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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
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. especially the physical consistency of the diet. (3) lengthened condylar neck. specifically: (1) some posterior bone deposition. inbreeding. (2) placing devices in the mouth to effect alterations in mastication. we believe that environmental factors. While the younger. and (4) varying the consistency of the diet from hard to soft. These are usually designed to force protrusion of the mandible 1. 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. (3) restricting animals to calorie-deprivation diets. racial outcrossing. A cornman habit in childhood. affect the developing masticatory apparatus and create variations in the occlusal phenotype. (2) bone resorption of the anterior edges of the glenoid fossa. 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. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. Experimen Is in effects of function Apart from radical surgery.vSuch effects are common in humans.0 mm in order to achieve molar occlusion. or accumulation of genetic mutations. Based on the available anthropological and experimental evidence. mouth breathing is probably a factor in the development of human malocclusions. Similar differences were also found in soft-diet rats.5-2. The maxillae of these animals showed reduced vertical and prognathic growth.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. abnormally-directed muscle tension. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. 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. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . 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) had mandibles that were smaller. (5) had smaller masseter and temporalis muscles. In all studies. Nutrition Experiments concerning the contribution of nutrition to normal occlusal development have taken the form of calorie deprivation of pigs. where toothfood-tooth contact takes place. More recently. there is no evidence to correlate this deficiency with the occlusal problems found in so many urbanized peoples of the western world. maintaining them for one year on a severe calorie-deficient diet. was less.32 Moore. they do not create conditions likely to be encountered by the developing human masticatory system. and (6) had skulls consistently smaller in mass and in linear dimensions. were smaller. The condyles. Watt and Williams. The smaller and less-active muscles applied less stress to the bones. 1 January. were smaller. in older animals. While calorie deprivation is undoubtedly a factor to consider in severely undernourished people. which responded with less deposition and growth."29 The consequences were tooth crowding and abnormal tooth-tooth contacts. Green. the dentition migrates to accommodate. the animals were maintained for approximately four months. While these experiments are important in assessing the response of the masticatory system to unusual mechanical stress. and COX. Attachment areas for the jawclosing muscles. was less in the soft-diet animals. (4) had less width of the maxillary dental arch. Tonge and McCance. The smaller masticatory muscles in soft-diet animals were almost certainly a result of lower work requirement in mastication. Sagne and Thilander. (2) exhibited no molar wear.29 McCance.28.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.Diet Consistency jOcclusal the teeth adapted and once again occluded adequately. which resist bite (especially incisal) reaction force. 51 No. Wear of the molar." and Henrikson. In young animals. They . which is essential to resist transverse bending. The authors found that their regime resulted in: (I) delayed dental development and eruption. before attempting to "rehabilitate" some with proper foods. Beecher and Corruccini35 examined a small population of rhesus macaques which had spent a short period during adolescence on contrasting hard/soft diets. Vol. the soft-diet animals: (I) were slightly smaller in body mass. Dietary consistency Although many workers have commented on the probable influence of the hardness or softness of the diet on the developing masticatory system." Barber. where muscle force is transmitted to the mandible. and (2) "greater delay in the development and growth of the jaws. the growing skeletal system adjusts. These studies demonstrate that forced realignments of occlusal relationships and mandibular movements lead to responses by the most flexible elements of the masticatory apparatus. Rehabilitation was only partiall y successful. Even buccolingual thickness of the mandible. although with no significant differences in shape. with condyles smaller and radiographically less dense. 1981 Development 63 four studies have examined this idea experimentally. When compared to the harddiet rats. especially in the mandible. Owens and Tonge= used weanling animals. All mandibular structures related to chewing were affected by diet hardness.
It was to fill these gaps in the experimental record that the following study was carried out. 5. Fresh mass of the entire masseter. Further. the same length of time used in previous experiments of this type. Body mass. A principal components analysis was performed on the data in order to discover t. Least-squares theory is used to rotate orthogonal axes to a position where the major axis subsumes a maximized amount of the total variance. 6. After sacrifice. with only two groups. 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. 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. Projection of indiThe Angle Orthodontist found significant narrowing of the maxillary arch with no lessening in length in the soft-diet monkeys. We have no way of knowing the consistency of rat diets used in earlier experiments of this type. This indicates that the "hard" diet was minimally harder . 3.ors (also known as directional cosines or latent vectors) summarize the multiple interplay among variables. The three populations were maintained on their respective diets for four months. 2. Anteroposterior length of condylar articular surface. MATERIALS AND METHODS The experimental animals were Sprague-Dawley rats. incisor to distal edge of last molar. Group II I was fed the soft diet six days. it could not be determined whether the masticatory system responded in direct proportion to differences in dietary consistency. and so on. 4. 'While a number of measurements have been taken from animals in dietary consistency experiments. Maxillary arch breadth across buccal points of ?vII. related to factor analysis. or whether no differences should be expected until a threshold in dietary consistency difference was reached. Mandibular length. resulting in a lj uantum change in the morphology of the masticatory system. evenly divided between males and females. acquired at 21 days of age. 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 Ml. This technique. finds the major axes of the between-variable correlation matrix. hard and soft diet. with dry pellets provided every seventh day only. Croup 11 was fed a gruel-like porridge consisting of ground chow moistened with water. 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.he correlations among the several variables. We found the lab chow to be crumbly. The eigenvect.64 Beecher and Corruccini than the gruel. Maxillary arch length. Animals were also examined for visible differences in tooth alignments and attritional differences between the three groups. easily breaking down into small granules. engendering only subtle differences in bite force.
the maxillary breadth is markedly increased in the hard-diet reared animals (Group I).58) ( 0.72 1.88) (0. The gross differences in the growth of most structures are not very marked. All correlations are higher in the animals that had more masticatory resistance. but not significantly so. and functional interpretation of the morphological integration among variables.. For instance. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals..7 I 9. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance. Mandibular Length (mm) (mm) .05. signifying that all structures are more tightly integrated in their growth and covariation.1 January. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II). Maxillary Breadth (mm) . Dispersion.64 8. Table 3 lists key paramo eters of this analysis.26 38. The major (coordinated growth) axis is largest in the hard-diet sample. although more similar to the soft group.36) (9.40) om AoV with 2 and 87 d.03 1.92 3. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples.d. Again.. a result attributable to the small differences in dietary consistency.56 ( 1. The strongest difference is in the growth relation of the masseter muscle.21' 1.18 lAO Intermittent Mean (s. indicating a greater range of sizes. Group III is intermediate in standard deviation. 51 No. • 3-sample TABLE I and Significance of Difference Hard Mean (s. Condyle Length Body weight (gm) .57 4.Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples... In particular. Masseter weight (gm) .38) ( 0.. Thus the residual from Mean. Variable Maxillary Length (mm) .d. Vol.89 3.41) (12.48) Soft Mean (s.39 8.20) ( 0.d.43 43.01) 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. but usually closer to Group I.28) (or Measured Traits F 0.) ~:I.17) ( 0. and the differences are often significant..50) (0.31 39.32) (0..) 23.39) (11.65) ( 0. Inversely related to this.46) ( 0.~6) (0.00 9.38 2.) 24..f. 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.45 8. RESULTS Development 65 more variable than Group I in the maxillary arch measurements. 1981 . the second axis is largest among the softdiet animals.40) ( 0.44 at p (0. Group I is more variable than Group II in every measurement.91 3. significant < .43 ( 1.79 9.
the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2. IMaxillary Length .93· 0. No caries were observed by us in any animals. We noticed no displaced or rotated teeth in any of the animals... Condyle Length ..74 0.55 0. which is significantly greater than expected by random error at P 2.. The Angle Orthodontist . the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample. Body weight .. Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes. In all respects..89 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.. 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.."] ""hile they noted easily observable differences in attrition between soft and hard diet groups.....79· 0. Thus.75· 0. • Significantly larger Max.. a decrease in correlation of masticatory structures is more noteworthy than de- = '.39 diet sample at p :c . and masseter weight..19.. Ii Mand.. ~ e: 0...83· 0. Perhaps a decrease in buffering among the growing components is the first step in their decrease in size..72 Condo 0. TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max.79· 0..85 Groups Mussel.5%...L 0... we could find none...66 Body 0.60 0. Masseter weight .05. 0.66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation..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. at the stage of growrh.differentiation represented by this difference in dietary consistency. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child.78 0.. . Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals. This phenomenon has also been noted in characters that are diminishing in size through natural selection.. Mandibular Length .. condyle length..71 Soft diet ~ r in hard < .. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth. Maxillary Breadth .......91· 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.54 0.37... Condyle length alone shows a notable residual on the second axis in Group I and III.H3 0... .
06 -0.41•42 Some of these may be of minimal or no importance.08 -0.14 0.90 0. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.89 0.04 0. prenatal insult. oral habits. Though the chewing explanation is most predictive of observed occlusal differences in the rats. follows different rates and peaks at different ages in the maxilla and mandible. experimental evidence which might weight such factors is absent.67 (78) 0.w Moorrees and Reed. trauma.57 -0.68 0. endocrine disturbance.97 0.16 -0.94 0. including mouth breathing.96 0. premature deciduous tooth loss.12 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.96 (83) 0.59 -0. disturbance of synchronous development in separately growing parts.95 4. 1981 .91 0.80 0.77 0. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles.05 -0.41 -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 4.51 0.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion.89 0.23 0. the mandible may depend on muscular function to grow to average size.68 0. 51 No. 1 January. hormonal intervention.82 0.95 (variation in growth) Soft Intermittent 0. especially of anterior teeth.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion. Tooth spacing and crowding.18 -0.90 0. Because of its widespread occurrence in western society. In particular.17 (69) 0.92 0.:" Maxillary growth may be under closer genetic programming. 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.10 Vol. Thus. with the maxillary teeth being more conservative and controlled by a fewer number of genes.93 0.80 One Principal Component Hard Two growth) Intermittent 4.60 (10) -0.06 -0. The aging process also yields different results in upper and lower arches. posture.29 0. but does not give the source of the growth-correlation breakdown.08 0.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle. and independence of tooth formation.50 (8) -0. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans.42-45 The growth independence of mandible and maxilla in humans has been frequently noted.73 -0.79 (13) -0.
: The face in profile. W. Orthod. 2. Oral Bioi.C. M. Washington. Res. B. 7. and muscle function. M.. Orthod.: The occlusion of the Australian aborigine. J. Kargar. 1937. Res. Soc. V. 47:811815. 19. Wood. 8. 1937. 16. 1Dent. 11. L. Res.v-se Thus. J. 22:466-467. and continual incisor function in rodents maintains a base level of function regardless of diet that has no parallel in man. The prevailing explanation for such lowering of growth correspondence has been the effect of racial admixture between populations. Angle Orthod. C. W. 1977. The amount of maxillary arch narrowing and collapse can be predicted by the amount of time the animals were chewing on hard food.. Clinch. Klatsky. 14:34. Lavelle. Suensk . 1947. 47:406-422. 1972. Niswander. Takano. and Thilander. The Angle Orthodontist . Waugh.: Bone.present convincing genetic arguments against this explanation. A.. J. J. Eur .: Tooth-Size and Occlusion in Twins. S. Tiss. 5. Dent. P. 1959. 3. 1968. Dent. Ortluul. A.: Hered· ity and the craniofacial complex. H. CONCLUSION We believe our results suggest future directions for fruitful research based on the following observations. 15. M. H. 45: 172-207. USPHS. W. has apparently resulted from lower chewing stress required by a soft dietary consistency. 1943. Solomon Islands. Bjork. 10. Kraus. Am. J. 162.-A. 5b). Trans. No. 18. Cotton.: Influence of the diet on the jaws and the face of the American Eskimo. and Bailit. Henrikson. Arch. I'. Res. 36:385.: Investigations concerning the dentitions of the Eskimos of Canada's Eastern Arctic. E. J. 1948.: Malocclusion in the modern Alaskan Eskimo.1936. ~ 1. In this animal population the disassociation in size of occlusal features during their growth. H. K. Ser.: Eskimo and Indian field studies in Alaska and Canada. VI!. Am.: Mal· occlusion in the Kwaio. A. IG:355-356. Dent. 40 (Suppl. L..: Dental observations among the Eskimos. W. 1977. J. 23:417-437. a Melanesian group on Mailaito. Price.. S. II. J. 13. 1961. Anthrop. 1. Am.1977. Basel.: Dental condition of two tribes of Taiwan aborigines-Ami and Atayal. J. Orthod. Assoc.: Further studies on the Xavantc Indians. teeth. 1976.: An assessment of occlusion of youths 12·17 years. Vital and Health Statistics. Am.. Williams. lium. W. 1972.°o and Lombardi and Bailit. L. 1951. D. 60:344354.. Lundstrom. R. Tand. 6. C. 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. and Frei. Lombardi. Calci]. Periodont . the inheritance of different-sized parts that must occlude is attributed to the independent segregation of disharmonious genes from disparate ancestors. and Wong. Dent.: Malocclusion and civilization. A iii. Kelly. Assoc. previously noted in certain groups of maloccluded humans and attributed to racial admixture. Wise.: The Downs analysis applied to three ethnic groups. Assoc. J. D. Hunt. Phys. Waugh. J. and Harvey. 17. E. 12. L.: Anglo-Saxon and modern British teeth. 21 :213-220. L. Goose. J. R. 9.68 Beecher and Corruccini REFERENCES occlusion in humans.. Lu. Alii. 56:117-126. Supp. Am. D. 4. A. B. 1966. 80-93. 36:283·294. Mediolateral maxillary growth seems dependent upon the muscular stimulation provided by rough elements in the diet. Dent. The unfused mandibular symphysis in the rat alters force distribution. 1951. Am. Chung et al. The ora! status of the Xavantes of Simoes Lopes. 19:543-553. B. 1948. 17:231-233.: Studies in the dietaries of contemporary primitive peoples. C. 14. Genet. 24:1640· 1647.: Maxillary dental arch width in Chinese living in Liverpool. Sagne.
: Principal factors controlling development of the mandible and maxilla. 45.: Biometrics of crowding and spacing of teeth in the mandible. and Weinstein. Clin. 39. Orthod. S.. Morph. 1975. C... Barber. 1973. Smith. 23: 471-495. B.. Am. W. 1976. 34. Am.: A twin study of dental dimensions. R.: Normal development of the jaws and teeth in pigs. J. 1975. II. Nutr. 24:171. Orthod. Rhinology XVI:191-196. Am. J. 48. M. 15. Potter. C. S. 1966. C. Res. 46. M. 1966.: Severe undernutrition in growing and adult animals. F. The mouth. J. Development 69 26. Br. J. J. Charlier. Am. C. Independent genetic determinants.: The cause and effect of mouthbreathing as related to malocclusion. 1978.: Neuromuscular and skeletal adaptations to altered function in the orofacial region. H. Orthod. Mourrees. K. 1963. Green. and the delay and malocclusions produced by calorie deficiencies. R. Yu. L. Zool. 1968. 1965. Am. 1'. 32. 58:565-577. Orthod. J. and Hylander. 73:895-928. H. S. Dent.: A radiographic cephalometric study of the central Australian Aborigine.. L.. C.. Angle Orthod. Orthod. Orthod. 41.: The effects of the physical consistency of food on the growth and development of the mandible and maxilla of the rat. J..: Problems and methods in research on the genetics of dental occlusion.. D.. Nance. G.: Primate experiments on oral sensation and dental malocclusions. Niswander. C.: Masticatory function and skull growth. 38. and Ramsfjord. Genet. R. Methods. 1972. 47:6577. Ackerman. F. 1971. 11. Sci. A III. 1965. 1973.. 23.. J.. 30. A. A. and Corruccini. 59:1-IR.. Am. J. January. K.. R.: Variability in characters undergoing rapid evolution.. University of Adelaide. 22. W.. Isaacson. R. L. Bouvier. 36. J. jaws. Florida Acad. E. D. 22:257-268. New Guinea. T. Orth od. Am. Gam.. 1. Anthrop.. Tonge. 5:419-421. H. 47:661-673. in press. 1958. Master's thesis. H. H. 146:1-34. H. S. B. ]. J. B. Bilben. Beecher. 1970. 42. Moorrees. and Williams. 1. 55:71-74. 50. L. Orthod.. 51 No. 1951. Vargervik. [rosthet. Anthrop. 1961. Nutr. Chung. and Reed. P. Phys. J. in press. W. 146: 123-131. H. 44:397412. 1965. Curro Anthrop. 43. 18. 19:361-372. Bader. M.: A genetic study of class III malocclusion. Watt. W.: Effects of the ph ysical consistency of the diet on the condylar growth of the rat mandible.: Electromyography and mechanics of mastication in the albino rat. 1938. C. J . R. N. Hixon. Lavelle. R. R. J. C.: Tool use and crowded teeth in Australopithecinae. W.. C. Oppenheimer. 31. 42:848-851. II. 12:77-88. ]. Runck. J. 33. 1966. 47. Vol. 115: 1-22. Orthod. 29. J.. and teeth of pigs.: Reply. A. McCance. J. J. Petrovic. S.. R. 1978. 35. M. McNamara.' 1. Evolution 19:214-233. Niswander. J. J. and Tonge.J. Craven. Dent. 64: 578-606.. J. 1. P. J. 25.: Research and malocclusion. Res. Phys. Am. and Dantuma. and BaiIit. ]. Brown. E. 16:503-512. 24.: Cenctic and epidemiologic studies of oral characteristics in Hawaii's schoolchildren. Malocclusion. Weijs. Am.: Genetics of common dental disorders.: Severe undernutrition in growing and adult animals. E. 1973. Angle Orthod.. C. 1973. R. Am. D. Schultz. 43:412-421. and Cox...: Anterior displacement of the mandible in adult rhesus monkeys. J. J. Dent.: Variation in dental occlusion and arches among Melanesians of Bougainville Island. Owens. R. Guthrie. V. S. and Davis. Br. and McCance.: Research related to malocclusion. A. 1971. Smith.Diet Consistency jOcclusal 20. P. C. K. 1965. 63:494-508. J. A. Papua. J.. 19: 197-206.. 6:356357. Lavelle. and Kau. 28. 1. J. Am.. and nailit.: Variability in wild and inbred mammalian populations. 1981 . A. R. age changes. L. Christiansen. D.-P... 28: 12·35. J. 1954. 44. Anat.: Craniofacial variation in a central Australian tribe. Hum. Maxillary and mandibular tooth size in different racial groups and in different occlusal categories. G. n. Curro Anthrop. 61:29-37.: Effects of dietary consistency on maxillary arch breadth in macaques. L. The effects of rehabilitation on the teeth and jaws of pigs.: Effects of mandibular hyperpropulsion on the prechondroblast ic zone of young rat condyle. Phvs. J. Am. M. J. R. 21. Morph. J. and McCance. and Chierici. Angle Orthod. 37.: Variation in the secular changes in the teeth and dental arch. Moore. L. an analysis of Microtus molars. Burstonc. 19: 14-34. 1977. 27. S. L. 40. Harvold. sex differences and population comparisons. Anthrop. A. 47: 195·208. Tonge. D. W. M. and HermannStutzmann.. 1956. Litton. W. C. S. Thoma. 51. Strnad. G. and Shapiro. Alii.: Effects of bone strain on cortical bone structure in macaques. Hiniker. Dent. 49. R. 1969..