Sexual dimorphism in the Atapuerca-SH hominids: the evidence from the mandibles

The pattern of sexual dimorphism in 15 mandibles from the Atapuerca-SH Middle Pleistocene site, attributed to Homo heidelbergensis, is explored. Two modern human samples of known sex are used as a baseline for establishing sexing criteria. The mandible was divided for analysis into seven study regions and differential expression of sexual dimorphism in these regions is analysed. A total of 40 continuous and 32 discrete variables were scored on the mandibles. The means method given in Regh & Leigh (Am. J. phys. Anthrop.110, 95-104, 1999) was followed for evaluating the potential of correct sex attribution for each variable.On average, the mandibles from the Atapuerca-SH site present a degree of sexual dimorphism about eight points higher than in H. sapiens samples. However, mandibular anatomy of the European Middle Pleistocene hominid records sexual dimorphism differentially. Different areas of the Atapuerca-SH mandibles exhibit quite distinct degrees of sexual dimorphism. For instance, variables of the alveolar arcade present very low or practically no sexual dimorphism. Variables related to overall size of the mandible and symphysis region present a medium degree of sex differences. Finally, ramus height, and gonion and coronoid process present a high degree of sexual dimorphism (indexes of sexual dimorphism are all above 130%). Whether this marked sexual dimorphism in specific anatomical systems affects sexual differences in body size is not completely clear and further studies are needed.Sexual differences detected in the mandible of modern humans have at least two components: differences related to musculo-skeletal development and differences related to a different growth trajectory in males and females (relative development of some of the basal border features). The Atapuerca-SH mandibles display little variation in the basal border, however. The limited variation of this mandibular region may indicate that the pattern of sexual variation in H. heidelbergensis is different enough to that of H. sapiens to caution against simple extrapolation of criteria from one pattern to the other.     

Ontogenetic approach to variation in Middle Pleistocene hominids. Evidence from the Atapuerca-SH mandibles

The ontogenetic processes underlying the variation detected in the European Middle Pleistocene hominids are explored. Regulation of growth parameters of the ontogenetic trajectory of these hominid is discussed in the context of heterochrony schemes. The sample of mandibles coming from the Atauuerca-SH site is used as a study case. The variability shown by these mandibles can be arranged along a morphological gradient, and study of growth directions has shown that the remodelling patterns are in line with the existence of a single morphogenetic pattern controlling the process. In addition, the spectrum of morphology detected in the Atapuerca sample virtually encompasses the entire range of variability detected in European Middle Pleistocene samples. From a morphological viewpoint, a hypermorphic pattern seems to be the responsible for the individual variation detected in the sample. Yet, when this information is related to mandibular growth and craniofacial interactions, evidence of an interindividual gradient of somatic development for the Atapuerca hominids is provided. Several hypotheses concerning growth parameters, however, may account for this gradient. Time and rate hypermorphosis hypothesis are discussed. Coincidence of the gradient of variation detected in this sample and the general mandibular growth trajectory of living species, support the hypothesis that is “time of maturation” of the organism the key factor.     

Morphological analysis of the neanderthal calotte from Ochtendung,Germany

In 1997 one of the authors (AvB) found three pieces of a calotte of a Neanderthal in a crater depression within the slag-cone volcanic-group called “Wannenköpfe” near the village of Ochtendung in the state of Rhineland-Palatinate, Germany. Three stone tools of the Mousterian culture were directly associated with the hominid fossil. The individual can be securely stratigraphically and absolutely chronologically placed within the early glacial phase of the second last glaciation, the Saale. A comparative-morphological analysis confirms earlier analyses that the remains were those of an adult male. The relatively old age establishes it as another important find of an early Neanderthal in Europe. In absolute years it dates roughly to the transitional time period of the latest EuropeanHomo erectus to the earliest Neanderthal. Morphological analysis confirms that the individual is close to a typical Neanderthal with also some additional erectoid characters. This observation supports the most widely accepted view that the Neanderthal of Europe evolved from an autochthonousHomo erectus group.     

Interproximal grooving in the Atapuerca-SH hominid dentitions

The dental sample recovered from the Sima de los Huesos (SH) Middle Pleistocene cave site of the Sierra de Atapuerca (Spain) includes 296 specimens. Interproximal wear grooves have been observed in 20 maxillary and mandibular posterior teeth belonging to at least five of the 32 individuals identified so far in the SH hypodigm. Interproximal grooving affected only the adults, and at an age between 25 and 40 years. The appearance, morphology, and location pattern of the SH wear grooves are similar to those reported in other fossil hominids and in more recent human populations. Two alternative proposals, the toothpicking and the fiber or sinew processing hypotheses, compete for explaining the formation of this anomalous wear. The characteristics observed in the wear grooves of the SH teeth are compatible only with the habitual probing of interdental spaces by means of hard and inflexible objects. Dietary grit may also have contributed to the abrasion of the root walls during the motion of the dental probes. Am. J. Phys. Anthropol. 102:369–376, 1997. © 1997 Wiley-Liss, Inc.     

Dental size variation in the Atapuerca-SH Middle Pleistocene hominids

The Middle Pleistocene Atapuerca-Sima de los Huesos (SH) site in Spain has yielded the largest sample of fossil hominids so far found from a single site and belonging to the same biological population. The SH dental sample includes a total of 452 permanent and deciduous teeth, representing a minimum of 27 individuals. We present a study of the dental size variation in these hominids, based on the analysis of the mandibular permanent dentition: lateral incisors, n=29; canines, n=27; third premolars, n=30; fourth premolars, n=34; first molars, n=38; second molars, n=38. We have obtained the buccolingual diameter and the crown area (measured on occlusal photographs) of these teeth, and used the bootstrap method to assess the amount of variation in the SH sample compared with the variation of a modern human sample from the Museu Antropologico of the Universidade of Coimbra (Portugal). The SH hominids have, in general terms, a dental size variation higher than that of the modern human sample. The analysis is especially conclusive for the canines. Furthermore, we have estimated the degree of sexual dimorphism of the SH sample by obtaining male and female dental subsamples by means of sexing the large sample of SH mandibular specimens. We obtained the index of sexual dimorphism (ISD=male mean/female mean) and the values were compared with those obtained from the sexed modern human sample from Coimbra, and with data found in the literature concerning several recent human populations. In all tooth classes the ISD of the SH hominids was higher than that of modern humans, but the differences were generally modest, except for the canines, thus suggesting that canine size sexual dimorphism in Homo heidelbergensis was probably greater than that of modern humans. Since the approach of sexing fossil specimens has some obvious limitations, these results should be assessed with caution. Additional data from SH and other European Middle Pleistocene sites would be necessary to test this hypothesis.     

Obstetric implications of neanderthal robusticity and bone density

Neanderthal pelvic morphology is not well understood, despite the recent find and analysis of the Kebara 2 pelvis. Many of the proposed hypotheses focus on the possible need for a larger birth canal. A previously unexplored aspect involves possible direct obstetric implications of bone robusticity and density. These characteristics ocan affect obstetrics in modern humans, especially the molding of the neonate's head during parturition: engineering studies have shown that denser neonate cranial bones undergo less deformation, and thicker (more robust) cranial bones would also be expected to deform less during the birth process. These bone characteristics may also result in a less flexible birth canal. Thus, more robust or denser bones could result in the need for a larger birth canal or a smaller neonate head, due to decreased flexibility. Examples from modern populations are discussed and the conclusions applied to Neanderthals, who are known to have had high bone robusticity and may have had high bone density, given their heavy musculature. (A positive association between muscle mass and bone density has been observed repeatedly in modern humans.) We conclude that bone robusticity and density may have obstetrical implications for Neanderthals, with particular importance for neonate head molding during birth.     

Structural features of the binding site of cholera toxin inferred from fluorescence measurements

The dependence on pH of the fluorescence of cholera toxin and its A and B subunits has been studied at 25 degrees C. The fluorescence intensity of cholera toxin is highly pH-dependent. In the pH range 7-9.5 it reaches a maximum corresponding to a quantum yield of 0.076. In the pH range 4-7 a strong increase in fluorescence intensity is observed (delta Q/Qmax = 0.64). Evaluation of the pH sensitivity of the fluorescence intensity of the A and B subunits reveals that the B subunit is mainly responsible for the observed pH effect (delta Q/Qmax for B subunit = 0.64). The intensity changes are paralleled by similar although less pronounced changes in the average fluorescence excited state life-time tau (delta tau/tau max = 0.33 for cholera toxin). Fluorimetric titration of the B subunit, which is related to the indole fluorescence of the lone Trp-88, reveals that the fluorescence intensity changes in the pH range 4-7 are due to reaction of two types of ionizable quencher displaying apparent pKa values of 4.4 and 6.2, respectively. It is suggested that the increase in fluorescence intensity with a midpoint at pH 6.2 is the result of deionization of the imidazolium side-chain of one or two out of the four histidine residues present in each beta-polypeptide chain, whereas a deionized carboxyl group is responsible for the quenching with midpoint at pH 4.4. Complex formation of cholera toxin or B subunit with the monosialoganglioside GM1 or the oligosaccharide moiety of GM1 (oligo-GM1) completely prevents the quenching by both quenchers. Addition of 6 M urea also eliminates the pH effect. The quenching is not the result of the dissociation of the B subunit into its constituent monomers. Upon fluorimetric titration of cholera toxin or B subunit above pH 9, a progressive drop in both fluorescence intensity and tau occurs. This decrease could be due to energy transfer from the indole moiety of Trp-88 to ionized tyrosines or by quenching through an unprotonated epsilon-amino group of lysine. Fluorimetric titration of the A subunit indicates that the tryptophan fluorescence is only moderately altered by ionizable groups displaying a pKa in the range 4 to 9. Activation of A subunit does not affect this lack of pH sensitivity. Above pH 9, however, a much more significant drop in the fluorescence intensity of activated A subunit occurs. The structural implications of the results are discussed.     

Bone remodeling of the Homo heidelbergensis mandible; the Atapuerca-SH sample.

Growth by bone remodeling is one of the key mechanisms responsible for skeletal morphology. This mechanism consists of the coordinated activity of two cellular groups: osteoblasts and osteoclasts, which are responsible for bone deposition and resorption, respectively. Information obtained from the study of these remodeling growth fields allows us to understand how species-specific craniofacial form is achieved. These data can help to explain the facial growth differences among Primates, both extinct and extant. The aim of this study was to obtain the distribution of growth remodeling fields of the Homo heidelbergensis mandible (Atapuerca-SH sample), and to infer the growth processes responsible for its specific morphology. A Reflected Light Microscope (RLM) was used to identify the microfeatures of the bone surface related to bone deposition and resorption. Results show that H. heidelbergensis presents a specific growth field distribution, which differs slightly between immature and adult individuals. Interpretation of these maps indicates that the mandible of H. heidelbergensis presents noteworthy variability in the symphyseal region. Two distinct patterns of growth are seen, one of those unique for this species and the other similar to that of Homo sapiens. The lingual side of the mandibular corpus has a resorption area found only in this species and one that includes a variable extension in immature and adult individuals. Finally, the mandibular ramus is characterized, among other aspects, by a large resorption field on its buccal surface. Considering the mandible as a whole, the bone remodeling pattern obtained in this work shows that lower facial growth in H. heidelbergensis is dominated mainly by forward growth, illustrated by the strong inward displacement of the ramus, which is in agreement with the Enlow's “V” growth principle.     

Calcite in the mandibles of a marine polychaete

Calcite forms a thick pad on the anterior-ventral surface of the mandibles of Eunice aphroditois (Pallas) (Annelida, Polychaeta, Eunicida, Eunicea). The calcite pads apparently serve as an abrasive in feeding; they are possibly renewed by continuous calcite deposition. Mandibular calcite pads are not usually recognized in the fossil record because they are destroyed during the leaching of limestones for the recovery of scolecodonts.     

Physicochemical features of the HERG channel drug binding site

Blockade of hERG K(+) channels in the heart is an unintentional side effect of many drugs and can induce cardiac arrhythmia and sudden death. It has become common practice in the past few years to screen compounds for hERG channel activity early during the drug discovery process. Understanding the molecular basis of drug binding to hERG is crucial for the rational design of medications devoid of this activity. We previously identified 2 aromatic residues, Tyr-652 and Phe-656, located in the S6 domain of hERG, as critical sites of interaction with structurally diverse drugs. Here, Tyr-652 and Phe-656 were systematically mutated to different residues to determine how the physicochemical properties of the amino acid side group affected channel block by cisapride, terfenadine, and MK-499. The potency for block by all three drugs was well correlated with measures of hydrophobicity, especially the two-dimensional approximation of the van der Waals hydrophobic surface area of the side chain of residue 656. For residue 652, an aromatic side group was essential for high affinity block, suggesting the importance of a cation-pi interaction between Tyr-652 and the basic tertiary nitrogen of these drugs. hERG also lacks a Pro-Val-Pro motif common to the S6 domain of most other voltage-gated K(+) channels. Introduction of Pro-Val-Pro into hERG reduced sensitivity to drugs but also altered channel gating. Together, these findings assign specific residues to receptor fields predicted by pharmacophore models of hERG channel blockers and provide a refined molecular understanding of the drug binding site.     

The surface-dwelling from layer I of Mira site: Main features and interpretations

Situated in Dnieper valley, in the central part of continental Ukraine, the site of Mira yields two well-preserved Palaeolithic occupation levels, possessing features of true living floors. The uppermost layer I, presenting remains of autumn-winter seasonal occupation, is dated to between 32,000 and 31,000 cal BP. Layer I contains EUP assemblage combining the technological and morphological features of local Middle and Upper Palaeolithic. Layer I includes many various objects, like pits, postholes, hearths, bone and ashes accumulations etc. Remains of sub-circular surface-dwelling construction with an area of about 14.5 m² were recognised in the uppermost layer due to numerous postholes and specific characteristics of the living floor. Micro-stratigraphic and spatial features recognise two distinct construction elements, namely the external irregular spherical contour and the inscribed slightly asymmetrical rectangular contour. An entrance, associated with four ashy lenses of likely smoke hearths, was recognised oriented east-wards toward a current river channel. This outer, close to the entrance, zone of construction contains numerous flints, in particular, flint tools, thousands tiny debris and waste-flakes of tool resharpening and reshaping, as well as bone ornaments and ornamented bone pieces, and also a fragment of a human molar. On the contrary, the practically free lithics though containing plenty of burned organic material, the back part of the construction, likely separated from the outer zone by a special partition, was seemingly served as a sleeping zone. It is possible to conclude about the recovery of remains of permanent carcass surface cylindrical dwelling that found analogies in ethnographical records. Keeping in mind the age and geographical position, we deal with the earliest instance of complicate surface-dwelling construction currently known in the steppe area of the East European plain.     

Age estimation of the roe deer (Capreolus capreolus) mandibles from the Mesolithic site of Star Carr, Yorkshire, based on radiographs of mandibular tooth development

A scheme developed by Brown & Chapman, (1991a), which allocates scores to stages of tooth development from radiographs, has been applied to 12 of the 28 roe deer ( Capreolus capreolus ) mandibular rami recovered from the early Mesolithic site of Star Carr, N. Yorkshire ( c. 7500 BC). The mandibles of 35 known-age roe deer kids from Britain and Denmark were also radiographed. The developmental stages of all premolars and molars were analysed and assigned a score using the scheme. After radiographing, four of the Star Carr rami could be paired with a fair degree of confidence, reducing the sample size from 12 to 10 roe deer. Comparisons of scores between the archaeological and modem populations revealed that seven of the Star Carr mandibular rami are aged approximately 10 to 11 months. The youngest is nine months, and the oldest 12 months. These results have significant implications for determining the seasonal presence of humans at the site. Contrary to the received view of a purely late spring/summer occupation of the site, it is now apparent that Star Carr was also visited during the late winter and early spring.     

Occurrence of 19p- in an infant with multiple dysmorphic features

A case with partial deletion on the short arm of one chromosome 19 and multiple dysmorphic features is reported.     

Multivariate analysis of Gigantopithecus mandibles

Multivariate analysis of measurements of the teeth and mandibles of Gigantopithecus species has been conducted, using several methods. Results indicate Gigantopithecus is an aberrant form, less related to australopithecines and gorillas than the latter are to each other. Gracile and robust australopithecines differ considerably more than do male and female gorillas.