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BEECHER AND ROBERT
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.
MODERN DIET AND MALOCCLUSION
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
vSuch effects are common in humans. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. and (4) varying the consistency of the diet from hard to soft. affect the developing masticatory apparatus and create variations in the occlusal phenotype.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased.5-2. the experimental attempts to induce developmental change in the mammalian masticatory apparatus have used four methods': (I) forcing mouth breathing in animals. While the younger. Similar differences were also found in soft-diet rats. abnormally-directed muscle tension.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. racial outcrossing. we believe that environmental factors. 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. (2) placing devices in the mouth to effect alterations in mastication. A cornman habit in childhood. older animals responded with tooth movements to adjust to the new occlusal alignments. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. Based on the available anthropological and experimental evidence. The maxillae of these animals showed reduced vertical and prognathic growth. specifically: (1) some posterior bone deposition. inbreeding. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. 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. (2) bone resorption of the anterior edges of the glenoid fossa. Experimen Is in effects of function Apart from radical surgery. (3) restricting animals to calorie-deprivation diets. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist .0 mm in order to achieve molar occlusion. 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. (3) lengthened condylar neck. mouth breathing is probably a factor in the development of human malocclusions. Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch. These are usually designed to force protrusion of the mandible 1. or accumulation of genetic mutations. especially the physical consistency of the diet. growing animals responded to this stress by skeletal adaptations. Using mature animals and a splint on the anterior teeth to force mandibular protrusion.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals.
Attachment areas for the jawclosing muscles. the animals were maintained for approximately four months. which responded with less deposition and growth. Sagne and Thilander. the growing skeletal system adjusts. When compared to the harddiet rats. which resist bite (especially incisal) reaction force. Green. the dentition migrates to accommodate. with condyles smaller and radiographically less dense. and (6) had skulls consistently smaller in mass and in linear dimensions. More recently. where muscle force is transmitted to the mandible. Tonge and McCance. The condyles. although with no significant differences in shape. (5) had smaller masseter and temporalis muscles. Wear of the molar." Barber. All mandibular structures related to chewing were affected by diet hardness. were smaller. Nutrition Experiments concerning the contribution of nutrition to normal occlusal development have taken the form of calorie deprivation of pigs. and (2) "greater delay in the development and growth of the jaws. which is essential to resist transverse bending. They . While calorie deprivation is undoubtedly a factor to consider in severely undernourished people. in older animals. (4) had less width of the maxillary dental arch. Beecher and Corruccini35 examined a small population of rhesus macaques which had spent a short period during adolescence on contrasting hard/soft diets.28. (2) exhibited no molar wear. Dietary consistency Although many workers have commented on the probable influence of the hardness or softness of the diet on the developing masticatory system. before attempting to "rehabilitate" some with proper foods. These studies demonstrate that forced realignments of occlusal relationships and mandibular movements lead to responses by the most flexible elements of the masticatory apparatus. While these experiments are important in assessing the response of the masticatory system to unusual mechanical stress. Rehabilitation was only partiall y successful. the soft-diet animals: (I) were slightly smaller in body mass. there is no evidence to correlate this deficiency with the occlusal problems found in so many urbanized peoples of the western world. In young animals. Watt and Williams. they do not create conditions likely to be encountered by the developing human masticatory system. especially in the mandible. The smaller masticatory muscles in soft-diet animals were almost certainly a result of lower work requirement in mastication.32 Moore.29 McCance. was less. Even buccolingual thickness of the mandible. Vol. 1 January.Diet Consistency jOcclusal the teeth adapted and once again occluded adequately. were smaller. Owens and Tonge= used weanling animals. was less in the soft-diet animals."29 The consequences were tooth crowding and abnormal tooth-tooth contacts. where toothfood-tooth contact takes place. (3) had mandibles that were smaller. maintaining them for one year on a severe calorie-deficient diet. In all studies. 1981 Development 63 four studies have examined this idea experimentally. The authors found that their regime resulted in: (I) delayed dental development and eruption." and Henrikson. 51 No. The smaller and less-active muscles applied less stress to the bones.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.
Animals were also examined for visible differences in tooth alignments and attritional differences between the three groups. incisor to Ml. 3. 5. MATERIALS AND METHODS The experimental animals were Sprague-Dawley rats. evenly divided between males and females. the heads were randomly numbered so that the measurer would not be aware of their group membership.he correlations among the several variables. related to factor analysis. Anteroposterior length of condylar articular surface. finds the major axes of the between-variable correlation matrix. The three populations were maintained on their respective diets for four months. Mandibular length.64 Beecher and Corruccini than the gruel. The second axis is the axis perpendicular to the first that subsumes the largest possible amount of residual variance. This technique. acquired at 21 days of age. incisor to distal edge of last molar. it could not be determined whether the masticatory system responded in direct proportion to differences in dietary consistency. 6. A principal components analysis was performed on the data in order to discover t. It was to fill these gaps in the experimental record that the following study was carried out. with only two groups. Ninety animals were divided into three groups: Group I was fed pelletted rat chow (Purina Formulab). The eigenvect. 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. 4. easily breaking down into small granules. Projection of indiThe Angle Orthodontist found significant narrowing of the maxillary arch with no lessening in length in the soft-diet monkeys. engendering only subtle differences in bite force. 2. with dry pellets provided every seventh day only. 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. and so on.ors (also known as directional cosines or latent vectors) summarize the multiple interplay among variables. After sacrifice. This indicates that the "hard" diet was minimally harder . Least-squares theory is used to rotate orthogonal axes to a position where the major axis subsumes a maximized amount of the total variance. Fresh mass of the entire masseter. resulting in a lj uantum change in the morphology of the masticatory system. Croup 11 was fed a gruel-like porridge consisting of ground chow moistened with water. Measurements Crable I) were taken as follows: 1. Body mass. We have no way of knowing the consistency of rat diets used in earlier experiments of this type. Maxillary arch breadth across buccal points of ?vII. We found the lab chow to be crumbly. Further. the same length of time used in previous experiments of this type. 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. Maxillary arch length. Group II I was fed the soft diet six days. hard and soft diet.
All correlations are higher in the animals that had more masticatory resistance.89 3. The strongest difference is in the growth relation of the masseter muscle.~6) (0.18 lAO Intermittent Mean (s. significant < . signifying that all structures are more tightly integrated in their growth and covariation. 51 No. The gross differences in the growth of most structures are not very marked.57 4.72 1. Maxillary Breadth (mm) .44 at p (0.) 23. but not significantly so.40) ( 0.05. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix.. Group I is more variable than Group II in every measurement.20) ( 0. Thus the residual from Mean.48) Soft Mean (s. the second axis is largest among the softdiet animals. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance. a result attributable to the small differences in dietary consistency.45 8. Table 3 lists key paramo eters of this analysis.) 24. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples. Inversely related to this. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures.1 January.31 39. indicating a greater range of sizes.64 8. Condyle Length Body weight (gm) .65) ( 0.d. Variable Maxillary Length (mm) . Dispersion.21' 1.39) (11.40) om AoV with 2 and 87 d. although more similar to the soft group.01) lAO ( 0. 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. Mandibular Length (mm) (mm) . For instance. but usually closer to Group I.91 3.36) (9.92 3.38) ( 0.39 8.43 43.50) (0..88) (0.79 9.32) (0.58) ( 0. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I). Vol. Masseter weight (gm) . and functional interpretation of the morphological integration among variables.43 ( 1. 1981 .26 38.d. and the differences are often significant.00 9..7 I 9.38 2..f.... Group III is intermediate in standard deviation.41) (12.56 ( 1.28) (or Measured Traits F 0. • 3-sample TABLE I and Significance of Difference Hard Mean (s... 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. The major (coordinated growth) axis is largest in the hard-diet sample.d..17) ( 0. RESULTS Development 65 more variable than Group I in the maxillary arch measurements. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II).46) ( 0. Again. In particular.Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples.03 1. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals.) ~:I.
.L 0..... 0. Body weight ..83· 0.. Perhaps a decrease in buffering among the growing components is the first step in their decrease in size....39 diet sample at p :c .. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth.differentiation represented by this difference in dietary consistency. at the stage of growrh.60 0. Thus... we could find none. Maxillary Breadth ..5%.79· 0.. and masseter weight.. which is significantly greater than expected by random error at P 2.85 Groups Mussel..H3 0. Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes.05.55 0..... the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2...74 0.. Condyle Length . Mandibular Length . No caries were observed by us in any animals.72 Condo 0. We noticed no displaced or rotated teeth in any of the animals..78 0. condyle length.79· 0. This phenomenon has also been noted in characters that are diminishing in size through natural selection. TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max. The Angle Orthodontist . Condyle length alone shows a notable residual on the second axis in Group I and III..91· 0.89 0. IMaxillary Length . 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.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....66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation.19. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child.54 0. In all respects.66 Body 0. Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals."] ""hile they noted easily observable differences in attrition between soft and hard diet groups....75· 0.. • Significantly larger Max.93· 0.. the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample... . Ii Mand.37. 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... ..71 Soft diet ~ r in hard < .. a decrease in correlation of masticatory structures is more noteworthy than de- = '.. Masseter weight . ~ e: 0.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...
16 -0.96 (83) 0.50 (8) -0. 1981 .10 Vol. The aging process also yields different results in upper and lower arches.95 (variation in growth) Soft Intermittent 0. 1 January. with the maxillary teeth being more conservative and controlled by a fewer number of genes.73 -0.29 0. oral habits.41•42 Some of these may be of minimal or no importance.w Moorrees and Reed.:" Maxillary growth may be under closer genetic programming. Because of its widespread occurrence in western society. experimental evidence which might weight such factors is absent. endocrine disturbance.23 0.80 One Principal Component Hard Two growth) Intermittent 4. trauma. premature deciduous tooth loss.60 (10) -0.93 0.79 (13) -0.06 -0. especially of anterior teeth.68 0.96 0. Though the chewing explanation is most predictive of observed occlusal differences in the rats.82 0.90 0. Thus.57 -0." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. hormonal intervention. and independence of tooth formation. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.68 0. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles.97 0. 51 No.06 -0.92 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.89 0.04 0.94 0.91 0.67 (78) 0.08 -0.42-45 The growth independence of mandible and maxilla in humans has been frequently noted. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control. disturbance of synchronous development in separately growing parts. but does not give the source of the growth-correlation breakdown.95 4.90 0. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion.41 -0.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.18 -0. follows different rates and peaks at different ages in the maxilla and mandible.77 0.12 0. prenatal insult.51 0. In particular.80 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 4.89 0.14 0. posture. including mouth breathing.08 0.05 -0.59 -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. Tooth spacing and crowding.17 (69) 0.
: The Downs analysis applied to three ethnic groups. P. Res. L. C. Kelly. 22:466-467. Dent. The Angle Orthodontist . Res.. Klatsky. J. Waugh. S. 5b). 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. K. Cotton.: The occlusion of the Australian aborigine. Kargar. S. Calci]. 21 :213-220.: Anglo-Saxon and modern British teeth. Dent. B. Arch. 12.: Further studies on the Xavantc Indians. 15. Clinch. 16. I'. Lombardi. L. J. W. and Frei. A.: Mal· occlusion in the Kwaio. Am. Lundstrom. B. E. 8. Niswander.: An assessment of occlusion of youths 12·17 years.1936. The unfused mandibular symphysis in the rat alters force distribution. Chung et al. L.: Eskimo and Indian field studies in Alaska and Canada. 3. 47:811815. 1948. C.. J. W.: Dental observations among the Eskimos. II. a Melanesian group on Mailaito. Phys. Tiss. 1951. Basel. Periodont . 1977. Vital and Health Statistics. 1966. 1Dent. ~ 1. J. Bjork. Supp. Kraus. J. and muscle function. teeth. Res. Lavelle. Am. 1. Ser. 1961. L. H. B. Dent. 13. In this animal population the disassociation in size of occlusal features during their growth.. J. Lu.. J. Orthod. 1959. Alii. Orthod. C. J. Assoc. 1976.. and Harvey. J. 2. H.v-se Thus. Sagne. W. 40 (Suppl.. Dent. 7. W. 18.: The face in profile.: Tooth-Size and Occlusion in Twins. The amount of maxillary arch narrowing and collapse can be predicted by the amount of time the animals were chewing on hard food. 1947. Dent. Angle Orthod. has apparently resulted from lower chewing stress required by a soft dietary consistency. The ora! status of the Xavantes of Simoes Lopes. 80-93. 14:34. M. 4. E. Eur . 19.: Studies in the dietaries of contemporary primitive peoples. J. A. Orthod. Price.: Influence of the diet on the jaws and the face of the American Eskimo.-A.: Investigations concerning the dentitions of the Eskimos of Canada's Eastern Arctic. 1937. D. Am. 1972. Hunt. 1977. H.: Malocclusion and civilization. J. 10. Am. IG:355-356. 1948. Wise.1977. A. and Thilander. Trans. Res.. the inheritance of different-sized parts that must occlude is attributed to the independent segregation of disharmonious genes from disparate ancestors. 17. Assoc. A iii.. 11. and Bailit. lium. M. D. Goose. V. Washington. Waugh. 24:1640· 1647. Genet. R. VI!. R. L.present convincing genetic arguments against this explanation. 1972. 1937.: Hered· ity and the craniofacial complex. 1968. Mediolateral maxillary growth seems dependent upon the muscular stimulation provided by rough elements in the diet. CONCLUSION We believe our results suggest future directions for fruitful research based on the following observations. Am. Assoc. 56:117-126. 23:417-437. 5. and continual incisor function in rodents maintains a base level of function regardless of diet that has no parallel in man. Williams. Am. 45: 172-207. Oral Bioi. 47:406-422.68 Beecher and Corruccini REFERENCES occlusion in humans.: Malocclusion in the modern Alaskan Eskimo. No.°o and Lombardi and Bailit. 9. Wood. 36:283·294. 19:543-553. Henrikson.C. Suensk . USPHS. 60:344354. 14. 6. M. 17:231-233.: Maxillary dental arch width in Chinese living in Liverpool. W. previously noted in certain groups of maloccluded humans and attributed to racial admixture. The prevailing explanation for such lowering of growth correspondence has been the effect of racial admixture between populations. 162. A. J. 1951. Tand. and Wong. Soc. Takano. Anthrop. 1943. 36:385. Solomon Islands.: Dental condition of two tribes of Taiwan aborigines-Ami and Atayal. D.: Bone. Ortluul.
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