周口店直立人3号头骨的性别和年龄:南京直立人头骨的启示

周口店直立人3号头骨是1929年发现的, 被步达生鉴定为一青春期或成年期的女性个体的头骨。后来, 步达生改变了观点, 认为该头骨是代表青春期早期的男性个体。魏敦瑞同意步达生的意见, 但认为该头骨应属于8岁或9岁的男性个体。此后,周口店直立人3号头骨的性别和年龄的鉴定一直困扰着许多古人类学家。1993年发现的南京直立人成年女性头骨在形态上和尺寸上与周口店直立人3号头骨有相近之处。本文对周口店直立人3号头骨与南京直立人等头骨作了形态和测量上的比较, 结果表明: 从头骨的尺寸大小, 眶上圆枕、枕圆枕和肌嵴的发育程度以及头骨骨壁厚度等来看, 周口店3号头骨有理由被认为是女性个体的。该头骨的鼓骨裂、下颌骨关节窝、泪腺窝、额窦、骨缝等都无法证明其是幼年个体。周口店直立人3号头骨应代表成年女性个体。     

中国直立人牙齿特征变异及其演化意义

以往研究发现,中国直立人化石呈现较大的形态变异。对于这种变异程度及造成变异的原因,在古人类学界有不同的认识。有学者提出在直立人阶段,中国古人类已经呈现明显的区域性差异,但也有人认为这些差异似乎没有规律性。近年,本文作者采用不同方法对若干地点的中国直立人牙齿特征及其变异进行了系列研究,取得了一些新的发现和认识。本文在回顾总结这些研究的基础上,结合对其它一些地点中国直立人牙齿特征的观测对比,对中国直立人牙齿特征表现特点及变异作了进一步的分析。本研究发现,中国直立人牙齿特征具有较大的变异范围。这些变异似可分为两种主要类型,元谋、建始、郧县梅铺、和县牙齿呈现出较多的原始特点,代表一种原始类型;周口店、沂源等地点的标本特征相对进步,表现出更多的典型直立人特征。其它一些地点的直立人化石呈现出混合或中间状态。值得注意的是部分呈现出原始牙齿特征类型的中国直立人的生存年代相对较晚,其牙齿特征的原始性与生存时代不具有对应关系。作者认为中国直立人牙齿特征类型反映了更新世早期和中期不同直立人群的演化状态。原始类型牙齿特征的形成不仅与演化时序性和地理分布有关,还在一定程度上反映了一些中国直立人群的演化隔离。     

南京直立人与印尼、周口店直立人的面颅形态比较

本文对南京直立人1号头骨与印尼Sangiran17头骨和周口店11号头骨作面颅形态上的比较。结果表明,南京1号头骨的面颅与周口店11号头骨的有许多共同的形态:较小的面颅尺寸、眶下区呈扁平状、较扁平的上部颜面、眶上圆枕较纤细、中眶型的眼眶、发育有颧切迹、上颌骨颧突基部位置较高等。南京1号头骨的面颅在另外一些方面显示出与Sangiran17头骨的形态上相近:阔上面型的面型、眶上圆枕内侧部比外侧部稍靠前、眶上圆枕下缘略呈平直状、鼻梁发育有中矢锐嵴、鼻梁横向呈峰状拱起、颧骨下缘外展、颧结节相对位置稍偏外侧、颧骨较高等。南京直立人面颅兼具周口店直立人11号头骨的和印尼直立人Sangiran17头骨的形态。从南京直立人的面颅形态可以推测我国南方的与北方的直立人群之间在体质形态上已经出现一定程度的地理变异。南京1号头骨面颅上的某些形态特征,如颧骨下缘外展等,可能反映了特征的梯度变异现象。这些特征的梯度变异的成因目前还很难确定。     

周口店直立人头骨创伤与人工切割痕迹辨析

自20世纪40年代魏敦瑞提出北京猿人（周口店直立人）可能存在“暴力争斗”和“同类残食”的观点之后,引发了一些学者和科普大众对北京猿人是否为“食人族”的争论。由于原始化石标本在第二次世界大战期间丢失,本文以保存下来的周口店（ZKD）直立人头骨原始模型和素描图为研究材料,对魏敦瑞提出的具有创伤和人工切割痕迹的5件标本（ZKD II、ZKD VI、ZKD X、ZKD XI和ZKD XII）进行辨析。结果显示：1）ZKD X、ZKD XI、ZKD XII这3件标本的头盖部,有7处可以确定为生前遭受非致死性撞击导致的局部创伤;2）ZKD VI头盖骨残片周边的及中央区域的断痕,为个体生前遭受致死性的暴力打击导致;3）ZKD II和ZKD VI顶骨位置疑似人工切割痕迹的沟槽实际上是动物啃咬或自然因素导致的,北京猿人“同类残食”的观点在本文研究中没有得到证实。北京猿人头骨的创伤痕迹都位于头盖部,以顶骨居多,其次为额骨,符合人群之间暴力冲突产生创伤的位置。     

关于北京猿人用火的证据:研究历史、争议与新进展

周口店第1地点古人类用火证据是该遗址一系列重大科学发现中的一项重要内容,在很长时期内,作为同类遗存中最早的记录及其分析论证结果被国际学术界广泛接受。但随着少数西方学者的质疑和挑战,从上世纪八十年代中期开始在这一问题上出现争议,其后开展的埋藏学和地球化学分析又得出进一步否定的结论。但周口店遗址的洞穴地层十分复杂,目前残存的堆积与古人类生存时期的状态有重大差别,与当初大规模发掘时见证的遗物遗迹的分布与埋藏状况也有很大不同,在剖面表层做局部有限的采样分析并不足以推翻以前的系统性研究结论,何况很多否定性的意见源自学术理论思潮的转变和缺乏具体分析的常规性推理。从以前的发掘记录和各种分析结果看,周口店遗址埋藏着丰富的古人类用火证据,这些证据不是孤立的,是可以相互验证和支持的。2009年以来在遗址开展的新的发掘与研究获得重要进展,揭示出具有结构的火塘、烧骨、石灰化的灰岩块等原地用火产生的遗物与遗迹,对相关材料的现代科技分析进一步确定这些遗存的人类用火性质。这样,遗址上文化层的用火证据变得明确无误,相关争议终可尘埃落定。对于下部地层中的用火证据,尚需做同样的发掘、分析和研究工作。     

南京1号直立人头骨与肯尼亚KNM-ER 3733人类头骨化石的形态比较

KNM-ER 3733人类头骨化石的年代为距今1.78百万年,1975年发现于肯尼亚。Walker和Leakey注意到这具头骨与周口店直立人的在脑颅形态上很相近,但二者在年代上相差大约1百万年,故认为直立人形态在这1百万年期间是稳定的。长期来此观点缺乏更多的人类化石证据来支持。1993年在中国发现了南京1号人类头骨化石。该头骨与KNM-ER 3733头骨一样兼具脑颅和面颅,且都属于成年女性个体,但南京1号人类头骨化石的年代比KNM-ER 3733人类头骨化石的要晚大约1百万年。因此,南京1号人类头骨是目前所知的可用来验证直立人头骨形态是否在1百万年期间保持稳定的唯一合适的人类头骨化石材料。形态比较表明,这两个人类头骨化石的脑颅虽然在眶上圆枕上沟的发育程度、眶后收缩的程度、额骨横向隆起的程度、角圆枕和乳后突的发育与否、顶骨形状以及骨壁厚度的表现上有所差异,但有更多的形态性状显示出相近。这些相近表现在脑颅的长、宽、高值上;颅容量上;脑颅的低矮性上;脑颅最大宽之位置上;额骨、顶骨、枕骨之矢弧值的比例上;眶上圆枕的纤细上;顶骨的大小和矢向扁平性上;颞线位置和颞鳞顶缘的形状上;枕鳞的低宽形状上;上枕鳞与下枕鳞之间的转折形状和比例上;枕骨圆枕和枕骨圆枕上沟的发育程度上等。这两具头骨的面颅虽然有同属突颌型的面角、皆发育有鼻骨间嵴、两鼻骨组成的上部宽度与下部宽度皆差别很大,但有更多的形态性状显示出差别。这些差别表现在面型上、颜面上部扁平度上、眶形和眶型上、上颌额突外侧面的朝向上、鼻骨横向隆起程度上、鼻梁外突程度上、鼻型上、颧骨下缘外展程度上、颊高上、颧上颌下缘的形状上、上颌颧突基部的位置上以及颧结节的位置上等。因此,南京1号头骨与KNM-ER 3733头骨之间在脑颅上显示出较多的相近性状,在面颅上则显示出较多的相异性状。脑颅方面的相近性状大多具有分类上的鉴别价值。这两个头骨脑颅形态的相近支持把KNM-ER 3733头骨鉴定为"直立人"的观点;也提示了南京1号头骨的脑颅似乎保持着1百多万年前的"祖先"形态。如果直立人的某些成员在至少1百万年期间保持着形态稳定的话,则这种形态上的稳定主要是表现在脑颅形态上。这两具头骨的面颅形态上较大差异的意义,目前尚不清楚。     

Developmental age and taxonomic affinity of the Mojokerto child,Java, Indonesia

An increasing number of claims place hominids outside Africa and deep in Southeast Asia at about the same time that Homo erectus first appears in Africa. The most complete of the early specimens is the partial child's calvaria from Mojokerto (Perning I), Java, Indonesia. Discovered in 1936, the child has been assigned to Australopithecus and multiple species of Homo, including H. modjokertensis, and given developmental ages ranging from 1–8 years. This study systematically assesses Mojokerto relative to modern human and fossil hominid growth series and relative to adult fossil hominids. Cranial base and vault comparisons between Mojokerto and H. sapiens sapiens (Hss) (n = 56), Neandertal (n = 4), and H. erectus (n = 4) juveniles suggest a developmental age range between 4 and 6 years. This range is based in part on new standards for assessing the relative development of the glenoid fossa. Regression analyses of vault arcs and chords indicate that H. erectus juveniles have more rounded frontals and less angulated occipitals than their adult counterparts, whereas Hss juveniles do not show these differences relative to adults. The growth of the cranial superstructures and face appear critical to creating differences in vault contours between H. erectus and Hss. In comparison with adult H. erectus and early Homo (n = 27) and adult Hss (n = 179), the Mojokerto child is best considered a juvenile H. erectus on the basis of synapomorphies of the cranial vault, particularly a metopic eminence and occipital torus, as well as a suite of characters that describe but do not define H. erectus, including obelion depression, supratoral gutter, postorbital constriction, mastoid fissure, lack of sphenoid contribution to glenoid fossa, and length and breadth ratios of the temporomandibular joint. Mojokerto is similar to other juvenile H. erectus in the degree of development of its cranial superstructures and its vault contours relative to adult Indonesian specimens. The synapomorphies which Mojokerto shares with H. erectus are often considered autapomorphies of Asian H. erectus and confirm the early establishment and long-term continuity of the Asian H. erectus bauplan. This continuity does not, however, necessarily reflect on the pattern of origin of modern humans in the region. Am J Phys Anthropol 102:497–514, 1997. © 1997 Wiley-Liss, Inc.     

Accelerated Evolutionary Rate May Be Responsible for the Emergence of Lineage-Specific Genes in Ascomycota

The evolutionary origin of “orphan” genes, genes that lack sequence similarity to any known gene, remains a mystery. One suggestion has been that most orphan genes evolve rapidly so that similarity to other genes cannot be traced after a certain evolutionary distance. This can be tested by examining the divergence rates of genes with different degrees of lineage specificity. Here the lineage specificity (LS) of a gene describes the phylogenetic distribution of that gene’s orthologues in related species. Highly lineage-specific genes will be distributed in fewer species in a phylogeny. In this study, we have used the complete genomes of seven ascomycotan fungi and two animals to define several levels of LS, such as Eukaryotes-core, Ascomycota-core, Euascomycetes-specific, Hemiascomycetes-specific, Aspergillus-specific, and Saccharomyces-specific. We compare the rates of gene evolution in groups of higher LS to those in groups with lower LS. Molecular evolutionary analyses indicate an increase in nonsynonymous nucleotide substitution rates in genes with higher LS. Several analyses suggest that LS is correlated with the evolutionary rate of the gene. This correlation is stronger than those of a number of other factors that have been proposed as predictors of a gene’s evolutionary rate, including the expression level of genes, gene essentiality or dispensability, and the number of protein-protein interactions. The accelerated evolutionary rates of genes with higher LS may reflect the influence of selection and adaptive divergence during the emergence of orphan genes. These analyses suggest that accelerated rates of gene evolution may be responsible for the emergence of apparently orphan genes. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Martin Kreitman]     

Evolutionary implications of error amplification in the self-replicating and protein-synthesizing machinery

Summary Evolutionary constraints operating on animal mitochondrial tRNA were estimated to be reduced to about 1/30 of those that apply to cytoplasmic tRNA. In the nuclear-cytoplasmic system, an effect of a mutation tRNA is likely to be amplified through positive feedback loops consisting of DNA polymerases, RNA polymerases, ribosomal proteins, aminoacyl-tRNA synthetases, tRNA processing enzymes, and others. This amplification phenomenon is called an error cascade and the loops that cause it are called error loops. The freedom of evolutionary change of cytoplasmic tRNA is expected to be severely restricted to avoid the error cascade. In fact, cytoplasmic tRNA is highly conserved during evolution. On the other hand, in the animal mitochondrial system, all of the proteins involved in error loops are coded for in the nuclear genome and imported from the cytoplasm, and accordingly the system is free from the error cascade. The difference in constraints operating on animal tRNA between cytoplasm and mitochondria is attributed to the presence or absence of error loops. It is shown that the constraints on mitochondrial tRNA in fungi are not as relaxed as those in animals. This observation is attributed to the presence of an error loop in fungal mitochondria, since at least one protein of the mitochondrial ribosome is coded for in the mitochondrial genome of fungi. The evolutionary rates of proteins involved in the processing of genetic information are discussed in relation to the error cascade.A preliminary version of this paper was presented at the International tRNA Workshop (Hakone, Japan, March 1983) and at the Second International Colloquium on Endocytobiology (Tübingen, FRG, April 1983)