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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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. older animals responded with tooth movements to adjust to the new occlusal alignments. and (4) varying the consistency of the diet from hard to soft. we believe that environmental factors. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. the experimental attempts to induce developmental change in the mammalian masticatory apparatus have used four methods': (I) forcing mouth breathing in animals. abnormally-directed muscle tension. (2) bone resorption of the anterior edges of the glenoid fossa. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . or accumulation of genetic mutations.vSuch effects are common in humans. 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. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. The maxillae of these animals showed reduced vertical and prognathic growth. 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. specifically: (1) some posterior bone deposition.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. A cornman habit in childhood. affect the developing masticatory apparatus and create variations in the occlusal phenotype. (3) lengthened condylar neck. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. racial outcrossing. While the younger. especially the physical consistency of the diet. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. These are usually designed to force protrusion of the mandible 1.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. inbreeding. 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) restricting animals to calorie-deprivation diets. Similar differences were also found in soft-diet rats. 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 2.5-2. (2) placing devices in the mouth to effect alterations in mastication. Experimen Is in effects of function Apart from radical surgery. growing animals responded to this stress by skeletal adaptations. mouth breathing is probably a factor in the development of human malocclusions. Based on the available anthropological and experimental evidence. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships.0 mm in order to achieve molar occlusion.

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

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

For instance..) 24..31 39.64 8.d. but usually closer to Group I.. Again. Table 3 lists key paramo eters of this analysis..32) (0. the second axis is largest among the softdiet animals.89 3. Thus the residual from Mean. indicating a greater range of sizes. 1981 .. Mandibular Length (mm) (mm) . and the differences are often significant.46) ( 0. Maxillary Breadth (mm) .56 ( 1.03 1. a result attributable to the small differences in dietary consistency.d. and functional interpretation of the morphological integration among variables.41) (12.57 4.~6) (0.43 ( 1. The strongest difference is in the growth relation of the masseter muscle. significant < .00 9. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance.39 8.) 23.88) (0. The gross differences in the growth of most structures are not very marked. Condyle Length Body weight (gm) . 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. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals.38 2. Inversely related to this. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II)..) ~:I.92 3.44 at p (0.45 8.36) (9. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures..21' 1. • 3-sample TABLE I and Significance of Difference Hard Mean (s.58) ( 0.f. 51 No.91 3. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples.. The major (coordinated growth) axis is largest in the hard-diet sample.01) lAO ( 0.28) (or Measured Traits F 0.7 I 9.38) ( 0. Dispersion..17) ( 0.d.20) ( 0.05.79 9. All correlations are higher in the animals that had more masticatory resistance.18 lAO Intermittent Mean (s. RESULTS Development 65 more variable than Group I in the maxillary arch measurements.48) Soft Mean (s.. Vol. signifying that all structures are more tightly integrated in their growth and covariation. In particular. Group I is more variable than Group II in every measurement.Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples. Variable Maxillary Length (mm) .39) (11.72 1.65) ( 0.50) (0.40) om AoV with 2 and 87 d. 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. although more similar to the soft group.43 43.1 January. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I). but not significantly so. Masseter weight (gm) . Group III is intermediate in standard deviation.40) ( 0.26 38.

and masseter weight... .. No caries were observed by us in any animals.. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child..91· 0.......5%.H3 0...93· 0. Maxillary Breadth .78 0. IMaxillary Length . . Body weight .72 Condo 0. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth..66 Body 0. 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.71 Soft diet ~ r in hard < .... ~ e: 0. condyle length. We noticed no displaced or rotated teeth in any of the animals.... Mandibular Length .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.83· 0. Thus......74 0. Masseter weight . which is significantly greater than expected by random error at P 2.. at the stage of growrh. Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals. we could find none..54 0. Ii Mand."] ""hile they noted easily observable differences in attrition between soft and hard diet groups.79· 0. This phenomenon has also been noted in characters that are diminishing in size through natural selection.66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation.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.85 Groups Mussel.. 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. the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2. Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes. Condyle Length . Condyle length alone shows a notable residual on the second axis in Group I and III...37. • Significantly larger Max. Perhaps a decrease in buffering among the growing components is the first step in their decrease in size.79· 0.. 0. a decrease in correlation of masticatory structures is more noteworthy than de- = '.19.05..... In all respects.60 0..89 0... TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max..75· 0.. the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample.55 0. The Angle Orthodontist ...differentiation represented by this difference in dietary consistency.39 diet sample at p :c .L 0..

89 0.67 (78) 0. and independence of tooth formation.80 0.97 0." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. The aging process also yields different results in upper and lower arches.96 0.06 -0.50 (8) -0. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control. follows different rates and peaks at different ages in the maxilla and mandible.16 -0.41 -0.10 Vol.82 0.18 -0. the mandible may depend on muscular function to grow to average size.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion. 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.93 0.95 4. posture.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion. but does not give the source of the growth-correlation breakdown.29 0.68 0. Thus.80 One Principal Component Hard Two growth) Intermittent 4.05 -0.95 (variation in growth) Soft Intermittent 0.91 0.68 0.59 -0. hormonal intervention.79 (13) -0.89 0.73 -0.77 0.08 0. with the maxillary teeth being more conservative and controlled by a fewer number of genes. prenatal insult. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans. Though the chewing explanation is most predictive of observed occlusal differences in the rats. 1981 . experimental evidence which might weight such factors is absent. disturbance of synchronous development in separately growing parts.51 0. trauma.:" Maxillary growth may be under closer genetic programming.w Moorrees and Reed.42-45 The growth independence of mandible and maxilla in humans has been frequently noted.57 -0.60 (10) -0.96 (83) 0.92 0.90 0.41•42 Some of these may be of minimal or no importance. Because of its widespread occurrence in western society. oral habits.04 0.08 -0.06 -0. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally. 1 January. premature deciduous tooth loss.12 0. including mouth breathing. Tooth spacing and crowding. endocrine disturbance. 51 No. In particular. 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.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.17 (69) 0.90 0.94 0. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles.23 0.14 0. especially of anterior teeth.

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