Ellen Schulz-Kornas

Morphology is not Destiny: Discrepancy between Form, Function and Dietary Adaptation in Bovid Cheek Teeth

Mammal teeth have evolved morphologies that allow for the efficient mechanical processing of different foods, therefore increasing dietary energy uptake for maintenance of high metabolic demands. However, individuals masticate foods with biomechanical properties at odds with the optimal function of a given tooth morphology. Here, we investigate tooth form and function using two quantitative 3D methods at different scales on the same individuals of nine bovid species. Dental topometry quantifies the gross morphology, and therefore, reflects evolutionary adaptive patterns. Surface texture analysis infers mechanical occlusal events, which reflect the actual tooth function, and is free from the influence of morphology. We found that tough foods can be satisfactorily exploited by grazing species with enamel ridge morphologies not more complex than those found in intermediate feeders and browsers. Thus, the evolution of enamel complexity is likely determined by a balance between adaptation and constraints. Wider enamel ridges seem to be a common functional trait in bovids to compensate for severe wear from abrasive foods and/or chipping from hard foods. Our results demonstrate that supposedly essential functional adaptations in tooth morphology may not be required to process food efficiently. This emphasizes the large plasticity between “optimal” morphology and the potential function of the tooth, and underscores the need to appreciate (apparently) maladaptive structures in mammalian evolution as nevertheless effective functioning units.

A set of hypotheses on tribology of mammalian herbivore teeth

Once erupted, mammal cheek teeth molars are continuously worn. Contact of molar surfaces with ingesta and with other teeth contribute to this wear. Microscopic wear features (dental surface texture) change continuously as new wear overprints old texture features. These features have been debated to indicate diet. The general assumption in relating occlusal textures to diet is that they are independent of masticatory movements and forces. If this assumption is not accepted, one needs to propose that occlusal textures comprise signals not only from the ‘last supper’ but also from masticatory events that represent ecological, species- or taxon-specific adaptations, and that occlusal textures therefore give a rather unspecific, somehow diet-related signal that is functionally inadequately understood. In order to test for mechanical mechanisms of wear, we created a hypothesis matrix that related sampled individuals with six tribological variables. Three variables represent mechanically relevant ingesta properties, and three represent animal-specific characteristics of the masticatory system. Three groups of mammal species (free ranging Cetartiodactyla and Perissodactyla, free ranging primates, and artificially fed rabbits) were investigated in terms of their 3D dental surface textures, which were quantified employing ten ISO 25178 surface texture parameters.Wefirst formulated a set of specific predictions based on theoretical reflections on the effects of diet properties and animal characteristics, and subsequently performed discriminant analysis to test which parameters actually followed these predictions. We found that parameters Vvc, Vmc, Sp, Sq allowed the prediction of both, ingesta properties and properties of the masticatory system, if combined with other parameters. Sha, Sda and S5v had little predictive power in our dataset. Spd seemed rather unrelated to ingesta properties and made this parameter a suitable indicator of masticatory system properties.

Tooth wear as a means to quantify intra-specific variations in diet and chewing movements

In mammals, tooth function, and its efficiency, depends both on the mechanical properties of the food and on chewing dynamics. These aspects have rarely been studied in combination and/or at the intra-specific level. Here we applied 3D dental surface texture analysis to a sample of field voles (Microtus agrestis) trapped from Finnish Lapland at different seasons and localities to test for inter-population variations. We also explored intra-individual variation in chewing dynamics by analysing two facets on the second upper molars. Our results confirm that the two localities have similar environments and that the voles feed on the same items there. On the other hand, the texture data suggest that diets are seasonally variable, probably due to varying concentrations of abrasives. Lastly, the textures on the buccal facets are more isotropic and their direction deviates more from the mesial chewing direction than the lingual facets. We interpret these results as reflecting food, rather than chewing, movements, where food particles are more guided on the lingual side of the molars. This has implications for the application of dental microwear analysis to fossils: only homologous facets can be compared, even when the molar row seems to constitute a functional unit.

Tooth wear patterns in black rats (Rattus rattus) of Madagascar differ more in relation to human impact than to differences in natural habitats

Abstract Dietary characteristics and environmental variables are important selective factors directing ecological diversification in rodents. On Madagascar, the introductions and spread of the commensal black rat (Rattus rattus) can be seen as example cases to study dietary niche occupation and dietary adaptation in an insular environment. We investigate how tooth wear as a measure of dietary adaptation of black rats differs between four distinct habitats (village, manioc fields, spiny forest, and rainforest) with different dietary resources. We use the 3D surface texture analysis (3DST, using 30 parameters according to ISO 25178) as a measure of dietary abrasiveness. 3DST is applied on the occlusal surface of the upper first molar of 37 black rat specimens. The rainforest sample displays less rough and less voluminous surface textures compared to the village samples as indicated by smaller values for height parameters (Sa, Sp, Sq), inverse areal material ratio (Smc), and volume parameters (Vm, Vmc, Vmp, Vv, and Vvc). We therefore rank sampling areas from highest to lowest abrasiveness (village>manioc fields/spiny forest>rainforest). The rats from villages and rainforest differ to such an extent that one could have interpreted them to belong to different species. This indicates a high degree of variability in terms of ingesta abrasiveness. Furthermore, the pronounced difference between rats from human habitations compared to rats from associated fields or natural vegetation is interpreted to clearly indicate shifts in dietary niche occupation in relation to human impact.

Comparative analyses of tooth wear in free-ranging and captive wild equids

REASONS FOR PERFORMING STUDY Captive breeding has played a crucial role in the conservation of threatened equid species. Grazing ruminants and rhinoceros in captivity have less abrasion-dominated tooth wear than their free-ranging conspecifics, with potential negative consequences for their health. However, a similar study on wild equids in captivity is missing. OBJECTIVES The aim was to establish if different tooth wear patterns are exhibited by free-ranging and captive equids. STUDY DESIGN Cross-sectional study of museum specimens comparing free-ranging and captive equids. METHODS Dental casts of maxillary cheek teeth of 228 museum specimens (122 from free-ranging and 106 from captive individuals) of 7 wild equid species were analysed using the extended mesowear method. Although teeth showing specific abnormalities were not scored, the presence of focal overgrowths (hooks) of the rostral premolars (106, 206) was recorded. RESULTS Captive Equus ferus przewalskii, E. grevyi, E. hemionus, E. quagga boehmi and E. zebra hartmannae have less abrasion-dominated tooth wear on their premolars than their free-ranging conspecifics (P<0.001). Fewer differences were exhibited between populations in the molars. No differences were exhibited in the distal cusp of the molars (110, 210) between populations, except in a small sample of E. kiang. Captive equids exhibited more homogeneous wear along the tooth row whereas free-ranging equids exhibited a tooth wear gradient, with more abrasion on premolars than molars. There were more rostral hooks on the premolars (106, 206) in the captive than the free-ranging population (P = 0.02). CONCLUSIONS Captive equids did experience less abrasion-dominated tooth wear than their free-ranging conspecifics, but the differences in tooth wear were less pronounced than those between captive and free-ranging wild ruminant and rhinoceros species. This indicates that feeding regimes for captive equids deviate less from natural diets than those for captive ruminants and rhinoceros but that factors leading to hook formation, in particular feeding height, should receive special attention. The Summary is available in Chinese - see Supporting information.

Functional anatomy of the equine temporomandibular joint: Collagen fiber texture of the articular surfaces

In the last decade, the equine masticatory apparatus has received much attention. Numerous studies have emphasized the importance of the temporomandibular joint (TMJ) in the functional process of mastication. However, ultrastructural and histological data providing a basis for biomechanical and histopathological considerations are not available. The aim of the present study was to analyze the architecture of the collagen fiber apparatus in the articular surfaces of the equine TMJ to reveal typical morphological features indicating biomechanical adaptions. Therefore, the collagen fiber alignment was visualized using the split-line technique in 16 adult warmblood horses without any history of TMJ disorders. Within the central two-thirds of the articular surfaces of the articular tubercle, the articular disc and the mandibular head, split-lines ran in a correspondent rostrocaudal direction. In the lateral and medial aspects of these articular surfaces, the split-line pattern varied, displaying curved arrangements in the articular disc and punctual split-lines in the bony components. Mediolateral orientated split-lines were found in the rostral and caudal border of the articular disc and in the mandibular fossa. The complex movements during the equine chewing cycle are likely assigned to different areas of the TMJ. The split-line pattern of the equine TMJ is indicative of a relative movement of the joint components in a preferential rostrocaudal direction which is consigned to the central aspects of the TMJ. The lateral and medial aspects of the articular surfaces provide split-line patterns that indicate movements particularly around a dorsoventral axis.

The phylogenetic signal in tooth wear: What does it mean?

A new study by Fraser et al (2018) urges the use of phylogenetic comparative methods, whenever possible, in analyses of mammalian tooth wear. We are concerned about this for two reasons. First, this recommendation may mislead the research community into thinking that phylogenetic signal is an artifact of some sort rather than a fundamental outcome of the evolutionary process. Secondly, this recommendation may set a precedent for editors and reviewers to enforce phylogenetic adjustment where it may unnecessarily weaken or even directionally alter the results, shifting the emphasis of analysis from common patterns manifested by large clades to rare cases.

Functional anatomy of the equine temporomandibular joint: Histological characteristics of the articular surfaces and underlining tissues

It has been assumed that dental conditions cause disorders of the equine temporomandibular joint (TMJ), due to biomechanical overload or aberrant loading. However, the incidence of published TMJ disorders in horses is low and this leads to the question whether the equine TMJ is adapted well to its biomechanical requirements or is able to remodel its articular surfaces in response to modified loading conditions. The aim of this study was to determine the histological characteristics of healthy equine TMJs. The tissue components of the articular surfaces of 10 TMJs obtained from horses without any clinical history of dental or TMJ disorders were analysed. Apart from the mandibular fossa of the temporal bone, the osseous aspects of the TMJ exhibited a uniform zoning pattern. The articular surfaces were composed of three tissue layers: (1) a superficial cell-rich dense connective tissue layer; (2) a middle fibrocartilage layer; and (3) a deep hyaline-like cartilage layer. The articular disc was composed of an inner core of fibrocartilage and hyaline-like cartilage meshwork covered with both cell-rich dense connective tissue and fibrocartilage at its dorsal and ventral aspects. In contrast, the mandibular fossa was only covered by a dense connective tissue, frequently supplemented by a synovial membrane, suggesting low biomechanical stress. Glycosaminoglycans, which are indicative of compressive loads, were predominantly present within the rostral part of the articular tubercle and the retroarticular process, the dorsal part of articular disc and the entire mandibular head, but were absent within the mandibular fossa. The results of this study suggest the presence of different biomechanical demands in the dorsal and ventral compartment of the equine TMJ.

Controlled feeding experiments with diets of different abrasiveness reveal slow development of mesowear signal in goats (Capra aegagrus hircus)

ABSTRACT Dental mesowear is applied as a proxy to determine the general diet of mammalian herbivores based on tooth-cusp shape and occlusal relief. Low, blunt cusps are considered typical of grazers and high, sharp cusps typical of browsers. However, how internal or external abrasives impact mesowear, and the time frame the wear signature takes to develop, still need to be explored. Four different pelleted diets of increasing abrasiveness (lucerne, grass, grass and rice husks, and grass, rice husks and sand) were fed to four groups of a total of 28 adult goats in a controlled feeding experiment over a 6-month period. Tooth morphology was captured by medical CT scans at the beginning and end of the experiment. These scans, as well as the crania obtained post mortem, were scored using the mesowear method. Comparisons between diet groups showed few significant differences after 6 months, irrespective of whether CT scans or the real teeth were scored. Only when assessing the difference in signal between the beginning and the end of the experiment did relevant, significant diet-specific effects emerge. Diets containing lower phytolith content caused a more pronounced change in mesowear towards sharper cusps/higher reliefs, while the feed containing sand did not result in more extreme changes in mesowear when compared with the same feed without sand. Our experiment suggests that the formation of a stable and hence reliable mesowear signal requires more time to develop than 6 months. Summary: A controlled feeding experiment in goats shows that the mesowear signal represents a stable dietary period longer than 6 months.

Examination of the interproximal wear mechanism: Facet morphology and surface texture analysis (advance online)

Dentition is considered a dynamic system with forces that directly affect dental treatment stability and success. Understanding the biomechanical forces that influence tooth alignment is essential for both planning and performing dental treatments, as well as for anthropological and evolutionary studies. While there is currently an abundance of research on the mechanics of dental wear at the occlusal surface, the mechanics of interproximal dental wear is largely unexplored. The fretting mechanism, a wear process resulting from small-amplitude cyclic motion of 2 solid contacting surfaces, was refuted as a possible mechanism for occlusal wear but has never been considered for interproximal wear. Therefore, the aim of the current study was to reveal the biomechanical process of the interproximal wear and to explore whether the fretting mechanism could be associated with this process. Premolar teeth with interproximal wear facets were examined by 3-dimensional surface texture analysis using a high-resolution confocal disc-scanning measuring system. The unique texture topography of 3 areas in the proximal surface of each tooth was analyzed by applying 3D dental surface texture analysis. Each area showed unique texture characteristics, presenting statistically significant differences between the inner area of the facet and its margins or the surface outside the facets borders. Based on these results, we concluded that fretting is a key mechanism involved in interproximal wear.