禄丰古猿牙齿釉质生长线与个体发育问题研究

运用扫描电子显微镜,对４枚禄丰古猿牙齿（恒齿）釉质结构进行了观察研究,发现：禄丰古猿牙齿釉质表面有明显的釉面横纹结构;釉面横纹的密度向牙颈方向逐渐增大;观察记数了４枚牙齿的釉面横纹数,进而推算出牙冠的形成时间和年龄。与化石人科成员,现代人及现生大猿比较,禄丰古猿牙冠发育模式及时间,与南方古猿纤细种比较接近或相似,明显长于南方古猿粗壮种,有别于现生大猿。     

华南化石猩猩前部牙齿釉面横纹与牙冠形成时间研究

釉面横纹的分布与数目可以反映牙齿生长发育的时间和速率变化, 在化石研究中能为复原个体生活史提供重要依据。本研究运用扫描电子显微镜观察华南化石猩猩门齿、犬齿釉面横纹分布与数目, 并估算门齿和犬齿牙冠形成时间, 结果如下: 牙冠从牙尖至牙颈方向釉面横纹分布密度有疏密变化, 牙尖釉面横纹密度小于10条/mm, 中间至牙颈釉面横纹密度较尖部增大, 大约10-15条/mm; 犬齿釉面横纹数目多于门齿, 雄性犬齿釉面横纹数目多于雌性; 根据釉面横纹计数及其生长周期的组织切片观察结果, 估算门齿牙冠形成时间大约为2.97-6.66年, 犬齿雄性长于雌性, 分别为6.25-11.31年和4.28-7.29年。与一些古猿、早期人类、现代人以及现生大猿比较, 华南化石猩猩釉面横纹整体密度稍大于南方古猿和傍人, 小于黑猩猩、大猩猩、现代人和禄丰古猿; 除侧门齿外, 华南化石猩猩釉面横纹数目明显多于南方古猿、傍人和现代人, 与大猩猩接近; 华南猩猩前部牙齿牙冠形成时间与现生大猿、禄丰古猿差别不大, 与现生猩猩最相近, 长于南方古猿和傍人。     

禄丰古猿前部牙齿的釉面横纹与牙冠形成时间

釉面横纹的数目可用于推断个体牙齿的牙冠形成时间,在生长发育研究中具有重要的意义。本研究运用数码体视显微镜和扫描电镜观察了云南石灰坝禄丰禄丰古猿(简称禄丰古猿)30枚齿冠完整的前部牙齿,包括上下颌中门齿6枚、侧门齿10枚和犬齿14枚。根据唇侧面釉面横纹计数的观察结果,分别以7天和9天芮氏线生长周期,估算各齿型的牙冠形成时间,结果显示:以生长周期7天计算,中门齿牙冠形成时间约为3.6-4.1年,侧门齿牙冠形成时间约为2.7-3.7年,犬齿牙冠形成时间约为4.2-7.0;以生长周期9天计算,中门齿牙冠形成时间约为4.4-5.2年,侧门齿牙冠形成时间约为3.4-4.7年,犬齿牙冠形成时间约为5.2-8.8年。为更深入地了解禄丰古猿牙冠形成时间在不同齿型及性别间足否存在明显差异,本文用SPSS软件对其进行显著性差异检验。采用小样本平均值的t值假设检验(置信区间为95%),结果如下:禄丰古猿前部牙齿的牙冠形成时间在各类牙齿的上下颌中不存在显著性差异;犬齿牙冠形成时间存在非常显著的性别差异,雄性牙冠形成时间明显长于雌性,侧门齿也存在显著的性别差异,而中门齿性别间则无显著性差异。此外对禄丰古猿中门齿,侧门齿和犬齿的牙冠形成时间进行单因素方差分析并两两对比,结果显示中门齿与侧门齿的牙冠形成时间不存在显著性差异,而犬齿与中门齿和侧门齿均存在显著性差异,犬齿牙冠形成时间明显长于门齿。同时也对禄丰古猿前部牙齿的牙冠形成时间与齿冠高进行相关性分析,其结果表明两者有显著的正相关性。将禄丰古猿与其他古猿和现生大猿、南方古猿以及人属成员进行对比,结果显示其前部牙齿牙冠形成时间长于原修康尔猿、南方古猿、傍人、人属成员,接近于蝴蝶禄丰古猿和大猩猩,而明显小于黑猩猩、华南化石猩猩及现生猩猩。     

禄丰古猿带状牙釉质发育不全的再观察

牙釉质发育不全是反映牙齿发育过程中遭遇生理性刺激的有效指标。本文对来自云南禄丰石灰坝的禄丰古猿(Lufengpithecus lufengensis)261枚单颗恒齿进行观察分析,发现223枚牙齿普遍罹患带状牙釉质发育不全(LEH),比例为85.4%,高于其它已报道过的中新世古猿、化石人科成员及现生大猿。另一方面,用扫描电镜和数码显微镜着重分析了12枚犬齿,均有多条LEH,且雄性犬齿的LEH条数多于雌性;相邻LEH间的釉面横纹数在14-25条之间。结合禄丰古猿釉面横纹的生长周期(9天),估算相邻LEH之间的时间间隔为4.8-6.6月,季节性营养不良可能是禄丰古猿釉质发育不全的主要原因。     

元谋古猿下颌臼齿三维立体特征

采用欧氏距离矩阵分析（EDMA）方法对72枚元谋古猿及作为对比样本的10枚禄丰古猿、现生大猿类和人类下颌臼齿齿冠13个测量标志点三维测量数据的统计分析显示：元谋古猿在下颌臼齿齿冠三维形态测量特征上与禄丰古猿最为接近。与现生大猿类及人类相比,元谋古猿和禄丰古猿均与人类之间呈现出非常显著的差异,而与猿类较为接近。它们两者及生大猿类均与人类之间具有许多共同的差异表现特点。元谋古猿在下颌臼齿三维测量特征方面与三种现生大猿类各自之间的差别表现相似。其中,元谋古猿与猩猩之间的牙齿形态特征上似乎更为接近。但目前对这些特征相似差异的含义尚难以确定。     

禄丰古猿牙齿釉质发育不全的观察研究

对出自禄丰石灰坝的26个禄丰古猿下颌齿列的246枚恒齿进行了观察研究,发现禄丰古猿具有普遍的带状釉质发育不全（LEH）现象,个体LEH比例为100%,恒齿LEH比例为85%。乳齿几乎没有LEH现象,第一恒臼齿的LEH比例也很低仅57%。根据牙齿萌出顺序及现代大猿的牙齿发育年龄特征,作者推断2—3岁之前的幼儿古猿很少出现釉质发育不全现象,这可能与母体的营养关照有关。禄丰古猿的LEH的发生频率具有明显的季节性,结合中新世晚期气候变化特征、古猿的生态环境、生活习性及食性特征分析,作者推测：季节性营养不良可能是造成禄丰古猿釉质发育不全的主要原因。     

禄丰古猿幼年下颌骨的研究

本文记述了１９８０年在云南省禄丰县石灰坝村禄丰古猿地点发现的最晚中新世禄丰古猿的幼年下颌骨ＰＡ８６９。作者将此标本与西瓦古猿的,南方古猿的和现生大猿的幼年下颌进行了对比,记述了禄丰古猿幼下年颌的若干特征,这些特征表明禄丰古猿幼年下颌体各部主要尺寸的比例关系本种成年的十分相似。     

巨猿牙齿釉质厚度及对食性适应与系统演化的意义

步氏巨猿(Gigantopithecus blacki)是更新世时期生活于我国华南地区的一种超大型猿类, 它的体态特征和演化分类倍受关注。牙齿釉质厚度在探讨灵长类食性、环境适应以及系统演化方面具有重要意义。本文利用显微CT技术构建18颗巨猿臼齿虚拟模型, 测量其釉质厚度。将巨猿釉质厚度与现代人、现生类人猿、古人类、中新世古猿及其他现生灵长类进行比较, 从牙齿釉质厚度探讨巨猿的食性适应和系统演化问题。结果发现巨猿的实测釉质厚度是目前所有已知现生和化石灵长类中最厚的, 只有傍人、南非早期人属及奥兰诺古猿三种化石灵长类与之接近; 如果考虑不同物种牙齿与身体大小的关联因素, 相对釉质厚度指数显示巨猿属于"厚"釉质类型, 但非"超厚"类型, 低于奥兰诺古猿、傍人、南非早期人属; 巨猿与某些中新世古猿 (如原康修尔猿尼安萨种、非洲古猿)、南方古猿、东非早期人属、亚洲直立人以及现代人、现生卷尾猴的相对釉质厚度指数相近。巨猿的厚釉质特征与其食性和环境适应密切相关, 使得牙齿具有非常强的抗磨损功能, 能够适应长时间的咀嚼和研磨食物。从釉质厚度的系统演化角度推测, 厚釉质应该是人类祖先的特征性状, 巨猿有可能是早期人类支系演化过程中的一个特化旁支, 同时也不排除巨猿是从某种具有厚釉质的中新世古猿旁支平行演化而来的可能性。     

云南元谋小河地区古猿地点的小型猿类化石

本文记述的云南元谋小河地区古猿地点发现的一种小型猿类。它的牙齿形态比晚中新世的禄丰粗壮池猿进步。而牙齿的某些形态介于粗壮池猿和现生长臂猿之间,它的发现为探讨现生长臂猿的起源与进货提供了新的化石依据。依哺乳动物群的初步研究,其时代稍晚于禄丰古猿地点的时代。鉴于它的形态特征和地史分布,作者将它订为一新属新种：进步滇猿Ｄｉａｎｏ ｐｉｔｈｅｃｕｓ ｐｒｏｇｒｅｓｓｕｓ ｇｅｎ．ｅｔ ｓｐ．ｎｏｖ。     

广西化石猩猩牙齿釉质厚度研究

华南和东南亚发现大量更新世的猩猩牙齿化石。本研究应用CT扫描三维重建的技术方法研究了广西更新世化石猩猩牙齿釉质厚度,并与现生类人猿、现代人、化石类人猿以及早期人类进行比较分析。结果显示:广西猩猩同类牙齿的釉质厚度与牙齿大小相关性很小;臼齿和前臼齿釉质厚度在上下颌之间不存在显著性差异;来自广西不同地区的猩猩化石牙釉质厚度无显著差异。与早期人科成员相比,广西猩猩的牙釉质相对较薄,平均与相对釉质厚度值都明显小于南方古猿、傍人。与早期人属相比,小于直立人、尼人以及非洲和欧洲的早期人属化石。与现代人和现生灵长类相比,广西化石猩猩釉质厚度明显大于大部分猴类和非洲大猿;平均釉质厚度稍大于现生猩猩,而与现代人更为接近;相对釉质厚度小于现代人,而与现生猩猩差异不大,都属于偏厚型釉质。本文讨论了釉质厚度与系统分类演化、食性适应的相关问题,作者推测釉质厚度可能是物种的特征属性,与牙齿功能适应有密切关联。     

Age at first molar emergence in Lufengpithecus lufengensis and its implications for life-history evolution

The late Miocene hominoid Lufengpithecus from Yunnan Province, China, is crucial for understanding hominoid evolution in Asia. Given that age at first permanent molar emergence is a key life-history trait in primates, the present study determined the age at death of the Lufengpithecus lufengensis juvenile PA868, which was in the process of erupting its first molar. Using a perikymata periodicity of 7-11 days, along with estimation of cusp formation time and the postnatal delay of crown mineralization, perikymata counts obtained from the permanent central incisor and canine germs indicate that the age at death of PA868 was 2.4-4.5 years based on the central incisor germ, and 2.5-4.7 years based on the canine germ. The age at the first molar emergence was actually slightly younger (by about 0.3 years), as demonstrated by tiny wear facets on this tooth, which indicate that gingival emergence had occurred sometime before death. The average age at first molar emergence of Lufengpithecus lufengensis PA868 is estimated to be 3.2-3.3 years, with a range of 2.1-4.4 years. In comparison to extant primates and other fossil hominoids, the life history of Lufengpithecus lufengensis is similar to that of extant great apes and the Miocene hominoids Afropithecus turkanensis and Sivapithecus parvada, as well as Plio-Pleistocene Australopithecus, and different from monkeys, gibbons, and modern humans.     

Perikymata spacing and distribution on hominid anterior teeth

We documented the spacing and distribution of perikymata on the buccal enamel surface of fossil hominin anterior teeth with reference to a sample of modern human and modern great ape teeth. A sample of 27 anterior teeth attributed to Australopithecus (5 to A. afarensis, 22 to A. africanus) and of 33 attributed to Paranthropus (6 to P. boisei, and 27 to P. robustus) were replicated and sputter-coated with gold to enable reflected light microscopy of their surface topography. Anterior teeth were then divided into 10 equal divisions of buccal crown height. The total perikymata count in each division of crown height was recorded using a binocular microscope fitted with a vernier micrometer eyepiece. Then the mean number of perikymata per millimeter was calculated for each division. Similar comparative data for a modern sample of 115 unworn human anterior teeth and 30 African great ape anterior teeth were collected from ground sections. Perikymata counts in each taxon (together with either known or presumed periodicities of perikymata) were then used to estimate enamel formation times in each division of crown height, for all anterior tooth types combined. The distributions of these estimates of time taken to form each division of crown height follow the same trends as the actual perikymata counts and differ between taxa in the same basic way. The distinction between modern African great apes and fossil hominins is particularly clear. Finally, we calculated crown formation times for each anterior tooth type by summing cuspal and lateral enamel formation times. Estimates of average crown formation times in australopiths are shorter than those calculated for both modern human and African great ape anterior teeth. The data presented here provide a better basis for exploring differences in perikymata spacing and distribution among fossil hominins, and provide the first opportunity to describe four specimens attributed to Homo in this context. Preliminary data indicate that differences may exist among the species attributed to early Homo, especially between Homo ergaster and Homo rudolfensis on the one hand, and Homo habilis sensu strico on the other.     

Morphometric analysis of hominoid lower molars from Yuanmou of Yunnan Province, China

Shape analyses of cross-sectional mandibular molar morphology, using Euclidean Distance Matrix Analysis, were performed on 79 late Miocene hominoid lower molars from Yuanmou of Yunnan Province, China. These molars were compared to samples of chimpanzee, gorilla, orangutan,Lufengpithecus lufengensis, Sivapithecus, Australopithecus afarensis, and human mandibular molars. Our results indicate that the cross-sectional shape of Yuanmou hominoid lower molars is more similar to the great apes that to humans. There are few differences between the Yuanmou,L. lufengensis, andSivapithecus molars in cross-sectional morphology, demonstrating strong affinities between these three late Miocene hominoids. All three of the fossil samples show strong similarities to orangutans. From this, we conclude that these late Miocene hominoids are more closely related to orangutants than to either the African great apes or humans.     

Enamel thickness,microstructure and development in Afropithecus turkanensis

Afropithecus turkanensis, a 17-17.5 million year old large-bodied hominoid from Kenya, has previously been reported to be the oldest known thick-enamelled Miocene ape. Most investigations of enamel thickness in Miocene apes have been limited to opportunistic or destructive studies of small samples. Recently, more comprehensive studies of enamel thickness and microstructure in Proconsul, Lufengpithecus, and Dryopithecus, as well as extant apes and fossil humans, have provided information on rates and patterns of dental development, including crown formation time, and have begun to provide a comparative context for interpretation of the evolution of these characters throughout the past 20 million years of hominoid evolution. In this study, enamel thickness and aspects of the enamel microstructure in two A. turkanensis second molars were quantified and provide insight into rates of enamel apposition, numbers of cells actively secreting enamel, and the time required to form regions of the crown. The average value for relative enamel thickness in the two molars is 21.4, which is a lower value than a previous analysis of this species, but which is still relatively thick compared to extant apes. This value is similar to those of several Miocene hominoids, a fossil hominid, and modern humans. Certain aspects of the enamel microstructure are similar to Proconsul nyanzae, Dryopithecus laietanus, Lufengpithecus lufengensis, Graecopithecus freybergi and Pongo pygmaeus, while other features differ from extant and fossil hominoids. Crown formation times for the two teeth are 2.4-2.6 years and 2.9-3.1 years respectively. These times are similar to a number of extant and fossil hominoids, some of which appear to show additional developmental similarities, including thick enamel. Although thick enamel may be formed through several developmental pathways, most Miocene hominoids and fossil hominids with relatively thick enamel are characterized by a relatively long period of cuspal enamel formation and a rapid rate of enamel secretion throughout the whole cusp, but a shorter total crown formation time than thinner-enamelled extant apes.     

Investigation into the relationship between perikymata counts and crown formation times

This study reports on a sample of 12 modern human incisors (from two archaeological sites) that were viewed with a scanning electron microscope and whose perikymata were counted. These 12 incisors more than doubles the previously published sample size of modern human incisors that have served as the published standard for perikymata number in human incisors and have been employed to define taxonomic relationships in fossil hominids. All previously published fossil specimens fall within the expanded range of modern human perikymata counts and can no longer be considered distinctively nonhuman in dental formation time. Five neandertal incisors from the Krapina site in Croatia, Yugoslavia, were also examined. These incisors substantially expand the previous data base for counts of perikymata in Homo sapiens neanderthalensis, likewise overlapping the previously published modern human range. Finally, the validity of methods that have been employed for deriving crown formation times from perikymata counts in fossil hominines is called into question. Utilizing the presently known perikymata ranges for modern humans, these methods do not predict the range of known crown formation times of modern humans as assessed from studies of living children.     

Macroscopic and microscopic analyses of linear enamel hypoplasia in Plio-Pleistocene South African hominins with respect to aspects of enamel development and morphology

This study uses macroscopic and microscopic methods to analyze the expression of linear enamel hypoplasia (LEH) in Plio-Pleistocene South African hominins. LEH is a developmental defect of enamel that is used in many anthropological contexts as a physiological stress indicator. Previous research has not settled the question as to whether differences in LEH expression exist between Paranthropus and Australopithecus and if they exist, to what extent these differences might be explained simply by taxonomic differences in enamel development and morphology rather than by differential stress experience. In this study, the analysis of LEH is conducted with respect to differences between Paranthropus and Australopithecus in aspects of enamel development and morphology that are thought to influence LEH expression. Two factors impacting LEH expression are considered: the duration of enamel formation, and the spacing of perikymata. It is predicted that if the first factor strongly influences the expression of LEH, then there should be fewer defects per tooth in Paranthropus because of its abbreviated crown formation spans (and fast extension rates) relative to Australopithecus. It is also predicted that because Australopithecus has more densely packed perikymata in comparable regions of the crown than Paranthropus, this taxon should, on average, have narrower defects than Paranthropus. To address these questions, 200 Australopithecus and 137 Paranthropus teeth were examined for LEH, and the analysis of defect width with respect to perikymata spacing was conducted on tooth impressions examined under a scanning electron microscope using INCA (Oxford Instruments) measurement software. Data support the first prediction: Australopithecus does have significantly more defects per canine tooth than Paranthropus. Data do not support the second prediction in large part because several Australopithecus specimens have wide groove defects in which perikymata are not visible and enamel is irregular. Such wide grooves are not predicted by perikymata spacing such that alternative explanations, including taxonomic differences in ameloblast sensitivity and the duration/severity of disruptions to enamel growth, must be considered.     

Variation in modern human premolar enamel formation times: implications for Neandertals

A recent study demonstrated that variation in enamel cap crown formation in the anterior teeth is greater than that in the molars from two geographically distinct populations: native indigenous southern Africans and northern Europeans. Eighty southern African and 69 northern European premolars (P3 and P4) were analyzed in the present study. Cuspal, lateral, and total enamel formation times were assessed. Although cuspal enamel formation times were not consistently different between the two populations, both lateral and total enamel formation times generally were. Bonferroni-corrected t-tests showed that southern Africans had significantly shorter lateral enamel formation time for five of the six cusps, as well as significantly shorter total enamel formation time for these same cusps. An analysis of covariance performed on the lingual cusps of the upper third and fourth premolars showed that differences in enamel formation times between these populations remained when crown height was statistically controlled. A further goal of this study was to ascertain, based on perikymata counts, what Neandertal periodicities would have to be in order for their teeth to have lateral enamel formation times equivalent to either southern Africans or northern Europeans. To this end, perikymata were counted on 32 Neandertal premolars, and the counts were inserted into regression formulae relating perikymata counts to periodicity for each population and each tooth type. Neandertal enamel formation times could be equivalent to those of southern Africans or northern Europeans only if their hypothetical periodicities fall within the range of periodicities for African apes and modern humans (i.e., 6-12 days). The analysis revealed that both populations could encompass Neandertal timings, with hypothetical periodicities based on the southern African population necessitating a lower range of periodicity (6-8 days) than those based on the northern European population (8-11 days).