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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
Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch.vSuch effects are common in humans. While the younger. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint. older animals responded with tooth movements to adjust to the new occlusal alignments. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . we believe that environmental factors. The maxillae of these animals showed reduced vertical and prognathic growth.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. These are usually designed to force protrusion of the mandible 1. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. 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. especially the physical consistency of the diet. 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. 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. Similar differences were also found in soft-diet rats. and (4) varying the consistency of the diet from hard to soft. specifically: (1) some posterior bone deposition.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.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. (3) lengthened condylar neck. (3) restricting animals to calorie-deprivation diets. (2) bone resorption of the anterior edges of the glenoid fossa. growing animals responded to this stress by skeletal adaptations. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. abnormally-directed muscle tension.0 mm in order to achieve molar occlusion.5-2. or accumulation of genetic mutations. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. mouth breathing is probably a factor in the development of human malocclusions. Based on the available anthropological and experimental evidence. (2) placing devices in the mouth to effect alterations in mastication. 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. inbreeding. racial outcrossing. affect the developing masticatory apparatus and create variations in the occlusal phenotype.
and COX. When compared to the harddiet rats. before attempting to "rehabilitate" some with proper foods. While calorie deprivation is undoubtedly a factor to consider in severely undernourished people. The smaller and less-active muscles applied less stress to the bones. The smaller masticatory muscles in soft-diet animals were almost certainly a result of lower work requirement in mastication. The authors found that their regime resulted in: (I) delayed dental development and eruption. and (6) had skulls consistently smaller in mass and in linear dimensions. They ."29 The consequences were tooth crowding and abnormal tooth-tooth contacts. Green. While these experiments are important in assessing the response of the masticatory system to unusual mechanical stress. especially in the mandible. 1 January." Barber. Rehabilitation was only partiall y successful. Beecher and Corruccini35 examined a small population of rhesus macaques which had spent a short period during adolescence on contrasting hard/soft diets. was less in the soft-diet animals.28. Nutrition Experiments concerning the contribution of nutrition to normal occlusal development have taken the form of calorie deprivation of pigs. where muscle force is transmitted to the mandible. 1981 Development 63 four studies have examined this idea experimentally. the dentition migrates to accommodate. 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. Even buccolingual thickness of the mandible. the soft-diet animals: (I) were slightly smaller in body mass. (2) exhibited no molar wear. 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. In young animals. Dietary consistency Although many workers have commented on the probable influence of the hardness or softness of the diet on the developing masticatory system. the growing skeletal system adjusts. in older 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. were smaller. with condyles smaller and radiographically less dense. More recently. 51 No. and (2) "greater delay in the development and growth of the jaws. In all studies. Tonge and McCance. Sagne and Thilander. which resist bite (especially incisal) reaction force. (4) had less width of the maxillary dental arch. they do not create conditions likely to be encountered by the developing human masticatory system. (5) had smaller masseter and temporalis muscles. where toothfood-tooth contact takes place." and Henrikson. All mandibular structures related to chewing were affected by diet hardness. Wear of the molar. The condyles.32 Moore. maintaining them for one year on a severe calorie-deficient diet.Diet Consistency jOcclusal the teeth adapted and once again occluded adequately. although with no significant differences in shape. which is essential to resist transverse bending.29 McCance. Vol. (3) had mandibles that were smaller. was less. which responded with less deposition and growth. Owens and Tonge= used weanling animals.
2. 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. hard and soft diet.64 Beecher and Corruccini than the gruel. easily breaking down into small granules. Mandibular length. Further. acquired at 21 days of age. Animals were also examined for visible differences in tooth alignments and attritional differences between the three groups. finds the major axes of the between-variable correlation matrix. Croup 11 was fed a gruel-like porridge consisting of ground chow moistened with water. 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. This indicates that the "hard" diet was minimally harder . 'While a number of measurements have been taken from animals in dietary consistency experiments. related to factor analysis.ors (also known as directional cosines or latent vectors) summarize the multiple interplay among variables. It was to fill these gaps in the experimental record that the following study was carried out. Group II I was fed the soft diet six days. or whether no differences should be expected until a threshold in dietary consistency difference was reached. Anteroposterior length of condylar articular surface. 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. the same length of time used in previous experiments of this type. 3. Measurements Crable I) were taken as follows: 1. The three populations were maintained on their respective diets for four months. 5.he correlations among the several variables. the heads were randomly numbered so that the measurer would not be aware of their group membership. with only two groups. Maxillary arch length. Projection of indiThe Angle Orthodontist found significant narrowing of the maxillary arch with no lessening in length in the soft-diet monkeys. We found the lab chow to be crumbly. Fresh mass of the entire masseter. it could not be determined whether the masticatory system responded in direct proportion to differences in dietary consistency. After sacrifice. This technique. evenly divided between males and females. 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. The eigenvect. resulting in a lj uantum change in the morphology of the masticatory system. with dry pellets provided every seventh day only. Maxillary arch breadth across buccal points of ?vII. A principal components analysis was performed on the data in order to discover t. engendering only subtle differences in bite force. Ninety animals were divided into three groups: Group I was fed pelletted rat chow (Purina Formulab). and so on. 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. incisor to Ml. Body mass. 4.
Variable Maxillary Length (mm) .. Condyle Length Body weight (gm) . For instance.) 24..45 8.) ~:I.1 January. Group I is more variable than Group II in every measurement. Again..39) (11. Group III is intermediate in standard deviation.7 I 9.65) ( 0. The gross differences in the growth of most structures are not very marked.48) Soft Mean (s. Breakdown in growth coordination Mcxillary breadth affected most The basic descriptive statistics in Table I show the hard-diet population (Group 1) to be larger in all dimensions and disproportionately so in some. All correlations are higher in the animals that had more masticatory resistance.20) ( 0.79 9.58) ( 0. the second axis is largest among the softdiet animals.. a result attributable to the small differences in dietary consistency. signifying that all structures are more tightly integrated in their growth and covariation.17) ( 0. RESULTS Development 65 more variable than Group I in the maxillary arch measurements.Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples.03 1. Table 3 lists key paramo eters of this analysis.64 8.. Inversely related to this.57 4. Dispersion. Vol. Masseter weight (gm) .50) (0.f. In particular.56 ( 1. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals.41) (12. but usually closer to Group I.43 ( 1. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix.92 3..) 23. Thus the residual from Mean.32) (0. 51 No.88) (0.89 3.. The strongest difference is in the growth relation of the masseter muscle. although more similar to the soft group.d.72 1.38) ( 0.43 43.91 3. significant < . The major (coordinated growth) axis is largest in the hard-diet sample.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. 1981 . Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples.39 8.44 at p (0.21' 1. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures. but not significantly so..01) lAO ( 0. Mandibular Length (mm) (mm) .46) ( 0. Maxillary Breadth (mm) .05.. and functional interpretation of the morphological integration among variables.36) (9.28) (or Measured Traits F 0.18 lAO Intermittent Mean (s. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II). the maxillary breadth is markedly increased in the hard-diet reared animals (Group I).38 2.31 39. • 3-sample TABLE I and Significance of Difference Hard Mean (s.d. and the differences are often significant.40) om AoV with 2 and 87 d.40) ( 0.. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance.00 9.26 38. indicating a greater range of sizes.~6) (0.
Ii Mand.. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth. Thus...H3 0. Perhaps a decrease in buffering among the growing components is the first step in their decrease in size. Mandibular Length . which is significantly greater than expected by random error at P 2.differentiation represented by this difference in dietary consistency. and masseter weight...71 Soft diet ~ r in hard < ... We noticed no displaced or rotated teeth in any of the animals...39 diet sample at p :c ."] ""hile they noted easily observable differences in attrition between soft and hard diet groups.60 0. ...91· 0.. a decrease in correlation of masticatory structures is more noteworthy than de- = '...54 0.. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child.. Condyle Length . Masseter weight .79· 0...66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation. the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2...37.. Body weight . The Angle Orthodontist .93· 0.75· 0. Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals...5%... Maxillary Breadth .78 0.. the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample. . In all respects.89 0.. 0.38 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...... No caries were observed by us in any animals.85 Groups Mussel. 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. TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max...79· 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.19. at the stage of growrh. 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.. IMaxillary Length .72 Condo 0.. Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes.83· 0. • Significantly larger Max.66 Body 0. ~ e: 0.. condyle length...05.55 0.74 0. we could find none..L 0... Condyle length alone shows a notable residual on the second axis in Group I and III.
82 0. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.80 0. the mandible may depend on muscular function to grow to average size.51 0.96 (83) 0.97 0. posture. 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. endocrine disturbance. especially of anterior teeth.89 0.05 -0. follows different rates and peaks at different ages in the maxilla and mandible.57 -0. disturbance of synchronous development in separately growing parts.41•42 Some of these may be of minimal or no importance.29 0.95 (variation in growth) Soft Intermittent 0.90 0. 1 January.06 -0. experimental evidence which might weight such factors is absent. including mouth breathing.16 -0.23 0.91 0.73 -0. The aging process also yields different results in upper and lower arches.42-45 The growth independence of mandible and maxilla in humans has been frequently noted. 51 No.79 (13) -0. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles. 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. Because of its widespread occurrence in western society.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.93 0.68 0.92 0. trauma.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion.77 0.:" Maxillary growth may be under closer genetic programming.89 0.08 0.80 One Principal Component Hard Two growth) Intermittent 4.08 -0. premature deciduous tooth loss.68 0.12 0. prenatal insult.10 Vol.18 -0.67 (78) 0.59 -0.04 0.90 0. with the maxillary teeth being more conservative and controlled by a fewer number of genes. oral habits.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion.w Moorrees and Reed. 1981 . and independence of tooth formation. Tooth spacing and crowding.06 -0. In particular.17 (69) 0. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control. Thus." 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. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans.14 0.94 0.96 0. hormonal intervention.41 -0.50 (8) -0. Though the chewing explanation is most predictive of observed occlusal differences in the rats.60 (10) -0.95 4.
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