化石人类脑演化研究概况

脑演化是人类演化的一个重要组成部分,其研究可以为人类起源、演化、人群关系及语言、智力等方面提供重要的信息。脑演化的主要证据是通过研究颅内模（endocast）及颅骨的形态得到的。颅内模是从颅骨内表面得到的脑的外部形态。有时颅骨的内腔充满泥沙,并且为钙质所结固,可以自然形成颅内模。也可以人工制作颅内模。颅内模和颅骨的内表面能够提供人类脑髓及神经进化方面的直接证据。对化石人类脑演化的研究主要包括以下几个方面的内容：测量或估计脑量的大小及其和身体大小之间的关系;研究脑量随时间的变化过程;通过对早期人类颅内模表面沟回形态特征的研究,探索脑功能区在早期人类和猿类的区别及在演化上的变化;左右大脑不对称性与一侧优势关系,探讨语言的起源和惯用手的脑功能基础等;脑膜中动脉系统、静脉窦系统及与血液循环相关的排泄孔的变化,探讨大脑各部分比例的变化和功能的日益复杂对供血需求的影响;通过对人类脑演化的研究,探讨人类进化的原因。本文通过对以上几个方面及其中国化石脑演化研究的介绍,对化石人类脑演化的研究概况作综合论述和简单回顾。     

The biological and chronological maturation of early hominids

Recent studies on the rate and pattern of dental development indicate that the growth and maturation of early hominids were more similar to the extant apes than to modern humans. This contrasts with the previously held opinion derived from combined dental development, pattern and attrition studies claiming that early hominids were more hominine in their development (Mann, 1975). This paper explores the origin of this difference of opinion and reviews immature hominid dentitions with the benefit of improved radiographs and new data on the pattern and rate of pongid dental development. Paranthropus and Australopithecus specimens are shown to possess an ape-like development pattern but incisor development is specialized in the former and superficially human-like in pattern. The present and recent studies on dental development rate and pattern justify the position that early hominids were more ape-like in their growth and development. Therefore, ages at death calculated from pongid dental development schedules are provided for most immature early hominids. More detailed studies of early hominid developmental biology are now possible. It is suggested that divergent heterochronic processes characterize changes in brain/body proportions during hominid evolution. Relative rates of bone remodeling processes can now be identified on early hominid skeletons. The paleodemographic analysis of early hominids is little changed by the developmental model one chooses.     

Patterns of evolution in Middle Pleistocene hominids

This paper reviews the chronology and morphological variability of Middle Pleistocene H. erectus. specimens. Functional complexes are delineated within the skull and dentition, and their total morphological patterns quantified using univariate and multivariate statistical analysis. Statistical distances are calculated between H. erectus and other hominid samples for each complex, compared to illustrate patterns of mosaic evolution within the skull and dentition of middle Quaternary hominids, and estimated evolution rates are derived. An attempt is made to relate the observed morphological patterns to ecological shifts by early hominid communities, and to assess their significance for hominid taxonomy.     

Hominid paleoecology and competitive exclusion: Limits to similarity,niche differentiation,and the effects of cultural behavior

Fossil evidence from the Plio-Pleistocene of Africa apparently has confirmed a multi-lineage interpretation of early hominid evolution. Empirical refutation of the single species hypothesis must now be matched to the evolutionary ecology theory, which can underwrite taxonomic assessment and help to explain sympatric hominid coexistence. This paper contributes to that goal by reassessing the ecological rationale provided for the single-species hypothesis. Limiting similarity concepts indicate that the allowable ecological overlap between sympatric competitors is greater than the degrees of metric overlap often advanced as standards for identifying fossil species. Optimal foraging theory and the compression hypothesis show that the initial ecological reaction of a hominid to a sympatric competitor would likely be micro-habitat divergence and possibly also temporal differentiation of resource use. The long-term, evolutionary response is niche divergence, probably involving diet as well. General niche partitioning studies suggest that diet and habitat are the most common dimensions of niche separation, although temporal separation is unusually frequent in carnivores. The equation of niche with culture, basic to the single-species hypothesis, has no analytic meaning. Finally, four minor points are discussed, suggesting that (a) extinction is not unlikely, even for a long-lived and competitively competent hominid lineage, (b)parsimony is fickle, (c)interspecific mutualism may jeopardize survival, and (d)generalists are subordinate competitors, but for hominids, seemingly, successful ones. I argue that analog models of hominid paleoecology should be replaced by the use of zoological and anthropological observations to assess the generality and reliability of ecological theory and comcepts that may encompass early hominids.     

Hominid cultural transmission and the evolution of language

This paper presents the hypothesis that linguistic capacity evolved through the action of natural selection as an instrument which increased the efficiency of the cultural transmission system of early hominids. We suggest that during the early stages of hominization, hominid social learning, based on indirect social learning mechanisms and true imitation, came to constitute cumulative cultural transmission based on true imitation and the approval or disapproval of the learned behaviour of offspring. A key factor for this transformation was the development of a conceptual capacity for categorizing learned behaviour in value terms - positive or negative, good or bad. We believe that some hominids developed this capacity for categorizing behaviour, and such an ability allowed them to approve or disapprove of their offsprings- learned behaviour. With such an ability, hominids were favoured, as they could transmit to their offspring all their behavioural experience about what can and cannot be done. This capacity triggered a cultural transmission system similar to the human one, though pre-linguistic. We suggest that the adaptive advantage provided by this new system of social learning generated a selection pressure in favour of the development of a linguistic capacity allowing children to better understand the new kind of evaluative information received from parents.     

Society and territory in human evolution

A survey of recent data on the socio-territorial organization of primates, carnivores and human hunter-gatheres discloses some striking similarities among them. These common features are integrated into a theory of hominid social evolution. It is postulated that the hominids, throughout most of their evolution, were organized into stable groups with the capacity to disperse into largely independent subgroups that remain affiliated with each other. During the course of hominid evolution, territoriality became an increasingly important function of the larger, stable units. The analysis illustrates the value of combining data from primates, carnivores, and human hunter-gatherers in the reconstruction of early hominid behavior.     

Dispersal and colonisation, long and short chronologies: how continuous is the Early Pleistocene record for hominids outside East Africa?

This paper examines the evidence for hominids outside East Africa during the Early Pleistocene. Most attention has focused recently on the evidence for or against a late Pliocene dispersal, ca. 1.8 Ma., of hominids out of Africa into Asia and possibly southern Europe. Here, the focus is widened to include North Africa as well as southern Asia and Europe, as well as the evidence in these regions for hominids after their first putative appearance ca. 1.8 Ma. It suggests that overall there is very little evidence for hominids in most of these regions before the Middle Pleistocene. Consequently, it concludes that the colonising capabilities of Homo erectus may have been seriously over-rated, and that even if hominids did occupy parts of North Africa, southern Europe and southern Asia shortly after 2 Ma, there is little evidence of colonisation. Whilst further fieldwork will doubtless slowly fill many gaps in a poorly documented Lower Pleistocene hominid record, it appears premature to conclude that the appearance of hominids in North Africa, Europe and Asia was automatically followed by permanent settlement. Rather, current data are more consistent with the view that Lower Pleistocene hominid populations outside East Africa were often spatially and temporally discontinuous, that hominid expansion was strongly constrained by latitude, and that occupation of temperate latitudes north of latitude 40 degrees was largely confined to interglacial periods.     

广西柳江土博甘前洞的铀系年代

本文报道晚期智人化石地点广西柳江博甘前洞新生碳酸盐岩和骨化石样的铀系测年结果。该地点表层钙板在约94Ka前开始形成,含化石粘土堆积叠压的钙板年代为约20ka,人牙化石应位于二者之间。二个动物化石样的二种铀系法年代范围为85－139ka,表明该地点与含化石堆积与表层钙板间无地层倒序现象,支持人牙化石大于100ka的结论。邻近的柳江晚期智人化石地点和柳州白莲洞人类遗址铀系测年的结果与本文一致。具现代解剖特征智人在中国南方出现的时间,很可能不晚于西非和非洲。在现代人类起源方面,中国不应是远离中心、滞后和被取代的地区。     

The rise of the hominids as an adaptive shift in fallback foods: plant underground storage organs (USOs) and australopith origins

We propose that a key change in the evolution of hominids from the last common ancestor shared with chimpanzees was the substitution of plant underground storage organs (USOs) for herbaceous vegetation as fallback foods. Four kinds of evidence support this hypothesis: (1) dental and masticatory adaptations of hominids in comparison with the African apes; (2) changes in australopith dentition in the fossil record; (3) paleoecological evidence for the expansion of USO-rich habitats in the late Miocene; and (4) the co-occurrence of hominid fossils with root-eating rodents. We suggest that some of the patterning in the early hominid fossil record, such as the existence of gracile and robust australopiths, may be understood in reference to this adaptive shift in the use of fallback foods. Our hypothesis implicates fallback foods as a critical limiting factor with far-reaching evolutionary effects. This complements the more common focus on adaptations to preferred foods, such as fruit and meat, in hominid evolution.     

Taxonomic affinities and evolutionary history of the Early Pleistocene hominids of Java: dentognathic evidence

Temporal changes, within-group variation, and phylogenetic positions of the Early Pleistocene Javanese hominids remain unclear. Recent debate focused on the age of the oldest Javanese hominids, but the argument so far includes little morphological basis for the fossils. To approach these questions, we analyzed a comprehensive dentognathic sample from Sangiran, which includes most of the existing hominid mandibles and teeth from the Early Pleistocene of Java. The sample was divided into chronologically younger and older groups. We examined morphological differences between these chronological groups, and investigated their affinities with other hominid groups from Africa and Eurasia. The results indicated that 1) there are remarkable morphological differences between the chronologically younger and older groups of Java, 2) the chronologically younger group is morphologically advanced, showing a similar degree of dentognathic reduction to that of Middle Pleistocene Chinese H. erectus, and 3) the chronologically older group exhibits some features that are equally primitive as or more primitive than early H. erectus of Africa. These findings suggest that the evolutionary history of early Javanese H. erectus was more dynamic than previously thought. Coupled with recent discoveries of the earliest form of H. erectus from Dmanisi, Georgia, the primitive aspects of the oldest Javanese hominid remains suggest that hominid groups prior to the grade of ca. 1.8-1.5 Ma African early H. erectus dispersed into eastern Eurasia during the earlier Early Pleistocene, although the age of the Javanese hominids themselves is yet to be resolved. Subsequent periods of the Early Pleistocene witnessed remarkable changes in the Javanese hominid record, which are ascribed either to significant in situ evolution or replacement of populations.     

Climate change and evolving human diversity in Europe during the last glacial

A link between climate change and human evolution during the Pleistocene has often been assumed but rarely tested. At the macro-evolutionary level Foley showed for hominids that extinction, rather than speciation, correlates with environmental change as recorded in the deep sea record. Our aim is to examine this finding at a smaller scale and with high-resolution environmental and archaeological archives. Our interest is in changing patterns of human dispersal under shifting Pleistocene climates during the last glacial period in Europe. Selecting this time frame and region allows us to observe how two hominid taxa, Neanderthals and Crô-Magnons, adapted to climatic conditions during oxygen isotope stage 3. These taxa are representative of two hominid adaptive radiations, termed terrestrial and aquatic, which exhibited different habitat preferences but similar tolerances to climatic factors. Their response to changing ecological conditions was predicated upon their ability to extend their societies in space and time. We examine this difference further using a database of all available radiocarbon determinations from western Europe in the late glacial. These data act as proxies for population history, and in particular the expansion and contraction of regional populations as climate changed rapidly. Independent assessment of these processes is obtained from the genetic history of Europeans. The results indicate that climate affects population contraction rather than expansion. We discuss the consequences for genetic and cultural diversity which led to the legacy of the Ice Age: a single hominid species, globally distributed.     

From apes to humans: locomotion as a key feature for phylogeny

If bipedalism has often been considered to be of a crucial interest for understanding hominid evolution, the acceptance of locomotor features to build phylogenies is still far from being a reality in the field. Especially for hominid evolution, it still seems to be difficult to accept that traits, other than craniodental ones, can be useful for defining the major dichotomies. The recent discovery of Australopithecus anamensis suggests a challenging view of the major dichotomy between apes and humans. Whilst it is widely accepted that Ardipithecus ramidus is ancestral to Australopithecus anamensis, which in its turn is ancestral to Australopithecus afarensis and then to later hominids, the postcranial adaptations, which should be taken into account, suggest another branching pattern. Based on the fact that by 4.0 million years two different locomotor patterns can be identified in hominids, two lineages would appear to be present: the "Australopithecine" lineage (with Australopithecus afarensis or Ardipithecus ramidus if the latter is really a hominid sensu stricto) and the "Hominine" lineage (with Australopithecus anamensis = Praeanthropus africanus).     

Observed and expected variation in hominid evolution

All of the major groups of fossil hominids (australopithecines, pithecanthropines, Neandertals, and early sapiens) were discovered by 1925, and therefore prior to the formulation of the synthetic theory of evolution that revolutionized the concept of the species in systematics. While these fossil finds were being made the framework for their interpretation included several assumptions: (1) that the number of living hominoid species was great, and that intraspecific variation was slight (authoritative sources recognized as many as 14 separate species of chimpanzees and 15 species of gorillas); (2) that the timescale of human evolution was brief (measured in tens or hundreds of thousands of years). As a result of these premises the consensus that hominid evolution was characterized by a large number of sympatric and synchronic species was virtually inevitable.In contrast, recent molecular studies demonstrate that genetic diversity among recent hominoids is so slight that even humans and chimpanzees differ at only about 1% of the loci that have been sampled so far; evidently, very small genetic differences can produce rather great contrasts in morphology. At the same time, geological break-throughs have increased the timescale for human evolution to several million years.It is concluded that morphological differences among fossil hominids, even if very appreciable and complex, do not necessarily reflect a great degree of either genetic or taxonomic diversity. Potential effects of evolutionary change through time should be incorporated into models of hominid evolution as a means of assessing the minimum number of lineages required to account for observed variations among hominid specimens.     

Natural selection in brain evolution of early hominids

Directional effect of natural selection on the arrangement of brain of anthropoids and man is reviewed. It is demonstrated that the evolution of the human nervous system is an integrated result of several multidirectional processes. At the early stages of the evolution of primates, the general biological principles of survival of the fittest, i.e., natural selection of the most adapted variants of the brain structure prevailed. During the period of hominid specialization, natural selection led to the formation of the neocortical control of voluntary movements, memory, and mental associations. At later stages of human morphological evolution, biological mechanisms of natural selection of the brain arrangement were replaced by social mechanisms. This process initiated hominid migrations and the growth of the brain size and individual variability in human ancestors. A model of cerebral sorting is proposed to explain the mechanisms of multidirectional selection leading to an increase in brain size of early hominids.     

Estimating hominid life history: the critical interbirth interval

Unlike any great apes, humans have expanded into a wide variety of habitats during the course of evolution, beginning with the transition by australopithecines from forest to savanna habitation. Novel environments are likely to have imposed hominids a demographic challenge due to such factors as higher predation risk and scarcer food resources. In fact, recent studies have found a paucity of older relative to younger adults in hominid fossil remains, indicating considerably high adult mortality in australopithecines, early Homo, and Neanderthals. It is not clear to date why only human ancestors among all hominoid species could survive in these harsh environments. In this paper, we explore the possibility that hominids had shorter interbirth intervals to enhance fertility than the extant apes. To infer interbirth intervals in fossil hominids, we introduce the notion of the critical interbirth interval, or the threshold length of birth spacing above which a population is expected to go to extinction. We develop a new method to obtain the critical interbirth intervals of hominids based on the observed ratios of older adults to all adults in fossil samples. Our analysis suggests that the critical interbirth intervals of australopithecines, early Homo, and Neanderthals are significantly shorter than the observed interbirth intervals of extant great apes. We also discuss possible factors that may have caused the evolutionary divergence of hominid life history traits from those of great apes.     

An Inquiry Safari: What Can We Learn From Skulls?

In this article, we describe an 8- to 10-day inquiry safari designed for middle/high school students to investigate hominid evolution using replica skulls of extant and extinct vertebrates. Students begin the unit using their own skulls and proceed to use the replica skulls of extant vertebrates to construct an understanding of how skulls can be used to interpret and infer diets, dentition, dental formulae, bipedal or quadrupedal locomotion, and the social structure of animals. They are then able to use this knowledge to construct similar inferences for extinct fossil hominids. Using radiometric dating data, the students develop possible phylogenetic pathways for hominid evolution. The lessons promote the use of inquiry skills including journaling, observing, drawing, puzzle-making, using taxonomic keys, and investigating into deep geological time.     

Early paleolithic of China and eastern Asia

In attempting to understand the course of human evolution and the nature of hominid adaptation over the past few million years, it is necessary to consider prevailing evidence from all parts of the world. Eastern Asia provides a range of important questions and challenges with regard to this evolutionary puzzle. Although evidence for earlier ape evolution is present in China (for example, at Lufeng in Yunnan Province), the earliest evidence for hominid presence appears to be in the Early Pleistocene, apparently the result of a migration of hominids to and subsequent adaptation within Eastern Asia. The archeological record provides a closer look at some technological aspects of this adaptation during the Early and Middle Pleistocene, showing both distinctive contrasts and intriguing continuities relative to the rest of the Old World.     

Foramen magnum position variation in Pan troglodytes, Plio-Pleistocene hominids, and recent Homo sapiens: implications for recognizing the earliest hominids

The anteroposterior position of the foramen magnum distinguishes living Homo sapiens from apes, and has been used as evidence for the hominid status of numerable fossils in the history of human paleontology. During the past decade, foramen magnum position has been cited as evidence of the hominid status of Ardipithecus and Sahelanthropus. Specifically, the basion of Ardpithecus is reported to be inline with the bicarotid chord, while the basion of Sahelanthropus is reported to both touch the biporion chord and intersect the bicarotid chord. In order to assess the effectiveness of anteroposterior foramen magnum position in distinguishing hominids from nonhominid apes, this study examined whether or not the positions of biporion and bicarotid relative to basion sufficiently distinguished Pan troglodytes from recent Homo sapiens and Plio-Pleistocene hominids. The distances from basion to the biporion chord (BSBIP) and from basion to the bicarotid chord (BSBIC) were measured on samples of chimpanzee (n = 69) and recent human (n = 42) crania and a sample of Plio-Pleistocene hominid fossils (n = 8). The data were used to test the hypothesis that BSBIP and BSBIC measurements do not sufficiently distinguish P. troglodytes from hominids. While basion to biporion (BSBIP) does not effectively distinguish P. troglodytes from Plio-Pleistocene hominids and humans when used univariately, basion to bicarotid (BSBIC), when used univariately or bivariately with BSBIP, can be used to test whether or not an unknown specimen is a hominid. These results are used to evaluate the hominid status of Ardipithecus and Sahelanthropus.     

The evolution and functions of laughter and humor: a synthetic approach

A number of recent hypotheses have attempted to explain the ultimate evolutionary origins of laughter and humor. However most of these have lacked breadth in their evolutionary frameworks while neglecting the empirical existence of two distinct types of laughter--Duchenne and non-Duchenne--and the implications of this distinction for the evolution of laughter as a signal. Most of these hypotheses have also been proposed in relative isolation of each other and remain disjointed from the relevant empirical literature. Here we attempt to remedy these shortcomings through a synthesis of previous laughter and humor research followed by (i) a reevaluation of this research in light of theory and data from several relevant disciplines, and (ii) the proposal of a synthetic evolutionary framework that takes into account phylogeny and history as well as proximate mechanisms and adaptive significance. We consider laughter to have been a preadaptation that was gradually elaborated and co-opted through both biological and cultural evolution. We hypothesize that Duchenne laughter became fully ritualized in early hominids between 4 and 2 mya as a medium for playful emotional contagion. This mechanism would have coupled the emotions of small hominid groups and promoted resource-building social play during the fleeting periods of safety and satiation that characterized early bipedal life. We further postulate that a generalized class of nonserious social incongruity would have been a reliable indicator of such safe times and thereby came to be a potent distal elicitor of laughter and playful emotion. This class of stimuli had its origins in primate social play and was the foundation for formal human humor. Within this framework, Duchenne laughter and protohumor were well established in the hominid biobehavioral repertoire when more cognitively sophisticated traits evolved in the hominid line between 2 mya and the present. The prior existence of laughter and humor allowed them to be co-opted for numerous novel functions, and it is from this process that non-Duchenne laughter and the "dark side" of laughter emerged. This perspective organizes the diversified forms and functions that characterize laughter and humor today and clarifies when and how laughter and humor evolved during the course of human evolution.