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






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.

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

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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. or accumulation of genetic mutations. and (4) varying the consistency of the diet from hard to soft. mouth breathing is probably a factor in the development of human malocclusions. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. the experimental attempts to induce developmental change in the mammalian masticatory apparatus have used four methods': (I) forcing mouth breathing in animals. attributed to altered muscle tension as the mandible is held slightly depressed during respiration.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. (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. inbreeding.0 mm in order to achieve molar occlusion. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . we believe that environmental factors. abnormally-directed muscle tension. 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. These are usually designed to force protrusion of the mandible 1. racial outcrossing. growing animals responded to this stress by skeletal adaptations.5-2. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. Experimen Is in effects of function Apart from radical surgery. Similar differences were also found in soft-diet rats. (2) bone resorption of the anterior edges of the glenoid fossa. (2) placing devices in the mouth to effect alterations in mastication. A cornman habit in childhood.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. While the younger. older animals responded with tooth movements to adjust to the new occlusal alignments.vSuch effects are common in humans. Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch. The maxillae of these animals showed reduced vertical and prognathic growth. especially the physical consistency of the diet. affect the developing masticatory apparatus and create variations in the occlusal phenotype. (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. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint.

Watt and Williams. When compared to the harddiet rats. Vol. there is no evidence to correlate this deficiency with the occlusal problems found in so many urbanized peoples of the western world. 51 No.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. (3) had mandibles that were smaller. and (2) "greater delay in the development and growth of the jaws. Sagne and Thilander. Rehabilitation was only partiall y successful.28. Attachment areas for the jawclosing muscles. They . Owens and Tonge= used weanling animals. especially in the mandible. and (6) had skulls consistently smaller in mass and in linear dimensions. the growing skeletal system adjusts." and Henrikson. The authors found that their regime resulted in: (I) delayed dental development and eruption. (2) exhibited no molar wear. the dentition migrates to accommodate. (4) had less width of the maxillary dental arch. (5) had smaller masseter and temporalis muscles.29 McCance.Diet Consistency jOcclusal the teeth adapted and once again occluded adequately. maintaining them for one year on a severe calorie-deficient diet. More recently. were smaller. 1981 Development 63 four studies have examined this idea experimentally. The condyles. although with no significant differences in shape. in older animals. where toothfood-tooth contact takes place. was less. which resist bite (especially incisal) reaction force. they do not create conditions likely to be encountered by the developing human masticatory system. the animals were maintained for approximately four months. was less in the soft-diet animals. All mandibular structures related to chewing were affected by diet hardness. which responded with less deposition and growth. where muscle force is transmitted to the mandible. In all studies. Beecher and Corruccini35 examined a small population of rhesus macaques which had spent a short period during adolescence on contrasting hard/soft diets. The smaller masticatory muscles in soft-diet animals were almost certainly a result of lower work requirement in mastication. Wear of the molar. before attempting to "rehabilitate" some with proper foods. the soft-diet animals: (I) were slightly smaller in body mass. Even buccolingual thickness of the mandible. In young animals. Green. Dietary consistency Although many workers have commented on the probable influence of the hardness or softness of the diet on the developing masticatory system. Tonge and McCance."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. While calorie deprivation is undoubtedly a factor to consider in severely undernourished people. which is essential to resist transverse bending. Nutrition Experiments concerning the contribution of nutrition to normal occlusal development have taken the form of calorie deprivation of pigs. and COX. The smaller and less-active muscles applied less stress to the bones." Barber.32 Moore. with condyles smaller and radiographically less dense. were smaller. While these experiments are important in assessing the response of the masticatory system to unusual mechanical stress. 1 January.

Body mass. evenly divided between males and females. Anteroposterior length of condylar articular surface. 5. engendering only subtle differences in bite force. This technique. The eigenvect. Ninety animals were divided into three groups: Group I was fed pelletted rat chow (Purina Formulab). it could not be determined whether the masticatory system responded in direct proportion to differences in dietary consistency. Group II I was fed the soft diet six days. MATERIALS AND METHODS The experimental animals were Sprague-Dawley rats. 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. 4.ors (also known as directional cosines or latent vectors) summarize the multiple interplay among variables. finds the major axes of the between-variable correlation matrix. Maxillary arch breadth across buccal points of ?vII. 2.64 Beecher and Corruccini than the gruel. It was to fill these gaps in the experimental record that the following study was carried out. with dry pellets provided every seventh day only. Croup 11 was fed a gruel-like porridge consisting of ground chow moistened with water. acquired at 21 days of age. 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. the same length of time used in previous experiments of this type. 6. We found the lab chow to be crumbly. related to factor analysis. The second axis is the axis perpendicular to the first that subsumes the largest possible amount of residual variance. 3. resulting in a lj uantum change in the morphology of the masticatory system. 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. Further. or whether no differences should be expected until a threshold in dietary consistency difference was reached. Animals were also examined for visible differences in tooth alignments and attritional differences between the three groups. with only two groups. A principal components analysis was performed on the data in order to discover t. 'While a number of measurements have been taken from animals in dietary consistency experiments. incisor to Ml.he correlations among the several variables. the heads were randomly numbered so that the measurer would not be aware of their group membership. Fresh mass of the entire masseter. 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. Maxillary arch length. This indicates that the "hard" diet was minimally harder . Mandibular length. The three populations were maintained on their respective diets for four months. easily breaking down into small granules. Measurements Crable I) were taken as follows: 1. 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.

. 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. Group I is more variable than Group II in every measurement.31 39.58) ( 0. 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. Dispersion.65) ( 0. Variable Maxillary Length (mm) .89 3. In particular.18 lAO Intermittent Mean (s.26 38. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance.. indicating a greater range of sizes. the second axis is largest among the softdiet animals.39 8.) 23.43 43.d. a result attributable to the small differences in dietary consistency.36) (9..43 ( 1.64 8.45 8.40) ( 0. All correlations are higher in the animals that had more masticatory resistance. Maxillary Breadth (mm) .Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures.d.. Group III is intermediate in standard deviation. Again. but not significantly so. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I).f. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix. Vol. The gross differences in the growth of most structures are not very marked. Thus the residual from Mean. Table 3 lists key paramo eters of this analysis.46) ( 0.20) ( 0.48) Soft Mean (s.. The strongest difference is in the growth relation of the masseter muscle. Masseter weight (gm) .39) (11.56 ( 1. • 3-sample TABLE I and Significance of Difference Hard Mean (s...38 2.) 24.21' 1.01) lAO ( 0.17) ( 0. Mandibular Length (mm) (mm) . 1981 .40) om AoV with 2 and 87 d. The major (coordinated growth) axis is largest in the hard-diet sample..1 January.~6) (0. Inversely related to this. For instance.. 51 No.57 4.03 1.44 at p (0.92 3.28) (or Measured Traits F 0.05.50) (0.91 3.7 I 9.00 9.. and functional interpretation of the morphological integration among variables. which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals. RESULTS Development 65 more variable than Group I in the maxillary arch measurements.72 1. 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). Condyle Length Body weight (gm) .79 9.) ~:I. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples.32) (0.41) (12.d. significant < . but usually closer to Group I.38) ( 0. although more similar to the soft group.88) (0. and the differences are often significant.

Body weight . TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max.55 0. at the stage of growrh... Thus.. Condyle length alone shows a notable residual on the second axis in Group I and III..5%.60 0.. 0..85 Groups Mussel.. The Angle Orthodontist . a decrease in correlation of masticatory structures is more noteworthy than de- = '.. IMaxillary Length ...72 Condo 0. This phenomenon has also been noted in characters that are diminishing in size through natural selection. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child....79· 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. Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals. which is significantly greater than expected by random error at P 2.. • Significantly larger Max..91· 0.93· 0. Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes.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...66 Body 0...37. Mandibular Length ..74 0.19. We noticed no displaced or rotated teeth in any of the animals. .. Perhaps a decrease in buffering among the growing components is the first step in their decrease in size. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth.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.39 diet sample at p :c ..54 0..78 0. the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample. Masseter weight ..L 0.89 0. 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.. No caries were observed by us in any animals.H3 0.83· 0.."] ""hile they noted easily observable differences in attrition between soft and hard diet groups.75· 0.. the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2.66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation. ~ e: 0..... condyle length...differentiation represented by this difference in dietary consistency.. Ii Mand... and masseter weight.79· 0. Condyle Length ..... Maxillary Breadth .. we could find none..71 Soft diet ~ r in hard < .. . In all respects.05.

experimental evidence which might weight such factors is absent.10 Vol.06 -0.82 0.92 0. 1 January.80 One Principal Component Hard Two growth) Intermittent 4.29 0. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control.12 0. with the maxillary teeth being more conservative and controlled by a fewer number of genes.95 (variation in growth) Soft Intermittent 0. premature deciduous tooth loss. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.68 0.50 (8) -0. Though the chewing explanation is most predictive of observed occlusal differences in the rats.67 (78) 0. oral habits.06 -0. trauma. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans. and independence of tooth formation.51 0.89 0.89 0.16 -0.77 0. Tooth spacing and crowding.91 0.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle. In particular.68 0. including mouth breathing.42-45 The growth independence of mandible and maxilla in humans has been frequently noted. follows different rates and peaks at different ages in the maxilla and mandible.57 -0.79 (13) -0. posture.93 0. prenatal insult." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. 1981 .59 -0.96 0.97 0.95 4. endocrine disturbance. 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 4.80 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.23 0. Because of its widespread occurrence in western society.90 0.18 -0.90 0. The aging process also yields different results in upper and lower arches.94 0.41•42 Some of these may be of minimal or no importance.w Moorrees and Reed.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion.96 (83) 0.08 0.05 -0.60 (10) -0. the mandible may depend on muscular function to grow to average size. disturbance of synchronous development in separately growing parts.73 -0.08 -0. especially of anterior teeth.04 0.14 0.17 (69) 0. hormonal intervention. 51 No.41 -0. Thus.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion.:" Maxillary growth may be under closer genetic programming. but does not give the source of the growth-correlation breakdown.

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