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Nature's technical ceramic: the avian eggshell.

ABSTRACT: Avian eggshells may break easily when impacted at a localized point; however, they exhibit impressive resistance when subjected to a well-distributed compressive load. For example, a common demonstration of material strength is firmly squeezing a chicken egg along its major axis between one's hands without breaking it. This research provides insight into the underlying mechanics by evaluating both macroscopic and microstructural features. Eggs of different size, varying from quail (30 mm) to ostrich (150 mm), are investigated. Compression experiments were conducted along the major axis of the egg using force-distributing rubber cushions between steel plates and the egg. The force at failure increases with egg size, reaching loads upwards of 5000 N for ostrich eggs. The corresponding strength, however, decreases with increasing shell thickness (intimately related to egg size); this is rationalized by a micro-defects model. Failure occurs by axial splitting parallel to the loading direction-the result of hoop tensile stresses due to the applied compressive load. Finite-element analysis is successfully employed to correlate the applied compressive force to tensile breaking strength for the eggs, and the influence of geometric ratio and microstructural heterogeneities on the shell's strength and fracture toughness is established.

SUBMITTER: Hahn EN

PROVIDER: S-EPMC5310731 | biostudies-literature | 2017 Jan

REPOSITORIES: biostudies-literature

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Nature's technical ceramic: the avian eggshell.

Hahn Eric N EN Sherman Vincent R VR Pissarenko Andrei A Rohrbach Samuel D SD Fernandes Daniel J DJ Meyers Marc A MA

Journal of the Royal Society, Interface 20170101 126

Avian eggshells may break easily when impacted at a localized point; however, they exhibit impressive resistance when subjected to a well-distributed compressive load. For example, a common demonstration of material strength is firmly squeezing a chicken egg along its major axis between one's hands without breaking it. This research provides insight into the underlying mechanics by evaluating both macroscopic and microstructural features. Eggs of different size, varying from quail (30 mm) to ost ...[more]

PMID: 28123095

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Project description:BACKGROUND: The exceptional diversity of coloration found in avian eggshells has long fascinated biologists and inspired a broad range of adaptive hypotheses to explain its evolution. Three main impediments to understanding the variability of eggshell appearance are: (1) the reliable quantification of the variation in eggshell colours; (2) its perception by birds themselves, and (3) its relation to avian phylogeny. Here we use an extensive museum collection to address these problems directly, and to test how diversity in eggshell coloration is distributed among different phylogenetic levels of the class Aves. METHODOLOGY AND RESULTS: Spectrophotometric data on eggshell coloration were collected from a taxonomically representative sample of 251 bird species to determine the change in reflectance across different wavelengths and the taxonomic level where the variation resides. As many hypotheses for the evolution of eggshell coloration assume that egg colours provide a communication signal for an avian receiver, we also modelled reflectance spectra of shell coloration for the avian visual system. We found that a majority of species have eggs with similar background colour (long wavelengths) but that striking differences are just as likely to occur between congeners as between members of different families. The region of greatest variability in eggshell colour among closely related species coincided with the medium-wavelength sensitive region around 500 nm. CONCLUSIONS: The majority of bird species share similar background eggshell colours, while the greatest variability among species aligns with differences along a red-brown to blue axis that most likely corresponds with variation in the presence and concentration of two tetrapyrrole pigments responsible for eggshell coloration. Additionally, our results confirm previous findings of temporal changes in museum collections, and this will be of particular concern for studies testing intraspecific hypotheses relating temporal patterns to adaptation of eggshell colour. We suggest that future studies investigating the phylogenetic association between the composition and concentration of eggshell pigments, and between the evolutionary drivers and functional impacts of eggshell colour variability will be most rewarding.

Project description:Under the shell of a chicken egg are two opposed proteinaceous disulfide-rich membranes. They are fabricated in the avian oviduct using fibers formed from proteins that are extensively coupled by irreversible lysine-derived crosslinks. The intractability of these eggshell membranes (ESM) has slowed their characterization and their protein composition remains uncertain. In this work, reductive alkylation of ESM followed by proteolytic digestion led to the identification of a cysteine rich ESM protein (abbreviated CREMP) that was similar to spore coat protein SP75 from cellular slime molds. Analysis of the cysteine repeats in partial sequences of CREMP reveals runs of remarkably repetitive patterns. Module a contains a C-X(4)-C-X(5)-C-X(8)-C-X(6) pattern (where X represents intervening non-cysteine residues). These inter-cysteine amino acid residues are also strikingly conserved. The evolutionarily-related module b has the same cysteine spacing as a, but has 11 amino acid residues at its C-terminus. Different stretches of CREMP sequences in chicken genomic DNA fragments show diverse repeat patterns: e.g. all a modules; an alternation of a-b modules; or an a-b-b arrangement. Comparable CREMP proteins are found in contigs of the zebra finch (Taeniopygia guttata) and in the oviparous green anole lizard (Anolis carolinensis). In all these cases the long runs of highly conserved modular repeats have evidently led to difficulties in the assembly of full length DNA sequences. Hence the number, and the amino acid lengths, of CREMP proteins are currently unknown. A 118 amino acid fragment (representing an a-b-a-b pattern) from a chicken oviduct EST library expressed in Escherichia coli is a well folded, highly anisotropic, protein with a large chemical shift dispersion in 2D solution NMR spectra. Structure is completely lost on reduction of the 8 disulfide bonds of this protein fragment. Finally, solid state NMR spectra suggest a surprising degree of order in intact ESM fibers.

Dataset Information

Nature's technical ceramic: the avian eggshell.

Publications

Nature's technical ceramic: the avian eggshell.

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