Publications

1995
AJ Thexton, RZ German, and AW Crompton. 1995. “Latency of the jaw-opening reflex in the premature, preweaning and adolescent pig (hanford strain miniature pig, Sus scrofa).” Arch Oral BiolArchives of Oral Biology, 40, Pp. 1133-5.Abstract
The jaw-opening (digastric) reflex was elicited by electrical stimulation of oral mucosa in miniature pigs (Sus scrofa) varying in age from 5 days premature to 101 days post-term. The latency of reflex electromyographic activity varied between 12-14 ms in the most immature animals and 9-11 ms in the oldest animals. The very long-latency digastric responses found in the immature young of nesting mammals were not seen in the relatively precocious young of this species.
AW Crompton. 1995. “Masticatory function in nonmammalian cynodonts and early mammals.” In Functional Morphology in Vertebrate Paleontology, edited by J Thomason. Cambridge: Cambridge University Press.
RZ German, AW Crompton, and AJ Thexton. 1995. “Rhythmic response to milk delivery during mammalian suckling.” American ZoologistAmerican Zoologist, 35, Pp. 59A.
AW Crompton and DE Lieberman. 1995. “The role of a mobile mandibular symphysis in mammals (abstract).” Edited by RG Radlanski and Herbert Renz. Proceedings of the 10th International Symposium on Dental Morphology.
D. Stern and A. W. Crompton. 1995. “A study of enamel organization, from reptiles to mammals.” In Aspects of Dental Biology: Paleontology, Anthropology and Evolution, edited by J Moggi-Cecchi, Pp. 1-25. Florence: International Institute for the Study of Man. stern_crompton_enamelorg.pdf
1994
AW Crompton, CB Wood, and DN Stern. 1994. “Differential wear of enamel: a mechanism for maintaining sharp cutting edges.” In Biomechanics of feeding in Vertebrates, edited by VL Bels, Chardon M, and P Vandewalle, 1st ed., 18: Pp. 321-346. Belgium: Springer-Verlag.
RZ German and AW Crompton. 1994. “Integration of swallowing and respiration in infant mammals.” Journal of MorphologyJournal of Morphology, 220, Pp. 348.
JF Bonaparte and AW Crompton. 1994. “A juvenile propainognathid cynodont skull from Ischigualasto formation and the origin of mammals.” Revista del Museo de Ciencias NaturalesRevista del Museo de Ciencias Naturales, 5, Pp. 1-12.
Karen Hiiemae, JB Palmer, and AW Crompton. 1994. “Tongue, palate, hyoid and the capacity for speech.” American Journal of Physical AnthropologyAmerican Journal of Physical Anthropology, 0(Suppl 18), Pp. 107.
Z. X. Luo and A. W. Crompton. 1994. “Transformation of the Quadrate (Incus) through the Transition from Nonmammalian Cynodonts to Mammals.” J Vert Paleo, 14, Pp. 341-374.Abstract

The quadrate (incus) bone underwent important evolutionary transformations through the cynodont-mammal transition. The following character transformations played crucial roles in modifying the cynodont quadrate into the mammalian incus: 1) progressively greater rotation of the dorsal plate relative to the trochlea; 2) the contact facet of the dorsal plate becomes concave; 3) development of a constricted neck between the dorsal plate and the trochlea; 4) simplification of the quadrate-cranium joint, resulting in better mobility of the joint; and 5) introduction of a stapedial process (crus longum). The dorsal plate rotation, the concave contact facet, the constricted neck, the mobile joint of the quadrate and the cranium are also present in some advanced non-mammalian cynodonts. Broad phylogenetic distributions of these features suggest that the major features of the incus of early mammals, as represented by Morganucodon, originated much earlier in phylogenetic history among non-mammalian cynodonts. Apomorphies of the quadrate (incus) among the advanced non-mammalian cynodonts favor a sister-group relationship of tritheledontids and mammals. The hypothesis on the postdentary origin of the mammalian tympanic membrane is favored by transformation of the quadrate through the cynodont-mammal transition. Three most important modifications of the quadrate (incus) through the cynodont-mammal transition are: formation of the concave contact facet, progressively greater rotation of the dorsal plate, and decrease in the number of cranial bones articulating with the quadrate. These modifications would simplify the quadrate-cranial joint and increase the mobility of the quadrate (incus) relative to the cranium while a functioning tympanic membrane was maintained on the mandible, improving the sensitivity of the postdentary tympanum. Probainognathus is among the earliest known non-mammalian cynodonts with a concave contact facet and a rotated dorsal plate in the quadrate. Thus we hypothesize that it represents a critical step in the phylogenetic transformation that led to the origin of the modern mammalian middle ear and tympanic membrane.

TransformationQuadrate1994.pdf
1993
AW Crompton. 1993. “Bone remodeling in response to naturally occurring mechanical stress.” American ZoologistAmerican Zoologist, 33, Pp. 103A.
AW Crompton and ZX Luo. 1993. “Relationships of the Liassic mammals, Sinoconodon, Morganucodonoehleri and Dinnetherium.” In Phylogeny of Mammals, edited by Frederick S Szalay, Michael J Novacek, and Malcolm C McKenna, Pp. 4-44. New York: Springer-Verlag.
1992
JB Palmer, NJ Rudin, G Lara, and AW Crompton. 1992. “Coordination of mastication and swallowing.” DysphagiaDysphagia, 7, Pp. 187-200.Abstract
The coordination of mastication, oral transport, and swallowing was examined during intake of solids and liquids in four normal subjects. Videofluorography (VFG) and electromyography (EMG) were recorded simultaneously while subjects consumed barium-impregnated foods. Intramuscular electrodes were inserted in the masseter, suprahyoid, and infrahyoid muscles. Ninety-four swallows were analyzed frame-by-frame for timing of bolus transport, swallowing, and phases of the masticatory gape cycle. Barium entered the pharynx a mean of 1.1 s (range -0.3 to 6.4 s) before swallow onset. This interval varied significantly among foods and was shortest for liquids. A bolus of food reached the valleculae prior to swallow onset in 37% of sequences, but most of the food was in the oral cavity at the onset of swallowing. Nearly all swallows started during the intercuspal (minimum gape) phase of the masticatory cycle. Selected sequences were analyzed further by computer, using an analog-to-digital convertor (for EMG) and frame grabber (for VFG). When subjects chewed solid food, there were loosley linked cycles of jaw and hyoid motion. A preswallow bolus of chewed food was transported from the oral cavity to the oropharynx by protraction (movement forward and upward) of the tongue and hyoid bone. The tongue compressed the food against the palate and squeezed a portion into the pharynx one or more cycles prior to swallowing. This protraction was produced by contraction of the geniohyoid and anterior digastric muscles, and occurred during the intercuspal (minimum gape) and opening phases of the masticatory cycle. The mechanism of preswallow transport was highly similar to the oral phase of swallowing. Alternation of jaw adductor and abductor activity during mastication provided a framework for integration of chewing, transport, and swallowing.
RZ German, AW Crompton, LC Levitch, and AJ Thexton. 1992. “The mechanism of suckling in two species of infant mammal: miniature pigs and long-tailed macaques.” J Exp ZoolJournal of Experimental Zoology, 261, Pp. 322-30.Abstract
Suckling is the form of feeding unique to infant mammals. The mechanism used by infant mammals to withdraw liquid from the nipple is the subject of considerable debate. Suckling has been examined in two species of infant mammals: miniature pigs and long-tailed macaques. In both species radio-opaque markers were inserted into the tongue and jaws; the movements of the jaw and tongue (and also of specific regions within the tongue) plus the movement of milk containing barium were studied by high-speed cineradiography (100 and 150 frames/sec). In the case of macaques, simultaneous pressure transducer recordings were also made. In both species, liquid moved out of the nipple as the intraoral space was expanded by a combination of tongue movement (negative pressure pumping) coupled with jaw opening. There was no evidence for expression (positive pressure on the nipple) in either species, strongly supporting the view that a suction mechanism is responsible for acquisition of milk from the nipple. Subsequent intraoral transport was different in the two species. The pigs used a second pump mechanism at the base of the tongue to transport liquid through the pillars of the fauces into the valleculae. The monkeys used a "squeeze-back" mechanism similar to the transport mechanism documented for adult macaques. Further work with other species can test our tentative hypothesis that all mammals use a negative pressure suction for acquisition, but, as is true for adult mammals, infants may use different transport mechanisms to form and move the bolus.
WL Hylander, KR Johnson, and AW Crompton. 1992. “Muscle force recruitment and biomechanical modeling: an analysis of masseter muscle function during mastication in Macaca fascicularis.” Am J Phys AnthropolAmerican Journal Physical Anthropology, 88, Pp. 365-87.Abstract
The main purpose of this study is to test the hypothesis that as subjects chew with increasing levels of force, the ratio of the working- to balancing-side jaw-muscle force (W/B) decreases and begins to approach 1.0. We did this by analyzing relative masseter force in Macaca fascicularis using both strain gage and surface electromyographic (EMG) techniques. In addition, we also analyzed: 1) the relationship between jaw position using cineradiographic techniques and relative masseter force, 2) the timing differences between relative masseter force from the working and balancing sides, and 3) the loading and unloading characteristics of the masseter muscle. Our findings indicate that when macaques increase the amount of overall masticatory force during chewing, the W/B ratio for masseter force frequently (but not always) decreases and begins to approach 1.0. Therefore, our working hypothesis is not completely supported because the W/B ratio does not decrease with increasing levels of force in all subjects. The data also demonstrate timing differences in masseter force. During apple-skin mastication, the average peak masseter force on the working side occurs immediately at or slightly after the initial occurrence of maximum intercuspation, whereas the average peak masseter force on the balancing side occurs well before maximum intercuspation. On average, we found that peak force from the balancing-side masseter precedes the working-side masseter by about 26 msec. The greater the asynchrony between working- and balancing-side masseter force, the greater the difference in the relative magnitude of these forces. For example, in the subject with the greatest asynchrony, the balancing-side masseter had already fallen to about one-half of peak force when the working-side masseter reached peak force. Our data also indicate that the loading and unloading characteristics of the masseter differ between the working and balancing sides. Loading (from 50 to 100% of peak force) and unloading (from 100 to 50% of peak force) for the balancing-side masseter tends to be rather symmetrical. In contrast, the working-side masseter takes much longer to load from 50 to 100% of peak force than it does to unload from 100 to 50% of peak force. Finally, it takes on average about 35 msec for the working-side zygoma and 42 msec for the balancing-side zygoma to unload from 100 to 50% of peak force during apple-skin mastication, indicating that the unloading characteristics of the macaque masseter during mastication closely approximates its relaxation characteristics (as determined by muscle stimulation).
1991
RZ German, C Suarez, and AW Crompton. 1991. “Coordination of Respiration and Swallowing in Infant Macaques (abstract),” 31, Pp. A19.
N. H. Shubin, A. W. Crompton, H. D. Sues, and P. E. Olsen. 1991. “New Fossil Evidence on the Sister-Group of Mammals and Early Mesozoic Faunal Distributions.” Science, 251, Pp. 1063-1065.Abstract

Newly discovered remains of highly advanced mammal-like reptiles (Cynodontia: Tritheledontidae) from the Early Jurassic of Nova Scotia, Canada, have revealed that aspects of the characteristic mammalian occlusal pattern are primitive. Mammals and tritheledontids share an homologous pattern of occlusion that is not seen in other cynodonts. The new tritheledontids represent the first definite record of this family from North America. The extreme similarity of North American and African tritheledontids supports the hypothesis that the global distribution of terrestrial tetrapods was homogeneous in the Early Jurassic. This Early Jurassic cosmopolitanism represents the continuation of a trend toward increased global homogeneity among terrestrial tetrapod communities that began in the late Paleozoic.

shubinsistergroup.pdf
1990
RZ German, AW Crompton, and AJ Thexton. 1990. “Biomechanics of Suckling in Macaca fascicularis (abstract).” Am Assoc Phys AnthroAmerican Association of Physical Anthropology, 81, Pp. 227.
AW Crompton and J Weijs-Boot. 1990. “Intra-oral food transport and swallowing in the American opossum. Part 1: Liquids.” Journal of MorphologyJournal of Morphology, in review.
RZ German, AW Crompton, and SL Filan. 1990. “Ontogeny of Feeding in Opossums (abstract).” Am Soc ZoolAmerican Society of Zoologists, 30, Pp. A21.

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