipedalism by mapping the neural representations in the brain of the fingers and toes of living people and monkeys. Contrary to the ‘hand-in-glove’ notion outlined above, our results suggest that adaptations underlying tool use evolved independently of those required for human bipedality. In both humans and monkeys, we found that each finger was represented separately in the primary sensorimotor cortex just as they are physically separated in the hand. This reflects the ability to use each digit independently, as required for the complex manipulation involved in tool use. The neural mapping of the subjects’ toes differed, however. In the monkeys, the somatotopic representation of the toes was fused, showing that the digits function predominantly as a unit in general grasping. Humans, by contrast, had an independent neurological representation of the big toe (hallux), suggesting association with bipedal locomotion. These observations suggest that the brain circuits for the hand had advanced beyond simple grasping, whereas our primate ancestors were still general arboreal quadrupeds. This early adaptation laid the foundation for the evolution of manual dexterity, which was preserved and enhanced in hominins. In hominins, a separate adaptation, involving the neural separation of the big toe, apparently occurred with bipedality. This accords with the known fossil evidence, including the recently reported hominin fossils which have been dated to 4.4 million years ago.

Cranial remains of Poposaurus gracilis (Pseudosuchia: Poposauroidea) from the Upper Triassic, the distribution of the taxon, and its implications for poposauroid evolution

Abstract The partial postcrania of Poposaurus gracilis , a bipedal poposauroid convergent with theropod dinosaurs, has been known for nearly a century, but the skull of P. gracilis has proven elusive. P. gracilis is part of a clade of morphologically divergent pseudosuchians (poposauroids) whose members are sometimes bipedal, lack dentition (i.e. beaks) and some have elongated neural spines (i.e. sails). However, the timing and acquisition of these character states is unknown given the uncertainty of the skull morphology of the ‘mid-grade’ poposauroid P. gracilis. Here, we present the first confirmed skull remains of P. gracilis directly associated with diagnostic pelvic elements that overlap with the holotype. The incomplete skeleton (PEFO 34865) from the Chinle Formation of Petrified Forest National Park (Arizona, USA) includes a left maxilla with a large, mediolaterally compressed tooth, left dentary, right prearticular and a partial postcranium. The character states of P. gracilis (bipedal, ‘sail-less’ and toothed) demonstrate that the evolution of bipedalism, the origin/loss of a dorsal ‘sail’ and the shift to an edentulous beak are complex in poposauroids. P. gracilis is widespread in the Upper Triassic formations in the western USA and is restricted temporally prior to the Adamanian–Revueltian faunal turnover during the Norian.

Pelvic and hindlimb myology of the basal archosaur Poposaurus gracilis (archosauria: Poposauroidea)

The discovery of a largely complete and well preserved specimen of Poposaurus gracilis has provided the opportunity to generate the first phylogenetically based reconstruction of pelvic and hindlimb musculature of an extinct nondinosaurian archosaur. As in dinosaurs, multiple lineages of basal archosaurs convergently evolved parasagittally erect limbs. However, in contrast to the laterally projecting acetabulum, or "buttress erect" hip morphology of ornithodirans, basal archosaurs evolved a very different, ventrally projecting acetabulum, or "pillar erect" hip. Reconstruction of the pelvic and hindlimb musculotendinous system in a bipedal suchian archosaur clarifies how the anatomical transformations associated with the evolution of bipedalism in basal archosaurs differed from that of bipedal dinosaurs and birds. This reconstruction is based on the direct examination of the osteology and myology of phylogenetically relevant extant taxa in conjunction with osteological correlates from the skeleton of P. gracilis. This data set includes a series of inferences (presence/absence of a structure, number of components, and origin/insertion sites) regarding 26 individual muscles or muscle groups, three pelvic ligaments, and two connective tissue structures in the pelvis, hindlimb, and pes of P. gracilis. These data provide a foundation for subsequent examination of variation in myological orientation and function based on pelvic and hindlimb morphology, across the basal archosaur lineage leading to extant crocodilians.

Comparative triceps surae morphology in primates: a review.

Primate locomotor evolution, particularly the evolution of bipedalism, is often examined through morphological studies. Many of these studies have examined the uniqueness of the primate forelimb, and others have examined the primate hip and thigh. Few data exist, however, regarding the myology and function of the leg muscles, even though the ankle plantar flexors are highly important during human bipedalism. In this paper, we draw together data on the fiber type and muscle mass variation in the ankle plantar flexors of primates and make comparisons to other mammals. The data suggest that great apes, atelines, and lorisines exhibit similarity in the mass distribution of the triceps surae. We conclude that variation in triceps surae may be related to the shared locomotor mode exhibited by these groups and that triceps surae morphology, which approaches that of humans, may be related to frequent use of semiplantigrade locomotion and vertical climbing.

Morphological integration and the evolution of knuckle-walking

The evolution of knuckle-walking has profound implications for our understanding of the emergence of bipedalism. The modern debate surrounding its evolution is concerned with whether or not it is homologous in chimpanzees and gorillas. Here, this problem is approached using the methods of morphological integration to test hypotheses of patterns and magnitudes of integration in the third manual ray and capitate. If knuckle-walking morphologies are highly integrated and evolve in a correlated bundle (i.e., comprising a functional complex), it seems reasonable that they could have been recruited independently relatively easily in gorillas and chimpanzees, thus increasing the likelihood of homoplasy. If, however, there is no evidence for a knuckle-walking complex, then it seems less likely that chimpanzees and gorillas would have evolved knuckle-walking independently. Results indicate that chimpanzees and gorillas are not characterized by high magnitudes of integration or unique patterns of integration that distinguish them from non-knuckle-walking taxa. This does not support the hypothesis of a knuckle-walking complex, nor does it support the contention that knuckle-walking could have been easily evolved independently in chimpanzees and gorillas. Implications for trait analysis and the evolution of bipedalism are discussed, as are recent analyses supporting the independent origins of knuckle-walking.

The lives of the brain

Humans are generally curious about their origins, and most cultures have stories relating to this concern. Now we know that our species originated in Africa, approximately 200,000 years ago. Six million years earlier, an ape-like ancestor, who walked on two legs and had new eating habits and reduced dentition, led to subsequent hominin species that gradually acquired a very large brain of unprecedented functional capacity. In The Lives of the Brain, John S. Allen — a neuroanthropologist at the Dana and David Dornsife Cognitive Neuroscience Imaging Center and Brain and Creativity Institute at the University of Southern California — captures this history and expands on what we know about the human brain. The scope and scholarship of this book is impressive. Allen discusses relevant contributions from the past, such as those of 100 years ago by W.E. LeGros Clark and G. Elliott Smith on comparative aspects of brain anatomy, covers the major contributions of researchers over the last 60 years, and includes the most recent results and conclusions of current investigators. There is much to learn, even by the experienced investigator, from reading this book, which is also a treat for any science-loving reader. Each chapter deals with a different topic and a surprising number of subtopics. For example, an early chapter introduces us to the brain and its basic neuroanatomy, but it also covers early, and now discarded, theories of brain evolution, the recognition that neocortex is not really new, and the debate over the value of the lunate sulcus as a marker of the border of the primary visual cortex in the endocasts of the brains of now extinct hominins. In this chapter, Allen is more optimistic than some of us would be in concluding that the classic Brodmann areas of the brain “can be readily identified as homologous units across primate species.” Progress in understanding brain evolution would have been much greater if this were the case, and many present day controversies would have been resolved. In the same chapter, Allen discusses the growth of comparative neuroscience over the past several decades, methods of determining the functional divisions of the brain, including informative studies of patients with lesions, and Wilder Penfield’s electrical stimulation studies of human cortex. He concludes with the reflection that most neuroscientists today do not study brains directly but representations of brains on computer screens. Such has been the impact of functional brain-imaging techniques. Other chapters are equally broad and varied. A chapter on brain size contains a nice history of early investigations, a review of the cranial capacity of fossil hominins, the problem of giving birth to a baby with a large brain, and a discussion of the relative roles of brain size increases compared with changes in brain organization in human brain evolution. Subsequent chapters on brain evolution range from discussion of the modified position of the foramen magnum with the evolution of bipedality, the degeneration of the human olfactory system, the recently published views on the functions of spindle cells of the human brain, the lunate sulcus debate, and frontal lobe evolution. Allen also draws examples of brain evolution from nonhuman species. Thus, he draws attention to the larger hippocampus for the memory of hidden food in food-storing birds. A chapter on the plastic brain considers the relative roles of genetic and environmental factors in brain development and brain plasticity as a result of learning and experience. Molecular and genetic aspects of brain development and evolution are covered further in a subsequent chapter. Elsewhere, we are told about the energy costs of the brain and how changes in diet were necessary in order to allow the evolution of big human brains. We are reminded that humans live a very long time compared with other primates and given reasons why a long life may have been selected for in human evolution. We are reassured by the evidence that human brains can be expected to function fairly well for at least 70 years, although some regressive changes are expected. A thoughtful chapter on the evolution of language covers the last 20 years of intensive research and debate, as well as earlier contributions. For those outside the field of linguistics, this chapter provides a good introductory background. At the book’s conclusion, Allen clearly feels that there has been great recent progress in understanding the evolution and organization of the human brain, but there is so much more to learn. He concludes that the large human brain reflects the evolution of many different cognitive adaptations. Perhaps most important is the ability to look into the future and reflect on the past, a cognitive ability that is said to be unique to humans. We can do both while reading this book.

15-P006 Hindlimb specialization and the evolution of bipedalism in the northern three-toed jerboa, Dipus sagitta

15-P006 Hindlimb specialization and the evolution of bipedalism in the northern three-toed jerboa, Dipus sagitta

The foot of Homo floresiensis

Homo floresiensis is an endemic hominin species that occupied Liang Bua, a limestone cave on Flores in eastern Indonesia, during the Late Pleistocene epoch. The skeleton of the type specimen (LB1) of H. floresiensis includes a relatively complete left foot and parts of the right foot. These feet provide insights into the evolution of bipedalism and, together with the rest of the skeleton, have implications for hominin dispersal events into Asia. Here we show that LB1's foot is exceptionally long relative to the femur and tibia, proportions never before documented in hominins but seen in some African apes. Although the metatarsal robusticity sequence is human-like and the hallux is fully adducted, other intrinsic proportions and pedal features are more ape-like. The postcranial anatomy of H. floresiensis is that of a biped, but the unique lower-limb proportions and surprising combination of derived and primitive pedal morphologies suggest kinematic and biomechanical differences from modern human gait. Therefore, LB1 offers the most complete glimpse of a bipedal hominin foot that lacks the full suite of derived features characteristic of modern humans and whose mosaic design may be primitive for the genus Homo. These new findings raise the possibility that the ancestor of H. floresiensis was not Homo erectus but instead some other, more primitive, hominin whose dispersal into southeast Asia is still undocumented.

Human balance, the evolution of bipedalism and dysequilibrium syndrome

A new model of the uniqueness, nature and evolution of human bipedality is presented in the context of the etiology of the balance disorder of dysequilibrium syndrome. Human bipedality is biologically novel in several remarkable respects. Humans are (a) obligate, habitual and diverse in their bipedalism, (b) hold their body carriage spinally erect in a multisegmental "antigravity pole", (c) use their forelimbs exclusively for nonlocomotion, (d) support their body weight exclusively by vertical balance and normally never use prehensile holds. Further, human bipedalism is combined with (e) upper body actions that quickly shift the body's center of mass (e.g. tennis serves, piggy-back carrying of children), (f) use transient unstable erect positions (dance, kicking and fighting), (g) body height that makes falls injurious, (h) stiff gait walking, and (i) endurance running. Underlying these novelties, I conjecture, is a species specific human vertical balance faculty. This faculty synchronizes any action with a skeletomuscular adjustment that corrects its potential destabilizing impact upon the projection of the body's center of mass over its foot support. The balance faculty depends upon internal models of the erect vertical body's geometrical relationship (and its deviations) to its support base. Due to the situation that humans are obligate erect terrestrial animals, two frameworks - the body- and gravity-defined frameworks - are in constant alignment in the vertical z-axis. This alignment allows human balance to adapt egocentric body cognitions to detect body deviations from the gravitational vertical. This link between human balance and the processing of geometrical orientation, I propose, accounts for the close link between balance and spatial cognition found in the cerebral cortex. I argue that cortical areas processing the spatial and other cognitions needed to enable vertical balance was an important reason for brain size expansion of Homo erectus. A novel source of evidence for this conjecture is the rare autosomal recessive condition of dysequilibrium syndrome. In dysequilibrium syndrome, individuals fail to learn to walk bipedally (with this not being due to sensory, vestibular nor motor coordination defects). Dysequilibrium syndrome is associated with severe spatial deficits that I conjecture underlie its balance dysfunction. The associated brain defects and gene mutations of dysequilibrium syndrome provide new opportunities to investigate (i) the neurological processes responsible for the human specific balance faculty, and (ii) through gene dating techniques, its evolution.