Eur J Oral Sci 2007; 115: 280287 Printed in Singapore.

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2007 The Authors. Journal compilation 2007 Eur J Oral Sci

European Journal of Oral Sciences

Early development of the lower deciduous dentition and oral vestibule in human embryos
Hovorakova M, Lesot H, Vonesch J-L, Peterka M, Peterkova R. Early development of the lower deciduous dentition and oral vestibule in human embryos. Eur J Oral Sci 2007; 115: 280287. 2007 The Authors. Journal compilation 2007 Eur J Oral Sci The aim of this work was to investigate the early development of the deciduous dentition and oral vestibule in the human embryonic lower jaw. Histological sections and three-dimensional reconstructions from prenatal weeks 69 were used. A continuous anlage for the oral vestibule did not exist in the mandible. In contrast to the upper jaw, where we previously observed that the dental and vestibular epithelia developed separately, two dento-vestibular bulges dierentiated in the incisor region of the mandible. The lingual parts of each bulge were found to give rise to the respective central and lateral incisors, whereas the labial parts dierentiated into the vestibular epithelium. In the canine and molar areas, the dental and vestibular epithelia originated separately. Later, the segments of the vestibular epithelium fused into the labial vestibular ridge, giving rise to the lower oral vestibule in the lip region. In the cheek region, the oral vestibule was found to originate in the mucosal inection between the developing jaw and the cheek. A similar heterogenous developmental base for the oral vestibule was also observed in the upper jaw. There is thus no general scheme for the early development of the dental and vestibular epithelia that applies to both the upper and lower jaws, and to both their anterior and posterior regions.

Maria Hovorakova1, Herve Lesot2,3, Jean-Luc Vonesch4, Miroslav Peterka1, Renata Peterkova1
1

Department of Teratology, Institute of Experimental Medicine, Academy of Sciences CR, Prague, Czech Republic; 2UMR INSERM 595, Strasbourg, France; 3Universite Louis Pasteur, Faculte de Chirurgie Dentaire, Strasbourg, France; 4Imaging Center Technology Platform, Institut de Genetique et de Biologie Moleculaire et Cellulaire, Illkirch, France

R. Peterkova, Department of Teratology, Institute of Experimental Medicine, Academy of Sciences CR, Videnska 1083, 142 20 Prague 4, Czech Republic Telefax: +420241062604 E-mail: repete@biomed.cas.cz Key words: deciduous tooth; dental epithelium; development; lower jaw; oral vestibule Accepted for publication May 2007

Tooth development in humans passes through the classical stages of odontogenesis: epithelial thickening; dental lamina; tooth bud; cap; and bell. Dierent authors have reported the rst morphological signs of odontogenesis in human embryos from embryonic day (ED) 28 (1) to ED 3440 (2). It is generally accepted that the upper and lower deciduous teeth develop from the respective horseshoe-shaped dental lamina (3, 4). Textbooks on human embryology present another horseshoeshaped epithelial structure that runs parallel and externally (labial or buccal) to the dental lamina in embryonic jaws (Fig. 1A). This is a vestibular lamina (also named the lip-furrow band or labio-gingival ridge), which develops between the dental lamina and the future lips and cheeks (58). The vestibular lamina is considered to be the developmental base of the oral vestibule, which is the free space between the alveolar portion of the jaws, including the teeth, and the lips or cheeks (6). Some textbooks do not mention the development of the oral vestibule at all (4, 9). Many authors have discussed the developmental relationship between the dental and vestibular laminae in humans, suggesting their independent development as two parallel epithelial anlagen (5, 6, 10), or that they arise from a common epithelial thickening (2, 7, 11), or that these laminae are subdivisions of a common thickened epithelial base in the anterior region and develop separately in the posterior region (12, 13).

However, three-dimensional (3D) imaging has documented that the formation of the oral vestibule is much more complex in the human upper embryonic jaw (14). No continuous horseshoe-shaped vestibular lamina exists. Instead, the upper vestibular epithelium includes a series of epithelial bulges and ridges (Fig. 1B). The dental and vestibular epithelia are regionalized in parallel along the mesio-distal axis. They reiteratively fuse together behind the developing upper deciduous canine, rst molar, and second molar (14). The aim of this study was to investigate the early morphogenesis and developmental relationships of the dental and vestibular epithelia in the human lower jaw, and to compare them with what has previously been observed in the upper jaw (14).

Material and methods

Embryos and fetuses The development of the dental epithelium was examined in a collection of serial histological sections held by the Department of Teratology, IEM AS CR, in Prague. The collection comprises 53 series of frontal and 7 series of sagittal histological sections of the heads of normal human embryos (from articially aborted embryos of unwanted pregnancies) compiled from the 1960s to the 1980s.

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Fig. 1. Schemes of the spatial relationship between the dental and vestibular epithelium in human embryos. Yellow, dental epithelium; orange, base of the oral vestibule. The remaining vestibular epithelium is green. (A) According to the generally accepted concept, the oral vestibule arises from a vestibular lamina (VL), which is a continuous structure running externally and parallel to the dental lamina (DL). (B) Data on the upper jaw (14) have shown epithelial bulges that initially emerge externally to the dental epithelium in the lip region. They fuse, being transformed into the canine vestibular ridge (CVR) in the posterior direction. The CVR fuses with the dental mound behind the deciduous canine primordium (c). In the cheek region, the vestibular epithelium forms the molar vestibular ridge (MVR) and the cheek-furrow ridge (the CFR is the epithelium lining the mucosal inection between the alveolus with teeth and cheeks). The MVR splits posteriorly into the medial (MMVR) and lateral (LMVR) branches. The fornix of the oral vestibule originates from the bulges and the CVR in the lip region and from the CFR in the cheek area (14). AC, accessory epithelial cap; m1 and m2, the upper rst and second deciduous molars, respectively. (C) Present data on the lower jaw. The level of the mouth corner (MC) is labeled. Dierent vestibular structures are shown: the area of irregularly thickened vestibular epithelium (ITVE), the labial vestibular ridge (LVR), and the mandibular cheek furrow ridge (dCFR). The deciduous lower tooth primordia are labeled: m1, rst molar; m2, second molar.

Table 1 The specimens used for three-dimensional reconstructions ranked according to the stage of tooth development.
Specimen number HU1 HU3 HU9 HU10 HU5 HU11 HU6 HU7 HU8 Secondary palate formation* /+ /+ /+ + + Developmental horizon (15) XX XXI XXII XXII XXII XXII XXII The upper and lower eyelids are not fused (2) The upper and lower eyelids are fused (2) Reconstructed lower jaw quadrant Right Right Left Right Left Right Right Right Left

Fixation Bouin-Hollande Formol 10% Bouin-Hollande Formol 10% Bouin-Hollande Bouin-Hollande Bouin-Hollande Bouin-Hollande Bouin-Hollande

Age ED 4042 ED 4244 ED 4446 ED 4446 ED 4446 ED 4446 ED 4446 Week 8 Week 9

Carnegie stage (16) 1617 17 18 18 18 18 18

ED, embryonic day *() Horizontalization of palatal shelves has not yet started; (/+) horizontalization started, one of the palatal shelves was horizontalized or both palatal shelves were horizontalized but not yet fused; (+) palatal shelves were fused. Only half of the embryonic head was available in which the palatal shelf was not yet horizontalized (it was not possible to determine the position of the second palatal shelf).

Determination of the embryonic stage As homogenous data allowing stage determination were not available for all specimens, the staging of embryos (younger than prenatal week 8) was checked and expressed on the basis of developmental horizons (15). The staging of fetuses (prenatal week 8 and older) was based on the morphological criteria proposed by Moore & Persaud (4). As a main criterion of staging, we used the development of the eye on frontal sections in all embryos and fetuses investigated. The staging was also correlated with the staging according to the Carnegie Collection (16). In addition, the

staging based on the classical methods was specied by the stage of tooth development. The tooth age allowed further ranking of a group of the embryos that exhibited the same stage determined by classical methods (see the alignment of embryos at ED 4446 in Table 1), as has been similarly documented in the mouse (17). Histology After xation in BouinHollande uid or in 10% formol, the heads were embedded in paran, cut in serial frontal sections (10 lm thickness), and stained by hematoxylin &

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Hovorakova et al. sional images were generated using a volume-rendering program (Sun Voxel Sun Microsystems, Santa Clara, CA, USA). In addition to dierent views on the 3D reconstructions, we also performed sagittal sections through the 3D reconstructions for complementary visualization of the relationships between the dental and vestibular epithelia.

eosin, alcian bluehematoxylin & eosin, or by a Periodic Acid Schi method. The early stages of tooth development (epithelial thickening, dental lamina, tooth bud) were identied according to criteria reported by Peterkova et al. (18). Computer-aided 3D reconstructions Three-dimensional reconstructions of the dental and adjacent oral epithelium of the lower jaw quadrant were performed in a representative sample of nine human embryos (Table 1). This sample showed a longitudinal adjustment of the successive steps of tooth development. In each of the embryos, the right or left jaw quadrant was randomly selected for 3D reconstruction. Contours of the dental and adjacent oral epithelium were drawn (magnication 120260, depending on the size of the specimen) at 10-lm intervals from each of the serial histological sections using a Leica DMLB (Leica Microsystems, Wetzlar, Germany) or a Jenaval (Carl Zeiss, Jena, Germany) microscope equipped with a drawing chamber. The superimposition of the drawings was performed by the best t method (19) with respect to the middle line and horizontal level for correct spatial positioning of the reconstructed structures. The digitalization of the serial drawings and the correlation of successive images have previously been described (20). Three-dimen-

Results
Morphology of the dental epithelium

In a jaw quadrant at ED 4042, two elds of thickened epithelium were present along the mesio-distal axis (Fig. 2A). These thickenings were situated in an end-toend orientation and separated by a deep notch (Fig. 2A). The adjacent ends showed a bulge shape corresponding to the regions of the deciduous lower lateral incisor (i2) and canine (c), respectively. Another bulge was observed in the region of the deciduous lower central incisor (i1) (Fig. 2A). The two bulges in the incisor region comprised the prospective dental and vestibular epithelia (dento-vestibular bulges). The lingual parts of these bulges protruded (prospective buds of deciduous i1 and i2), whereas the labial parts

Fig. 2. Computer-aided three-dimensional reconstructions of the epithelium on the oral surface of the lower jaw in HU1 at embryonic days (ED) 4042 (A), in HU3 at ED 4244 (B), and in HU10 at ED 4446 (C) at dierent stages of development (see Table 1) and their schematic interpretation. The deciduous lower tooth primordia are labeled: c, canine; i1, central incisor; i2, lateral incisor; and m1, rst molar. Dierent vestibular structures are shown: the labial vestibular ridge (LVR), the irregularly thickened vestibular epithelium (ITVE), and the epithelium lining the inection of the forming lower vestibule adjacent to the cheek region of the mandible (dCFR). The position of the mouth corner (MC) and ductus parotideus (DP) is indicated. The midline is shaded. The levels labeled hj indicate the positions of frontal histological sections shown in Fig. 4HJ, respectively.

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remained low (prospective vestibular epithelium). The bulge in the canine region corresponded to the primordium of c. From the canine region, a thickening of the dental epithelium extended in the distal direction (Fig. 2A), where the deciduous lower rst molar (m1) showed the bud stage. At ED 4244, the buds of i1 and i2 dierentiated from the lingual parts of the corresponding bulge. During further development, the labial parts of the two dentovestibular bulges in the incisor region gave rise to the vestibular epithelium. Further posteriorly along the course of the mandible, the dental epithelium formed a mound with swellings at the places of c and m1 (Fig. 2B). As tooth swellings, as well as the mound, showed a bud shape on frontal sections, it was not yet possible to delimitate tooth primordia at a bud stage. However, tooth primordia started to be distinct in developmentally more advanced embryos at ED 4446 (HU10 and older embryos; see Table 1, Fig. 2C), after the teeth reached a cap stage, whereas the mound epithelium in interdental spaces retained a bud-shape on frontal sections (Fig. 2C). During prenatal wk 8 the situation was similar to that at ED 4446. However, the tooth caps were more dierentiated (Figs 3A and 4). In the oldest examined
A

fetus (HU8), i1, i2, c, and m1 reached the bell stage of tooth development, and the cap of the deciduous second molar (m2) became distinct (Fig. 3B). At all stages, the dental epithelium divided into two parts behind the most distal developing tooth. The medial band disappeared after a short distance, whereas the lateral band continued in a disto-lateral direction (e.g. Figs 2B and 3B).
Morphology of the vestibular epithelium

The vestibular epithelium did not arise as a continuous vestibular lamina. While the bulges in the incisor area were formed by the dento-vestibular epithelium (ED 4044), the dental and vestibular epithelia developed separately in the caninemolar area. In the incisor area, the labial parts of the epithelial bulges progressively separated from i1 and i2 as the vestibular epithelium. In the canine and molar regions, the band of thickened vestibular epithelium was present externally to the dental epithelium at ED 4042 (Fig. 2A). In the cheek region, the area of the irregularly thickened vestibular epithelium (ITVE) dierentiated from ED 4244 (Figs 2, 3 and 4EG).

Fig. 3. Computer-aided three-dimensional reconstructions of the epithelium on the oral surface of the lower jaw in HU7 at prenatal wk 8 (A) and HU8 at prenatal wk 9 (B) at dierent stages of development (see Table 1) and their schematic interpretation. In (B) only the distal part of the long dental arch could be reconstructed because of the technical limits of the camera. The deciduous tooth primordia are labeled: c, canine; i1, central incisor; i2, lateral incisor; m1, rst molar; and m2, second molar. Dierent vestibular structures are shown: the labial vestibular ridge (LVR), the irregularly thickened vestibular epithelium (ITVE) and the epithelium lining the inection of the forming lower vestibule (dCFR). The position of the mouth corner (MC) and ductus parotideus (DP) is indicated. The midline is shaded. The levels labeled ag indicate the positions of frontal histological sections shown in Fig. 4AG, respectively.

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B

Fig. 4. Frontal histological sections showing the lower dental and vestibular epithelia at prenatal wk 8 (AG) and at embryonic days (ED) 4446 (HJ). The images shown in panels AJ were taken at the levels aj indicated in Figs 3A and 2C. The deciduous tooth primordia are labeled: c, canine; i1, central incisor; i2, lateral incisor; and m1, rst molar. The vestibular structures are shown: the labial vestibular ridge (LVR) and the irregularly thickened vestibular epithelium (ITVE). The branching (not visible on views on three-dimensional models in Figs 3A and 2C) at the posterior end of the dental epithelium is shown at prenatal wk 8 (EG) and at ED 4446 (HJ). Black arrowheads point to the dental epithelium. Bar, 100 lm.

At ED 4446, the vestibular and dental epithelia transiently fused in the canine region (Fig. 2C). Later, the vestibular epithelium again separated from the canine primordium and fused posteriorly with the ITVE and anteriorly with the vestibular epithelium in the incisor region (Fig. 3A). The fused vestibular epithelium in the incisor and canine regions and the medial part of the ITVE gave rise to the labial vestibular ridge (LVR) (Figs 2C, 3 and 4AE). In week 8, the LVR extended as far as to the m1 level. The LVR closely approached the dental mound anteriorly to m1 (Fig. 3A), but we did not observe a fusion. At ED 4042, an epithelial ridge dierentiated in the most lateral part of the cheek region of the mandible. This mandibular cheek furrow ridge (dCFR) remained located laterally to the ITVE at all subsequent stages under observation (Figs 2 and 3). The dCFR epithelium lined the mucosal inection (the prospective lower part of the oral vestibule) between the developing alveolus and the cheeks.

Discussion
Tooth development

On histological sections, the deciduous teeth passed through the typical stages of tooth development: the

thickening, lamina, bud, cap, and bell. However, the 3D reconstructions showed that deciduous tooth buds do not have the form of isolated buds protruding from a surrounding epithelium. At the so-called bud stage, the dental epithelium formed a dental mound, which showed a bud shape on frontal sections along the antero-posterior course of the mandible. At the places of developing canine and m1, only swellings were apparent on the mound in 3D reconstructions, corresponding to the increased size of the bud-shaped epithelium on frontal sections. This is why it was not possible to determine the exact limits of the deciduous tooth primordia before the cap stage (Figs 2 and 3A). Such morphology of the tooth primordia at the bud stage has also been documented in the human upper jaw (14) and during mouse molar development (18, 21). It has been reported that the epithelial anlage of the deciduous dentition originates at the same time in the upper and lower jaws (22). However, a comparison of our previous (14) and present data supports the suggestion that the development of the upper teeth lags behind the development of the lower dentition (5, 10). This lag has been related to the complex development in the upper portion of the face (brain, sense organs, nasal cavity), which is also in progress at the time of tooth development (5). The unbalanced development of the upper and lower dentitions can also be explained by the fact that the maxillary and mandibular branches of the rst branchial arch grow unequally in time and size (23). A method of reciprocal cell transplantations between maxillary and mandibular arch ectomesenchymal cells revealed intrinsic dierences between these populations of cranial neural crest-derived cells in the mouse, which respond dierently to epithelial signals. These dierences have been related to the dierent origins of the neural crest cells that populate these components of the rst branchial arch (24). In the axolotl, direct cell lineage analysis in vivo has shown that there are two independent condensations of neural crest cells owing into the maxillary and mandibular outgrowths, respectively (25). In human functional dentition, the deciduous teeth are located next to one another and the interdental spaces are negligible. However, signicant spaces separate individual tooth germs during development (Figs 2 and 3). This means that the growth rate will have to be higher in the tooth germs than in the interdental spaces until the tooth reaches the bell stage. Generally, the interdental spaces between the developing anterior teeth (incisors) are shorter in comparison to the interdental spaces in the distal regions (canine and molars) in both upper and lower jaws (26, 27). The most signicant interdental space was present between the canine and the rst molar in both the upper (14) and lower (Figs 2 and 3) jaws. This can be explained by the reduction in tooth number during human evolution. The basic mammalian tooth formula comprises 3 incisors, 1 canine, 4 premolars, and 3 molars (28). The human deciduous rst and second molars have been homologized with the respective deciduous third and

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fourth premolars from the basic mammalian dentition (29). The missing rst and second premolars might explain the presence of a signicant interdental space between the canine and the rst deciduous molar ( third deciduous premolar) in the human embryonic dental arch.
The vestibular epithelium and the prospective vestibule of the mouth

There was always a protrusion of the vestibular epithelium externally to the dental epithelium on histological sections (Fig. 4). Without correlation with the corresponding 3D views, this protrusion could be considered as a component of a continuous horseshoe-shaped vestibular lamina, giving rise to the prospective oral vestibule (Fig. 1A). The concept of the development of the oral vestibule from a continuous vestibular lamina has been presented by many authors (1,5) and is still maintained in embryological textbooks (68). However, the present 3D reconstructions did not show any continuous vestibular lamina in the lower jaw (Figs 2 and 3). In its place, there were several discontinuous epithelial structures (Fig. 1C) as already observed in the upper jaw (14) (Fig. 1B). The vestibular epithelium adjacent to the incisor primordia was segmented in parallel to individual teeth in both the upper (14) and lower jaws (Fig. 1). However, there was an apparent dierence: in the upper incisor

region, the segments of the dental and vestibular epithelia arise separately (14), whereas in the lower incisor region, there were two segments (bulges), each comprising precursors of the prospective incisor and its adjacent vestibule (Figs 1 and 5). In the canine and cheek region of both the upper (14) and lower jaws, the dental and vestibular epithelia arose independently (Figs 1, 2 and 5). The lower oral vestibule had dierent developmental origins in the lip and cheek regions (Fig. 1C). In the lip region, it developed from the labial vestibular ridge, which presumably corresponds to the lip-furrow-band of, for example Schour (5), or with the labio-gingival sheet of Bolk (11). However, in the cheek region of the mandible (located behind the level of the mouth corner), the oral vestibule originated from the epithelial ridge that lines the inection between the cheeks and the prospective alveolus (Fig. 1C). The distinct origins of the fornix of the oral vestibule in the lip and cheek regions have also been reported in the human upper jaw (14) (Fig. 1B), in mice (30), and in sheep (31). The developmental relationship between the dentition (dental lamina) and the oral vestibule (vestibular lamina) has not been completely explained in the literature. It has been stated that these structures: (i) evolve separately (5, 6, 10); (ii) have a common origin (2, 7, 11); or (iii) are subdivisions of a common thickened epithelial base anteriorly, whereas they develop as two separate epithelial structures in the posterior regions of the jaw (12, 13).

Fig. 5. Comparison of the upper and lower dental (DE) and vestibular (VE) epithelia on three-dimensional (3D) reconstructions. The DE and VE of the upper (A, B) and lower (C, D) jaws of the embryo HU2 [embryonic days (ED) 4244] are presented as an aerial view on the 3D reconstruction (A, C) and on a sagittal section through the 3D reconstruction (B, D). In the upper incisor region, DE and VE dierentiate apart from each other, being separated by a groove indicated by a blue arrow (A, B). However, in the lower jaw, one bulge formed by the dento-vestibular epithelium (DVE) is present in each of the central and lateral incisor regions (C, D). At later stages, the lingual parts of the two bulges give rise to i1 and i2, respectively. The labial parts dierentiate into the vestibular epithelium. The midline is shaded.

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These dierent concepts might result from observations performed by dierent authors on dierent jaws (upper or lower) and jaw regions (lip or cheek region) in embryos at dierent developmental stages. Our data clearly show that each of these concepts correctly reects only one particular situation, depending on the region and the time of observation (Figs 1 and 5). In the upper jaw, there are two separate epithelial thickenings: the inner dental and outer vestibular epithelia (14). This observation corresponds with the rst concept mentioned above, proposing the independent development of the dentition and oral vestibule (5, 6, 10). Nevertheless, the parallel segmentation of the dental and vestibular epithelia in the upper jaw suggests their close developmental relationship (14). The situation was completely dierent in the lower jaw (Fig. 1C) compared with the upper jaw (Fig. 1B). The dental and vestibular epithelia developed from a common dento-vestibular bulge, which was present in both the central and lateral incisor regions (Figs 2 and 5). This situation ts with the second concept that suggests a common origin of the dentition and the oral vestibule (2, 7, 11). In the posterior (canine and cheek) region of the lower jaw, the dental and vestibular epithelia originated separately (in accordance with the rst concept above) (Figs 1C and 2). So, the present data on the lower jaw correspond with the third above-mentioned concept regarding the common origin of the dentition and the oral vestibule in the anterior region and their separate origin in the posterior region of the jaw (12, 13). The dental and vestibular epithelia develop separately on the fused facial processes forming the upper jaw arch (14). However, if traced to earlier stages, they might also have a common origin in the epithelial plate that has been observed on each of not-yet merged facial outgrowths in human embryos (1, 32). If suciently young mouse embryos are investigated, a similar common developmental origin can be traced out, giving rise not only to teeth and oral vestibule, but even to the palatal ridges. Based on this ontogenetic relationship, the palatal ridges and the oral vestibule have been presented as odontogenous structures from a phylogenetic aspect (30). The latent odontogenous potential of the vestibular epithelium can be further supported by the integration of the vestibular epithelium into the dental lamina in the mouse (30) and in humans (14). It has been suggested that the structures of the vestibular epithelium in mammals might have an evolutionary relationship to the earliest generations of teeth in reptiles (33). The complex development of the oral vestibule might help to explain some of the pathologies located externally to the functional dentition.
Acknowledgements We thank Prof. J. Slipka (Institute Histol. Embryol., Medical Fac., Charles University, Pilsen, CR) for providing a part of human embryological material and Mr J. Dutt for critical reading of the manuscript. This study was supported by the Grant Agency of the Czech Republic (304/05/ 2665), Ministry of Education, Youth, and Sports of the Czech Republic (project COST B23.002) and by the Academy of Sciences of the Czech Republic (project AV0Z 50390512).

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