Figure - available via license: Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International
Content may be subject to copyright.
-The effect of egg shell formation on plasma calcium, and on intestinal and uterine CaBP of laying quails l
Source publication
A vitamin D3-dependent calcium-binding protein (CaBP) has been found in the intestinal mucosa of Japanese quail (Coturnix coturnix japonica). This protein is similar, if not identical to that of the chick (Gallus domesticus). A similar protein fraction appears also in uterine mucosa of laying quail. Both intestinal and uterine CaBP levels are
highe...
Context in source publication
Citations
... UV-B plays a major role in the synthesis of Vitamin D3. Vitamin D3 activates the calcium-binding proteins in the intestine (Bar et al., 1976). Although dietary Vitamin D3 is supplemented, UV-B radiation has been shown to have a better effect on the bone health of young chicks compared to dietary supplementation of Vitamin D3 (Edwards, 2003). ...
Keel bone damage is an important welfare issue in laying hens and can occur with a high prevalence of up to 100% of hens within one flock. Affected hens suffer from pain. Although multiple factors contribute to the prevalence and severity of keel bone damage, selection for high laying performance appears to play a key role. With up to 300 eggs/year, Japanese quails show a high laying performance, too, and, thus, may also show keel bone damage. However, to our knowledge, there are no scientific results on keel bone damage in Japanese quails to date. Therefore, the aim of this study was to assess whether keel bone fractures and deviations occur in Japanese quails and to obtain more detailed information about the development of their keel bone during the production cycle. A group of 51 female quails were radiographed at 8, 10, 15, 19, and 23 weeks of age. The X-rays were used to detect fractures and deviations and to measure the lateral surface area, length, and radiographic density of the keel bone. In addition, the length of the caudal cartilaginous part of the keel bone was measured to learn more about the progress of ossification. At 23 weeks of age, quails were euthanized and their macerated keel bones assessed for fractures and deviations. Both keel bone deviations and keel bone fractures were detected in the Japanese quails. In the 23rd week of age, 82% of the quails had a deviated keel bone as assessed after maceration. Furthermore, there was a decrease in radiographic density, lateral surface area, and length of the keel bone between weeks of age 8 and 19. This could indicate a general loss of bone substance and/or demineralization of the keel bone. Our study shows that keel bone damage is not only a problem in laying hens but also affects female Japanese quails.
... Therefore, two mechanisms are enhanced: (i) enhancement for net absorption of Ca 2+ during the dark period, especially early in that period when feed is still present in the gut; and (ii) the activation of an efficient process of bone resorption, mostly of the medullary bone. Restoration of the circadian bone loss of Ca 2+ in birds that lay long sequential clutches occurs during the subsequent daylight period, when the intestinal absorption of Ca 2+ is enabled by the renewal of calcium intake (Bar et al., 1976). The bone accumulates rapidly during sexual maturation and the onset of egg production, and slow accumulation of its components continues throughout the laying period. ...
Experiments were performed in 110 ISA Brown egg production hens (Gallus gallus), kept from 15 to 26 weeks of age in enriched (furnished) housing technology. The present objective was to investigate the presence of single nucleotide polymorphisms (SNPs) of the ATP2B1 gene and their effects on calcium homeostasis in laying hens. The plasma membrane calcium-transporting ATPase 1 gene (ATP2B1) in hens is located on chromosome 1, region 43 273 706 – 43 305 815 bp. The ATP2B1 gene has 21 exons, and in this study three were genotyped. In each experimental group of animals, only alleles without deletions in exon 10 and only allele A in exon 12 were found. In exon 8, only genotypes CC/CC, TT/CC and TT/TT were found. These genotypes are associated with femur breaking strength, bone diameter, bone marrow diameter and compact bone thickness. No significant effects of SNPs in exon 8 on bone characteristics were found.
... Mean ± SE. Means designated by different letter are significantly different (P b 0.05) (with permission of: (Bar et al., 1976a;Montecuccoli et al., 1977b;Bar et al., 1992a;Bar et al., 1992d). Quelo et al., 1994). ...
... In the female bird intestinal calbindin mRNA (Striem and Bar, 1991;Bar et al., 1992a) and calbindin Montecuccoli et al., 1977b;Bar and Norman, 1981;Bar et al., 1990bBar et al., , 1978aBar et al., , 1992aBar et al., , 1996Nys et al., 1992a;Wu et al., 1994) are moderately increased during sexual maturation. Onset of laying markedly increased calbindin mRNA (Nys et al., 1989;Bar et al., 1990bBar et al., , 1992aStriem and Bar, 1991;Nys et al., 1992a) and calbindin (Wasserman and Taylor, 1968;Bar and Hurwitz 1973b;Bar et al., 1976aBar et al., , 1978aBar et al., ,b, 1992a) (Sugiyama et al., 2007) synthesis in the intestine and ESG. Molting (Yoshimura et al., 1997;Yosefi et al., 2003) and any other factor that arrests egg production (Bar and Hurwitz 1973b,a;Bar et al., 1992a) markedly reduce the intestinal and ESG calbindin contents. ...
Birds that lay long clutches (series of eggs laid sequentially before a "pause day"), among them the high-producing, strongly-calcifying Gallus gallus domesticus (domestic hen) and Coturnix coturnix japonica (Japanese quail), transfer about 10% of their total body calcium daily. They appear, therefore, to be the most efficient calcium-transporters among vertebrates. Such intensive transport imposes severe demands on ionic calcium (Ca2+) homeostasis, and activates at least two extremely effective mechanisms for Ca2+ transfer from food and bone to the eggshell. This review focuses on the development, action and regulation of the mechanisms associated with paracellular and transcellular Ca2+ transport in the intestine and the eggshell gland (ESG); it also considers some of the proteins (calbindin, Ca2+ATPase, Na+/Ca2+ exchange, epithelial calcium channels (TRPVs), osteopontin and carbonic anhydrase (CA) associated with this phenomenon. Calbindins are discussed in some detail, as they appear to be a major component of the transcellular transport system, and as only they have been studied extensively in birds. The review aims to gather old and new knowledge, which could form a conceptual basis, albeit not a completely accepted one, for our understanding of the mechanisms associated with this phenomenon. In the intestine, the transcellular pathway appears to compensate for low Ca2+ intake, but in birds fed adequate calcium the major drive for calcium absorption remains the electrochemical potential difference (ECPD) that facilitates paracellular transport. However, the mechanisms involved in Ca2+ transport into the ESG lumen are not yet established. In the ESG, the presence of Ca2+-ATPase and calbindin--two components of the transcellular transport pathway--and the apparently uphill transport of Ca2+ support the idea that Ca2+ is transported via the transcellular pathway. However, the positive (plasma with respect to mucosa) electrical potential difference (EPD) in the ESG, among other findings, indicates that there may be major alternative or complementary paracellular passive transport pathways. The available evidence hints that the flow from the gut to the ESG, which occurs during a relatively short period (11 to 14 h out the 24- to 25.5-h egg cycle), is primarily driven by carbonic anhydrase (CA) activity in the ESG, which results in high HCO3(-) content that, in turn, "sucks out" Ca2+ from the intestinal lumen via the blood and ESG cells, and deposits it in the shell crystals. The increased CA activity appears to be dependent on energy input, whereas it seems most likely that the Ca2+ movement is secondary, that it utilizes passive paracellular routes that fluctuate in accordance with the appearance of the energy-dependent CA activity, and that the level of Ca2+ movement mimics that of the CA activity. The on-off signals for the overall phenomenon have not yet been identified. They appear to be associated with the circadian cycle of gonadal hormones, coupled with the egg cycle: it is most likely that progesterone acts as the "off" signal, and that the "on" signal is provided by the combined effect of an as-yet undefined endocrine factor associated with ovulation and with the mechanical strain that results from "egg white" formation and "plumping". This strain may initially trigger the formation of the mammillae and the seeding of shell calcium crystals in the isthmus, and thereafter initiate the formation of the shell in the ESG.
... Mean ± SE. Means designated by different letter are significantly different (P b 0.05) (with permission of: (Bar et al., 1976a;Montecuccoli et al., 1977b;Bar et al., 1992a;Bar et al., 1992d). Quelo et al., 1994). ...
... In the female bird intestinal calbindin mRNA (Striem and Bar, 1991;Bar et al., 1992a) and calbindin Montecuccoli et al., 1977b;Bar and Norman, 1981;Bar et al., 1990bBar et al., , 1978aBar et al., , 1992aBar et al., , 1996Nys et al., 1992a;Wu et al., 1994) are moderately increased during sexual maturation. Onset of laying markedly increased calbindin mRNA (Nys et al., 1989;Bar et al., 1990bBar et al., , 1992aStriem and Bar, 1991;Nys et al., 1992a) and calbindin (Wasserman and Taylor, 1968;Bar and Hurwitz 1973b;Bar et al., 1976aBar et al., , 1978aBar et al., ,b, 1992a) (Sugiyama et al., 2007) synthesis in the intestine and ESG. Molting (Yoshimura et al., 1997;Yosefi et al., 2003) and any other factor that arrests egg production (Bar and Hurwitz 1973b,a;Bar et al., 1992a) markedly reduce the intestinal and ESG calbindin contents. ...
Egg laying and shell calcification impose severe extra demands on ionic calcium (Ca2+) homeostasis; especially in birds characterized by their long clutches (series of eggs laid sequentially before a "pause day"). These demands induce vitamin D metabolism and expression. The metabolism of vitamin D is also altered indirectly, by other processes associated with increased demands for calcium, such as growth, bone formation and egg production. A series of intestinal, renal or bone proteins are consequently expressed in the target organs via mechanisms involving a vitamin D receptor. Some of these proteins (carbonic anhydrase, calbindin and calcium-ATPase) are also found in the uterus (eggshell gland) or are believed to be involved in calcium transport in the intestine or kidney (calcium channels). The present review deals with vitamin D metabolism and the expression of the above-mentioned proteins in birds, with special attention to the strongly calcifying laying bird.
... Calcium absorption in the small intestine occurs mainly by active transport. Higher calcium absorption at the level of the intestines has been observed during shell formation than before this process (Bar et al, 1976). The Ca 2+ -ATPase influences absorption of calcium in the chick intestine (Haussler et al., 1970). ...
Plasma calcium concentration and uterine and duodenal adenosine triphosphatase (ATPase) activities were determined during shell formation for high (H) and low (L) shell strength lines of hens selected from the last of four consecutive generations. The H and L lines were divided into three groups according to shell formation at 0, 15, and 22 h following oviposition. Plasma total calcium was determined from blood samples collected from the common carotid artery. Activity of ATPase was determined in uterine and duodenal mucosa.
Shell strength, shell weight, percentage of shell per egg and shell thickness of the H line hens significantly exceeded those of the L line. During shell formation, no significant fluctuation in plasma calcium levels was observed within a line, but overall mean plasma calcium concentrations were higher in the H line than L line. Uterine ATPase activity increased with time after oviposition in both lines, with that of the H line being greater. Duodenal ATPase activity of H line hens remained fairly constant throughout the period, but this value showed fluctuations in the L line hens. It thus appears that laying hens with high and low shell strength may vary in their ability to use calcium for shell formation.
... When injected intramuscularly with estradiol, both lines exhibited a significant increase in both vitellogenin binding and protein-bound calcium concentrations (Grunder et al, 1980b). Calcium-binding protein in the intestine and uterus can be increased significantly by vitamin D 3 administration (Corradino et al, 1968;Bar et al, 1976). Although a positive correlation with estradiol was not found, Soares and Ottinger (1980) found a positive correlation 'Journal Article No. 4518 between the percentage of the egg that was shell and plasma 1,25-dihydroxyvitamin D 3 concentration when blood samples were taken at 6 and 20 hr postoviposition. ...
Laying hens were divided into high and low shell quality groups on the basis of egg specific gravity during the 5th, 7th, and 9th months of egg production. Concentrations of progesterone and estradiol-17 beta in the plasma were determined by radioimmunoassay in blood samples taken by cardiac puncture from hens at either 18, 21, or 24 hr postoviposition. Egg production and egg weight were not significantly different between shell quality groups; however, hens in the low shell quality group were heavier (P less than .05), had longer clutches (P less than .05), and lower egg specific gravity (P less than .0001) than hens in the high shell quality group. Egg production and clutch size declined (P less than .01) in both groups with increased age. Plasma estradiol and progesterone concentrations were not different between shell quality groups or among periods of production, though progesterone and estradiol concentrations were greater (P less than .005) at 21 hr postoviposition than at 18 or 24 hr. The correlation coefficient between plasma concentrations of estradiol and progesterone was significant. There was no significant association between the plasma concentrations of these hormones and egg shell quality. These data suggest that concentrations of estradiol and progesterone in plasma, during the 6 hr before ovulation, are not highly related to shell quality in the laying hen.
... Mean levels of quail duodenal CaBP reported by Bar et al. (1976) are slightly higher than the CaBP levels found in the turkey poult by Musser etal. (1976). ...
... protein. No difference was found between calcifying and inactive uterus samples as was found by Bar et al. (1976). This may be due to the small sample size in this study. ...
Detection of immunologically similar calcium binding proteins (CaBP) in the duodenum and the uterus of both laying turkey hens and laying Japanese quail by a 45 Ca binding assay yielded results highly correlated with the actual concentration of CaBP measured by radial immunodiffusion. Levels of duodenal CaBP in laying turkeys maintained on normal laying hen rations are lower than those found in hens just starting egg production and than laying hens fed a Ca-restricted diet for one month.
Calcium and phosphorus are critical elements in the maintenance of living systems. Thus, elegant regulatory mechanisms have evolved by which calcium and phosphorus homeostasis is maintained. In vertebrates, the intestine, kidney, and bone are intimately involved in this process and in fish, the gills appear also to regulate mineral homeostasis. A number of endocrine organs function in the orchestration of mineral homeostasis, including the parathyroid gland, parafollicular cells (C-cell) of the thyroid, and the endocrine kidney. In addition, endocrine factors from the adrenals, gonads, thyroid, and pituitary appear, either directly or indirectly, to influence calcium and phosphorus homeostasis. The purpose of this chapter is to focus primarily on the current views of the mechanisms involved in the intestinal transport of calcium and inorganic phosphate in relation to mineral absorption, with particular emphasis on the physiologic and biochemical mechanisms of calcium and phosphate transport.
In this paper, we present a review of the current knowledge of calcium and phosphorus metabolism as it relates to homeostatic mechanisms in poultry; contrasts with mammalian systems, as well as information that is available for mammalian systems but not for avian systems, are included. The importance of understanding these homeostatic mechanisms and their effect on Ca and P absorption are discussed briefly in the context of correctly formulating poultry diets. Clearly deficiencies, excesses, or imbalances in Ca and P result in a cascade of changes, including increases or decreases in the absorption of these 2 minerals from the intestinal lumen. The effect of these changes on the actual digestibility of Ca and P from ingredients or diets has not been determined yet; most of the methodologies we used for the determination of P digestibility are done under deficient conditions. The methods used to determine the release of Ca and P from diets when phytases were used were also done under deficient Ca and P conditions. given the effect of Ca and P homeostatic mechanisms on absorption, methodologies and, possibly, the quantitative the effect of these homeostatic mechanisms on values generated for Ca and P should be reviewed before they can confidently be applied.
Zusammenfassung
Mittels indirekter Immunofluoreszenz‐ bzw. Immunoperoxidasetechnik gelang ein licht‐ und elektronenoptischer Nachweis von Vitamin D‐abhängigem CaBP im Uterus legereifer Wachteln, die nach 17tägiger Depletion wieder mit 4000 i. E. Vitamin D 3 bzw. 10% lyophilisiertem Goldhafer angefüttert worden waren. Die Präparation der Schnitte mit dem Gefrier‐Auftau‐Verfahren ergab eine Lokalisation von CaBP im Zytoplasma der Drüsenepithelzellen. Unter Verwendung der Gefriertrocknung und der Gefriersubstitution hingegen zeigte sich eine CaBP‐spezifische Reaktion im Zytoplasma sowohl der Drüsen‐ als auch der Oberflächenepithelzellen.
Elektronenmikroskopisch erfolgte mit allen drei Verfahren ein Nachweis von CaBP in den Polysomen und gelegentlich auch in den Mitochondrien der Drüsenepithelzellen sowie in unterschiedlichen elektronendichten Gebilden in den Tropfen von Drüsenepithelzellen, die in der Lamina propria situiert waren. Während mit dem Gefrier‐Auftau‐Verfahren — möglicherweise infolge eines “Auswascheffektes” — im Oberflächenepithel kein CaBP festzustellen war, fand sich mit den beiden anderen Präparationsmethoden eine CaBP‐spezifische Färbung in den Polysomen und in den Mitochondrien der Becherzellen und der Flimmerepithelzellen des Oberflächenepithels.
Diese Befunde weisen CaBP im Uterus als zytoplasmatisches Protein aus. Seine Beteiligung am intrazellulären Kalziumtransport während der Eischalenbildung wird diskutiert.
Summary
Experimental studies on extraosseous calcification in hypervitaminosis D 3 V. Light and electronoptical localization of calcium‐binding protein (CaBP) in the uterus of egg‐laying quail
Using immune‐peroxidase or immunofluorescence techniques a light‐ and EM‐demonstration of vitamin D‐dependent CaBP in the uterus of laying quail which after 17 days depletion were fed with 4000 i. u. of vitamin D 3 or 10% lyophilized “goldhafer”, was successfully carried out. Preparation of sections with the freeze‐thaw method enabled localization of CaBP to be identified in the cytoplasm of glandular epithelial cells. By using freeze‐drying and freeze‐substitution a CaBP‐specific reaction was detected in the cytoplasm of both glandular and surface epithelial cells.
With the EM it was possible to demonstrated CaBP in the polysomes and occasionally also in the mitochondria of glandular epithelial cells as well as in various electron‐dense structures in the droplets of glandular epithelial cells situated in the lamina propria. Although in the freeze‐thaw method — possibly as a result of a ‘washing‐out’ effect — no CaBP could be found in the surface epithelium, the two other techniques gave a CaBP‐specific staining in the polysomes and in the mitochondria of goblet cells in the ciliated epithelial cells of the surface epithelium.
These findings show that CaBP is present in the uterus as a cytoplasmic protein. Its involvement in intracellular Ca transport during shell formation is discussed.
Résumé
Recherches expérimentales sur la calcification extraosseuse lors d'hypervinaminose D 3 V. Localisation au microscope optique et électronique de la protéine liant le calcium (CaBP) dans l'utérus de cailles pondeuses .
Il a été possible au moyen de la technique par immunofluorescence et immunooperoxydase de mettre en évidence par microscopie optique et électronique la présence de CaBP dépendant de la vitamine D dans l'utérus de cailles pondeuses qui après une depletion de 17 jours avaient reçu à nouveau 4000 UI de vitamins D3 et respectivement 10% d'avoine dorée lyophilisée. Les préparations des coupes par la technique congélation‐décongélation a permis une localisation de CaBP dans le cytoplasme des cellules épithéliales glandulaires. L'emploi de la lyophilisation et de la subsitution par congélation a montré par contre une réaction CaBP spécifique aussi bien dans le cytoplasme des cellules épithéliales glandulaires que superficielles. Le microscope électronique a permis avec les trois procédés de mettre en évidence CaBP dans les polysomes, de temps en temps également dans les mitochondries des cellules epitheliales glandulaires ainsi que dans différentes images étanches aux électrone dans les gouttes des cellules epitheliales glandulaires qui étaient situées dans Lamina propria. Avec la méthode congélation‐décongélation probablement à la suite d'un effet de ≤lavage≥, CaBP n'a pas été mis en évidence dans l'épithélium de surface. Les deux autres méthodes de préparation ont permis de trouver une coloration spécifique de CaBP dans les polysomes et dans les mitochondries des cellules caliciformes et les cellules épithéliales à cils vibratifs de l'épithélium de surface.
Ces résultats montrent que CaBP est une protéine cytoplasmique dans l'utérus. On discute sa participation au transport intracellulaire du calcium durant la formation de la coquille de l'œuf.
Resumen
Estudios experimentales sobre la calcificacion extraósea en la hipervitaminosis D 3 V. Localización foto y electrónicoóptica de la proteína fijadora de calcio (PFCa) en el útero de codornices ponedoras
Mediante la técnica de la inmunofluorescencia indirecta resp. de la inmunoperoxidasa, se logró la puesta en evidencia foto y electronicoóptica de PFCa dependiente de la vitamina D en el útero de codornices maduras para la puesta, las cuales habían sido vueltas a alimentar, tras una depleción de 17 días, con 4000 u. i. de vitamina D 3 resp. un 10% de avena dorada liofilizada. La preparación de los cortes con el procedimiento de congelación‐descongelación reveló una localización de PFCa en el citoplasma de las células del epitelio glandular. Sin embargo, utilizando la liofilización y la criosubstitución se reveló una reacción específica de PFCa en el citoplasma tanto de las células glandulares como de las del epitelio superficial.
Al microscopio electrónico se pudo poner en evidencia con todas las tres técnicas la PFCa en los polisomas y, en ocasiones también, en las mitocondrias de las células de los epitelios glandulares así como en formaciones opacas de forma diferente en las gotas de células de epitelios glandulares, que se hallaban situados en la lámina propia. Mientras que con el procedimiento de congelación‐descongelación — probablemente a causa de un ≤efecto de lavado≥ — no se pudo apreciar ninguna PFCa en el epitelio superficial, se halló con las otras dos técnicas de preparación una coloración específica de PFCa en los polisomas y en las mitocondrias de las células caliciformes y de las células de los epitelios vibrátiles del epitelio superficial.
Estos hallazgos justifican a la PFCa en el útero como proteína citoplasmática. Se discute su participación en el transporte intracelular del calcio durante la formación de la cáscara de huevo.
