Technical note: An in vitro study of dental microwear formation using the BITE Master II chewing machine

Dental microwear has been used for decades to reconstruct the diets of fossil hominins and bioarchaeological populations. The basic theory has been that hard‐brittle foods (e.g., nuts, bone) require crushing and leave pits as they are pressed between opposing cheek‐tooth surfaces, whereas soft‐tough foods (e.g., grass blades, meat) require shearing and leave scratches as they are dragged along opposing surfaces that slide past one another. However, recent studies have called into question the efficacy of microwear as an indicator of diet. One issue has been the limited number of in vitro studies providing empirical evidence for associations between microwear pattern and chewing behavior. We here describe a new study using a chewing simulator, the BITE Master II, to examine the effects of angle of approach between opposing teeth and food consistency on microwear surface texture. Results indicate that opposing teeth that approach one another: 1) perpendicular to the occlusal plane (crushing) result in pits; 2) parallel to the occlusal plane (shearing) result in striations in the direction of movement; and 3) oblique to the occlusal plane (45°) result in both striations and pits. Results further suggest that different food types and abrasive loads affect the propensity to accumulate microwear features independent of feature shapes. Am J Phys Anthropol 158:769–775, 2015. © 2015 Wiley Periodicals, Inc.     

Telemetry System for Assessing Jaw-Muscle Function in Free-ranging Primates

In vivo laboratory-based studies describing jaw-muscle activity and mandibular bone strain during mastication provide the empirical basis for most evolutionary hypotheses linking primate masticatory apparatus form to diet. However, the laboratory data pose a potential problem for testing predictions of these hypotheses because estimates of masticatory function and performance recorded in the laboratory may lack the appropriate ecological context for understanding adaptation and evolution. For example, in laboratory studies researchers elicit rhythmic chewing using foods that may differ significantly from the diets of wild primates. Because the textural and mechanical properties of foods influence jaw-muscle activity and the resulting strains, chewing behaviors studied in the laboratory may not adequately reflect chewing behaviors of primates feeding in their natural habitats. To circumvent this limitation of laboratory-based studies of primate mastication, we developed a system for recording jaw-muscle electromyograms (EMGs) from free-ranging primates so that researchers can conduct studies of primate jaw-muscle function in vivo in the field. We used the system to record jaw-muscle EMGs from mantled howlers (Alouatta palliata) at Hacienda La Pacifica, Costa Rica. These are the first EMGs recorded from a noncaptive primate feeding in its natural habitat. Further refinements of the system will allow long-term EMG data collection so that researchers can correlate jaw-muscle function with food mechanical properties and behavioral observations. In addition to furthering understanding of primate feeding biology, our work will foster improved adaptive hypotheses explaining the evolution of primate jaw form.     

Effects of the replacement of concentrate and fibre-rich hay by high-quality hay on chewing,rumination and nutrient digestibility in non-lactating Holstein cows

The aim of this study was to investigate the effects of high-quality hay with an elevated sugar content alone or with graded amounts of concentrate feed on chewing and ruminating activity, apparent total tract digestibility (ATTD) and ruminal pH at different time points after feeding in the free ruminal liquid (FRL) and the particle-associated ruminal liquid (PARL). Eight rumen cannulated non-lactating Holstein cows were arranged in a Latin square design in four experimental runs lasting 25 d each. The four diets tested were 60NQ (60% normal-quality hay + 40% concentrate), 60HQ (60% high-quality hay + 40% concentrate), 75HQ (75% high-quality hay + 25% concentrate) and 100HQ (100% high-quality hay). Normal and high-quality hays differed in sugar contents (11.3% vs. 18.7% in dry matter [DM]), neutral detergent fibre (NDF; 57.7% vs. 46.3% in DM), acid detergent fibre (ADF, 35.0% vs. 23.5% in DM) and crude protein (CP, 11.3% vs. 23.5% in DM). Data showed that ATTD of DM, CP, NDF and ADF was higher with the high-quality hay diets. Time spent eating was reduced with high-quality hay. However, time spent ruminating was longest in Group 100HQ. In all groups, ruminal pH of FRL and PARL decreased with time after the morning feeding. But 10 h later, pH of Group 100HQ was higher again compared with the other groups. Considering the average pH in FRL over all measured time points, cows in Groups 60NQ and 100HQ had higher pH values of 6.85 and 6.83, respectively. Regarding pH values in PARL, animals of Group 60NQ displayed the highest pH value (6.68), whereas the lowest value of 6.21 was found in Group 60HQ. Overall, results suggest that high-quality hay maintains the diet’s structural effectiveness by stimulating rumination and stabilising ruminal pH while greatly improving ATTD. However, the structural effectiveness of the high-quality hay gets impaired with increasing proportion of concentrate feed in the diet.     

Characteristics of boli formed by dairy cows upon ingestion of fresh ryegrass,lucerne or chicory

This study examined the comminution of fresh herbage, subsequent nutrient release, and the characteristics of swallowed boli from three physically and chemically contrasting forages during ingestive mastication by dairy cows. The extent and pattern of nutrient release will determine their availability to rumen microflora, and potentially influence their efficiency of use. The forages evaluated were perennial ryegrass (ryegrass, Lolium perenne L., cv Alto AR37), lucerne (Medicago sativa L., cv Torlesse) and chicory (Cichorium intybus L., cv Choice). Experimental design was a 3×3 cross-over with three forages and three consecutive 1-day measurement periods, conducted twice. Six non-lactating, pregnant, multiparous Holstein-Friesian×Jersey cows (Bos taurus) were used, with the first cross-over applied to three mature (10.1±0.61 years old; BW 631±64 kg) cows, and the second to three young (4.8±0.02 years; BW 505±19 kg) cows. Fresh cut forage was offered to the cows following partial rumen evacuation. Swallowed boli were collected directly at the cardia at the commencement, middle and end of the first feeding bout of the first meal of the day. Forage species did not affect the fresh weight of ingested boli (mean 169 g, P=0.605) but the proportion of saliva in boli varied between forage. Boli of chicory contained the greatest amount of herbage material and least amount of saliva, whereas ryegrass boli were the opposite. Boli fresh weight tended to increase as time in the meal progressed, but the age of the cow was not shown to affect any boli characteristics or nutrient release. Particle size reduction was affected by forage, with 31%, 38% and 35% of chicory, lucerne and ryegrass herbage reduced to <2 mm. There was little evidence of relationship between comminution and any physical or chemical characteristic of the forage, except in ryegrass where extent of comminution was moderately correlated with herbage strength. Proportional release of herbage soluble carbohydrate exceeded that of N during mastication. Differences in loss of N were moderately correlated with the amount of N in the herbage (R²=0.53) but herbage comminution was not strongly correlated with release of either N or carbohydrate. These findings illustrate the complex animal×forage interactions that occur during mastication, and that it is not possible to infer nutrient loss from herbage based on herbage characteristics as the driver for this differ between species.     

Experimental stress analysis of topographic diversity in early hominid gnathic morphology

Reconstructing the biomechanics of early hominid mastication is a key element in most models of hominid differentiation. Traditionally, ostelogical features marking muscle attachment surfaces have served as a reference system from which the vector geometry of the masticatory force system and resultant force distributions could be predicted. To augment traditional morphological and computational approaches, we developed a simulation system capable of replicating human and non-human primate chewing motions. The forces of occlusion are recorded as photoelastic fringes in a urethane alveolar process. Simulation experiments evaluating the functional correlates of topographic diversity in zygomatic root position and mandibular ramus height in early hominids indicated that the mandibles and dentitions of robust australopithecines are well adapted to sustain high magnitude, low gradient load distributions.     

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.     

Changes in orientation of attritional wear facets with implications for jaw motion in a mixed longitudinal sample of Propithecus edwardsi from Ranomafana National Park, Madagascar

In many mammalian species, the progressive wearing down of the teeth that occurs over an individual's lifetime has the potential to change dental function, jaw movements, or even feeding habits. The orientation of phase-I wear facets on molars reveals the direction of jaw movement during the power stroke of mastication. We investigated if and how molar wear facets change with increasing wear and/or age by examining a mixed longitudinal dataset of mandibular tooth molds from wild Propithecus edwardsi (N = 32 individuals, 86 samples). Measurements of the verticality of wear facets were obtained from three-dimensional digital models generated from μCT scans. Results show that verticality decreases over the lifetime of P. edwardsi, a change that implies an increasingly lateral translation of the jaw as the teeth move into occlusion. A more transverse phase-I power stroke supports the hypothesis that these animals chew to maximize longevity and functionality of their teeth, minimizing the "waste" of enamel, while maintaining sharp shearing crests. Results of this study indicate that wear facet verticality is more closely correlated with age than overall amount of tooth wear, measured as area of exposed dentin, suggesting that age-related changes in cranial morphology may be more responsible for adjustments in jaw motion over the lifetimes of Propithecus than wear-related changes inthe shape of occluding teeth. Finally, the rate of decrease in wear facet verticality with age is greater in males than in females suggesting differences in development and/or access to resources between the sexes in this species.     

Functional and evolutionary significance of the recruitment and firing patterns of the jaw adductors during chewing in Verreaux's sifaka (Propithecus verreauxi)

Jaw-muscle electromyographic (EMG) patterns indicate that compared with thick-tailed galagos and ring-tailed lemurs, anthropoids recruit more relative EMG from their balancing-side deep masseter, and that this muscle peaks late in the power stroke. These recruitment and firing patterns in anthropoids are thought to cause the mandibular symphysis to wishbone (lateral transverse bending), resulting in relatively high symphyseal stresses. We test the hypothesis that living strepsirrhines with robust, partially fused symphyses have muscle recruitment and firing patterns more similar to anthropoids, unlike those strepsirrhines with highly mobile unfused symphyses. Electromyographic (EMG) activity of the superficial and deep masseter, anterior and posterior temporalis, and medial pterygoid muscles were recorded in four dentally adult Verreaux's sifakas (Propithecus verreauxi). As predicted, we find that sifaka motor patterns are more similar to anthropoids. For example, among sifakas, recruitment levels of the balancing-side (b-s) deep masseter are high, and the b-s deep masseter fires late during the power stroke. As adult sifakas often exhibit nearly complete symphyseal fusion, these data support the hypothesis that the evolution of symphyseal fusion in primates is functionally linked to wishboning. Furthermore, these data provide compelling evidence for the convergent evolution of the wishboning motor patterns in anthropoids and sifakas.     

Technical note: Dental microwear textures of "Phase I" and "Phase II" facets

The power stroke of mastication has been traditionally divided into two parts, one which precedes centric occlusion, and the other which follows it-"Phase I" and "Phase II," respectively. Recent studies of primate mastication have called into question the role of Phase II in food processing, as they have found little muscle activity or accompanying bone strain following centric occlusion. That said, many researchers today look to Phase II facets to relate diet to patterns of dental microwear. This suggests the need to reevaluate microwear patterns on Phase I facets. Here we use texture analysis to compare and contrast microwear on facets representing both phases in three primate species with differing diets (Alouatta palliata, Cebus apella, and Lophocebus albigena). Results reaffirm that microwear patterns on Phase II facets better distinguish taxa with differing diets than do those on Phase I facets. Further, differences in microwear textures between facet types for a given taxon may themselves reflect diet. Some possible explanations for differences in microwear textures between facet types are proposed.