{"paper_id":"0d4b77bb-6209-461c-a7e7-07aa8840b1f7","body_text":"New findings on Developmental Studies of the oropharyngeal Salivary Glands in Japanese Quails (Coutrinx coutrinx japonica) | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article New findings on Developmental Studies of the oropharyngeal Salivary Glands in Japanese Quails (Coutrinx coutrinx japonica) Mahmoud Osman Khalifa, Mahmoud Abd-Elkareem, Wafaa Gaber, Abdelmohaimen Mostafa Saleh This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3958585/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 12 Nov, 2024 Read the published version in BMC Veterinary Research → Version 1 posted 14 You are reading this latest preprint version Abstract The oropharyngeal floor of the lower beak in Japanese quails ( Coutrinx coutrinx japonica ) contains the sublingual and mandibular salivary glands . Although few literatures spoke about the oropharyngeal glands, our study demonstrated the best for full morphological and cytochemical illustration. The morphological and cytochemical analysis were done on 20 healthy Japanese quail embryos with ages of the 6 th , 10 th , 11 th , and 13 th days of incubation, and 25 healthy quail chicks at ages zero (hatching day old), 7 th , 14 th , 30 th , and 60 th days old. The primordia of the sublingual and mandibular salivary glands were noticed at the 6 th and 10 th days of the prehatching old respectively as an epithelial bud. After hatching, both primordia were elongated and differentiated into secretory units. These glands are mucous polystomatic tubulo-alveolar paired glands which were situated in the submucosa. The sublingual glands consisted of 3-5 lobes extended from two ceratobranchial caudally by their wide ends beyond the median sulcus of the prefrenular part of sublingual space rostrally; where they opened by their constricted part. The taste buds are variable in size and position; the associated salivary glands type was the largest, which all taste pores varied from 8.2-12 um. The mandibular glands lay on the paralingual groove which arose at 10 days old embryo. Furthermore, the mandibular glands were located dorsomedial to the sublingual glands and extended longitudinally from the rostral border of the frenulum linguae to the caudal tips of sublingual glands. Notably, the taste buds decreased in the volume and number with advancing age. Both gland secretions showed various histochemical reactions that ended with highly alcinophilic (acidic) materials in advanced ages. Sublingual salivary glands mandibular gland mucous tubuloalveolar Taste bud quail Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Figure 9 Figure 10 Figure 11 Figure 12 Figure 13 Figure 14 Highlights The oropharyngeal floor contains both the sublingual and mandibular salivary glands. The sublingual and mandibular salivary glands were compound tubuloalveolar paired glands. The sublingual salivary glands consist of 3-5 elongated lobes non-branched with common duct. The primordia of the sublingual salivary gland arose at 6 days old embryo as an epithelial bud. The primordia of the mandibular salivary gland arose at 10 days old embryo Both glands’ secretion ended with strong alcinophilic (acidic) contents. The surface taste buds decreased in volume and number with age advancing. Taste buds differed in size and position with taste pores from 8.2-12 um. Background Salivary glands are the accessory structure of the lingual apparatus in birds, exhibiting numerous cytochemical features related to the dietary habits of the species. The function, general morphological, and the histological structures have been discussed in several studies [ 1 , 2 , 3 , 4 ]. However, the glands had a minor role compared to mammals which secreted saliva. The saliva had both an antimicrobial role and a moisture function to food [ 5 ]. Many authors had poor imagination on either salivary glands cytochemistry nor epithelial taste buds in comparison to our imagination here. Surprisingly, woodpecker secretes sticky saliva that helps in sticking ants and insects [ 6 ]. Interestingly, many birds had glands that secrete mucous saliva [ 7 , 8 , 4 ], although few birds had either serous or seromucous glands [ 9 ]. The floor of the oropharyngeal cavity composed of mucosa internally and thin skin externally, in between lay a loose connective tissue layer. Whereas they were containing the paired rostral sublingual glands and caudal paired mandibular salivary glands in chickens [ 10 ], and Muscovy ducks 11 . In our imagination different reports distinguished between various types of food and diet which the taste buds along the oropharynx distributed [ 12 , 13 ]. Additionally, it was approved that the Sanderlings bird was able to discriminate among different sugar concentrations, and types [ 14 ]. Notably, many investigations have been done on the histology and histochemistry of the avian salivary glands [ 15 , 16 ]. Although, little was known about salivary glands of the oropharyngeal floor, their histogenesis, and histochemistry during the pre- and post-hatching period in male and female Japanese quail. The present study aims to investigate the developmental changes of the sublingual and mandibular salivary glands of Japanese quail, morphologically and histochemically by using different techniques; grossly, light microscopy, scanning electron microscopy, and histochemical methods. Materials & Methods Sampling: The quails were obtained from a poultry farm of the Faculty of Agriculture, Assiut University, Assiut, Egypt. In this study, we used forty-five healthy Japanese quails (Coturnix coturnix japonica) divided into twenty prehatching embryos starting from the 6th day pre-hatching till hatching day (zero days old = 17 days old), then twenty-five chick quails at the 7, 14, 30, and 60th day post-hatching old. The materials were collected from the Research Farm of Faculty of Agriculture Assiut University. Whereas fertilized eggs were put on a forced-draft incubator (37.5° ±0.3°C / 60% RH). To be noticed, the eggs were collected within 1 week of laying and preserved in a refrigerator at 4°C for ensuring the symmetrical aging of the specimens before placing into the incubator. A- All methods were performed in accordance with the relevant guidelines and regulations with arriveguidelines instructions (https://arriveguidelines.org). The institutional review board of the Ethics Committee of the faculty of Veterinary Medicine, Assiut University, Egypt; approved this study License No. (06/2023/0051). The oropharyngeal floor was incised and exposed, accordingly to 4 . B- Gross double staining visualization: Three specimens from each age from the seventh and forteenth day-old quail chicks were double stained with either each alcian blue and alizarin red alone or both following [4] Then, the photos were photographed using the stereomicroscope (LEICAS6D). The measurement procedures were done by ImageJ software (https://fiji.sc/). C- Histological and Histochemical Examinations: For paraffin sections and staining, three specimens of each age were used for the fixation, histological process, and decalcification which were done following 4 ; After proper fixation, the samples were kept in 10% formic acid/formol saline for the process of decalcification to ensure of adequate decalcification of the bony and cartilaginous contents of the specimens. After proper decalcification, the specimens dehydrated in ascending degrees of ethanol (70–100%), cleared in methyl benzoate and embedded in paraffin wax stages I, II, and III. The specimens were embedded in paraplast blocks (Sigma Aldrich). Serial 5–6 µm cross, longitudinal, and frontal sections from the oropharyngeal floor were cut by a LEICA 2155rm automatic microtome. Thereafter, the sections were stained for: 1- General morphological studies - Harris hematoxylin and eosin stain [17]. - Crossmon's triple technique [18]. 2- Histochemical studies - Periodic acid-Schiff (PAS) technique for detection of neutral mucopolysaccharides [19]. - Alcian blue technique for demonstration of acidic mucopolysaccharides [20]. - Combined Alcian blue-PAS technique [19]. Almost staining techniques were performed according to Khalifa et al. The histological slides were examined by the OLYMPUS BX51 microscope, and the photos were taken by the OLYMPUS DP72 camera adapted into the microscope. The image staining analysis was done by using ImageJ software (https://fiji.sc/). 3- Drawing a diagram: Drawing an illustrated diagram to show the secretory units of the compound tubuloalveolar sublingual salivary glands. We used Paint program in Windows 7, with Microsoft power point 2010 to draw this diagram. D- Scanning electron microscope (SEM): For scanning electron microscope (SEM) investigation of 10 and 13-day prehatching & 60-day post-hatching old quail: Three samples from each age were used. The components of the floor of the oropharynx were washed several times in 0.1 M phosphate buffer at pH (7.2 ± 0.1). Post-hatching samples were rinsed with acetic acid 2%, then fixed in 4% glutaraldehyde solution for 24 hours. Post-fixation was made in 1% sodium tetroxide solution for two hours at 4°C. After that, the fixed samples were washed in 0.1 M phosphate buffer at pH = (7.2 ± 0.1), then dehydrated in ascending grades of ethanol followed by critical point-dried in liquid carbon dioxide. All specimens were mounted on aluminum stubs covered with carbon tabs and sputtered with gold. The prepared specimens were examined and photographed using JEOL scanning electron microscopy (JSM-5400) at an accelerating voltage of 15kv in the electron microscope unit of Assiut University [21]. The nomenclature used in the present study was coped with the (Nomina Anatomica avium) as well as that was synonymized and homologized with names in previous and recent studies of the chicken and other avian species by different authors. The measurement procedures were done by ImageJ software (https://fiji.sc/). E- Statical analysis: The measurement procedures were done by ImageJ software (https://fiji.sc/). The statical data were represented mean ± SE. The statistics were done by SPSS software and Excell sheet for drawing the figure. Results In this study, we focused on both the sublingual and mandibular salivary glands, where located within the oropharyngeal floor submucosa. A. The sublingual salivary glands: The first sign of the sublingual salivary gland primordia could be seen in the 6-day-old quail embryo as an epithelial thickening of compacted cellular mass (epithelial placode and prebud) within the soft sublingual floor epithelium (Fig. 1A & 1B). Consequently, we noticed that the highly proliferated epithelial masses extended from the bud form to the elongated cords at the 10-day-old quail embryo. Interestingly, these cords have two ends; a rostromedial wide end and a caudolateral narrow end. Canalization was observed in some cords (Fig. 1C &1C*). By advancing age, the lobar cords of the sublingual salivary glands increased in their length and run more caudally without branching at the 11-day-old quail embryo. The rostral end of the cord was broad, less proliferative, and encased by a concentric layer of mesenchyme, whereas the narrow caudolateral proliferative end was still uncovered, suggesting further growth and expansion. Surprisingly, the cord was constricted nearly mid-distance. Due to cell growth competition and apoptosis, many halo vacuoles were obviously present within the rostral part of the cords indicating the beginning of canalization (Fig. 1D). We observed that canalization started rostrally (proximally) and progressed distally (caudally), and the glands extended without branching. By the 13th day of incubation, we found the many vacuoles within the glandular endopieces mass coalesced together to form a single common canal rostrally. Consequently, the lining of the glandular epithelium showed a transformation process from stratified epithelium to monolayer (simple glandular epithelium). Thus, the lumen filled with a secretory-like substance representing the sloughed lining epithelium (Fig. 2A & 2B). Moreover, our study found that the exfoliated substance gave PAS/Alcian blue negative results, at this age. Then, at the maturation stage, the secretory terminals were presented with more lining corrugation and cellular compaction. The lining epithelium showed low columnar cells with foamy cytoplasm and basal vesicular nuclei. Conceptively, near the opening, the lining epithelium transformed from high columnar to low columnar. Thereafter, it ends by stratified squamous epithelium whereas the opening of the taste bud-associating salivary glands found in the hatching chick (Fig. 2C & 2D). Collectively, the first few days of embryo incubation showed highly developmental glandular events. Our gross anatomical examination of 14-day-old chicks revealed that the sublingual glands were paired, consisting of 5–7 parallel elongated lobes with broad blind ends which, directed rostromedially, and narrow ends directed caudolaterally (Figs. 4). The glandular lobes extended longitudinally from the mandibular symphysis rostrally within the sublingual floor to beyond the level of the proximal part of the ceratobranchial. As we found the lobes were constricted at the level of the basihyal (Figs. 3), where they supported the laryngeal mound therefore, it was suggested the significant size of the larynx. Hence, the lobes decreased in the lobation number at that portion (Figs. 4). Concequently, the lobes were numerous rostrally and few caudally whereas are variable from 3–4 lobes caudally (Figs. 3) and (Figs. 4). We conclude that there was an asymmetrical bilateral glandular lobes number along the same side. Our post-hatching histological studies at 14-and 30-days old quail chicks, suggested that the sublingual salivary glands were non-branched compound tubuloalveolar mucous glands that were supported ventrolaterally by muscle mylohyoideus, and a thin connective tissue stroma. Interestingly, the median sulcus was separated between two glandular masses rostrally. The secretory endpieces of each lobule drained their secretion directly into a common secretory duct that opened into the soft sublingual floor separately or collectively with other canals (polystomatic). The ductal epithelial lining was simple tall columnar cells. The ductal canal was narrow while, it passed via the epithelium to end by the excretory duct (Fig. 5A-D). Notably, our histological data confirmed here that the lobation at the level of the sublingual floor were 5–7 bilaterally (Fig. 5A). But, caudally the lobation was five on the right side as follow; two larger, two medium-sized, and one smaller. However, the left side had three lobes; two larger and one smaller. As we noticed that the lobes were numerous rostrally and few caudally. So that the common canals of the sublingual glands were narrower rostrally than the caudal lobes (Fig. 6A-C & Fig. 7A-B). Also, we found an autonomic ganglion that embedded among secretory lobules (Fig. 6D & 6E). Interestingly, the mucous of the sublingual salivary glands showing the apocrine mode of secretion with cellular and nuclear contents detaching within the lumen(Fig. 7C-D). In this study, we found two types of taste buds that have a barrel shape as following; the taste buds with taste pores which was called taste buds-associated salivary glands, and surface epithelial taste buds (Fig. 8A-B). Surface epithelial taste-buds were characterized by deep dermal papillae within the epithelium and deeply stained central cells assumed as gustatory cells (Fig. 8C-D). While the taste buds-associated salivary glands opening, were surrounded by highly mitotic polygonal basal cells within the barrel shape structure (Fig. 8E-F) Our SEM (Figs. 9) and morphometrical measurements (Table 1) revealed that at 8th days old quail embryo the soft sublingual floor was full of numerous domes, mushrooms or spherical like taste buds with various sizes; defined into three different sized. We categorized them into Larger, moderate, and smaller sized taste buds which were ranging from (40, 12, 6) um 2 respectively. Also, the data showed some large sized taste buds nearly fused together (Fig. 9A). By the age of the 10th day old embryo, we noticed that the taste buds had surface area of 36 um 2 and sublingual salivary glands opening diameter was 5 um. Notably, the taste buds decreased in the number and size. However, some of them fused together to form a large one of 144 um 2 (Fig. 9B). With age progression (13 days pre-hatching), the sublingual glands opening increased in the diameter and reached to 28 um with no secretion could be seen. However, the taste bud-associated salivary glands could be seen, with surface area equal to 35 um 2 (Fig. 9C). At 30- and 60-days old chick, various sized taste buds could be measured as follow; the larger (32 um 2 ), moderate (12 um 2 ), and smaller (9 um 2 ) sized, and their taste pores were varied from 8.2–12 um. These taste buds were emerging through the scales of the surface of the sublingual floor epithelium and mucous secretion exited via the salivary gland opening (Fig. 9D-F). Consequently, our data indicated that the taste buds were different in size and position. Of note, our histochemical analysis of the sublingual glands (Table 2A) showed a strong positive reaction with PAS and combined AB/PAS stains, while weakly reaction to AB stain in the newly hatched quail chick (Fig. 10A-C). Moreover, in the seventh day old chick, the sublingual glands were shown moderate positive reaction to alcian blue and strong to PAS and combined AB/PAS stains (Figs. 10D-F). However, at 14 days old quail chick, these glands showed moderate positive reaction to AB, very strong positive reaction to PAS and to combined AB/PAS stains (Fig. 11A-C). In 30 days old quail, the sublingual glands showed strong positive reaction to AB stain and very strong positive to combined AB/PAS stains (purple coloration), but negative results to PAS stain (Fig. 11D-E). However, in the 60th days old quails, the glands showed negative results to PAS stain and very strong positive reaction to AB stain and very strong alcianophilic reaction to AB/PAS stains (greenish blue coloration) (Fig. 11F). B- The mandibular salivary glands : Developmentally, we found that the mandibular glands appeared at 10 days old embryo arising from the epithelium of the paralingual groove. We observed that the developmental characterization of the gland was like the sublingual glands in almost stages. However, the mandibular glands were different in branching of the glandular endpieces (Fig. 12A-F). By gross studies, we found the glands extended caudal to the frenulum linguae, lying within the floor of the paralingual grooves (Fig. 3B). They are paired compound tubuloalveolar glands lay in the tunica submucosa of the paralingual grooves of the oropharynx. The mandibular glands extend longitudinally from the rostral border of the frenulum linguae to the caudal tips of sublingual glands. They lay on the dorsomedial border of the sublingual glands. They open into the ventral or medial aspect of the paralingual groove (Fig. 13A-B). Once again, the mode of the secretion resembles the sublingual gland. Moreover, the glandular acini separated with thick connective tissue septa (interlobular C.T). (Fig. 7E-F). The histochemical study (Table 2B) suggested that the mandibular salivary glands in 30 days old quails showed very strong positive reaction to all of the alcian blue, PAS and combined AB/PAS stains (Fig. 13C-F). By the 60-day old quail, these glands showed a strong positive reaction to AB stain and combined AB/PAS stains (greenish blue coloration only), but negative results to PAS stain. Figures (14) is a schematic diagram showing the secretory units of the compound tubuloalveolar sublingual salivary glands. Discussion Salivary glands are unique specialized structures, which play a vital role in mammals and birds [ 22 ]. In this study, we focused on the paired oropharyngeal floor-related salivary glands of the Japanese quail. Both glands called the sublingual and mandibular salivary glands as anatomical nomenclature followed [ 23 ] in chicken. However, [ 23 , 24 , 25 ] suggested that Glandula sublingualis synonymies to Glandula submandibularis rostralis or mandibularis rostralis, while the Glandula mandibularis Synonymies to (Gl. Mandibularis caudalis) or (Gl. Submandibularis caudalis). Also,[ 3 ] divided them into lateral and medial mandibular glands in emu. Of note, [ 23 , 24 ] discussed both glands deeply in chicken. But few histological and histochemical studies have been done on sublingual and mandibular salivary glands in quails. From our prehatching data, we found that the sublingual and mandibular salivary glands firstly appeared as a thick invagination (a bud stage) of the sublingual floor and paralingual epithelium, at the 6th & 10th day pre-hatching old respectively in agreement to [ 4 , 26 ]. Although [ 25 ] assumed that the primordia arose at 8-day quail from the oral floor epithelium that was extended without branching in agreement with us. Additionally, as we revealed that the canalization process of the sublingual gland started and became obviously detected at the 10th and 11th days of the incubation respectively. Hence, the transformation process of the glandular lining from stratified epithelium to monolayer (simple glandular epithelium). Moreover, the branching of the sublingual gland primordia occurred at 13th-day pre-hatching. To find that [ 25 ] noticed also the lingual salivary gland had a branching mechanism that was different from the sublingual salivary glands. However, it liked as the mandibular salivary glands as we approved. Interestingly, the murine antenatal morphogenesis of the salivary glands branching was a clear example in prebud, initial, canalicular, and terminal bud stages [ 27 , 28 ]. All the prenatal developmental stages of the sublingual salivary glands passed with cellular proliferation, quiescence, apoptotic, and cytodifferentiation events, in which were directed by special growth factors, cytokines, and signaling pathways that target transcription factors, to be upregulated spatially and temporally [ 29 , 30 , 31 , 28 , 32 , 33 ]. Obviously, it has been shown that sublingual glands morphogenesis was dependent on the cross-talking between the epithelial-mesenchymal sides [ 34 ]. Our prehatching results revealed the presence of the secretory-like product and cell fragment within the lumina at the 13-day-old embryo, which gave negative results with AB or PAS stains in agreement with [ 35 ] who assumed that the secretory products appear on the 15th day old chick embryo. Also,[ 36 ] detected that acidic secretion was released from the developing proventriculus at the 13th days old chick embryo because of the swallowed albumin stimulation. A similar finding was found in quail 4 . Hence, our results within the sublingual salivary glands indicated that the acini opened directly into a large common wide secretory lumen of the lobule which thus serves as a common excretory duct to open into the oral cavity either may be fused with another lobular common duct or opened separately. The secretory system was lined with the same cells type of the secretory end pieces; simple columnar cells that were transformed to low columnar to finally become stratified epithelium in agreement to [ 35 , 36 ]. Moreover, the secretory end pieces of the other salivary glands have drained their secretion within intralobuar ducts and then into interlobar duct to the common secretory canal. [ 37 ] found that the columnar cells lining of the glandular duct surrounding the lumen and with microvilli with the apical cellular portion in posterior lingual salivary gland in quail similar to our findings. Moreover, 3 supported the presence of a central canal, but the glands are simple branched tubular in emu and the lining was simple ciliated columnar cells or pseudostratified columnar epithelium. While in the present study it was compound tubuloalveolar gland in agreement with 38 in rock dove. In support to the previous results, the sublingual salivary glands of mice consisted of large and small ducts that ended in a single lumen-containing secretion [ 31 , 27 , 28 ]. Surprising, it was clear from gross and histological data that the sublingual glands lobation was different from [ 23 ] in chicken who assumed that the glands consisted of 5–7 lobes. However, we found that the lobation number was different bilaterally and unilaterally, and depended on the location. Importantly, no author defined the details or pointed out to that even if in quails or chicken. The current study showed that the mandibular gland was paired compound branched tubulo-alveolar glands within the submucosa of the paralingual floor of the mouth. Whereas, it extended from the caudal of the frenulum linguae to the tips of the gland sublingualis. Their orifices were opened into the bottom of the paralingual groove. These results were like that of [ 37 , 10 , 43 , 23 , 3 ]. However, [ 39 , 40 ] assumed the mandibular salivary glands to be a part of the gland sublingualis referred to that, as being the posterior part of a single gland, of which the gland sublingualis is the anterior part. Notably, we observed that the glands were supported with thick connective tissue stroma compatible with [ 3 ]. The present SEM and histological studies about taste buds in Japanese quails were similar to our previous studies. Whereas taste buds were two types; the larger one is associated with the salivary glands opening and the smaller one called surface taste buds lay free on the oral surface mucosa corresponding to the findings in chicken [ 41 , 43 , 61 ], and in emu, [ 3 ]. However, our past morphometric studies mentioned that the taste pore of taste buds associated with preglottal gland had no change in diameter (8.17 µm) at 14- and 30-days old quail chick. While the preglottal salivary gland openings widths were 25.5 µm, and 80.4 µm at 14- and 30-days old quail chick respectively [ 4 ]. However,[ 61 ] found that taste buds parameters in chicken were (40–70 µm) in width and (70–120 µm) in height. While taste pore was (3–7 µm) in width. Although, the openings of the salivary glands are greater than (10 µm) in diameter and were observed near the taste bud. In addition to that, other authors’ data revealed that taste pores width was equal to (5–10 µm)[ 41 ] or (6 µm) [ 42 ]. Accordingly, in the present study (Table 1) showed the average diameter in (um) of the sublingual salivary gland openings (SGO), the taste buds openings (TBO), and the taste buds surface area (TBA) represented by (um 2 ) in quails. Conclusively, we found out that, taste buds were categorized into three sized groups; larger, moderate, and smaller, and according to the position of taste buds associated with salivary glands opening and surface epithelial one. Also, taste pores varied from (8.2–12 µm). These confusing results of previous literatures were because the measurement of many authors trials neglected the fact of different sized, and the positioned taste buds as we did. Notably, we assumed that salivary glands act as a flusher by mucous secretion for taste buds associated with its opening, like Ebner’s glands in mammals. Finally, we did not think that many literatures spoke about the taste buds in detail in quail as we did, suggesting that they have a vital role in food discrimination and prehension. Our histological data indicated that the sublingual and mandibular salivary glands secreted mucous in agreement with [ 44 , 45 , 46 ] who surveyed salivary glands of some birds to have a mucous secretion type. Referring to our histochemical results, the glands were strongly positive to PAS stain, but they were weakly to AB stain at the hatching day of chicks. Then reacting negatively to PAS, but strongly to AB stain at 30- and 60-days post-hatching old. Moreover, [ 47 , 48 ] found that the posterior lingual salivary glands were robustly positive for neutral mucopolysaccharides than the anterior one in both Common Myna and red jungle fowl. However, there no mucosubstance in lingual salivary glands of the little erget [ 49 ]. In addition, [ 37 , 16 ] supported the presence of sialo glycoconjugates within the preglottal and the lingual salivary glands in Japanese quails. Also, this was stated that in the salivary glands of chicken by [ 50 , 51 ]. In addition, [ 37 , 16 ] supported the presence of sialo glycoconjugates within the preglottal and the lingual salivary glands in Japanese quails. Also, this was stated in the salivary glands of chicken by [ 50 , 51 ]. However, palatine salivary glands of chicken produced glycoproteins, sulphomucins, and carboxymucins, which were rich in sialic acid showed more PAS and alciniophilic containing metachromatic granules [ 52 ]. Of note,[ 53 ] suggested that the lingual glands were more acidic with sulphated or acidic sialomucin. Accordingly, the double differential staining, of alcian blue (AB) and PAS stains differentiated between acidic glycoconjugates stained (blue) with AB and neutral one stained (magenta) with PAS in agreement to [ 50 ] in chicken. From that point, we concluded that the secretory mucous took the fate from neutral to acidic result. A tentative interpretation of the type of the lingual gland secretions in little egret could be made according to the classification of mucosubstances were proposed by other workers [ 54 , 55 , 56 ]. The lingual salivary gland of little egret was exhibited to be alcianophilia and reacted positively to the different techniques employed for proteins detection. Accordingly, these glands are confirmed to be of mucoserous type. The lingual glands of chicken [ 57 ], penguin [ 58 ] (Samar et al. 1995), white-cheeked bulbul [ 59 ], and Eurasian collared dove [ 60 ] (Taib and Jarrar, 2001), were reported to be of mucous type, however the quail [ 16 ] and the little [ 49 ] had a mixture of serous and mucous type salivary glands. The secretory products of the lingual salivary glands of these species of birds as seen in the present study form a blend of mucoserous secretions that contained sialomucins, sulfomucins, and proteins 49 . To illustrate that our result found the glands were well developed in quails; and took acidic reaction with age advancing as [ 58 ] finding. As our findings showing the secretory units of the compound tubuloalveolar sublingual salivary glands (Fig. 14). Conclusion In conclusion, we assumed that the sublingual salivary glands and mandibular salivary glands were the basic feedings adapting units among the mucous salivary glands system. Consequently, such developmental and histochemical structures of the sublingual salivary glands in the quail play a role in various food intakes and are responsible for the softening and formation of food chows. These findings of oropharyngeal glands survey give a good future imagination in the quail and other birds for for further study. Declarations Competing interests The author(s) declare no competing interests. Availability of materials and data The data that support the findings of this study are available on request from the corresponding author. Author contributions M.O.K. designed the research; M.O.K. acquired the data. M.O.K., M.A., W.G., and A.M.S. analyzed the data, and M.O.K. wrote the manuscript. All authors discussed the results and commented on the manuscript. Ethics Declarations Ethical approval Our experiment was done as well as all quail bird experiments were performed following the guidance and administration of the Animal Research Committee at the Faculty of Veterinary Medicine, Assiut University, Assiut, Egypt. License No. (06/2023/0051). Acknowledgments This work is a collaborative research program between Assiut University and Aswan University. The funders play no role in the study design, data collection, analysis, decision to publish, or preparation of the manuscript. References King, A. 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Embryol. 30, 367±373. https://doi.org/10.1046/j.1439-0264.2001.00353.x Harris, H. (1900). On the rapid conversion of haematoxylin into haematein in staining reactions. Journal of Applied Microscopic Laboratory Methods, 3, 777. https://www.scienceopen.com/document?vid=c2c7d328-e08c429d-b85c-fa8c3371e3b8. Crossmon, G. (1937). A modification of Mallory's connective tissue stainwith a discussion of principles involved.The Anatomical Record,69(1),33–38. Mc Manus, J. (1946). Histological demonstration of Mucin after periodic acid. Nature, 158(4006), 158–202. Bancroft, J. D., & Stevens, A. S. (1982).Theory and practice of histological techniques. 2. Edinburgh: Churchill Livingstone Bancroft, J. D., & Gamble, M. (2002). Theory and practice of histological techniques (5th ed., PP. 593–620). Edinburgh: Churchill Livingstone. Bock W. J. Salivary glands in the Gary Jays (Perisoreus). (1961). 1;78(3):355 65.https://doi.org/10.2307/4082273. Homberger, D. G., & Meyers, R. A. (1989). 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G., Agas, D., Materazzi, S., Menghi, G., and Marchetti, L., (2009). Ultrastructure and lectin cytochemistry of secretory cells in lingual glands of the Japanese quail (Coturnix coturnix japonica). Histol. Histopathol., 24: 1087-1096. Al-Nefeiy, F. A. and Alahmary, B. A. (2015): Morphological, histological and histochemical studies of the lingual salivary glands of the rock dove, Patagioenas livia (Columbidae). International Journal of current research and academic review. ISSN: 2347-3215 Volume 3 Number 8 pp. 280-289. Mcleond, W. M.; Trotter, D. M. and Lumb, J. W. (1964): Avian anatomy Burgess, Minneapolis. Hodges, R. D. (1974). The histology of the fowl. London; New York: Academic Press. Gentle M. J. (1971): The lingual taste buds of Gallus domesticus L. British Poultry Science 12, 245–248. Saito, I. (1965). Comparative anatomical studies of the oral organs of the poultry. IV. Macroscopical observation of the salivary glands. Bulletin of the Faculty of Agriculture, 12, 110–120. 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Morphological and histochemical observations of the red jungle fowl tongue Gallus gallus. African Journal of Biotechnology, 10(48), 9969–9977. Al-Mansour, M.I., Jarrar, B.M. 2007. Morphological, histological and histochemical study of the lingual salivary glands of the little egret, Egretta garzetta. Saudi J. Biological Sci., 14: 75 81. Arthitvong s.; Makmee, N. and Suprasert, A. (1999): Histochemical detection of glycoconjugates in the anterior lingual salivary glands of the domestic fowl. Kasetsart, J. (Nat. Sci.), 33: 243-250. Suprasert A., Arthitvong S., Koonjaenak S., (2000) Lectin histochemistry of glycoconjugates in the mandibular gland of chicken. Kasetsart J. (Nat. Sci.) 34.85-90. Samar ME, Ávila RE, Esteban FJ, Olmedo L, Dettin L, Massone A, et al. Histochemical and ultrastructural study of the chicken salivary palatine glands. Acta Histochem. 2002:104(2):199–207. https://www.sciencedirect.com/science/article/pii/S0065128104701166 El-Bakry, A.M., & Iwasaki, S. (2014). Ultrastructure and histochemical study of the lingual salivary glands of some bird species. Pakistan Journal of Zoology, 46 , 553-559. Pearse A. G. E., 1972: Histochemistry: Theoritical and Applied. 3rd ed. J. & A. Churchill, London, pp. 1303-1444. Scott D. E., Dorling J., (1965): Differential staining of acid glycosaminoglycans (mucopolysaccharides) by Alcian blue in salt solutions. Histochemie, 5: 221-33. Spicer S. S., Meyer D. B., Histochemical differentiation of acid mucopolysaccharides by means of combined aldehyde fuchsin-alcian blue staining. (1960). Tech Bull Regist Med Technol Am Soc Clin Pathol Regist Med Technol.30:53-60. Fujii, S., Tamura, T. 1966. Histochemical studies on the mucins of the chicken salivary glands. J. Fac. Fish. Anim. Husb, Hiroshima Univ., 6: 345 355. Samar, M.E., Avila, R.E., De Fabro, S.P., Centurion, C. 1995. Structural and cytochemical study of salivary glands in the magellanic penguin Spheniscus magellanicus and the kelp gull Larus domincanus. Marine Ornithol., 23: 154 156. Al-Mansour, M.I., Jarrar, B.M. (2004). Structure and secretions of the lingual salivary glands of the white- cheecked bulbul, Pycnonotus leucogenys, (Pycnontidae). Saudi. J. Sci., 11(2): 119 126. Taib, N.T., Jarrar, B.M. 2001. Histological characterization of the lingual salivary glands of the Eurasian collared dove, Streptopelia decaocta. Pak. J. Bio. Sci., 4(11): 1425 1428. Kudo, K., Nishimura, S., & Tabata, S. (2008). Distribution of taste buds in layer-type chickens: scanning electron microscopic observations. Animal Science Journal, 79(6), 680–685. https://doi.org/10.1111/j.1740- 0929.2008.00580.x. Tables Table (1): Showing the average diameter (um) of the sublingual salivary gland openings (SGO), the taste buds openings (TBO) and taste buds surface area (TBA) (um 2 ) in quails Item SGO (um) TBA (um 2 ) TBO (um) Age 8 days embryo - Small sized (6±0.01) Medium sized (12±0.05) Large sized (40±0.015) - 10 days embryo 5±0.02 Decreased in number and some fused to be 144±0.03 - 13 days embryo 28±0.15 35±0.012 - Hatching chick 30±0.1 - 11±0.023 7 days chick 36±0.04 - 12±0.1 30- & 60-days chick 45±0.08 Small sized (9±0.06) Medium sized (12±0.013) Large sized (32±0.02) 8.2±0.35 Table (2): Showing the histochemical properties of the sublingual salivary gland (Gs) & mandibular salivary gland (Gm) in quail chicks. (A) Sublingual salivary gland (B) Mandibular salivary gland Zero day 7 days 14 days 30 days 60 days 30 days AB +/- ++ ++ +++ ++++ ++++ PAS (Magenta) +++ +++ ++++ - - ++++ AB/PAS Purple +++ +++ ++++ ++++ ++++ Alcinophilic (greenish-purple) ++++ Alcinophilic (greenish-purple) Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 12 Nov, 2024 Read the published version in BMC Veterinary Research → Version 1 posted Editorial decision: Revision requested 16 May, 2024 Reviews received at journal 04 May, 2024 Reviews received at journal 30 Apr, 2024 Reviewers agreed at journal 26 Apr, 2024 Reviewers agreed at journal 25 Apr, 2024 Reviews received at journal 23 Apr, 2024 Reviews received at journal 05 Apr, 2024 Reviewers agreed at journal 02 Apr, 2024 Reviewers agreed at journal 28 Mar, 2024 Reviewers invited by journal 28 Mar, 2024 Editor invited by journal 19 Feb, 2024 Editor assigned by journal 19 Feb, 2024 Submission checks completed at journal 15 Feb, 2024 First submitted to journal 15 Feb, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {\"props\":{\"pageProps\":{\"initialData\":{\"identity\":\"rs-3958585\",\"acceptedTermsAndConditions\":true,\"allowDirectSubmit\":false,\"archivedVersions\":[],\"articleType\":\"Research Article\",\"associatedPublications\":[],\"authors\":[{\"id\":273034604,\"identity\":\"264d0b84-8eb7-42b0-a2fa-1d6afb7e8521\",\"order_by\":0,\"name\":\"Mahmoud Osman Khalifa\",\"email\":\"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABAUlEQVRIiWNgGAWjYFACHhDBDGPYQEXZiNHCBmakka7lMGEtuu1nj2742GbNwC/fe3TDzx3n5eVn5BgwfCg7zMDPvwCrFrMzeWk3Z7alM0i28aXd7D1z23DDjRwDxhnnDjNIzniAXcuBHLPbvG2HGQyO8Zjd4G27zbhBOseAGSxy4wB2LeffILTc/Nt2zn7+bKCWv0ARe1xabiDZAmQcSGy4DdTCCBLhb8Ch5Y3ZzRnn0nkk24B6ZduSkzfcf1ZwsAcoInEDe4iZnc8xu/GhzFqOn/mM2c23bXa283sOb3zwAyTSj91hMMCDwjsAFpFIwKsFG+DHb8soGAWjYBSMGAAASq5j+2OQJAMAAAAASUVORK5CYII=\",\"orcid\":\"\",\"institution\":\"Faculty of Veterinary Medicine, Aswan university\",\"correspondingAuthor\":true,\"prefix\":\"\",\"firstName\":\"Mahmoud\",\"middleName\":\"Osman\",\"lastName\":\"Khalifa\",\"suffix\":\"\"},{\"id\":273034605,\"identity\":\"b9679112-64e3-4cb0-b5f7-36a811fe08f2\",\"order_by\":1,\"name\":\"Mahmoud Abd-Elkareem\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"Faculty of Veterinary Medicine, Assiut University\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Mahmoud\",\"middleName\":\"\",\"lastName\":\"Abd-Elkareem\",\"suffix\":\"\"},{\"id\":273034606,\"identity\":\"93476515-3f0a-4399-b7e0-d101f7b3ab4a\",\"order_by\":2,\"name\":\"Wafaa Gaber\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"Faculty of Veterinary Medicine, Assiut University\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Wafaa\",\"middleName\":\"\",\"lastName\":\"Gaber\",\"suffix\":\"\"},{\"id\":273034607,\"identity\":\"38de7491-5a91-45b7-9fa7-a19ab4f08a5d\",\"order_by\":3,\"name\":\"Abdelmohaimen Mostafa Saleh\",\"email\":\"\",\"orcid\":\"\",\"institution\":\"Faculty of Veterinary Medicine, Assiut University\",\"correspondingAuthor\":false,\"prefix\":\"\",\"firstName\":\"Abdelmohaimen\",\"middleName\":\"Mostafa\",\"lastName\":\"Saleh\",\"suffix\":\"\"}],\"badges\":[],\"createdAt\":\"2024-02-15 12:04:07\",\"currentVersionCode\":1,\"declarations\":\"\",\"doi\":\"10.21203/rs.3.rs-3958585/v1\",\"doiUrl\":\"https://doi.org/10.21203/rs.3.rs-3958585/v1\",\"draftVersion\":[],\"editorialEvents\":[{\"content\":\"https://doi.org/10.1186/s12917-024-04355-7\",\"type\":\"published\",\"date\":\"2024-11-12T15:57:35+00:00\"}],\"editorialNote\":\"\",\"failedWorkflow\":false,\"files\":[{\"id\":51386301,\"identity\":\"52f0caec-d85a-45fb-b9b6-4d5adf08e537\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:29\",\"extension\":\"jpg\",\"order_by\":1,\"title\":\"Figure 1\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":1768374,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of the sublingual floor: (A \\u0026amp; B): Sagittal sections of a 6-day old quail embryo, showing the primordia of the sublingual salivary glands (Rectangular shape, Fig. 1A). Note: the epithelial thickening of the sublingual floor mucosa (epithelial placode) (black arrowheads, Fig. 1B). (SLF) sublingual floor, (T) tongue, (MR) mandibular ramus, muscle mylohyoideus (Mmh) (C): Sagittal sections of 10-day old quail embryo showing the glands sublingualis (Gs) have a canalized cord like shape with two ends; rostromedial wide part (black star), and caudolateral narrow part (red star). Notice: the muscle mylohyoideus (Mmh). (C*): Cross section showing canalization (blue star) within the gland and stratified lining epithelium of the same age. (D): \\u0026nbsp;11-day old quail embryo showing the extension of the cord of the gland sublingualis (Gs) with its rostromedial wide part (black star) covered by concentric layer of mesenchyme and caudolateral narrow part (red star) uncovered by mesenchyme, and central constricted part (red arrowhead) small spaces have been shown (black arrowhead). H \\u0026amp; E, Scale bars; A: 500 µm, B: 100 µm, C: 200 µm, C\\u003csup\\u003e*\\u003c/sup\\u003e: 50 µm, D: 200 µm.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture1.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/0e9a85fdcd3dca7e1dd4b4a4.jpg\"},{\"id\":51386297,\"identity\":\"71a2daae-ba5d-4418-96e5-c7087388b79e\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:29\",\"extension\":\"jpg\",\"order_by\":2,\"title\":\"Figure 2\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":1658633,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of frontal sections in the sublingual floor of 13-day old quail embryo (A): Showing straight common canal (red star) along the gland sublingualis (Gs). Note: the muscle myelohyoid (Mmh) and Meckelian cartilage (MC). B: Showing the presence of cellular exfoliation (red star) of the gland sublingualis. (C \\u0026amp; D): Cross sections of a hatching quail chick; C: Showing the well-organized gland sublingualis (Gs) opening in the surface epithelium (Ep; stratified squamous epithelium non cornified). D: Showing the simple columnar glandular epithelium (red arrowhead), more infoldings in secretory end pieces and the secretory materials (Blue arrowhead) of the gland sublingualis. Note the taste bud (TB) with taste pore (black arrowhead). H \\u0026amp; E, Scale bars; A: 500 µm, B: 100 µm, C: 100 µm, D: 50 µm.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture2.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/411557e5b0fc0e2976c5ca3c.jpg\"},{\"id\":51386294,\"identity\":\"968d7394-ac69-4304-8198-e38d5e7c0c3c\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:28\",\"extension\":\"jpg\",\"order_by\":3,\"title\":\"Figure 3\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":444305,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotographs of a dorsal view of the oropharyngeal floor of a 14-day old quail chick: (A) Showing sublingual floor (SLF) after tongue (T) reflection, and prefrenular median sulcus where sublingual salivary glands open (black star). Note, frenulum linguae (FL). (X 6.3). (B) Photograph showing pharyngeal floor parts of paralingual grooves (black arrows) and distribution of mandibular salivary glands (blue dots). Note, Ramus mandibularis (RM), lingual apex (L.Apex), lingual body (LB), Lingual root (LR), and laryngeal mound (LM). (X 6.3).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture3.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/5b38f80e4f0fccef60fcb681.jpg\"},{\"id\":51386302,\"identity\":\"372fa72d-b6ed-4544-b94e-72eaa80adb9f\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:30\",\"extension\":\"jpg\",\"order_by\":4,\"title\":\"Figure 4\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":699948,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotographs of the gross staining (after maceration) of 14-day old quail chick (A) dorsal view of the oral floor after tongue (T) reflection showing sublingual floor (SLF), and prefrenular median sulcus where sublingual salivary gland open (red star). Note, frenulum linguae (FL). (Alzarin red \\u0026amp; alcian blue stains, X 6.3). (B) ventral view of the lower beak of a 14-day old quail chick showing gland sublingualis (Gs) extends from symphysis mandibularis (S) beyond the first parts of the ceratobranchialia (CB), ramus mandibularis (RM). Note the basihyoid (BH), Paraglossal (PG), trachea (TR), urohyale (UH), and central constricted part (red asterisks). Sublingual groove (black asterisk). (Alizarin red \\u0026amp; alcian blue X 6.3).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture4.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/ebae996cb7f3c0a89543994c.jpg\"},{\"id\":51386290,\"identity\":\"7bad9158-0c1c-4a3a-b3e0-fd99a3190f26\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:27\",\"extension\":\"jpg\",\"order_by\":5,\"title\":\"Figure 5\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":2034646,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of oral floor of a 30-day old quail: (A \\u0026amp; B): Cross sections showing the paired gland sublingualis (Gs) which composed of variable 5-7 lobules in each side at the oral part; lying in submucosa and supported ventro-laterally by muscle mylohyoideus (Mmh). Note: median sulcus (black star), ramus mandibularis (RM), sublingual floor (SLF) and tongue (T). (Crossmon’s trichrome stain, Scale bars; A: 1000 µm, B: 500 µm). (C \\u0026amp; D): Frontal sections showing the lobular structure of gland sublingualis composed of compound tubuloalveolar secretory units open into common secretory duct (CSD) which lined by simple columnar epithelium (black arrowhead). (H \\u0026amp; E stain, scale bars; C: 200 µm, D: 100 µm).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture5.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/6dca47226cf9552930fc89bd.jpg\"},{\"id\":51386292,\"identity\":\"49147b9b-c33e-402e-aef6-633d87db2eab\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:28\",\"extension\":\"png\",\"order_by\":6,\"title\":\"Figure 6\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":506504,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of pharyngeal floor of a 30-day old quail. The left side showing gland sublingualis (Gs) consist of 3 lobes (A \\u0026amp; B) while the right side showing 5 different sized lobes; 2 larger ones, 2 moderate ones and a smaller one in (A \\u0026amp; C). Note: muscle mylohyoideus (Mmh), paralingual groove (Plg), gland mandibularis (Gm) and ramus mandibularis (RM), laryngeal mound (LM). (Crossmon’s trichrome stain, scale bars; A: 1000 µm, B \\u0026amp; C: 500 µm). (D \\u0026amp; E): Autonomic ganglion (AG) adjust to the glands sublingualis. (H \\u0026amp; E stain, scale bars; D: 100 µm; E: 50 µm).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"6.png\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/27abe95a9b2af069dd161555.png\"},{\"id\":51386300,\"identity\":\"375d36a2-4193-4f6c-98a5-80669f55f262\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:29\",\"extension\":\"jpg\",\"order_by\":7,\"title\":\"Figure 7\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":653395,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of different sections in the oropharyngeal floor of a 30-day old quail chick. (A): Frontal section showing many taste buds within the epithelium (blue arrowheads), the gland sublingualis (black arrowheads), the lobar structures of the gland sublingualis (black dash rectangular shape). (B): The lobar structure of the gland sublingualis with a central lumen (Black star) (C): Cross sections of the gland sublingualis (Gs) at the pharyngeal level with wide central lumen (red star) filled with secretion. (D): Apocrine secretion, the cellular content depris (red color) and secretion (greenish color). (E): Apocrine secretion of the gland mandibularis (Gm) poured into paralingual groove (Plg) with cellular contents and secretions. Note the submucosa (SM). (F): Secretory endpieces with thick connective tissue septa (red star) of the gland mandibularis (Gm). (Crossmon’s trichrome stains, scale bars; A: 500 µm, B, D \\u0026amp; F: 100 \\u0026amp;µm C \\u0026amp; E: 200 µm).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"7.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/797d40c748a98eb7662f5a7b.jpg\"},{\"id\":51386303,\"identity\":\"3a3b5d03-e005-4adf-b870-2bc8ffc16851\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:30\",\"extension\":\"jpg\",\"order_by\":8,\"title\":\"Figure 8\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":521398,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of different sagittal sections in the quail oropharyngeal floor. (A-D): Photomicrograph of a sagittal section oropharyngeal floor of 14-days old quail chick showing the types of the barrel shaped taste buds with taste pores. (A): Taste buds associated salivary glands opening (black rectangular shaped) and surface epithelial taste buds (blue rectangular shaped). Note the tongue (T) and sublingual floor (SLF). (B): Showing taste pores (black dotted circle), dark stained cells (black arrowhead) and epithelium (Ep). (C \\u0026amp; D): Surface epithelial taste buds (blue dotted circle), with deep dermal papillae (black headarrows). (Crossmon’s trichrome stains, scale bars; A: 500 µm, B, D: 50 µm \\u0026amp; C: 100 µm). (E \\u0026amp; F): Photomicrograph of a sagittal section oropharyngeal floor of a 60-days old quail showing the opening of the gland sublingualis (red arrow) arose from the epithelium of the sublingual floor, taste bud (red rectangle), taste pores (black arrow), the basal epithelial highly mitotic cells (red star) and the surface epithelium (Ep). (PAS \\u0026amp; Hx stain, scale bars, E: 100 µm \\u0026amp; F: 50 µm).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"8.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/13ca19753ebfd7f55ed69d58.jpg\"},{\"id\":51386298,\"identity\":\"9ae2587d-68d4-44a5-b1af-f864172923fd\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:29\",\"extension\":\"jpg\",\"order_by\":9,\"title\":\"Figure 9\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":567743,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eScanning electron micrographs of sublingual floor surface (A): 8 days prehatching, showing dome-shaped mushroom-like taste buds (black arrowheads) along the sublingual floor (red star) (X 1500). (B): 10 days pre-hatching, showing the opening of the sublingual salivary glands (black arrowheads) with mushroom-like taste buds associated with salivary glands (red arrowheads). (X 750). (C): 13 days pre-hatching showing the opening of the sublingual salivary glands with no secretion could be observed. Note taste bud associated salivary gland (yellow short arrow), surface epithelium with elongated flat cells (yellow star) (X 1500). (D): 30 days old chick showing the different sized taste buds along the sublingual floor epithelium arose among the epithelial scales; larger ones stuck with a little mucous and guarded the salivary gland opening (red arrowhead) and smaller ones, were the surface taste buds (green arrowhead). (E \\u0026amp; F): 60 days post-hatching quail; (E): Showing the sublingual salivary gland opening (red star), mucous debris adjacent to the opening's rim (blue arrowhead), large-sized taste buds associated with salivary gland (white arrowhead) (X 1000). (F): Showing the taste buds associated with the sublingual surface surrounded with pits (red dotted circle and red arrowhead), others associated with mucous (black stars). Note the decrease in the number of the small surface taste buds type.\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"9.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/45d662ee3cd3a3682e553cef.jpg\"},{\"id\":51386599,\"identity\":\"bd608797-9677-489b-a203-6e07c4089b39\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:49:27\",\"extension\":\"jpg\",\"order_by\":10,\"title\":\"Figure 10\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":2709676,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of sagittal sections in the oropharyngeal floor of a newly hatching quail chick. (A): Showing the gland sublingualis strong positive to PAS stain. (PAS stain, scale bar 200 µm). (B): Showing the gland sublingualis strong positive to combined AB/PAS stains. (AB/PAS stains, scale bar 200 µm). (C): Showing the gland sublingualis weakly positive to AB stain, scale bar 100 µm). (D): 7-day old quail chick showing the gland sublingualis strong positive to PAS stain. (PAS stain, scale bar 200 µm). (E): A frontal section of a 7-day old quail chick showing the gland sublingualis moderate positive to AB stain. (AB stain, scale bar 200 µm). (F): \\u0026nbsp;A sagittal section of a 7-day old quail chick showing the gland sublingualis strong positive to combined AB/PAS stains (purple coloration). (AB/PAS stains, scale bar 100 µm).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture10.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/93d04d77444ea7a345c89089.jpg\"},{\"id\":51386293,\"identity\":\"844917f2-3eec-4c5a-99dc-c5292c1d7147\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:28\",\"extension\":\"jpg\",\"order_by\":11,\"title\":\"Figure 11\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":3837930,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of the oropharyngeal floor of a 14 day (A \\u0026amp; B \\u0026amp; C); a 30-day (D \\u0026amp; E) and a 60-day (F) old quail chick. (A): A frontal section showing the gland sublingualis very strong positive to PAS stain. (PAS stain, scale bars 500 µm). (B): Showing the gland sublingualis moderate positive to AB stain (AB stain, scale bars 200 µm). (C): Showing very strong positive to combined AB/PAS stains. (AB/PAS stain, scale bars 500 µm). (D \\u0026amp; E): Photomicrograph of a cross-section in the oropharyngeal floor showing the gland sublingualis strong positive to AB stain \\u0026amp; very strong positive to combined AB/PAS stains (purple coloration) respectively. (D: AB, scale bars 100 µm E: AB/PAS, scale bars 500 µm). (F): Cross section showing the gland sublingualis very strong alcinophilic to AB stain and the same result with combined AB/PAS stains. (AB stain, scale bar 200 µm).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture11.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/d200c03beac5e5aa270057c1.jpg\"},{\"id\":51386295,\"identity\":\"2d5dfe89-c9e8-4f69-a705-4ceea1917cb2\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:28\",\"extension\":\"jpg\",\"order_by\":12,\"title\":\"Figure 12\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":1863713,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of frontal sections in the oro-pharyngeal floor: (A \\u0026amp; B): Showing the gland mandibularis (Gm) buds and cords of 10-days old quail embryo originated from epithelial of the paralingual groove (Plg). Note the Meckelian cartilage (MC). (C): Expansion of the cord of 11-days old quail embryo. (D): Canalization of the expanded cord of 12-days old embryo and thin mesenchymal interaction (red arrowheads). (E \\u0026amp; F): Thick concentric mesenchymal layers surrounded the branched cord of 13-days old embryo with more branched endpieces (black arrowheads). (H \\u0026amp; E stains, scale bars; A: 500 µm, B, C, D \\u0026amp; F: 100 µm, E: 200 µm).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture12.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/f7e485c56b9ca2a97a9ca3d1.jpg\"},{\"id\":51386299,\"identity\":\"4d3aa301-3e9e-4a06-9d0c-fbf69b5308fb\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:29\",\"extension\":\"jpg\",\"order_by\":13,\"title\":\"Figure 13\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":1700344,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003ePhotomicrographs of transverses sections in the oro-pharyngeal floor of a 30-day old quail (A): Showing the gland mandibularis (Gm) lies dorsomedially to gland sublingualis (Gs) and opens ventrally into paralingual groove (Plg). \\u0026nbsp;(Crossmon’s trichrome, scale bar 500 µm). (B): Showing the gland mandibularis (Gm) composes of compound tubule-alveolar secretory endopices lies in the submucosa. \\u0026nbsp;(H \\u0026amp; E, scale bar 200 µm). (C \\u0026amp; D): Showing the gland mandibularis (Gm) and the gland sublingualis (Gs) very strong positive to PAS stain (PAS, scale bar; C: 500 µm, D: 100 µm). (E): Showing the gland mandibularis (Gm) and the gland sublingualis (Gs) very strong positive to combined AB/PAS stains. (AB/PAS stains, scale bar: 500 µm). (F): Showing the gland mandibularis (Gm) very strong positive to AB stain. (AB stain, scale bar: 200 µm).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture13.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/2b36460f9f1684ee9acb9225.jpg\"},{\"id\":51386296,\"identity\":\"fa04639e-e3f2-41ba-8831-c779219cbdd8\",\"added_by\":\"auto\",\"created_at\":\"2024-02-20 17:41:28\",\"extension\":\"jpg\",\"order_by\":14,\"title\":\"Figure 14\",\"display\":\"\",\"copyAsset\":false,\"role\":\"figure\",\"size\":1496495,\"visible\":true,\"origin\":\"\",\"legend\":\"\\u003cp\\u003eSchematic diagram showing the secretory units of the compound tubuloalveolar sublingual salivary glands. Note, the common secretory duct (CSD), the interlobar connective tissue (ILC), and the glandular lobe (Lob).\\u003c/p\\u003e\",\"description\":\"\",\"filename\":\"Picture14.jpg\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/526db5af93bad9719b3da432.jpg\"},{\"id\":69286073,\"identity\":\"f225fee2-ca81-4434-a779-fa4fb0a144de\",\"added_by\":\"auto\",\"created_at\":\"2024-11-18 19:29:19\",\"extension\":\"pdf\",\"order_by\":0,\"title\":\"\",\"display\":\"\",\"copyAsset\":false,\"role\":\"manuscript-pdf\",\"size\":21081104,\"visible\":true,\"origin\":\"\",\"legend\":\"\",\"description\":\"\",\"filename\":\"manuscript.pdf\",\"url\":\"https://assets-eu.researchsquare.com/files/rs-3958585/v1/e6c3ab1f-0d6f-477d-905b-52920c1aaf63.pdf\"}],\"financialInterests\":\"No competing interests reported.\",\"formattedTitle\":\"New findings on Developmental Studies of the oropharyngeal Salivary Glands in Japanese Quails (Coutrinx coutrinx japonica)\",\"fulltext\":[{\"header\":\"Highlights\",\"content\":\"\\u003cul class=\\\"decimal_type\\\"\\u003e\\n \\u003cli\\u003eThe oropharyngeal floor contains both the sublingual and mandibular salivary glands.\\u0026nbsp;\\u003c/li\\u003e\\n \\u003cli\\u003eThe sublingual and mandibular salivary glands were compound tubuloalveolar paired glands.\\u003c/li\\u003e\\n \\u003cli\\u003eThe sublingual salivary glands consist of 3-5 elongated lobes non-branched with common duct.\\u003c/li\\u003e\\n \\u003cli\\u003eThe primordia of the sublingual salivary gland arose at 6 days old embryo as an epithelial bud.\\u003c/li\\u003e\\n \\u003cli\\u003eThe primordia of the mandibular salivary gland arose at 10 days old embryo\\u0026nbsp;\\u003c/li\\u003e\\n \\u003cli\\u003eBoth glands\\u0026rsquo; secretion ended with strong alcinophilic (acidic) contents.\\u0026nbsp;\\u003c/li\\u003e\\n \\u003cli\\u003eThe surface taste buds decreased in volume and number with age advancing.\\u003c/li\\u003e\\n \\u003cli\\u003eTaste buds differed in size and position with taste pores from 8.2-12 um.\\u003c/li\\u003e\\n\\u003c/ul\\u003e\"},{\"header\":\"Background\",\"content\":\"\\u003cp\\u003eSalivary glands are the accessory structure of the lingual apparatus in birds, exhibiting numerous cytochemical features related to the dietary habits of the species. The function, general morphological, and the histological structures have been discussed in several studies [\\u003cspan citationid=\\\"CR1\\\" class=\\\"CitationRef\\\"\\u003e1\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR2\\\" class=\\\"CitationRef\\\"\\u003e2\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e3\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e4\\u003c/span\\u003e]. However, the glands had a minor role compared to mammals which secreted saliva. The saliva had both an antimicrobial role and a moisture function to food [\\u003cspan citationid=\\\"CR5\\\" class=\\\"CitationRef\\\"\\u003e5\\u003c/span\\u003e]. Many authors had poor imagination on either salivary glands cytochemistry nor epithelial taste buds in comparison to our imagination here. Surprisingly, woodpecker secretes sticky saliva that helps in sticking ants and insects [\\u003cspan citationid=\\\"CR6\\\" class=\\\"CitationRef\\\"\\u003e6\\u003c/span\\u003e]. Interestingly, many birds had glands that secrete mucous saliva [\\u003cspan citationid=\\\"CR7\\\" class=\\\"CitationRef\\\"\\u003e7\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR8\\\" class=\\\"CitationRef\\\"\\u003e8\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e4\\u003c/span\\u003e], although few birds had either serous or seromucous glands [\\u003cspan citationid=\\\"CR9\\\" class=\\\"CitationRef\\\"\\u003e9\\u003c/span\\u003e]. The floor of the oropharyngeal cavity composed of mucosa internally and thin skin externally, in between lay a loose connective tissue layer. Whereas they were containing the paired rostral sublingual glands and caudal paired mandibular salivary glands in chickens [\\u003cspan citationid=\\\"CR10\\\" class=\\\"CitationRef\\\"\\u003e10\\u003c/span\\u003e], and Muscovy ducks\\u003csup\\u003e11\\u003c/sup\\u003e. In our imagination different reports distinguished between various types of food and diet which the taste buds along the oropharynx distributed [\\u003cspan citationid=\\\"CR12\\\" class=\\\"CitationRef\\\"\\u003e12\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR13\\\" class=\\\"CitationRef\\\"\\u003e13\\u003c/span\\u003e]. Additionally, it was approved that the Sanderlings bird was able to discriminate among different sugar concentrations, and types [\\u003cspan citationid=\\\"CR14\\\" class=\\\"CitationRef\\\"\\u003e14\\u003c/span\\u003e]. Notably, many investigations have been done on the histology and histochemistry of the avian salivary glands [\\u003cspan citationid=\\\"CR15\\\" class=\\\"CitationRef\\\"\\u003e15\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e16\\u003c/span\\u003e]. Although, little was known about salivary glands of the oropharyngeal floor, their histogenesis, and histochemistry during the pre- and post-hatching period in male and female Japanese quail. The present study aims to investigate the developmental changes of the sublingual and mandibular salivary glands of Japanese quail, morphologically and histochemically by using different techniques; grossly, light microscopy, scanning electron microscopy, and histochemical methods.\\u003c/p\\u003e\"},{\"header\":\"Materials \\u0026 Methods\",\"content\":\"\\u003cdiv id=\\\"Sec3\\\"\\u003e\\n \\u003ch2\\u003eSampling:\\u003c/h2\\u003e\\n \\u003cp\\u003eThe quails were obtained from a poultry farm of the Faculty of Agriculture, Assiut University, Assiut, Egypt. In this study, we used forty-five healthy Japanese quails (Coturnix coturnix japonica) divided into twenty prehatching embryos starting from the 6th day pre-hatching till hatching day (zero days old\\u0026thinsp;=\\u0026thinsp;17 days old), then twenty-five chick quails at the 7, 14, 30, and 60th day post-hatching old. The materials were collected from the Research Farm of Faculty of Agriculture Assiut University. Whereas fertilized eggs were put on a forced-draft incubator (37.5\\u0026deg; \\u0026plusmn;0.3\\u0026deg;C\\u003cstrong\\u003e/\\u003c/strong\\u003e60% RH). To be noticed, the eggs were collected within 1 week of laying and preserved in a refrigerator at 4\\u0026deg;C for ensuring the symmetrical aging of the specimens before placing into the incubator.\\u003c/p\\u003e\\n \\u003cp\\u003e\\u003cstrong\\u003eA- \\u003c/strong\\u003eAll methods were performed in accordance with the relevant guidelines and regulations with arriveguidelines instructions (https://arriveguidelines.org). The institutional review board of the Ethics Committee of the faculty of Veterinary Medicine, Assiut University, Egypt; approved this study License No. (06/2023/0051). The oropharyngeal floor was incised and exposed, accordingly to\\u003csup\\u003e4\\u003c/sup\\u003e.\\u003c/p\\u003e\\n\\u003c/div\\u003e\\n\\u003cdiv id=\\\"Sec4\\\"\\u003e\\n \\u003ch2\\u003eB- Gross double staining visualization:\\u003c/h2\\u003e\\n \\u003cdiv\\u003e\\n \\u003cp\\u003eThree specimens from each age from the seventh and forteenth day-old quail chicks were double stained with either each alcian blue and alizarin red alone or both following [4] Then, the photos were photographed using the stereomicroscope (LEICAS6D). The measurement procedures were done by ImageJ software (https://fiji.sc/).\\u003c/p\\u003e\\n \\u003c/div\\u003e\\n\\u003c/div\\u003e\\n\\u003cdiv id=\\\"Sec5\\\"\\u003e\\n \\u003ch2\\u003eC- Histological and Histochemical Examinations:\\u003c/h2\\u003e\\n \\u003cp\\u003eFor paraffin sections and staining, three specimens of each age were used for the fixation, histological process, and decalcification which were done following\\u003csup\\u003e4\\u003c/sup\\u003e; After proper fixation, the samples were kept in 10% formic acid/formol saline for the process of decalcification to ensure of adequate decalcification of the bony and cartilaginous contents of the specimens. After proper decalcification, the specimens dehydrated in ascending degrees of ethanol (70\\u0026ndash;100%), cleared in methyl benzoate and embedded in paraffin wax stages I, II, and III. The specimens were embedded in paraplast blocks (Sigma Aldrich). Serial 5\\u0026ndash;6 \\u0026micro;m cross, longitudinal, and frontal sections from the oropharyngeal floor were cut by a LEICA 2155rm automatic microtome. Thereafter, the sections were stained for:\\u003c/p\\u003e\\n \\u003cp\\u003e1- General morphological studies\\u003c/p\\u003e\\n \\u003cp\\u003e- Harris hematoxylin and eosin stain [17].\\u003c/p\\u003e\\n \\u003cp\\u003e- Crossmon\\u0026apos;s triple technique [18].\\u003c/p\\u003e\\n \\u003cp\\u003e2- Histochemical studies\\u003c/p\\u003e\\n \\u003cp\\u003e- Periodic acid-Schiff (PAS) technique for detection of neutral mucopolysaccharides [19].\\u003c/p\\u003e\\n \\u003cp\\u003e- Alcian blue technique for demonstration of acidic mucopolysaccharides [20].\\u003c/p\\u003e\\n \\u003cp\\u003e- Combined Alcian blue-PAS technique [19].\\u003c/p\\u003e\\n \\u003cp\\u003eAlmost staining techniques were performed according to Khalifa et al. The histological slides were examined by the OLYMPUS BX51 microscope, and the photos were taken by the OLYMPUS DP72 camera adapted into the microscope. The image staining analysis was done by using ImageJ software (https://fiji.sc/).\\u003c/p\\u003e\\n \\u003cp\\u003e3- Drawing a diagram:\\u003c/p\\u003e\\n \\u003cdiv\\u003e\\n \\u003cp\\u003eDrawing an illustrated diagram to show the secretory units of the compound tubuloalveolar sublingual salivary glands. We used Paint program in Windows 7, with Microsoft power point 2010 to draw this diagram.\\u003c/p\\u003e\\n \\u003c/div\\u003e\\n\\u003c/div\\u003e\\n\\u003cdiv id=\\\"Sec6\\\"\\u003e\\n \\u003ch2\\u003eD- Scanning electron microscope (SEM):\\u003c/h2\\u003e\\n \\u003cp\\u003eFor scanning electron microscope (SEM) investigation of 10 and 13-day prehatching \\u0026amp; 60-day post-hatching old quail:\\u003c/p\\u003e\\n \\u003cp\\u003eThree samples from each age were used. The components of the floor of the oropharynx were washed several times in 0.1 M phosphate buffer at pH (7.2\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.1). Post-hatching samples were rinsed with acetic acid 2%, then fixed in 4% glutaraldehyde solution for 24 hours. Post-fixation was made in 1% sodium tetroxide solution for two hours at 4\\u0026deg;C. After that, the fixed samples were washed in 0.1 M phosphate buffer at pH = (7.2\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;0.1), then dehydrated in ascending grades of ethanol followed by critical point-dried in liquid carbon dioxide. All specimens were mounted on aluminum stubs covered with carbon tabs and sputtered with gold. The prepared specimens were examined and photographed using JEOL scanning electron microscopy (JSM-5400) at an accelerating voltage of 15kv in the electron microscope unit of Assiut University [21]. The nomenclature used in the present study was coped with the (Nomina Anatomica avium) as well as that was synonymized and homologized with names in previous and recent studies of the chicken and other avian species by different authors. The measurement procedures were done by ImageJ software (https://fiji.sc/).\\u003c/p\\u003e\\n\\u003c/div\\u003e\\n\\u003cdiv id=\\\"Sec7\\\"\\u003e\\n \\u003ch2\\u003eE- Statical analysis:\\u003c/h2\\u003e\\n \\u003cp\\u003eThe measurement procedures were done by ImageJ software (https://fiji.sc/). The statical data were represented mean\\u0026thinsp;\\u0026plusmn;\\u0026thinsp;SE. The statistics were done by SPSS software and Excell sheet for drawing the figure.\\u003c/p\\u003e\\n\\u003c/div\\u003e\"},{\"header\":\"Results\",\"content\":\"\\u003cp\\u003eIn this study, we focused on both the sublingual and mandibular salivary glands, where located within the oropharyngeal floor submucosa.\\u003c/p\\u003e \\u003cdiv id=\\\"Sec9\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003eA. The sublingual salivary glands:\\u003c/h2\\u003e \\u003cp\\u003eThe first sign of the sublingual salivary gland primordia could be seen in the 6-day-old quail embryo as an epithelial thickening of compacted cellular mass (epithelial placode and prebud) within the soft sublingual floor epithelium (Fig.\\u0026nbsp;1A \\u0026amp; 1B). Consequently, we noticed that the highly proliferated epithelial masses extended from the bud form to the elongated cords at the 10-day-old quail embryo. Interestingly, these cords have two ends; a rostromedial wide end and a caudolateral narrow end. Canalization was observed in some cords (Fig.\\u0026nbsp;1C \\u0026amp;1C*). By advancing age, the lobar cords of the sublingual salivary glands increased in their length and run more caudally without branching at the 11-day-old quail embryo. The rostral end of the cord was broad, less proliferative, and encased by a concentric layer of mesenchyme, whereas the narrow caudolateral proliferative end was still uncovered, suggesting further growth and expansion. Surprisingly, the cord was constricted nearly mid-distance. Due to cell growth competition and apoptosis, many halo vacuoles were obviously present within the rostral part of the cords indicating the beginning of canalization (Fig.\\u0026nbsp;1D). We observed that canalization started rostrally (proximally) and progressed distally (caudally), and the glands extended without branching.\\u003c/p\\u003e \\u003cp\\u003eBy the 13th day of incubation, we found the many vacuoles within the glandular endopieces mass coalesced together to form a single common canal rostrally. Consequently, the lining of the glandular epithelium showed a transformation process from stratified epithelium to monolayer (simple glandular epithelium). Thus, the lumen filled with a secretory-like substance representing the sloughed lining epithelium (Fig.\\u0026nbsp;2A \\u0026amp; 2B). Moreover, our study found that the exfoliated substance gave PAS/Alcian blue negative results, at this age. Then, at the maturation stage, the secretory terminals were presented with more lining corrugation and cellular compaction. The lining epithelium showed low columnar cells with foamy cytoplasm and basal vesicular nuclei. Conceptively, near the opening, the lining epithelium transformed from high columnar to low columnar. Thereafter, it ends by stratified squamous epithelium whereas the opening of the taste bud-associating salivary glands found in the hatching chick (Fig.\\u0026nbsp;2C \\u0026amp; 2D). Collectively, the first few days of embryo incubation showed highly developmental glandular events.\\u003c/p\\u003e \\u003cp\\u003eOur gross anatomical examination of 14-day-old chicks revealed that the sublingual glands were paired, consisting of 5\\u0026ndash;7 parallel elongated lobes with broad blind ends which, directed rostromedially, and narrow ends directed caudolaterally (Figs.\\u0026nbsp;4). The glandular lobes extended longitudinally from the mandibular symphysis rostrally within the sublingual floor to beyond the level of the proximal part of the ceratobranchial. As we found the lobes were constricted at the level of the basihyal (Figs.\\u0026nbsp;3), where they supported the laryngeal mound therefore, it was suggested the significant size of the larynx. Hence, the lobes decreased in the lobation number at that portion (Figs.\\u0026nbsp;4). Concequently, the lobes were numerous rostrally and few caudally whereas are variable from 3\\u0026ndash;4 lobes caudally (Figs.\\u0026nbsp;3) and (Figs.\\u0026nbsp;4). We conclude that there was an asymmetrical bilateral glandular lobes number along the same side.\\u003c/p\\u003e \\u003cp\\u003eOur post-hatching histological studies at 14-and 30-days old quail chicks, suggested that the sublingual salivary glands were non-branched compound tubuloalveolar mucous glands that were supported ventrolaterally by muscle mylohyoideus, and a thin connective tissue stroma. Interestingly, the median sulcus was separated between two glandular masses rostrally. The secretory endpieces of each lobule drained their secretion directly into a common secretory duct that opened into the soft sublingual floor separately or collectively with other canals (polystomatic). The ductal epithelial lining was simple tall columnar cells. The ductal canal was narrow while, it passed via the epithelium to end by the excretory duct (Fig.\\u0026nbsp;5A-D). Notably, our histological data confirmed here that the lobation at the level of the sublingual floor were 5\\u0026ndash;7 bilaterally (Fig.\\u0026nbsp;5A). But, caudally the lobation was five on the right side as follow; two larger, two medium-sized, and one smaller. However, the left side had three lobes; two larger and one smaller. As we noticed that the lobes were numerous rostrally and few caudally. So that the common canals of the sublingual glands were narrower rostrally than the caudal lobes (Fig.\\u0026nbsp;6A-C \\u0026amp; Fig.\\u0026nbsp;7A-B). Also, we found an autonomic ganglion that embedded among secretory lobules (Fig.\\u0026nbsp;6D \\u0026amp; 6E). Interestingly, the mucous of the sublingual salivary glands showing the apocrine mode of secretion with cellular and nuclear contents detaching within the lumen(Fig.\\u0026nbsp;7C-D).\\u003c/p\\u003e \\u003cp\\u003eIn this study, we found two types of taste buds that have a barrel shape as following; the taste buds with taste pores which was called taste buds-associated salivary glands, and surface epithelial taste buds (Fig.\\u0026nbsp;8A-B). Surface epithelial taste-buds were characterized by deep dermal papillae within the epithelium and deeply stained central cells assumed as gustatory cells (Fig.\\u0026nbsp;8C-D). While the taste buds-associated salivary glands opening, were surrounded by highly mitotic polygonal basal cells within the barrel shape structure (Fig.\\u0026nbsp;8E-F)\\u003c/p\\u003e \\u003cp\\u003eOur SEM (Figs.\\u0026nbsp;9) and morphometrical measurements (Table\\u0026nbsp;1) revealed that at 8th days old quail embryo the soft sublingual floor was full of numerous domes, mushrooms or spherical like taste buds with various sizes; defined into three different sized. We categorized them into Larger, moderate, and smaller sized taste buds which were ranging from (40, 12, 6) um\\u003csup\\u003e2\\u003c/sup\\u003e respectively. Also, the data showed some large sized taste buds nearly fused together (Fig.\\u0026nbsp;9A). By the age of the 10th day old embryo, we noticed that the taste buds had surface area of 36 um\\u003csup\\u003e2\\u003c/sup\\u003e and sublingual salivary glands opening diameter was 5 um. Notably, the taste buds decreased in the number and size. However, some of them fused together to form a large one of 144 um\\u003csup\\u003e2\\u003c/sup\\u003e (Fig.\\u0026nbsp;9B). With age progression (13 days pre-hatching), the sublingual glands opening increased in the diameter and reached to 28 um with no secretion could be seen. However, the taste bud-associated salivary glands could be seen, with surface area equal to 35 um\\u003csup\\u003e2\\u003c/sup\\u003e (Fig.\\u0026nbsp;9C). At 30- and 60-days old chick, various sized taste buds could be measured as follow; the larger (32 um\\u003csup\\u003e2\\u003c/sup\\u003e), moderate (12 um\\u003csup\\u003e2\\u003c/sup\\u003e), and smaller (9 um\\u003csup\\u003e2\\u003c/sup\\u003e) sized, and their taste pores were varied from 8.2\\u0026ndash;12 um. These taste buds were emerging through the scales of the surface of the sublingual floor epithelium and mucous secretion exited via the salivary gland opening (Fig.\\u0026nbsp;9D-F). Consequently, our data indicated that the taste buds were different in size and position.\\u003c/p\\u003e \\u003cp\\u003eOf note, our histochemical analysis of the sublingual glands (Table\\u0026nbsp;2A) showed a strong positive reaction with PAS and combined AB/PAS stains, while weakly reaction to AB stain in the newly hatched quail chick (Fig.\\u0026nbsp;10A-C). Moreover, in the seventh day old chick, the sublingual glands were shown moderate positive reaction to alcian blue and strong to PAS and combined AB/PAS stains (Figs.\\u0026nbsp;10D-F). However, at 14 days old quail chick, these glands showed moderate positive reaction to AB, very strong positive reaction to PAS and to combined AB/PAS stains (Fig.\\u0026nbsp;11A-C). In 30 days old quail, the sublingual glands showed strong positive reaction to AB stain and very strong positive to combined AB/PAS stains (purple coloration), but negative results to PAS stain (Fig.\\u0026nbsp;11D-E). However, in the 60th days old quails, the glands showed negative results to PAS stain and very strong positive reaction to AB stain and very strong alcianophilic reaction to AB/PAS stains (greenish blue coloration) (Fig.\\u0026nbsp;11F).\\u003c/p\\u003e \\u003c/div\\u003e \\u003cdiv id=\\\"Sec10\\\" class=\\\"Section2\\\"\\u003e \\u003ch2\\u003e\\u003cb\\u003eB- The mandibular salivary glands\\u003c/b\\u003e:\\u003c/h2\\u003e \\u003cp\\u003eDevelopmentally, we found that the mandibular glands appeared at 10 days old embryo arising from the epithelium of the paralingual groove. We observed that the developmental characterization of the gland was like the sublingual glands in almost stages. However, the mandibular glands were different in branching of the glandular endpieces (Fig.\\u0026nbsp;12A-F). By gross studies, we found the glands extended caudal to the frenulum linguae, lying within the floor of the paralingual grooves (Fig.\\u0026nbsp;3B). They are paired compound tubuloalveolar glands lay in the tunica submucosa of the paralingual grooves of the oropharynx. The mandibular glands extend longitudinally from the rostral border of the frenulum linguae to the caudal tips of sublingual glands. They lay on the dorsomedial border of the sublingual glands. They open into the ventral or medial aspect of the paralingual groove (Fig.\\u0026nbsp;13A-B). Once again, the mode of the secretion resembles the sublingual gland. Moreover, the glandular acini separated with thick connective tissue septa (interlobular C.T). (Fig.\\u0026nbsp;7E-F).\\u003c/p\\u003e \\u003cp\\u003eThe histochemical study (Table\\u0026nbsp;2B) suggested that the mandibular salivary glands in 30 days old quails showed very strong positive reaction to all of the alcian blue, PAS and combined AB/PAS stains (Fig.\\u0026nbsp;13C-F). By the 60-day old quail, these glands showed a strong positive reaction to AB stain and combined AB/PAS stains (greenish blue coloration only), but negative results to PAS stain. Figures\\u0026nbsp;(14) is a schematic diagram showing the secretory units of the compound tubuloalveolar sublingual salivary glands.\\u003c/p\\u003e \\u003c/div\\u003e\"},{\"header\":\"Discussion\",\"content\":\"\\u003cp\\u003eSalivary glands are unique specialized structures, which play a vital role in mammals and birds [\\u003cspan citationid=\\\"CR22\\\" class=\\\"CitationRef\\\"\\u003e22\\u003c/span\\u003e]. In this study, we focused on the paired oropharyngeal floor-related salivary glands of the Japanese quail. Both glands called the sublingual and mandibular salivary glands as anatomical nomenclature followed [\\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e23\\u003c/span\\u003e] in chicken. However, [\\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e23\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR24\\\" class=\\\"CitationRef\\\"\\u003e24\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e25\\u003c/span\\u003e] suggested that Glandula sublingualis synonymies to Glandula submandibularis rostralis or mandibularis rostralis, while the Glandula mandibularis Synonymies to (Gl. Mandibularis caudalis) or (Gl. Submandibularis caudalis). Also,[\\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e3\\u003c/span\\u003e] divided them into lateral and medial mandibular glands in emu. Of note, [\\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e23\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR24\\\" class=\\\"CitationRef\\\"\\u003e24\\u003c/span\\u003e] discussed both glands deeply in chicken. But few histological and histochemical studies have been done on sublingual and mandibular salivary glands in quails.\\u003c/p\\u003e \\u003cp\\u003eFrom our prehatching data, we found that the sublingual and mandibular salivary glands firstly appeared as a thick invagination (a bud stage) of the sublingual floor and paralingual epithelium, at the 6th \\u0026amp; 10th day pre-hatching old respectively in agreement to [\\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e4\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR26\\\" class=\\\"CitationRef\\\"\\u003e26\\u003c/span\\u003e]. Although [\\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e25\\u003c/span\\u003e] assumed that the primordia arose at 8-day quail from the oral floor epithelium that was extended without branching in agreement with us. Additionally, as we revealed that the canalization process of the sublingual gland started and became obviously detected at the 10th and 11th days of the incubation respectively. Hence, the transformation process of the glandular lining from stratified epithelium to monolayer (simple glandular epithelium). Moreover, the branching of the sublingual gland primordia occurred at 13th-day pre-hatching. To find that [\\u003cspan citationid=\\\"CR25\\\" class=\\\"CitationRef\\\"\\u003e25\\u003c/span\\u003e] noticed also the lingual salivary gland had a branching mechanism that was different from the sublingual salivary glands. However, it liked as the mandibular salivary glands as we approved. Interestingly, the murine antenatal morphogenesis of the salivary glands branching was a clear example in prebud, initial, canalicular, and terminal bud stages [\\u003cspan citationid=\\\"CR27\\\" class=\\\"CitationRef\\\"\\u003e27\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR28\\\" class=\\\"CitationRef\\\"\\u003e28\\u003c/span\\u003e].\\u003c/p\\u003e \\u003cp\\u003eAll the prenatal developmental stages of the sublingual salivary glands passed with cellular proliferation, quiescence, apoptotic, and cytodifferentiation events, in which were directed by special growth factors, cytokines, and signaling pathways that target transcription factors, to be upregulated spatially and temporally [\\u003cspan citationid=\\\"CR29\\\" class=\\\"CitationRef\\\"\\u003e29\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR30\\\" class=\\\"CitationRef\\\"\\u003e30\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR31\\\" class=\\\"CitationRef\\\"\\u003e31\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR28\\\" class=\\\"CitationRef\\\"\\u003e28\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR32\\\" class=\\\"CitationRef\\\"\\u003e32\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR33\\\" class=\\\"CitationRef\\\"\\u003e33\\u003c/span\\u003e]. Obviously, it has been shown that sublingual glands morphogenesis was dependent on the cross-talking between the epithelial-mesenchymal sides [\\u003cspan citationid=\\\"CR34\\\" class=\\\"CitationRef\\\"\\u003e34\\u003c/span\\u003e].\\u003c/p\\u003e \\u003cp\\u003eOur prehatching results revealed the presence of the secretory-like product and cell fragment within the lumina at the 13-day-old embryo, which gave negative results with AB or PAS stains in agreement with [\\u003cspan citationid=\\\"CR35\\\" class=\\\"CitationRef\\\"\\u003e35\\u003c/span\\u003e] who assumed that the secretory products appear on the 15th day old chick embryo. Also,[\\u003cspan citationid=\\\"CR36\\\" class=\\\"CitationRef\\\"\\u003e36\\u003c/span\\u003e] detected that acidic secretion was released from the developing proventriculus at the 13th days old chick embryo because of the swallowed albumin stimulation. A similar finding was found in quail\\u003csup\\u003e4\\u003c/sup\\u003e.\\u003c/p\\u003e \\u003cp\\u003eHence, our results within the sublingual salivary glands indicated that the acini opened directly into a large common wide secretory lumen of the lobule which thus serves as a common excretory duct to open into the oral cavity either may be fused with another lobular common duct or opened separately. The secretory system was lined with the same cells type of the secretory end pieces; simple columnar cells that were transformed to low columnar to finally become stratified epithelium in agreement to [\\u003cspan citationid=\\\"CR35\\\" class=\\\"CitationRef\\\"\\u003e35\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR36\\\" class=\\\"CitationRef\\\"\\u003e36\\u003c/span\\u003e]. Moreover, the secretory end pieces of the other salivary glands have drained their secretion within intralobuar ducts and then into interlobar duct to the common secretory canal. [\\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e37\\u003c/span\\u003e] found that the columnar cells lining of the glandular duct surrounding the lumen and with microvilli with the apical cellular portion in posterior lingual salivary gland in quail similar to our findings. Moreover,\\u003csup\\u003e3\\u003c/sup\\u003e supported the presence of a central canal, but the glands are simple branched tubular in emu and the lining was simple ciliated columnar cells or pseudostratified columnar epithelium. While in the present study it was compound tubuloalveolar gland in agreement with \\u003csup\\u003e38\\u003c/sup\\u003e in rock dove. In support to the previous results, the sublingual salivary glands of mice consisted of large and small ducts that ended in a single lumen-containing secretion [\\u003cspan citationid=\\\"CR31\\\" class=\\\"CitationRef\\\"\\u003e31\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR27\\\" class=\\\"CitationRef\\\"\\u003e27\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR28\\\" class=\\\"CitationRef\\\"\\u003e28\\u003c/span\\u003e].\\u003c/p\\u003e \\u003cp\\u003eSurprising, it was clear from gross and histological data that the sublingual glands lobation was different from [\\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e23\\u003c/span\\u003e] in chicken who assumed that the glands consisted of 5\\u0026ndash;7 lobes. However, we found that the lobation number was different bilaterally and unilaterally, and depended on the location. Importantly, no author defined the details or pointed out to that even if in quails or chicken.\\u003c/p\\u003e \\u003cp\\u003eThe current study showed that the mandibular gland was paired compound branched tubulo-alveolar glands within the submucosa of the paralingual floor of the mouth. Whereas, it extended from the caudal of the frenulum linguae to the tips of the gland sublingualis. Their orifices were opened into the bottom of the paralingual groove. These results were like that of [\\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e37\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR10\\\" class=\\\"CitationRef\\\"\\u003e10\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e43\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR23\\\" class=\\\"CitationRef\\\"\\u003e23\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e3\\u003c/span\\u003e]. However, [\\u003cspan citationid=\\\"CR39\\\" class=\\\"CitationRef\\\"\\u003e39\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR40\\\" class=\\\"CitationRef\\\"\\u003e40\\u003c/span\\u003e] assumed the mandibular salivary glands to be a part of the gland sublingualis referred to that, as being the posterior part of a single gland, of which the gland sublingualis is the anterior part. Notably, we observed that the glands were supported with thick connective tissue stroma compatible with [\\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e3\\u003c/span\\u003e].\\u003c/p\\u003e \\u003cp\\u003eThe present SEM and histological studies about taste buds in Japanese quails were similar to our previous studies. Whereas taste buds were two types; the larger one is associated with the salivary glands opening and the smaller one called surface taste buds lay free on the oral surface mucosa corresponding to the findings in chicken [\\u003cspan citationid=\\\"CR41\\\" class=\\\"CitationRef\\\"\\u003e41\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR43\\\" class=\\\"CitationRef\\\"\\u003e43\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR61\\\" class=\\\"CitationRef\\\"\\u003e61\\u003c/span\\u003e], and in emu, [\\u003cspan citationid=\\\"CR3\\\" class=\\\"CitationRef\\\"\\u003e3\\u003c/span\\u003e]. However, our past morphometric studies mentioned that the taste pore of taste buds associated with preglottal gland had no change in diameter (8.17 \\u0026micro;m) at 14- and 30-days old quail chick. While the preglottal salivary gland openings widths were 25.5 \\u0026micro;m, and 80.4 \\u0026micro;m at 14- and 30-days old quail chick respectively [\\u003cspan citationid=\\\"CR4\\\" class=\\\"CitationRef\\\"\\u003e4\\u003c/span\\u003e]. However,[\\u003cspan citationid=\\\"CR61\\\" class=\\\"CitationRef\\\"\\u003e61\\u003c/span\\u003e] found that taste buds parameters in chicken were (40\\u0026ndash;70 \\u0026micro;m) in width and (70\\u0026ndash;120 \\u0026micro;m) in height. While taste pore was (3\\u0026ndash;7 \\u0026micro;m) in width. Although, the openings of the salivary glands are greater than (10 \\u0026micro;m) in diameter and were observed near the taste bud.\\u003c/p\\u003e \\u003cp\\u003eIn addition to that, other authors\\u0026rsquo; data revealed that taste pores width was equal to (5\\u0026ndash;10 \\u0026micro;m)[\\u003cspan citationid=\\\"CR41\\\" class=\\\"CitationRef\\\"\\u003e41\\u003c/span\\u003e] or (6 \\u0026micro;m) [\\u003cspan citationid=\\\"CR42\\\" class=\\\"CitationRef\\\"\\u003e42\\u003c/span\\u003e].\\u003c/p\\u003e \\u003cp\\u003eAccordingly, in the present study (Table\\u0026nbsp;1) showed the average diameter in (um) of the sublingual salivary gland openings (SGO), the taste buds openings (TBO), and the taste buds surface area (TBA) represented by (um\\u003csup\\u003e2\\u003c/sup\\u003e) in quails. Conclusively, we found out that, taste buds were categorized into three sized groups; larger, moderate, and smaller, and according to the position of taste buds associated with salivary glands opening and surface epithelial one. Also, taste pores varied from (8.2\\u0026ndash;12 \\u0026micro;m). These confusing results of previous literatures were because the measurement of many authors trials neglected the fact of different sized, and the positioned taste buds as we did. Notably, we assumed that salivary glands act as a flusher by mucous secretion for taste buds associated with its opening, like Ebner\\u0026rsquo;s glands in mammals. Finally, we did not think that many literatures spoke about the taste buds in detail in quail as we did, suggesting that they have a vital role in food discrimination and prehension.\\u003c/p\\u003e \\u003cp\\u003eOur histological data indicated that the sublingual and mandibular salivary glands secreted mucous in agreement with [\\u003cspan citationid=\\\"CR44\\\" class=\\\"CitationRef\\\"\\u003e44\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR45\\\" class=\\\"CitationRef\\\"\\u003e45\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR46\\\" class=\\\"CitationRef\\\"\\u003e46\\u003c/span\\u003e] who surveyed salivary glands of some birds to have a mucous secretion type. Referring to our histochemical results, the glands were strongly positive to PAS stain, but they were weakly to AB stain at the hatching day of chicks. Then reacting negatively to PAS, but strongly to AB stain at 30- and 60-days post-hatching old. Moreover, [\\u003cspan citationid=\\\"CR47\\\" class=\\\"CitationRef\\\"\\u003e47\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR48\\\" class=\\\"CitationRef\\\"\\u003e48\\u003c/span\\u003e] found that the posterior lingual salivary glands were robustly positive for neutral mucopolysaccharides than the anterior one in both Common Myna and red jungle fowl. However, there no mucosubstance in lingual salivary glands of the little erget [\\u003cspan citationid=\\\"CR49\\\" class=\\\"CitationRef\\\"\\u003e49\\u003c/span\\u003e]. In addition, [\\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e37\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e16\\u003c/span\\u003e] supported the presence of sialo glycoconjugates within the preglottal and the lingual salivary glands in Japanese quails. Also, this was stated that in the salivary glands of chicken by [\\u003cspan citationid=\\\"CR50\\\" class=\\\"CitationRef\\\"\\u003e50\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e51\\u003c/span\\u003e].\\u003c/p\\u003e \\u003cp\\u003eIn addition, [\\u003cspan citationid=\\\"CR37\\\" class=\\\"CitationRef\\\"\\u003e37\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e16\\u003c/span\\u003e] supported the presence of sialo glycoconjugates within the preglottal and the lingual salivary glands in Japanese quails. Also, this was stated in the salivary glands of chicken by [\\u003cspan citationid=\\\"CR50\\\" class=\\\"CitationRef\\\"\\u003e50\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR51\\\" class=\\\"CitationRef\\\"\\u003e51\\u003c/span\\u003e]. However, palatine salivary glands of chicken produced glycoproteins, sulphomucins, and carboxymucins, which were rich in sialic acid showed more PAS and alciniophilic containing metachromatic granules [\\u003cspan citationid=\\\"CR52\\\" class=\\\"CitationRef\\\"\\u003e52\\u003c/span\\u003e]. Of note,[\\u003cspan citationid=\\\"CR53\\\" class=\\\"CitationRef\\\"\\u003e53\\u003c/span\\u003e] suggested that the lingual glands were more acidic with sulphated or acidic sialomucin. Accordingly, the double differential staining, of alcian blue (AB) and PAS stains differentiated between acidic glycoconjugates stained (blue) with AB and neutral one stained (magenta) with PAS in agreement to [\\u003cspan citationid=\\\"CR50\\\" class=\\\"CitationRef\\\"\\u003e50\\u003c/span\\u003e] in chicken. From that point, we concluded that the secretory mucous took the fate from neutral to acidic result.\\u003c/p\\u003e \\u003cp\\u003eA tentative interpretation of the type of the lingual gland secretions in little egret could be made according to the classification of mucosubstances were proposed by other workers [\\u003cspan citationid=\\\"CR54\\\" class=\\\"CitationRef\\\"\\u003e54\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR55\\\" class=\\\"CitationRef\\\"\\u003e55\\u003c/span\\u003e, \\u003cspan citationid=\\\"CR56\\\" class=\\\"CitationRef\\\"\\u003e56\\u003c/span\\u003e]. The lingual salivary gland of little egret was exhibited to be alcianophilia and reacted positively to the different techniques employed for proteins detection. Accordingly, these glands are confirmed to be of mucoserous type. The lingual glands of chicken [\\u003cspan citationid=\\\"CR57\\\" class=\\\"CitationRef\\\"\\u003e57\\u003c/span\\u003e], penguin [\\u003cspan citationid=\\\"CR58\\\" class=\\\"CitationRef\\\"\\u003e58\\u003c/span\\u003e] (Samar et al. 1995), white-cheeked bulbul [\\u003cspan citationid=\\\"CR59\\\" class=\\\"CitationRef\\\"\\u003e59\\u003c/span\\u003e], and Eurasian collared dove [\\u003cspan citationid=\\\"CR60\\\" class=\\\"CitationRef\\\"\\u003e60\\u003c/span\\u003e] (Taib and Jarrar, 2001), were reported to be of mucous type, however the quail [\\u003cspan citationid=\\\"CR16\\\" class=\\\"CitationRef\\\"\\u003e16\\u003c/span\\u003e] and the little [\\u003cspan citationid=\\\"CR49\\\" class=\\\"CitationRef\\\"\\u003e49\\u003c/span\\u003e] had a mixture of serous and mucous type salivary glands. The secretory products of the lingual salivary glands of these species of birds as seen in the present study form a blend of mucoserous secretions that contained sialomucins, sulfomucins, and proteins\\u003csup\\u003e49\\u003c/sup\\u003e. To illustrate that our result found the glands were well developed in quails; and took acidic reaction with age advancing as [\\u003cspan citationid=\\\"CR58\\\" class=\\\"CitationRef\\\"\\u003e58\\u003c/span\\u003e] finding. As our findings showing the secretory units of the compound tubuloalveolar sublingual salivary glands (Fig.\\u0026nbsp;14).\\u003c/p\\u003e\"},{\"header\":\"Conclusion\",\"content\":\"\\u003cp\\u003eIn conclusion, we assumed that the sublingual salivary glands and mandibular salivary glands were the basic feedings adapting units among the mucous salivary glands system. Consequently, such developmental and histochemical structures of the sublingual salivary glands in the quail play a role in various food intakes and are responsible for the softening and formation of food chows. These findings of oropharyngeal glands survey give a good future imagination in the quail and other birds for for further study.\\u003c/p\\u003e\"},{\"header\":\"Declarations\",\"content\":\"\\u003cp\\u003e\\u003cstrong\\u003eCompeting interests\\u003c/strong\\u003e\\u003c/p\\u003e\\n\\u003cp\\u003eThe author(s) declare no competing interests.\\u003c/p\\u003e\\n\\u003cp\\u003e\\u003cstrong\\u003eAvailability of materials and data\\u0026nbsp;\\u003c/strong\\u003e\\u003c/p\\u003e\\n\\u003cp\\u003eThe data that support the findings of this study are available on request from the corresponding author.\\u003c/p\\u003e\\n\\u003cp\\u003e\\u003cstrong\\u003eAuthor contributions\\u003c/strong\\u003e\\u003c/p\\u003e\\n\\u003cp\\u003eM.O.K. \\u0026nbsp;designed the research; M.O.K. acquired the data. M.O.K., M.A., W.G., and A.M.S. analyzed the data, and M.O.K. wrote the manuscript. All authors discussed the results and commented on the manuscript.\\u003c/p\\u003e\\n\\u003cp\\u003e\\u003cstrong\\u003eEthics Declarations\\u003c/strong\\u003e\\u003c/p\\u003e\\n\\u003cp\\u003e\\u003cstrong\\u003eEthical approval\\u003c/strong\\u003e\\u003c/p\\u003e\\n\\u003cp\\u003eOur experiment was done as well as all quail bird experiments were performed following the guidance and administration of the Animal Research Committee at the Faculty of Veterinary Medicine, Assiut University, Assiut, Egypt. License No. (06/2023/0051).\\u003c/p\\u003e\\n\\u003cp\\u003e\\u003cstrong\\u003eAcknowledgments\\u003c/strong\\u003e\\u003c/p\\u003e\\n\\u003cp\\u003eThis work is a collaborative research program between Assiut University and Aswan University. The funders play no role in the study design, data collection, analysis, decision to publish, or preparation of the manuscript.\\u003c/p\\u003e\"},{\"header\":\"References\",\"content\":\"\\u003col\\u003e\\n\\u003cli\\u003eKing, A. S., \\u0026amp; Mclelland, J. (1984). Birds their structure and function (2nd ed.). London: Bailliere Tindall.\\u003c/li\\u003e\\n\\u003cli\\u003eNalavade, M. N. and Varute, A. T. (1977): Histochemical analysis of mucosubstances in the lingual glands and taste buds of some birds. Acta Histochem. 60: 18-31.\\u003c/li\\u003e\\n\\u003cli\\u003eCrole, M.R, Soley, J. T. 2011. Distribution and structure of the glandular tissue in the oropharynx and proximal esophagus of the emu (Dromaius novaehollandiae). Acta Zoologica (Stockholm). 92: 206-215.\\u003c/li\\u003e\\n\\u003cli\\u003eKhalifa M. O., Abd-Elkareem M., Gaber W., Li T-S., Saleh A. M. Developmental morphological analyses on the preglottal salivary gland in Japanese quails (Coturnix japonica) (2022). Microsc Res Tech. 85(1):156\\u0026ndash;68.\\u003c/li\\u003e\\n\\u003cli\\u003eGill, F.B. Ornithology. 2nd edition. New York: W.H. Freeman and Company, 1994.\\u003c/li\\u003e\\n\\u003cli\\u003eOatley TB, Earl\\u0026eacute; RA, Prins AJ. The diet and foraging behavior of the ground. Ostrich. 1989.1;60(2):75\\u0026ndash;84.\\u003c/li\\u003e\\n\\u003cli\\u003eHomberger, D.G. (1986): The lingual apparatus of the African grey parrot, Psittacus erithacus Linn\\u0026eacute; (Aves: Psittacidae): description and theoretical mechanical analysis. Washington, D.C.: Ornithological Monographs No. 39.\\u003c/li\\u003e\\n\\u003cli\\u003eKhalifa M. O., Abd-Elkareem M., Gaber W., Li T-S., Saleh A. M. Developmental morphological analyses on the preglottal salivary gland in Japanese quails (Coturnix japonica) (2022). Microsc Res Tech. 85(1):156\\u0026ndash;68.\\u003c/li\\u003e\\n\\u003cli\\u003eToryu, Y.; Hoshino, T. and Tamate, H. (1960): Histological study of the lingual salivary glands in the chicken with special reference to the occurrence of the glycogen in the gland cell nuclei. Tohoku J. agr. Res., 11: 309.\\u003c/li\\u003e\\n\\u003cli\\u003eMclelland, J. (1975). Aves digestive system. In Sission and Grossman\\u0026apos;s, the anatomy of the domestic animals (Vol. 2, 5th ed.). Philadelphia: Getty R. Saunders. \\u003c/li\\u003e\\n\\u003cli\\u003eMohamed, A. A. (2010): Some developmental studies on the floor of the oropharynx of the duck. M.Sc. thesis, South Valley University, Egypt.\\u003c/li\\u003e\\n\\u003cli\\u003ePortman, A. \\u0026quot;Sensory Organs: Skin, Taste and Olfaction.\\u0026quot; In Biology and Comparative Physiology of Birds, Vol. II, by A.J. Marshall, 37-48. New York and London: Academic Press, 1961.\\u003c/li\\u003e\\n\\u003cli\\u003eProctor, N.S., Lynch, P.J. Manual of Ornithology. New Haven and London: Yale University Press, 1993.\\u003c/li\\u003e\\n\\u003cli\\u003eBeason, R.C. \\u0026quot;Through a Bird\\u0026apos;s Eye - Exploring Avian Sensory Perception. (2003). Bird Strike Committee USA/Canada, 5th Joint Annual Meeting. Toronto: Internet Center for Wildlife Damage Management.\\u003c/li\\u003e\\n\\u003cli\\u003eMenghi, G.; Scocco, P. and Ceccarelli, P. (1993): Basic and lectin histochemistry for studying glycoconjugates in the lingual salivary glands of the Japanese quail (Coturnix coturnix japonica). Arch. Oral. Biol., 38: 649-655.\\u003c/li\\u003e\\n\\u003cli\\u003eLiman, N., Bayram, G. and Koceak, M. (2001): Histological and histochemical studies on the lingual, preglottal and laryngeal salivary glands of the Japanese quail (Coturnix coturnix japonica) at the post-hatching period. Anat. Histol. Embryol. 30, 367\\u0026plusmn;373. https://doi.org/10.1046/j.1439-0264.2001.00353.x\\u003c/li\\u003e\\n\\u003cli\\u003eHarris, H. (1900). On the rapid conversion of haematoxylin into haematein in staining reactions. Journal of Applied Microscopic Laboratory Methods, 3, 777. https://www.scienceopen.com/document?vid=c2c7d328-e08c429d-b85c-fa8c3371e3b8. \\u003c/li\\u003e\\n\\u003cli\\u003eCrossmon, G. (1937). 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Dev., 74:179-183.\\u003c/li\\u003e\\n\\u003cli\\u003eMelnick M, Chen H, Zhou Y-M, Jaskoll T.(2001): Embryonic mouse submandibular salivary gland morphogenesis and the TNF/TNF-R1 signal transduction pathway. Anat. Rec.262:318-320.\\u003c/li\\u003e\\n\\u003cli\\u003eJaskoll T, Zhou Y-M, Chai Y, Makarenkova HP, Collinson JM, West JD, Hajihosseini MK, Lee J, Melnick M Embryonic submandibular gland morphogenesis: stage-specific protein localization of FGFs, BMPs, Pax 6 and Pax 9 and abnormal SMG phenotypes in Fgf/R2- IIIc, BMP7 and Pax6 mice. (2002): Cells, Tissues, Organs. 270:83-98.\\u003c/li\\u003e\\n\\u003cli\\u003eKashimata M, Gresik E: Epidermal growth factor system is a physiological regulator of development of the mouse fetal submandibular gland and regulates expression of the \\u0026quot;6- integrin subunit.(1997). Develop. Dyn.208:149-161. \\u003c/li\\u003e\\n\\u003cli\\u003eJaskoll T, Melnick M (1999b): Submandibular gland morphogenesis: stage-specific expression of TGFalpha, EGF, IGF, TGF-beta, TNF and IL-6 signal transduction in normal mice and the phenotypic effects of TGF-beta2, TGF-beta3, and EGF-R null mutations. Anat. Rec., 256:252-268. \\u003c/li\\u003e\\n\\u003cli\\u003eKashimata M, Sayeed S, Ka A, Onetti-Muda A, Sakagami H, Faraggiana T, Gresik E: The ERK1/2 signaling pathway is involved in the stimulation of branching morphogenesis of fetal mouse submandibular glands by EGF.(2000). Developmental Biology. 220:183-196.\\u003c/li\\u003e\\n\\u003cli\\u003eHardman P, Landels E, Woolfe A, Spooner B: TGF-beta1 inhibits growth and branching morphogenesis in embryonic mouse submandibular and sublingual glands in vitro. (1994). Dev. Growth Diff.36:567-577.\\u003c/li\\u003e\\n\\u003cli\\u003eCutler LS, Gremski W.(1991):Epithelial-mesenchymal interactions in the development of salivary glandsCrit Rev Oral Biol Med.2:1-12.\\u003c/li\\u003e\\n\\u003cli\\u003eShih, L. F. (1966): A histological and histochemical study of the developing salivary gland in the chick (Gallus domesticus).M. SC. Thesis, Smcgill University, Montreal, Canada.\\u003c/li\\u003e\\n\\u003cli\\u003eTivane C., Histological features and surface morphology of the oropharyngeal cavity \\u0026amp; proximal oesophygus. (2008). Master dissertation, Chapter 3, University of Pretoria, Pretoria, South Africa.\\u003c/li\\u003e\\n\\u003cli\\u003eCapacchletti, M., Sabbiet, M. G., Agas, D., Materazzi, S., Menghi, G., and Marchetti, L., (2009). Ultrastructure and lectin cytochemistry of secretory cells in lingual glands of the Japanese quail (Coturnix coturnix japonica). Histol. Histopathol., 24: 1087-1096.\\u003c/li\\u003e\\n\\u003cli\\u003eAl-Nefeiy, F. A. and Alahmary, B. A. (2015): Morphological, histological and histochemical studies of the lingual salivary glands of the rock dove, Patagioenas livia (Columbidae). International Journal of current research and academic review. ISSN: 2347-3215 Volume 3 Number 8 pp. 280-289.\\u003c/li\\u003e\\n\\u003cli\\u003eMcleond, W. M.; Trotter, D. M. and Lumb, J. W. (1964): Avian anatomy Burgess, Minneapolis.\\u003c/li\\u003e\\n\\u003cli\\u003eHodges, R. D. (1974). The histology of the fowl. London; New York: Academic Press.\\u003c/li\\u003e\\n\\u003cli\\u003eGentle M. J. (1971): The lingual taste buds of Gallus domesticus L. British Poultry Science 12, 245\\u0026ndash;248.\\u003c/li\\u003e\\n\\u003cli\\u003eSaito, I. (1965). Comparative anatomical studies of the oral organs of the poultry. IV. Macroscopical observation of the salivary glands. Bulletin of the Faculty of Agriculture, 12, 110\\u0026ndash;120.\\u003cspan dir=\\\"LTR\\\"\\u003e\\u003c/span\\u003e\\u003c/li\\u003e\\n\\u003cli\\u003eNickel, R., Schummer, A., \\u0026amp; Seiferle, E. (1977). Anatomy of the domestic birds. Berlin, Germany: Verlag Paul Parey.\\u003c/li\\u003e\\n\\u003cli\\u003eSchauder, W. (1923): Anatomie der Hausvogel. In: Martints Lehrbuch der Anatomie der Haustiere. Stuttgart.\\u003c/li\\u003e\\n\\u003cli\\u003eGrossman, J. D. (1927). Some anatomical features of the fowl. Ohio Veterinary Alumni Quarterly, 15, 27\\u0026ndash;35.\\u003c/li\\u003e\\n\\u003cli\\u003eCalhoun, M. L. (1954). Microscopic anatomy of the digestive system of the chicken. Ames, IA: Iowa State University Press. Crossmon, G. (1937). A modification of Mallory\\u0026apos;s connective tissue stain with a discussion of principles involved. The Anatomical Record, 69(1), 33\\u0026ndash;38.\\u003c/li\\u003e\\n\\u003cli\\u003eKadhim, K. K., Hameed, A.-T., \\u0026amp; Abass, T. A. (2013). Histomorphological and Histochemical observations of the common myna (Acridotheres tristis) tongue. ISRN Veterinary Science, 2013, 980465. https://doi.org/ 10.1155/2013/980465.\\u003c/li\\u003e\\n\\u003cli\\u003eKadhim, K. K., Zuki, A. B. Z., Babjee, S. M. A., Noordin, M. M., \\u0026amp; ZamriSaad, M. (2011). Morphological and histochemical observations of the red jungle fowl tongue Gallus gallus. African Journal of Biotechnology, 10(48), 9969\\u0026ndash;9977.\\u003c/li\\u003e\\n\\u003cli\\u003eAl-Mansour, M.I., Jarrar, B.M. 2007. Morphological, histological and histochemical study of the lingual salivary glands of the little egret, Egretta garzetta. Saudi J. Biological Sci., 14: 75 81.\\u003c/li\\u003e\\n\\u003cli\\u003eArthitvong s.; Makmee, N. and Suprasert, A. (1999): Histochemical detection of glycoconjugates in the anterior lingual salivary glands of the domestic fowl. Kasetsart, J. (Nat. Sci.), 33: 243-250.\\u003c/li\\u003e\\n\\u003cli\\u003eSuprasert A., Arthitvong S., Koonjaenak S., (2000) Lectin histochemistry of glycoconjugates in the mandibular gland of chicken. Kasetsart J. (Nat. Sci.) 34.85-90.\\u003c/li\\u003e\\n\\u003cli\\u003eSamar ME, \\u0026Aacute;vila RE, Esteban FJ, Olmedo L, Dettin L, Massone A, et al. Histochemical and ultrastructural study of the chicken salivary palatine glands. Acta Histochem. 2002:104(2):199\\u0026ndash;207. https://www.sciencedirect.com/science/article/pii/S0065128104701166\\u003c/li\\u003e\\n\\u003cli\\u003eEl-Bakry, A.M., \\u0026amp; Iwasaki, S. (2014). Ultrastructure and histochemical study of the lingual salivary glands of some bird species. \\u003cem\\u003ePakistan Journal of Zoology, 46\\u003c/em\\u003e, 553-559.\\u003c/li\\u003e\\n\\u003cli\\u003ePearse A. G. E., 1972: Histochemistry: Theoritical and Applied. 3rd ed. J. \\u0026amp; A. Churchill, London, pp. 1303-1444.\\u003c/li\\u003e\\n\\u003cli\\u003eScott D. E., Dorling J., (1965): Differential staining of acid glycosaminoglycans (mucopolysaccharides) by Alcian blue in salt solutions. Histochemie, 5: 221-33.\\u003c/li\\u003e\\n\\u003cli\\u003eSpicer S. S., Meyer D. B., Histochemical differentiation of acid mucopolysaccharides by means of combined aldehyde fuchsin-alcian blue staining. (1960). Tech Bull Regist Med Technol Am Soc Clin Pathol Regist Med Technol.30:53-60.\\u003c/li\\u003e\\n\\u003cli\\u003eFujii, S., Tamura, T. 1966. Histochemical studies on the mucins of the chicken salivary glands. J. Fac. Fish. Anim. Husb, Hiroshima Univ., 6: 345 355.\\u003c/li\\u003e\\n\\u003cli\\u003eSamar, M.E., Avila, R.E., De Fabro, S.P., Centurion, C. 1995. Structural and cytochemical study of salivary glands in the magellanic penguin Spheniscus magellanicus and the kelp gull Larus domincanus. Marine Ornithol., 23: 154 156.\\u003c/li\\u003e\\n\\u003cli\\u003eAl-Mansour, M.I., Jarrar, B.M. (2004). Structure and secretions of the lingual salivary glands of the white- cheecked bulbul, Pycnonotus leucogenys, (Pycnontidae). Saudi. J. Sci., 11(2): 119 126.\\u003c/li\\u003e\\n\\u003cli\\u003eTaib, N.T., Jarrar, B.M. 2001. Histological characterization of the lingual salivary glands of the Eurasian collared dove, Streptopelia decaocta. Pak. J. Bio. Sci., 4(11): 1425 1428.\\u003c/li\\u003e\\n\\u003cli\\u003eKudo, K., Nishimura, S., \\u0026amp; Tabata, S. (2008). Distribution of taste buds in layer-type chickens: scanning electron microscopic observations. Animal Science Journal, 79(6), 680\\u0026ndash;685. https://doi.org/10.1111/j.1740- 0929.2008.00580.x.\\u003c/li\\u003e\\n\\u003c/ol\\u003e\"},{\"header\":\"Tables\",\"content\":\"\\u003cp\\u003eTable (1): Showing the average diameter\\u0026nbsp;(um)\\u0026nbsp;of the sublingual salivary gland openings (SGO), the taste buds openings (TBO) and taste buds surface\\u0026nbsp;area (TBA) (um\\u003csup\\u003e2\\u003c/sup\\u003e)\\u0026nbsp;in quails\\u0026nbsp;\\u003c/p\\u003e\\n\\u003ctable border=\\\"1\\\" cellspacing=\\\"0\\\" cellpadding=\\\"0\\\"\\u003e\\n \\u003ctbody\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"24.95479204339964%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eItem\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"18.44484629294756%\\\" rowspan=\\\"2\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eSGO (um) \\u0026nbsp;\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"39.059674502712475%\\\" rowspan=\\\"2\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eTBA (um\\u003csup\\u003e2\\u003c/sup\\u003e)\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"17.540687160940326%\\\" rowspan=\\\"2\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eTBO (um) \\u0026nbsp;\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"100%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eAge\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"24.95479204339964%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e8 days embryo\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"18.44484629294756%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e-\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"39.059674502712475%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eSmall sized (6\\u0026plusmn;0.01)\\u003c/p\\u003e\\n \\u003cp\\u003e\\u0026nbsp; \\u0026nbsp; Medium sized (12\\u0026plusmn;0.05)\\u003c/p\\u003e\\n \\u003cp\\u003eLarge sized (40\\u0026plusmn;0.015)\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"17.540687160940326%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e-\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"24.95479204339964%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e10 days embryo\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"18.44484629294756%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e5\\u0026plusmn;0.02\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"39.059674502712475%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eDecreased in number and some fused to be 144\\u0026plusmn;0.03\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"17.540687160940326%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e-\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"24.95479204339964%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e13 days embryo\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"18.44484629294756%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e28\\u0026plusmn;0.15\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"39.059674502712475%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e35\\u0026plusmn;0.012\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"17.540687160940326%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e-\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"24.95479204339964%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eHatching chick\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"18.44484629294756%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e30\\u0026plusmn;0.1\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"39.059674502712475%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e-\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"17.540687160940326%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e11\\u0026plusmn;0.023\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"24.95479204339964%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e7 days chick\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"18.44484629294756%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e36\\u0026plusmn;0.04\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"39.059674502712475%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e-\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"17.540687160940326%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e12\\u0026plusmn;0.1\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"24.95479204339964%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e30- \\u0026amp; 60-days \\u0026nbsp; chick\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"18.44484629294756%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e45\\u0026plusmn;0.08\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"39.059674502712475%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e\\u0026nbsp;Small sized (9\\u0026plusmn;0.06)\\u003c/p\\u003e\\n \\u003cp\\u003e\\u0026nbsp;Medium sized (12\\u0026plusmn;0.013)\\u003c/p\\u003e\\n \\u003cp\\u003e\\u0026nbsp;Large sized (32\\u0026plusmn;0.02)\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"17.540687160940326%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e8.2\\u0026plusmn;0.35\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003c/tbody\\u003e\\n\\u003c/table\\u003e\\n\\u003cp\\u003e\\u003cbr\\u003e\\u003c/p\\u003e\\n\\u003cp\\u003eTable (2): Showing the histochemical properties of the sublingual salivary gland (Gs) \\u0026amp; mandibular salivary gland (Gm) in quail chicks.\\u003c/p\\u003e\\n\\u003ctable border=\\\"1\\\" cellspacing=\\\"0\\\" cellpadding=\\\"0\\\" width=\\\"624\\\"\\u003e\\n \\u003ctbody\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"65.38461538461539%\\\" colspan=\\\"6\\\" valign=\\\"top\\\"\\u003e\\u003cstrong\\u003e(A) Sublingual salivary gland\\u003c/strong\\u003e\\u003c/td\\u003e\\n \\u003ctd width=\\\"34.61538461538461%\\\" valign=\\\"top\\\"\\u003e\\u003cstrong\\u003e(B) Mandibular salivary gland\\u003c/strong\\u003e\\u003cbr\\u003e\\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003ctr\\u003e\\n \\u003ctd width=\\\"13.804173354735152%\\\" valign=\\\"top\\\"\\u003e\\u003cbr\\u003e\\u003c/td\\u003e\\n \\u003ctd width=\\\"8.98876404494382%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003eZero day\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"8.186195826645266%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e7 days\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"9.630818619582664%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e14 days\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"9.309791332263242%\\\" valign=\\\"top\\\"\\u003e\\n 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\\u003cp\\u003e\\u003cstrong\\u003eAB/PAS\\u003c/strong\\u003e\\u003c/p\\u003e\\n \\u003cp\\u003e\\u003cstrong\\u003ePurple\\u003c/strong\\u003e\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"8.98876404494382%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e\\u003cstrong\\u003e+++\\u003c/strong\\u003e\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"8.186195826645266%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e\\u003cstrong\\u003e+++\\u003c/strong\\u003e\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"9.630818619582664%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e\\u003cstrong\\u003e++++\\u003c/strong\\u003e\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"9.309791332263242%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e\\u003cstrong\\u003e++++\\u003c/strong\\u003e\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"15.409309791332264%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e\\u003cstrong\\u003e++++\\u003c/strong\\u003eAlcinophilic (greenish-purple)\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003ctd width=\\\"34.670947030497594%\\\" valign=\\\"top\\\"\\u003e\\n \\u003cp\\u003e\\u003cstrong\\u003e++++\\u003c/strong\\u003e\\u003c/p\\u003e\\n \\u003cp\\u003eAlcinophilic\\u0026nbsp;\\u003c/p\\u003e\\n \\u003cp\\u003e(greenish-purple)\\u003c/p\\u003e\\n \\u003c/td\\u003e\\n \\u003c/tr\\u003e\\n \\u003c/tbody\\u003e\\n\\u003c/table\\u003e\"}],\"fulltextSource\":\"\",\"fullText\":\"\",\"funders\":[],\"hasAdminPriorityOnWorkflow\":false,\"hasManuscriptDocX\":true,\"hasOptedInToPreprint\":true,\"hasPassedJournalQc\":\"\",\"hasAnyPriority\":false,\"hideJournal\":false,\"highlight\":\"\",\"institution\":\"\",\"isAcceptedByJournal\":true,\"isAuthorSuppliedPdf\":false,\"isDeskRejected\":\"\",\"isHiddenFromSearch\":false,\"isInQc\":false,\"isInWorkflow\":false,\"isPdf\":false,\"isPdfUpToDate\":true,\"isWithdrawnOrRetracted\":false,\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"bmc-veterinary-research\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":false,\"externalIdentity\":\"\",\"sideBox\":\"Learn more about [BMC Veterinary Research](http://bmcvetres.biomedcentral.com/)\",\"snPcode\":\"12917\",\"submissionUrl\":\"https://submission.nature.com/new-submission/12917/3?\",\"title\":\"BMC Veterinary Research\",\"twitterHandle\":\"@BMC_series\",\"acdcEnabled\":true,\"dfaEnabled\":true,\"editorialSystem\":\"stoa\",\"reportingPortfolio\":\"BMC Series\",\"inReviewEnabled\":true,\"inReviewRevisionsEnabled\":true},\"keywords\":\"Sublingual salivary glands, mandibular gland, mucous, tubuloalveolar, Taste bud, quail\",\"lastPublishedDoi\":\"10.21203/rs.3.rs-3958585/v1\",\"lastPublishedDoiUrl\":\"https://doi.org/10.21203/rs.3.rs-3958585/v1\",\"license\":{\"name\":\"CC BY 4.0\",\"url\":\"https://creativecommons.org/licenses/by/4.0/\"},\"manuscriptAbstract\":\"\\u003cp\\u003eThe oropharyngeal floor of the lower beak in Japanese quails (\\u003cem\\u003eCoutrinx coutrinx japonica\\u003c/em\\u003e)\\u003cstrong\\u003e \\u003c/strong\\u003econtains the sublingual and mandibular salivary glands\\u003cstrong\\u003e. Although few literatures spoke about the oropharyngeal glands, our study demonstrated the best for full morphological and cytochemical illustration. The\\u003c/strong\\u003e morphological and cytochemical analysis were done on 20 healthy Japanese quail embryos with ages of the 6\\u003csup\\u003eth\\u003c/sup\\u003e, 10\\u003csup\\u003eth\\u003c/sup\\u003e, 11\\u003csup\\u003eth\\u003c/sup\\u003e, and 13\\u003csup\\u003eth\\u003c/sup\\u003e days of incubation, and 25 healthy quail chicks at ages zero (hatching day old), 7\\u003csup\\u003eth\\u003c/sup\\u003e, 14\\u003csup\\u003eth\\u003c/sup\\u003e, 30\\u003csup\\u003eth\\u003c/sup\\u003e, and 60\\u003csup\\u003eth\\u003c/sup\\u003e days old. The primordia of the sublingual and mandibular salivary glands were noticed at the 6\\u003csup\\u003eth\\u003c/sup\\u003e and 10\\u003csup\\u003eth\\u003c/sup\\u003e days of the prehatching old respectively as an epithelial bud. After hatching, both primordia were elongated and differentiated into secretory units. These glands are mucous polystomatic tubulo-alveolar paired glands which were situated in the submucosa. The sublingual glands consisted of 3-5 lobes extended from two ceratobranchial caudally by their wide ends beyond the median sulcus of the prefrenular part of sublingual space rostrally; where they opened by their constricted part. The taste buds are variable in size and position; the associated salivary glands type was the largest, which all taste pores varied from 8.2-12 um. The mandibular glands lay on the paralingual groove which arose at 10 days old embryo. Furthermore, the mandibular glands were located dorsomedial to the sublingual glands and extended longitudinally from the rostral border of the frenulum linguae to the caudal tips of sublingual glands. Notably, the taste buds decreased in the volume and number with advancing age. Both gland secretions showed various histochemical reactions that ended with highly alcinophilic (acidic) materials in advanced ages.\\u003c/p\\u003e\",\"manuscriptTitle\":\"New findings on Developmental Studies of the oropharyngeal Salivary Glands in Japanese Quails (Coutrinx coutrinx japonica)\",\"msid\":\"\",\"msnumber\":\"\",\"nonDraftVersions\":[{\"code\":1,\"date\":\"2024-02-20 17:41:21\",\"doi\":\"10.21203/rs.3.rs-3958585/v1\",\"editorialEvents\":[{\"type\":\"communityComments\",\"content\":0},{\"type\":\"decision\",\"content\":\"Revision requested\",\"date\":\"2024-05-17T01:23:55+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"editorInvitedReview\",\"content\":\"\",\"date\":\"2024-05-04T06:03:15+00:00\",\"index\":\"hide\",\"fulltext\":\"\"},{\"type\":\"editorInvitedReview\",\"content\":\"\",\"date\":\"2024-04-30T07:22:04+00:00\",\"index\":\"hide\",\"fulltext\":\"\"},{\"type\":\"reviewerAgreed\",\"content\":\"6520878388327470635397545332514101472\",\"date\":\"2024-04-26T12:38:45+00:00\",\"index\":\"hide\",\"fulltext\":\"\"},{\"type\":\"reviewerAgreed\",\"content\":\"74136541352515895004265275421960961847\",\"date\":\"2024-04-25T06:19:24+00:00\",\"index\":\"hide\",\"fulltext\":\"\"},{\"type\":\"editorInvitedReview\",\"content\":\"\",\"date\":\"2024-04-23T06:56:14+00:00\",\"index\":\"hide\",\"fulltext\":\"\"},{\"type\":\"editorInvitedReview\",\"content\":\"\",\"date\":\"2024-04-05T10:13:52+00:00\",\"index\":\"hide\",\"fulltext\":\"\"},{\"type\":\"reviewerAgreed\",\"content\":\"3bd98cee-c2e1-4378-8807-bff705eb57e0\",\"date\":\"2024-04-02T14:42:16+00:00\",\"index\":\"hide\",\"fulltext\":\"\"},{\"type\":\"reviewerAgreed\",\"content\":\"22a6b01a-203c-4cd6-ad94-a0ee2639ee28\",\"date\":\"2024-03-28T13:48:18+00:00\",\"index\":\"hide\",\"fulltext\":\"\"},{\"type\":\"reviewersInvited\",\"content\":\"\",\"date\":\"2024-03-28T13:29:44+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"editorInvited\",\"content\":\"\",\"date\":\"2024-02-19T11:42:46+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"editorAssigned\",\"content\":\"\",\"date\":\"2024-02-19T11:14:19+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"checksComplete\",\"content\":\"\",\"date\":\"2024-02-15T14:11:23+00:00\",\"index\":\"\",\"fulltext\":\"\"},{\"type\":\"submitted\",\"content\":\"BMC Veterinary Research\",\"date\":\"2024-02-15T12:01:13+00:00\",\"index\":\"\",\"fulltext\":\"\"}],\"status\":\"published\",\"journal\":{\"display\":true,\"email\":\"info@researchsquare.com\",\"identity\":\"bmc-veterinary-research\",\"isNatureJournal\":false,\"hasQc\":true,\"allowDirectSubmit\":false,\"externalIdentity\":\"\",\"sideBox\":\"Learn more about [BMC Veterinary Research](http://bmcvetres.biomedcentral.com/)\",\"snPcode\":\"12917\",\"submissionUrl\":\"https://submission.nature.com/new-submission/12917/3?\",\"title\":\"BMC Veterinary Research\",\"twitterHandle\":\"@BMC_series\",\"acdcEnabled\":true,\"dfaEnabled\":true,\"editorialSystem\":\"stoa\",\"reportingPortfolio\":\"BMC Series\",\"inReviewEnabled\":true,\"inReviewRevisionsEnabled\":true}}],\"origin\":\"\",\"ownerIdentity\":\"56140412-a34f-472c-993d-7112b1cc886b\",\"owner\":[],\"postedDate\":\"February 20th, 2024\",\"published\":true,\"recentEditorialEvents\":[],\"rejectedJournal\":[],\"revision\":\"\",\"amendment\":\"\",\"status\":\"published-in-journal\",\"subjectAreas\":[],\"tags\":[],\"updatedAt\":\"2024-11-18T19:24:36+00:00\",\"versionOfRecord\":{\"articleIdentity\":\"rs-3958585\",\"link\":\"https://doi.org/10.1186/s12917-024-04355-7\",\"journal\":{\"identity\":\"bmc-veterinary-research\",\"isVorOnly\":false,\"title\":\"BMC Veterinary Research\"},\"publishedOn\":\"2024-11-12 15:57:35\",\"publishedOnDateReadable\":\"November 12th, 2024\"},\"versionCreatedAt\":\"2024-02-20 17:41:21\",\"video\":\"\",\"vorDoi\":\"10.1186/s12917-024-04355-7\",\"vorDoiUrl\":\"https://doi.org/10.1186/s12917-024-04355-7\",\"workflowStages\":[]},\"version\":\"v1\",\"identity\":\"rs-3958585\",\"journalConfig\":\"researchsquare\"},\"__N_SSP\":true},\"page\":\"/article/[identity]/[[...version]]\",\"query\":{\"redirect\":\"/article/rs-3958585\",\"identity\":\"rs-3958585\",\"version\":[\"v1\"]},\"buildId\":\"qtupq5eGEP_6zYnWcrvyt\",\"isFallback\":false,\"isExperimentalCompile\":false,\"dynamicIds\":[84888],\"gssp\":true,\"scriptLoader\":[]}","source_license":"CC-BY-4.0","license_restricted":false}