Prolonged use of a soft diet during early growth and development alters feeding behavior and chewing kinematics in a young animal model.

In infants and children with feeding and swallowing issues, modifying solid foods to form a liquid or puree is used to ensure adequate growth and nutrition. However, the behavioral and neurophysiological effects of prolonged use of this intervention during critical periods of postnatal oral skill development have not been systematically examined, although substantial anecdotal evidence suggests that it negatively impacts downstream feeding motor and coordination skills, possibly due to immature sensorimotor development. Using an established animal model for infant and juvenile feeding physiology, we leverage X-ray reconstruction of moving morphology to compare feeding behavior and kinematics between 12-week-old pigs reared on solid chow (control) and an age- and sex-matched cohort raised on the same chow softened to a liquid. When feeding on two novel foods, almond and apple, maintenance on a soft diet decreases gape cycle duration, resulting in a higher chewing frequency. When feeding on almonds, pigs in this group spent less time ingesting foods compared to controls, and chewing cycles were characterized by less jaw rotation about a dorsoventral axis (yaw) necessary for food reduction. There was also a reduced tendency to alternate chewing side with every chew during almond chewing, a behavioral pattern typical of pigs. These more pronounced impacts on behavior and kinematics during feeding on almonds, a tougher and stiffer food than apples, suggest that food properties mediate the behavioral and physiological impacts of early texture modification and that the ability to adapt to different food properties may be underdeveloped. In contrast, the limited effects of food texture modification on apple chewing indicate that such intervention/treatment does not alter feeding behavior of less challenging foods. Observed differences cannot be attributed to morphology because texture modification over the treatment period had limited impact on craniodental growth. Short-term impacts of soft-texture modification during postweaning development on feeding dynamics should be considered as potential negative outcomes of this treatment strategy.

Symphyseal morphology and jaw muscle recruitment levels during mastication in musteloid carnivorans.

In studies of mammalian mastication, a possible relationship has been proposed between bilateral recruitment of jaw adductor muscle force during unilateral chewing and the degree of fusion of the mandibular symphysis. Specifically, species that have unfused, mobile mandibular symphyses tend to utilize lower levels of jaw adductor force on the balancing (nonchewing) than the working (chewing) side of the head, when compared to related species with fused symphyses. Here, we compare jaw adductor recruitment levels in two species of musteloid carnivoran: the carnivorous ferret (unfused symphysis), and the frugivorous kinkajou (fused symphysis). During forceful chewing, we observe that ferrets recruit far more working-side muscle force than kinkajous, regardless of food toughness and that high working-to-balancing side ratios are the result of increased working-side force, often coupled with reduced balancing-side force. We propose that in carnivorans, high working-to-balancing side force ratios coupled with an unfused mandibular symphysis are necessary to rotate the hemimandible for precise unilateral occlusion of the carnassial teeth and to sustain laterally oriented force on the jaw to engage the carnassial teeth during shearing of tough foods. In contrast, the kinkajou's flattened cheekteeth permit less precise occlusion and require medially-oriented forces for grinding, thus, a fused symphysis is mechanically beneficial.

Characterizing tongue deformations during mastication using changes in planar components of three-dimensional angles.

Understanding of tongue deformations during mammalian mastication is limited, but has benefited from recent developments in multiplanar imaging technology. Here, we demonstrate how a standardized radiopaque marker implant configuration and biplanar fluoroscopy can quantify three-dimensional shape changes during chewing in pigs. Transverse and sagittal components of the three-dimensional angle between markers enable characterizing deformations in anatomically relevant directions. The transverse component illustrates bending to the left or to the right, which can occur symmetrically or asymmetrically, the latter sometimes indicating regional widening. The sagittal component reflects 'arching' or convex deformations in the dorsoventral dimension symmetrically or asymmetrically, the latter characteristic of twisting. Trends are detected in both the transverse and sagittal planes, and combinations thereof, to modify tongue shape in complex deformations. Both the transverse and sagittal components were also measured at key jaw and tongue positions, demonstrating variability particularly with respect to maximum and minimum gape. This highlights the fact that unlike tongue position, tongue deformations are more independent of jaw position, likely in response to the ever-changing bolus shape and position. From a methodological perspective, our study showcases advantages of a repeatable three-marker implant configuration suitable for animals of different sizes and highlights considerations for different implant patterns. This article is part of the theme issue 'Food processing and nutritional assimilation in animals'.

Regional Tongue Deformations During Chewing and Drinking in the Pig.

As a muscular hydrostat, the tongue undergoes complex deformations during most oral behaviors, including chewing and drinking. During thesebehaviors, deformations occur in concert with tongue and jaw movements to position and transport the bolus. Moreover, the various parts of the tongue may move and deform at similar timepoints relative to the gape cycle or they may occur at different timepoints, indicating regional biomechanical and functional variation. The goal of this study is to quantify tongue deformations during chewing and drinking in pigs by characterizing intrinsic changes in tongue dimensions (i.e., length and width) across multiple regions simultaneously. Tongue deformations are generally larger during chewing cycles compared to drinking cycles. Chewing and drinking also differ in the timing, relative to the gape cycle, of regional length and width, but not total length, deformations. This demonstrates functional differences in the temporal dynamics of localized shape changes, whereas the global properties of jaw-tongue coordination are maintained. Finally, differences in the trade-off between length and width deformations demonstrate that the properties of a muscular hydrostat are observed at the whole tongue level, but biomechanical variation (e.g., changes in movements and deformations) at the regional level exists. This study provides new critical insights into the regional contributions to tongue deformations as a basis for future work on multidimensional shape changes in soft tissues.

En tant qu'hydrostat musculaire, la langue subit des déformations complexes pendant la plupart des comportements oraux, en particulier au cours de la mastication et de l'ingestion de liquide. Au cours de ces comportements, les déformations se produisent de concert avec les mouvements de la langue et des mâchoires pour positionner et transporter le bolus. De plus, les différentes parties de la langue peuvent se déplacer et se déformer à des moments similaires ou différents par rapport au cycle d'ouverture de la bouche, indiquant une variation biomécanique et fonctionnelle régionale de la langue. L'objectif de cette étude est de quantifier les déformations de la langue pendant la mastication et l'ingestion d'eau chez le porc en caractérisant les changements intrinsèques des dimensions de la langue (i.e., longueur et largeur) des différentes régions de la langue simultanément. Les déformations de la langue sont généralement plus importantes pendant les cycles de mastication que pendant les cycles d'ingestion d'eau. La mastication et l'ingestion d'eau diffèrent également dans le timing (par rapport au cycle d'ouverture de la bouche) des déformations régionales de la langue en longueur et en largeur, mais pas en longueur totale. Cela démontre des différences fonctionnelles dans la dynamique temporelle des changements localisés de la forme de la langue alors que les propriétés globales de la coordination mâchoire-langue sont maintenues. Enfin, les différences dans le compromis mettant en jeu les déformations en longueur et en largeur démontrent que les propriétés d'un hydrostat musculaire sont observées au niveau de la langue entière, mais qu'il existe une variation biomécanique (par exemple, des changements dans les mouvements et les déformations) au niveau régional. Cette étude fournit de nouvelles informations essentielles sur les contributions régionales des déformations de la langue, qui serviront de base aux travaux futurs sur les changements multidimensionnels de forme dans les tissus mous.

Jaw kinematics and tongue protraction-retraction during chewing and drinking in the pig.

Mastication and drinking are rhythmic and cyclic oral behaviors that require interactions between the tongue, jaw and a food or liquid bolus, respectively. During mastication, the tongue transports and positions the bolus for breakdown between the teeth. During drinking, the tongue aids in ingestion and then transports the bolus to the oropharynx. The objective of this study was to compare jaw and tongue kinematics during chewing and drinking in pigs. We hypothesized there would be differences in jaw gape cycle dynamics and tongue protraction-retraction between behaviors. Mastication cycles had an extended slow-close phase, reflecting tooth-food-tooth contact, whereas drinking cycles had an extended slow-open phase, corresponding to tongue protrusion into the liquid. Compared with chewing, drinking jaw movements were of lower magnitude for all degrees of freedom examined (jaw protraction, yaw and pitch), and were bilaterally symmetrical with virtually no yaw. The magnitude of tongue protraction-retraction (Txt), relative to a mandibular coordinate system, was greater during mastication than during drinking, but there were minimal differences in the timing of maximum and minimum Txt relative to the jaw gape cycle between behaviors. However, during drinking, the tongue tip is often located outside the oral cavity for the entire cycle, leading to differences between behaviors in the timing of anterior marker maximum Txt. This demonstrates that there is variation in tongue-jaw coordination between behaviors. These results show that jaw and tongue movements vary significantly between mastication and drinking, which hints at differences in the central control of these behaviors.

Jaw kinematics and tongue protraction-retraction during chewing and drinking in the pig.

Mastication and drinking are rhythmic and cyclic oral behaviors that require interactions between the tongue, jaw and a food or liquid bolus, respectively. During mastication, the tongue transports and positions the bolus for breakdown between the teeth. During drinking, the tongue aids in ingestion and then transports the bolus to the oropharynx. The objective of this study was to compare jaw and tongue kinematics during chewing and drinking in pigs. We hypothesized there would be differences in jaw gape cycle dynamics and tongue protraction-retraction between behaviors. Mastication cycles had an extended slow-close phase, reflecting tooth-food-tooth contact, whereas drinking cycles had an extended slow-open phase, corresponding to tongue protrusion into the liquid. Compared with chewing, drinking jaw movements were of lower magnitude for all degrees of freedom examined (jaw protraction, yaw and pitch), and were bilaterally symmetrical with virtually no yaw. The magnitude of tongue protraction-retraction (Txt), relative to a mandibular coordinate system, was greater during mastication than during drinking, but there were minimal differences in the timing of maximum and minimum Txt relative to the jaw gape cycle between behaviors. However, during drinking, the tongue tip is often located outside the oral cavity for the entire cycle, leading to differences between behaviors in the timing of anterior marker maximum Txt. This demonstrates that there is variation in tongue-jaw coordination between behaviors. These results show that jaw and tongue movements vary significantly between mastication and drinking, which hints at differences in the central control of these behaviors.

Getting Humans Off Monkeys' Backs: Using Primate Acclimation as a Guide for Habitat Management Efforts.

Wild primates face grave conservation challenges, with habitat loss and climate change projected to cause mass extinctions in the coming decades. As large-bodied Neotropical primates, mantled howling monkeys (Alouatta palliata) are predicted to fare poorly under climate change, yet are also known for their resilience in a variety of environments, including highly disturbed habitats. We utilized ecophysiology research on this species to determine the morphological, physiological, and behavioral mechanisms howlers employ to overcome ecological challenges. Our data show that howlers at La Pacifica, Costa Rica are capable of modifying body size. Howlers displayed reduced mass in warmer, drier habitats, seasonal weight changes, frequent within-lifetime weight fluctuations, and gradual increases in body mass over the past four decades. These within-lifetime changes indicate a capacity to modify morphology in a way that can impact animals' energetics and thermodynamics. Howlers are also able to consume foods with a wide variety of food material properties by altering oral processing during feeding. While this capability suggests some capacity to cope with the phenological shifts expected from climate change and increased habitat fragmentation, data on rates of dental microwear warn that these acclimations may also cost dental longevity. Lastly, we found that howlers are able to acclimate to changing thermal pressures. On shorter-term daily scales, howlers use behavioral mechanisms to thermoregulate, including timing activities to avoid heat stress and utilizing cool microhabitats. At the seasonal scale, animals employ hormonal pathways to influence heat production. These lines of evidence cumulatively indicate that howlers possess morphological, physiological, and behavioral mechanisms to acclimate to environmental challenges. As such, howlers' plasticity may facilitate their resilience to climate change and habitat loss. While habitat loss in the tropics is unlikely to abate, our results point to a potential benefit of active management and selective cultivation to yield large, interconnected forest fragments with targeted phenology that provides both a complex physical structure and a diversity of food sources. These steps could assist howlers in using their natural acclimation potential to survive future conservation threats.

Unilateral lingual nerve transection alters jaw-tongue coordination during mastication in pigs.

During chewing, movements and deformations of the tongue are coordinated with jaw movements to manage and manipulate the bolus and avoid injury. Individuals with injuries to the lingual nerve report both tongue injuries due to biting and difficulties in chewing, primarily because of impaired bolus management, suggesting that jaw-tongue coordination relies on intact lingual afferents. Here, we investigate how unilateral lingual nerve (LN) transection affects jaw-tongue coordination in an animal model (pig, Sus scrofa). Temporal coordination between jaw pitch (opening-closing) and 1) anteroposterior tongue position (i.e., protraction-retraction), 2) anteroposterior tongue length, and 3) mediolateral tongue width was compared between pre- and post-LN transection using cross-correlation analyses. Overall, following LN transection, the lag between jaw pitch and the majority of tongue kinematics decreased significantly, demonstrating that sensory loss from the tongue alters jaw-tongue coordination. In addition, decrease in jaw-tongue lag suggests that, following LN transection, tongue movements and deformations occur earlier in the gape cycle than when the lingual sensory afferents are intact. If the velocity of tongue movements and deformations remains constant, earlier occurrence can reflect less pronounced movements, possibly to avoid injuries. The results of this study demonstrate that lingual afferents participate in chewing by assisting with coordinating the timing of jaw and tongue movements. The observed changes may affect bolus management performance and/or may represent protective strategies because of altered somatosensory awareness of the tongue.NEW & NOTEWORTHY Chewing requires coordination between tongue and jaw movements. We compared the coordination of tongue movements and deformation relative to jaw opening-closing movements pre- and post-lingual nerve transection during chewing in pigs. These experiments reveal that the timing of jaw-tongue coordination is altered following unilateral disruption of sensory information from the tongue. Therefore, maintenance of jaw-tongue coordination requires bilateral sensory information from the tongue.

Effects of food properties on chewing in pigs: Flexibility and stereotypy of jaw movements in a mammalian omnivore.

Chewing is a rhythmic oral behavior that requires constant modifications of jaw movements in response to changes in food properties. The food-specific kinematic response is dependent on the potential for kinematic flexibility allowed by morphology and modulation of motor control. This study investigates the effects of food toughness and stiffness on the amplitude and variability of jaw movements during chewing in a typical omnivorous mammalian model (pigs). Jaw movements were reconstructed using X-ray Reconstruction Of Moving Morphology (XROMM) and kinematic data associated with the amplitude of jaw pitch (opening-closing) and jaw yaw (mediolateral rotation) were extracted for each cycle. Between-food differences were tested for the amplitude of jaw movements during each phase of the gape cycle, as well as in their respective within-food variability, or stereotypy, as indicated by coefficients of variation. With increasing toughness, jaw pitch amplitude is decreased during fast close, larger and more stereotyped during slow close, smaller but more variable during slow open, and more variable during fast open. In addition, when chewing on tougher foods, the amplitude of jaw yaw during slow close only increases in a subset of individuals, but all become less variable (i.e., more stereotyped). In contrast, increasing food stiffness has no effect on the amplitude or the variability of jaw pitch, whereas jaw yaw increases significantly in the majority of individuals studied. Our data demonstrate that food stiffness and toughness both play a role in modulating gape cycle dynamics by altering the trajectory of jaw movements, especially during the slow-close phase and tooth-food-tooth contact, albeit differently. This highlights how a generalist oral morphology such as that of pigs (e.g., bunodont teeth lacking precise occlusion, permissive temporomandibular joint allowing extensive condylar displacements in 3 dimensions) enables organisms to not only adjust chewing movements in their amplitude, but also in their variability.

The effect of unilateral lingual nerve injury on the kinematics of mastication in pigs.

OBJECTIVE: This study evaluates the effect of unilateral lingual sensory loss on the spatial and temporal dynamics of jaw movements during pig chewing. DESIGN: X-ray Reconstruction of Moving Morphology (XROMM) was used to reconstruct the 3-dimensional jaw movements of 6 pigs during chewing before and after complete unilateral lingual nerve transection. The effect of the transection were evaluated at the temporal and spatial level using Multiple Analysis of Variance. Temporal variables include gape cycle and phase durations, and the corresponding relative phase durations. Spatial variables include the amplitude of jaw opening, jaw yaw, and mandibular retraction-protraction. RESULTS: The temporal and spatial dynamics of jaw movements did not differ when chewing ipsilateral versus contralateral to the transection. When compared to pre-transection data, 4 of the 6 animals showed significant changes in temporal characteristics of the gape cycle following the transection, irrespective of chewing side, but the specific response to the lesion was highly dependent on the animal. On the other hand, in affected individuals the amplitude of jaw movements was altered similarly in all 3 dimensions: jaw opening and protraction-retraction increased whereas jaw yaw decreased. CONCLUSION: The variable impact of this injury in this animal model suggests that individuals use different compensatory strategies to adjust or maintain the temporal dynamics of the gape cycle. Because the amplitude of jaw movements are more adversely affected than their timing, results suggest that maintaining the tongue-jaw coordination is critical and this can come at the expense of bolus handling and masticatory performance.

Flexibility of feeding movements in pigs: effects of changes in food toughness and stiffness on the timing of jaw movements.

In mammals, chewing movements can be modified, or flexible, in response to changes in food properties. Variability between and within food in the temporal characteristics of chewing movements can impact chewing frequency and rhythmicity, which in turn may affect food breakdown, energy expenditure and tooth wear. Here, we compared total chewing cycle duration and intra-cycle phase durations in pigs chewing on three foods varying in toughness and stiffness: apples (low toughness, low stiffness), carrots (high toughness, low stiffness), and almonds (high toughness, high stiffness). We also determined whether within-food variability in timing parameters is modified in response to changes in food properties. X-ray Reconstruction Of Moving Morphology (XROMM) demonstrates that the timing of jaw movements are flexible in response to changes in food properties. Within each food, pigs also exhibited flexibility in their ability to vary cycle parameters. The timing of jaw movements during processing of high-toughness foods is more variable, potentially decreasing chewing rhythmicity. In contrast, low-toughness foods result in jaw movements that are more stereotyped in their timing parameters. In addition, the duration of tooth-food-tooth contact is more variable during the processing of low-stiffness foods compared with tough or stiff foods. Increased toughness is suggested to alter the timing of the movements impacting food fracture whereas increased stiffness may require a more cautious control of jaw movements. This study emphasizes that flexibility in biological movements in response to changes in conditions may not only be observed in timing but also in the variability of their timing within each condition.

Thyroid hormone fluctuations indicate a thermoregulatory function in both a tropical (Alouatta palliata) and seasonally cold-habitat (Macaca fuscata) primate.

Thyroid hormones boost animals' basal metabolic rate and represent an important thermoregulatory pathway for mammals that face cold temperatures. Whereas the cold thermal pressures experienced by primates in seasonal habitats at high latitudes and elevations are often apparent, tropical habitats also display distinct wet and dry seasons with modest changes in thermal environment. We assessed seasonal and temperature-related changes in thyroid hormone levels for two primate species in disparate thermal environments, tropical mantled howlers (Alouatta palliata), and seasonally cold-habitat Japanese macaques (Macaca fuscata). We collected urine and feces from animals and used ELISA to quantify levels of the thyroid hormone triiodothyronine (fT3 ). For both species, fT3 levels were significantly higher during the cooler season (wet/winter), consistent with a thermoregulatory role. Likewise, both species displayed greater temperature deficits (i.e., the degree to which animals warm their body temperature relative to ambient) during the cooler season, indicating greater thermoregulatory pressures during this time. Independently of season, Japanese macaques displayed increasing fT3 levels with decreasing recently experienced maximum temperatures, but no relationship between fT3 and recently experienced minimum temperatures. Howlers increased fT3 levels as recently experienced minimum temperatures decreased, although demonstrated the opposite relationship with maximum temperatures. This may reflect natural thermal variation in howlers' habitat: wet seasons had cooler minimum and mean temperatures than the dry season, but similar maximum temperatures. Overall, our findings support the hypothesis that both tropical howlers and seasonally cold-habitat Japanese macaques utilize thyroid hormones as a mechanism to boost metabolism in response to thermoregulatory pressures. This implies that cool thermal pressures faced by tropical primates are sufficient to invoke an energetically costly and relatively longer-term thermoregulatory pathway. The well-established relationship between thyroid hormones and energetics suggests that the seasonal hormonal changes we observed could influence many commonly studied behaviors including food choice, range use, and activity patterns.

An ontogenetic perspective on symphyseal fusion, occlusion and mandibular loading in alpacas (Vicugna pacos).

A primary hypothesis for the evolution of mandibular symphyseal fusion in some mammals is that it functions to resist loads incurred during routine mastication. Anecdotal support for this hypothesis is based on the fact that when the symphysis fuses, it typically does so early during postnatal ontogeny prior to or around the time of weaning. However, little is known about the process of fusion, particularly relative to feeding behaviors and the dynamics of mastication, including occlusion and masticatory loading. In the present study, we investigate the timing and process of symphyseal fusion in alpacas (Vicugna pacos) in the context of maturation of the oral apparatus and oral behavior. We also report on in vivo strains from the symphysis and corpus in young alpacas prior to and following full fusion and M1 occlusion. Results show that fusion begins rostrally by 1 month and is complete by 6-7 months whereas all deciduous premolars and M1 come into occlusion by 6 months. Although symphyseal loading patterns are maintained throughout ontogeny, in young alpacas symphyseal strain magnitudes are low compared with adults but corpus strain magnitudes are comparable to those found in adults. Reduced symphyseal loading in young individuals is contrary to what might be predicted given that the symphysis is still fusing. When considered in light of the development of occlusion and rumination, strain magnitudes may be necessarily low and reflect an overall delay in the maturation of masticatory dynamics.

An assessment of skin temperature gradients in a tropical primate using infrared thermography and subcutaneous implants.

Infrared thermography has become a useful tool to assess surface temperatures of animals for thermoregulatory research. However, surface temperatures are an endpoint along the body's core-shell temperature gradient. Skin and fur are the peripheral tissues most exposed to ambient thermal conditions and are known to serve as thermosensors that initiate thermoregulatory responses. Yet relatively little is known about how surface temperatures of wild mammals measured by infrared thermography relate to subcutaneous temperatures. Moreover, this relationship may differ with the degree that fur covers the body. To assess the relationship between temperatures and temperature gradients in peripheral tissues between furred and bare areas, we collected data from wild mantled howling monkeys (Alouatta palliata) in Costa Rica. We used infrared thermography to measure surface temperatures of the furred dorsum and bare facial areas of the body, recorded concurrent subcutaneous temperatures in the dorsum, and measured ambient thermal conditions via a weather station. Temperature gradients through cutaneous tissues (subcutaneous-surface temperature) and surface temperature gradients (surface-ambient temperature) were calculated. Our results indicate that there are differences in temperatures and temperature gradients in furred versus bare areas of mantled howlers. Under natural thermal conditions experienced by wild animals, the bare facial areas were warmer than temperatures in the furred dorsum, and cutaneous temperature gradients in the face were more variable than the dorsum, consistent with these bare areas acting as thermal windows. Cutaneous temperature gradients in the dorsum were more closely linked to subcutaneous temperatures, while facial temperature gradients were more heavily influenced by ambient conditions. These findings indicate that despite the insulative properties of fur, for mantled howling monkeys surface temperatures of furred areas still demonstrate a relationship with subcutaneous temperatures. Given that most mammals possess dense fur, this provides insight for using infrared imaging in thermoregulatory studies of wild animals lacking bare skin.

The influence of diet on masticatory motor patterns in musteloid carnivorans: An analysis of jaw adductor activity in ferrets (Mustela putorius furo) and kinkajous (Potos flavus).

Broad similarities in the timing of jaw adductor activity driving jaw movements across distantly related and morphologically disparate species have led to the hypothesis that mammalian masticatory motor patterns are conserved. However, some quantitative analyses also suggest that masticatory motor patterns have evolved in concert with dietary and/or morphological specialization. Here, we assess this relationship in two closely related carnivoran species with divergent diets and morphology: carnivorous ferrets and frugivorous kinkajous. Using electromyography to characterize jaw adductor activity during rhythmic mastication, we test the hypotheses that (1) carnivoran masticatory motor patterns differ from those of non-carnivorans based on previously published data, and (2) differences between ferret and kinkajou motor patterns are associated with dietary and morphological differences. We find that both species exhibit highly synchronous jaw adductor activity that is likely typical of most carnivorans. Kinkajous differ from ferrets, however, in having a balancing-side zygomaticomandibularis that is active later than all other adductors. The significance of these different masticatory motor patterns may relate to morphological differences in the dentition of ferrets and kinkajous. Whereas ferret cheek teeth have vertical occlusal surfaces that limit jaw closing to a primarily dorsally directed movement, kinkajous have relatively flat occlusal surfaces that allow more transverse movement, which may be essential for processing fruits. Our results suggest that some aspects of masticatory motor patterns are highly conserved yet some components are modified in concert with functional and morphological evolution of the masticatory apparatus.

Muscle Logic: New Knowledge Resource for Anatomy Enables Comprehensive Searches of the Literature on the Feeding Muscles of Mammals.

BACKGROUND: In recent years large bibliographic databases have made much of the published literature of biology available for searches. However, the capabilities of the search engines integrated into these databases for text-based bibliographic searches are limited. To enable searches that deliver the results expected by comparative anatomists, an underlying logical structure known as an ontology is required. DEVELOPMENT AND TESTING OF THE ONTOLOGY: Here we present the Mammalian Feeding Muscle Ontology (MFMO), a multi-species ontology focused on anatomical structures that participate in feeding and other oral/pharyngeal behaviors. A unique feature of the MFMO is that a simple, computable, definition of each muscle, which includes its attachments and innervation, is true across mammals. This construction mirrors the logical foundation of comparative anatomy and permits searches using language familiar to biologists. Further, it provides a template for muscles that will be useful in extending any anatomy ontology. The MFMO is developed to support the Feeding Experiments End-User Database Project (FEED, https://feedexp.org/), a publicly-available, online repository for physiological data collected from in vivo studies of feeding (e.g., mastication, biting, swallowing) in mammals. Currently the MFMO is integrated into FEED and also into two literature-specific implementations of Textpresso, a text-mining system that facilitates powerful searches of a corpus of scientific publications. We evaluate the MFMO by asking questions that test the ability of the ontology to return appropriate answers (competency questions). We compare the results of queries of the MFMO to results from similar searches in PubMed and Google Scholar. RESULTS AND SIGNIFICANCE: Our tests demonstrate that the MFMO is competent to answer queries formed in the common language of comparative anatomy, but PubMed and Google Scholar are not. Overall, our results show that by incorporating anatomical ontologies into searches, an expanded and anatomically comprehensive set of results can be obtained. The broader scientific and publishing communities should consider taking up the challenge of semantically enabled search capabilities.

Effect of Postnatal Myostatin Inhibition on Bite Mechanics in Mice.

As a negative regulator of muscle size, myostatin (Mstn) impacts the force-production capabilities of skeletal muscles. In the masticatory system, measures of temporalis-stimulated bite forces in constitutive myostatin KOs suggest an absolute, but not relative, increase in jaw-muscle force. Here, we assess the phenotypic and physiologic impact of postnatal myostatin inhibition on bite mechanics using an inducible conditional KO mouse in which myostatin is inhibited with doxycycline (DOX). Given the increased control over the timing of gene inactivation in this model, it may be more clinically-relevant for developing interventions for age-associated changes in the musculoskeletal system. DOX was administered for 12 weeks starting at age 4 months, during which time food intake was monitored. Sex, age and strain-matched controls were given the same food without DOX. Bite forces were recorded just prior to euthanasia after which muscle and skeletal data were collected. Food intake did not differ between control or DOX animals within each sex. DOX males were significantly larger and had significantly larger masseters than controls, but DOX and control females did not differ. Although there was a tendency towards higher absolute bite forces in DOX animals, this was not significant, and bite forces normalized to masseter mass did not differ. Mechanical advantage for incisor biting increased in the DOX group due to longer masseter moment arms, likely due to a more anteriorly-placed masseter insertion. Despite only a moderate increase in bite force in DOX males and none in DOX females, the increase in masseter mass in males indicates a potentially positive impact on jaw muscles. Our data suggest a sexual dimorphism in the role of mstn, and as such investigations into the sex-specific outcomes is warranted.

In Vivo Measurement of Mesokinesis in Gekko gecko: The Role of Cranial Kinesis during Gape Display, Feeding and Biting.

Cranial kinesis refers to movements of skeletal sub-units relative to one another at mobile sutures within the skull. The presence and functional significance of cranial kinesis has been investigated in various vertebrates, with much of our understanding coming from comparative studies and manipulation of ligamentous specimens. Drawing on these studies, cranial kinesis in lizards has been modeled as a four-bar linkage system involving streptostyly (rotation of the quadrate), hypokinesis (dorsoventral flexion and extension of the palato-maxillary sub-unit), mesokinesis (dorsoventral flexion and extension of the snout at the fronto-parietal suture) and metakinesis (sliding movements between parietal and supraocciptal bones). In vivo studies, although limited, suggest that cranial kinesis serves an important role during routine behaviors such as feeding. Here, we use X-ray Reconstruction Of Moving Morphology to further quantify mesokinesis in vivo in Gekko gecko during three routine behaviors: gape display, biting and post-ingestion feeding. During gape display, the snout rotates dorsally above rest position, with mesokinesis accounting for a 10% increase in maximum gape over that achieved solely by the depression of the lower jaw. During defensive biting, the snout rotates ventrally below rest position to participate in gape closure. Finally, ventroflexion of the snout also occurs during post-ingestion feeding, accounting for 42% of gape closure during intra-oral transport, 86% during puncture-crushing, and 61% during pharyngeal packing. Mesokinesis thus appears to facilitate prey puncturing by allowing the snout to rotate ventrally so that the upper teeth pierce the prey item, thus limiting the need for large movements of the lower jaw. This is suggested to maintain a firm grip on the prey and reduce the possibility of prey escape. More generally, this study demonstrates that mesokinesis is a key component of defensive biting and gape display behaviors, as well as post-ingestion feeding, all of which are linked to organismal fitness.

Using biplanar fluoroscopy to guide radiopaque vascular injections: a new method for vascular imaging.

Studying vascular anatomy, especially in the context of relationships with hard tissues, is of great interest to biologists. Vascular studies have provided significant insight into physiology, function, phylogenetic relationships, and evolutionary patterns. Injection of resin or latex into the vascular system has been a standard technique for decades. There has been a recent surge in popularity of more modern methods, especially radiopaque latex vascular injection followed by CT scanning and digital "dissection." This technique best displays both blood vessels and bone, and allows injections to be performed on cadaveric specimens. Vascular injection is risky, however, because it is not a standardizable technique, as each specimen is variable with regard to injection pressure and timing. Moreover, it is not possible to view the perfusion of injection medium throughout the vascular system of interest. Both data and rare specimens can therefore be lost due to poor or excessive perfusion. Here, we use biplanar video fluoroscopy as a technique to guide craniovascular radiopaque latex injection. Cadaveric domestic pigs (Sus scrofa domestica) and white-tailed deer (Odocoileus virginianus) were injected with radiopaque latex under guidance of fluoroscopy. This method was found to enable adjustments, in real-time, to the rate, location, and pressure at which latex is injected in order to avoid data and specimen loss. In addition to visualizing the injection process, this technique can be used to determine flow patterns, and has facilitated the development of consistent markers for complete perfusion.

Body temperature and thermal environment in a generalized arboreal anthropoid, wild mantled howling monkeys (Alouatta palliata).

Free-ranging primates are confronted with the challenge of maintaining an optimal range of body temperatures within a thermally dynamic environment that changes daily, seasonally, and annually. While many laboratory studies have been conducted on primate thermoregulation, we know comparatively little about the thermal pressures primates face in their natural, evolutionarily relevant environment. Such knowledge is critical to understanding the evolution of thermal adaptations in primates and for comparative evaluation of humans' unique thermal adaptations. We examined temperature and thermal environment in free-ranging, mantled howling monkeys (Alouatta palliata) in a tropical dry forest in Guanacaste, Costa Rica. We recorded subcutaneous (Tsc ) and near-animal ambient temperatures (Ta ) from 11 animals over 1586.5 sample hours during wet and dry seasons. Howlers displayed considerable variation in Tsc , which was largely attributable to circadian effects. Despite significant seasonal changes in the ambient thermal environment, howlers showed relatively little evidence for seasonal changes in Tsc . Howlers experienced warm thermal conditions which led to body cooling relative to the environment, and plateaus in Tsc at increasingly warm Ta . They also frequently faced cool thermal conditions (Ta < Tsc ) in which Tsc was markedly elevated compared with Ta . These data add to a growing body of evidence that non-human primates have more labile body temperatures than humans. Our data additionally support a hypothesis that, despite inhabiting a dry tropical environment, howling monkeys experience both warm and cool thermal pressures. This suggests that thermal challenges may be more prevalent for primates than previously thought, even for species living in nonextreme thermal environments.