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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
we believe that environmental factors. older animals responded with tooth movements to adjust to the new occlusal alignments. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. specifically: (1) some posterior bone deposition. inbreeding. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. growing animals responded to this stress by skeletal adaptations. racial outcrossing. While the younger.0 mm in order to achieve molar occlusion.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased.vSuch effects are common in humans. 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. Similar differences were also found in soft-diet rats. (2) bone resorption of the anterior edges of the glenoid fossa. and (4) varying the consistency of the diet from hard to soft. especially the physical consistency of the diet.5-2. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . (3) lengthened condylar neck. 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. Based on the available anthropological and experimental evidence. A cornman habit in childhood. (2) placing devices in the mouth to effect alterations in mastication. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. The maxillae of these animals showed reduced vertical and prognathic growth. 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 experimental attempts to induce developmental change in the mammalian masticatory apparatus have used four methods': (I) forcing mouth breathing in animals.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. mouth breathing is probably a factor in the development of human malocclusions. or accumulation of genetic mutations. affect the developing masticatory apparatus and create variations in the occlusal phenotype. Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch. abnormally-directed muscle tension. (3) restricting animals to calorie-deprivation diets. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. These are usually designed to force protrusion of the mandible 1.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. Experimen Is in effects of function Apart from radical surgery.
1 January. was less. which responded with less deposition and growth. and (2) "greater delay in the development and growth of the jaws. Tonge and McCance. Watt and Williams. (3) had mandibles that were smaller. All mandibular structures related to chewing were affected by diet hardness. Rehabilitation was only partiall y successful. was less in the soft-diet animals. (2) exhibited no molar wear. the soft-diet animals: (I) were slightly smaller in body mass. the growing skeletal system adjusts. and (6) had skulls consistently smaller in mass and in linear dimensions. Beecher and Corruccini35 examined a small population of rhesus macaques which had spent a short period during adolescence on contrasting hard/soft diets. and COX.Diet Consistency jOcclusal the teeth adapted and once again occluded adequately. Dietary consistency Although many workers have commented on the probable influence of the hardness or softness of the diet on the developing masticatory system. maintaining them for one year on a severe calorie-deficient diet. 1981 Development 63 four studies have examined this idea experimentally. While these experiments are important in assessing the response of the masticatory system to unusual mechanical stress. Green. Owens and Tonge= used weanling animals.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. in older animals. before attempting to "rehabilitate" some with proper foods. The authors found that their regime resulted in: (I) delayed dental development and eruption. Vol. They . When compared to the harddiet rats. In young animals. the dentition migrates to accommodate. In all studies. which is essential to resist transverse bending. especially in the mandible. where toothfood-tooth contact takes place. These studies demonstrate that forced realignments of occlusal relationships and mandibular movements lead to responses by the most flexible elements of the masticatory apparatus. with condyles smaller and radiographically less dense."29 The consequences were tooth crowding and abnormal tooth-tooth contacts.32 Moore. More recently. Sagne and Thilander. The condyles. although with no significant differences in shape. Attachment areas for the jawclosing muscles." Barber. were smaller. they do not create conditions likely to be encountered by the developing human masticatory system. where muscle force is transmitted to the mandible. Nutrition Experiments concerning the contribution of nutrition to normal occlusal development have taken the form of calorie deprivation of pigs. (4) had less width of the maxillary dental arch. the animals were maintained for approximately four months. there is no evidence to correlate this deficiency with the occlusal problems found in so many urbanized peoples of the western world. The smaller masticatory muscles in soft-diet animals were almost certainly a result of lower work requirement in mastication. The smaller and less-active muscles applied less stress to the bones." and Henrikson. Wear of the molar. 51 No. (5) had smaller masseter and temporalis muscles.28.29 McCance. Even buccolingual thickness of the mandible. While calorie deprivation is undoubtedly a factor to consider in severely undernourished people. were smaller. which resist bite (especially incisal) reaction force.
Fresh mass of the entire masseter. Maxillary arch breadth across buccal points of ?vII. related to factor analysis. it could not be determined whether the masticatory system responded in direct proportion to differences in dietary consistency. 3. Mandibular length. the heads were randomly numbered so that the measurer would not be aware of their group membership. finds the major axes of the between-variable correlation matrix. 6. Maxillary arch length. A principal components analysis was performed on the data in order to discover t. The eigenvect. The second axis is the axis perpendicular to the first that subsumes the largest possible amount of residual variance. 5. with dry pellets provided every seventh day only. After sacrifice. 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. Animals were also examined for visible differences in tooth alignments and attritional differences between the three groups. This technique. Group II I was fed the soft diet six days. Croup 11 was fed a gruel-like porridge consisting of ground chow moistened with water. Body mass. acquired at 21 days of age. easily breaking down into small granules. Least-squares theory is used to rotate orthogonal axes to a position where the major axis subsumes a maximized amount of the total variance. This indicates that the "hard" diet was minimally harder . 'While a number of measurements have been taken from animals in dietary consistency experiments. 2.he correlations among the several variables. The three populations were maintained on their respective diets for four months. MATERIALS AND METHODS The experimental animals were Sprague-Dawley rats. We have no way of knowing the consistency of rat diets used in earlier experiments of this type. 4. and so on. incisor to Ml. resulting in a lj uantum change in the morphology of the masticatory system. engendering only subtle differences in bite force. Anteroposterior length of condylar articular surface. the same length of time used in previous experiments of this type. 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.ors (also known as directional cosines or latent vectors) summarize the multiple interplay among variables. with only two groups. Ninety animals were divided into three groups: Group I was fed pelletted rat chow (Purina Formulab). We found the lab chow to be crumbly.64 Beecher and Corruccini than the gruel. Further. incisor to distal edge of last molar. hard and soft diet. Measurements Crable I) were taken as follows: 1. It was to fill these gaps in the experimental record that the following study was carried out. evenly divided between males and females. Projection of indiThe Angle Orthodontist found significant narrowing of the maxillary arch with no lessening in length in the soft-diet monkeys. or whether no differences should be expected until a threshold in dietary consistency difference was reached.
92 3.00 9.38) ( 0. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix. Maxillary Breadth (mm) .39) (11.40) ( 0. Again. 1981 .. Variable Maxillary Length (mm) .26 38. In particular. All correlations are higher in the animals that had more masticatory resistance.64 8.38 2. Inversely related to this. significant < . Thus the residual from Mean.) ~:I.43 ( 1.58) ( 0.. Condyle Length Body weight (gm) .39 8. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance.05. For instance. Group I is more variable than Group II in every measurement.28) (or Measured Traits F 0.17) ( 0.d.72 1.) 23.56 ( 1. Masseter weight (gm) . Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples. signifying that all structures are more tightly integrated in their growth and covariation. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II).... 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.03 1.46) ( 0.18 lAO Intermittent Mean (s. the second axis is largest among the softdiet animals. Mandibular Length (mm) (mm) .88) (0.21' 1. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I).1 January.. Table 3 lists key paramo eters of this analysis. although more similar to the soft group. Dispersion.36) (9.f. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals. Group III is intermediate in standard deviation. 51 No. and functional interpretation of the morphological integration among variables.32) (0. Vol.Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples.d. • 3-sample TABLE I and Significance of Difference Hard Mean (s.89 3... The strongest difference is in the growth relation of the masseter muscle.01) lAO ( 0..31 39.57 4.44 at p (0.7 I 9.48) Soft Mean (s.43 43. but usually closer to Group I.40) om AoV with 2 and 87 d. 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. and the differences are often significant. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures.d.~6) (0. The gross differences in the growth of most structures are not very marked.41) (12.91 3.65) ( 0.) 24.79 9. indicating a greater range of sizes. but not significantly so. The major (coordinated growth) axis is largest in the hard-diet sample.50) (0.. RESULTS Development 65 more variable than Group I in the maxillary arch measurements.45 8.20) ( 0. a result attributable to the small differences in dietary consistency.
91· 0. In all respects. a decrease in correlation of masticatory structures is more noteworthy than de- = '.. at the stage of growrh. Ii Mand.. TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child. Masseter weight .72 Condo 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. This phenomenon has also been noted in characters that are diminishing in size through natural selection.39 diet sample at p :c .55 0.. ~ e: 0.79· 0...L 0. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth.75· 0. the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample... and masseter weight..54 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. which is significantly greater than expected by random error at P 2.66 Body 0.93· 0..... condyle length.differentiation represented by this difference in dietary consistency. Body weight . Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes.."] ""hile they noted easily observable differences in attrition between soft and hard diet groups...66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation.... Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals...74 0..19. the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2.89 0....71 Soft diet ~ r in hard < . Condyle length alone shows a notable residual on the second axis in Group I and III.H3 0..5%. IMaxillary Length .60 0. Mandibular Length . This increased multivariate variability in soft-diet animals is all the more striking in view of the fact that they were less variable than the hardfood animals in every univariate trait...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. Condyle Length . The Angle Orthodontist .78 0. We noticed no displaced or rotated teeth in any of the animals.. Thus. ... Maxillary Breadth . • Significantly larger Max.83· 0. we could find none.....05....85 Groups Mussel.79· 0... Perhaps a decrease in buffering among the growing components is the first step in their decrease in size.. 0. . No caries were observed by us in any animals.37...
08 -0.10 Vol. 1981 .97 0.68 0.08 0.29 0.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.68 0.06 -0. oral habits. and independence of tooth formation.57 -0.82 0.17 (69) 0. the mandible may depend on muscular function to grow to average size." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. hormonal intervention. experimental evidence which might weight such factors is absent.91 0.73 -0. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles.16 -0.94 0.93 0.w Moorrees and Reed.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion.67 (78) 0.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion.41•42 Some of these may be of minimal or no importance.05 -0. premature deciduous tooth loss.42-45 The growth independence of mandible and maxilla in humans has been frequently noted.:" Maxillary growth may be under closer genetic programming.89 0. prenatal insult.51 0.77 0. The aging process also yields different results in upper and lower arches. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.41 -0.12 0.96 (83) 0.96 0. especially of anterior teeth. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans.79 (13) -0. posture. endocrine disturbance. In particular. with the maxillary teeth being more conservative and controlled by a fewer number of genes.95 (variation in growth) Soft Intermittent 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. but does not give the source of the growth-correlation breakdown. 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. Though the chewing explanation is most predictive of observed occlusal differences in the rats. including mouth breathing.80 0. follows different rates and peaks at different ages in the maxilla and mandible. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control.92 0.60 (10) -0. Tooth spacing and crowding.50 (8) -0.18 -0.90 0.95 4.90 0. trauma.59 -0. 1 January.89 0. Because of its widespread occurrence in western society. 51 No. disturbance of synchronous development in separately growing parts.14 0. Thus.23 0.06 -0.80 One Principal Component Hard Two growth) Intermittent 4.04 0.
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