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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
> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. especially the physical consistency of the diet. (3) lengthened condylar neck. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. abnormally-directed muscle tension. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. Similar differences were also found in soft-diet rats. The maxillae of these animals showed reduced vertical and prognathic growth.5-2. While the younger.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. (2) placing devices in the mouth to effect alterations in mastication. or accumulation of genetic mutations.vSuch effects are common in humans. affect the developing masticatory apparatus and create variations in the occlusal phenotype. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. 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. the experimental attempts to induce developmental change in the mammalian masticatory apparatus have used four methods': (I) forcing mouth breathing in animals. A cornman habit in childhood. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . and (4) varying the consistency of the diet from hard to soft. Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch. older animals responded with tooth movements to adjust to the new occlusal alignments. mouth breathing is probably a factor in the development of human malocclusions.0 mm in order to achieve molar occlusion. 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. Experimen Is in effects of function Apart from radical surgery. specifically: (1) some posterior bone deposition. (3) restricting animals to calorie-deprivation diets. 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.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. we believe that environmental factors. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. racial outcrossing. growing animals responded to this stress by skeletal adaptations. These are usually designed to force protrusion of the mandible 1. (2) bone resorption of the anterior edges of the glenoid fossa. inbreeding. Based on the available anthropological and experimental evidence.
the animals were maintained for approximately four months.Diet Consistency jOcclusal the teeth adapted and once again occluded adequately. were smaller. in older animals. The authors found that their regime resulted in: (I) delayed dental development and eruption. Attachment areas for the jawclosing muscles. They . The smaller masticatory muscles in soft-diet animals were almost certainly a result of lower work requirement in mastication. Nutrition Experiments concerning the contribution of nutrition to normal occlusal development have taken the form of calorie deprivation of pigs. with condyles smaller and radiographically less dense. the growing skeletal system adjusts.28. 51 No. Even buccolingual thickness of the mandible. Tonge and McCance. All mandibular structures related to chewing were affected by diet hardness.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. (4) had less width of the maxillary dental arch. The smaller and less-active muscles applied less stress to the bones. which responded with less deposition and growth." Barber. although with no significant differences in shape. they do not create conditions likely to be encountered by the developing human masticatory system. In young animals. where muscle force is transmitted to the mandible. Dietary consistency Although many workers have commented on the probable influence of the hardness or softness of the diet on the developing masticatory system.29 McCance. which is essential to resist transverse bending. The condyles. the soft-diet animals: (I) were slightly smaller in body mass. Beecher and Corruccini35 examined a small population of rhesus macaques which had spent a short period during adolescence on contrasting hard/soft diets. Watt and Williams."29 The consequences were tooth crowding and abnormal tooth-tooth contacts. These studies demonstrate that forced realignments of occlusal relationships and mandibular movements lead to responses by the most flexible elements of the masticatory apparatus. before attempting to "rehabilitate" some with proper foods. was less. (3) had mandibles that were smaller. While these experiments are important in assessing the response of the masticatory system to unusual mechanical stress.32 Moore. maintaining them for one year on a severe calorie-deficient diet. was less in the soft-diet animals. Wear of the molar. especially in the mandible." and Henrikson. Rehabilitation was only partiall y successful. (5) had smaller masseter and temporalis muscles. the dentition migrates to accommodate. (2) exhibited no molar wear. When compared to the harddiet rats. where toothfood-tooth contact takes place. Vol. and COX. 1981 Development 63 four studies have examined this idea experimentally. While calorie deprivation is undoubtedly a factor to consider in severely undernourished people. 1 January. More recently. and (2) "greater delay in the development and growth of the jaws. and (6) had skulls consistently smaller in mass and in linear dimensions. Sagne and Thilander. Owens and Tonge= used weanling animals. were smaller. Green. In all studies. there is no evidence to correlate this deficiency with the occlusal problems found in so many urbanized peoples of the western world. which resist bite (especially incisal) reaction force.
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. Group II I was fed the soft diet six days. The three populations were maintained on their respective diets for four months. Croup 11 was fed a gruel-like porridge consisting of ground chow moistened with water. Animals were also examined for visible differences in tooth alignments and attritional differences between the three groups. The second axis is the axis perpendicular to the first that subsumes the largest possible amount of residual variance. 5. Least-squares theory is used to rotate orthogonal axes to a position where the major axis subsumes a maximized amount of the total variance. 4. hard and soft diet.64 Beecher and Corruccini than the gruel. evenly divided between males and females. incisor to distal edge of last molar. finds the major axes of the between-variable correlation matrix. Body mass. incisor to Ml. MATERIALS AND METHODS The experimental animals were Sprague-Dawley rats. 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. The eigenvect. We found the lab chow to be crumbly.ors (also known as directional cosines or latent vectors) summarize the multiple interplay among variables. 3. Projection of indiThe Angle Orthodontist found significant narrowing of the maxillary arch with no lessening in length in the soft-diet monkeys. with only two groups. Anteroposterior length of condylar articular surface. Maxillary arch breadth across buccal points of ?vII. A principal components analysis was performed on the data in order to discover t. Measurements Crable I) were taken as follows: 1. Fresh mass of the entire masseter. acquired at 21 days of age. and so on. the same length of time used in previous experiments of this type. the heads were randomly numbered so that the measurer would not be aware of their group membership. This indicates that the "hard" diet was minimally harder . 2. Further. 6. resulting in a lj uantum change in the morphology of the masticatory system. it could not be determined whether the masticatory system responded in direct proportion to differences in dietary consistency. easily breaking down into small granules.he correlations among the several variables. related to factor analysis. Maxillary arch length. with dry pellets provided every seventh day only. or whether no differences should be expected until a threshold in dietary consistency difference was reached. Mandibular length. Ninety animals were divided into three groups: Group I was fed pelletted rat chow (Purina Formulab). '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. This technique. It was to fill these gaps in the experimental record that the following study was carried out. After sacrifice.
88) (0. The strongest difference is in the growth relation of the masseter muscle. • 3-sample TABLE I and Significance of Difference Hard Mean (s.56 ( 1. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples. 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.21' 1.1 January...64 8.. signifying that all structures are more tightly integrated in their growth and covariation. Mandibular Length (mm) (mm) ... significant < . Dispersion.79 9. For instance.92 3.28) (or Measured Traits F 0. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance. Again.38 2.05. Group III is intermediate in standard deviation.. Maxillary Breadth (mm) . The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix..36) (9. Thus the residual from Mean. Group I is more variable than Group II in every measurement. Masseter weight (gm) .f.d.39 8. The major (coordinated growth) axis is largest in the hard-diet sample.46) ( 0. and the differences are often significant.31 39.45 8.. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures.Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples.38) ( 0. The gross differences in the growth of most structures are not very marked. the second axis is largest among the softdiet animals.. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals.50) (0.d.00 9.41) (12. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I).43 43.) 23. Inversely related to this. 1981 .72 1.57 4. indicating a greater range of sizes.17) ( 0.91 3.40) ( 0. a result attributable to the small differences in dietary consistency.18 lAO Intermittent Mean (s.~6) (0.43 ( 1.20) ( 0. Condyle Length Body weight (gm) . although more similar to the soft group.89 3. and functional interpretation of the morphological integration among variables. Table 3 lists key paramo eters of this analysis.65) ( 0.44 at p (0.39) (11.d. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II).26 38.32) (0.40) om AoV with 2 and 87 d.03 1. In particular. Variable Maxillary Length (mm) . but not significantly so. RESULTS Development 65 more variable than Group I in the maxillary arch measurements. but usually closer to Group I.) ~:I.) 24. 51 No. All correlations are higher in the animals that had more masticatory resistance.48) Soft Mean (s. Vol..01) lAO ( 0.58) ( 0.7 I 9.
54 0....78 0. This phenomenon has also been noted in characters that are diminishing in size through natural selection. No caries were observed by us in any animals.. This increased multivariate variability in soft-diet animals is all the more striking in view of the fact that they were less variable than the hardfood animals in every univariate trait. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth. .79· 0....74 0. We noticed no displaced or rotated teeth in any of the animals. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child. Mandibular Length ..."] ""hile they noted easily observable differences in attrition between soft and hard diet groups. Condyle Length ..5%. at the stage of growrh. .. condyle length..... Ii Mand.79· 0.66 Body 0.71 Soft diet ~ r in hard < . Perhaps a decrease in buffering among the growing components is the first step in their decrease in size. Masseter weight . the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample. Condyle length alone shows a notable residual on the second axis in Group I and III..75· 0. we could find none.83· 0... which is significantly greater than expected by random error at P 2. Thus. Body weight .. 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.55 0.. the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2.differentiation represented by this difference in dietary consistency..89 0..H3 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.66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation.. The Angle Orthodontist .85 Groups Mussel.. and masseter weight.. • Significantly larger Max.19. In all respects.39 diet sample at p :c .93· 0.05.... Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes... TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max..... 0.60 0....91· 0. ~ e: 0. IMaxillary Length ..72 Condo 0. a decrease in correlation of masticatory structures is more noteworthy than de- = '.. 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..L 0..37. Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals.
08 -0. experimental evidence which might weight such factors is absent. trauma.06 -0.68 0.41 -0.06 -0. with the maxillary teeth being more conservative and controlled by a fewer number of genes. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans. 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.:" Maxillary growth may be under closer genetic programming.94 0." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. Though the chewing explanation is most predictive of observed occlusal differences in the rats.96 (83) 0. 51 No. 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 4.23 0.12 0. including mouth breathing. The aging process also yields different results in upper and lower arches. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control.14 0. premature deciduous tooth loss. Because of its widespread occurrence in western society. 1 January. In particular.57 -0.08 0.42-45 The growth independence of mandible and maxilla in humans has been frequently noted.95 4. Tooth spacing and crowding.90 0. disturbance of synchronous development in separately growing parts.77 0.50 (8) -0.41•42 Some of these may be of minimal or no importance.51 0.29 0. 1981 . and independence of tooth formation. but does not give the source of the growth-correlation breakdown.80 One Principal Component Hard Two growth) Intermittent 4.79 (13) -0.92 0.93 0. hormonal intervention.04 0.97 0.w Moorrees and Reed.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion. posture. follows different rates and peaks at different ages in the maxilla and mandible.82 0.68 0.96 0. the mandible may depend on muscular function to grow to average size.89 0. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.89 0.91 0.60 (10) -0. especially of anterior teeth.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.59 -0. Thus.73 -0.10 Vol.05 -0.18 -0.16 -0.95 (variation in growth) Soft Intermittent 0.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion.80 0. endocrine disturbance. prenatal insult. oral habits.90 0.67 (78) 0.17 (69) 0. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles.
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