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PubMed1986

Eggshell conductance--Fick's or Stefan's law?

The diffusion of gases through the avian eggshell has usually been interpreted as an example of Fick's law, and this appears to be the best explanation for gas exchange through narrow long pores. Many eggshells, however, contain more elaborate pores, of which the funnel-shaped type is the most common. It is suggested that most of the resistance to diffusion through these types of pores is best explained by Stefan's law of diffusion through apertures. The implications of this theory to the understanding of pore functions are discussed.

Animals↗

PubMed1988

Diffusive resistance of avian eggshell pores.

Resistance to gas diffusion through the avian eggshell resides in the microscopic pores which penetrate the shell. We calculated the resistance to water vapor diffusion of individual pores in the shells of 23 species of avian eggs, based on measurements of pore dimensions taken from drawings of 321 pore casts published by Tyler (1962, 1964, 1965) and Tyler and Simkiss (1959). Diffusive resistances were calculated from Fick's first law, using a 100-segment model of each pore. In addition, we added 2 series resistances, calculated from Stefan's law, to account for boundary layer resistances at the inner and outer pore apertures. Convective resistances for the same 100-segment model were computed from Poiseuille's law. A special, symmetrically branching model is presented for the diffusive resistance of the branched pores of ostrich eggshells, based on the drawings of Tyler and Simkiss (1959). The total aperture resistance was less than 6.2% of total pore resistance, while the outside aperture effect was on average only 1.5%. The calculated average pore conductance for all species was 5.4 micrograms (day Torr)-1, about three times higher than the average value of 1.6 micrograms (day Torr)-1 obtained by dividing measured shell conductance by the number of pores (Ar and Rahn, 1985). A possible explanation for this discrepancy is advanced. However, it is to be noted that in spite of the discrepancy, both calculated and functional values of pore conductance appear to be independent of egg mass.

Animals↗

PubMed1989

The relationship between eggshell porosity and air space gas tensions measured before and during the parafoetal period and their effects on the hatching process in the domestic fowl.

The partial pressure of gases in the air space were measured before and during the parafoetal period for domestic fowl eggs of the same initial weight but varying in eggshell conductance. Embryos developed and hatched normally from eggs with a wide range of shell conductance and resultant air space gas partial pressures. Air space PO2 levels measured just before pipping ranged from 55 to 65 mm Hg in low and high conductance eggs, respectively, whilst PCO2 levels ranged from 75 to 55 mm Hg. With increasing shell conductance embryos membrane-penetrated and pipped the shell later, but hatch time was unaffected. Membrane penetration and external pipping appeared to occur in response to the partial pressures of gas in the air space but not in response to one particular level of air space PO2 or PCO2. During the parafoetal stage air space PO2 decreased at about 3.8 mm Hg/h and air space PCO2 increased at about 1.8 mm Hg/h and these rates of change were unaffected by eggshell conductance.

Animals↗

PubMed1993

Oxygen and avian eggshell formation at high altitude.

Many birds at high altitude lay eggs with reduced eggshell diffusive conductance to water vapor. Disagreement exists about the cause, but hypotheses include physiological acclimatization and genetic adaptation. To investigate this, we tested for the occurrence of physiological acclimatization and quantified the effect of relieving hypoxia at high altitude. Ten laying hens (Gallus domesticus) were exposed to elevations of 3800 m (PIO2 90 Torr), 3800 m with supplementary oxygen (PIO2 140 Torr), and 1200 m (PIO2 125 Torr). 573 eggs were collected and analyzed during the 17-week experiment. Shell conductance, aggregate pore area, and shell thickness were reduced at 3800 m compared to 1200 m or 3800 m with supplementary oxygen. There was a lag in the response to changes in altitude or PIO2 that corresponded to a time constant of approximately 2.5 weeks. We conclude that physiological acclimatization of eggshell conductance occurs in some chickens, and that it is probably stimulated by hypoxia.

Acclimatization↗

PubMed1991

Three-dimensional reconstruction of innermost chorion layer of Drosophila grimshawi and Drosophila melanogaster eggshell mutant fs(1)384.

A low-resolution three-dimensional structure of the crystalline innermost chorionic layer (ICL) of the Hawaiian species Drosophila grimshawi and the Drosophila melanogaster eggshell mutant fs(1)384 has been calculated from electron microscope images of tilted negatively stained specimens. The isolated ICL of Drosophila grimshawi is a three-layer structure, about 36 nm thick, whereas the ICL of Drosophila melanogaster eggshell mutant fs(1)384 is a single layer, about 12 nm thick. Each unit in both crystalline structures includes octamers made up of four heterodimers. Crosslinks between the structural elements, both within and between unit cells form an interconnecting network, apparently important in maintaining the integrity of the layer. A model which may account for the ICL self-assembly formation in vivo and the ICL observed lattice polymorphism is proposed, combining data from the three-dimensional reconstruction work and secondary structure features of the ICL component proteins s36 and s38.

Animals↗

PubMed1989

A preliminary study of the translocation of aldicarb across the duck eggshell.

The duck eggshell has the reputation of being more permeable than that of the domestic hen. If this is true, the developing embryo could be at greater risk from xenobiotic agents, since toxicants picked up on the feathers could be transferred to the embryo during incubation. This study looked for such an effect on the developing embryo after the application of aldicarb to the eggshell. At 72 hr, the eggs were painted with 3, 7, 11, or 15 microM aldicarb in 500 microliters water. The eggs were then incubated to Day 24. The gross morphological measurements were then recorded. A similar study was made using domestic hen eggs; these were treated after 36 hr incubation and incubated to Day 17. Direct injection into the yolk sac of both species was used for further comparison. There was a statistically significant reduction (P less than 0.01) in the middle web toe length with 11 and 15 microM aldicarb and the tarsometatarsus length with 7, 11, and 15 microM. Compared with the duck control group, the group given 15 microM aldicarb had reductions of approximately 8% in the tarsometatarsus and approximately 9% in the middle web toe. No statistically significant changes were produced in the chick embryos.

Aldicarb↗

PubMed1987

Possible eggshell protein gene from Schistosoma mansoni.

We have identified and sequenced a cDNA clone of a mRNA found only in mature female schistosomes. This mRNA is not detectably synthesized by female worms from single sex infections (unisexual females), by males or by the developing miracidia in the eggs. The clone hybridises to a highly abundant polyadenylated mRNA of approximately 1500 nucleotides. The nucleotide sequence of the clone predicts a polypeptide comprising two repetitive regions. A pentapeptide repeat with the consensus sequence Gly-Tyr-Asp-Lys-Tyr, and a region rich in histidine residues. Hybrid selected mRNA translated in vitro with [3H]tyrosine as labelled amino acid yields a polypeptide of 48 kDa (p48) that corresponds to the major [3H]tyrosine labelled translation product of female worm total mRNA. p48 does not label with [35S]methionine and is absent from the translation products of male and unisexual female mRNAs. The amino acid sequence of p48 has significant homologies to silk moth chorion proteins and we suggest that it is one of the major components of the schistosome eggshell probably accounting for the high level of [3H]tyrosine incorporation into the vitellaria of Schistosoma mansoni. The tyrosine content of the polypeptide suggests that it may play a role in phenol oxidase mediated cross-linking of the schistosome eggshell and in support of this we find that mushroom phenol oxidase will cause the specific cross-linking of p48 in in vitro translation products.

Animals↗

PubMed1988

Identification and localisation of the products of a putative eggshell precursor gene in the vitellarium of Schistosoma mansoni.

An abundant 0.9 kb female-specific mRNA in Schistosoma mansoni is thought to code for an egg-shell precursor protein [Bobek et al. (1986) Proc. Natl. Acad. Sci. USA 83, 5544-5548]. This gene contains two ORFs. A recombinant plasmid was constructed that expresses a fusion protein containing a glycine- and tyrosine-rich polypeptide coded for by one of these ORFs. Antisera raised against homogenates of female, but not of male, S. mansoni recognise this fusion protein, providing direct evidence that this ORF is used by S. mansoni. In comparative Western blots of S. mansoni homogenates from males and females affinity purified antibodies that react with the fusion protein react exclusively with proteins from females, recognising a 28 kDa polypeptide and a smear of immunoreactive material probably caused by oxidative crosslinking. In immunohistology, the affinity purified antibodies react with mature vitelline cells in female schistosomes. The immunoreactive material is localised in the so-called 'vitelline droplets' that are morphologically very similar to 'shell globules', known to contain egg-shell precursors, that are found in Fasciola hepatica. In situ hybridisation shows that the eggshell precursor gene is only transcribed in immature vitelline cells and has a short half-life. Taken together, these observations provide persuasive evidence that the 0.9 kb mRNA codes for an eggshell precursor.

Animals↗

PubMed1989

Identification, expression and in situ hybridization of an eggshell protein gene from Fasciola hepatica.

The molecular basis of egg formation in the parasitic liver fluke, Fasciola hepatica, was investigated by isolating and characterizing an abundant cDNA from a female genital complex cDNA library. It was expressed in Escherichia coli as a beta-galactosidase fusion protein, which was purified and used to produce polyclonal antibodies. Using immunoblots, the antiserum recognized two soluble constituents of isolated egg shells, both significantly larger than predicted from cDNA sequencing. Using in situ hybridization, the message was detected in cells in the adult vitelline follicles. Eggshell protein mRNA expressed in E. coli will provide a source of precursor protein for further studies of parasite eggshell formation.

Animals↗

PubMed1992

Eggshell precursor proteins of Fasciola hepatica, II. Microheterogeneity in vitelline protein B.

At least 3 structural protein precursors of the eggshell are synthesized and stockpiled in the extensive vitelline cells of the liver fluke Fasciola hepatica L. One of these, vitelline protein B, consists of a closely related family of proteins that owes its apparent electrophoretic heterogeneity to variations in the Tyr to DOPA conversion as well as to subtle variations in the primary sequence. The efficiency of the Tyr to DOPA conversion ranges from a maximum of about 90% to a minimum of 55% in the protein. Trypsin digestion in borate buffer at pH 8 was used to produce DOPA-peptides for sequencing. Notably, trypsin does not cleave Arg/Lys-DOPA sequences at borate concentrations greater than 0.15 M. Peptides with DOPA-containing sequences most frequently have flanking amino acids such as Lys, Ser, or Asp on the N-terminal side and Gly or Asp on the C-terminal side. All protein variants fall within a narrow molecular weight range (30-33 kDa), a pI range of 6.9 to 8.3, and the collective majority would appear to share a common N-terminal sequence up to residue 28. The results suggest some combination of the following: variations in post-translational hydroxylation, alternative post-transcriptional splicing and/or the existence of multiple gene copies of eggshell precursors. The latter have been shown to occur in the blood fluke Schistosoma mansoni [15].

Amino Acid Sequence↗

PubMed1984

Effects of suppression of eggshell calcification and of 1,25(OH)2D3 on Mg2+, Ca2+ and Mg2+HCO-3 ATPase, alkaline phosphatase, carbonic anhydrase and CaBP levels--I. The laying hen uterus.

Stimulation of Mg2+, Ca2+ and Mg2+HCO-3 dependent ATPase activity in mitochondrial and microsomal fractions from the uteri of laying hens is demonstrated. ATPase activity was greatest with 5 mM concentrations of Mg2+ at pH 8.5, and at pH 7.4-7.8 following the addition of bicarbonate. Suppression of eggshell calcification, induced by insertion of a thread into the uterus, did not alter Mg2+, Ca2+ and Mg2+HCO-3 ATPase activities. Alkaline phosphatase activity was generally low, and was unaffected by suppression of eggshell calcification. Levels of carbonic anhydrase and calcium binding protein were lower in the uteri of hens laying shell-less eggs. Injections of 1,25(OH)2D3 in hens laying shell-less eggs did not alter CaBP levels or enzyme activities. It is concluded that factors other than 1,25(OH)2D3 and gonadal hormones are involved in the regulation of uterine CaBP levels.

Adenosine Triphosphatases↗

PubMed1985

The effects of eggshell porosity on blood-gas and acid-base status of domestic fowl embryos within eggs of the same weight.

Eggshell porosity affects the metabolism and growth of the developing embryo and is likely, therefore, to influence blood-gas and acid-base status. PO2 in the airspace and in blood from the allantoic vein, PCO2 in the airspace and in blood from the allantoic artery and allantoic vein, and pH of blood from the allantoic artery and vein are all affected by shell porosity. Low porosity eggshells result in an increased retention of carbon dioxide within the egg, in partial compensation for which an increase in the level of bicarbonate in the blood is predicted. It is concluded that some of the wide variation in blood physiology data between eggs at the same stage of incubation, which has been recorded in the scientific literature, is the result of variation in shell porosity between eggs.

Acid-Base Equilibrium↗

PubMed2004

Soluble eggshell membrane protein: preparation, characterization and biocompatibility.

The preparation, characterization and biocompatibility of soluble eggshell membrane (SEP) are reported. The dissolution process, which is the key step of the preparation of SEP, has been followed by scanning electron microscopy (SEM) to observe the changes of the surfaces and thickness of the eggshell membrane (ESM). The composition of SEP has been investigated by amino acid analysis and elemental analysis. Based on the fact that SEP losses significantly cystine, and that SEP has a higher content of sulfur, an assumption involving combination with 3-mercaptopropionic acid (the reagent used for reductive cleavage of disulfide bonds) following the cleavage of the original disulfide bonds has been proposed, which explains the solubility of SEP. The thermal and surface properties have been studied by thermogravimetric analysis (TGA) and contact angle measurement. The biocompatibility of SEP, as demonstrated by cell culture of NIH3T3, is comparable to collagen type I and superior to raw ESM either inside or outside surface.

Animals↗

PubMed2006

Development and analytical application of an uric acid biosensor using an uricase-immobilized eggshell membrane.

An uric acid biosensor fabricated from a uricase-immobilized eggshell membrane and an oxygen electrode was presented. The detection schemes involve the enzymatic reactions of the uricase leading to the depletion of dissolved oxygen level upon exposure to uric acid solution. The decrease in oxygen level was monitored and related to the uric acid concentration. The scanning electron micrographs show the microstructure of the eggshell membrane within which the uricase is successfully immobilized. The effects of enzyme loading, pH, temperature, and phosphate buffer concentration on the response of the biosensor were investigated in detail. The uric acid biosensor has a linear response range of 4.0-640 microM with a detection limit of 2.0 microM (S/N=3). The response time was less than 100 s. The biosensor exhibited good repeatable response to a 0.10mM uric acid solution with a relative standard deviation of 3.1% (n=7). The reproducibility of fabrication of the biosensors using four different membranes was good with a R.S.D. of 3.2%. The biosensor showed extremely good stability with a shelf-life of at least 3 months. Some common potential interferents in samples such as glucose, urea, ascorbic acid, lactic acid, glycine, DL-alpha-alanine, DL-cysteine, KCl, NaCl, CaCl2, MgSO4, and NH4Cl showed no interferences on the response of the uric acid biosensor. The biosensor was successfully applied to determine the uric acid level in some human serum and urine samples, and the results agreed well with those obtained by a commercial colorimetric assay kit.

Animals↗

PubMed1999

Isolation of a glycosylated form of the chicken eggshell protein ovocleidin and determination of the glycosylation site. Alternative glycosylation/phosphorylation at an N-glycosylation sequon.

Ovocleidin, a major protein of the avian eggshell calcified layer, occurs in the eggshell soluble organic matrix in at least two forms. The major form is a phosphoprotein with two phosphorylated serines (OC-17) which was sequenced recently. A minor form is a glycosylated protein with identical sequence and only one phosphorylated serine (OC-23). The site of glycosylation is Asn(59), the only asparagine in the amino acid sequence contained in the N-glycosylation site consensus sequence, N-A-S. Ser(61), which is part of this site, is phosphorylated in OC-17 but not in OC-23 indicating that the two modifications are mutually exclusive. This is the first example of alternative glycosylation/phosphorylation occurring at an N-glycosylation site.

Animals↗

PubMed2000

Comparison of pericranium and eggshell as space fillers used in combination with guided bone regeneration: an experimental study.

PURPOSE: The purpose of the study was to determine the potential adjunctive role of 2 space fillers, used in combination with guided tissue regeneration, on bone regeneration in rat skull defects. MATERIALS AND METHODS: The study was conducted on 45 adult Wistar rats. A bilateral 6-mm-wide full-thickness skull defect was created in the parietal region of each animal. The right defect was chosen as the experiment site, and the left defect was left empty as a control. Each experiment site was covered by an inner and outer polytetrafluoroethylene membrane. The 45 rats were divided into 3 groups; no space filler between the 2 membranes was used in group I (n = 15), a free pericranial autograft was used as a space filler between the 2 membranes in group II (n = 15), and purified eggshell powder was used as space filler between the 2 membranes in group III (n = 15). Five animals in each group were killed at 15, 30, and 90 days. The harvested specimens were subjected to contact radiography and standard microscopic examination, and the rates of osteogenesis were assessed by a semiquantitative method. RESULTS: No evidence of bone regeneration was seen in any animals of the three groups at 15 days. At 30 days, bone regeneration only appeared in group I (P > .05). At 90 days, complete bone regeneration was observed in the group I in 3 of 5 animals (P < .05). In group II, the pericranial graft showed no osteogenic properties. In group III, the eggshell powder showed no resorption, but no osteoconduction was noticed. CONCLUSIONS: Although the osteogenic mechanism of guided tissue regeneration is not clear, this study suggests that the physical properties of the membrane are more important than the use of an adjunctive space filler.

Animals↗

PubMed2003

Species-specific activation of EGF receptor signaling underlies evolutionary diversity in the dorsal appendage number of the genus Drosophila eggshells.

In Drosophila melanogaster, the patterning of dorsal appendages on the eggshell is strictly controlled by EGFR signaling. However, the number of dorsal appendages is remarkably diverse among Drosophila species. For example, D. melanogaster and D. virilis have two and four dorsal appendages, respectively. Here we show that during oogenesis the expression patterns of rhomboid (rho) and argos (aos), positive and negative regulators of EGFR signaling, respectively, were substantially different between D. melanogaster and D. virilis. Importantly, the number and position of both the rho expression and MAPK activation were consistent with those of the dorsal appendages in each species. Despite the differences in the spatial expression, these results suggest that the function of EGFR signaling in dorsal appendage formation is largely conserved between these two species. Thus, our results link the species-specific activation of EGFR signaling and the evolution of eggshell morphology in Drosophila.

Acute-Phase Proteins↗

PubMed2003

The Drosophila embryonic patterning determinant torsolike is a component of the eggshell.

The development of the head and tail regions of the Drosophila embryo is dependent upon the localized polar activation of Torso (Tor), a receptor tyrosine kinase that is uniformly distributed in the membrane of the developing embryo. Trunk (Trk), the proposed ligand for Tor, is secreted as an inactive precursor into the perivitelline fluid that lies between the embryonic membrane and the vitelline membrane (VM), the inner layer of the eggshell. The spatial regulation of Trk processing is thought to be mediated by the secreted product of the torsolike (tsl) gene, which is expressed during oogenesis by a specialized population of follicle cells present at the two ends of the oocyte. We show here that Tsl protein is specifically localized to the polar regions of the VM in laid eggs. We further demonstrate that although Tsl can associate with nonpolar regions of the VM, the activity of polar-localized Tsl is enhanced, suggesting the existence of another spatially restricted factor acting in this pathway. The incorporation of Tsl into the VM provides a mechanism for the transfer of spatial information from the follicle cells to the developing embryo. To our knowledge, Tsl represents the first example of an embryonic patterning determinant that is a component of the eggshell.

Animals↗