Effects of Dietary Consistency on Craniofacial and Occlusal Development in the Rat






Dr. Beecher is Assistant Professor, Department of Anatomy at Wright State University in Dayton, Ohio. He is a graduate of the University of Virginia, and holds a Ph.D. degree in anthropology from Duke University. Dr. Corruccini is Assistant Professor, Department of Anthropology, Southern Illinois University in Carbondale. He is a graduate of the University of Colorado, and holds a Ph.D. degree in anthropology and paleontology from the University of California at Berkeley. Address: Dr. Robert M. Beecher Department of Anatomy Wright State University Medicine Dayton, OH 45435

Moderate differences in the hardness of diet are related to significant differences in maxillary width and other measures of facial size. Probably even more important is a relationship to the coordination of growth of different parts of the dentofacial complex. Muscular stimulation mediated through occlusal function seems to play a significant role in the coordinated development of facial structures.

School of

This study was supported by a Biomedical Research Support Grant from Wright State University.

The prevalence of malocclusions in an urbanized society such as the United States is so high (50%1) that one is led to suspect the involvement of environmental factors in the etiology. Non-western, hunter-gatherer peoples are characterized by much lower frequencies of malocclusion." However, the transition of peoples from rural or aboriginal to urban or industrial living has been rapidly followed by an increase in the incidence of malocclusion. This increase is found whether comparing medieval Europeans to recent ones--- or nonwestern peoples to their immediate descendents living in a westernized environmen t. 2,5-15 There is a strong genetic factor in the development of occlusion. Occlusal patterns within families have been demonstrated to be heritable to some degree,16,17 and ethnic groups have been characterized by the frequencies of particular occlusal patterns.18-21 61

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especially the physical consistency of the diet. we believe that environmental factors. growing animals responded to this stress by skeletal adaptations. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. Similar differences were also found in soft-diet rats. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch. (3) lengthened condylar neck.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. (2) bone resorption of the anterior edges of the glenoid fossa. Based on the available anthropological and experimental evidence. racial outcrossing. abnormally-directed muscle tension. affect the developing masticatory apparatus and create variations in the occlusal phenotype. Schultz= documents that dental crowding is quite rare in primates} and neither any of the specimens he presents nor any of the 400 wild primate dentitions that one of us (RSC) has examined would rate a Treatment Priority Index (TPI) score higher than 2. or accumulation of genetic mutations. mouth breathing is probably a factor in the development of human malocclusions. Experimen Is in effects of function Apart from radical surgery. inbreeding. The maxillae of these animals showed reduced vertical and prognathic growth. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. and (4) varying the consistency of the diet from hard to soft.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. specifically: (1) some posterior bone deposition.vSuch effects are common in humans.sv-" A crown on the maxillary buccal teeth of rhesus monkeys was used by MeNamara-" to force the mandibular molars to occlude slightly lower than the normal occlusal surface of the uppers. (2) placing devices in the mouth to effect alterations in mastication.0 mm in order to achieve molar occlusion. the experimental attempts to induce developmental change in the mammalian masticatory apparatus have used four methods': (I) forcing mouth breathing in animals. and ('1) an increase in the angle formed by a line running from the condyle down the ramus with the line representing the occlusal plane. older animals responded with tooth movements to adjust to the new occlusal alignments.5-2. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . While the younger. A cornman habit in childhood. (3) restricting animals to calorie-deprivation diets. Several workers have applied stress to the masticatory apparatus or have altered occlusal relationships using appliances fastened to the teeth of rats or rhesus monkeys. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. These are usually designed to force protrusion of the mandible 1.

Beecher and Corruccini35 examined a small population of rhesus macaques which had spent a short period during adolescence on contrasting hard/soft diets.28. Green. in older animals. (5) had smaller masseter and temporalis muscles. While these experiments are important in assessing the response of the masticatory system to unusual mechanical stress. with condyles smaller and radiographically less dense. 1981 Development 63 four studies have examined this idea experimentally. (2) exhibited no molar wear. Nutrition Experiments concerning the contribution of nutrition to normal occlusal development have taken the form of calorie deprivation of pigs. Sagne and Thilander. and (6) had skulls consistently smaller in mass and in linear dimensions. Dietary consistency Although many workers have commented on the probable influence of the hardness or softness of the diet on the developing masticatory system. the soft-diet animals: (I) were slightly smaller in body mass. was less in the soft-diet animals. especially in the mandible. which responded with less deposition and growth. The authors found that their regime resulted in: (I) delayed dental development and eruption. While calorie deprivation is undoubtedly a factor to consider in severely undernourished people.29 McCance. and COX. More recently." Barber. Even buccolingual thickness of the mandible. Attachment areas for the jawclosing muscles. They . 1 January. were smaller. The smaller and less-active muscles applied less stress to the bones. In young animals. Tonge and McCance. Owens and Tonge= used weanling animals. which is essential to resist transverse bending.used weanling rats divided into populations eating either pelle ted rat chow as a hard diet or crushed or water-softened chow as a soft diet. the growing skeletal system adjusts. All mandibular structures related to chewing were affected by diet hardness. When compared to the harddiet rats. Watt and Williams. (4) had less width of the maxillary dental arch. they do not create conditions likely to be encountered by the developing human masticatory system." and Henrikson. The condyles. before attempting to "rehabilitate" some with proper foods. was less. These studies demonstrate that forced realignments of occlusal relationships and mandibular movements lead to responses by the most flexible elements of the masticatory apparatus. there is no evidence to correlate this deficiency with the occlusal problems found in so many urbanized peoples of the western world.Diet Consistency jOcclusal the teeth adapted and once again occluded adequately. Vol. were smaller. the dentition migrates to accommodate. Wear of the molar.32 Moore. where muscle force is transmitted to the mandible. where toothfood-tooth contact takes place. In all studies. 51 No. which resist bite (especially incisal) reaction force. maintaining them for one year on a severe calorie-deficient diet. and (2) "greater delay in the development and growth of the jaws. the animals were maintained for approximately four months. The smaller masticatory muscles in soft-diet animals were almost certainly a result of lower work requirement in mastication. although with no significant differences in shape. Rehabilitation was only partiall y successful. (3) had mandibles that were smaller."29 The consequences were tooth crowding and abnormal tooth-tooth contacts.

incisor to Ml. 2. Maxillary arch length. 6. resulting in a lj uantum change in the morphology of the masticatory system.he correlations among the several variables. or whether no differences should be expected until a threshold in dietary consistency difference was reached. The three populations were maintained on their respective diets for four months. finds the major axes of the between-variable correlation matrix. it could not be determined whether the masticatory system responded in direct proportion to differences in dietary consistency. Body mass. Anteroposterior length of condylar articular surface. Projection of indiThe Angle Orthodontist found significant narrowing of the maxillary arch with no lessening in length in the soft-diet monkeys. After sacrifice. We have no way of knowing the consistency of rat diets used in earlier experiments of this type. evenly divided between males and females.ors (also known as directional cosines or latent vectors) summarize the multiple interplay among variables. Measurements Crable I) were taken as follows: 1. the heads were randomly numbered so that the measurer would not be aware of their group membership. MATERIALS AND METHODS The experimental animals were Sprague-Dawley rats. Ninety animals were divided into three groups: Group I was fed pelletted rat chow (Purina Formulab). The second axis is the axis perpendicular to the first that subsumes the largest possible amount of residual variance. there has been no attempt to integrate these data in such a way that interactions between different parts of the growing masticatory systems could be measured. Fresh mass of the entire masseter. This indicates that the "hard" diet was minimally harder . hard and soft diet. This technique. A principal components analysis was performed on the data in order to discover t. Croup 11 was fed a gruel-like porridge consisting of ground chow moistened with water. Mandibular length. Group II I was fed the soft diet six days. This correlated with the histological findings by Bouvier and Hylander-" that significantly fewer secondary Haversian systems were present in the mandibular corpus of soft-diet monkeys than in the hard-diet monkeys of the same population. incisor to distal edge of last molar. acquired at 21 days of age. 4. Further. We found the lab chow to be crumbly. easily breaking down into small granules. with dry pellets provided every seventh day only. The eigenvect. It was to fill these gaps in the experimental record that the following study was carried out. and so on.64 Beecher and Corruccini than the gruel. Maxillary arch breadth across buccal points of ?vII. related to factor analysis. 3. 5. Least-squares theory is used to rotate orthogonal axes to a position where the major axis subsumes a maximized amount of the total variance. with only two groups. 'While a number of measurements have been taken from animals in dietary consistency experiments. the same length of time used in previous experiments of this type. Animals were also examined for visible differences in tooth alignments and attritional differences between the three groups. engendering only subtle differences in bite force.

Variable Maxillary Length (mm) .39 8. RESULTS Development 65 more variable than Group I in the maxillary arch measurements.92 3.89 3. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II). significant < .38) ( 0. All correlations are higher in the animals that had more masticatory resistance.26 38.d.d. Condyle Length Body weight (gm) . The gross differences in the growth of most structures are not very marked..Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples..65) ( 0.44 at p (0. Thus the residual from Mean.f.d.64 8. greater variation away [rom a major principal components axis describing general growth could be interpreted as meaning that there is poorer coordination of regional growth rates in those individuals.40) ( 0. 1981 .00 9. but usually closer to Group I. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I)..79 9. signifying that all structures are more tightly integrated in their growth and covariation..41) (12. Masseter weight (gm) . The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix.45 8.. In particular. Table 3 lists key paramo eters of this analysis..17) ( 0.18 lAO Intermittent Mean (s.58) ( 0.21' 1. 51 No. but not significantly so.43 ( 1. For instance. indicating a greater range of sizes.. Vol. Dispersion. although more similar to the soft group. • 3-sample TABLE I and Significance of Difference Hard Mean (s.91 3. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals. Again. Group III is intermediate in standard deviation. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance. The strongest difference is in the growth relation of the masseter muscle. Breakdown in growth coordination Mcxillary breadth affected most The basic descriptive statistics in Table I show the hard-diet population (Group 1) to be larger in all dimensions and disproportionately so in some.) 23.48) Soft Mean (s.~6) (0.28) (or Measured Traits F 0.31 39.05.39) (11. and functional interpretation of the morphological integration among variables. Mandibular Length (mm) (mm) .7 I 9.) ~:I. and the differences are often significant.50) (0. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples.32) (0.1 January.88) (0. Maxillary Breadth (mm) ..20) ( 0..43 43.38 2.03 1.. the second axis is largest among the softdiet animals.01) lAO ( 0.46) ( 0. The major (coordinated growth) axis is largest in the hard-diet sample.56 ( 1. Group I is more variable than Group II in every measurement. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures.) 24.72 1.36) (9. a result attributable to the small differences in dietary consistency. Inversely related to this.57 4.40) om AoV with 2 and 87 d.

the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2. ... Mandibular Length .83· 0. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child.75· 0.....37..66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation. No caries were observed by us in any animals.38 Attrition not affected the major growth aXIS (which could be interpreted as error in growth or variation in growth rate) is easily largest in the soft diet sample......05.L 0..... This increased multivariate variability in soft-diet animals is all the more striking in view of the fact that they were less variable than the hardfood animals in every univariate trait. Thus.. a decrease in correlation of masticatory structures is more noteworthy than de- = '...93· 0."] ""hile they noted easily observable differences in attrition between soft and hard diet groups. Perhaps a decrease in buffering among the growing components is the first step in their decrease in size.. TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max.85 Groups Mussel.....60 0. The Angle Orthodontist . we could find none. This phenomenon has also been noted in characters that are diminishing in size through natural selection..55 0..79· 0..79· 0.. Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes.. which is significantly greater than expected by random error at P 2. and masseter weight.72 Condo 0. We noticed no displaced or rotated teeth in any of the animals.H3 0....74 0. In all respects.. Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals.54 0..differentiation represented by this difference in dietary consistency.71 Soft diet ~ r in hard < .. . condyle length.89 0.19. DISCUSSION Present-day insults to occlusion Oppenheimer" and Lavell-v document the better occlusion that existed in human populations prior to the onset of the industrial revolution. Condyle length alone shows a notable residual on the second axis in Group I and III.. Body weight .5%.78 0. at the stage of growrh. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth.. IMaxillary Length . • Significantly larger Max. ~ e: 0... Masseter weight .91· 0. Ii Mand...Ve examined the occlusal surfaces of the rat molars as a further test of the comparability of our hard chow and that used thirty years ago by Watt and Williams. 0.39 diet sample at p :c . Maxillary Breadth . Condyle Length . the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample.66 Body 0...

Because of its widespread occurrence in western society.90 0. especially of anterior teeth. premature deciduous tooth loss.50 (8) -0. the mandible may depend on muscular function to grow to average size. the reduction of chewing resistance and related growth stimulation provided by "civilized" man's refined diet may be the strongest causal factor in developmental malocclusions.12 0.67 (78) 0.96 (83) 0.92 0.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion.95 (variation in growth) Soft Intermittent 0.97 0.96 0.05 -0.68 0.06 -0.08 -0.57 -0. trauma.60 (10) -0. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.w Moorrees and Reed.80 One Principal Component Hard Two growth) Intermittent 4.41•42 Some of these may be of minimal or no importance.:" Maxillary growth may be under closer genetic programming.04 0.41 -0.94 0. Thus. 51 No.89 0. including mouth breathing.06 -0. 1981 .17 (69) 0.93 0.10 Vol.08 0. In particular.82 0. but does not give the source of the growth-correlation breakdown.18 -0.73 -0. considerable evidence ties upper-lower correlation diminution with development of mal- TABLE 3 Eigen Values (latent roots) and Eigenvectors (latent vectors) of the Correlation Matrices in Three Dietary Groups Principal Component (coordinated Hard Soft EigenValue ('70 variance) Eigenvector correlations: Maxillary Length Maxillary Breadth Mandibular Length Condyle Length Body weight Masseter weight 4. follows different rates and peaks at different ages in the maxilla and mandible. hormonal intervention. disturbance of synchronous development in separately growing parts. posture. Though the chewing explanation is most predictive of observed occlusal differences in the rats. and independence of tooth formation. Tooth spacing and crowding.59 -0.89 0.95 4.29 0.14 0. oral habits. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles.91 0.16 -0. experimental evidence which might weight such factors is absent.90 0.23 0. 1 January.79 (13) -0." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control.80 0. The aging process also yields different results in upper and lower arches. with the maxillary teeth being more conservative and controlled by a fewer number of genes.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion. endocrine disturbance.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.68 0. prenatal insult.51 0.77 0.42-45 The growth independence of mandible and maxilla in humans has been frequently noted.

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