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Effect of Diet on Growth

Effect of Diet on Growth

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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

or accumulation of genetic mutations. This movement resulted in short-term periodontal traumas which abated when The Angle Orthodontist . A cornman habit in childhood.> Disrupted occlusion The prevalence of an occlusion that varies from the ideal is not the norm among mammals. Mandibular protrusion in young animals resulted in changes in the temporomandibular joint.vSuch effects are common in humans. racial outcrossing.0 mm in order to achieve molar occlusion. While the younger. growing animals responded to this stress by skeletal adaptations. (3) lengthened condylar neck. older animals responded with tooth movements to adjust to the new occlusal alignments. specifically: (1) some posterior bone deposition. Mouth breathing Mouth breathing has been implicated experimentally (using macaques) in the development of open bite and or an abnormally narrow maxillary dental arch. Using mature animals and a splint on the anterior teeth to force mandibular protrusion. especially the physical consistency of the diet. the experimental attempts to induce developmental change in the mammalian masticatory apparatus have used four methods': (I) forcing mouth breathing in animals. 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. Hiniker and RamsIjord= also found dentitional changes as teeth were forced to move to maintain occlusal relationships. mouth breathing is probably a factor in the development of human malocclusions. The maxillae of these animals showed reduced vertical and prognathic growth. inbreeding. affect the developing masticatory apparatus and create variations in the occlusal phenotype.62 Beecher and Corruccini probably due to a reduction in muscle use rather than increased. The rapid increase in the incidence of malocclusion (often within one generation) gives cause to look beyond genetic explanations such as relaxed selection pressures. (2) placing devices in the mouth to effect alterations in mastication. Experimen Is in effects of function Apart from radical surgery. 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.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. (2) bone resorption of the anterior edges of the glenoid fossa. attributed to altered muscle tension as the mandible is held slightly depressed during respiration. abnormally-directed muscle tension. Similar differences were also found in soft-diet rats. and (4) varying the consistency of the diet from hard to soft. These are usually designed to force protrusion of the mandible 1. 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 2.5-2. we believe that environmental factors. Based on the available anthropological and experimental evidence. (3) restricting animals to calorie-deprivation diets.

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

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

) 23.43 43. Group III is slightly A much more obvious feature of difference is manifested in the COl'relation structure between measurements within samples. this measurement alone shows a differentiation that is too large (relative to within-sample variation) to be reasonably ascribed to chance. 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. but usually closer to Group I. signifying that all structures are more tightly integrated in their growth and covariation..~6) (0. The comparison of correlation structure can be more easily accomplished through examination of the principal components of the correlation matrix. RESULTS Development 65 more variable than Group I in the maxillary arch measurements. although more similar to the soft group. the maxillary breadth is markedly increased in the hard-diet reared animals (Group I). The gross differences in the growth of most structures are not very marked.41) (12. Masseter weight (gm) . which is much more integrated with growth in the condyle and upper arch breadth in hard-diet than in soft-diet animals. Again.89 3.21' 1.. Maxillary Breadth (mm) .Diet Consistency jOcclusal vidual cases onto the principal component axes allows interrpretation of multivariate variabilitv within samples. Group III is intermediate in standard deviation. Table 3 lists key paramo eters of this analysis. significant < .50) (0. The major (coordinated growth) axis is largest in the hard-diet sample.) ~:I.18 lAO Intermittent Mean (s..39 8.. Condyle Length Body weight (gm) .28) (or Measured Traits F 0.d. Thus the residual from Mean. Vol.58) ( 0.91 3.d.64 8.03 1. Mandibular Length (mm) (mm) .17) ( 0..44 at p (0.26 38. Dispersion. 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.05. The strongest difference is in the growth relation of the masseter muscle.46) ( 0.39) (11.65) ( 0. Variable Maxillary Length (mm) .20) ( 0.38 2.38) ( 0. For instance. Table 2 indicates the pairwise correlations in the hard and soft diet groups (I and II).36) (9. • 3-sample TABLE I and Significance of Difference Hard Mean (s... All correlations are higher in the animals that had more masticatory resistance... 1981 .88) (0..72 1.57 4. and functional interpretation of the morphological integration among variables.7 I 9.f.01) lAO ( 0.) 24. Inversely related to this. The population reared on an intermittently hard diet (Group III) is generally intermediate in the size of structures. and the differences are often significant.56 ( 1.32) (0.79 9.1 January.43 ( 1. the second axis is largest among the softdiet animals.00 9.40) om AoV with 2 and 87 d.d.40) ( 0. In particular.45 8. a result attributable to the small differences in dietary consistency.48) Soft Mean (s. 51 No.92 3.31 39. indicating a greater range of sizes. Group I is more variable than Group II in every measurement. but not significantly so.

.. Only in the 19th and 20th centuries (il apIJcars) has food become so processed tha t for practical purposes..75· 0. ~ e: 0... we could find none..91· 0...L 0...72 Condo 0.. TABLE 2 Product-moment Correlations Among Variables Within the Soft (lower triangular half of matrix) and Hard (upper triangular half) Dietary Max. Maxillary Breadth . which is significantly greater than expected by random error at P 2..93· 0. condyle length....... the intermittent-harddiet group (III) is intermediate when compared with Groups I and 11 but more similar to the hard-diet sample....79· 0. . Masseter weight .05. IMaxillary Length . 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.5%. . Masseter weight tends to be inversely correlated with maxillary breadth and condyle length on the second axis in soft animals.....37.66 Beecher and Corruccini crease in size of the structures that seems to follow with a stronger dietary differentiation.. Perhaps a decrease in buffering among the growing components is the first step in their decrease in size..54 0..39 diet sample at p :c .85 Groups Mussel.66 Body 0.. most chewing stress has been removed and little bite force is now called into play in the jaws of the growing child..differentiation represented by this difference in dietary consistency. No caries were observed by us in any animals.. The correlations of variables with the axes show that the breakdown in growth coordination in soft-diet animals is especially marked in maxillary breadth. Body weight .. Ii Mand. This phenomenon has also been noted in characters that are diminishing in size through natural selection. Condyle length alone shows a notable residual on the second axis in Group I and III. Condyle Length .83· 0.. In all respects.."] ""hile they noted easily observable differences in attrition between soft and hard diet groups. a decrease in correlation of masticatory structures is more noteworthy than de- = '.H3 0.60 0.79· 0.. the F-ratio of variation on axis two (dispersion from axis one) between soft and hard samples is 2. • Significantly larger Max..74 0. and masseter weight.. 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.89 0...19. Mandibular Length .78 0.. at the stage of growrh.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.71 Soft diet ~ r in hard < .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.. Thus. 0...55 0. The Angle Orthodontist . We noticed no displaced or rotated teeth in any of the animals.

41•42 Some of these may be of minimal or no importance.94 0.06 -0.05 -0. and independence of tooth formation.50 (8) -0. Potter et alY demonstrate that upper and lower dental arches are under very different sorts of genetic control.:" Maxillary growth may be under closer genetic programming.95 (variation in growth) Soft Intermittent 0.79 (13) -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 4.18 -0.12 0.57 -0. disturbance of synchronous development in separately growing parts.42-45 The growth independence of mandible and maxilla in humans has been frequently noted. especially of anterior teeth.82 0.w Moorrees and Reed.w Carrr" lists breakdown of synchronous growth in lower and upper jaws as a possible cause of malocclusion. Thus. it clearly is not favored by many orthodontists as an explanation of the rising rate of occlusal variation in humans. as shown by the tendency of some indi- Development 67 viduals with anodontia to have average-size mandibles.97 0. but does not give the source of the growth-correlation breakdown.23 0. posture. 51 No. with the maxillary teeth being more conservative and controlled by a fewer number of genes.96 0. Tooth spacing and crowding. endocrine disturbance.e-v Lower correlation between upper and lower teeth in maloccluded individuals (as compared with normal) has been noted by Lavelle.68 0. The aging process also yields different results in upper and lower arches." also stress the importance of high correlation between tooth and arch size to the development of good occlusion. the mandible may depend on muscular function to grow to average size.60 (10) -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.59 -0. 1 January.67 (78) 0.89 0. Though the chewing explanation is most predictive of observed occlusal differences in the rats. follows different rates and peaks at different ages in the maxilla and mandible.90 0.16 -0.04 0.08 0. 1981 . trauma.91 0.Diet Consistency/Occlusal A number of other environmental factors have been suggested as influences on developing occlusion.92 0. since cases of anodontia show restricted growth while cases with muscular paralysis may develop normally.77 0. Because of its widespread occurrence in western society.73 -0. including mouth breathing.89 0.80 One Principal Component Hard Two growth) Intermittent 4.90 0. oral habits.10 Vol.06 -0.14 0.68 0.93 0.80 0. In particular.96 (83) 0. prenatal insult.17 (69) 0.51 0.08 -0. premature deciduous tooth loss. hormonal intervention. experimental evidence which might weight such factors is absent.29 0.41 -0.95 4.

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