中国古人类颅内模及脑演化研究进展

颅内模保存有脑表面的形态结构,是脑演化研究的直接证据。中国最早复原和研究的颅内模来自20世纪20年代北京周口店遗址发现的3号猿人头骨;此后虽然中国境内也相继发现了一些古人类的头骨化石,但由于古人类标本非常珍贵,不允许对其进行实体解剖,加上多数头骨破碎或者内部附有地层胶结物,导致颅内模无法成功复原。受技术水平及研究手段的限制,研究者一般只是侧重于化石外表形态结构的研究。高分辨率工业CT和3D软件的应用,可以在不损坏标本的情况下,虚拟复原出化石的内部解剖结构,使得一些重要的古人类化石标本的颅内模被复原出来,促进了脑演化的研究。近年来,本文第一作者采用新技术、新方法复原了南京直立人、柳江人等一些重要的中国古人类头骨的颅内模,通过对其颅容量、脑沟回特征、脑不对称性、脑表面的动、静脉血管压迹、各脑叶的大小、形状及比例的研究,获取了中国古人类脑形态特征变化的数据,为探讨东亚地区古人类的演化提供了参考信息。     

脑化石说明了什么?

大自然经常通过化石暴露了自己的秘密,脑化石就在揭示大自然的奥妙中扮演了一个发人深思的角色。脊椎动物死后,脑的软组织被泥砂或其它沉积物替换、矿化成为脑的遗骸,把原来脑子的构造与外型栩栩如生地保存下来。只要我们掌握了大自然留下的脑子的立体摄影——脑化石,就可以测量出动物生前的脑量(图1)。通过对脑量的深入研究,就可以进一步探索动物形态学以及动物进化史上的一些有意义的问题。那么,脑化石能告诉我们一些甚么呢? 早在1871年,达尔文在他的一本名著《人类起源与性的选择》中就曾经明确地指出:脑子的大小与身体的大小是密切相关的。但是受当时科学发展水平的     

化石人类脑进化研究与进展

化石人类脑演化主要是研究在人类进化过程中脑的变化过程,它是随着古神经学和古人类学的发展而发展起来的。古神经学(Paleoneurology),是神经学和占生物学的交叉学科,通过对化石脊椎动物和现牛动物的神经系统的比较,探索脊椎动物神经系统的演化过程。     

魏敦瑞对北京猿人化石的研究及其人类演化理论

魏敦瑞对中国猿人（现为直立人北京亚种）,化石作了详尽而深入的描述并指出了直立人的典型特征。他正确地推测, 在从猿到人的演化过程中首先是直立姿势的采用, 跟着是头骨的变化, 脑的扩张是头骨变化的动因。本世纪前半叶, 魏敦瑞第一次用一个图将所有已知的人类化石集合在一起, 作系统排列的尝试, 来表示整个人类的演化过程。他相信所有化石人类属于一个种, 能够互相杂交, 人类不是起源于一个地方, 而是起源于几个地方。他反对南方古猿曾经参与人类进化的说法, 代之以认为人类进化的早先各期是巨人阶段, 但是他的巨人理论迄今没有化石的证据。关于现代人的起源, 多地区起源说和非洲起源说目前正在激烈争论之中。     

柳江人身体大小和形状——体重、身体比例及相对脑量的分析

发现于广西柳江的更新世晚期人类化石除1具完整的头骨外,还包含有右侧髋骨、骶骨、两段股骨及若干件椎骨。根据各方面的特征分析,初步认定这些化石属于同一个体。这一有利条件为我们比较准确地获取与该个体身体大小和形状有关的指标数据提供了可能。本文通过对柳江人头骨及复原骨盆的测量,计算了柳江人的身高、体重、身体比例、相对脑量等。在此基础上分析了柳江人的身体大小和形状。本研究发现:柳江人化石所代表的个体具有适应温暖气候环境的纤细型身体比例,代表相对脑量的EQ指数5.602大于金牛山、山顶洞等中国更新世中、晚期化石人类,而与包括港川人在内的更新世末期及现代人类的EQ指数接近。柳江人体重52.0kg小于金牛山、山顶洞、尼安德特人等生活在高纬度地区的化石人类,而与港川、非洲的KNM-ER3883、KNM-ER3733等生活在温暖环境的古人类接近。作者认为这些发现除说明柳江人生活的气候环境外,还提示柳江人身体大小、比例及相对脑量与更新世末期及现代人类接近。     

人类颅骨厚度研究的概述与展望

颅骨厚度是重要的解剖学特征，也是常见的测量项目，在体质人类学研究中具有重要意义。目前关于颅骨厚度的研究主要集中在颅骨厚度的年龄与时代变化、人群间和性别间的差异、颅骨厚度的影响因素和与力学性能的关系等几个方面。颅骨厚度的相关研究有助于我们更为全面地了解人类颅骨的形态变异和进化情况。国内对颅骨厚度的关注相对较少，本文通过梳理和归纳已有关于颅骨厚度研究的相关文献数据与结论，对颅骨厚度相关研究进展及其测量方法进行了概述，并对国内颅骨厚度研究作了简要回顾和展望。     

现代人的出现与扩散——中国的化石证据北大核心CSCDCSSCI

自2002年在周口店附近的田园洞发现大约4万年前的现代人化石以来,相继在湖北郧西黄龙洞、广西崇左智人洞等地点发现了早期现代人化石。这些化石发现证实大约10万年前早期现代人在华南地区已经出现。最近在湖南道县福岩洞发现的人类牙齿化石及相关研究进一步揭示具有完全现代形态的人类8万-12万年前在华南局部地区已经出现;而在这个时间段的华北地区,以许家窑人为代表的人类化石形态仍较原始,其演化尚未进入早期现代人阶段。这些研究发现提示,在中国地区,华南是现代人形成与扩散的中心区域,早期现代人以及完全现代类型的人类都可能首先在华南地区出现,然后向华北地区扩散。现有的化石形态证据显示,更新世晚期华南地区人类具有较大的演化变异,可能同时生存有几种不同的演化类群。智人洞属于从古老型智人向现代人演化的过渡类型,而道县则代表着演化进入完全现代类型的人类。基于前人研究及本文的分析,作者认为柳江、资阳、丽江、田园洞等更新世晚期人类化石特征比较进步,在演化上属于与道县相似的现代类型人类。值得注意的是,这些研究进展在引起对现代人在东亚地区出现和扩散关注的同时,古人类学界对其中涉及的许多问题还存在争论。本文在回顾分析这些研究进展的基础上,就相关问题进行了讨论。     

中更新世非洲Bodo人类头骨化石与周口店直立人的比较--中国与非洲人类头骨特征对比之三

发现于埃塞俄比亚MiddleAwash地区Bodo地点距今60万年的人类头骨化石是迄今发现的最为古老和完整的非洲中更新世人类化石。由于Bodo头骨化石在形态特征上兼有直立人与智人的特点,多年来学术界对其分类地位一直存在争议。Rightmire认为Bodo头骨化石与BrokenHill及Petralona等在分类上属于古老型智人的中更新世人类更为接近,是非洲直立人向古老型智人过渡的代表。至少在距今60万年的中更新世早期直立人向古老型智人转变的成种事件在非洲就已经发生。以Bodo头骨为代表的一批更新世中期非洲和欧洲人类化石构成了可能是后期人类祖先的人属海德堡种。这些观点导致了近年学术界对古老型智人在非洲及欧亚出现时间以及更新世中期非洲和欧亚地区古人类相互之间演化关系的关注。基于这样的背景,本文对年代与Bodo化石接近的周口店直立人头骨特征与Bodo头骨的相似及差异表现情况进行了对比研究。结果发现Bodo头骨在一系列特征上与周口店直立人相似,同时在包括颅容量在内的其它一些特征上呈现出后期智人的特点,但总体形态上似乎与直立人更为相似。作者认为尽管这种进化上的镶嵌现象在中国古人类化石记录上也广泛存在,但由于中国人类化石标本在年代上的不确定性,目前还没有可靠的证据说明这种集直立人与智人化石特征为一体的镶嵌性在中国古人类化石出现的时间接近或早于非洲。考虑到中国与非洲直立人生存年代的巨大差异及人类演化的不同步或地区间差异,具有较多后期人类特征表现的人类首先出现在非洲是完全可能的。根据这些研究对比,作者就人类演化的镶嵌现象、更新世中期非洲与亚洲地区人类演化上的差异等问题进行了讨论。     

由环境关系揭示物种演化与人类进化性质及进步实质

综合一些新研究和人类进化历史,阐明人类进化与物种演化性质不同：物种演化是适应特定环境,与特定环境产生依存关系,人类进化是适应不同环境或不再依存特定环境,出现与环境关系性质改变。说明人类进步实质是脑等变化带来与环境关系性质改变,使人类生存和进步出现由偶然性到必然性变化,进步标准是适应不同环境,即出现与特定环境依存度减小的趋势性变化。以环境关系为衡量尺度,对物种进步标准、进化定义和进化树结构提出修正意见。     

现代人的出现与扩散——中国的化石证据

自2002年在周口店附近的田园洞发现大约4万年前的现代人化石以来,相继在湖北郧西黄龙洞、广西崇左智人洞等地点发现了早期现代人化石。这些化石发现证实大约10万年前早期现代人在华南地区已经出现。最近在湖南道县福岩洞发现的人类牙齿化石及相关研究进一步揭示具有完全现代形态的人类8万-12万年前在华南局部地区已经出现;而在这个时间段的华北地区,以许家窑人为代表的人类化石形态仍较原始,其演化尚未进入早期现代人阶段。这些研究发现提示,在中国地区,华南是现代人形成与扩散的中心区域,早期现代人以及完全现代类型的人类都可能首先在华南地区出现,然后向华北地区扩散。现有的化石形态证据显示,更新世晚期华南地区人类具有较大的演化变异,可能同时生存有几种不同的演化类群。智人洞属于从古老型智人向现代人演化的过渡类型,而道县则代表着演化进入完全现代类型的人类。基于前人研究及本文的分析,作者认为柳江、资阳、丽江、田园洞等更新世晚期人类化石特征比较进步,在演化上属于与道县相似的现代类型人类。值得注意的是,这些研究进展在引起对现代人在东亚地区出现和扩散关注的同时,古人类学界对其中涉及的许多问题还存在争论。本文在回顾分析这些研究进展的基础上,就相关问题进行了讨论。     

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.     

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.     

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.     

Brief communication: Possible third molar impactions in the hominid fossil record

Impacted third molars affect 15%–20% of modern Americans and Western Europeans. In contrast, third molar impactions have not been reported in the early hominid fossil record. It is uncertain whether the lack of reports reflects an absence of impactions or a failure to recognize them. This communication is intended to raise awareness of the possibility of impactions by describing the appearance of impacted teeth and by noting two possible instances of impaction in early hominids. Specifically, the mandibular third molars of the Sterkfontein specimen, STS52b (Australopithecus africanus), and the left maxillary third molar of the Lake Turkana specimen, KNM-WT17400 (Australopithecus boisei), are positioned in a manner which suggests that they would not have erupted normally. Both specimens also exhibit strong crowding of the anterior dentition, providing further support for the view that these individuals lacked sufficient space for normal eruption of the third molars. Other published reports of dental crowding in the hominid fossil record are noted, and it is suggested that more attention be paid to dental impaction and dental crowding in hominid evolution. © 1993 Wiley-Liss, Inc.     

Hominid evolution and the concept of techno-cultural dynamism in Africa

Archaeology treats hominid evlution as a bio-cultural phenomenon and reveals the behavioral activities of man as far back as 3 to 5 millions years ago, with different genera and species of hominids and their cultural endowment. Hominid evolution is also considered as exhibiting some technocultural dynamism. The relevance of this concept in the struggle for survival is focused upon in this paper. The ability of the hominids to explore and exploit the environment formed the basis of their survival. The paper concoludes that Africa as the cradle of hominid evolution has however lagged behind in global techno- cultural development.     

The evolution of brain size and intelligence in man

The brain size of hominids has increased approximately threefold during the evolution of the hominids fromAustralopithecus toHomo sapiens. It is proposed that the principal reason for this increase is that larger brains conferred greater intelligence, and greater intelligence conferred a selection advantage. A number of anthropologists have difficulty accepting this thesis because they believe that brain size is not associated with intelligence in man. Evidence is reviewed, and new evidence from two studies is presented, to show that brain size as measured by head size is positively correlated with intelligence as measured by intelligence tests. On two recent samples statistically significant correlations of .21 and .30 were obtained between estimates of brain size and IQ. It is considered that brain size is positively associated with intelligence in man and that this is the major reason for the increase in brain size of the hominids during the last 3.2 million years.     

Hallucial convergence in early hominids

There is a richly documented fossil record of the evolutionary transition from ape-sized brains that are less that one-third the size of modern humans through a series of intermediate-sized brains up to the modern range. The first report on the discovery of the foot of the Stw 573 skeleton emphasized the apparent transitional nature of its great toe [Clarke, R.J., Tobias, P.V., 1995. Sterkfontein Member 2 foot bones of the oldest South African hominid. Science 269, pp. 521-524]. The hallux appeared to be intermediate in its divergence between human-like adduction and ape-like abduction. A major part of this evidence is the medial encroachment of the metatarsal I facet on the medial cuneiform. This study quantifies the variability of this feature in extant hominoids and fossil hominids. The results are consistent with the view that all currently known hominids were specialized for bipedality and lacked the ape-like ability to oppose the great toe.     

Increased breathing control: Another factor in the evolution of human language

Investigation into the evolution of human language has involved evidence of many different kinds and approaches from many different disciplines. For full modern language, humans must have evolved a range of physical abilities for the production of our complex speech sounds, as well as sophisticated cognitive abilities. Human speech involves free‐flowing, intricately varied, rapid sound sequences suitable for the fast transfer of complex, highly flexible communication. Some aspects of human speech, such as our ability to manipulate the vocal tract to produce a wide range of different types of sounds that form vowels and consonants, have attracted considerable attention from those interested in the evolution of language. 1 , 2 However, one very important contributory skill, the human ability to attain very fine control of breathing during speech, has been neglected. Here we present evidence of the importance of breathing control to human speech, as well as evidence that our capabilities greatly exceed those of nonhuman primates. Human speech breathing demands fine neurological control of the respiratory muscles, integrated with cognitive processes and other factors. Evidence from comparison of the vertebral canals of fossil hominids and those of extant primates suggests that a major increase in thoracic innervation evolved in later hominid evolution, providing enhanced breathing control. If that is so, then earlier hominids would have had quite restricted speech patterns, whereas more recent hominids, with human‐like breath control abilities, would have been capable of faster, more varied speech sequences.     

Brain expansion and rotation during hominid evolution

The question of how an endocast (or brain) is oriented within a skull that is positioned in the Frankfurt plane is investigated for African great apes, early hominids STS 71, KNM-ER 1813 and KNM-ER 1470, and modern humans using a 3SPACE digitizer. Our results suggest that, rather than being positioned in the orientation in which isolated brains (endocasts) are conventionally illustrated, brains within skulls that are oriented in the Frankfurt plane tend to be inclined so that the frontal pole is higher than the occipital pole, especially inHomo. These preliminary findings have implications for interpreting early hominid endocasts such as that of AL 162-28.     

Rates of evolution: Is there a conflict between neo-darwinian evolutionary theory and the fossil record?

Neo-darwinian and population genetics theory assumes that the necessary and sufficient set of conditions for all genetic, therefore evolutionary, change has been identified. Punctuationalists have assumed the opposite and cite the fossil record as evidence for change too rapid to be explained in neo-darwinian theory. Data is given here to provide estimates of the rate of evolution in hominid fossils, in living populations, and of that rate which would qualify as punctuational in the hominid fossil record. Evolution in living populations is orders of magnitude greater than that found in the fossil record and far greater than necessary to create apparently instantaneous saltations in the fossil record. It is suggested that such saltations may not represent more rapid rates of evolution but, rather, the persistence of evolutionary change in a given direction for a longer than normal period.