The intestine of poultry plays a significant role in the health and production through enzymatic and microbial digestion of feed as well as absorption of dietary nutrients. The current study was designed to explore the gross and histological alterations in the broiler duodenum and cecum triggered by dietary dexamethasone (DEX). The study was conducted on four homogenous groups of one-day-old (20 chicks/group) i.e. one control Non-DEX and three treatment groups (DEX-1, DEX-2, and DEX-3). The broilers were fed commercial broiler feed containing DEX at the rate of 0, 3, 5, and 7mg/kg feed in the Non-DEX, DEX-1, DEX-2, and DEX-3 groups respectively. The gross morphologic and morphometric data were recorded immediately after the collection of samples on days 7, 14, 21, and 28. Then, the tissue samples were processed for histological investigation. In the gross morphometric study, the weight, length, and width of the intestine were found significantly less in the DEX groups. Histopathological study results showed degeneration of intestinal glands (duodenum), mucosa, and lymphatic nodules with loss of lymphatic nodules (cecum). The percentage of the degenerated nodule was also decreased. The length, width, and surface area of the duodenal villi, thickness of the mucosal layer of the cecum, and diameter of the cecal lymphatic nodules were significantly decreased in all the DEX groups. The magnitude of the alterations was associated with both the dose and duration of DEX treatment. However, the current study results imply that DEX treatment significantly alters the morphologic and morphometric characteristics of the broiler intestine.
Research Article
Evaluation of the effects of steroid growth promoter on the intestinal morphology and morphometry in broiler chicken.
https://doi.org/10.21203/rs.3.rs-1358412/v1
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Broiler
Duodenum
Cecum
Dexamethasone
Morphology
Morphometry
Poultry meat is a popular source of protein in the human diet which has made the poultry sector economically valuable over the last few decades. In 2020, the poultry sector predicted a global production of 130 million tons of meat which will make it the most consumed animal meat worldwide (Borda-Molina et al., 2018). In comparison to the traditional husbandry techniques, modern poultry strains are genetically modified for rapid growth in a shorter period (Schmidt et al., 2009). The available strains of broilers such as Cobb-500, cobb-700, arbor acress, lohman meat, etc. are the outcome of genetic modification (Nasrin et al., 2012). In this context, understanding the role of the gastrointestinal tract is crucial, as it may have an impact on the growth performance and health of broiler.
In the field of veterinary research, broiler “Gut health” is a widely discussed topic due to its role in health and growth performance. Duodenum and cecum are the major gastrointestinal parts for the digestion and absorption of nutrients from the diet. The duodenum receives the digestive enzymes and bicarbonates produced by the pancreas as well as the bile produced by the liver, which plays a major role in the digestion of protein and fat. The cecum plays a significant role in its growth as it harbors a diversified bacterial community and can ferment a wide variety of feed ingredients like starch, cellulose, hemicellulose, polysaccharides, fructo-oligosaccharides, and lignin to different final products (Clench and Mathias, 1995 and Svihus, 2014). Hence, the microbial community of the cecum also plays an important role in the fermentation of carbohydrates (Lovegrove et al., 2017). The undigested compounds which are not absorbed in the small intestine, finally end up in the cecum, where they are digested by an anaerobic fermentation process (de Carvalho et al., 2020). As the proximal region of the cecum is continuously exposed to extra cecal bacterial or other antigenic invasions, the lymphatic nodules of the cecum play a critical function against these foreign invaders (Majeed et al., 2009). Thus the cecum impedes the colonization of pathogenic microorganisms and detoxifies harmful elements to maintain its optimum digestive and absorptive functions (Tan et al., 2019). So, the importance of the duodenum and cecum in broiler growth and development is indisputable.
A wide variety of growth promoters are used in the broiler industry as feed additives to improve the genetic potentiality, feed efficiency, and ultimately, the growth rate of broilers. Antibiotics and anabolic steroids such as steroid hormones, resorcylic acid lactones, stilbenes, antithyroid agents, b-agonists, and glucocorticoids (dexamethasone and corticosterone) are commonly used as growth promoters (Hussein and Khalil, 2013; Toldra and Reig, 2016). Supplying antibiotics as feed additives to poultry has long been practiced for its beneficial effects on growth rate (Castanon, 2007). However, the use of these drugs as growth promoters has unintended consequences like antibiotic resistance, accumulation of residues in the edible tissues, environmental contamination, etc. Though the efficacy of glucocorticoids is irrefutable, they also possess several side effects which are associated with dose and duration of administration (Ramamoorthy and Cidlowski, 2016).
According to the previous study reports, supplementation of steroid growth promoters like DEX with diet do not improve the growth rate of broiler (Afrose et al., 2018; Ademu et al., 2018). Besides inducing stress, corticosteroids can also influence digestive function by markedly decreasing the digestibility of protein and carbohydrates (Puvadolpirod and Thaxton, 2000). DEX reportedly damages the kidney by vacuolation of kidney tissue, shrinking the tubules, and decreasing the number of nephrons (Moisiadis et al., 2014; Salman and Hassooni, 2020 and Sultana et al., 2021). A high dose of DEX alters the breast and thigh meat morphology (Akter et al., 2020) and also results in the developmental arrest of immune organs in the broiler (Sultana et al., 2020a). DEX also alters the liver morphology according to the previous report (Sultana et al., 2020b). The effects of prednisolone related to gastrointestinal motility, intestinal histology, and mucosal mast cells were studied in rats (Lima et al., 2017).
The enteric system is a major body system that is responsible for the digestion and absorption of dietary nutrients. So, any alteration in its morphology affects the functions of other organs or body systems. Nonetheless, the adverse effects of DEX on morphology and morphometry of the enteric systems of broiler are not yet well-documented. So, we designed the current study to investigate the effects of different doses of DEX on the gross morphology and morphometry (i.e. color, weight, length, and width) as well as the histomorphology and histomorphometry (i.e. Villi length, width and villi surface area of duodenum, mucosal height, the greater and lesser diameter of lymphatic nodules of cecum) of broiler duodenum and cecum.
2.1. Experimental animals
In this research, 80 one-day-old broiler chicks of Cobb-500 strain were purchased from a local commercial hatchery (Provita Hatchery Limited, Mymensingh, Bangladesh) and randomly housed in separate cages. The chicks were clinically healthy and free from any kind of visible deformity.
2.2. Housing and feeding management
The DOCs were then randomly assigned to four groups i.e. One control (Non-DEX) and three experimental or treated groups as DEX-1, DEX-2, and DEX-3. The DOCs were fed (ad libitum) commercial broiler feed (Nourish Poultry and Hatchery Ltd., Bangladesh) containing corticosteroid (Dexamethasone, BP 0.5 mg, Opsonin Limited) at the rate of 3mg/kg, 5mg/kg, 7mg/kg in group DEX-1, DEX-2 and DEX-3 respectively. Starter and grower feed composition (Sultana et al., 2020a) and pellet size were chosen to meet the recommended nutrient requirements. A constant supply of an adequate amount of feed and fresh drinking water was ensured. Standard temperature and relative humidity were maintained throughout the experiment (Sultana et al., 2021).
2.3. Gross morphology and morphometry
Samples were collected on 7, 14, 21, and 28 days of the experiment. Five birds from each group were sacrificed manually on each sampling day by the cervical subluxation method. The duodenum and right cecum were dissected immediately. Color, weight, length, and width of the cecum were considered for the gross morphologic and morphometric study. The color of the duodenum and cecum was compared between the Non-DEX and DEX groups by visual inspection. Weight (gm) was measured using a high precision balance (FGH Series, AND Company Limited, Korea), and length, the width was measured by a graded scale (cm). The width of the cecum was measured from the middle portion of the blind sac.
2.4. Histopathology
About one cm of the duodenal and cecal segment (from the blind sac) was collected and fixed in 10% buffered formalin and then routinely processed into slides with Hematoxylin and Eosin (H&E) stain according to the protocol described by Sultana et al., 2020b. All the stained tissue sections were examined and analyzed blindly to avoid any biases. The histomorphological attributes of the duodenal and cecal tissues were studied under a light microscope (Leica DMR, Leica Microsystems, Wetzlar, Germany) at 100X and 400X magnification. Length (measurement was taken from the tips of the villi to the villi-crypt junctions) and width (measured from the mid-region of the villi) of duodenal villi, mucosal thickness, and diameter of lymphatic nodules of cecum were measured in micrometer (µm) using a calibrated stage micrometer. The surface area of the villi was calculated using the following formula-
, where π = 3:1416 (Moghaddam and Alizadeh-Ghamsari, 2013). The number of intact and degenerated lymphatic nodules in the cecum was counted and the percentage was calculated. Measurement of all the variables was done from 10 randomly selected focuses at 100X magnification.
Necessary photographs were captured from ten randomly selected focuses at 100X and 400X magnifications for better illustration of the obtained results. All the images were captured by photomicroscope (Model: CX41U-LH50HG, Olympus Corporation, Tokyo, Japan).
2.5. Statistical analyses:
All the data obtained in this study were analyzed using IBM SPSS Statistics 22. Differences among the groups of birds were compared using one-way ANOVA with posthoc Duncan’s multiple range test where P<0.05 was considered significant. In all trials, data were expressed as mean ± standard error of the mean (SEM).
3.1. Gross morphologic and morphometric profile of duodenum
The gross attributes and alterations of the duodenum were shown in figure 1. The duodenum of the Non-DEX group appeared light greyish color whereas the DEX groups revealed slightly darker color with a reddish tinge.
The gross morphometric parameters of the duodenum were shown in figure 2. The weight of the duodenum was significantly (P<0.05) less in the DEX treated groups compared to the Non-DEX group. Among the DEX groups, the highest weight was found in the DEX-1 group and the lowest weight was found in the DEX-3 group on all days of the experiment. Similarly, the length and width were also significantly (P<0.05) less in the DEX groups. On day 7, the length was found significantly (P<0.05) less in the DEX-2 and DEX-3 groups compared to the Non-DEX group. The highest length was found in the DEX-1 group and the lowest length was seen in the DEX-3 group. On day 28, there was no significant (P>0.05) difference in length between the DEX groups. There was a significant (P<0.05) difference in width on day 7 among the DEX groups. However, a significant (P<0.05) difference in width was found from days 14 to 28 among the DEX groups.
3.2. Histopathological profile of duodenum
The histo-architecture of the duodenum of Non-DEX and DEX groups was shown in figure 3. The Non-DEX group revealed general histological architecture. However, the DEX groups showed marked microscopic alterations, especially in the duodenal glands (crypt of Lieberkuhn) and villi. In the DEX-1 group, the intestinal glands showed no alterations though they started to degenerate in the DEX-2 group. In the DEX-3 group, almost all the intestinal glands were degenerated, resulting in the reduction of the intestinal gland population.
The histomorphometric data of duodenum were shown in table 1. The histomorphometric study revealed that both the length and width of duodenal villi were significantly (P<0.05) less in the DEX groups as compared to the Non-DEX group. However, among the DEX groups, the highest length and width of duodenal villi was found in the DEX-1 group and the lowest value was found in the DEX-3 group. The surface area of the villi was also significantly (P<0.05) decreased in the DEX groups which were also dose-dependent.
3.3. Gross morphologic and morphometric profile of cecum
The gross attributes and alterations of the cecum were shown in figure 4. The cecum was a paired organ with a lower diameter at the origin but gradually increased in size until it formed a blind end terminally. The gross morphometry distinguished three distinct regions in the cecum. The cecum of the Non-DEX group was greenish-grey in color whereas the cecum of the DEX groups was greyish color with a slight reddish tinge.
The weight of the left cecum in an individual group of broilers at different days of the experiment was shown in Figure 5. The weight was significantly (P<0.05) less in the DEX groups as compared to the Non-DEX group. Significant (P<0.05) differences among the DEX groups were seen only at day 28 where the DEX-2 group was significantly (P<0.05) different from the DEX-1 and DEX-3 groups. The length and width of the left cecum in the individual group of broilers at different days of the experiment were shown in Fig. 3. Both the length and width were significantly (P<0.05) less in the DEX groups as compared to the Non-DEX group. There were also significant (P<0.05) differences among the DEX groups.
3.4. Histopathological profile of cecum
The histo-architecture of the cecum of Non-DEX and DEX groups was shown in figure 6. The Non-DEX group revealed general histological architecture. On the other hand, marked disruption of mucosal epithelial cells was seen in all the DEX groups. Mucosal degeneration was also seen which was more extensive in the high dose groups. Degeneration of mucosa started from the tip of the mucosal layer to the base. The lymphatic nodules showed degenerative changes in the treatment groups from day 7 which increased afterward. Degenerative changes were increased significantly in the high-dose groups. On day 28, the histological architecture was almost completely lost in the DEX-3 group. The thickness of the muscular layer also decreased in the DEX treated broilers. Some extent of muscular degeneration was also seen.
The histomorphometric data of mucosal height, the greater and lesser diameter of lymphatic nodules were shown in table 2. Mucosal height was significantly (P<0.05) less in the DEX groups as compared to the Non-DEX group. Both the greater and lesser diameter of the lymphatic nodules was also significantly (P<0.05) less in the DEX treated broilers. There were also significant (P<0.05) differences among the DEX groups as the values were decreased gradually with the increased dose of DEX.
The percentage of degenerated lymphatic nodules was shown in figure 7. The percentage was increased with the progression of both age and DEX dose. The maximum percentage (92.31%) of degenerated lymphatic nodules was seen on day 28 in the DEX-3 group.
The morphology and morphometric attributes of the intestine possess a significant role in maintaining gut health, nutrient digestibility, and ultimately in the growth performance of the broiler. The duodenum of poultry contributes to the protein and fat digestion whereas the cecum plays a significant role in preventing the colonization of pathogens, detoxifying substances detrimental to gut health, processing nutrients from ingested feed, and harvesting of the ingestion by harboring complex and dynamic microbial communities (Tan et al., 2019). Height and width of villi, the thickness of the mucosa, crypt depth, etc. are commonly measured to assess the gut health and digestibility (Montagne et al., 2003, Berrocoso et al., 2017) So, the present study aimed to investigate the effects of different doses of dietary GC, DEX on the gross morphology and morphometry (i.e. color, weight, length, and width) as well as the histomorphology and histomorphometry (i.e. mucosal thickness, the greater and lesser diameter of lymphatic nodules) of broiler duodenum and cecum. Our results indicate that dietary DEX altered the morphological and morphometric characteristics of the broiler intestine.
4.1. Gross morphology and morphometry
In the current study, the color of the intestine was almost similar in the Non-DEX and DEX groups which were greenish-grey to reddish-grey in color. These findings are in accordance with the findings described by Getty (1975). The findings of gross morphometric measurements of the Non-DEX group at different days of the experiment are also almost similar to the results of Nasrin et al., (2012). However, in the DEX groups, all the gross morphometric parameters were decreased significantly compared to the control. Intestinal sizes are closely related to body sizes (Clench and Mathias, 1995). Dietary DEX reduces the growth performance and weight gain in the broiler (Ademu et al., 2018). The previous study reports also suggest that corticosteroids like DEX negatively affect organ weight gain and reduce organ size (Vahdatpour, 2009 and Tschanz et al., 2003). However, the effects of DEX on intestinal volume and weight gain had not been documented previously. Feed efficiency and the growth of broilers greatly depend on the enzymatic activities in the duodenum and the diversified microbial community that inhabits the caeca. Dietary lipid particles are mostly digested and emulsified in the duodenum by pancreatic enzymes and hepatic bile acids (Bauer et al., 2005). Nutrients, such as undigested starch, protein, and fiber enter into the caeca bypassing the small intestine (Svihus et al., 2013). The blind end of the cecum helps to retain the digest for longer periods and thus enhances nutrient absorption (Clench and Mathias, 1995). So, the reduced intestinal size may adversely affect the overall growth rate of the broiler.
4.2. Histopathological profile of duodenum and cecum
In the histomorphological study, the duodenum and cecum of the Non-DEX groups at different days of the experiment revealed general histological characteristics as described in the previous studies (Majeed et al., 2009 and Nasrin et al., 2012). On the other hand, significant alterations were seen in the DEX groups. The length of the villi was found significantly less in the DEX groups which match the findings of the previous study report (Carvalho et al., 2018). The length of the villi is one of the prominent indices of the intestine's capacity to absorb nutrition (Li et al., 2008). The decrease of the surface area of the villi might also explain the lower weights of the duodenum and cecum due to reduced nutrient absorption. The crypts of Liberkuhn lie between the intestinal villi and assist in the protein digestion as well as protection of the host from enteric pathogens (Clevers and Bevins, 2013). Crypt depth is considered an indicator of intestinal epithelium maturation (Li et al., 2008). In the current study, the crypts have almost degenerated in the higher dose group which may affect the overall gut health and performance.
Marked disruption of the mucosal surface epithelial cells along with mucosal degeneration was seen in the cecum of DEX treated broilers. Continuity of the intestinal lining epithelium is crucial for maintaining intestinal permeability which is mainly regulated by tight junction distribution and integrity. The epithelial cells overlying the mucosa are derived from progenitors like stem cells residing within the crypt (Barker et al., 2007). The undifferentiated epithelial cells exit the base of the crypt and migrate luminally and mature into highly specialized absorptive enterocytes (Edelblum and Turner, 2015). If the mucosal epithelial cells get damaged or destroyed due to adverse conditions in the intestine, the stem cells located in the crypt repair the epithelial layer by reproducing new undifferentiated epithelial cells (Shen, 2009). So, in case of degeneration or loss of crypt, the damaged villous epithelial layer can’t regenerate which may greatly alter the intestinal functionality. Mucosal morphological features correspond with increased feed efficiency and growth rate in broiler as the alterations in the mucosal layer is related to intestinal function (Yamauchi, 2002). Histomorphometric investigation showed that the mucosal height declined significantly in the treated broilers. This finding is similar to the results reported by Lima et al., 2017 and Berenjian et al., 2020. The major role of the caeca is to separate the intestinal contents into a nutrient-rich fluid fraction that enters the caeca for digestion and absorption (King and McLelland 1984). The villi of the mucosa extend into the lumen of the intestine and thus increase the absorptive surface area (Shen, 2009). So, the decrease of the mucosal surface area will lead to the reduction of absorptive surface area in the cecum which may negatively affect the feed efficiency and ultimately the growth rate and health of the broiler.
Besides these, the degeneration and loss of lymphatic nodules were also seen in the current study. Lymphatic nodules are among the gut-associated lymphoid tissues in poultry, which is one of the main components of the lateral immune system. These gut-associated lymphoid tissues react with the gut microflora and play a pivotal role in controlling the incidence of poultry enteric disorders through its immunological functions (Callaway et al., 2008). Hence, the structural integrity of lymphatic nodules is crucial for proper functioning and to achieve optimal production performance through digestion, absorption of nutrients, and immunity.
The thickness of the muscularis layer was also seen to be decreased in the treated broilers with scattered degenerative changes. DEX decreases protein synthesis, increases protein catabolism and proteolytic activities in the muscularis layer, and thus leads to muscular atrophy (Schakman et al., 2013). However, the exact mechanism of the muscle degeneration and reduction of the thickness of the muscularis layer of the broiler cecum is ambiguous.
Looking at an overview of the obtained results from the current study, it is evident that DEX treatment affects the gross and histological morphology and morphometry of the broiler intestine. The adverse effects of DEX on the intestinal mucosa, glands, and lymphatic nodules may lead to impaired digestion of food particles, reduced absorption of nutrients, and diminished gut immunity which may affect the health and production of broiler. However, further study is recommended to investigate the effect of DEX on the enzymatic activity in the duodenum as well as the microbial population in the broiler cecum.
Acknowledgments
We appreciate the research assistance from the Department of Parasitology, Faculty of veterinary science, Bangladesh Agricultural University, Bangladesh. We also acknowledge the technical support from the Department of Anatomy and Histology, Faculty of Veterinary Science, Bangladesh Agricultural University, Bangladesh.
Author contributions
Nasrin Sultana conceptualized and supervised the experiment. Rafiqul Islam performed the experiment. Rekha Rani Das assisted in the sample collection and gross data recording. Anisuzzaman provided research assistance for histomorphometric data collection. Sonali Bhakta and Rafiqul Islam analyzed the data and interpreted the results. Nasrin Sultana and Rafiqul Islam drafted the manuscript. Mohammad Rafiqul Islam and Ziaul Haque critically revised and edited the manuscript. All authors gave final approval and agreed to be accountable for all aspects of work in ensuring that questions relating to the accuracy or integrity of any part of the work.
Funding
This work was supported by the Bangladesh Agricultural University Research System (BAURES, Grant number-2020/57/BAU), Bangladesh Agricultural University, Bangladesh.
Ethical approval
This experiment was performed based on approval by Animal Welfare and Experimentation Ethics Committee, Bangladesh Agricultural University (BAU), Bangladesh [AWEEC/BAU/2021(3)].
Conflict of Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Data availability
Data will be made available on reasonable request.
- Ademu, L.A., Erakpatobor-Iyeghe, G.T., Barje, P.P., Daudu, O.M., and Wafar, R.J., 2018. Response of broiler chickens under dexamethasone induced stress conditions. Asian Journal of Research in Animal and Veterinary Sciences, 1, 1-9. https://doi.org/ 10.9734/AJRAVS/2018/39782
- Afrose, M., Sultana, N., Islam, M.R., 2018. Physiological responses of corticosteroid, dexamethasone in broiler chicken. International Journal of Scientific Research, 7, 1459-1463. https://doi.org/ 10.14202/vetworld.2020.2330-2337
- Barker, N., Van Es, J.H., Kuipers, J., Kujala, P., Van Den Born, M., Cozijnsen, M., Haegebarth, A., Korving, J., Begthel, H., Peters, P.J., Clevers, H., 2007. Identification of stem cells in small intestine and colon by marker gene Lgr5. Nature. 449, 1003-1007. https://doi.org/10.1038/nature06196
- Bauer, E., Jakob, S., and Mosenthin, R., 2005. Principles of physiology of lipid digestion. Asian-Australasian Journal of Animal Sciences, 18, 282-295. https://doi.org/10.5713/ajas.2005.282
- Berenjian, A., Sharifi, S.D., Mohammadi-Sangcheshmeh, A., and Bakhtiarizadeh, M.R., 2021. Omega-3 fatty acids reduce the negative effects of dexamethasone-induced physiological stress in laying hens by acting through the nutrient digestibility and gut morphometry. Poultry Science, 100, 100889. https://doi.org/10.1016/j.psj.2020.12.002
- Berrocoso, J. D., Kida, R., Singh, A. K., Kim, Y. S., and Jha, R., 2017. Effect of in ovo injection of raffinose on growth performance and gut health parameters of broiler chicken. Poultry science, 96, 1573-1580. https://doi.org/10.3382/ps/pew430
- Borda-Molina, D., Seifert, J., and Camarinha-Silva, A., 2018. Current Perspectives of the Chicken Gastrointestinal Tract and Its Microbiome. Computational and Structural Biotechnology Journal, 16, 131–139. https://doi.org/10.1016/j.csbj.2018.03.002
- Callaway, T.R., Edrington, T.S., Anderson, R.C., Harvey, R.B., Genovese, K.J., Kennedy, C.N., Venn, D.W., Nisbet, D.J., 2008. Probiotics, prebiotics and competitive exclusion for prophylaxis against bacterial disease. Animal Health and Research Reviews, 9, 217-225. https://doi.org/10.1017/S1466252308001540
- Carvalho, D., Herpich, J.I., Chitolina, G.Z., Gava, M.S., de Moraes, L.B., Furian, T.Q., Borges, K.A., Fortes, F.B.B., de Souza Moraes, H.L. and Salle, C.T.P., 2018. Characterization of immune and enteric systems of broilers after imunosupression with dexamethasone. Acta Scientiae Veterinariae, 46, 1606.
- Castanon, J.I.R., 2007. History of the use of antibiotic as growth promoters in European poultry feeds. Poultry science, 86, 2466-2471. https://doi.org/10.3382/ps.2007-00249
- Clench, M.H., and Mathias, J.R., 1995. The avian cecum: a review. Wilson Bulletin, 107, 93-121. https://www.jstor.org/stable/i393596
- Clevers, H.C., and Bevins, C.L., 2013. Paneth cells: maestros of the small intestinal crypts. Annual review of physiology, 75, 289-311. https://doi.org/10.1146/annurev-physiol-030212-183744
- de Carvalho, N.M., Oliveira, D.L., Saleh, M.A.D., Pintado, M.E., and Madureira, A.R., 2020. Importance of gastrointestinal in vitro models for the poultry industry and feed formulations. Animal Feed Science and Technology, 271, 114730. https://doi.org/10.1016/j.anifeedsci.2020.114730
- Edelblum, K.L., and Turner, J.R., 2015. Epithelial Cells: Structure, Transport, and Barrier Function. Mucosal Immunology, 1, 187-210. http://dx.doi.org/10.1016/B978-0-12-415847-4.00012-4
- Getty, R., 1975. Sisson and Grossman's the Anatomy of the Domestic Animals. Fifth ed. W.B. Saunders Company, London.
- Hussein, M.A., and Khalil, S., 2013. Screening of some antibiotics and anabolic steroids residues in broiler fillet marketed in El-Sharkia governorate. Life Science Journal, 10, 2111-2118.
- King, A.S., and McLelland, J., 1984. Birds, their structure and function. Second ed. Bailliere Tindall, 1 St. Annes Road, pp. 131-136.
- Li Y., Cai H.Y., Liu G.H., Dong X.L., Chang W.H., Zhang S., Zheng A.J. and Chen G.L., 2008. Effects of stress simulated by dexamethasone on jejunal glucose transport in broilers. Poultry Science, 88, 330-337.
- Lima, M.B.D., Gama, L.A., Hauschildt, A.T., Dall’Agnol, D.J.R., Corá, L.A., and Americo, M.F., 2017. Gastrointestinal motility, mucosal mast cell, and intestinal histology in rats: effect of prednisone. BioMed Research International, 2017, 1-8. https://doi.org/10.1155/2017/4637621
- Lovegrove, A., Edwards, C.H., De Noni, I., Patel, H., El, S.N., Grassby, T., Zielke, C., Ulmius, M., Nilsson, L., and Butterworth, P.J., 2017. Role of polysaccharides in food, digestion, and health. Critical Reviews in Food Science and Nutrition, 57, 237–253. http://dx.doi.org/10.1080/10408398.2014.939263
- Majeed, M.F., Al-Asadi, F.S., Nassir, A.A., and Rahi, E.H., 2009. The morphological and histological study of the cecum in broiler chicken. Basrah Journal of Veterinary research, 8, 19-25. https://doi.org/10.33762/bvetr.2009.55203
- Moghaddam, H.N, and Alizadeh-Ghamsari, A.H., 2013. Improved performance and small intestinal development of broiler chickens by dietary L-glutamine supplementation. Journal of Applied Animal Research, 41, 1-7. https://doi.org/10.1080/09712119.2012.738214
- Moisiadis, V.G., and Matthews, S.G., 2014. Glucocorticoids and fetal programming part 2: mechanisms. Nature Reviews Endocrinology, 10, 403-411. https://doi.org/10.1038/nrendo.2014.74
- Montagne, L., Pluske, J. R., and Hampson, D.J., 2003. A review of interactions between dietary fibre and the intestinal mucosa, and their consequences on digestive health in young non-ruminant animals. Animal feed science and technology, 108, 95-117. https://doi.org/10.1016/S0377-8401(03)00163-9.
- Nasrin, M., Siddiqi, M.N.H., Masum, M.A., and Wares, M.A., 2012. Gross and histological studies of digestive tract of broilers during postnatal growth and development. Journal of Bangladesh Agricultural University, 10, 69-77. https://doi.org/10.3329/jbau.v10i1.12096
- Puvadolpirod, S., and Thaxton, J.P., 2000. Model of physiological stress in chickens 4. Digestion and metabolism. Poultry Science, 79, 383-390. https://doi.org/10.1093/ps/79.3.383
- Ramamoorthy, S., and Cidlowski, J.A., 2016. Corticosteroids: mechanisms of action in health and disease. Rheumatic Disease Clinics of North America, 42, 15-31. https://doi.org/10.1016/j.rdc.2015.08.002.
- Salman, R.J., and Hassooni, Z.A., 2020. Morphological and Histological effect induced by each of Cefotaxime, Dexamethason and mixture of them to the stomach, liver, kidney and lung in rats. Drug Invention Today, 11, 3313.
- Schakman, O., Kalista, S., Barbé, C., Loumaye, A., and Thissen, J. P., 2013. Glucocorticoid-induced skeletal muscle atrophy. International Journal of Biochemistry & Cell Biology, 45, 2163-2172. https://doi.org/10.1016/j.biocel.2013.05.036.
- Schmidt, C.J., Persia, M.E., Feierstein, E., Kingham, B., and Saylor, W.W., 2009. Comparison of a modern broiler line and a heritage line unselected since the 1950s. Poultry Science, 88, 2610–2619. https://doi.org/10.3382/ps.2009-00055
- Shen, L., 2009. Functional morphology of the gastrointestinal tract. Molecular Mechanisms of Bacterial Infection via the Gut, 337, 1-35. https://doi.org/10.1007/978-3-642-01846-6_1
- Akter, F., Sultana, N., Afrose, M., Kabir, A., Islam, R., Sikder, M.H., 2021. Adaptations of muscular biology in response to potential glucocorticoid treatment in broiler chicken. Journal of Advanced Biotechnology and Experimental Therapeutics, 4, 1-8. https://doi.org/10.5455/jabet.2021.d100
- Sultana, N., Islam, R., Akter, A., Ayman, U., Bhakta, S., Rony, S.A., Nahar, A. and Alam, R., 2021. Biochemical and morphological attributes of broiler kidney in response to dietary glucocorticoid, dexamethasone. Saudi journal of biological sciences, 28, 6721-6729. https://doi.org/10.1016/j.sjbs.2021.07.047
- Sultana, N, Afrose, M., and Rafiq, K., 2020b. Effects of steroid growth promoter on morphological and biochemical adaptations in liver of broiler. Vet. World. 13, 2330-2337. https://doi.org/10.14202/vetworld.2020.2330-2337
- Sultana, N., Amin, T., Afrose, M., Rony, S.A., and Rafiq, K., 2020a. Effects of Dexamethasone on the Morphometry and Biometry of Immune Organs of Broiler Chicken. International Journal Morphology, 38, 1032-1038. https://doi.org/10.4067/S0717-95022020000401032
- Svihus, B., 2014. Function of the digestive system. Journal of Applied Poultry Research, 23, 306–314. https://doi.org/10.3382/japr.2014-00937
- Svihus, B., Choct, M., Classen, H.L., 2013. Function and nutritional roles of the avian caeca: a review. World's Poultry Science Journal, 69, 249-264. https://doi.org/10.1017/S0043933913000287
- Tan, Z., Luo, L., Wang, X., Wen, Q., Zhou, L., and Wu, K., 2019. Characterization of the cecal microbiome composition of Wenchang chickens before and after fattening. PloS One. 14, e0225692. https://doi.org/10.1371/journal.pone.0225692
- Toldrá, F., and Reig, M., 2016. Growth Promoters: Characteristics and Determination. Encyclopedia of Food and Health. 266–269. https://doi.org/10.1016/b978-0-12-384947-2.00362-7
- Tschanz, S.A., Makanya, A.N., Haenni, B., and Burri, P.H., 2003. Effects of neonatal high-dose short-term glucocorticoid treatment on the lung: a morphologic and morphometric study in the rat. Pediatric Research, 53, 72-80. https://doi.org/10.1203/01.PDR.0000041513.93422.C8
- Vahdatpour, T., Nazer Adl, K., Ebrahim Nezhad, Y., Mahery Sis, N., Riyazi, S.R., and Vahdatpour, S., 2009. Effects of corticosterone intake as stress-alternative hormone on broiler chickens: performance and blood parameters. Asian Journal of Animal and Veterinary Advances, 4, 16-21. https://doi.org/10.3923/ajava.2009.16.21
- Yamauchi, K.E., 2002. Review on chicken intestinal villus histological alterations related with intestinal function. Journal of Poultry Science, 39, 229-242. https://doi.org/10.2141/jpsa.39.229
Table 1 Histomorphometric data on duodenal villi length, width and surface area in Non-DEX and DEX groups at different days of the experiment
|
Parameters |
Groups |
Day 7 |
Day 14 |
Day 21 |
Day 28 |
|
Villi length (µm) |
Non-DEX |
799.30 ± 10.35a |
961.80 ± 15.52a |
1082.25 ± 12.43a |
1211.85 ± 14.19a |
|
DEX-1 |
624.20 ± 11.99b |
668.50 ± 11.67b |
692.75 ± 9.43b |
747.30 ± 8.49b |
|
|
DEX-2 |
609.10 ± 10.65b |
637.45 ± 10.79bc |
678.30 ± 10.14b |
712.35 ± 7.66c |
|
|
DEX-3 |
596.60 ± 14.64b |
624.70 ± 7.88c |
648.95 ± 7.55c |
656.60 ± 10.20d |
|
|
Villi width (µm) |
Non-DEX |
128.90 ± 4.96a |
162.75 ± 6.49a |
196.15 ± 6.47a |
212.30 ± 6.91a |
|
DEX-1 |
112.35 ± 4.12b |
126.90 ± 4.65b |
146.45 ± 5.41b |
172.20 ± 6.02b |
|
|
DEX-2 |
107.7 ± 5.07b |
122.75 ± 5.72bc |
138.60 ± 4.30b |
159.10 ± 4.58b |
|
|
DEX-3 |
92.85 ± 3.68c |
107.85 ± 4.51c |
119.10 ± 4.63c |
133.25 ± 4.98c |
|
|
Villi surface area (×10-3 , µm2) |
Non-DEX |
322.83 ± 10.62a |
492.48 ± 23.63a |
667.24 ± 25.86a |
807.59 ± 25.71a |
|
DEX-1 |
220.15 ± 8.91b |
266.82 ± 11.31b |
319.37 ± 14.14b |
404.55 ± 15.38b |
|
|
DEX-2 |
206.17 ± 10.39b |
246.11 ± 12.78b |
294.76 ± 7.88b |
356.60 ± 12.69c |
|
|
DEX-3 |
174.03 ± 8.01c |
211.22 ± 7.83c |
242.17 ± 7.97c |
274.67 ± 10.57d |
a,b,c,d Values in the same column with different alphabetic superscripts are significantly different from each other (P < 0.05)
Table 2 Histomorphometric data on mucosal height and diameter of lymphatic nodule in control and DEX groups at different days of the experiment
|
Parameters |
Groups |
Day 7 |
Day 14 |
Day 21 |
Day 28 |
||||
|
Mucosal thickness (µm) |
Non-DEX |
200.75 ± 15.22a |
261.10 ± 22.94a |
307.50 ± 18.04a |
397.20 ± 18.27a |
||||
|
DEX-1 |
174.80 ± 6.75a |
193.00 ± 3.79b |
144.00 ± 9.68c |
225.45 ± 15.66b |
|||||
|
DEX-2 |
137.40 ± 11.13b |
152.35 ± 5.09c |
179.55 ± 9.92b |
222.10 ± 18.95b |
|||||
|
DEX-3 |
134.55 ± 33.77b |
138.90 ± 6.72d |
154.10 ± 6.39c |
144.90 ± 8.84c |
|||||
|
Greater and lesser diameter of Lymphatic nodule (µm) |
Non-DEX |
110.30 ± 5.35a |
76.85 ± 3.70a |
119.65 ± 3.69a |
81.20 ± 4.81a |
131.65 ± 6.72a |
88.45 ± 3.55a |
148.00 ± 3.80a |
92.80 ± 2.71a |
|
DEX-1 |
105.85 ± 5.43b |
56.55 ± 4.23b |
92.10 ± 3.91b |
59.65 ± 2.71b |
47.95 ± 1.98d |
63.80 ± 2.71b |
102.25 ± 3.69b |
89.90 ± 6.72a |
|
|
DEX-2 |
87.00 ± 3.24c |
42.05 ± 4.23c |
61.40 ± 3.99c |
49.30 ± 2.71c |
83.90 ± 3.15b |
56.55 ± 2.61b |
88.00 ± 3.63c |
63.80 ± 2.69b |
|
|
DEX-3 |
58.00 ± 5.13d |
26.10 ± 1.78d |
43.45 ± 2.71d |
44.95 ± 2. 51c |
67.55 ± 2.73c |
59.45 ± 4.81b |
81.70 ± 6.23c |
49.30 ± 2.71c |
|
a,b,c,d Values in the same column with different alphabetic superscripts are significantly different from each other (P < 0.05)