Primary sensorimotor cortex exhibits complex dependencies of spike-field coherence on neuronal firing rates, field power, and behavior.

Spike-field coherence (SFC) is widely used to assess cortico-cortical interactions during sensorimotor behavioral tasks by measuring the consistency of the relative phases between the spike train of a neuron and the concurrent local field potentials (LFPs). Interpretations of SFC as a measure of functional connectivity are complicated by theoretical work suggesting that estimates of SFC depend on overall neuronal activity. We evaluated the dependence of SFC on neuronal firing rates, LFP power, and behavior in the primary motor (MIo) and primary somatosensory (SIo) areas of the orofacial sensorimotor cortex of monkeys ( Macaca mulatta) during performance of a tongue-protrusion task. Although we occasionally observed monotonically increasing linear relationships between coherence and firing rate, we most often found highly complex, nonmonotonic relationships in both SIo and MIo and sometimes even found that coherence decreased with increasing firing rate. The lack of linear relationships was also true for both LFP power and tongue-protrusive force. Moreover, the ratio between maximal firing rate and the firing rate at peak coherence deviated significantly from unity, indicating that MIo and SIo neurons achieved maximal SFC at a submaximal level of spiking. Overall, these results point to complex relationships of SFC to firing rates, LFP power, and behavior during sensorimotor cortico-cortical interactions: coherence is a measure of functional connectivity whose magnitude is not a mere monotonic reflection of changes in firing rate, LFP power, or the relevantly controlled behavioral parameter. NEW & NOTEWORTHY The concern that estimates of spike-field coherence depend on the firing rates of single neurons has influenced analytical methods employed by experimental studies investigating the functional interactions between cortical areas. Our study shows that the overwhelming majority of the estimated spike-field coherence exhibited complex relations with firing rates of neurons in the orofacial sensorimotor cortex. The lack of monotonic relations was also evident after testing the influence of local field potential power and force on spike-field coherence.

Bone strain magnitude is correlated with bone strain rate in tetrapods: implications for models of mechanotransduction.

Hypotheses suggest that structural integrity of vertebrate bones is maintained by controlling bone strain magnitude via adaptive modelling in response to mechanical stimuli. Increased tissue-level strain magnitude and rate have both been identified as potent stimuli leading to increased bone formation. Mechanotransduction models hypothesize that osteocytes sense bone deformation by detecting fluid flow-induced drag in the bone's lacunar-canalicular porosity. This model suggests that the osteocyte's intracellular response depends on fluid-flow rate, a product of bone strain rate and gradient, but does not provide a mechanism for detection of strain magnitude. Such a mechanism is necessary for bone modelling to adapt to loads, because strain magnitude is an important determinant of skeletal fracture. Using strain gauge data from the limb bones of amphibians, reptiles, birds and mammals, we identified strong correlations between strain rate and magnitude across clades employing diverse locomotor styles and degrees of rhythmicity. The breadth of our sample suggests that this pattern is likely to be a common feature of tetrapod bone loading. Moreover, finding that bone strain magnitude is encoded in strain rate at the tissue level is consistent with the hypothesis that it might be encoded in fluid-flow rate at the cellular level, facilitating bone adaptation via mechanotransduction.

Directional information from neuronal ensembles in the primate orofacial sensorimotor cortex.

Neurons in the arm and orofacial regions of the sensorimotor cortex in behaving monkeys display directional tuning of their activity during arm reaching and tongue protrusion, respectively. While studies on population activity abound for the arm motor cortex, how populations of neurons from the orofacial sensorimotor cortex represent direction has never been described. We therefore examined and compared the directional information contained in the spiking activity of populations of single neurons recorded simultaneously from chronically implanted microelectrode arrays in the orofacial primary motor (MIo, N = 345) and somatosensory (SIo, N = 336) cortices of monkeys (Macaca mulatta) as they protruded their tongue in different directions. Differential modulation to the direction of tongue protrusion was found in >60% of task-modulated neurons in MIo and SIo and was stronger in SIo (P < 0.05). Moreover, mutual information between direction and spiking was significantly higher in SIo compared with MIo at force onset and force offset (P < 0.01). Finally, the direction of tongue protrusion was accurately predicted on a trial-by-trial basis from the spiking activity of populations of MIo or SIo neurons by using a discrete decoder (P < 0.01). The highly reliable decoding was comparable between MIo and SIo neurons. However, the temporal evolution of the decoding performance differed between these two areas: MIo showed late-onset, fast-rising, and phasic performance, whereas SIo exhibited early-onset, slow-rising, and sustained performance. Overall, the results suggest that both MIo and SIo are highly involved in representing the direction of tongue protrusion but they differ in the amplitude and temporal processing of the directional information distributed across populations of neurons.

Preference mapping of frozen and fresh raspberries.

The purpose of the study was to identify key sensory attributes that influence consumer liking for frozen and fresh red raspberries using preference mapping. Sensory profiling of different raspberry cultivars and selections from the Washington State Univ. and Oregon State Univ. breeding programs was carried out using a trained panel (frozen, n = 12 and fresh, n = 10). In addition, a subset of frozen and fresh raspberries of each cultivar was assessed by consumers for sensory acceptability (n = 105 and n = 100, respectively). Based on overall hedonic ratings, cluster analysis identified 3 clusters of frozen raspberry consumers from day 1 (41% "nondistinguishers," 34% "likers," and 25% "nonlikers") and day 2 (41% "group 1 likers," 26% "nonlikers," and 34% and 33% group 2 likers"). For fresh raspberry consumers, 2 clusters were detected from day 1 (54% "likers" and 46% nondistinguishers") and day 2 (54% "group 1 likers" and 46% "group 2 likers"). Preference mapping was applied on the descriptive sensory and acceptability of clustered consumer data. Partial least squares regression results showed that liking of frozen raspberries was driven by high raspberry flavor, firmness, and sweetness. Conversely, disliking of frozen raspberries was related to high sour and aftertaste intensity. In the case of fresh raspberries, high color uniformity, raspberry aroma, raspberry flavor, floral aroma, green flavor, bitter, astringency, and aftertaste increased the acceptability, whereas high color intensity and green aroma were associated with negative drivers of liking. The information obtained in this study can be a useful guide for breeders in the selection of characteristics for growing superior quality raspberries.

Short communication: low-fat ice cream flavor not modified by high hydrostatic pressure treatment of whey protein concentrate.

The purpose of this study was to examine flavor binding of high hydrostatic pressure (HHP)-treated whey protein concentrate (WPC) in a real food system. Fresh Washington State University (WSU, Pullman) WPC, produced by ultrafiltration of separated Cheddar cheese whey, was treated at 300 MPa for 15 min. Commercial WPC 35 powder was reconstituted to equivalent total solids as WSU WPC (8.23%). Six batches of low-fat ice cream were produced: A) HHP-treated WSU WPC without diacetyl; B) and E) WSU WPC with 2 mg/L of diacetyl added before HHP; C) WSU WPC with 2 mg/L of diacetyl added after HHP; D) untreated WSU WPC with 2 mg/L of diacetyl; and F) untreated commercial WPC 35 with 2 mg/L of diacetyl. The solution of WSU WPC or commercial WPC 35 contributed 10% to the mix formulation. Ice creams were produced by using standard ice cream ingredients and processes. Low-fat ice creams containing HHP-treated WSU WPC and untreated WSU WPC were analyzed using headspace-solid phase microextraction-gas chromatography. Sensory evaluation by balanced reference duo-trio test was carried out using 50 untrained panelists in 2 sessions on 2 different days. The headspace-solid phase microextraction-gas chromatography analysis revealed that ice cream containing HHP-treated WSU WPC had almost 3 times the concentration of diacetyl compared with ice cream containing untreated WSU WPC at d 1 of storage. However, diacetyl was not detected in ice creams after 14 d of storage. Eighty percent of panelists were able to distinguish between low-fat ice creams containing untreated WSU WPC with and without diacetyl, confirming panelists' ability to detect diacetyl. However, panelists were not able to distinguish between low-fat ice creams containing untreated and HHP-treated WSU WPC with diacetyl. These results show that WPC diacetyl-binding properties were not enhanced by 300-MPa HHP treatment for 15 min, indicating that HHP may not be suitable for such applications.

Chewing variation in lepidosaurs and primates.

Mammals chew more rhythmically than lepidosaurs. The research presented here evaluated possible reasons for this difference in relation to differences between lepidosaurs and mammals in sensorimotor systems. Variance in the absolute and relative durations of the phases of the gape cycle was calculated from kinematic data from four species of primates and eight species of lepidosaurs. The primates exhibit less variance in the duration of the gape cycle than in the durations of the four phases making up the gape cycle. This suggests that increases in the durations of some gape cycle phases are accompanied by decreases in others. Similar effects are much less pronounced in the lepidosaurs. In addition, the primates show isometric changes in gape cycle phase durations, i.e. the relative durations of the phases of the gape cycle change little with increasing cycle time. In contrast, in the lepidosaurs variance in total gape cycle duration is associated with increases in the proportion of the cycle made up by the slow open phase. We hypothesize that in mammals the central nervous system includes a representation of the optimal chew cycle duration maintained using afferent feedback about the ongoing state of the chew cycle. The differences between lepidosaurs and primates do not lie in the nature of the sensory information collected and its feedback to the feeding system, but rather the processing of that information by the CNS and its use feed-forward for modulating jaw movements and gape cycle phase durations during chewing.

High hydrostatic pressure modification of whey protein concentrate for use in low-fat whipping cream improves foaming properties.

Whey is the inevitable by-product of cheese production. Whey can be incorporated into a variety of foods, but little has been done to investigate its suitability in whipping cream. The objective of this work was to evaluate the foaming properties of selected low-fat whipping cream formulations containing whey protein concentrate (WPC) that did or did not undergo high hydrostatic pressure (HHP) treatment. Fresh whey was concentrated by ultrafiltration, pasteurized, and standardized to 8.23% total solids and treated with HHP at 300 MPa for 15 min. Viscosity, overrun, and foam stability were determined to assess foaming properties. Sensory evaluation was conducted with 57 panelists using a duo-trio difference test. The optimal whipping time for the selected formulations was 3 min. Whipping cream containing untreated WPC and HHP-treated WPC resulted in greater overrun and foam stability than the control whipping cream without WPC. Panelists distinguished a difference between whipping cream containing untreated WPC and whipping cream containing HHP-treated WPC. High hydrostatic pressure-treated WPC can improve the foaming properties of low-fat whipping cream, which may justify expansion of the use of whey in whipping cream and application of HHP technology in the dairy industry.

High hydrostatic pressure modification of whey protein concentrate for improved body and texture of lowfat ice cream.

Previous research demonstrated that application of high hydrostatic pressure (HHP), particularly at 300 MPa for 15 min, can enhance foaming properties of whey protein concentrate (WPC). The purpose of this research was to determine the practical impact of HHP-treated WPC on the body and texture of lowfat ice cream. Washington State University (WSU)-WPC was produced by ultrafiltration of fresh separated whey received from the WSU creamery. Commercial whey protein concentrate 35 (WPC 35) powder was reconstituted to equivalent total solids as WSU-WPC (8.23%). Three batches of lowfat ice cream mix were produced to contain WSU-WPC without HHP, WSU-WPC with HHP (300 MPa for 15 min), and WPC 35 without HHP. All lowfat ice cream mixes contained 10% WSU-WPC or WPC 35. Overrun and foam stability of ice cream mixes were determined after whipping for 15 min. Ice creams were produced using standard ice cream ingredients and processing. The hardness of ice creams was determined with a TA-XT2 texture analyzer. Sensory evaluation by balanced reference duo-trio test was carried out using 52 volunteers. The ice cream mix containing HHP-treated WSU-WPC exhibited the greatest overrun and foam stability, confirming the effect of HHP on foaming properties of whey proteins in a complex system. Ice cream containing HHP-treated WSU-WPC exhibited significantly greater hardness than ice cream produced with untreated WSU-WPC or WPC 35. Panelists were able to distinguish between ice cream containing HHP-treated WSU-WPC and ice cream containing untreated WPC 35. Improvements of overrun and foam stability were observed when HHP-treated whey protein was used at a concentration as low as 10% (wt/wt) in ice cream mix. The impact of HHP on the functional properties of whey proteins was more pronounced than the impact on sensory properties.

Effects of supplemental fat on growth performance and quality of beef from steers fed corn finishing diets.

To measure the effects of dietary fat on feedlot performance and carcass characteristics, and on beef appearance, moisture binding, shelf life, palatability, and fatty acid content, 126 crossbred beef steers (321.1 +/- 0.57 kg of BW) were allotted to a randomized complete block (3) design with a 3 x 2 + 1 factorial arrangement of dietary treatments. The main effects were level of yellow grease (0, 3, or 6%) and alfalfa hay (3.5 or 7%) in corn-based diets containing 15% potato by-product (PB). The added treatment was 6% tallow and 7% alfalfa in a barley-based diet containing 15% PB. Dry matter intake and ADG were not affected by diet; however, G:F and diet NE content increased linearly (P < 0.10) with yellow grease. Kidney, pelvic, and heart fat (2.0 to 2.3 +/- 0.07) and yield grade (2.8 to 3.1 +/- 0.09) increased linearly (P < or = 0.05) with yellow grease. Steers fed corn plus 6% yellow grease had lower (P < 0.05) beef firmness and beef texture scores but greater (P < 0.01) fat color score than those fed barley plus 6% tallow. Moisture retention of beef was not affected by dietary treatment, except purge score during retail storage, which was decreased linearly (P < 0.01) from 2.1 to 1.6 +/- 0.06 by level of yellow grease. Steaks from steers fed barley plus 6% tallow had greater (P < 0.05) shear force than those from steers fed corn plus 6% yellow grease, and beef flavor increased linearly (P < 0.05) from 6.2 to 6.7 +/- 0.11 as the level of yellow grease increased. Level of yellow grease linearly increased (P < 0.01) transvaccenic acid (TVA) by 61% and CLA content of beef by 48%. Beef from steers fed corn plus yellow grease had lower (P < 0.05) palmitoleic and oleic acids and greater (P < 0.05) linoleic, TVA, and CLA than beef from steers fed the barley-tallow diet. Feeding yellow grease increased diet energy content, which increased carcass fatness, and altered beef fatty acid content, which increased beef flavor without affecting moisture retention, shelf life, or cooking properties of the beef. Additionally, beef from steers fed corn plus 6% yellow grease was more tender and had more polyunsaturated fatty acid content and CLA than beef from steers fed barley plus 6% tallow.

Influence of visual masking technique on the assessment of 2 red wines by trained and consumer assessors.

During sensory evaluation assessments, visual masking techniques are frequently employed to disguise color differences between samples and minimize perceptual bias. Particularly in wine, the impact of these masking techniques on panelist evaluations has not been well studied. The objective of this study was to study the influence of visual masking techniques on the aroma and flavor assessment of 2 red wines and observe the impact of these techniques on trained and consumer sensory panels. Specific masking techniques included (1) blue wine glass/white illumination; (2) clear glass/red illumination; and (3) clear glass/white illumination. Ten panelists were trained to recognize 7 aroma and flavor attributes, while consumer panelists (n=80) evaluated attributes and liking. For the trained panel, the visual masking technique affected only perceived spicy flavor of Syrah (P < or = 0.05), with the clear glass/red illumination resulting in more intense spicy flavor compared to the other 2 conditions. Principal components analysis showed that for the 2 red wines evaluated by the trained panel, red illumination resulted in higher spicy attributes and perceived astringency while wines served in blue wine glasses were higher in perceived astringency. For the consumer panel, red illumination resulted in wines higher in perceived astringency and blue wine glasses resulted in wines higher in perceived flavor liking. These results indicated that the visual masking techniques may influence both trained and consumer panel evaluation of aroma and flavor attributes of red wine. However, beyond red wine, this study makes the larger point that the choice of masking technique does impact sensory evaluations.

In vivo function of the craniofacial haft: the interorbital "pillar".

The craniofacial haft resists forces generated in the face during feeding, but the importance of these forces for the form of the craniofacial haft remains to be determined. In vivo bone strain data were recorded from the medial orbital wall in an owl monkey (Aotus), rhesus macaques (Macaca mulatta), and a galago (Otolemur) during feeding. These data were used to determine whether: the interorbital region can be modeled as a simple beam under bending or shear; the face is twisting on the brain case during unilateral biting or mastication; the interorbital "pillar" is being axially compressed during incisor loading and both axially compressed and laterally bent during mastication; and the interorbital "pillar" transmits axial compressive forces from the toothrow to the braincase. The strain data reveal that the interorbital region cannot be modeled as a anteroposteriorly oriented beam bent superiorly in the sagittal plane during incision or mastication. The strain orientations recorded in the majority of experiments are concordant with those predicted for a short beam under shear, although the anthropoids displayed evidence of multiple loading regimes in the medial orbital wall. Strain orientation data corroborate the hypothesis that the strepsirrhine face is twisted during mastication. The hypothesis that the interorbital region is a member in a rigid frame subjected to axial compression during mastication receives some support. The hypothesis that the interorbital region is a member in a rigid frame subjected to lateral bending during mastication is supported by the epsilon1/absolute value epsilon2 ratio data but not by the strain orientation data. The timing of peak shear strains in the medial orbital wall of anthropoids does not bear a consistent relationship to the timing of peak shear strain in the mandibular corpus, suggesting that bite force is not the only external force influencing the medial orbital wall. Strain orientation data suggest the existence of two distinct loading regimes, possibly associated with masseter or medial pterygoid contraction. Regardless of the loading regime, all taxa showed low strain magnitudes in the medial orbital wall relative to the anterior root of the zygoma and the mandibular corpus. The strain gradients documented here and elsewhere suggest that, in anthropoids at least, local effects of external forces are more important than a single global loading regime. The low strain magnitudes in the medial orbital wall and in other thin bony plates around the orbit suggest that these structures are not optimally designed for resisting feeding forces. It is hypothesized that their function is to provide rigid support and protection for soft-tissue structures such as the nasal epithelium, the brain, meninges, and the eye and its adnexa. In contrast with the face of Otolemur, which appears to be subjected to a single predominant loading regime, anthropoids may experience different loading regimes in different parts of the face. This implies that the anthropoid and strepsirrhine facial skulls might be optimized for different functions.

Evolution of activity patterns and chromatic vision in primates: morphometrics, genetics and cladistics.

Hypotheses for the adaptive origin of primates have reconstructed nocturnality as the primitive activity pattern for the entire order based on functional/adaptive interpretations of the relative size and orientation of the orbits, body size and dietary reconstruction. Based on comparative data from extant taxa this reconstruction implies that basal primates were also solitary, faunivorous, and arboreal. Recently, primates have been hypothesized to be primitively diurnal, based in part on the distribution of color-sensitive photoreceptor opsin genes and active trichromatic color vision in several extant strepsirrhines, as well as anthropoid primates (Tan & Li, 1999 Nature402, 36; Li, 2000 Am. J. phys. Anthrop. Supple.30, 318). If diurnality is primitive for all primates then the functional and adaptive significance of aspects of strepsirrhine retinal morphology and other adaptations of the primate visual system such as high acuity stereopsis, have been misinterpreted for decades. This hypothesis also implies that nocturnality evolved numerous times in primates. However, the hypothesis that primates are primitively diurnal has not been analyzed in a phylogenetic context, nor have the activity patterns of several fossil primates been considered. This study investigated the evolution of activity patterns and trichromacy in primates using a new method for reconstructing activity patterns in fragmentary fossils and by reconstructing visual system character evolution at key ancestral nodes of primate higher taxa. Results support previous studies that reconstruct omomyiform primates as nocturnal. The larger body sizes of adapiform primates confound inferences regarding activity pattern evolution in this group. The hypothesis of diurnality and trichromacy as primitive for primates is not supported by the phylogenetic data. On the contrary, nocturnality and dichromatic vision are not only primitive for all primates, but also for extant strepsirrhines. Diurnality, and possibly X-linked polymorphic trichromacy, evolved at least in the stem lineage of Anthropoidea, or the stem lineage of all haplorhines.

The primate cranial base: ontogeny, function, and integration.

Understanding the complexities of cranial base development, function, and architecture is important for testing hypotheses about many aspects of craniofacial variation and evolution. We summarize key aspects of cranial base growth and development in primates that are useful for formulating and testing hypotheses about the roles of the chondrocranium and basicranium in cranial growth, integration, and function in primate and human evolution. We review interspecific, experimental, and ontogenetic evidence for interactions between the cranial base and brain, and between the cranial base and the face. These interactions indicate that the cranial base plays a key role in craniofacial growth, helping to integrate, spatially and functionally, different patterns of growth in various adjoining regions of the skull such as components of the brain, the eyes, the nasal cavity, the oral cavity, and the pharynx. Brain size relative to cranial base length appears to be the dominant influence on many aspects of basicranial variation, especially the angle of the cranial base in the midsagittal plane, but other factors such as facial size, facial orientation, and posture may also be important. Major changes in cranial base shape appear to have played crucial roles in the evolution of early primates, the origin of anthropoids, and the origin of Homo sapiens.

The petrosal of Omomys carteri and the evolution of the primate basicranium.

The first omomyine petrosals, those of Omomys carteri, are described. Omomys probably had a tympanic bulla and canals for the intratympanic carotid circulation derived from the petrosal bone. The stapedial and promontory canals were complete, large and subequal. The posterior carotid foramen entered the bulla posteromedially. The intratympanic portion of the facial nerve was fully enclosed in bone, the stapedius fossa is extrabullar and the parotic fissure is patent. The mastoid was pneumatized from the epitympanic recess and a supracochlear cavity may have been present. The Omomys petrosals exhibit a generic omomyiform morphology, exhibiting no features that can be interpreted as autapomorphies and only one feature shared with adapiforms. The monophyly of Omomyiformes is based on other cranial characters, dental and postcranial characters assessed elsewhere. The similarity of the Shanghuang petrosal to the petrosals of omomyiforms, as well as the ambiguous evidence of its association, suggest that an omomyiform affinity for that petrosal cannot be ruled out.

Electromyography of the anterior temporalis and masseter muscles of owl monkeys (Aotus trivirgatus) and the function of the postorbital septum.

Anthropoids and tarsiers are distinguished from all other vertebrates by the possession of a postorbital septum, which is formed by the frontal, alisphenoid, and zygomatic bones. Cartmill [(1980) In: Evolutionary Biology of the New World Monkeys and Continental Drift. New York: Plenum, p 243-274] suggested that the postorbital septum evolved in the stem lineage of tarsiers and anthropoids to insulate the eye from movements arising in the temporal fossa. Ross [(1996) Am J Phys Anthropol 91:305-324] suggested that the septum insulates the orbital contents from incursions by the line of action of the anterior temporal muscles caused by the unique combination of high degrees of orbital frontation and convergence. Both of these hypotheses must explain why insulation of the orbital contents could not be achieved by decreasing the size of the anterior temporal musculature with a corresponding increase in size of the remaining jaw adductors, rather than evolving a postorbital septum. One possibility is that the anterior temporalis is an important contributor to vertically directed bite forces during all biting and chewing activities. Another possibility is that reduction in anterior temporal musculature would compromise the ability to produce powerful bite forces, either at the incisors or along the postcanine toothrow. To evaluate these hypotheses, electromyographic (EMG) recordings were made from the masseter muscle and the anterior and posterior portions of the temporalis muscles of two owl monkeys, Aotus trivirgatus. The EMG data indicate that anterior temporalis activity relative to that of the superficial masseter is lower during incision than mastication. In addition, activity of the anterior temporalis is not consistently higher than the posterior temporalis during incision. The data indicate relatively greater activity of anterior temporalis compared to other muscles during isometric biting on the postcanine toothrow. This may be due to decreased activity in superficial masseter and posterior temporalis, rather than elevated anterior temporalis activity. The anterior temporalis is not consistently less variable in activity than the superficial masseter and posterior temporalis. The EMG data gathered here indicate no reason for suggesting that the anterior temporal muscles in anthropoids are utilized especially for incisal preparation of hard fruits. Maintenance of relatively high EMG activity in anterior temporalis across a wide range of biting behaviors is to be expected in a vertically oriented and rostrally positioned muscle such as this because, compared to the posterior temporalis, superficial masseter and medial pterygoid, it can contribute relatively larger vertical components of force to bites along the postcanine toothrow. The in vivo data do not support this hypothesis, possibly because of effects of bite point and bite force orientation.

Symphyseal fusion and jaw-adductor muscle force: an EMG study.

The purpose of this study is to test various hypotheses about balancing-side jaw muscle recruitment patterns during mastication, with a major focus on testing the hypothesis that symphyseal fusion in anthropoids is due mainly to vertically- and/or transversely-directed jaw muscle forces. Furthermore, as the balancing-side deep masseter has been shown to play an important role in wishboning of the macaque mandibular symphysis, we test the hypothesis that primates possessing a highly mobile mandibular symphysis do not exhibit the balancing-side deep masseter firing pattern that causes wishboning of the anthropoid mandible. Finally, we also test the hypothesis that balancing-side muscle recruitment patterns are importantly related to allometric constraints associated with the evolution of increasing body size. Electromyographic (EMG) activity of the left and right superficial and deep masseters were recorded and analyzed in baboons, macaques, owl monkeys, and thick-tailed galagos. The masseter was chosen for analysis because in the frontal projection its superficial portion exerts force primarily in the vertical (dorsoventral) direction, whereas its deep portion has a relatively larger component of force in the transverse direction. The symphyseal fusion-muscle recruitment hypothesis predicts that unlike anthropoids, galagos develop bite force with relatively little contribution from their balancing-side jaw muscles. Thus, compared to galagos, anthropoids recruit a larger percentage of force from their balancing-side muscles. If true, this means that during forceful mastication, galagos should have working-side/balancing-side (W/B) EMG ratios that are relatively large, whereas anthropoids should have W/B ratios that are relatively small. The EMG data indicate that galagos do indeed have the largest average W/B ratios for both the superficial and deep masseters (2.2 and 4.4, respectively). Among the anthropoids, the average W/B ratios for the superficial and deep masseters are 1.9 and 1.0 for baboons, 1.4 and 1.0 for macaques, and both values are 1.4 for owl monkeys. Of these ratios, however, the only significant difference between thick-tailed galagos and anthropoids are those associated with the deep masseter. Furthermore, the analysis of masseter firing patterns indicates that whereas baboons, macaques and owl monkeys exhibit the deep masseter firing pattern associated with wishboning of the macaque mandibular symphysis, galagos do not exhibit this firing pattern. The allometric constraint-muscle recruitment hypothesis predicts that larger primates must recruit relatively larger amounts of balancing-side muscle force so as to develop equivalent amounts of bite force. Operationally this means that during forceful mastication, the W/B EMG ratios for the superficial and deep masseters should be negatively correlated with body size. Our analysis clearly refutes this hypothesis. As already noted, the average W/B ratios for both the superficial and deep masseter are largest in thick-tailed galagos, and not, as predicted by the allometric constraint hypothesis, in owl monkeys, an anthropoid whose body size is smaller than that of thick-tailed galagos. Our analysis also indicates that owl monkeys have W/B ratios that are small and more similar to those of the much larger-sized baboons and macaques. Thus, both the analysis of the W/B EMG ratios and the muscle firing pattern data support the hypothesis that symphyseal fusion and transversely-directed muscle force in anthropoids are functionally linked. This in turn supports the hypothesis that the evolution of symphyseal fusion in anthropoids is an adaptation to strengthen the symphysis so as to counter increased wishboning stress during forceful unilateral mastication. (ABSTRACT TRUNCATED)

Kinematic data on primate head and neck posture: implications for the evolution of basicranial flexion and an evaluation of registration planes used in paleoanthropology.

Kinematic data on primate head and neck posture were collected by filming 29 primate species during locomotion. These were used to test whether head and neck posture are significant influences on basicranial flexion and whether the Frankfurt plane can legitimately be employed in paleoanthropological studies. Three kinematic measurements were recorded as angles relative to the gravity vector, the inclination of the orbital plane, the inclination of the neck, and the inclination of the Frankfurt plane. A fourth kinematic measurement was calculated as the angle between the neck and the orbital plane (the head-neck angle [HNA]). The functional relationships of basicranial flexion were examined by calculating the correlations and partial correlations between HNA and craniometric measurements representing basicranial flexion, orbital kyphosis, and relative brain size (Ross and Ravosa [1993] Am. J. Phys. Anthropol. 91:305-324). Significant partial correlations were observed between relative brain size and basicranial flexion and between HNA and orbital kyphosis. This indicates that brain size, rather than head and neck posture, is the primary influence on flexion, while the degree of orbital kyphosis may act to reorient the visual field in response to variation in head and neck posture. Regarding registration planes, the Frankfurt plane was found to be horizontal in humans but inclined in all nonhuman primates. In contrast, nearly all primates (including humans) oriented their orbits such that they faced anteriorly and slightly inferiorly. These results suggest that for certain functional craniometric studies, the orbital plane may be a more suitable registration plane than Frankfurt "Horizontal."

Mandibular corpus strain in primates: further evidence for a functional link between symphyseal fusion and jaw-adductor muscle force.

Previous work indicates that compared to adult thick-tailed galagos, adult long-tailed macaques have much more bone strain on the balancing-side mandibular corpus during unilateral isometric molar biting (Hylander [1979a] J. Morphol. 159:253-296). Recently we have confirmed in these same two species the presence of similar differences in bone-strain patterns during forceful mastication. Moreover, we have also recorded mandibular bone strain patterns in adult owl monkeys, which are slightly smaller than the galago subjects. The owl monkey data indicate the presence of a strain pattern very similar to that recorded for macaques, and quite unlike that recorded for galagos. We interpret these bone-strain pattern differences to be importantly related to differences in balancing-side jaw-adductor muscle force recruitment patterns. That is, compared to galagos, macaques and owl monkeys recruit relatively more balancing-side jaw-adductor muscle force during forceful mastication. Unlike an earlier study (Hylander [1979b] J. Morphol. 160:223-240), we are unable to estimate the actual amount of working-side muscle force relative to balancing-side muscle force (i.e., the W/B muscle force ratio) in these species because we have no reliable estimate of magnitude, direction, and precise location of the bite force during mastication. A comparison of the mastication data with the earlier data recorded during isometric molar biting, however, supports the hypothesis that the two anthropoids have a small W/B jaw-adductor muscle force ratio in comparison to thick-tailed galagos. These data also support the hypothesis that increased recruitment of balancing-side jaw-adductor muscle force in anthropoids is functionally linked to the evolution of symphyseal fusion or strengthening. Moreover, these data refute the hypothesis that the recruitment pattern differences between macaques and thick-tailed galagos are due to allometric factors. Finally, although the evolution of symphyseal fusion in primates may be linked to increased stress associated with increased balancing-side muscle force, it is currently unclear as to whether the increased force is predominately vertically directed, transversely directed, or is a near equal combination of these two force components (cf. Ravosa and Hylander [1994] In Fleagle and Kay [eds.]: Anthropoid Origins. New York: Plenum, pp. 447-468).

In vivo and in vitro bone strain in the owl monkey circumorbital region and the function of the postorbital septum.

Anthropoids and tarsiers are the only vertebrates possessing a postorbital septum. This septum, formed by the frontal, alisphenoid, and zygomatic bones, separates the orbital contents from the temporal muscles. Three hypotheses suggest that the postorbital septum evolved to resist stresses acting on the skull during mastication or incision. The facial-torsion hypothesis posits that the septum resists twisting of the face about a rostrocaudal axis during unilateral mastication; the transverse-bending hypothesis argues that the septum resists caudally directed forces acting at the lateral orbital margin during mastication or incision; and the tension hypothesis suggests that the septum resists ventrally directed components of masseter muscle force during mastication and incision. This study evaluates these hypotheses using in vitro and in vivo bone strain data recorded from the circumorbital region of owl monkeys. Incisor loading of an owl monkey skull in vitro bends the face upward in the sagittal plane, compressing the interorbital region rostrocaudally and "buckling" the lateral orbital walls. Unilateral loading of the toothrow in vitro also bends the face in the sagittal plane, compressing the interorbital region rostrocaudally and buckling the working side lateral orbital wall. When the lateral orbital wall is partially cut, so as to reduce the width of its attachment to the braincase, the following changes in circumorbital bone strain patterns occur. During loading of the incisors, lower bone strain magnitudes are recorded in the interorbital region and lateral orbital walls. In contrast, during unilateral loading of the P3, higher bone strain magnitudes are observed in the interorbital region, and generally lower bone strain magnitudes are observed in the lateral orbital walls. During unilateral loading of the M2, higher bone strain magnitudes are observed in both the interorbital region and in the lateral orbital wall ipsilateral to the loaded molar. Comparisons of the in vitro results with data gathered in vivo suggest that, during incision and unilateral mastication, the face is subjected to upward bending in the sagittal plane resulting in rostrocaudal compression of the interorbital region. Modeling the lateral orbital walls as curved plates suggests that during mastication the working side wall is buckled due to the dorsally directed component of the maxillary bite force which causes upward bending of the face in the sagittal plane. The balancing side lateral orbital wall may also be buckled due to upward bending of the face in the sagittal plane as well as being twisted by the caudoventrally directed components of the superficial masseter muscle force. The in vivo data do not exclude the possibility that the postorbital septum functions to improve the structural integrity of the postorbital bar during mastication. However, there is no reason to believe that a more robust postorbital bar could not also perform this function. Hypotheses stating that the postorbital septum originally evolved to reinforce the skull against routine masticatory loads must explain why, rather than evolving a postorbital septum, the stem anthropoids did not simply enlarge their postorbital bars.

Basicranial flexion, relative brain size, and facial kyphosis in nonhuman primates.

Numerous hypotheses explaining interspecific differences in the degree of basicranial flexion have been presented. Several authors have argued that an increase in relative brain size results in a spatial packing problem that is resolved by flexing the basicranium. Others attribute differences in the degree of basicranial flexion to different postural behaviors, suggesting that more orthograde animals require a ventrally flexed pre-sella basicranium in order to maintain the eyes in a correct forward-facing orientation. Less specific claims are made for a relationship between the degree of basicranial flexion and facial orientation. In order to evaluate these hypotheses, the degree of basicranial flexion (cranial base angle), palate orientation, and orbital axis orientation were measured from lateral radiographs of 68 primate species and combined with linear and volumetric measures as well as data on the size of the neocortex and telencephalon. Bivariate correlation and partial correlation analyses at several taxonomic levels revealed that, within haplorhines, the cranial base angle decreases with increasing neurocranial volume relative to basicranial length and is positively correlated with angles of facial kyphosis and orbital axis orientation. Strepsirhines show no significant correlations between the cranial base angle and any of the variables examined. It is argued that prior orbital approximation in the ancestral haplorhine integrated the medial orbital walls and pre-sella basicranium into a single structural network such that changes in the orientation of one necessarily affect the other. Gould's ("Ontogeny and Phylogeny." Cambridge: Belknap Press, 1977) hypothesis, that the highly flexed basicranium of Homo may be due to a combination of a large brain and a relatively short basicranium, is corroborated.