Association of relatively delayed emergence of mandibular molars with molar reduction and molar position

Among 234 children examined annually from age three to 20 years at the Burlington Growth Centre, there was statistically significant cooccurrence of early and late emergence sequences of the permanent first and second molars relative to the central incisors and second premolars in the same jaw and in both jaws. Alternatively, mandibular molar delay was not accompanied by corresponding maxillary molar delay, and the mandibular molars emerged later than the maxillary molars. This was strongly associated with Angle Class II malocclusion, indicating a relationship between relative time of emergence and relative position of opposing molars. Delay of the mandibular molar relative to the successional teeth or maxillary molars was associated with increased frequency of four cusped first and second molars and agenesis of third molars, indicating a tendency for co-occurrence of delay in timing of molar emergence with reduction in structure of the molars. These relationships were evident even though emergences were affected by early loss of a deciduous second molar which increased M1I1 and M2P2 sequences by earlier emergence of M1 and delayed emergence of P2.     

Interspecific allometry of the mandible,dental arch,and molar area in anthropoid primates: Functional morphology of masticatory components

Allometric equations relating the lengths and widths of the mandible and dental arch, and of molar area, were obtained in a wide range of anthropoid primates grouped into four subsets, pongids, cercopithecids, nonmarmoset platyrrhines, and marmosets. Mandibular width is negatively allometric against length across anthropoids but cercopithecids had relatively wider mandibles than nonmarmosets of the same size class. Mandibular length relative to dental arch length was isometric within and between the four groups but dental arch width scaled negatively against all the other dimensions examined in this study, indicating a functional dissociation between the dental arcade and the bony mandible. Molar area showed various scaling patterns relative to mandibular length (isometry) and width (positive). There were no parameters that scaled positively against body weight across groups, except for molar area in cercopithecids (strongly) and nonmarmoset (moderately). Notable functional specializations include relatively long dental arches in cercopithecoids, related to large, elongate bilophodont molars, and the tendency to increase relative jaw length across the range of anthropoid sizes, reflecting negative allometry of the brain (cranial bicondylar width). We caution that various allometry and functional patterns may be masked by generalizing from broad taxonomic comparison involving a large sweep of adaptative patterns.     

A cinefluorographic study of mandibular movement during feeding in the rat (Rattus norvegicus)

The anatomy of the masticatory apparatus, and particularly of the mandibular joints, has led to the view that mandibular movement in the Rodentia is predominantly propalinal, or forwards and backwards in direction. As part of an investigation into the mechanism of function of the mandibular joints in these animals, the feeding behaviour of "August" strain and "Wistar" rats was examined by cinephotography and cinefluorography. The rats were trained to feed on barium sulphate impregnated biscuit and animal cake and to drink radio-opaque liquids. Cinefluorographic recordings of ingestion, mastication, deglutition and of drinking were taken in both the lateral and dorso-ventral projections.
Analysis of the recordings has shown a fundamental separation of ingestive and masticatory activity in the rat, which can be attributed to the morphology of the jaws and particularly to the disparity in the lengths of the mandibular and maxillary diastemas. To bring the incisor teeth into occlusion for ingestion, the mandible is brought forward through the rest position and the condyle into articulation with the anterior part of the fossa. In mastication the condyle is moved backwards to bring the molar teeth into occlusion and the condyle into articulation with the posterior articular facet on the fossa. Once the mandible has been moved into the appropriate position for either ingestion or mastication and deglutition, the movements involved in the separation or chewing of the food are cyclical with combined horizontal and transverse movements as well as the fundamental vertical movement acting to open and close the mouth. The basic movement of ingestion carries the mandibular incisors upwards and forwards across the lingual surfaces of the maxillary incisors, so separating the bite. The grinding stroke of mastication is a horizontal movement carrying the mandibular molars anteriorly across the maxillary.     

Genetic structure of a tribal population, the Yanomama Indians. XIII. Dental microdifferentiation.

Data are presented on the frequency of the following eight dental traits in 635 Yanomama and 65 Makiritare Indians: upper central incisor rotation or winging, shoveling of maxillary incisors, maxillary molar hypocone reduction, Carabelli's trait, mandibular molar cusp number, mandibular molar cusp pattern rotation of second lower premolar, and pattern of second lower premolar cusps. Yanomama dentition is unusual in the high frequency of six cusps on the mandibular molars. There is marked dental microdifferentiation between villages; significant agreement was observed between a matrix of pairwise "dental distances" based on six morphological traits and corresponding matrices based on 11 genetic systems and on geographic location.     

The British dental arch

Thirty-two dimensions of the maxillary and mandibular dental arches were measured on 320 adult skulls derived from Anglo-saxon to modern times. Multivariate analysis showed varying patterns of contrast between the samples, depending upon which arch dimensions were considered. The results suggest that dental arch changes occur independently of jaw changes.     

The role of effectors of the activin signalling pathway, activin receptors IIA and IIB, and Smad2, in patterning of tooth development.

The gene for activin betaA is expressed in the early odontogenic mesenchyme of all murine teeth but mutant mice show a patterning defect where incisors and mandibular molars fail to develop but maxillary molars develop normally. In order to understand why maxillary molar tooth development can proceed in the absence of activin, we have explored the role of mediators of activin signalling in tooth development. Analysis of tooth development in activin receptor II and Smad2 mutants shows that a similar tooth phenotype to activin betaA mutants can be observed. In addition, we identify a novel downstream target of activin signalling, the Iroquois-related homeobox gene, Irx1, and show that its expression in activin betaA mutant embryos is lost in all tooth germs, including the maxillary molars. These results strongly suggest that other transforming growth factor beta molecules are not stimulating the activin signalling pathway in the absence of activin. This was confirmed by a non-genetic approach using exogenous soluble receptors to inhibit all activin signalling in tooth development, which reproduced the genetic phenotypes. Activin, thus, has an essential role in early development of incisor and mandibular molar teeth but this pathway is not required for development of maxillary molars.     

Electromyography and mechanics of mastication in the albino rat.

The masticatory apparatus in the albino rat was studied by means of electromyography and subsequent estimation of muscular forces. The activity patterns of the trigeminal and suprahyoid musculature and the mandibular movements were recorded simultaneously during feeding. The relative forces of the individual muscles in the different stages of chewing cycles and biting were estimated on the basis of their physiological cross sections and their activity levels, as measured from integrated electromyograms. Workinglines and moment arms of these muscles were determined for different jaw positions. In the anteriorly directed masticatory grinding stroke the resultants of the muscle forces at each side are identical; they direct anteriorly, dorsally and slightly lingually and pass along the lateral side of the second molar. Almost the entire muscular resultant force is transmitted to the molars while the temporo-mandibular joint remains unloaded. A small transverse force, produced by the tense symphyseal cruciate ligaments balances the couple of muscle resultant and molar reaction force in the transverse plane. After each grinding stroke the mandible is repositioned for the next stroke by the overlapping actions of three muscle groups: the pterygoids and suprahyoids produce depression and forward shift, the suprahyoids and temporal backward shift and elevation of the mandible while the subsequent co-operation of the temporal and masseter causes final closure of the mouth and starting of the forward grinding movement. All muscles act in a bilaterally symmetrical fashion. The pterygoids contract more strongly, the masseter more weakly during biting than during chewing. The wide gape shifts the resultant of the muscle forces more vertically and moreposteriorly. The joint then becomes strongly loaded because the reaction forces are applied far anteriorly on the incisors. The charateristic angle between the almost horizontal biting force and the surface of the food pellet indicates that the lower incisors produce a chisel-like action. Tooth structure reflects chewing and biting forces. The transverse molar lamellae lie about parallel to the chewing forces whereas perpendicular loading of the occlusal surfaces is achieved by their inclination in the transverse plane. The incisors are loaded approximately parallel to their longitudinal axis, placement that avoids bending forces during biting. It is suggested that a predominantly protrusive musculature favors the effective force transmission to the lower incisors, required for gnawing. By grinding food across transversely oriented molar ridges the protrusive components of the muscles would be utilized best. From the relative weights of the masticatory muscles in their topographical relations with joints, molars and incisors it may be concluded that the masticatory apparatus is a construction adapted to optimal transmission of force from muscles to teeth.     

Morphological features of Jat dentition

Observations on morphological characters of milk and permanent teeth, based on 648 pairs of dental casts of 356 male and 292 female Jat children of Haryana (India) are reported. Deciduous teeth show high frequencies of bilateral winging of maxillary central incisor, Carabelli's cusp of maxillary second molar, and deflecting wrinkle of mandibular second molar. Reduction of maxillary molar cusps is more marked in males than in females. Y pattern is very common in deciduous molars. Permanent teeth have high frequencies of grooved cingulum of incisors, cingular nodule of lateral incisors and canines, and distal accessary ridge of canines. Low frequencies of Carabelli's cusp and winging are also common. The tendency towards faintly developed shovelling in milk incisors occurs more often than in the permanent teeth.     

Development of the helicoidal plane

In this study of thebelicoidal occlusal plane the relationships between tooth wear, the transverse slopes of mandibular molars and dental arch breadths were examined in 74 pre-contemporary Australian Aboriginal skulls. With increasing age and tooth wear the orientation of the mandibular occlusal surfaces increased towards the buccal. The differential occlusal orientation from first to third molars, present at eruption, tended to increase progressively with tooth wear. These functionally induced changes, together with regional differences in relative breadths of the maxillary and mandibular dental arches, are important in the development of abelicoidal occlusal plane.     

Incremental enamel development in modern human deciduous anterior teeth

This study reconstructs incremental enamel development for a sample of modern human deciduous mandibular (n = 42) and maxillary (n = 42) anterior (incisors and canines) teeth. Results are compared between anterior teeth, and with previous research for deciduous molars (Mahoney: Am J Phys Anthropol 144 (2011) 204-214) to identify developmental differences along the tooth row. Two hypotheses are tested: Retzius line periodicity will remain constant in teeth from the same jaw and range from 6 to 12 days among individuals, as in human permanent teeth; daily enamel secretion rates (DSRs) will not vary between deciduous teeth, as in some human permanent tooth types. A further aim is to search for links between deciduous incremental enamel development and the previously reported eruptionsequence. Retzius line periodicity in anterior teeth ranged between 5 and 6 days, but did not differ between an incisor and molar of one individual. Intradian line periodicity was 12 h. Mean cuspal DSRs varied slightly between equivalent regions along the tooth row. Mandibular incisors initiated enamel formation first, had the fastest mean DSRs, the greatest prenatal formation time, and based upon prior studies are the first deciduous tooth to erupt. Relatively rapid development in mandibular incisors in advance of early eruption may explain some of the variation in DSRs along the tooth row that cannot be explained by birth. Links between DSRs, enamel initiation times, and the deciduous eruption sequence are proposed. Anterior crown formation times presented here can contribute toward human infant age-at-death estimates. Regression equations for reconstructing formation time in worn incisors are given.     

Postural stability of the human mandible during locomotion

Movements of the head and of the mandible relative to the head were measured in human subjects walking and running on a treadmill at various speeds and inclinations. A miniature magnet and piezo-electric accelerometer assembly was mounted on the mandibular incisors, and a Hall-effect sensor along with a second accelerometer mounted on a maxillary incisor along a common vertical axis. Signals from these sensors provided continuous records of vertical head and mandible acceleration, and relative jaw position. Landing on the heel or on the toe in different forms of locomotion was followed by rapid deceleration of the downward movement of the head and slightly less rapid deceleration of the downward movement of the mandible, i.e., the mandible moved downwards relative to the maxilla, then upwards again to near its normal posture within 200 ms. No tooth contact occurred in any forms of gait at any inclination. The movement of the mandible relative to the maxilla depended on the nature and velocity of the locomotion and their effects on head deceleration. The least deceleration and hence mandibular displacement occurred during toe-landing, for example, during "uphill" running. The maximum displacement of the mandible relative to the head was less than 1mm, even at the fastest running speed. The mechanisms that limit the vertical movements of the jaw within such a narrow range are not known, but are likely to include passive soft-tissue visco-elasticity and stretch reflexes in the jaw-closing muscles.     

On the maxillary nerve

The trigeminal, the fifth cranial nerve of vertebrates, represents the rostralmost component of the nerves assigned to pharyngeal arches. It consists of the ophthalmic and maxillomandibular nerves, and in jawed vertebrates, the latter is further divided into two major branches dorsoventrally. Of these, the dorsal one is called the maxillary nerve because it predominantly innervates the upper jaw, as seen in the human anatomy. However, developmentally, the upper jaw is derived not only from the dorsal part of the mandibular arch, but also from the premandibular primordium: the medial nasal prominence rostral to the mandibular arch domain. The latter component forms the premaxillary region of the upper jaw in mammals. Thus, there is an apparent discrepancy between the morphological trigeminal innervation pattern and the developmental derivation of the gnathostome upper jaw. To reconcile this, we compared the embryonic developmental patterns of the trigeminal nerve in a variety of gnathostome species. With the exception of the diapsid species studied, we found that the maxillary nerve issues a branch (nasopalatine nerve in human) that innervates the medial nasal prominence derivatives. Because the trigeminal nerve in cyclostomes also possesses a similar branch, we conclude that the vertebrate maxillomandibular nerve primarily has had a premandibular branch as its dorsal element. The presence of this branch would thus represent the plesiomorphic condition for the gnathostomes, implying its secondary loss within some lineages. The branch for the maxillary process, more appropriately called the palatoquadrate component of the maxillary nerve (V₂), represents the apomorphic gnathostome trait that has evolved in association with the acquisition of an upper jaw. J. Morphol. 275:17–38, 2014. © 2013 Wiley Periodicals, Inc.     

Superficial,suprahyoid, and infrahyoid neck musculature in naked mole-rats (Heterocephalus glaber): Relative size and potential contributions to independent movement of the lower incisors

Naked mole-rats (Heterocephalus glaber) are fossorial, eusocial rodents that exhibit the unusual capability of moving their lower incisors independently in lateral and rostroventral directions. The evolution of this trait would presumably also involve concurrent alterations in neck musculature to support and control movements of the lower incisors. In order to assess morphological adaptations that might facilitate these movements, we performed detailed dissections of the neck musculature of adult naked mole-rats. In addition to characterizing attachment sites of superficial, suprahyoid, and infrahyoid musculature, we also quantified muscle mass and mandibular features thought to be associated with gape (condyle height, condyle length, and jaw length). Based on muscle attachment sites, the platysma myoides may contribute to lateral movement of the lower incisor and hemi-mandible in naked mole-rats. The large digastric muscle is likely to be a main contributor to rostroventral movement of each lower incisor. The geniohyoid and mylohyoid muscles also likely contribute to rostroventral movements of the lower incisors, and the mylohyoid may also produce lateral spreading of the hemi-mandibles. The transverse mandibular (intermandibularis) muscle likely serves to reposition the lower incisors back to a midline orientation following a movement.     

Dental morphology and odontometry of the Tibetan immigrants

Dental features such as size, shape, cusp number and groove pattern, deflecting wrinkle, protostylid, form and size of dental arch and occlusion pattern, as well as crown diameters of maxillary and mandibular dentition of the Tibetan immigrants in India are described. Sex differences in various measurements are also noted. Overall reduction in size, presence of hypodontia of the third molar, and absence of Carabelli's trait are a distinct progressive/evolutionary trend in the Tibetan dentition, while presence of a shovel-shaped lingual surface of central and lateral incisors is a retrogressive/primitive condition. In some of the dental features Tibetans resemble Caucasoid and modern populations, in certain others they resemble other Mongoloid populations, and in still other features they resemble aboriginal populations and fossil hominids.     

Morphoregulation of teeth: modulating the number, size, shape and differentiation by tuning Bmp activity

During development and evolution, the morphology of ectodermal organs can be modulated so that an organism can adapt to different environments. We have proposed that morphoregulation can be achieved by simply tilting the balance of molecular activity. We test the principles by analyzing the effects of partial downregulation of Bmp signaling in oral and dental epithelia of the keratin 14-Noggin transgenic mouse. We observed a wide spectrum of tooth phenotypes. The dental formula changed from 1.0.0.3/1.0.0.3 to 1.0.0.2(1)/1.0.0.0. All mandibular and M3 maxillary molars were selectively lost because of the developmental block at the early bud stage. First and second maxillary molars were reduced in size, exhibited altered crown patterns, and failed to form multiple roots. In these mice, incisors were not transformed into molars. Histogenesis and differentiation of ameloblasts and odontoblasts in molars and incisors were abnormal. Lack of enamel caused misocclusion of incisors, leading to deformation and enlargement in size. Therefore, subtle differences in the level, distribution, and timing of signaling molecules can have major morphoregulatory consequences. Modulation of Bmp signaling exemplifies morphoregulation hypothesis: simple alteration of key signaling pathways can be used to transform a prototypical conical-shaped tooth into one with complex morphology. The involvement of related pathways and the implication of morphoregulation in tooth evolution are discussed.     

Mandibular movement patterns relative to food types in common tree shrews (Tupaia glis)

Although common tree shrews have long been considered a model system for early eutherian mastication, little information on mandibular movement patterns relative to specific food types has been reported. Detailed analysis of mandibular movement patterns when related to resulting attrition facets may permit more accurate extrapolations regarding the dietary habits of primitive mammals. Marker beads were sewn to chins of five animals that were placed in a restraint system and filmed while they fully masticated mealworm larvae and standardized pieces of banana, almond, and commercial cat chow. These sequences were divided into early, middle, and late thirds of food reduction. Mandibular positions from both frontal and lateral perspectives were digitized frame by frame to yield plots of orbits in three dimensions as well as graphic display of displacements, velocities, and accelerations. Plot coordinates were averaged to generate composite orbital shapes. Significant (p < 0.01) findings included: (1) shortest orbital durations and greatest peak closing velocities and accelerations in early third of reduction; (2) smallest maximum gape, smallest maximum lateral excursion from midline, and longest duration of powerstroke relative to orbital duration in late third of reduction; (3) shortest orbital durations and smallest maximum gape during mastication of chow; (4) greatest maximum lateral excursion during mastication of mealworm larvae; and (5) smallest peak closing accelerations during mastication of banana. Significant differences were also found among subjects for all parameters examined. Capacity for complex jaw movement may have been critical for allowing primitive molars to be used for trituration of a variety of food types, and may have preceded evolution of more specialized molar forms.     

Fishing for jaws in early vertebrate evolution: a new hypothesis of mandibular confinement

The evolutionary origin of the vertebrate jaw persists as a deeply puzzling mystery. More than 99% of living vertebrates have jaws, but the evolutionary sequence that ultimately gave rise to this highly successful innovation remains controversial. A synthesis of recent fossil and embryological findings offers a novel solution to this enduring puzzle. The Mandibular Confinement Hypothesis proposes that the jaw evolved via spatial confinement of the mandibular arch (the most anterior pharyngeal arch within which the jaw arose). Fossil and anatomical evidence reveals: (i) the mandibular region was initially extensive and distinct among the pharyngeal arches; and (ii) with spatial confinement, the mandibular arch acquired a common pharyngeal pattern only at the origin of the jaw. The confinement occurred via a shift of a domain boundary that restricted the space the mesenchymal cells of the mandibular arch could occupy. As the surrounding domains replaced mandibular structures at the periphery, this shift allowed neural crest cells and mesodermal mesenchyme of the mandibular arch to acquire patterning programs that operate in the more posterior arches. The mesenchymal population within the mandibular arch was therefore no longer required to differentiate into specialized feeding and ventilation structures, and was remodelled into a jaw. Embryological evidence corroborates that the mandibular arch must be spatially confined for a jaw to develop. This new interpretation suggests neural crest as a key facilitator in correlating elements of the classically recognized vertebrate head ‘segmentation’.     

A study of the taxonomic significance of the dental arch.

Sixteen dimensions were measured from the maxillary and mandibular dental arches of different ethnic groups of man, apes and monkeys. Multivariate analysis showed that discrimination was possible among the ethnic groups of man on the one hand and between the ape and monkey samples on the other. Nevertheless, the actual degree of discrimination between the primate samples depended upon whether the maxillary or mandibular arch dimensions were analysed. Furthermore, subsequent inclusion of the dental arch dimensions of fossil hominoid samples into the analysis confirmed the taxonomic significance of the dental arch, although its importance must await the acquisition of the more non-human primate data.     

Three-dimensional dynamical capabilities of the human masticatory muscles

Many habitual human jaw movements are non-symmetrical. Generally, it is observed that when the lower incisors move to one side the contralateral condyle moves forwards onto the articular eminence, whereas the ipsilateral condyle stays in the mandibular fossa, moving slightly to the ipsilateral side. These jaw movements are the result of contractions of active masticatory muscles and guided by the temporomandibular joints, their ligaments and passive elastic properties of the muscles. It is not known whether the movements are primarily dependent on passive guidance, active muscle control or both. Therefore, the objective of this study was to analyse the interplay between these factors during non-symmetrical jaw movements. A six-degrees-of-freedom dynamical biomechanical model of the human masticatory system was used. The movements were not restricted to a priori defined joint axes. Jaw movement simulations were performed by unilateral activity of the muscles. The ligaments or the passive elastic properties of the muscles could be removed during these simulations. Laterodeviations conform to naturally observed ones could be generated by unilateral muscle contractions. The movement of the lower incisors was hardly affected by the absence of passive elastic muscle properties or temporomandibular ligaments. The latter, however, influenced the movement of the condyles. The movements could be understood by analysing the combination of forces and torques with respect to the centre of gravity of the lower jaw. In addition, the loading of the condyles appeared to be an important determinant for the movement. This analysis emphasizes that the movements of the jaw are primarily dependent on the orientation of the contributing muscles with respect to this centre of gravity and not on the temporomandibular ligaments or passive elastic muscle properties.