Histomorphology and ultrastructure of the proventriculus of the broad breasted white turkey (Meleagris gallopavo, Linnaeus 1758).

This study aims to examine the functional morphology of the proventriculus of the broad breasted white turkey (BBWT) (Meleagris gallopavo, Linnaeus 1758) using gross anatomy, light microscopy, gross/histomorphometric analysis, and scanning and transmission electron microscopy. The proventriculus was characterized internally by many elevated papillae with a mound, leafy flower, and lotus flower-shapes. Each papilla was enclosed by concentrically organized mucosal folds with distinct or indistinct proventricular gland openings on its top. Longitudinal folds and grooves at the junction of the proventriculus with the esophagus exhibited various sized and shaped openings of esophageal glands with irregular outlines. Histologically, the surface epithelium of the proventriculus was covered by a thin layer of cuticle, particularly evident at its junction with the gizzard. The lamina epithelialis and propria, and secretory units of proventricular lobules were infiltrated by aggregations of lymphocytes and lymphoid follicles (nodules). Variably shaped glandular lobules of proventricular glands occupied the submucosa, surrounded by thin strands of smooth muscle fibers derived from muscularis mucosa. Triangular, cuboidal, or columnar-shaped secretory oxyntico-peptic cells lined the secretory units. Many telocytes (pyramidal or fusiform-shaped cell bodies with lengthy telopodes) were observed in interstitial tissue. Further, two types of argyrophilic endocrine cells were identified within the glandular epithelium. The morphology of the M. gallopavo proventriculus reflects its dietary habits and behavior.

The foregut and its manipulation via feeding practices in the chicken.

The digestive system of a chicken is simple, short, and extremely efficient. In the upper gastro-intestinal tract (GIT) or foregut, in particular when feed is not continuously available, moistening of feed occurs in the crop followed by chemical and physical digestion in the proventriculus and gizzard, respectively, as a preparation for nutrient absorption which occurs in the distal portions of the GIT-duodenum, jejunum, ileum, and to a lower extent in the ceca and colon. Broiler husbandry practices in the past have focused on achieving high intakes of readily digestible nutrients and largely failed to consider the ancestral purpose of the foregut. Besides holding important digestive functions, the rich bacterial communities in both the crop and the gizzard transform these organs in barriers preventing the invasion of the GIT by pathogens. However, the amount of time feed spends in the foregut-and therefore the environment in this gut segment-is extremely variable and can be manipulated by different feeding practices, such as feed structure and composition, frequency of feeding, and/or light regimes.With further progress in terms of performance and health of broiler chickens being sought, and following decades of research focusing on distal GIT segments, it is about time to explore how the foregut can contribute to achieve these goals. This review revisits morphological, physiological, and microbiological characteristics of the foregut; explores the importance of this GIT portion as feed intake regulator; and discusses how husbandry and feeding practices such as lighting regimes and feed structure may be adapted to activate the crop and the gizzard, which results in performance and health improvements. Finally, interactions of these feeding practices with exogenous enzymes are discussed and suggestions for further research are made.

Developmental morphology study on the stomach of African ostrich chicks.

This study investigates the developmental morphology changes on the proventriculus and gizzard of African ostrich chicks using gross anatomy, and light and transmission electronic microscopy. The body weight; the weight of the proventriculus and gizzard; the thickness of simple tubular glands, muscularis mucosae, compound tubular glands, and muscular layer of the proventriculus; and the thickness of the gizzard glands and muscularis mucosae were measured on postnatal d one, 45, 90, and 334. Under transmission electronic microscopy, the oxynticopeptic cells, mucous cells, and endocrine cells were observed in the proventriculus on postnatal d one, 45, and 90. Our results revealed that the weights of the proventriculus and gizzard (relative to the body weight) both peaked on d 90, respectively, and declined thereafter. The thicknesses of the simple tubular glands, muscularis mucosae, compound tubular glands, muscular layer of the proventriculus, and gizzard glands increased with increasing bird age. However, the thickness of the muscularis mucosae in the gizzard peaked on d 90 and gradually decreased thereafter. The result of transmission electron microscopy revealed that the glands in the submucosa layer of the proventriculus did not differ from other cell types, aside from an increase in cell diameter and an increase in cytoplasmic content. A significant increase in the number of mitochondria and smooth endoplasmic reticulum was noted, as well as an in granules secreted by endocrine cells. Therefore, to improve brood rates of reared African ostrich chicks, feed management ought to be enhanced between postnatal d one and day 90.

Molecular cloning of motilin and mechanism of motilin-induced gastrointestinal motility in Japanese quail.

Motilin, a peptide hormone produced in the upper intestinal mucosa, plays an important role in the regulation of gastrointestinal (GI) motility. In the present study, we first determined the cDNA and amino acid sequences of motilin in the Japanese quail and studied the distribution of motilin-producing cells in the gastrointestinal tract. We also examined the motilin-induced contractile properties of quail GI tracts using an in vitro organ bath, and then elucidated the mechanisms of motilin-induced contraction in the proventriculus and duodenum of the quail. Mature quail motilin was composed of 22 amino acid residues, which showed high homology with chicken (95.4%), human (72.7%), and dog (72.7%) motilin. Immunohistochemical analysis showed that motilin-immunopositive cells were present in the mucosal layer of the duodenum (23.4±4.6cells/mm(2)), jejunum (15.2±0.8cells/mm(2)), and ileum (2.5±0.7cells/mm(2)), but were not observed in the crop, proventriculus, and colon. In the organ bath study, chicken motilin induced dose-dependent contraction in the proventriculus and small intestine. On the other hand, chicken ghrelin had no effect on contraction in the GI tract. Motilin-induced contraction in the duodenum was not inhibited by atropine, hexamethonium, ritanserin, ondansetron, or tetrodotoxin. However, motilin-induced contractions in the proventriculus were significantly inhibited by atropine and tetrodotoxin. These results suggest that motilin is the major stimulant of GI contraction in quail, as it is in mammals and the site of action of motilin is different between small intestine and proventriculus.

[Knockdown of InR gene in ventral nephrocytes promotes resistance to toxic stress in Drosophila melanogaster females].

Hemolymph filtration in insects is performed by nephrocytes, additional cells of the circulatory system that are not connected to Malpighian vessels. Drosophila has two types of nephrocytes: the ventral ("garland"), which are situated around the connection site of the esophagus and proventriculus, and the pericardial, which are localized around the heart. In this study, we examined the role of the of insulin-like receptor (InR)gene in regulation of the function of ventral nephrocytes (VNC) in D. melanogaster females. Immunofluorescent analysis of female VNC with anti-InR antibodies revealed for the first time that the InR gene is expressed in VNC cells. To determine whether a change in the level of InR expression has an effect on VNC function in Drosophila females, we implemented an antisense suppressor of the InR gene, together with a driver that is expressed specifically in VNC. VNC function was evaluated by survival of the females exposed to toxic stress (treatment with AgNO3). This study has shown for the first time that suppression of InR expression in VNC leads to a rise in the survival of flies under conditions of toxic stress.

Evaluation of litter type and dietary coarse ground corn inclusion on broiler live performance, gastrointestinal tract development, and litter characteristics.

Two 49 d floor pen studies were conducted to evaluate the effects of litter type and dietary coarse ground corn (CC) inclusion on broiler live performance, gastrointestinal tract (GIT) development, and litter characteristics. Experiment 1 was a 2×2 factorial arrangement of 2 genders (male or female) and 2 CC levels (0 or 50%). From 15 to 35 d, the addition of CC decreased feed intake (P<0.01) and BW gain (P<0.05) of males but not females. The inclusion of CC decreased feed intake (P<0.01) and BW gain (P<0.01) from 0 to 49 d but improved adjusted feed conversion ratio (AdjFCR) from 35 to 49 d (P<0.05). Male broilers exhibited better live performance than females during the study as evidenced by greater feed intake (P<0.01) and BW gain (P<0.01), and improved FCR (P<0.01), but with increased mortality (P<0.05). The inclusion of CC increased relative gizzard weight (P<0.01) and decreased relative proventriculus weight (P<0.01) at 49 d. Experiment 2 was a 2×2 factorial arrangement of 2 CC levels (0 or 50%) and 2 litter types (ground old litter or new wood shavings litter). The inclusion of CC decreased feed intake throughout the experiment without affecting final BW when only males were used and improved FCR after 25 d (P<0.01). New litter improved FCR from 1 to 14 d (P<0.01). At 49 d, the birds fed the CC diet had reduced excreta nitrogen (P<0.05) and litter moisture (P<0.05). In conclusion, 50% CC inclusion initially produced negative effects on live performance that became positive as BW increased. The effects of CC became evident at an earlier age for males. New litter had only a marginal benefit on broiler live performance.

Peripheral "chicken" obestatin administration does not affect feed intake and gut muscle contractility of meat-type and layer-type chicks (Gallus gallus domesticus).

Obestatin has recently been discovered in the rat stomach. As for ghrelin, the 23-amino acid obestatin is also derived from post-translational processing of the prepro-ghrelin gene but seems to have opposite effects on feed intake. In avian species, ghrelin is mainly present in the proventriculus and decreases feed intake, as opposed to its orexigenic properties in mammals. An obestatin-like sequence was also found in the avian ghrelin precursor protein but the potential involvement of this peptide in appetite regulation of chickens is unclear. We therefore investigated the effects of a single peripheral administration of this predicted "chicken" obestatin peptide on voluntary feed intake of 7- to 9-day-old meat-type and layer-type chicks. "Chicken" obestatin was injected intraperitoneally or intravenously at a dose of 1 nmol or 10 nmol/100 g body weight and feed intake was measured up to 4 h post injection. None of these treatments did reveal any effect of the putative "chicken" obestatin on appetite of either meat-type of layer-type chicks. Furthermore, "chicken" obestatin also failed to affect the in vitro contractility of muscle strips from crop and proventriculus. In conclusion, in the given experimental settings, the putative "chicken" obestatin has indistinctive physiological effects on feed intake and in vitro muscle contractility of gut segments, and hence its functional properties in ingestive behavior of avian species remain obscure.

The differences between the localizations of MUC1, MUC5AC, MUC6 and osteopontin in quail proventriculus and gizzard may be a reflection of functional differences of stomach parts.

Mucins are high molecular weight glycoproteins which constitute the major component of the mucus layer and are produce by many epithelial tissues in vertebrates. Osteopontin (OPN) is an adhesive phosphorylated glycoprotein that is expressed by a broad range of tissues and cells. Although gastric mucins MUC1, MUC5AC, MUC6 and OPN have been widely used in histological studies and in diagnostic pathology in order to diagnose gastric carcinomas, their localizations in the stomach of quail have not yet been studied. In this study, the localizations of MUC1, MUC5AC, MUC6 and OPN in the proventriculus and gizzard of Japanese quail during the post-hatching period were compared at light microscope levels by applying immunohistochemical methods. In all ages studied, the immunoreactivity of MUC5AC was present in the lining epithelium of both folds and superficial proventricular glands in the proventriculus, whereas MUC1, MUC6 and OPN reactivity was found in the oxynticopeptic cells of profound proventricular glands. In addition, some cells in the fold epithelium of the proventriculus showed a positive reaction to OPN. The immunoreactivity of MUC1 in gizzard was different from that of MUC5AC. Although MUC5AC was expressed in the cells of both the surface epithelium and profound glands of the gizzard, MUC1 was only localized in the profound glands of the gizzard. However, MUC6 and OPN immunoreactivity was absent in the gizzard. The results indicated that the differences between the localizations of MUC1, MUC5AC, MUC6 and OPN in quail proventriculus and gizzard may be a reflection of functional differences of stomach parts. Although the biological significances of the expressions of MUC1, MUC5AC, MUC6 and OPN in the quail stomach remains unknown, these notable glycoproteins may be associated with barrier function, host defence, and/or secretion.

Correlation between Musashi-1 and c-hairy-1 expression and cell proliferation activity in the developing intestine and stomach of both chicken and mouse.

Musashi-1 (Msi-1) is an RNA-binding protein that plays key roles in the maintenance of neural stem cell states and in their differentiation into neural cells. Msi-1 has also been proposed as a candidate marker gene of mammalian intestinal stem cells and their immediate lineages. In this study, we examined Msi-1 expression in the small intestine and the stomach of both chicken and mouse during embryonic, fetal and postnatal development. In addition, we analyzed the expression of c-hairy-1, a chicken homologue of mouse Hes1, and assessed the proliferative activity of the cells expressing both of these factors. Significantly, during the development of these digestive organs in both species Msi-1 expression showed dynamic changes, suggesting that it is important for digestive organ development, particularly for epithelial differentiation. Based on our observations of the expression patterns of Msi-1 and c-hairy-1 in the adult small intestine, we speculate that Msi-1 is also a stem cell marker of the chicken small intestinal epithelium.

The anatomy, physiology, and diseases of the avian proventriculus and ventriculus.

Diseases affecting the proventriculus and ventriculus often present with similar clinical signs. It is important for the avian practitioner to be familiar with these diseases, their prevalence, and the species most commonly affected to judiciously prioritize the appropriate diagnostic techniques. A basic understanding of the anatomy and physiology of the proventriculus and ventriculus is useful in integrating the pathophysiology and clinical signs associated with variable disease processes. It is also essential to evaluate radiographs and endoscopic images, perform diagnostic techniques, make a diagnosis, and provide appropriate therapy.

Potentiation of motilin-induced contraction by nitric oxide synthase inhibition in the isolated chicken gastrointestinal tract.

The present experiments were designed to determine whether or not endogenous nitric oxide (NO) modifies the contractile response to chicken motilin (ch-MT) in the gastrointestinal (GI) tract (proventriculus and small intestine) of the chicken. ch-MT (1 nmol L(-1)-1 micromol L(-1)) caused contractions of longitudinal muscle strips of the proventriculus through both myogenic and neurogenic (mostly cholinergic) mechanisms. On the other hand, ch-MT (0.1 nmol L(-1)-100 nmol L(-1)) contracted the small intestine (duodenum, jejunum and ileum) only through a myogenic mechanism. L-Nitroarginine methylester (L-NAME) potentiated, and L-arginine inhibited, the ch-MT- induced contraction without affecting the responsiveness of acetylcholine (ACh) or 5-hydroxytryptamine in the proventriculus. Electrical field stimulation (EFS)- and 1,1-dimethyl-4-phenylpiperazinium (DMPP)- induced contractions were also potentiated by L-NAME. The potentiation by L-NAME was prevented by L-arginine but not by D-arginine. However, in the presence of atropine or tetrodotoxin, neither L-NAME nor L-arginine modified the responses to ch-MT and DMPP. In contrast to the proventriculus, L-NAME and L-arginine were both ineffective in modifying the ch-MT-induced contraction in the small intestine. These results indicate that NO synthase inhibition potentiates the contractile response of ch-MT, EFS and DMPP in the chicken proventriculus through reduction of endogenous NO-mediated presynaptic inhibition on neural ACh release. However, NOS inhibition did not modify the myogenic (direct) action of ch-MT in gastric and intestinal smooth muscles of the chicken.

Performance and gastro-intestinal response of broiler chickens fed on cereal grain-based foods soaked in water.

1. Two experiments were carried out to investigate the addition of 1 3 kg water per kg air-dry mash diets containing high proportions (600 to 700 g/kg) of ground cereal grains (wheat, barley or oats) on broiler performance and the structure and function of the gastro-intestinal tract. 2. Chicks at the age of 7 d were fed on the wheat-, barley- or oats-based diets in the dry or wet forms for 35 d. Food and water intakes were recorded daily while body weight was measured weekly. Two birds from each treatment were killed each week to measure gut size and the viscosity of gut contents. Tissue samples from various digestive segments were histo-morphologically examined to determine the thickness of tissue layers, size of tissue glands, villa heights, crypt depths and thickness of tunica muscularis. Crypt cell proliferation rate (CCPR) for each segment was also determined using a metaphase arrest technique. 3. The results from both experiments showed that wetting food significantly (P<0.05) increased food intake, total water intake and body weight gain of broiler chickens. The body weight gains of birds were proportional to their food intakes so that the efficiency of food utilisation was similar for all treatments. Dry matter retention of food tended to increase in birds given wet food from 7 to 21 d but not thereafter, compared to the dry-fed birds. Although water intake from the water bottle was significantly (P<0.05) reduced in birds given wet food, total water intakes from the water bottle plus that from food were significantly (P<0.05) higher in the wet-fed birds than in the dry-fed birds. The ratio of total water to dry food intake was, however, similar in both feeding regimens. 4. The fresh empty weight of the gut was increased by wet-feeding while its relative weight to body weight and the length of gut was not affected by dietary treatments. Significantly greater development of the tissue glands in the proventriculus and gizzard was observed in the birds given wet food; this was associated with the reduced thickness of the muscular layer of these segments. An increase in villus height was also observed in duodenum, small intestine, caeca and colon of birds given wet food, compared to those given dry food. CCPR was significantly (P<0.05) reduced by wet-feeding throughout the digestive tract. This was associated with a significant decrease in the mean viscosity of the gut contents and the concentration of volatile fatty acids (VFAs) in the caeca. 5. Wetting diets based on cereal grains caused a significant improvement in the performance of broiler chickens. The mechanism of the beneficial effects of wet feeding could be attributed to the decreased viscosity of gut contents; the greater development of the layer of villi in the digestive segments and the reduced CCPR in the crypts of the epithelium.

The influence of environmental temperature on in vivo limestone solubilization, feed passage rate, and gastrointestinal pH in laying hens.

Several studies were conducted to determine whether suppression of in vivo limestone solubilization was partially responsible for the reduction of shell quality under conditions of high environmental temperatures. In Experiment 1, excreta from hyperthermic and thermoneutral hens fed three levels of Ca (3.5, 4.3, and 5.2%) at two cycling environmental temperatures (averaging 28.3 and 26.1 C) were analyzed for percentage in vivo limestone solubilization. Hens in Experiment 2 received diets containing 3.9% Ca at 32.8 and 18.3 C. Rate of feed passage and gastrointestinal pH were also measured in Experiment 2. Experiment 3 evaluated the influence of temperature (22.2 and 30.0 C) and feed consumption on percentage limestone solubilization when Ca intake was held constant. In vivo limestone solubilization was influenced by Ca level in Experiment 1, but not by temperature. Hyperthermic hens solubilized a higher percentage of limestone than thermoneutral hens in Experiment 2, but it was concluded that this was due to a difference in Ca consumption and not due to temperature directly. When Ca intake was held constant in Experiment 3, there was no difference in limestone solubilization at the temperatures tested. Rate of feed passage was 16.6% slower in the hyperthermic hens. Crop, proventriculus, and upper small intestine pH were similar at each temperature, but gizzard pH was significantly lower in the hyperthermic hens. It was concluded that the high environmental temperatures used in the present studies did not suppress in vivo limestone solubilization.

Stunting syndrome in broilers: effect of age and exogenous amylase and protease on performance, development of the digestive tract, digestive enzyme activity, and apparent digestibility.

Day-old male, meat-type chicks raised in brooder batteries were infected by orally administering an inoculum prepared from intestines of broiler chicks infected with stunting syndrome (SS). Naive controls were kept in a parallel room. The chicks were fed a commercial starter diet supplemented with two levels of enzyme preparations to 14 d of age. The experiment was continued to the age of 6 wk in order to estimate compensatory feed intake and growth. In a parallel study, digestibility of the feed was determined from 1 to 3 wk of age with control or inoculated chicks. The enzymes amylase and proteases were produced by Bacillus subtilis and Penicillium emersonii. Enzyme supplementation had no effect on feed intake, growth, or feed utilization, or on digestibility of fat, starch, protein, or energy. Because enzyme supplementation did not consistently affect performance of chicks and no interactions were observed between enzyme supplementation and infection status, data are presented for effects of infection only. Inoculation of SS-infective material reduced performance to 4 wk. Compensatory growth and feed intake were observed from the age of 4 wk onward. At the age of 6 wk the slight retardation of the inoculated chicks was not significant. On Week 1, retention of fat, starch, protein, and energy was significantly depressed in the inoculated chicks. At the age of 2 wk, retention of starch was not depressed, and at the age of 3 wk, the only consistent depression was that observed for fat. The proventriculus weight and content were consistently higher in inoculated chicks, as were the small intestine and intestinal content. The pH of the gizzard content was higher, and that of the small intestine content was lower, in the inoculated birds than in their control counterparts. Stunting syndrome infection was accompanied by a significant depression of trypsin activity in the pancreas at the age of 1 and 2 wk. At these periods, amylase and chymotrypsin were not affected. At 6 wk of age, the activities of amylase, trypsin, and chymotrypsin in the pancreas were higher in the inoculated than in the control birds. In the intestinal chime, amylase, trypsin, and chymotrypsin activities were lower in the inoculated birds on Week 1 and 2 (NS for amylase on Week 1). On Week 6, the activity of all enzymes assayed was higher in the inoculated birds (NS for amylase). It is suggested that the main factors depressing feed intake and growth in SS-infected birds are most probably beyond those of digestion.

Hormonal control of avian pancreas by gastrin-releasing peptide from the proventriculus.

Endocrine cells in the acid-secreting part of the avian stomach, the proventriculus, contain two forms of gastrin-releasing peptide (GRP) of 27 and 6 residues, respectively. We have examined the actions of exogenous GRP-27 and GRP-6 and endogenously released GRP in the control of pancreatic secretion in urethan-anesthetized turkeys. Chicken GRP-27 and the structurally related amphibian peptide bombesin were potent stimulants of fluid and protein output from the pancreas (at 6-100 pmol/kg, iv). GRP-6 had no significant effect at doses up to 1,000 times higher. A bombesin antagonist, (CH3)2-CHCO-[D-Ala24]GRP-20--26-NHCH3, inhibited the action of exogenous chicken GRP-27 but did not inhibit intravenous cholecystokinin octapeptide (CCK-8). Distension of the proventriculus with a solution of peptone produced an increase in the flow of pancreatic juice and an increase in protein output, which was not reduced by atropine. The bombesin antagonist produced a reversible inhibition of this response. A CCK-gastrin antagonist, BOC-beta-Ala-Trp-Leu-Asp-O(CH2)2- phenyl(4F), which inhibited the action of exogenous CCK, had no effect on the pancreatic response to exogenous GRP-27 or to distension of the proventriculus with peptone. We suggest that protein-rich solutions in the proventriculus release GRP, which in turn acts directly on the pancreas to stimulate enzyme secretion.

Ability of cimetidine to increase intestinal pH of chicks.

Cimetidine was tested for its ability to increase the pH of intestinal contents of chicks. In Experiment 1, 18 mg cimetidine injected i.m. per kilogram body weight (BW) increased (P less than .05) proventricular pH within 1 hr, but proventricular pH began to decrease 4 hr postinjection. The pH of the contents of the duodenum was increased (P less than .05) 4 hr postinjection, whereas the pH of the contents of the jejunum was unaffected. In Experiment 2, 36 mg cimetidine injected i.m. per kilogram BW increased (P less than .05) the pH of the contents in the proventriculus and duodenum, but did not alter the pH in the jejunum and ileum. These results show that cimetidine can be used for experimental purposes to increase the pH of the proventricular and duodenal contents in chickens.

The endocrine cells of the chicken proventriculus.

The endocrine cells of the chicken proventriculus were investigated by selective staining techniques, immunohistochemistry and electron microscopy. The following endocrine cell types were identified: 1) Argyrophilic ECL-cells, of unknown function, were very numerous in the 21-day-old chick, but less numerous in the newborn chick; 2) somatostatin-producing D-cells; 3) GLI-cells producing glucagon-related peptides; 4) X-cells of unknown function; 5) BN-cells producing bombesin; and 6) relatively few 5-hydroxytryptamine-producing EC-cells. Each of these cell types show a distinct morphology, distribution and histochemical reactivity. With the exception of BN-cells, they resemble rather closely the corresponding endocrine cell types previously described in the oxyntic mucosa (EGL, D, X and EC cells) or in the intestinal mucosa (L-cells) of the mammalian gut.

Effect of bombesin on pancreatic secretion and gall bladder motility of the chicken.

Bombesin strongly stimulated the chicken pancreatic secretion. When given by i.v. infusion, the threshold dose was of the order of 7.5-45.0 ng/kg/min and maximum enzyme output was obtained at a rate of 60 ng/kg/min. In addition to total enzyme output, enzyme concentration was also increased. Caerulein displayed a more potent stimulant effect, but composition of juice produced by the two polypeptides was similar. Tachyphylaxis occurred only with bombesin. Neither atropine nor gastric acidification affected the response to bombesin. Bombesin was totally ineffective in promoting gall bladder emptying. It is suggested that in the chicken, bombesin acts on the exocrine pancreas indirectly through release of an endogenous pancreozymin possibly devoid of cholecystokinetic activity.