Odontogenic Research Articles

Unlocking Osseointegration: Surface Engineering Strategies for Enhanced Dental Implant Integration.

Tooth loss is a prevalent problem faced by individuals of all ages across the globe. Various biomaterials, such as metals, bioceramics, polymers, composites of ceramics and polymers, etc., have been used for the manufacturing of dental implants. The success of a dental implant primarily depends on its osseointegration rate. The current surface modification techniques fail to imbibe the basics of tooth development, which can impart better mineralization and osseointegration. This can be improved by developing an understanding of the developmental pathways of dental tissue. Stimulating the correct signaling pathways through inductive material systems can bring about a paradigm shift in dental implant materials. The current review focuses on the developmental pathway and mineralization process that happen during tooth formation and how surface modifications can help in biomimetic mineralization, thereby enhancing osseointegration. We further describe the effect of dental implant surface modifications on mineralization, osteoinduction, and osseointegration; both in vitro and in vivo. The review will help us to understand the natural process of teeth development and mineralization and how the surface properties of dental implants can be further improved to mimic teeth development, in turn increasing osseointegration.

Root Growth and Apical Foramen Closure Velocity of Maxillary Permanent Central Incisor in Korean Children

This study aimed to evaluate the growth velocity of root elongation and the timing of apical foramen closure in maxillary permanent central incisors using panoramic radiographs. The study also examined the timing of alveolar eruption and its relationship with root development while considering potential sex-specific differences. The study included 176 patients (94 males, 82 females) from the Seoul National University Dental Hospital, each with three or more panoramic radiographs taken between 2014 and 2023 and analyzed to measure the crown length, total tooth length, root length, and mesiodistal width of the apical foramen. Root lengths were expressed as percentages of the fully developed stage. Scatter plots and polynomial trend lines were used to evaluate the relationship between age and tooth development, with intraobserver reliability assessed using Cohen’ s kappa. Root growth velocity peaked during the early stages of development, with females generally exhibiting earlier and faster apical foramen closure than males. Significant correlations were found between the root length and apical foramen width (Pearson correlation coefficients, males, -0.907; females, -0.887, <i>p</i> < 0.0001). The mean age for alveolar eruption was 7.27 years for males (N = 16) and 7.10 for females (N = 17), predominantly at Demirjian stage F. The study presents the critical periods of rapid root elongation and apical foramen closure in the maxillary central incisors, underscoring their importance for clinical dental practice. Future research is warranted to provide a more comprehensive understanding of human dental development.

The Shh-p38-NFATc1 signaling pathway is essential for osteoclastogenesis during tooth eruption

Tooth eruption, a critical stage in tooth development, is related to osteoclastogenesis. Intraperitoneal injection of Shh agonists into neonatal mice promoted tooth eruption at postnatal day (PN) 15, whereas treatment with the Shh inhibitor (LDE225) suppressed this process. When RAW264.7 osteoclast precursor cells were treated with RANKL, NFATc1 translocated from the cytoplasm to the nucleus and induced cell differentiation into TRAP+ osteoclasts; this process was activated by Shh but inhibited by LDE225. Treating RAW264.7 cells with the p38 inhibitor, BIRB796, also inhibited NFATc1 nuclear localization. p-p38 expression in the alveolar bone of PN3 and PN5 mice was decreased by treatment with LDE225, and RAW264.7 cell differentiation was reduced by BIRB796, regardless of treatment with Shh. Furthermore, Shh activated p38 signaling pathway in RAW264.7 cells, while p38 phosphorylation was reduced by LDE225, which ultimately inhibited osteoclast precursor differentiation. Therefore, we concluded that Shh promotes osteoclast precursor differentiation via the p38-NFATc1 signaling pathway.

Polycomb protein Bmi1 promotes odontoblast differentiation by accelerating Wnt and BMP signaling pathways.

Bmi1 is a polycomb protein localized in stem cells and maintains their stemness. This protein is also reported to regulate the expression of various differentiation genes. In this study, to analyze the role of Bmi1 during dentinogenesis, we examined the immunohistochemical localization of Bmi1 during rat tooth development as well as after cavity preparation. Bmi1 localization was hardly detected in the dental mesenchyme at the bud and cap stages. After the bell stage, however, this protein became detectable in preodontoblasts and early odontoblasts just beginning dentin matrix secretion. As dentin formation progressed, Bmi1 immunoreactivity in the odontoblasts decreased in intensity. After cavity preparation, cells lining the dentin and some pulp cells under the cavity were immunopositive for Bmi1 at 4days. Odontoblast-like cells forming reparative dentin were immunopositive for Bmi1 at 1week, whereas their immunoreactivity was not detected after 8weeks. We further analyzed the function of Bmi1 using KN-3 cells, a dental mesenchymal cell line. Overexpression of Bmi1 in KN-3 cells promoted mineralized tissue formation. In contrast, siRNA knockdown of Bmi1 in KN-3 cells reduced alkaline phosphatase activity and the expression of odontoblast differentiation marker genes such as Runx2, osterix, and osteocalcin. Additionally, KN-3 cells transfected with siRNA against Bmi1 showed reduced nuclear transition of β-catenin and expression of phosphorylated-Smad1/5/8. Taken together, these findings suggest that Bmi1 was localized in the odontoblast-lineage cells in their early differentiation stages. Bmi1 might positively regulate their differentiation by accelerating Wnt and BMP signaling pathways.

Novel MSX1 Gene Variants in Chinese Children with Non-Syndromic Tooth Agenesis: A Clinical and Genetic Analysis

Background: Tooth agenesis is the most frequently occurring genetic developmental anomaly in clinical dentistry. The MSX1 gene, essential for tooth development, has been associated with non-syndromic tooth agenesis. This study aims to identify novel MSX1 variants associated with this condition and to understand their impact on tooth development. Methods: This study involved the genetic analysis of two children presenting with non-syndromic tooth agenesis. Conservation analysis and 3D structural modeling were conducted to assess the pathogenicity of these variants. Additionally, a review of 108 patients with known MSX1 variants was performed to identify patterns of tooth agenesis. Results: We discovered two novel MSX1 variants, c.823 T>G and c.890 A>G, located in the second exon of the MSX1 gene. The identified MSX1 variants, c.823 T>G and c.890 A>G, were predicted to be pathogenic. Conservation analysis showed that the impacted amino acids are highly conserved across species, and 3D structural analysis indicated potential disruptions to protein function. Among the 108 patients reviewed, a consistent pattern of tooth agenesis was observed, with the most frequently missing teeth being the maxillary second premolars, the mandibular second premolars, and the maxillary first premolars. Conclusions: This research broadens the known range of MSX1 gene variants and deepens our comprehension of the genetic foundations of non-syndromic tooth agenesis. The findings provide valuable insights for genetic counseling and future research into tooth development, emphasizing the importance of MSX1 in dental anomalies.

Localization of α-smooth muscle actin in osteoblast differentiation during periodontal development.

α-Smooth muscle actin (α-SMA) is an actin isoform commonly found within vascular smooth muscle cells. Moreover, α-SMA-positive cells are localized in the dental follicle (DF). DF is derived from alveolar bone (AB), cementum, and periodontal ligament (PDL). Therefore, α-SMA-positive cells in the periodontal tissue are speculated to be a marker for mesenchymal stem cells during tooth development. In particular, the mechanism of osteoblast differentiation is not clear. This study demonstrated the fate of α-SMA-positive cells around the tooth germ immunohistochemically. First, α-SMA- and Runx2-positive localization at embryonic days (E) 13, E14, postnatal days (P) 9, and P15 was demonstrated. α-SMA- and Runx2-positive cells were detected in the upper part of the DF at P1. At P9 and P15, α-SMA-positive cells in the PDL were detected in the upper and lower parts. The positive reaction of Runx2 was also localized in the PDL. Then, the distribution of α-SMA-positive cell progeny at P9 and P15 were clarified using α-SMA-CreERT2/ROSA26-loxP-stop-loxP-tdTomato (α-SMA/tomato) mice. It has known that Runx2-positive cells differentiate into osteoblasts. In this study, some Runx2 and α-SMA-positive cells were localized in the DF and PDL. The lineage-tracing analysis demonstrated that the α-SMA/tomato-positive cells expressing Runx2 or Osterix were detected on the AB surface at P15. α-SMA/tomato-positive cells expressing type I collagen were found in the AB matrix. These results indicate that the progeny of the α-SMA-positive cells in the DF could differentiate into osteogenic cells. In conclusion, α-SMA could be a potential marker of progenitor cells that differentiate into osteoblasts.

Functional and pathogenic insights into CNNM4 variants in Jalili syndrome

Jalili syndrome, an autosomal recessive disorder causing cone-rod dystrophy and amelogenesis imperfecta, is a rare genetic disorder impacting visual and dental development. Missense variants (c.1474G > T and c.1475G > A) previously identified in patients with Jalili syndrome have been linked to functional impairment of CNNM4, however, the biological consequences of these pathogenic variants remain largely unexplored. In this study, we investigated the functional implications of these CNNM4 missense variants, which correspond to p.(Gly492Cys) and p.(Gly492Asp) substitutions within the CBS domain of the CNNM4 protein. Our findings demonstrated that these variants exhibit significantly reduced protein stability and increased mRNA decay rates compared with wild type. Despite exhibiting normal Mg2+ localization, the mutant proteins demonstrated significantly reduced Mg²⁺ extrusion activity. This suggests that the pathogenic mechanism underlying Jalili syndrome associated with these variants likely involves decreased mRNA and/or protein stability, rather than mislocalization. Our study provides valuable insights into the interplay between genetic variations, molecular stability, and functional consequences in the context of CNNM4-related disorders, highlighting the importance of CNNM4-mediated Mg²⁺ transport in Jalili syndrome. Further investigation into the mechanisms regulating CNNM4 expression and protein stability may reveal potential therapeutic avenues.

Radiographic Assessment of Root/Crown Ratio and Short Root Anomaly: A Retrospective Study in Chinese Population.

Research suggested that racial and gender differences may exist in tooth root development and the prevalence of short root anomaly (SRA). However, data on Asian and specifically Chinese populations were limited. This retrospective study aimed to evaluate the prevalence of SRA and the characteristics of the root-to-crown ratio (R/C ratio) within the Chinese population. Modified Lind's criteria were used to measure the crown and root lengths in permanent dentition. A total of 27 142 teeth from 1000 subjects were evaluated using panoramic radiographs. Fisher's exact test was employed to analyse gender differences in SRA. Paired-sample and independent-sample t tests were used to assess the characteristics of R/C ratios. SRA was observed in 27 patients, with a prevalence of 2.7% and a tooth prevalence of 0.309%. Females had a significantly higher prevalence of SRA compared to males (p < 0.01). SRA was found exclusively in incisors (83.72%) and premolars (16.28%), with the maxillary central incisor being the most commonly affected tooth. Additionally, there were statistical differences in the R/C ratio based on gender and jaw position in the normal groups. The highest R/C ratios were observed in the mandibular second premolars, while the maxillary central incisors had the lowest ratios. SRA was relatively common in the Chinese population, and clinicians should be aware of its presence. It was more frequently observed in females and shows specificity in tooth position, with the maxillary central incisor being the most commonly affected tooth. Our study provides valuable insights for the early diagnosis and treatment of SRA in the Chinese population.

Assessing optimal timing for orthodontic intervention: A scoping review

There is still debate over the ideal time to begin orthodontic treatment. To optimize growth potential and patient cooperation, definitive treatment typically is delayed until the late mixed dentition and is used to resolve functional issues and anomalies of dental development. However, for specific types of malocclusion, some medical professionals advise beginning treatment earlier. The degree of the malocclusion, the patient's age and level of development at the time the patient sought treatment, and the timing of the treatment interventions all had an impact. This article explains over all review of the timing of the orthodontic treatment.

Sensors: future tools for detecting young patient's stress during a dental invasive versus a non-invasive dental treatment-a pilot study.

A reliable tool to visualise children's early stress signs to prevent dental fear development is needed. The aim was to evaluate the commercially available, CE marked, Shimmer3 GSR + unit's ability to indicate for stress as a reaction of fear or pain for a non-invasive dental treatment (NI) and an invasive dental treatment (I). Patients 14-16years old were invited to undergo an oral check-up (NI) or an orthodontic premolar extraction (I), respectively. Digital data, measured via electrodes and optical pulse probe, placed on the wrist and fingers, monitored by the Shimmer3 GSR + unit, was transferred via Bluetooth to the HP-laptop. The observed digital parameters were: heart rate based on photoplethysmography (PPG), galvanic skin response (GSR), and 3-axis gyroscope and accelerometer signals for hand movements. Protocols for patient self-report scales were used: coloured analogue scale for pain intensity, facial analogue scale for the mood, and a dental fear scale. Descriptive statistics was performed. The NI-group: 20 patients, (14.6 ± 0.5years), underwent 20 oral check-ups. The I-group: 14 patients, (15.3 ± 0.5years), underwent 28 premolar extractions. All patients tolerated the Shimmer3 GSR + unit well. The GSR signal increased significantly, at start and during the oral injection, in the I-group. The GSR amplitudes persisted throughout and post the dental injection. No general uniform pattern or high GSR amplitudes were produced regarding NI-group. Considering the limitations of this study, the following conclusions can be made: the invasive treatment resulted in a specific unison GSR pattern, while the non-invasive procedure showed individually scattered GSR reactions. The commercially available CE-marked Shimmer3 GSR + device indicated the patient's stress response triggered by the invasive anaesthetic procedure.

Dental evidence for extended growth in early Homo from Dmanisi

Human life history is characterized by an extended period of immaturity during which there is a disjunction between cerebral and somatic growth rates1. This mode of ontogeny is thought to be essential for the acquisition of advanced cognitive capabilities in a socially complex environment while the brain is still growing2. Key information about when and how this pattern evolved can be gleaned from the teeth of fossil hominins because dental development informs about the pace of life history3–5. Here we show that the first evolutionary steps towards an extended growth phase occurred in the genus Homo at least 1.77 million years ago, before any substantial increase in brain size. We used synchrotron phase-contrast tomography6 to track the microstructural development of the dentition of a subadult early Homo individual from Dmanisi, Georgia. The individual died at the age of 11.4 ± 0.6 years, shortly before reaching dental maturity. Tooth growth rates were high, similar to rates in living great apes. However, the Dmanisi individual showed a human-like delayed formation of the posterior relative to the anterior dentition, and a late growth spurt of the dentition as a whole. The unique combination of great-ape-like and human-like features of dental ontogeny suggests that early Homo had evolved an extended growth phase before a general slow-down in life history, possibly related to biocultural reproduction7 rather than brain growth.

Generating Tooth Organoids Using Defined Bioorthogonally Cross-Linked Hydrogels.

Generating teeth in vitro requires mimicking tooth developmental processes. Biomaterials are essential to support 3D tooth organoid formation, but their properties must be finely tuned to achieve the required biomimicry for tooth development. For the first time, we used bioorthogonally cross-linked hydrogels as defined 3D matrixes for tooth developmental engineering, and we highlighted how their properties play a pivotal role in enabling 3D tooth organoid formation in vitro. We prepared hydrogels by mixing gelatin precursors modified either with tetrazine (Tz) or norbornene (Nb) moieties. We tuned the hydrogel properties (E = 2-7 kPa; G' = 500-1500 Pa) by varying the gelatin concentration (8% vs 12% w/V) and stoichiometric ratio (Tz:Nb = 1 vs 0.5). We encapsulated dental epithelial-mesenchymal cell pellets in a library of hydrogels and identified a hydrogel formulation that enabled successful growth kinetics and morphogenesis of tooth germs, introducing a defined tunable platform for tooth organoid engineering and modeling.

Metalloproteinases are involved in the regulation of prenatal tooth morphogenesis.

During development, tooth germs undergo various morphological changes resulting from interactions between the oral epithelium and ectomesenchyme. These processes are influenced by the extracellular matrix, the composition of which, along with cell adhesion and signalling, is regulated by metalloproteinases. Notably, these include matrix metalloproteinases (MMPs), a disintegrin and metalloproteinases (ADAMs), and a disintegrin and metalloproteinases with thrombospondin motifs (ADAMTSs). Our analysis of previously published scRNAseq datasets highlights that these metalloproteinases show dynamic expression patterns during tooth development, with expression in a wide range of cell types, suggesting multiple roles in tooth morphogenesis. To investigate this, Marimastat, a broad-spectrum inhibitor of MMPs, ADAMs, and ADAMTSs, was applied to ex-vivo cultures of mouse molar tooth germs. The treated samples exhibited significant changes in tooth germ size and morphology, including an overall reduction in size and an inversion of the typical bell shape. The cervical loop failed to extend, and the central area of the inner enamel epithelium protruded. Marimastat treatment also disrupted proliferation, cell polarization and organization compared to control tooth germs. Additionally, a decrease in laminin expression was observed, leading to a disruption in continuity of the basement membrane at the epithelial-mesenchymal junction. Elevated Hif-1α expression correlated with a disruption to blood vessel development around the tooth germs. These results reveal the crucial role of metalloproteinases in tooth growth, shape, cervical loop elongation, and the regulation of blood vessel formation during prenatal tooth development.

Expression Of Nuclear Factor IC And Detection Of Calcium During Odontogenesis And Odontogenic Tumors

Background: Tooth development is a complex and well-coordinated developmental process that is achieved through a series of reciprocal interactions between dental epithelium and neural crest-derived ectomesenchyme. The dental epithelium gives rise to the outer and inner enamel epithelium from which ameloblasts differentiate, whereas ectomesenchymal cells differentiate into odontoblasts. Nfic has a specific function as a key regulator of dentin formation. Nfic signaling modulates late odontoblast differentiation and mineralization. Both dental papilla cells and dental follicle cells have extensive proliferation ability, expressed similar cell surface antigens and were capable of forming hard tissue. But dental papilla cells had diverse calcium expression than that in dental follicle cells. Most of the literatures have focused the influence of Nfic and OPN in tooth germs. There is no clear evidence to elicit their influence in odontogenic tumours. Aims and objective: The present study was intended to evaluate expression and elicit the influence of Nfic and calcium binding protein (OPN) in odontogenesis and odontogenic tumors. Materials and methods: The study comprised of 20 tooth germs from human foetuses of terminated pregnancies with age ranging between 10 to 36 weeks and 30 paraffin embedded blocks of histopathologically diagnosed odontogenic tumours. Five micrometer thick sections were taken and stained immunohistochemically with Nfic and OPN and evaluated for their expression in tooth germs and in odontogenic tumors. Results: In tooth germs, Nfic showed strong expression in odontoblasts and OPN showed strong expression in enamel matrix. Nfic showed positive expression in odontogenic tumours and OPN showed strong expression in calcification foci in AOT. Conclusion: The present study was an attempt to assess the influence of Nfic during crown formation. Expression of Nfic in tooth germs confirmed its influence in crown formation. Expression of OPN suggested that it promotes mineralization in tooth germs, bone formation and calcification focus in odontogenic tumours

Multidisciplinary interventions in ectodermal dysplasia: A clinical case report

ABSTRACT Ectodermal dysplasia (ED) is a spectrum of inherited disorders that compromise the development and function of ectodermal structures, like hair, nails, and teeth. This case report describes a 17-year-old male with sparse hair and cognitive difficulties who was diagnosed with ED in childhood. A multidisciplinary evaluation with dermatology, neurology, and dentistry revealed characteristic clinical features, and the histopathological diagnosis was confirmed via punch biopsy. Also, ED poses challenges beyond dermatologic manifestations, affecting cognitive function, psychological well-being, and nutrition. Multidisciplinary management, early diagnosis, and awareness are crucial for optimizing patient outcomes and exploring potential therapies.

Comparing tooth development timing between ethnic groups, excluding nutritional and environmental influences.

The timing of dental development in ethnic Finns and Somalis, who were born and living in Finland, was compared, with efforts to minimize environmental bias. The developmental status of seven lower left permanent teeth were staged according to Demirjian et al., using panoramic radiographs from 2,100 Finnish and 808 Somali females and males, aged 2 to 23 years. For each tooth, a continuation-ratio model was constructed to analyze the allocated stages as a function of sex and ethnic origin. Several statistically significant differences in mean age of certain tooth developmental stage transitions were revealed. While Somalis generally displayed stage transitions at younger age, none of the seven teeth consistently showed earlier stage transitions in Somalis compared to Finns. Within each tooth, the lowest (or highest) mean age of stage transition varied without any discernible pattern between the two ethnic groups. Overall, the observed differences in mean age of stage transition between the groups was minimal, suggesting a low impact on clinical and forensic age assessment practice. In conclusion, the studied ethnic Finn and Somali groups with equal nutritional and /or environmental conditions exhibit similar timing in the development of all lower left permanent teeth.

Multi-modal characterization of rodent tooth development.

Craniofacial tissues undergo hard tissue development through mineralization and changes in physicochemical properties. This study investigates the mechanical and chemical properties of developing enamel, dentin, and bone in the mouse mandible. We employ a multi-modal, multi-scale analysis of the developing incisor and first molar at postnatal day 12 by integrating micro-computed tomography (microCT), nanoindentation (NI), energy dispersive spectroscopy (EDS), and Raman spectroscopy. Our findings demonstrate distinct patterns of mechanical, elemental, and chemical changes across mineralized tissues. These results suggest that mineral composition drives mechanical properties across different craniofacial hard tissues. Integrating multi-modal characterization of mineralized tissues opens new opportunities for investigating structure-function relationships in craniofacial biology and genetics.

Nonsurgical endodontic treatment of Type II dens invaginatus by removal of the internal structure of the tooth : a case report

Dens invaginatus is a developmental anomaly that occurs when the enamel organ is invaginated into the dental papilla before the process of calcification during tooth development. Dens invaginatus exhibits various anatomical variations, and these structural changes can complicate treatment. For effective management of dens invaginatus, it is crucial to accurately understand the internal structure of the tooth and to establish an appropriate treatment plan. Cone-beam computed tomography (CBCT) has proven to be a valuable tool for three-dimensional reconstruction of the tooth. This case report presents a maxillary lateral incisor with pulpal necrosis and periapical periodontitis. The case involves dens invaginatus characterized by a separate tooth structure, diagnosed using CBCT, and a treatment plan was established accordingly. For treatment, a dental microscope was used to remove the internal structure of the tooth, followed by filling with mineral trioxide aggregate (MTA).

DPP an extracellular matrix molecule induces Wnt5a mediated signaling to promote the differentiation of adult stem cells into odontogenic lineage

Dentin phosphophoryn (DPP) an extracellular matrix protein activates Wnt signaling in DPSCs (dental pulp stem cells). Wnt/β catenin signaling is essential for tooth development but the role of DPP-mediated Wnt5a signaling in odontogenesis is not well understood. Wnt5a is typically considered as a non-canonical Wnt ligand that elicits intracellular signals through association with a specific cohort of receptors and co-receptors in a cell and context–dependent manner. In this study, DPP facilitated the interaction of Wnt5a with Frizzled 5 and LRP6 to induce nuclear translocation of β-catenin. β-catenin has several nuclear binding partners that promote the activation of Wnt target genes responsible for odontogenic differentiation. Interestingly, steady increase in the expression of Vangl2 receptor suggest planar cell polarity signaling during odontogenic differentiation. In vitro observations were further strengthened by the low expression levels of Wnt5a and β-catenin in the teeth of DSPP KO mice which exhibit impaired odontoblast differentiation and defective dentin mineralization. Together, this study suggests that the DPP-mediated Wnt5a signaling could be exploited as a therapeutic approach for the differentiation of dental pulp stem cells into functional odontoblasts and dentin regeneration.

Population Structure and Selection Signatures in Chinese Indigenous Zhaotong Pigs Revealed by Whole-Genome Resequencing.

Zhaotong pig (ZTP) is a Chinese indigenous pig breed in Yunnan Province, known for its unique body shape and appearance, good meat quality, strong foraging ability, and adaptability. However, there is still a lack of research on its genome. In order to investigate the genetic diversity, population structure, and selection signatures of the breed, we conducted a comprehensive analysis by resequencing on 30 ZTPs and comparing them with genomic data from 10 Asian wild boars (AWBs). A total of 45,514,452 autosomal SNPs were detected in the 40 pigs, and 23,649,650 SNPs were retained for further analysis after filtering. The HE, HO, PN, MAF, π, and Fis values were calculated to evaluate the genetic diversity, and the results showed that ZTPs had higher genetic diversity and lower inbreeding coefficient compared with AWBs. Population structure was analyzed using NJ tree, PCA, ADMIXTURE, and LD methods. It was found that ZTPs were population independent of AWBs and had a lower LD decay compared to AWBs. Moreover, the results of the IBS genetic distance and G matrix showed that most of the individuals had large genetic distances and distant genetic relationships in ZTPs. Selection signatures were detected between ZTPs and AWBs by using two methods, FST and π ratio. Totals of 1104 selected regions and 275 candidate genes were identified. Finally, functional enrichment analysis identified some annotated genes that might affect fat deposition (NPY1R, NPY5R, and NMU), reproduction (COL3A1, COL5A2, GLRB, TAC3, and MAP3K12), growth (STAT6 and SQOR), tooth development (AMBN, ENAM, and ODAM), and immune response (MBL2, IL1A, and DNAJA3). Our results will provide a valuable basis for the future effective protection, breeding, and utilization of ZTPs.