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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vSuch effects are common in humans. (2) placing devices in the mouth to effect alterations in mastication. inbreeding. Based on the available anthropological and experimental evidence. The maxillae of these animals showed reduced vertical and prognathic growth. Experimen Is in effects of function Apart from radical surgery. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures.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. 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. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. affect the developing masticatory apparatus and create variations in the occlusal phenotype. While the younger. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. A cornman habit in childhood. we believe that environmental factors.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. These are usually designed to force protrusion of the mandible 1. and (4) varying the consistency of the diet from hard to soft.0 mm in order to achieve molar occlusion. Similar differences were also found in soft-diet rats. or accumulation of genetic mutations. mouth breathing is probably a factor in the development of human malocclusions. growing animals responded to this stress by skeletal adaptations.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. (3) restricting animals to calorie-deprivation diets. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . (2) bone resorption of the anterior edges of the glenoid fossa. 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. (3) lengthened condylar neck. especially the physical consistency of the diet. older animals responded with tooth movements to adjust to the new occlusal alignments. 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. racial outcrossing. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. 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. specifically: (1) some posterior bone deposition.5-2. Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch.

were smaller. Attachment areas for the jawclosing muscles. When compared to the harddiet rats. which is essential to resist transverse bending. Nutrition Experiments concerning the contribution of nutrition to normal occlusal development have taken the form of calorie deprivation of pigs. 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. was less in the soft-diet animals.32 Moore. Owens and Tonge= used weanling animals. Rehabilitation was only partiall y successful." Barber. maintaining them for one year on a severe calorie-deficient diet. (4) had less width of the maxillary dental arch. Vol."29 The consequences were tooth crowding and abnormal tooth-tooth contacts. the dentition migrates to accommodate. More recently. where toothfood-tooth contact takes place. 1 January. they do not create conditions likely to be encountered by the developing human masticatory system. In all studies. and COX. They . Beecher and Corruccini35 examined a small population of rhesus macaques which had spent a short period during adolescence on contrasting hard/soft diets. which resist bite (especially incisal) reaction force. which responded with less deposition and growth. Even buccolingual thickness of the mandible. although with no significant differences in shape. The smaller and less-active muscles applied less stress to the bones.28.29 McCance. in older animals. The smaller masticatory muscles in soft-diet animals were almost certainly a result of lower work requirement in mastication. with condyles smaller and radiographically less dense. Tonge and McCance. The authors found that their regime resulted in: (I) delayed dental development and eruption. (3) had mandibles that were smaller. Green. there is no evidence to correlate this deficiency with the occlusal problems found in so many urbanized peoples of the western world. While these experiments are important in assessing the response of the masticatory system to unusual mechanical stress. before attempting to "rehabilitate" some with proper foods.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. All mandibular structures related to chewing were affected by diet hardness. (2) exhibited no molar wear. 1981 Development 63 four studies have examined this idea experimentally. These studies demonstrate that forced realignments of occlusal relationships and mandibular movements lead to responses by the most flexible elements of the masticatory apparatus. the soft-diet animals: (I) were slightly smaller in body mass. (5) had smaller masseter and temporalis muscles. and (6) had skulls consistently smaller in mass and in linear dimensions. the growing skeletal system adjusts. where muscle force is transmitted to the mandible. In young animals.Diet Consistency jOcclusal the teeth adapted and once again occluded adequately. especially in the mandible. was less. 51 No. Sagne and Thilander. the animals were maintained for approximately four months. were smaller. and (2) "greater delay in the development and growth of the jaws. While calorie deprivation is undoubtedly a factor to consider in severely undernourished people." and Henrikson. Wear of the molar. The condyles.

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

7 I 9.) 24.41) (12. Condyle Length Body weight (gm) .32) (0. Table 3 lists key paramo eters of this analysis. In particular. Variable Maxillary Length (mm) . and the differences are often significant. Thus the residual from Mean.64 8. The gross differences in the growth of most structures are not very marked. 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.f.39 8.03 1.~6) (0..01) lAO ( 0.48) Soft Mean (s.79 9..20) ( 0.45 8.72 1.26 38. the second axis is largest among the softdiet animals.) 23.50) (0.05. Group I is more variable than Group II in every measurement.40) om AoV with 2 and 87 d. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance. The major (coordinated growth) axis is largest in the hard-diet sample. 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.17) ( 0..58) ( 0.88) (0. although more similar to the soft group. Group III is intermediate in standard deviation.d.91 3. and functional interpretation of the morphological integration among variables. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples.1 January. but not significantly so. signifying that all structures are more tightly integrated in their growth and covariation.57 4.40) ( 0.) ~:I.46) ( 0.43 ( 1. For instance.92 3.44 at p (0. The strongest difference is in the growth relation of the masseter muscle.38) ( 0..89 3. Dispersion.d. Vol. 51 No. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I).18 lAO Intermittent Mean (s. All correlations are higher in the animals that had more masticatory resistance. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals..00 9.. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II).28) (or Measured Traits F 0.39) (11.36) (9. Inversely related to this. • 3-sample TABLE I and Significance of Difference Hard Mean (s.d.. significant < ..Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples.. Mandibular Length (mm) (mm) . indicating a greater range of sizes. a result attributable to the small differences in dietary consistency.31 39. Masseter weight (gm) . Maxillary Breadth (mm) . Again.56 ( 1. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures.43 43.65) ( 0.21' 1. but usually closer to Group I. RESULTS Development 65 more variable than Group I in the maxillary arch measurements.. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix. 1981 .38 2.

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

disturbance of synchronous development in separately growing parts. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally. prenatal insult. 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 4.14 0.95 (variation in growth) Soft Intermittent 0.08 0. Thus.:" Maxillary growth may be under closer genetic programming.73 -0.67 (78) 0. 51 No. 1981 .w Moorrees and Reed.68 0. and independence of tooth formation.93 0.41 -0. experimental evidence which might weight such factors is absent.06 -0.80 0.80 One Principal Component Hard Two growth) Intermittent 4. hormonal intervention.90 0.90 0.57 -0.68 0.89 0.94 0. Though the chewing explanation is most predictive of observed occlusal differences in the rats.42-45 The growth independence of mandible and maxilla in humans has been frequently noted. premature deciduous tooth loss.79 (13) -0. but does not give the source of the growth-correlation breakdown.04 0.41•42 Some of these may be of minimal or no importance. including mouth breathing.97 0. follows different rates and peaks at different ages in the maxilla and mandible.08 -0.89 0.18 -0. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles. especially of anterior teeth.91 0.60 (10) -0.29 0.96 (83) 0.96 0.17 (69) 0. 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.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion.51 0. In particular. oral habits.50 (8) -0.95 4.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.23 0.12 0. endocrine disturbance. Because of its widespread occurrence in western society.82 0. trauma.92 0.10 Vol." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. posture. The aging process also yields different results in upper and lower arches.05 -0. with the maxillary teeth being more conservative and controlled by a fewer number of genes.16 -0.77 0. the mandible may depend on muscular function to grow to average size. Tooth spacing and crowding.06 -0.59 -0. 1 January. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control.

W. 19:543-553. The unfused mandibular symphysis in the rat alters force distribution.1977. Takano. Arch. ~ 1. J.: Tooth-Size and Occlusion in Twins. M. Washington. M. 2. The ora! status of the Xavantes of Simoes Lopes.: Dental condition of two tribes of Taiwan aborigines-Ami and Atayal. Price. J. 1937. 8. 11.: Eskimo and Indian field studies in Alaska and Canada. Ser. Am.present convincing genetic arguments against this explanation. 45: 172-207. Lombardi. W. Calci]. Eur . J. R. 23:417-437. Res. Periodont . 40 (Suppl.: Hered· ity and the craniofacial complex. Tiss. B. Dent. a Melanesian group on Mailaito. The prevailing explanation for such lowering of growth correspondence has been the effect of racial admixture between populations. Assoc. Hunt. Am. K. teeth.: Mal· occlusion in the Kwaio. Anthrop. Trans. The Angle Orthodontist ..: Dental observations among the Eskimos.: Malocclusion and civilization. 18. USPHS. The amount of maxillary arch narrowing and collapse can be predicted by the amount of time the animals were chewing on hard food. 17:231-233. D.°o and Lombardi and Bailit. 80-93. Suensk .: Maxillary dental arch width in Chinese living in Liverpool.: Further studies on the Xavantc Indians. 7. 14. 12. A. Bjork. 1951. Wood. Oral Bioi. 1948.: Malocclusion in the modern Alaskan Eskimo. A iii. 5. Am.: The Downs analysis applied to three ethnic groups. M. A. and Wong.C. L. has apparently resulted from lower chewing stress required by a soft dietary consistency. Lundstrom. 47:811815. D.v-se Thus. Assoc. 1966. Angle Orthod. Orthod. I'. 1948. Tand. Res. 3. W. J.. S. J. and Frei. 5b). 14:34. E. and Bailit. 6. Williams. 1947. W.. R. 60:344354. S. Genet. E. 9. Niswander. B. previously noted in certain groups of maloccluded humans and attributed to racial admixture. 1. 1959.: Anglo-Saxon and modern British teeth. 10. Lavelle. 1968. 19. C. Goose. 47:406-422.. Wise. J. No. J. Kelly. Waugh. J. P. 1951. H. Dent. 1937. CONCLUSION We believe our results suggest future directions for fruitful research based on the following observations. L. 1977. V. 1943. 1972. Kraus.. Lu. Clinch. L. Assoc.1936.68 Beecher and Corruccini REFERENCES occlusion in humans. and Thilander. 17. Ortluul. 1977. 1Dent. H. Res. and muscle function. Dent. 36:283·294. J. J. Alii.. A. Henrikson. Chung et al. Klatsky. 1972. 1976. 16. Kargar. Replication of these experiments with a laboratory animal more closely related to man in terms of both biology and masticatory system is seen as a vital future need. IG:355-356. 24:1640· 1647. 15. Supp.-A.. L. Dent. VI!. II. Cotton. and Harvey. 56:117-126. D.: Influence of the diet on the jaws and the face of the American Eskimo. Soc. Solomon Islands. W. Phys. Waugh. Vital and Health Statistics. C. 1961. Basel.: An assessment of occlusion of youths 12·17 years. J. Am. 36:385. Am.: The occlusion of the Australian aborigine.. Orthod. Res. lium. Am.: Bone. B. 162. A.: Studies in the dietaries of contemporary primitive peoples. 4. Sagne. In this animal population the disassociation in size of occlusal features during their growth. 22:466-467. 13. Dent. C. and continual incisor function in rodents maintains a base level of function regardless of diet that has no parallel in man. Mediolateral maxillary growth seems dependent upon the muscular stimulation provided by rough elements in the diet. H. Orthod.: The face in profile. L. J. the inheritance of different-sized parts that must occlude is attributed to the independent segregation of disharmonious genes from disparate ancestors. 21 :213-220.: Investigations concerning the dentitions of the Eskimos of Canada's Eastern Arctic.

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