Biomechanical scaling of the hominoid mandibular symphysis

Experimental investigation of mandibular bone strain in cercopithecine primates has established that the mandible is bent in the transverse plane during the power stroke of mastication. Additional comparative work also supports the assumption that the morphology of the mandibular symphysis is functionally linked to the biomechanics of lateral transverse bending, or "wishboning" of the mandibular corpus. There are currently no experimental data to verify that lateral transverse bending constitutes an important loading regime among hominoid primates. There are, however, allometric models from cercopithecoid primates that allow prediction of scaling patterns in hominoid mandibular dimensions that would be consistent with a mechanical environment that includes wishboning as a significant component. This study uses computed tomography (CT) scans to visualize cortical bone distribution in the anterior corpus of a sample of four genera of extant hominoids. From the cortical bone contours, area properties of the mandibular symphysis are calculated, and these variables are subjected to an allometric analysis to detect whether scaling of jaw dimensions are consistent with a wishboning loading regime. Scaling of the hominoid symphysis recalls patterns observed in cercopithecoid monkeys, which lends indirect support for the hypothesis that wishboning is an integral part of the masticatory loading environment in living apes. Inclination of the symphysis, rather than changes in cross-sectional shape or development of the superior transverse torus, represents a morphological solution for minimizing the potentially harmful effects of wishboning in the jaws of these primates.     

The functional significance of primate mandibular form.

A stress analysis of the primate mandible suggests that vertically deep jaws in the molar region are usually an adaptation to counter increased sagittal bending stress about the balancing-side mandibular corpus during unilateral mastication. This increased bending stress about the balancing side is caused by an increase in the amount of balancing-side muscle force. Furthermore, this increased muscle force will also cause an increase in dorso-ventral shear stress along the mandibular symphysis. Since increased symphyseal stress can be countered by symphyseal fusion and as increased bending stress can be countered by a deeper jaw, deep jaws and symphyseal fusion are often part of the same functional pattern. In some primates (e.g., Cercocebus albigena), deep jaws are an adaptation to counter bending in the sagittal plane during powerful incisor biting, rather than during unilateral mastication. The stress analysis of the primate mandible also suggests that jaws which are transversely thick in the molar region are an adaptation to counter increased torsion about the long axis of the working-side mandibular corpus during unilateral mastication. Increased torsion of the mandibular corpus can be caused by an increase in masticatory muscle force, an increase in the transverse component of the postcanine bite force and/or an increase in premolar use during mastication. Patterns of masticatory muscle force were estimated for galagos and macaques, demonstrating that the ratio of working-side muscle force to balancing-side muscle force is approximately 1.5:1 in macaques and 3.5:1 in galagos during unilateral isometric molar biting. These data support the hypothesis that mandibular symphyseal fusion is an adaptative response to maximize unilateral molar bite force by utilizing a greater percentage of balancing-side muscle force.     

Mandibular function in Galago crassicaudatus and Macaca fascicularis: an in vivo approach to stress analysis of the mandible.

Single-element and/or rosette strain gages were bonded to mandibular cortical bone in Galago crassicaudatus and Macaca fascicularis. Five galago and eleven macaque bone strain experiments were performed and analyzed. In vivo bone strain was recorded from the lateral surface of the mandibular corpus below the postcanine tooth row during transducer biting and during mastication and ingestion of food objects. In macaques and galagos, the mandibular corpus on the balancing side is primarily bent in the sagittal plane during mastication and is both twisted about its long axis and bent in the sagittal plane during transducer biting. On the working side, it is primarily twisted about its long axis and directly sheared perpendicular to its long axis, and portions of it are bent in the sagittal plane during mastication and molar transducer biting. In macaques, the mandibular corpus on each side is primarily bent in the sagittal plane and twisted during incisal transducer biting and ingestion of food objects, and it is transversely bent and slightly twisted during jaw opening. Since galagos usually refused to bite the transducer or food objects with their incisors, an adequate characterization of mandibular stress patterns during these behaviors was not possible. In galagos the mandibular corpus experiences very little transverse bending stress during jaw opening, perhaps in part due to its unfused mandibular symphysis. Marked differences in the patterns of mandibular bone strain were present between galagos and macaques during the masticatory power stroke and during transducer biting. Galagos consistently had much more strain on the working side of the mandibular corpus than on the balancing side. These experiments support the hypothesis that galagos, in contrast to macaques, employ a larger amount of working-side muscle force relative to the balancing-side muscle force during unilateral biting and mastication, and that the fused mandibular symphysis is an adaption to use a maximal amount of balancing-side muscle force during unilateral biting and mastication. These experiments also demonstrate the effects that rosette position, bite force magnitudes, and types of food eaten have on recorded mandibular strain patterns.     

Growth-related changes in prehistoric Jomon and modern Japanese mandibles with emphasis on cortical bone distribution

Cortical bone distribution of the anthropoid mandibular symphysis has been addressed in relation to mechanical stress generated by mastication. To examine whether or not bone mass and distribution patterns of the human mandibular symphysis could be interpreted as an example of functional adaptation, we compared the skeletal growth series of two populations, prehistoric Jomon, considered to represent a "robust" mandibular morphology associated with a presumed heavier masticatory load, and modern Japanese. Results showed that the adult Jomon symphysis possessed significantly greater bone mass and thicker cortical bone compared to the modern Japanese condition. However, the second moments of area did not differ significantly between the two, indicating comparable rigidity against bending. Furthermore, the Jomon mandibles of the infant to juvenile stages exhibited most of the adult characteristics, in both bone mass/distribution of the symphysis and in mandibular corpus/ramus morphologies. The present study also demonstrated the presence of a growth pattern of symphyseal cortical thickness, common to both the Jomon and the modern Japanese series. In both populations, subsequent to deciduous molar occlusion, cortical bone tends to be thickest at the inferolingual symphysis, at the location where the highest tensile stresses presumably occur during mastication. These findings suggest that the "robust" characteristics of the Jomon mandible are initially manifested early in development, and that the effect of mechanical stimulus to bone mass formation in the human symphysis is largely confined to a regulatory role during growth modeling.     

Diet and Mandibular Morphology in African Apes

Investigations seeking to understand the relationship between mandibular form, function, and dietary behavior have focused on the mandibular corpus and symphysis. African apes vary along a gradient of folivory/frugivory, yet few studies have evaluated the morphology of the mandibular corpus and symphysis in these taxa, and the investigations have yielded mixed results. Specifically, studies using external metrics have identified differences in mandibular proportions that analysis of cortical bone distribution has not substantiated. I contribute to the ongoing debate on the relationship between jaw form and dietary behavior by comparing mandibular corporal and symphyseal shapes in African apes. Importantly, and in contrast to previous studies of African ape internal geometry, I include the Virunga mountain gorillas (Gorilla beringei beringei), the ape most specialized toward a folivorous diet. I test the hypotheses that 1) Gorilla beringei beringei always has significantly more robust mandibular corpora and symphyses, relative to mandibular length, than all other African apes and 2) all gorillas have significantly more robust mandibular corpora and symphyses, relative to mandibular length, than Pan. Results demonstrate that the folivorous mountain gorillas consistently exhibit a relatively wider mandibular symphysis and corpus than all other African apes. Furthermore, all gorillas consistently exhibit relatively more robust mandibular corporal and symphyseal dimensions than Pan. The results indicate that among African apes, mountain gorillas are better able to counter lateral transverse bending (wishboning) loads at the symphysis and torsional loads at the corpus. All gorillas are likewise better able to resist wishboning and vertical bending at the symphysis, and sagittal bending and torsion at the corpus, than Pan, findings that are consistent with masticating relatively tougher foods, repetitive loading of the jaws, or both. I offer possible explanations for the lack of concordance in results between studies that have analyzed the biomechanical properties of African ape mandibles and others that have relied on external metrics. More comprehensive study of the internal geometry of the mandible is needed to resolve whether African apes differ morphologically in ways predicted by dietary variation.

Mandibular morphology and diet in the genusCebus

The influence of hard-object feeding on the size and shape of the mandibular corpus was investigated through a comparative biomechanical analysis of the jaws of adult femaleCebus apella andCebus capucinus. Computed tomography (CT) was used to discern the amount and distribution of cortical bone at M₂ and symphyseal cross sections. From these data, the biomechanical properties of the mandibular corpus were determined to assess the structural rigidity of the jaw with respect to the bending, torsional, and shear stresses that occur during mastication and incision. The mandibles ofC. apella are demonstrably more robust than those ofC. capucinus in terms of biomechanical rigidity; differences in corporeal size rather than shape largely account for the enhanced robusticity in the sample ofC. apella. The differences that separate the two taxa probably represent a structural response to the mechanical demands of durophagy inC. apella. These observations suggest that specialization on a diet of hard objects may be expected to result in an overall hypertrophy of bony contours throughout the mandibular corpus.     

Do mandibular cross‐sectional properties and dental microwear give similar dietary signals?

Previous animal experimental work evaluating the effects of dietary consistency on mastication was generally limited to studies of either mandibular structure or rates and types of tooth wear. Control groups fed hard diets (HD) consistently exhibited increased cortical remodeling and/or bone strength when compared to groups fed soft diets (SD). Results of tooth-wear studies showed faster rates of tooth wear in HD animals. This study evaluates the effects of dietary differences on both mandibular structural morphology and postcanine dental microwear in the same animals. We examined mandibles and dentitions from eight miniature swine, raised from 4 weeks to 9 months of age on HD and SD (n = 4, each group). Mandibular structural properties were calculated from peripheral quantitative computed tomography slices at the dp3-dp4 and dp4-M1 junctions. Dental microwear analysis was performed on mandibular lingual crushing facets of dp4 and M1, using photomicrographs of high-resolution casts taken at 500x magnification in a scanning electron microscope. Our results suggest that between the dp3-dp4 contact, HD animals have mandibles that are stronger and more rigid mediolaterally than SD animals. At the dp4-M1 contact, HD animals have mandibles that are stronger and more rigid mediolaterally, dorsoventrally, and in torsion than SD animals. Dental microwear results indicate that SD pigs have higher incidences of pitting and more overall microwear features on their premolars than do HD pigs, yet there are no significant differences in molar microwear morphology between the dietary groups. Near-significant correlations exist between pit size and dorsoventral bending strength, but only for HD pigs. These results suggest that dietary consistency significantly affects both mandibular structure and dental microwear, yet direct correlations between the two are complicated by a number of factors.     

The structure of the mandibular corpus and its relationship to feeding behaviours in extant carnivorans

Assuming some optimization of bone structure to applied mechanical loadings in vivo , different killing and feeding behaviours in carnivores should be reflected in observed differences in cross-sectional shape of their mandibular corpora. Section moduli are used to gauge the magnitudes of bending moments in the mandibular corpus and, when dentary length is controlled, the magnitudes of forces applied to the corpus. Comparisons are made of section moduli at the P₃P₄ and P₄M₁ interdental gaps among canids, felids and hyaenids; in canids only, the M₁M₂ interdental gap was also studied. Local variations in loadings are identified by comparing the section moduli at adjacent loci along the corpus within each family.
The findings of this study show that the precarnassial corpora of canids and hyaenids have greater strength in bending than the corpora of felids of similar body weight. This is taken to reflect relatively greater bending moments under loading in the corpora of canids and hyaenids due, in part, to their elongate dentaries (relative to body weight). Relative to dentary length , however, the precarnassial corpora of felids and hyaenids have much greater strength in bending than the corpora of canids. These scaling relationships appear to reflect the high customary forces (i.e. not moments) applied to the precarnassial corpora of felids and hyaenids with sustained canine killing bites and with bone ingestion using the premolars, respectively. An increase in bending strength of the corpus caudal to the camassial blade in canids is interpreted to be an adaptation for bone-crushing with the postcarnassial molars.     

Biomechanics of cross-sectional size and shape in the hominoid mandibular corpus

Mandibular cross sections of Pan, Pongo, Gorilla, Homo, and two fossil specimens of Paranthropus were examined by computed tomography (CT) to determine the biomechanical properties of the hominoid mandibular corpus. Images obtained by CT reveal that while the fossil hominids do not differ significantly from extant hominoids in the relative contribution of compact bone to total subperiosteal area, the shape of the Paranthropus corpora indicates that the mechanical design of the robust australopithecine mandible is fundamentally distinct from that of modern hominoids in terms of its ability to resist transverse bending and torsion. It is also apparent that, among the modern hominoids, interspecific and sexual differences in corpus shape are not significant from a biomechanical perspective. While ellipse models have been used previously to describe the size, shape, and subsequent biomechanical properties of the corpus, the present study shows that such models do not predict the biomechanical properties of corpus cross-sectional geometry in an accurate or reliable manner. The traditional "robusticity" index of the mandibular corpus is of limited utility for biomechanical interpretations. The relationship of compact bone distribution in the corpus to dimensions such as mandibular length and arch width may provide a more functionally meaningful definition of mandibular robusticity.     

Effects of age and sex on intra-group spacing behaviour in juvenile savannah baboons,Papio cynocephalus cynocephalus

Twenty juvenile members of known genealogies in two baboon groups were studied over a 16-month period to evaluate a number of predictions about juvenile spacing behaviour based on the natural history of savannah baboons. Young juveniles (1–2·5 years old) approached more frequently and spent more time in proximity to other group members than did old juveniles (3–5·5 years old). In particular, young juveniles associated more closely with their mothers, particular adult males (possible fathers) and age-peers than did old juveniles. Approaches of young juveniles towards unrelated, high-ranking adults were more likely to occur during feeding than were those of old juveniles. Also, following such an approach, young juveniles were more likely than old juveniles to begin feeding immediately. The overall rates of feeding of old juveniles were depressed when they were in proximity to unrelated, high-ranking adults, whereas the feeding rates of young juveniles were not. Juvenile males approached adult males more often than did juvenile females. Juvenile females approached unrelated adult females more often than did juvenile males. Sex differences also existed in juveniles' choices of unrelated adult female neighbours. Juvenile females associated most often with lactating females, whereas juvenile males associated primarily with cycling females. During group resting, juvenile females approached adult females from higher-ranking matrilines more often than they approached adult females from lower-ranking matrilines. Juvenile males did not exhibit this attraction. Also, among old juveniles, females associated closely with their mothers, whereas males did not. Taken together, the results support the hypotheses that juvenile baboons associate with group members in ways that (1) enhance the probability of surviving an early period of high mortality, (2) create opportunities for social learning of sex-typical behaviours/skills, and, for females, (3) facilitate acquisition of familial dominance status.     

Spatial patterning of bone stiffness in the anterior mandibular corpus of Macaca fascicularis: Implications for models of bone adaptation

Elastic modulus of bone from the anterior mandibular corpus was determined via microindentation in a mixed-sex ontogenetic sample (N = 14) of Macaca fascicularis. This investigation focused on the hypothesis that material heterogeneity in the macaque mandibular symphysis—provided an accounting of age and sex variation—is explicable as a means to homogenize strains in this region. Experimental data and theoretical models of masticatory loading indicate that in the absence of material compensation, large strain gradients exist in the anterior mandibular corpus of macaques, particularly between lingual and labial cortical plates owing to the effects of lateral transverse bending. Microindentation data indicate that juvenile macaques possess less stiff bone than their subadult and adult counterparts; however, sex differences in elastic modulus are not apparent. Anisotropy variation is idiosyncratic; that is, there is not a common pattern of variation in stiffness sampled among orthogonal planes across individuals. Similarly, differences in stiffness between lingual and labial cortical plates, as well as differences among alveolar, midcorpus, and basal regions are inconsistently observed. Consequently, we find little evidence in support of the hypothesis that spatial variation in bone stiffness functions to homogenize strains in the anterior corpus; in fact, in some individuals, this spatial variation serves to exacerbate, rather than to minimize, strain gradients. The mechanical benefit of elastic modulus variation in the macaque mandibular symphysis is unclear; this variation may not confer adaptive benefit in terms of structural integrity despite the fact that such variation has discernible functional consequences. Am J Phys Anthropol 156:649–660, 2015. © 2014 Wiley Periodicals, Inc.     

The mechanical significance of morphological variation in the macaque mandibular symphysis during mastication

Catarrhine symphyseal morphology displays considerable variation. Although this has been related to dentition, phylogeny, sexual dimorphism, and facial orientation, most emphasis has been given to the functional significance of the symphysis to mechanical loading during mastication. The current state of knowledge regarding the mechanical significance of the symphysis is based on a combination of in vivo experimental and comparative studies on Macaca fascicularis. These approaches have provided considerable insight into the stereotypical patterns of loading in the symphyseal region during chewing and hypotheses related to the associated symphyseal morphologies. Finite element analysis (FEA) was used to assess how in silico manipulation translates into the mechanical loading hypotheses previously proposed experimentally. In particular, this study tests the form-function relationship of the symphysis of an adult M. fascicularis mandible during lateral transverse bending and dorsoventral shear of the mandibular symphysis, and a series of modified hypothetical morphologies including absence/presence of tori and variation in the inclination and depth of the symphysis. FEA results of this study support previous findings that stresses associated with lateral transverse bending and dorsoventral shear of the mandibular symphysis can be minimized via an increased labio-lingual thickness in the superior transverse torus, an oblique symphyseal inclination, and/or an increased symphyseal depth. The finding that reduction of strains related to lateral transverse bending and dorsoventral shear can be achieved through a number of different morphologies contributes to our understanding of the influence of morphological and/or developmental constraints, such as dental development, on symphyseal form.     

Factors affecting morphological development of the sagittal otolith in juvenile and adult small yellow croaker (Larimichthys polyactis Bleeker, 1877)

The morphology and morphometrics of the sagittal otoliths of small yellow croaker (Larimichthys polyactis) from the southern Yellow Sea were investigated. Study objectives were to evaluate the shape variability and morphometric variables of sagittae of juveniles and adults as related to developmental changes and habit shift. A total of 152 fish were sampled from April to June of 2012 and 2013, along the coastal waters of the Lüsi spawning ground in the southern Yellow Sea. Changes in otolith shapes from the juvenile to the adult were presented with the rim development through the entire‐lobed‐entire transition and with the curvature of the cauda toward the ventral margin. The otolith elongation in the juvenile stage occurred at 10–20 mm standard length (SL) and was likely associated with the formation of otolith accessory growth centers from larvae to juvenile. The L. polyactis sagittal otoliths acquired their definitive shape at 130 mm SL maturity. Ontogeny on otolith shape might be related, for example, to the factors of metamorphic development, feeding habitat, and ambient water salinity, which varied throughout the growth of L. polyactis.     

Patterns of growth of the mandibular corpus in spotted hyenas (Crocuta crocuta) and cougars (Puma concolor)

Differences in jaw morphology among adult carnivorans are well established, but the ontogenetic mechanisms by which these differences arise are largely unexplored. Mandibular ontogeny in Crocuta crocuta and Puma concolor is analysed biomechanically using principles of beam theory. In each species, the development of cross-sectional properties of the mandibular corpus associated with rigidity under loading follows a biphasic pattern of growth. In early postnatal growth, deposition of cortical bone appears to be constrained by the overall weaker tissue with which juvenile skeletons are constructed and by the need to volumelrically accommodate the developing teeth within their bony crypts. Thus, this stage of growth is characterized by a net periosteal deposition of bone and a swelling of the medullary cavity. In late postnatal growth, the constraints on endosteal deposition of bone are relieved as the permanent teeth erupt; thus, cortical thicknesses increase sharply by periosteal expansion as well as medullary contraction. Finally, it is noted that basic differences in jaw construction between Crocuta and Puma appear to develop prenatally as they are largely in place at birth. Hence, postnatal development enhances, but does not soley contribute to, the biomechanical differences in the jaws of these species.     

Development of the vertebral column and caudal complex in a flyingfish,Parexocoetus mento mento (Teleostei: Exocoetidae)

The developmental pattern of the vertebral column and caudal complex in juvenile (16.9 mm SL) to adult (112.2 mm SL)Parexocoetus mento mento is described Juvenile external caudal morphology was similar to the adult condition, although juveniles exhibited various internal ontogenetic changes. Osteological develoment was almost completed at 60–69 mm SL. Complete ossification of the vertebral column and caudal complex appeared to be the optimal condition giving strength for flight. Loss of perforations in the centra, neural and haemal arches may be consistent with the rigid and straightened body position during take-off. Some ontogenetic changes in the caudal complex were related to functional aspects. Ankylosis of the NPU2 spur to the uroneural notch, fusion of hypurals 3+4 and 5 and the elongated hypural 1+2 (lower hypural) were linked to the acquisition of stability and strength in the caudal complex.     

Cranial adaptation to a high attrition diet in Japanese macaques

Primates with diets that require greater occlusal forces to process exhibit anteroposteriorly shorter, vertically deeper faces, more anteriorly placed masseter attachment areas, and broader, taller mandibular corpora compared to closely related species/populations. Japanese macaques (Macaca fuscata)eat different, perhaps mechanically tougher to process, foods than other macaques do. Accordingly, they should exhibit structural features of the skull related to dissipating great occlusal loads. To test this hypothesis I compared cranial variables amongst wild-caught, adult female skulls (n = 85) of M. fuscataand three other macaque species (M. mulatta, M. fascicularis,and M. nemestrina)and applied least-squares and reduced-major-axis regression analysis and principal components analysis (PCA) to 17 cranial variables reflecting facial, vault, and mandibular dimensions. When scaled for size, the Japanese macaque has a vertically deeper and anteroposteriorly shorter face,a broader but not taller mandibular corpus, and a more anteriorly placed masseter muscle than the other three macaques do. The first PCA axis isolates variation due to a suite of characters related to mechanical efficiency in dissipating occlusal loads (vertically deep face and broad corpus) and differentiates the Japanese macaques from the other species. This, coupled with reported dietary differences among species, suggests that Japanese macaques are selected for dissipating greater occlusal loads than other macaques are. The presence of a narrow mandible relative to cranial breadth and a hyperrobust mandibular corpus width suggests that axial torsion is a significant influence in the masticatory regime of M. fuscata.The lack of an increase in corpus height indicates that parasagittal bending is not as significant an influence. Geographic and climatic influences cannot account for the patterns of variation between M. fuscataand the other macaques.     

Ontogenetic change of bone microstructures and its ethological implication in Champsosaurus (Diapsida,Choristodera)

Compact cortex in a Champsosaurus (Diapsida, Choristodera) femur is ontogenetically replaced with extensively developed cancellous bone. This histological shift, together with retention of calcified cartilage to late ontogenetic stage, was previously considered to show that adult champsosaurs were more adapted to aquatic environments than juveniles. However, the new histological examination reveals the nearly amedullar condition of a juvenile femur consisting of thick periosteal cortex and less cancellous bone tissue and the amedullar but more porous condition of adult femora. This likely demonstrates that the femoral inner structure of the juvenile is denser than those of the adults, and therefore, juveniles were more aquatic. It is suggested that morphological variations between two sympatric species of Champsosaurus reflect sexual dimorphism in a single species and limb bones with more robust morphology, showing better terrestrial adaptation for nesting on land, belong to females. The similarity of gross limb bone morphology between juveniles and inferred adult males indicates aquatic habitats for juveniles, coincident with the new interpretation of bone microstructures. No differences are, however, recognised in femoral microstructure between inferred sexes in adults. The possibly lowered density of femur in adults is considered as an adaptation to increase the mobility in water.     

Temporal trends in femoral diaphyseal torsional asymmetry among the Arikara associated with postural behavior

Average femoral torsion has been reported to differ among populations, and several studies have observed a relatively high prevalence of femoral anteversion asymmetry in Native Americans, especially females. This study investigates sexual dimorphism and temporal trends in femoral torsional asymmetry among the Arikara from the seventeenth to the early nineteenth century. To establish if there are population differences, femoral torsion was first measured using a direct method on a diverse comparative sample of Native Americans from the Southwest, Midwest, and Great Plains as well as American Whites and Blacks. To examine temporal trends among the Arikara, femoral torsion was examined using the orientation of the maximum bending rigidity at subtrochanteric in 154 females and 164 males from three temporal variants of the Arikara Coalescent tradition. There is significant sexual dimorphism in femoral torsional directional and absolute asymmetry among most Native American samples, but not among American Whites and Blacks. Among the Arikara there is significant sexual dimorphism in femoral torsional asymmetry in all three temporal variants, and asymmetry in femoral torsional asymmetry increased significantly from the protohistoric to the early historic period among females. The increased femoral torsional asymmetry is likely associated with a common side‐sitting posture observed in historic photographs of Great Plains females. Historic Arikara females may have habitually sat in this compulsory position for extended periods while conducting domestic chores. The dramatic change from the protohistoric to historic period suggests a cultural change in sitting posture among females that was widespread across the Northern Plains. Am J Phys Anthropol 154:512–524, 2014. © 2014 Wiley Periodicals, Inc.     

Oxygen uptake through gills and skin in relation to body weight of an air-breathing siluroid fish, Saccobranchus (=Heteropneustes) fossilis

Oxygen uptake through gills and skin has been measured in juvenile and adult Saccobranchus fossilis and its relationship represented by the equation Vo₂= aW^b. When air-breathing is allowed the O₂ uptake via the gills and skin together increases by powers of 1 -084, 0–986 and 1–328 in juvenile, adult and juvenile+adult respectively. When air-breathing is prevented the slope (b) for O₂ uptake via the gills appear to be less in juveniles (0–765) than in adults (0–784) and juveniles+adults together (0–814). Under the same experimental condition, the slope for O₂ uptake via the gills+skin is also less in juveniles (0–478) than in adults (0–799) and juveniles+adults (0–755). Further decrease in the exponent value is found for the oxygen uptake of skin in relation to body weight under surfacing-prevented conditions ( b = 0–542). Different exponent values for juvenile and adult fishes may be due to their different growth pattern and physiology.     

Relationship between canine dimorphism and mandibular morphology in the hamadryas baboon and the Japanese monkey

To examine morphological interrelationships between canine size and mandibular corpus shape, inter-sex comparisons were made in the hamadryas baboon and the Japanese monkey, known to display extreme and moderate canine dimorphism, respectively. Results of adult comparisons showed that all mandibular dimensions were significantly larger in the males than in the females in both species. In the hamadryas baboon, the males also exhibited a higher ratio of anterior to posterior corpus height than the females. This sex difference in corpus shape was not significant in the Japanese monkey, indicating lack of involvement of canine dimorphism. Analysis of mandibular growth patterns in the hamadryas baboon demonstrated that significant sexual size difference did not occur before incisor eruption, and that the anteriorly high corpus of the adult male mandible was associated with a rapidly increasing symphyseal height after incisor eruption. It was also shown that the female canine started to erupt shortly after incisor eruption, while the forming male canine continued to stay near the mandibular base and developed further in size until eruption. The relative positions of the incisors kept shifting upwards even after eruption in the males, while they hardly changed in the females. It is therefore suggested that the prolonged development and size increase of the male canine is accompanied by further enlargement of the symphysis, resulting in the higher anterior corpus of the adult males compared to the adult females. The results thus indicate the importance of understanding the spatial relationships of the developing teeth in interpreting mandibular morphology.