Suckling in therian mammals requires synchronous activity in the tensor veli palatini, palatoglossus,mylohyoideusand intrinsic tongue muscles that close the fauces. These muscles draw the dorsal surface of the tongue against a tensed soft palate to form a seal between the oral cavity and oropharynx. Depressing the tongue in front of the seal induces negative pressure and draws milk into the oral cavity.To trace the origin offauces region in mammals,we studied serial sections of a pouch young marsupialandCT scans of non-mammalian cynodonts, ictidosaurs, mammaliaforms, and extant mammals.In Late Triassic ictidosaurs (Brasilitherium) and mammaliaforms, the origin of a medial slip of the reptilian posterior pterygoideus migrated to the lateral surface of the pterygopalatine boss that supported the lateral edge of a non-muscular soft palate.Suckling arose in early mammals after fibers of this slip migrated into the soft palate to form the tensor veli palatini, and the palatoglossus separated from the pharyngeal constrictors. The added stress on the pterygopalatine boss led to the addition of a hamulus.Once the transverse process of the pterygoid was lost, the lateral posterior pterygoideus differentiated to form the medial pterygoid.Monotreme ancestorsmodified the fauces regionto break down invertebrates between keratinized padsontheposteriortongue andundertheventral surface of a long palatine.Theylost the tensor veli palatini and palatoglossus andlack the ability to suckle.
Ictidosaurian genera are allocated to two families, Tritheledontidae and Therioherpetidae. This paper provides a diagnosis for Ictidosauria. The previously named family Brasilodontidae is shown to be a junior synonym of a family, Therioherpetidae. It is concluded that Ictidosauria originated from Late Permian procynosuchid non-mammalian cynodonts rather than from Middle Triassic probainognathid non-mammalian cynodonts. The structure of the skull and jaws of a derived traversodontid Ischignathus sudamericanus from the early Late Triassic of Argentina supports an earlier view that tritylodontids are more closely related to traversodontid than probainognathid non-mammalian cynodonts. Tritylodontids should not be included in Ictidosauria, nor should they considered to be a sister group to mammaliaforms.
Nasal regions of the non-mammalian cynodont, Massetognathus, Probainognathus, and Elliotherium were reconstructed from micro-CT scans and compared with scans and published accounts of more derived forms, including Brasilitherium, Morganucodon, Haldanodon, and extant mammals. The basic structure of the modern mammalian nose, already present in non-mammalian cynodonts of the Early Triassic, underwent little modification during the Triassic. A respiratory chamber opened into a nasopharyngeal passage through an enlarged primary choana bordered posteriorly by a transverse lamina that formed the floor to a more posterior olfactory chamber. Cartilaginous respiratory turbinals initially provided a surface for evaporative cooling during periods of increased activity in the exceptionally high ambient temperatures of the Triassic. A similar mechanism for heat loss is present in extant crocodilians, squamates, and mammals. In the Late Triassic and Early Jurassic non-mammaliaform cynodonts (Elliotherium) and mammaliaforms (Morganucodon), the pterygopalatine ridges behind the hard secondary palate extended ventrally and formed the lateral walls to a narrow nasopharynx as pterygoid hamuli do in extant mammals. Ridges in this position suggest the presence of a palatopharyngeus muscle in late non-mammaliaform cynodonts that could hold the larynx in an intranarial position during rest or low activity levels to prevent inhaled air from entering the oral cavity, thus allowing cartilaginous respiratory turbinals to assume an additional role as temporal countercurrent exchange sites for heat and water conservation. Ossification of respiratory turbinals in mammals enhanced their efficiency for conserving heat and water at rest, as well as their ability to dissipate heat during thermal stress.
To interpret the fundamental differences in the structure and origin of the braincase sidewalls of monotremes, multituberculates, and therians, we examined MicroCT scans of a mammaliaform, Morganucodon; two non-mammalian cynodonts, Massetognathus and Probainognathus; a stem therian, Vincelestes; a juvenile and adult monotreme, Ornithorhynchus; and two marsupials, Monodelphis and Didelphis. The skull of Morganucodon resembles the pattern predicted for an early mammal: the descending flanges of the frontal and parietal cover the lateral surface of the orbitosphenoid and the palatine forms most of the medial wall of the orbit. In monotremes, the lateral region of the chondrocranium ossifies to form a long presphenoid/orbitosphenoid complex. During the transition from early mammals to extant mammals the height of the alisphenoid decreased drastically, the anterior lamina extended anteriorly to form part of the sidewall while the lateral surface of the orbitosphenoid was exposed by the dorsal withdrawal of the frontal and parietal. By contrast, in multituberculates and therians the lateral edges of the frontals extended further ventrally and the orbitosphenoid was reduced to a smaller orbital exposure below the frontals. In multituberculates the alisphenoid decreased in height, replaced by an anterior extension of the anterior lamina. The palatine withdrew from the orbital wall, replaced by a dorsally directed expansion of the maxilla. Extant therians have lost the anterior lamina. The inferior edges of the frontal followed the further ventral migration of the orbitosphenoid. The alisphenoid and parietal form most of the braincase sidewall.
It has recently been suggested that the mechan- ism for lifting liquid from a bowl into the oral cavity during lapping is fundamentally different in cats and dogs: cats use adhesion of liquid to the tongue tip while dogs ‘scoop’ with their backwardly curled tongue. High-speed light videos and X-ray videos show that on the con- trary, both cats and dogs use the mechanism of adhesion. Liquid is transported through the oral cavity to the oesophagus, against gravity, on the surface of the tongue as it is drawn upwards, then a tight contact between the tongue surface and palatal rugae traps liquid and prevents its falling out as the tongue is protruded. At least three cycles are needed for intraoral transport of liquid in the dog.
Synopsis All infant mammals make a transition from suckling milk to eating solid foods. Yet, the neuromuscular implications of the transition from a liquid-only diet to solid foods are unknown even though the transport and swallowing of liquids is different from that of solids. We used legacy electromyography (EMG) data to test hypotheses concerning the changes in motor pattern and neuromuscular control that occur during the transition from an all-liquid diet to consumption of solid food in a porcine model. EMG signals were recorded from five oropharyngeal muscles in pigs at three developmental stages (infants, juveniles, and adults) feeding on milk, on food of an intermediate consistency (porridge), and on dry chow (juveniles and adults only). We measured cycle frequency and its variation in "transport cycles" and "swallow cycles". In the swallow cycles, a measure of variation of the EMG signal was also calculated. Variation in cycle frequency for transport and swallow cycles was lowest in adults, as predicted, suggesting that maturation of feeding mechanisms occurs as animals reach adulthood. Infants had lower variation in transport cycle frequency than did juveniles drinking milk, which may be due to the greater efficiency of the infant's tight oral seal against the teat during suckling, compared to a juvenile drinking from a bowl where a tight seal is not possible. Within juveniles, variation in both transport and swallow cycle frequencies was directly related to food consistency, with the highest variation occurring when drinking milk and the lowest when feeding on solid food. There was no difference in the variation of the EMG activity between intact infants and juveniles swallowing milk, although when the latter swallow porridge the EMG signals were less variable than for milk. These results suggest that consistency of food is a highly significant determinant of the variation in motor pattern, particularly in newly weaned animals.
The pharyngeal swallow can be elicited as an isolated event but, in normal animals, it occurs within the context of rhythmic tongue and jaw movement (RTJM). The response includes activation of the multifunctional geniohyoid muscle, which can either protract the hyoid or assist jaw opening; in conscious nonprimate mammals, two bursts of geniohyoid EMG activity (GHemg) occur in swallow cycles at times consistent with these two actions. However, during experimentally elicited pharyngeal swallows, GHemg classically occurs at the same time as hyoglossus and mylohyoid activity (short latency response) but, when the swallow is elicited in the decerebrate in the absence of RTJM, GHemg occurs later in the swallow (long latency response). We tested the hypothesis that it was not influences from higher centers but a brain stem mechanism, associated with RTJM, which caused GHemg to occur earlier in the swallow. In 38 decerebrate piglets, RTJM occurred sporadically in seven animals. Before RTJM, GHemg had a long latency, but, during RTJM, swallow related GHemg occurred synchronously with activity in hyoglossus and mylohyoid, early in the swallow. Both early and late responses were present during the changeover period. During this changeover period, duplicate electrodes in the geniohyoid could individually detect either the early or the late burst in the same swallow. This suggested that two sets of geniohyoid task units existed that were potentially active in the swallow and that they were differentially facilitated or inhibited depending on the presence or absence of rhythmic activity originating in the brain stem.
The masticatory motor patterns of three tammar wallabies and two red kangaroos were determined by analyzing the pattern of electromyographic (EMG) activity of the jaw adductors and correlating it with lower jaw movements, as recorded by digital video and videoradiography. Transverse jaw movements were limited by the width of the upper incisal arcade. Molars engaged in food breakdown during two distinct occlusal phases characterized by abrupt changes in the direction of working-side hemimandible movement. Separate orthal (Phase I) and transverse (Phase II) trajectories were observed. The working-side lower jaw initially was drawn laterally by the balancing-side medial pterygoid and then orthally by overlapping activity in the balancing- and working-side temporalis and the balancing-side superficial masseter and medial pterygoid. Transverse movement occurred principally via the working-side medial pterygoid and superficial masseter. This pattern contrasted to that of placental herbivores, which are known to break down food when they move the working-side lower jaw transversely along a relatively longer linear path without changing direction during the power stroke. The placental trajectory results from overlapping activity in the working- and balancing-side adductor muscles, suggesting that macropods and placental herbivores have modified the primitive masticatory motor pattern in different ways.
Although backward folding of the epiglottis is one of the signal events of the mammalian adult swallow, the epiglottis does not fold during the infant swallow. How this functional change occurs is unknown, but we hypothesize that a change in swallow mechanism occurs with maturation, prior to weaning. Using videofluoroscopy, we found three characteristic patterns of swallowing movement at different ages in the pig: an infant swallow, a transitional swallow and a post-weaning (juvenile or adult) swallow. In animals of all ages, the dorsal region of the epiglottis and larynx was held in an intranarial position by a muscular sphincter formed by the palatopharyngeal arch. In the infant swallow, increasing pressure in the oropharynx forced a liquid bolus through the piriform recesses on either side of a relatively stationary epiglottis into the esophagus. As the infant matured, the palatopharyngeal arch and the soft palate elevated at the beginning of the swallow, so exposing a larger area of the epiglottis to bolus pressure. In transitional swallows, the epiglottis was tilted backward relatively slowly by a combination of bolus pressure and squeezing of the epiglottis by closure of the palatopharyngeal sphincter. The bolus, however, traveled alongside but never over the tip of the epiglottis. In the juvenile swallow, the bolus always passed over the tip of the epiglottis. The tilting of the epiglottis resulted from several factors, including the action of the palatopharyngeal sphincter, higher bolus pressure exerted on the epiglottis and the allometry of increased size. In both transitional and juvenile swallows, the subsequent relaxation of the palatopharyngeal sphincter released the epiglottis, which sprang back to its original intranarial position.
The currently accepted description of the pattern of electromyographic (EMG) activity in the pharyngeal swallow is that reported by Doty and Bosma in 1956; however, those authors describe high levels of intramuscle and of interindividual EMG variation. We reinvestigated this pattern, testing two hypotheses concerning EMG variation: 1) that it could be reduced with modern methodology and 2) that it could be explained by selective detection of different types of motor units. In eight decerebrate infant pigs, we elicited radiographically verified pharyngeal swallows and recorded EMG activity from a total of 16 muscles. Synchronization signals from the video-radiographic system allowed the EMG activity associated with each swallow to be aligned directly with epiglottal movement. The movements were highly stereotyped, but the recorded EMG signals were variable at both the intramuscle and interanimal level. During swallowing, some muscles subserved multiple functions and contained different task units; there were also intramuscle differences in EMG latencies. In this situation, statistical methods were essential to characterize the overall patterns of EMG activity. The statistically derived multimuscle pattern approximated to the classical description by Doty and Bosma (Doty RW, Bosma JF. J Neurophysiol 19: 44-60, 1956) with a leading complex of muscle activities. However, the mylohyoid was not active earlier than other muscles, and the geniohyoid muscle was not part of the leading complex. Some muscles, classically considered inactive, were active during the pharyngeal swallow.
It was proposed that the power stroke in primates has two distinct periods of occlusal contact, each with a characteristic motion of the mandibular molars relative to the maxillary molars. The two movements are called phase I and phase II, and they occur sequentially in that order (Kay and Hiiemae  Am J. Phys. Anthropol. 40:227-256, Kay and Hiiemae  Prosimian Biology, Pittsburgh: University of Pittsburgh Press, p. 501-530). Phase I movement is said to be associated with shearing along a series of crests, producing planar phase I facets and crushing on surfaces on the basins of the molars. Phase I terminates in centric occlusion. Phase II movement is said to be associated with grinding along the same surfaces that were used for crushing at the termination of phase I. Hylander et al. ( Am J. Phys. Anthropol. 72:287-312; see also Hiiemae  Food Acquisition and Processing, London: Academic Press, p. 257-281; Hylander and Crompton  Am J. Phys. Anthropol. 52:239-251,  Arch. Oral. Biol. 31:841-848) analyzed data on macaques and suggested that phase II movement may not be nearly as significant for food breakdown as phase I movement simply because, based on the magnitude of mandibular bone strain patterns, adductor muscle and occlusal forces are likely negligible during movement out of centric occlusion. Our goal is to better understand the functional significance of phase II movement within the broader context of masticatory kinematics during the power stroke. We analyze vertical and transverse mandibular motion and relative activity of the masseter and temporalis muscles during phase I and II movements in Papio anubis. We test whether significant muscle activity and, by inference, occlusal force occurs during phase II movement. We find that during phase II movement, there is negligible force developed in the superficial and deep masseter and the anterior and posterior temporalis muscles. Furthermore, mandibular movements are small during phase II compared to phase I. These results suggest that grinding during phase II movement is of minimal importance for food breakdown, and that most food breakdown on phase II facets occurs primarily at the end of phase I movement (i.e., crushing during phase I movement). We note, however, that depending on the orientation of phase I facets, significant grinding also occurs along phase I facets during phase I.