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

ROBERT

M.

BEECHER AND ROBERT

S.

CORRUCCINI

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

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

Vol. 51

No. I

January,

1981

Similar differences were also found in soft-diet rats. especially the physical consistency of the diet. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. Based on the available anthropological and experimental evidence. the experimental attempts to induce developmental change in the mammalian masticatory apparatus have used four methods': (I) forcing mouth breathing in animals. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. specifically: (1) some posterior bone deposition. and (4) varying the consistency of the diet from hard to soft. (2) bone resorption of the anterior edges of the glenoid fossa.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. 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. (2) placing devices in the mouth to effect alterations in mastication. While the younger. Experimen Is in effects of function Apart from radical surgery. (3) lengthened condylar neck. racial outcrossing. or accumulation of genetic mutations. growing animals responded to this stress by skeletal adaptations. older animals responded with tooth movements to adjust to the new occlusal alignments. affect the developing masticatory apparatus and create variations in the occlusal phenotype.vSuch effects are common in humans. A cornman habit in childhood. 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.0 mm in order to achieve molar occlusion.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. 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. (3) restricting animals to calorie-deprivation diets. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . The maxillae of these animals showed reduced vertical and prognathic growth. we believe that environmental factors.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. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. inbreeding. abnormally-directed muscle tension. These are usually designed to force protrusion of the mandible 1.5-2. mouth breathing is probably a factor in the development of human malocclusions. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures.

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

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

The major (coordinated growth) axis is largest in the hard-diet sample. 1981 .. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I). The strongest difference is in the growth relation of the masseter muscle. Condyle Length Body weight (gm) . RESULTS Development 65 more variable than Group I in the maxillary arch measurements.38) ( 0..) ~:I. Maxillary Breadth (mm) ..03 1. All correlations are higher in the animals that had more masticatory resistance.39 8..92 3.45 8.64 8.~6) (0. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals.40) ( 0.18 lAO Intermittent Mean (s.d.17) ( 0. Masseter weight (gm) . but not significantly so.26 38. indicating a greater range of sizes.01) lAO ( 0.d.38 2.48) Soft Mean (s.05. The gross differences in the growth of most structures are not very marked.d.00 9.56 ( 1. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures. In particular. Dispersion.89 3. Table 3 lists key paramo eters of this analysis.36) (9. and the differences are often significant. Thus the residual from Mean.50) (0. Again. signifying that all structures are more tightly integrated in their growth and covariation..57 4.43 ( 1.46) ( 0. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance.43 43..32) (0..39) (11.91 3. Group III is intermediate in standard deviation.31 39.. Inversely related to this. but usually closer to Group I.. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix.44 at p (0. significant < ..72 1.1 January. Group I is more variable than Group II in every measurement. Vol. Variable Maxillary Length (mm) . the second axis is largest among the softdiet animals. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples.40) om AoV with 2 and 87 d. a result attributable to the small differences in dietary consistency.7 I 9.f.41) (12. • 3-sample TABLE I and Significance of Difference Hard Mean (s.79 9. 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. 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.65) ( 0.28) (or Measured Traits F 0.88) (0.21' 1. although more similar to the soft group. and functional interpretation of the morphological integration among variables.Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples.20) ( 0. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II). For instance. Mandibular Length (mm) (mm) . 51 No.) 23.58) ( 0.) 24.

... Condyle Length .... Masseter weight .H3 0. In all respects.55 0.66 Body 0... Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes..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.72 Condo 0. IMaxillary Length . The Angle Orthodontist .. 0.differentiation represented by this difference in dietary consistency. Body weight .60 0.75· 0. Maxillary Breadth .05.. .... Mandibular Length . No caries were observed by us in any animals.79· 0. condyle length.91· 0.. we could find none.39 diet sample at p :c . the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2. at the stage of growrh. and masseter weight.37.. Condyle length alone shows a notable residual on the second axis in Group I and III.19.. We noticed no displaced or rotated teeth in any of the animals.. ~ e: 0.. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child.66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation....79· 0. which is significantly greater than expected by random error at P 2.. Thus. Ii Mand. the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample."] ""hile they noted easily observable differences in attrition between soft and hard diet groups..78 0.83· 0...89 0. This phenomenon has also been noted in characters that are diminishing in size through natural selection.5%. Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals. ..85 Groups Mussel.L 0..74 0.... • Significantly larger Max.. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth.....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..54 0..71 Soft diet ~ r in hard < . TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max.93· 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.... a decrease in correlation of masticatory structures is more noteworthy than de- = '. 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.. Perhaps a decrease in buffering among the growing components is the first step in their decrease in size.

trauma.06 -0.93 0.:" Maxillary growth may be under closer genetic programming.57 -0.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.80 0.60 (10) -0.96 0. posture.w Moorrees and Reed.06 -0." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. but does not give the source of the growth-correlation breakdown.42-45 The growth independence of mandible and maxilla in humans has been frequently noted.90 0.10 Vol. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control.94 0.91 0. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.41 -0.08 0. premature deciduous tooth loss.67 (78) 0.16 -0. Because of its widespread occurrence in western society.14 0. including mouth breathing.90 0. follows different rates and peaks at different ages in the maxilla and mandible. 51 No.95 (variation in growth) Soft Intermittent 0.80 One Principal Component Hard Two growth) Intermittent 4. endocrine disturbance. 1 January.68 0. with the maxillary teeth being more conservative and controlled by a fewer number of genes. and independence of tooth formation.89 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.96 (83) 0. Tooth spacing and crowding. In particular.82 0.59 -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 4.17 (69) 0. disturbance of synchronous development in separately growing parts.79 (13) -0.95 4.29 0.12 0. prenatal insult.08 -0.92 0.05 -0. the mandible may depend on muscular function to grow to average size. oral habits. especially of anterior teeth. hormonal intervention.68 0. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles. Though the chewing explanation is most predictive of observed occlusal differences in the rats.50 (8) -0. experimental evidence which might weight such factors is absent.04 0. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans.18 -0.73 -0.23 0.89 0.97 0. The aging process also yields different results in upper and lower arches. 1981 . Thus.41•42 Some of these may be of minimal or no importance.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion.51 0.77 0.

v-se Thus. a Melanesian group on Mailaito.. Supp. The amount of maxillary arch narrowing and collapse can be predicted by the amount of time the animals were chewing on hard food.: Eskimo and Indian field studies in Alaska and Canada. Washington. Soc. 7. Trans. and Wong. In this animal population the disassociation in size of occlusal features during their growth. A iii. Suensk . Hunt. previously noted in certain groups of maloccluded humans and attributed to racial admixture. 17. Goose. Klatsky.: Dental observations among the Eskimos.. and muscle function.. Price.. 1951. Ser. A. Cotton. has apparently resulted from lower chewing stress required by a soft dietary consistency.C. C. H. Henrikson.1936. 1961.: The face in profile. Vital and Health Statistics. ~ 1. C. Sagne. S. 1976. E. Dent. Assoc. W. 9. and Harvey. 12. J. 45: 172-207. 1951. J. D. USPHS.present convincing genetic arguments against this explanation. Anthrop. Angle Orthod. J. The Angle Orthodontist . 1947. 19:543-553. 24:1640· 1647. Orthod. Res. Bjork. Eur . H. Lundstrom. 6. Alii. Am. IG:355-356. Solomon Islands. 1977. II. 8. Orthod. The ora! status of the Xavantes of Simoes Lopes. B. Kraus. and continual incisor function in rodents maintains a base level of function regardless of diet that has no parallel in man. L. Dent. 1972. J. CONCLUSION We believe our results suggest future directions for fruitful research based on the following observations. L. VI!. 36:385.: Bone.-A. D. Dent. A. Res. Am. 1948. and Bailit. Lu. 14:34. J.: Hered· ity and the craniofacial complex. A. 36:283·294.: Malocclusion in the modern Alaskan Eskimo.: Investigations concerning the dentitions of the Eskimos of Canada's Eastern Arctic. Am. Kelly. Periodont . 1959. 11. 14.: Mal· occlusion in the Kwaio. R. 1Dent. Lavelle. Dent. E.: The occlusion of the Australian aborigine.. 40 (Suppl. 19. 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. Wise. Genet. L. 13. Res. J. 1966.: Malocclusion and civilization. Lombardi. B. 5b). A. 1968. 1. J. W. 162. 15. Tand. 1937. 1937. J. Wood. R. M. Phys. 1972. Waugh. W. 2. M. Am. Williams. No. Takano. and Frei. 56:117-126. Tiss.: Maxillary dental arch width in Chinese living in Liverpool. and Thilander. Mediolateral maxillary growth seems dependent upon the muscular stimulation provided by rough elements in the diet. W. 10. K. Clinch.°o and Lombardi and Bailit. C..: Anglo-Saxon and modern British teeth. Assoc. H. P. W. The prevailing explanation for such lowering of growth correspondence has been the effect of racial admixture between populations. Am. Calci]. Niswander. J. J. 47:406-422. I'.1977. B. teeth. Oral Bioi. the inheritance of different-sized parts that must occlude is attributed to the independent segregation of disharmonious genes from disparate ancestors. 16.: Influence of the diet on the jaws and the face of the American Eskimo.. 3. 4. Kargar.: Studies in the dietaries of contemporary primitive peoples. J..68 Beecher and Corruccini REFERENCES occlusion in humans. 17:231-233. Dent. Res. 60:344354. 1977.: An assessment of occlusion of youths 12·17 years. 1948. L. Basel. 18. Assoc. V. Ortluul. D. The unfused mandibular symphysis in the rat alters force distribution. 5. Arch. 47:811815.: The Downs analysis applied to three ethnic groups. 22:466-467. M. L. 23:417-437. Waugh. lium. Am. Chung et al. J.: Further studies on the Xavantc Indians. 80-93. Orthod.: Dental condition of two tribes of Taiwan aborigines-Ami and Atayal.: Tooth-Size and Occlusion in Twins. S. 1943. 21 :213-220.

R. Niswander. 51 No. Nance. 33. Orthod.: Craniofacial variation in a central Australian tribe. Christiansen. and Cox. Am. T. 1978.. R. J.. C. Smith. A. Am. 24. 27. J . 1. E. 42. G. J. 115: 1-22. J. P. Niswander. J. J. D. R. Br. J.. S.. Charlier. 1971. Anat. Malocclusion.: Severe undernutrition in growing and adult animals. 35.: Anterior displacement of the mandible in adult rhesus monkeys. Curro Anthrop. L. and Tonge. D.. E. The mouth. Bader. 146:1-34. 1961..: Biometrics of crowding and spacing of teeth in the mandible. Am. 1966. age changes. Chung. 1973. Mourrees. McCance. J. 19: 14-34. G.. 1972. 1954. R. Am. 1965. and Ramsfjord. Am. M.: A genetic study of class III malocclusion. P. J.: Electromyography and mechanics of mastication in the albino rat. 21. Res. W. 23: 471-495. J. M. Am. B. Potter. C.. 146: 123-131. 44:397412. A. Am.. B. 29. Angle Orthod. L. 11. G. 28: 12·35. 19:361-372.. and Hylander. jaws. 1973. J. Beecher. H. 31. J. Litton. 1973. C.: Research related to malocclusion. 73:895-928. Independent genetic determinants. P.. and Kau. J. V. 1971. Schultz. C. Orthod.: Principal factors controlling development of the mandible and maxilla. 1. 1965.: Effects of the ph ysical consistency of the diet on the condylar growth of the rat mandible. Orthod. A. 22. Bouvier. and Dantuma. J. McNamara. M. M. 1975. 12:77-88. 1976. an analysis of Microtus molars. C.. Papua. L. 34. 1. and nailit. Methods. H. Maxillary and mandibular tooth size in different racial groups and in different occlusal categories. and Shapiro. Angle Orthod. Isaacson. Barber. 6:356357.: Effects of mandibular hyperpropulsion on the prechondroblast ic zone of young rat condyle. Strnad. Curro Anthrop. J. Harvold. 1951.Diet Consistency jOcclusal 20. 18. J..: Cenctic and epidemiologic studies of oral characteristics in Hawaii's schoolchildren.: Effects of dietary consistency on maxillary arch breadth in macaques.. 1958. J. and teeth of pigs... 55:71-74. R. 64: 578-606. Am.: Effects of bone strain on cortical bone structure in macaques. A. Alii.. in press. Guthrie. Evolution 19:214-233. Nutr. n. Vol. B... J.: Variability in characters undergoing rapid evolution. Zool. L. 32.: Variability in wild and inbred mammalian populations.: Problems and methods in research on the genetics of dental occlusion.: Primate experiments on oral sensation and dental malocclusions. Burstonc. 45. C. J. 16:503-512.. R. and Weinstein. J. L. January. R.: The effects of the physical consistency of food on the growth and development of the mandible and maxilla of the rat. Development 69 26. 1973. W. S. 1966. Petrovic. S. J.. Hixon. Rhinology XVI:191-196. Dent. 49. Anthrop. R. Am.: Normal development of the jaws and teeth in pigs. Watt. Orthod. S. Brown. W. H. H. Moorrees. and Corruccini.. R. Dent. C. L. 25.. 1966. A.: A twin study of dental dimensions..: Severe undernutrition in growing and adult animals. 50. F. Owens. Phvs. 24:171. J. and Davis. 48.: Masticatory function and skull growth.: Genetics of common dental disorders. D. R. 1965. 47:6577. Tonge.: The cause and effect of mouthbreathing as related to malocclusion. J. 41.J. Moore. C.. D. R. E. S. 1970. 1968. 43:412-421. and McCance. F.. Hum. Phys. C. A. Genet. S. L. in press. J. Anthrop. C. N.: Reply. 63:494-508. W. Lavelle. and Reed. H. Dent. W. University of Adelaide. J. Br. 61:29-37. and Chierici. J..: A radiographic cephalometric study of the central Australian Aborigine. C. Orthod. ]. J. Oppenheimer. 40.: Tool use and crowded teeth in Australopithecinae. 1938. Morph. Orthod. M. 47:661-673. H. Res. Am. W. K. 1977. 30. Am. Gam. Yu. Orth od. Craven. 1'. and HermannStutzmann. 19: 197-206. Master's thesis. R. 39. New Guinea. S. 1956. 59:1-IR. Hiniker. D.. and the delay and malocclusions produced by calorie deficiencies. J. II. 36.. R. Vargervik. C. 44. Tonge. and McCance.-P. 38. L. S. Smith. 15. A III. K.. Runck. 5:419-421. 1978. 43. Thoma. Florida Acad.: Variation in dental occlusion and arches among Melanesians of Bougainville Island. Lavelle. Anthrop.: Research and malocclusion. 1. J. J. ]. ]. Morph. 1969. 1965. 47. and BaiIit. 42:848-851.. The effects of rehabilitation on the teeth and jaws of pigs. Sci. sex differences and population comparisons. Green... Phys. 23. 22:257-268. W. H. II.: Neuromuscular and skeletal adaptations to altered function in the orofacial region. A. 58:565-577. J. M. 1975. Angle Orthod. 37.' 1. Bilben.. R. 1981 . Orthod. 1963. Orthod.: Variation in the secular changes in the teeth and dental arch. Am. Ackerman. and Williams. Nutr. Weijs. Clin. Dent. [rosthet. 47: 195·208. 46. K. 28. 51.

Sign up to vote on this title
UsefulNot useful