Structure and relationship of the mammillary gore to membrane fibres and initial calcification of the avian Egg Shell 1

1. The interior of the mammillary knob is shown to be composed of: (1) outer shell membrane fibres that penetrate the basal portion of the knob and become associated with (2) The mammillary core, and (3) the bulk calcification of the main knob.

2. The mammillary knob consists of radial crystals which are arranged in a hemisphere on the outer shell membrane fibres, and are composed of a central zone which is a homogeneous solid, crystalline in nature, and an outer heterogeneous vesiculated zone.

3. The walls of the mammillary knob are continuous with the base and sides of the mammillary core radial crystals; the dorso‐central zone of the core crystals are exposed to a cavity that communicates with the upper zones of the mammillary knob and palisade layer via the central channel. The dorsal surface of the core cavity is formed by a layer of unorganised crystals of the mammillary knob proper.

4. Initial calcification of the knob begins at the base and sides of the mammillary core and expands over the organic core as the walls of the knob thicken during subsequent calcification. Incomplete fusion of the central expansions from peripheral knob areas results in the central channel.     

Whitening of brown shelled eggs: Individual variation and relationships with age,fearfulness, oviposition interval and stress

1. Fearfulness, shell colour, incidence and degree of shell whitening and the interval between ovipositions were studied in two populations of 30 brown egg laying hens with family histories of a low or a high incidence of egg shell whitening.

2. Hens of the population with the high incidence of whitening appeared to be more fearful than hens of the population with the low incidence of whitening.

3. Brown colouration of the egg shell and the incidence and degree of shell whitening declined as the hens aged.

4. Brown colouration and egg shell whitening were most pronounced on the blunt ends of the eggs.

5. A large part of the variation in egg shell whitening was attributable to the individual (hen) component of variance.

6. Differences in egg shell whitening, between the two populations, were detectable throughout the 26 weeks of the experiment.

7. Oviposition intervals were similar for normal and coated eggs when birds were not exposed to disturbance.

8. Disturbance of hens increased oviposition intervals and the incidence and degree of shell whitening, to a similar extent, in both populations.

9. It is concluded that stress‐related egg retention is not the sole factor responsible for abnormal egg shell whitening. Shell whitening may occur as a consequence of the premature termination of shell pigmentation as well as a consequence of the retardation of oviposition which occurs when hens are disturbed.     

Genotypic differences in speed and magnitude of response to ahemeral lighting

1. The paper compares between three genetically different lines in their egg weight and shell thickness response to a 28‐h cycle.

2. The response in shell thickness to ahemeral lighting was more rapid (2.2 d) than for egg weight (4.2 d).

3. Differences between genotypes were evident in both the speed and magnitude of response.

4. Overall, egg weight and shell thickness on the 28‐h cycle were 7% and 8% greater than on 24 h.

5. The allometric increase indicates that the increase in shell weight was on average 14% more than the increase in egg weight.     

Effect of cuticle removal on the water vapour conductance of egg shells of several species of domestic bird

1. The effect of egg shell cuticle removal, using a 14.7 mM solution of sodium hypochlorite, on the water vapour conductance of egg shells was studied in 5 species of domestic bird.

2. The water vapour conductance of fowl and Muscovy duck egg shells was not affected by the treatment.

3. Cuticle removal increased the conductance of turkey and goose shells, particularly those already relatively porous to water vapour.

4. Removal of cuticles from domestic duck egg shells increased water vapour conductance. In this case the effect was most marked for eggs with lower initial conductances.

5. The differing patterns of resistance to water vapour loss associated with egg shell cuticle are related to the ultrastructure of the cuticle itself, especially the number and size of cuticular fissures.

6. The results suggest that egg shell cuticle is a component in a series of resistances associated with the egg shell and that the use of Fick's Law is insufficient to explain water vapour diffusion across the shell.

7. Embryo survival may be improved by cuticle removal, though this may be limited to those species in which the cuticle restricts water vapour loss in initially low conductance egg shells.     

Traits influencing the hatching performance of Japanese quail eggs

1. Japanese quail eggs from moderately heavier sires showed superior fertility; while fertile eggs from moderately heavier dams hatched slightly better than the eggs from lighter dams.

2. Higher rates of fertility and hatchability of Japanese quail eggs were observed from parents of 10 to 19 weeks of age, with peak fertility and hatchability at 14 and 12 weeks of age, respectively.

3. Sex ratios of 1:2 to 1:5 gave comparable fertility and hatchability results.

4. The hatching performance of quail eggs from cage and deep litter reared breeders was comparable.

5. Fertility and hatchability were directly proportional to the egg weight.

6. Quail egg shell colour, tints and blotches were found to influence hatching performance.

7. Storing quail eggs at 16 ± 2°C and 75 ± 5% relative humidity for more than 4 d reduced hatchability.

8. Hatchability of eggs stored at room temperature was improved if they were sealed in polyethylene bags.

9. Provision of light during the first 14 d of incubation resulted in a photo‐acceleration of about 3.2 h.     

The use of the plasma chemistry unit as an aid to the scanning electron microscope study of avian egg‐shell structure

1. In order to study the mammillary layer of the avian egg shell by scanning electron microscopy, it is necessary to separate the outer shell membrane from the calcified shell.

2. Chemical methods of effecting membrane removal are difficult to standardise due to variations in the strength of the membrane‐shell bond.

3. The use of reactive gas plasma provides an alternative, more efficient method for removing membranes without the risk of damage to underlying crystalline structures.     

Studies on the avian shell gland during egg formation: Aqueous and electrolytic composition of the mucosa 1

1. The intracellular composition of the avian shell gland mucosa was studied at six stages of egg formation.

2. Total water content of the mucosa before shell calcification was 4.8 kg/kg dry weight (dw) and rose to 6.6 kg/kg dw during shell formation; 60% of the increase was intracellular and 40% extracellular.

3. The potassium concentration of the mucosa remained constant during egg formation.

4. Chloride was removed constantly from the mucosal cells during egg formation.

5. A model was constructed for the glandular cells which depicts them as primarily NaHCO₃‐secreting cells towards the luminal side and HC1‐secreting cells towards the serosal side, their main function being to provide CO^2‐ ₃ for shell formation. In this model, the columnar mucosal cells are responsible for the calcium used in shell formation.     

Explanted embryo culture: In vitro and in ovo techniques for domestic fowl

1. Embryos of the domestic fowl (72 h old) have been explanted into shell‐less cultures or ‘surrogate’ eggshells, in order to investigate the possibility of rearing these embryos to hatching.

2. Rocking embryo cultures during the first half of incubation enhanced embryo growth.

3. Embryos explanted into ‘surrogate’ eggshells of either other individuals or other species have been successfully ‘hatched’.

4. A normal chorioallantois is formed in these surrogate eggshells. This enables a functional albumen sac to form and eggshell resorption to be achieved.

5. Embryos grown in ‘surrogate’ eggshells are slightly smaller than controls but otherwise normal.

6. The technique provides opportunities for genetic engineering experiments.     

Effects of short intermittent lighting on food consumption and performance of dwarf broiler breeders and progeny

1. Two hundred and sixty four dwarf broiler breeder hens were subjected to ad libitum or restricted feeding and to four lighting patterns: 15L (bright light):9D (dark), 15L:9d (dim light), 2L:10d: 1L:11d, (0.5L:3.5d) × 6.

2. Breeder hens fed ad libitum and subjected to either conventional or intermittent lighting ingested respectively, 25 g and 14 to 17 g more than hens restricted to 115 g/d.

3. Body weight was greater in hens fed ad libitum irrespective of the lighting pattern and of the amount of food intake.

4. Persistency of egg production was impaired by intermittent lighting.

5. Ad libitum feeding reduced egg fertility. The decrease was larger in breeder hens on intermittent lighting.

6. Hatchability was increased in hens submitted to the symmetrical lighting pattern (0.5L:3.5d).

7. Egg and chick weights were higher in hens fed ad libitum.

8. Shell index and shell breaking strength increased in restricted hens on the intermittent lighting pattern (0.5L:3.5d) × 6.

9. The best performance was obtained in restricted hens on the conventional lighting pattern.     

Effect of beta‐aminoproprionitrile on eggshell formation

1. Eggshells are bioceramic‐biopolymer composites made by a cell‐mediated deposition of an extracellular matrix which drives the organisation of the inorganic phase. Ultrastructurally, eggshells are composed of shell membranes, mammillary knobs, palisade, and cuticle. Shell membranes are two nets of type X collagen‐containing fibrils. On to these membranes, the mammillary knobs, that is, the crystal nucleation sites, are deposited. Type X collagen is highly cross‐linked and insoluble.

2. In order to evaluate the role of type X collagen cross‐linking on eggshell formation, hens were injected with different doses of β‐aminoproprionitrile, which specifically interferes with cross‐link formation.

3. Changes in egg size and shape were observed. Scanning electron micrographs analysis of these eggs demonstrated marked changes in crystal growth and shell membrane structure and arrangement. A dot‐blot analysis, using a monoclonal antibody against chicken type X collagen, shows a dose‐dependent increase in shell membrane collagen extractability.

4. It is concluded that the formation of β‐aminoproprionitrile‐sensitive cross‐links among the type X collagen molecules of the shell membranes play an essential role in normal eggshell formation.     

Egg‐shell deposition and blood plasma inorganic phosphorus concentration in individual laying hens

1. The relationship between plasma inorganic phosphorus measured at the end of the dark period and the weight of the next egg shell produced was investigated in 30 Hisex Brown hens.

2. Hens received an adequate conventional layers’ diet and the lighting pattern was conventional (17 L:7 D). Observations were made early (31 to 40 weeks) and late (62 to 68 weeks of age) in the laying period.

3. Eggs were classified by position in the clutch sequence and significant negative correlations were found between shell weights of early eggs in the sequence and plasma phosphate at the end of the dark period.

4. No significant trends were found in plasma total calcium.

5. No significant differences were found in bone compositions of birds producing consistently more or less than average shell weight during the laying period.

6. The negative correlations between plasma phosphate and shell weight are consistent with the observations of Sauveur and Mongin (1983) and show that impairment of shell deposition is associated with skeletal mobilisation as indicated by increase in plasma phosphate. This is consistent with the observations on bone composition and indicates that selection for shell quality will tend to select birds which are not dependent on excessive skeletal mobilisation during shell formation.     

Shell strength of eggs from medium‐bodied hybrid hens housed in cages or on range in outside pens

1. On average, over the laying year, a higher proportion of eggs were cracked from cages (4.6%) than from range (2.0%); the difference tended to increase as the flocks aged.

2. There was evidence that eggs from range had stronger shells: they required more energy to crack, shell thickness was greater and shell deformation at 70 weeks was less (though the difference was not significant).

3. The differences between systems in shell strength, although real, were small. It is probable that environmental insults make a much larger contribution than shell strength to differences in crack incidence.

4. Eggs laid by hens on range were larger (+ 6.8% at 36 weeks and + 2.9% at 70 weeks) than those laid by the same stock housed in cages.     

Behaviour of egg shell under compression in relation to deformation measurements1

1. The egg does not completely obey Hook's law when compressed between flat surfaces.

2. Deformation was not completely reversible due to crumbling at the points of contact with the force.

3. There are three components of non‐destructive deformation: (a) local deformation at the points of contact, (b) local surface crumbling and (c) deformation of the over‐all shell structure.

4. The latter two are uncontrolled, contributing to errors in the prediction of fracture force and depending on the non‐destructive force applied.

5. These effects should be considered in test selection and interpretation.

6. The optimum non‐destructive force was found to be 1 kg.     

Differences in egg‐shell quality among the Sinai Bedouin fowl,the commercial white leghorn and their crossbreds

1. A comparison of egg‐shell characteristics was made among the Sinai breed (a desert inhabiting strain), the commercial White Leghorn and their reciprocal crossbreds.

2. The Sinai egg was smaller and its shell thicker and stronger than the Leghorn egg. All the differences were statistically significant.

3. Shell thickness and strength of Leghorn eggs were in accordance with values predicted on the basis of egg mass, while those of the Sinai and the two crosses were considerably higher than predicted.

4. The apparent differences in mean egg mass between Sinai and Leghorn breeds did not explain the differences in egg‐shell quality, neither did the shape index which was very similar among all breeds.

5. The findings suggest that the differences in egg‐shell properties are of a genetic origin.

6. The thick, strong and less permeable egg‐shell of the Sinai breed may reflect adaptations to its arid environmental origin and to incubation in the open.

7. These genetically‐determined egg‐shell characteristics might serve in a future selection for improved egg‐shell quality in poultry.     

Effect of strain and age on the relationship of oviposition time to shell strength

1. The relationships between oviposition time and specific gravity, shell thickeness, deformation and breaking strength of eggs from six commercial layer strains were examined after 6 and 12 months of production.

2. Shell strength, assessed by any of the methods, was relatively low in eggs collected at 10.00 h and was generally successively greater in those collected at 12.00, 14.00 or 16.00 h. Over the same period egg weight decreased.

3. These time‐related changes in shell strength were similar for the two production ages despite the difference in shell quality due to age.

4. No consistent significant differences were observed between strains in time of oviposition.     

The action of sulphanilamide on shell deposition

1. The action of sulphanilamide on egg shell deposition has been studied in the domestic fowl. The length of time that eggs remain in the shell gland was measured; the relative thickness and protoporphyrin pigment concentration for egg shells removed from the shell gland were determined; egg shell structure was examined using a replica technique.

2. Sulphanilamide appeared to act mainly by reducing the rate of shell deposition, time spent in the shell gland was unaffected.

3. The thickness of shell components was reduced in a roughly proportional manner in thin shells produced during sulphanilamide treatment.

4. It is proposed that other thin shells in which the component layers are all reduced to a similar extent are also the result of a decreased deposition rate.     

Relationships between age,shell quality and individual rate and duration of shell formation in domestic hens

1. The duration and rate of shell formation was measured individually in each of 44 hens. Comparisons were made between hens of different ages, and also between hens which laid eggs of differing shell qualities.

2. Shell formation began 9–5 to 11 h after the oviposition of the previous egg, increased linearly for 13 h and then plateaued at 1–5 h before oviposition.

3. The variation in egg shell quality observed in two experimental groups of hens was 75% attributable to differences in the rate of shell deposition and 25% attributable to difference in the duration of shell deposition.

4. The mean interval between ovipositions increased with age but neither shell weight, nor rate or duration of shell depositon were affected. The increase in egg weight with age accounted for the decrease in shell quality.

5. Intervals between ovipositions were positively correlated with the duration of shell formation, especially its latter stages.

6. Egg production was negatively correlated with the interval between ovipositions but not with the duration or rate of shell deposition. There was a negative correlation between duration and rate of shell deposition, except in the case of the hens producing eggs of poor shell quality.     

Studies of membrane,mammilary cores and cuticle of the hen egg shell

Mammillary cores and outer shell membrane have been studied by comparing the weight and chemical composition of outer membrane with cores still attached and of outer membrane free from cores, from the same shells. Inner membrane and cuticle have also been investigated.

The weights of inner and outer membrane are significantly correlated (P < 0.05). Both membranes contain galactosamine, glucosamine, sialic acid, glucose, mannose and fucose but in different proportions. Uronic acid is probably absent.

An average of 3.3 mg of organic mammillary core material per shell (52 μg/cm²) was present in the shells investigated, but there was a trend suggesting that eggs laid by more mature hens may contain rather more. Cores are rich in hexo‐samine, sialic acid and hexose, but the probable absence of uronic acid suggests that chondroitin sulphates play no part in shell initiation. There were no major differences in the amino acid spectrum of membrane samples with and without cores.

Galactosamine, glucosamine, sialic acid, galactose, glucose, mannose, fucose and an unidentified pentose are present in the cuticle. In some respects these sugars are combined differently from their combination in the adjacent outer levels of the matrix.     

The hen's egg: Incidence and inheritance of some egg abnormalities

1. More than 9000 eggs, each laid by a known hen, were examined and malformations of six types recorded: double‐yolked, A, B, rough, shell‐less and thin‐shelled.

2. A and B malformations are those seen in, respectively, an egg that is held in the shell gland for an abnormally long period and one that reaches the shell gland while an A‐egg is still there; many of the eggs recorded as rough‐shelled were probably A‐eggs for which the corresponding B‐eggs had been lost.

3. The birds were of three strains: a medium‐body‐weight brown‐egg‐laying strain and a White Leghorn strain, both of commercial origin, and a laboratory strain of Brown Leghorns.

4. The incidences of egg malformation in the three strains were 8.3, 9.2 and 0.5%.

5. A‐, B‐ and rough‐shelled eggs constituted more than two‐thirds of all malformed eggs.

6. Evidence from two generations suggests that these malformations have a high heritability.     

The importance of convection in the determination of the water vapour conductance of avian egg shells

1. It was confirmed that convection across the avian egg shell surface does not affect diffusive transport of water vapour through the shell.

2. Relative water vapour conductance, i.e. the ratio between the water vapour conductance as determined in a desiccator and the actual water vapour conductance, as well as relative humidity in the desiccator are influenced by the total rate of water loss of the eggs in the desiccator.

3. The total rate of water loss is influenced by the number of eggs in the batch, their average actual water vapour conductance, desiccator temperature and barometric pressure.

4. Therefore water vapour conductance is easily underestimated, in particular when determined in still air at a small surface area of desiccant.

5. All the above effects are considerably reduced by convection.

6. Further improvements in technique are suggested, particularly for practice where determinations have to be made for a relatively large batch of eggs.