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Many tropical bird species have restricted elevational distributions, potentially limited by how environmental conditions affect physiological processes. While some studies have examined adult physiology across elevations, relatively little attention has been given to the structure and function of eggshells despite their critical role in regulating gas exchange during the vulnerable embryonic stage. At high elevations, dry air is expected to increase water loss from the egg, and natural selection may favor lower gas conductance to reduce desiccation risk. Structural variation in eggshells, such as increased shell thickness or reduced pore size and density, could serve as a mechanism to regulate gas diffusion. To test for adaptive variation in eggshell traits along elevational gradients, we measured water vapor conductance and used scanning electron microscopy (SEM) to examine eggshell structure in 197 bird species from the Andes. We found that water vapor conductance declined at high elevations across avian communities. However, structural changes in eggshells varied among bird families and did not vary in a predictable way with elevation, suggesting no relationship or divergent adaptive responses to shared selective pressures, particularly in shell thickness, pore density, and pore size. We propose that examining functional and structural eggshell traits can offer insight into species’ elevational limits and inform predictions about their responses to climate change. Along mountain slopes, changes in humidity, air pressure, and temperature can affect physiological processes and constrain where species can live. As a result, plants and animals are often adapted to the specific conditions of their environments. These patterns have been widely studied across organisms, including birds, where researchers have examined metabolic rates and insulating features such as adult plumage. However, less is known about an even more vulnerable stage of life – when individuals cannot move, and development has only just begun. During embryonic development within an egg, a bird depends on a delicate balance with the external environment. Even small environmental changes can disrupt this balance by altering gas exchange through microscopic pores in the eggshell. Oxygen must diffuse inward, while carbon dioxide and water vapor diffuse outward. At high elevations, maintaining this balance becomes more challenging because dry air increases the risk of excessive water loss. Studying eggshell adaptations can provide refined insights into how environmental conditions during the nesting period may constrain species distributions. Ocampo et al. asked whether birds in tropical mountains cope with environmental challenges during nesting by adjusting gas exchange through the eggshell, particularly by reducing water loss at high elevations. They then investigated the mechanisms underlying these adjustments by examining variation in eggshell microstructure across many species, including traits such as thickness and porosity. First, the researchers documented several nests and eggs not previously described by science, particularly from species inhabiting remote regions of the Amazon and the Andes. By examining egg physiology along elevational gradients, they found that high-elevation species lose water at slower rates than lowland species, likely as an adaptation to reduce the risk of desiccation. These findings support the idea that egg physiology may constrain species distributions. They also identified microstructural variation in eggshells across lineages, although further work is needed to clarify the patterns and their functional significance. Overall, the study of Ocampo et al. highlights the importance of considering all life stages when evaluating physiological responses to climate change. Studies of species’ natural history remain essential and, when combined with hypothesis-driven evolutionary ecology, provide a powerful framework for understanding species distributions and improving predictions of their responses to environmental change. These findings may inform ecologists and physiologists, as well as conservation practitioners, land managers, policymakers, and researchers developing bioinspired materials. Realizing these benefits will require translating findings into predictive models and management strategies, alongside further advancing our understanding of eggshell structure.

The higher availability of zinc (Zn) from organic than inorganic sources is already established, but more assertive and cost-friendly protocols on the total replacement of inorganic with organic Zn sources for laying hens still need to be developed. Because some discrepancy in the effects of this replacement in laying hen diets is noticeable in the literature, the objective of this meta-analysis was to properly quantify the effect size of total replacing inorganic Zn with organic Zn in the diet of laying hens on their laying performance, egg quality, and Zn excretion. A total of 2340 results were retrieved from Pubmed, Scielo, Scopus, WOS, and Science Direct databases. Of these, 18 primary studies met all the eligibility criteria and were included in this meta-analysis. Overall, the replacement of inorganic Zn with organic Zn, regardless of other factors, improved ( p < 0.01) egg production by 1.46%, eggshell thickness by 0.01 mm, and eggshell resistance by 0.11 kgf/cm 2 . Positive results of the same nutritional strategy on egg weight and Zn excretion were only observed at specific conditions, especially when organic Zn was supplemented alone in the feed, not combined with other organic minerals. Therefore, there is evidence in the literature that the total replacement of inorganic Zn with organic Zn improves egg production, eggshell thickness, and eggshell resistance. Factors such as hen age and genetics, organic Zn source, concentration of Zn in the feed, and the strategy of its supplementation have to be more carefully considered in protocols designed to address egg weight and Zn excretion by the hen.

Background Egg-laying performance is economically important in poultry breeding programs. Crossbreeding between indigenous and elite commercial lines to exploit heterosis has been an upward trend in traditional layer breeding for niche markets. The objective of this study was to analyse the genetic background and to estimate the heterosis of longitudinal egg-laying traits in reciprocal crosses between an indigenous Beijing-You and an elite commercial White Leghorn layer line. Egg weights were measured for the first three eggs, monthly from 28 to 76 weeks of age, and at 86 and 100 weeks of age. Egg quality traits were measured at 32, 54, 72, 86, and 100 weeks of age. Egg production traits were measured from the start of lay until 43, 72, and 100 weeks of age. Heritabilities and phenotypic and genetic correlations were estimated. Heterosis was estimated as the percentage difference of performance of a crossbred from that of the parental average. Reciprocal cross differences were estimated as the difference between the reciprocal crossbreds as a percentage of the parental average. Results Estimates of heritability of egg weights ranged from 0.29 to 0.75. Estimates of genetic correlations between egg weights at different ages ranged from 0.72 to 1.00. Estimates of heritability for cumulative egg numbers until 43, 72, and 100 weeks of age were around 0.15. Estimates of heterosis for egg weight and cumulative egg number increased with age, ranging from 1.0 to 9.0% and from 1.4 to 11.6%, respectively. From 72 to 100 weeks of age, crossbreds produced more eggs per week than the superior parent White Leghorn (3.5 eggs for White Leghorn, 3.8 and 3.9 eggs for crossbreds). Heterosis for eggshell thickness ranged from 2.7 to 6.6% when using Beijing-You as the sire breed. No significant difference between reciprocal crosses was observed for the investigated traits, except for eggshell strength at 54 weeks of age. Conclusions The heterosis was substantial for egg weight and cumulative egg number, and increased with age, suggesting that non-additive genetic effects are important in crossbreds between the indigenous and elite breeds. Generally, the crossbreds performed similar to or even outperformed the commercial White Leghorns for egg production persistency.

Cleidoic eggs possess very efficient and orchestrated systems to protect the embryo from external microbes until hatch. The cuticle is a proteinaceous layer on the shell surface in many bird and some reptile species. An intact cuticle forms a pore plug to occlude respiratory pores and is an effective physical and chemical barrier against microbial penetration. The interior of the egg is assumed to be normally sterile, while the outer eggshell cuticle hosts microbes. The diversity of the eggshell microbiome is derived from both maternal microbiota and those of the nesting environment. The surface characteristics of the egg, outer moisture layer and the presence of antimicrobial molecules composing the cuticle dictate constituents of the microbial communities on the eggshell surface. The avian cuticle affects eggshell wettability, water vapor conductance and regulates ultraviolet reflectance in various ground-nesting species; moreover, its composition, thickness and degree of coverage are dependent on species, hen age, and physiological stressors. Studies in domestic avian species have demonstrated that changes in the cuticle affect the food safety of eggs with respect to the risk of contamination by bacterial pathogens such as Salmonella and Escherichia coli . Moreover, preventing contamination of internal egg components is crucial to optimize hatching success in bird species. In chickens there is moderate heritability (38%) of cuticle deposition with a potential for genetic improvement. However, much less is known about other bird or reptile cuticles. This review synthesizes current knowledge of eggshell cuticle and provides insight into its evolution in the clade reptilia. The origin, composition and regulation of the eggshell microbiome and the potential function of the cuticle as the first barrier of egg defense are discussed in detail. We evaluate how changes in the cuticle affect the food safety of table eggs and vertical transmission of pathogens in the production chain with respect to the risk of contamination. Thus, this review provides insight into the physiological and microbiological characteristics of eggshell cuticle in relation to its protective function (innate immunity) in egg-laying birds and reptiles.

The purpose of this study was to determine the effects of the dietary addition of zinc (Zn) in the form of sulphate (Zn-S), glycine (Zn-G), and hydroxychloride (Zn-H) on quail performance, eggshell quality, antioxidant status, mineral excretion, biomechanical properties and mineralization of bone. A total of 75 female quails (10-weeks-old) were randomly distributed into 3 groups with 5 replications, each with 5 female quails. Treatment groups were set up by adding Zn-S and Zn-H as the inorganic form and Zn-G as the organic form of zinc to the corn-soybean basal diet (34.14 mg/kg Zn) to obtain 50 mg/kg Zn and the feeding experiment lasted for 12 weeks. Performance parameters and egg production were not impaired by the Zn source ( P  > 0.05). The inclusion of Zn-S in the diet produced a reduction in eggshell thickness, while an improvement in yolk antioxidant capacity (measured as MDA concentration) was detected compared to the other Zn sources ( P  < 0.05). Shear strength and cortical bone area increased ( P  < 0.05) with Zn-G supplementation, however, the mineral concentration of the tibia was higher ( P  < 0.01) in those quails who had received Zn-H. Lower levels of mineral excretion were observed in both types of supplementations compared to Zn-S. Therefore, it can be stated that Zn-G or Zn-H supplementation in the diet of laying quails could be an interesting strategy to reduce mineral excretion and improve bone mineralization without affecting performance compared to Zn-S. However, further studies are needed to determine the differences between Zn-G and Zn-H.

The Gekkota is an important clade in the evolution of calcified eggshells in that some of its families lay rigid eggshells like archosaurs. However, the fundamental differences and similarities between the mechanism of rigid eggshell formation of the Gekkota and Archosauria have not been investigated thoroughly due to the lack of knowledge of gekkotan eggshells. Here, we report for the first time a comprehensive analysis of morphological, chemical compositional, and crystallographic features of rigid and soft gekkotan eggshells. Exhaustive morphological description provided common characters for gekkotan eggshells, as well as unique features of each species. We found that elemental distribution of rigid gekkotan eggshells is different from that of avian eggshells, especially in the case of Mg and P. In addition, the crystallographic features (size, shape, and alignment of calcite grains) of gekkotan eggshells are completely different from those of archosaur eggshells. The result of this study suggests that soft gekkotan eggshells are morphologically more similar to tuatara eggshells rather than soft eggshells of derived squamates. The chemical compositional analysis suggests that the eggshell may act as a mineral reservoir for P and F as well as Ca. More importantly, all chemical compositions and crystallographic features imply that the gekkotan eggshell formation may begin at the outer surface and growing down to the inner surface, which is opposite to the direction of the archosaur eggshell formation. This character would be crucial for identifying fossil gekkotan eggs, which are poorly known in paleontology. All these lines of evidence support that soft gekkotan and tuatara eggshells share the primitive characters of all lepidosaurid eggshells. Finally, gekkotan and archosaur rigid eggshells represent a typical example of convergent evolution in the lineage of the Sauropsida.

Calcified eggshells protect developing embryos against environmental stress and contribute to reproductive success 1 . As modern crocodilians and birds lay hard-shelled eggs, this eggshell type has been inferred for non-avian dinosaurs. Known dinosaur eggshells are characterized by an innermost membrane, an overlying protein matrix containing calcite, and an outermost waxy cuticle 2 – 7 . The calcitic eggshell consists of one or more ultrastructural layers that differ markedly among the three major dinosaur clades, as do the configurations of respiratory pores. So far, only hadrosaurid, a few sauropodomorph and tetanuran eggshells have been discovered; the paucity of the fossil record and the lack of intermediate eggshell types challenge efforts to homologize eggshell structures across all dinosaurs 8 – 18 . Here we present mineralogical, organochemical and ultrastructural evidence for an originally non-biomineralized, soft-shelled nature of exceptionally preserved ornithischian Protoceratops and basal sauropodomorph Mussaurus eggs. Statistical evaluation of in situ Raman spectra obtained for a representative set of hard- and soft-shelled, fossil and extant diapsid eggshells clusters the originally organic but secondarily phosphatized Protoceratops and the organic Mussaurus eggshells with soft, non-biomineralized eggshells. Histology corroborates the organic composition of these soft-shelled dinosaur eggs, revealing a stratified arrangement resembling turtle soft eggshell. Through an ancestral-state reconstruction of composition and ultrastructure, we compare eggshells from Protoceratops and Mussaurus with those from other diapsids, revealing that the first dinosaur egg was soft-shelled. The calcified, hard-shelled dinosaur egg evolved independently at least three times throughout the Mesozoic era, explaining the bias towards eggshells of derived dinosaurs in the fossil record. Molecular analyses of newly discovered, embryo-bearing ornithischian and sauropod dinosaur eggs suggest that the ancestral dinosaur egg was soft-shelled, and that hard-shelled eggs evolved independently at least three times in the major dinosaur lineages.

A meta-analysis was conducted to determine the effects of different types of oligosaccharides (OS) on the egg quality, intestinal profile, nutrient digestibility, and immunity of laying hens. A dataset was constructed from articles published between 2006 and 2023 via a literature search using several keywords related to oligosaccharides and laying hens' performance. After careful evaluation, the final dataset was developed from 25 in vivo studies comprising 165 comparisons. The meta-regression showed that levels of OS supplementation curvilinearly increased (p < 0.05) hen day egg production (HDEP), and crude protein digestibility (CPD). However, OS supplementation did not affect (p > 0.05) feed intake (FI), feed conversion ratio (FCR), egg mass, egg weight, eggshell thickness, shell strength, and Haugh unit. OS supplementation showed a positively improved antioxidant activity, as indicated by the curvilinear effect on superoxide dismutase (SOD) and malondialdehyde (MDA) concentration. In the categorical meta-analysis, mannan oligosaccharide (MOS), xylooligosaccharides (XOS), and chitosan oligosaccharide (COS) significantly increased (p < 0.01) HDEP and CPD. MOS was the only type of OS that decreased (p < 0.05) FCR while FOS showed to increase (p < 0.01) Haugh unit. To sum up, oligosaccharides could generally increase HDEP but MOS is superior to improving HDEP and FCR while FOS is better than other OS to increase eggshell thickness. HIGHLIGHTS Oligosaccharides increase egg production and protein digestibility in laying hens. Mannan oligosaccharide improves feed conversion ratio in laying hens. Elevated trends of IgG and IgA after supplementation with oligosaccharides. Fructo-oligosaccharides improve eggshell thickness and suppress Malondialdehyde (MDA) in laying hens.

This study aimed to determine the mechanism by which H9N2 avian influenza virus (AIV) affects eggshell quality. Thirty-week-old specific pathogen free egg-laying hens were inoculated with the chicken-origin H9N2 AIV strain (A/Chicken/shaanxi/01/2011) or with inoculating media without virus by combined intraocular and intranasal routes. The time course for the appearance of viral antigen and tissue lesions in the oviduct was coincident with the adverse changes in egg production in the infected hens. The viral loads of AIV have a close correlation with the changes in the uterus CaBP-D28k mRNA expression as well as the Ca concentrations in the eggshells in the infected hens from 1 to 7 days post inoculation (dpi). Ultrastructural examination of eggshells showed significantly decreased shell thickness in the infected hens from 1 to 5 dpi ( P  < 0.05). Furthermore, obvious changes in the structure of the external shell surface and shell membrane were detected in the infected hens from 1 to 5 dpi as compared with the control hens. In conclusion, this study confirmed that H9N2 AIV strain (A/Chicken/shaanxi/01/2011) infection is associated with severe lesions of the uterus and abnormal expression of CaBP-D28k mRNA in the uteri of the infected hens. The change of CaBP-D28k mRNA expression may contribute to the deterioration of the eggshell quality of the laying hens infected with AIV. It is noteworthy that the pathogenicity of H9N2 AIV strains may vary depending on the virus strain and host preference.

In this study, we compared the antibacterial effectivity of the eggs of six precocial and four altricial bird species using Escherichia coli, based on their eggshell traits. The ultrastructure of eggshell was observed using a scanning electron microscope (SEM). According to SEM results, eggs from precocial birds (chicken, turkey, quail, duck, ostrich, and goose) had cuticle on the eggshells, while eggs from altricial birds (pigeon, budgerigar, munia, and canary) did not. The environment/selection pressure may induce the divergent evolution process in eggs of precocial and altricial birds. The E. coli experiment results showed that chicken, turkey, quail, duck, and goose eggs, with a high cuticle opacity, exhibited a much lower E. coli penetration rate. In contrast, the eggs with poor (ostrich) or without (pigeon, budgerigar, munia, and canary) cuticle exhibited a higher penetration rate. It is suggested that cuticle is a main barrier against bacterial penetration in precocial birds' eggs. Turkey and quail eggs showed the lowest E. coli contamination rate (3.33% and 2.22%, respectively), probably because of the tightly connected nanosphere structure on their cuticle. As for altricial birds' eggs, the eggs of budgerigar, munia, and canary with small pore diameter (0.57 to 1.22 μm) had a lower E. coli penetration rate than pigeon eggs (45.56%, 66.67%, 50%, and 97.78%, respectively, P < 0.05), indicating that pore diameter played a significant role in defending against bacterial trans-shell invasion. We found that eggshell thickness and pore area decreased with egg size. The cuticle quality had no relationship with egg size, but was closely related to the bird species. The E. coli penetration rate of altricial birds' eggs was significantly higher than that of precocial birds' eggs, mainly because the pores are exposed on the eggshell surface and cuticle protection is absent. This study provides detailed information on the eggshell cuticle, which gives insight into the cuticle evolution process that occurred in precocial and altricial bird species. Moreover, the results of E. coli penetration may help understanding the antibacterial behavior in birds.