EGG (O.E. aeg, cf. Ger. Ei, Swed. aegg, and prob. Gr. , Lat. ovum), the female reproductive cell or ovum of animals, which gives rise generally only after fertilization to the young. The largest eggs are those of birds; and this because, to the minute essential portion of the egg, or germ, from which the young bird grows, there is added a large store of food-material - the yolk and white of the egg - destined to nourish the growing embryo while the whole is enclosed within a hard shell.
The relative sizes of eggs depend entirely on the amount of the food-yolk thus enclosed with the germ; while the form and texture of the outer envelope are determined by the nature of the environment to which the egg is exposed. Where the food material is infinitesimal in quantity the egg is either not extruded - the embryo being nourished by the maternal tissues, - or it passes out of the parental body and gives rise at once to a free-living organism or "larva" (see Larval Forms), as in the case of many lowly freshwater and marine animals. In such cases no "egg" in the usual sense of the term is produced.
The number of eggs periodically produced by any given individual depends on the risks of destruction to which they, and the young to which they give rise, are exposed: not more than a single egg being annually laid by some species, while with others the number may amount to millions.
Birds' Eggs. - The egg of the bird affords, for general purposes, the readiest example of the modifications imposed on eggs by the external environment. Since it must be incubated by the warmth of the parent's body, the outer envelope has taken the form of a hard shell for the protection of the growing chick from pressure, while the dyes which commonly colour the surface of this shell serve as a screen to hide it from egg-eating animals.
Carbonate of lime forms the principal constituent of this shell; but in addition phosphate of lime and magnesia are also present. In section, this shell will be found to be made up of three more or less distinct crystalline layers, traversed by vertical canals, whereby the shell is made porous so as to admit air to the developing chick.
The outermost, or third, layer of this shell often takes the form of a glaze, as of porcelain, as for example in the burnished egg of the ostrich: or it may assume the character of a thick, chalky layer as in some cuckoos (Guira, Crotophaga ani), cormorants, grebes and flamingoes: while in some birds as in the auks, gulls and tinamous, this outer layer is wanting; yet the tinamous have the most highly glazed eggs of all birds, the second layer of the shell developing a surface even more perfectly burnished than that formed by the outermost, third layer in the ostrich.
While the eggs of some birds have the shell so thin as to be translucent, e.g. kingfisher, others display considerable thickness, the maximum being reached in the egg of the extinct Aepyornis.
Though in shape differing but little from that of the familiar hen's egg, certain well-marked modifications of form are yet to be met with. Thus the eggs of the plover are pear-shaped, of the sand-grouse more or less cylindrical, of the owls and titmice spherical and of the grebes biconical.
In the matter of coloration the eggs of birds present a remarkable range. The pigments to which this coloration is due have been shown, by means of their absorption spectra (Sorby, Proc. Zool. Soc., 1875), to be seven in number. The first of these, oorhodeine, is brown-red in tone, and rarely absent: the second and third, oocyanin, and banded oocyanin, are of a beautiful blue, and though differing spectroscopically give rise to the same product when oxidized: the fourth and fifth are yellow, and rufous ooxanthine, the former combining with oocyanin gives rise to the wonderful malachite green of the emu's egg, while the latter occurs only in the eggs of tinamous: the sixth is lichenoxanthine, a pigment not yet thoroughly known but present in the shells of all eggs having a peculiar brick-red colour. Still less is known of the seventh pigment which is, as yet, nameless. It is a substance giving a banded absorption spectrum, and which, mixed with other pigments, imparts an abnormally browner tint. The origin of these pigments is yet uncertain, but it is probable that they are derived from the haemoglobin or red colouring matter of the blood. This being so, then the pigments of the egg-shell differ entirely in their nature from those which colour the yolk or the feathers.
While many eggs are either colourless or of one uniform tint, the majority have the surface broken up by spots or lines, or a combination of both, of varying tints: the pigment being deposited as the egg passes down the lower portion of the oviduct. That the egg during this passage turns slowly on its long axis is shown by the fact that the spots and lines have commonly a spiral direction; though some of the markings are made during periods of rest, as is shown by their sharp outlines, movement giving a blurred effect. Where the egg is pyriform, the large end makes way for the smaller. Many eggs display, in addition to the strongly marked spots, more or fewer fainter spots embedded in a deeper layer of the shell, and hence such eggs are said to be "double-spotted," e.g. rails and plovers.
Among some species, as in birds of prey, the intensity of this coloration is said to increase with age up to a certain point, when it as gradually decreases. Frequently, especially where but two eggs are laid (Newton), all the dye will be deposited, sometimes on the first, sometimes on the last laid, leaving the other colourless. But although of a number of eggs in a "clutch" - as the full complement of eggs in a nest is called - no two are exactly alike, they commonly bear a very close resemblance. Among certain species, however, which lay several eggs, one of the number invariably differs markedly from the rest, as for example in the eggs of the house-sparrow or in those of the sparrow-hawk, where, of a clutch of six, two generally differ conspicuously from the rest. Differing though these eggs do from the rest of the clutch, all yet present the characters common to the species. But the eggs of some birds, such as the Australian swamp quail, Synoecus australis, present a remarkably wide range of variation in the matter of coloration, no two clutches being alike, the extremes ranging from pure white to eggs having a greenish ground colour and rufous spots or blotches. But a still more interesting illustration of variation equally marked is furnished by the chikor partridge (Caccabis chukar), since here the variation appears to be correlated with the geographical distribution of the species. Thus eggs taken in Greece are for the most part cream-coloured and unspotted; those from the Grecian Archipelago are generally spotted and blotched; while more to the eastward spots are invariably present, and the blotches attain their maximum development.
But in variability the eggs of the guillemot (Lomvia troile) exceed all others: both in the hue of the ground colour and in the form of the superimposed markings, these eggs exhibit a wonderful range for which no adequate explanation has yet been given.
Individual peculiarities of coloration are commonly reproduced, not only with this species but also in others, year after year.
Significance of colour
The coloration of the egg bears no sort of relation to the coloration of the bird which lays it; but it bears on the other hand a more or less direct relation to the nature of the environment during incubation.
White eggs may generally be regarded as representing the primitive type of egg, since they agree in this particular with the eggs of reptiles. And it will generally be found that eggs of this hue are deposited in holes or in domed nests. So long indeed as nesting-places of this kind are used will the eggs be white. And this because coloured eggs would be invisible in dimly lighted chambers of this description, and therefore constantly exposed to the risk of being broken by the sitting bird, or rolling out of reach where the chamber was large enough to admit of this, whereas white eggs are visible so long as they can be reached by the faintest rays of light. Pigeons invariably lay white eggs; and while some deposit them in holes others build an open nest, a mere platform of sticks. These exceptions to the rule show that the depredations of egg-eating animals are sufficiently guarded against by the overhanging foliage, as well as by the great distance from the ground at which the nest is built. Birds which have reverted to the more ancient custom of nesting in holes after having developed pigmented eggs, have adopted the device of covering the shell with a layer of chalky matter (e.g. puffins), or, to put the case more correctly, they have been enabled to maintain survival after their return to the more ancient mode of nidification, because this reversion was accompanied by the tendency to cover the pigmented surface of the shell with this light-reflecting chalky incrustation.
Eggs which are deposited on the bare ground, or in other exposed situations, are usually protectively coloured: that is to say, the hue of the shell more or less completely harmonizes with the ground on which the egg is placed. The eggs of the plover tribe afford the most striking examples of this fact.
But the majority of birds deposit their eggs in a more or less elaborately constructed nest, and in such cases the egg, so far from being protectively coloured, often displays tints that would appear calculated rather to attract the attention of egg-stealing animals; bright blue or blue spotted with black being commonly met with. It may be, however, that coloration of this kind is less conspicuous than is generally supposed, but in any case the safety of the egg depends not so much on its coloration as on the character of the nest, which, where protective devices are necessary, must harmonize sufficiently with its surroundings to escape observation from prowling egg-stealers of all kinds.
The size of the egg depends partly on the number produced and partly on the conditions determining the state of the young bird at hatching: hence there is a great disparity in the relative sizes of the eggs of different birds. Thus it will be found that young birds which emerge in the world blind, naked and helpless are the product of relatively small eggs, while on the contrary young hatched from relatively large eggs are down-clad and active from birth.
The fact that the eggs must be brooded by the parent is also a controlling factor in so far as number is concerned, for no more can be hatched than can be covered by the sitting bird. Other factors, however, less understood, also exercise a controlling influence in this matter. Thus the ostrich lays from 12 to 16, the teal 15, the partridge 12-20, while among many other species the number is strictly limited, as in the case of the hornbills and guillemots, which lay but a single egg; the apteryx, divers, petrels and pigeons never lay more than 2, while the gulls and plovers never exceed 4. Tropical species are said to lay fewer eggs than their representatives in temperate regions, and further immature birds lay more and smaller eggs than when fully adult.
Partly owing to the uniformity of shape, size and texture of the shell, the eggs of birds are by no means easy to distinguish, except in so far as their family resemblances are concerned: that is to say, except in particular cases, they cannot be specifically distinguished, and hence they are of but little or no value for the purposes of classification.
Save only among the megapodes, all birds brood their eggs, the period of incubation varying from 13 days, as in small passerine birds, to 8 weeks, as in the cassowary, though eggs of the rhea and of Struthio hatch in from 5 to 6 weeks. But the megapodes deposit their eggs in mounds of decaying vegetable matter or in sand in the neighbourhood of hot springs, and there without further apparent care leave them. Where the nestling is active from the moment of hatching the eggs have a relatively longer incubation period than in cases where the nestlings are for a long while helpless.
Eggs of Mammals. - Only in the spiny ant-eater, or Echidna, and the duck-billed platypus, or Ornithorhynchus, among the Mammalia, are the eggs provided with a large store of yolk, enclosed within a shell, and extruded to undergo development apart from the maternal tissues. In the case of the echidna the eggs, two in number, are about as large as those of a sparrow, similar in shape, and have a white, parchment-like shell. After expulsion they are transferred by the beak of the mother to a pouch resembling that of the marsupial kangaroos, and there they undergo development. The Ornithorhynchus, on the other hand, lays from two to four eggs, which in size and general appearance resemble those of the echidna. They are, however, deposited in a loosely constructed nest at the end of a long burrow and there brooded. In Marsupials, the eggs are smaller than those of Echidna and Ornithorhynchus, and they contain a larger proportion of yolk than occurs in higher mammals.
Eggs of Reptiles. - The eggs of reptiles are invariably provided with a large amount of food yolk and enclosed with a firm test or shell, which though generally parchment-like in texture may be calcareous as in birds, as, for example, in many of the tortoises and turtles and in the crocodiles.
Among reptiles the egg is always white or yellowish, while the number laid often far exceeds that in the case of birds. The tuatara of New Zealand, however, lays but ten - white hard-shelled, long and oval - at intervals between November and January. The long intervals between the appearance of the successive eggs is a characteristic feature of the reptiles, but is met with among the birds only in the megapodes, which, like the reptiles, do not "brood" their eggs.
Among the Chelonia the number of eggs varies from two to four in some of the tortoises, to 200 in some of the turtles: while in the crocodiles between 20 and 30 are produced, hard-shelled and white.
The eggs of the lizards are always white or yellowish, and generally soft-shelled; but the geckos and the green lizard lay hard-shelled eggs. Many of the soft-shelled eggs are remarkable for the fact that they increase in size after extrusion, owing to the stretching of the membranous shell by the growing embryo. In the matter of number lizards are less prolific than many of the Chelonia, a dozen eggs being the general number, though as many as thirty may be produced at a time, as in the case of the common chameleon.
While as a general rule the eggs of lizards are laid in burrows or buried, some are retained within the body of the parent until the young are ready to emerge; or they may even hatch within the oviduct. This occurs with some chameleons and some lizards, e.g. the slow-worm. The common English lizard is also viviparous. Normally the young leaves the egg immediately after its extrusion, but if by any chance this extrusion is delayed they escape while yet in the oviduct.
The majority of the snakes lay eggs, but most of the vipers and the aquatic snakes are viviparous, as also are a few terrestrial species. The shell of the egg is always soft and parchment-like. As a rule the number of eggs produced among the snakes is not large, twenty or thirty being common, but some species of python lay as many as a hundred. Generally, among the oviparous snakes the eggs are buried, but some species of boas jealously guard them, enclosing them within the coils of the body.
Eggs of Amphibia. - Among the amphibia a greater variety obtains in the matter of the investment of the egg, as well as in the number, size and method of their disposal. The outer covering is formed by a toughening of the surface of a thick gelatinous coat which surrounds the essential parts of the egg. This coat in many species of salamander - using this name in the wide sense - is produced into threads which serve either to anchor the eggs singly or to bind them together in bunches.
Viviparity occurs both among the limbless and the tailed Amphibia, the eggs hatching before they leave the oviduct or immediately after extrusion. The number of young so produced is generally not large, but the common salamander (Salamandra maculosa) may produce as many as fifty at a birth, though fifteen is the more normal figure. When the higher number is reached the young are relatively small and weak.
As a rule among the Amphibia the young leave the egg in the form of larvae, generally known as "tadpoles"; but many species produce eggs containing a sufficient amount of food material to enable the whole of the larval phase to be completed before hatching.
Among the tailless Amphibia (frogs and toads) there are wide differences in the number of eggs produced, while the methods by which these eggs are disposed of present a marvellous variety.
As a rule vast quantities of eggs are shed by the female into the water in the form of "spawn." In the common toad as many as 7000 eggs may be extruded at a time. These leave the body in the form of two long strings - one from each oviduct - of translucent globules, gelatinous in texture, and enclosing a central Sphere of yolk, the upper pole of which is black. The spawn of the common frog differs from that of the toad in that the eggs all adhere to form a huge jelly-like mass. But in many species the number of eggs produced are few; and these may be sufficiently stored with food-yolk to allow of the tadpole stage being passed before hatching, as in frogs of the genus Hylodes. In many cases the eggs are deposited out of the water and often in quite remarkable ways.
Eggs of Fishes. - The eggs of fishes present an extremely wide range of form, and a no less extensive range in the matter of number. Both among the cartilaginous and bony fishes viviparity occurs. Most of the sharks and rays are viviparous, but in the oviparous species the eggs present some interesting and peculiar forms. Large in size, the outer coat or "shell" is in all cases horn-like and flexible, but differs greatly in shape. Thus in the egg of the larger spotted dog-fish it is oblong in shape, flattened from side to side, and has the angles produced into long, slender tendrils. As the egg is laid the lower tendrils project from the vent, and the mother rubs herself against some fixed body. The tendrils soon catch fast in some slight projection, when the egg is dragged forth there to remain till hatching takes place. A couple of narrow slits at each corner of the upper end serve to admit fresh water to the imprisoned embryo during the later stages of development; when development is complete escape is made through the end of the shell. In the rays or "skates," long spines take the place of tendrils, the egg simply resting at the bottom of the sea. The empty egg-cases of the rays are often found on the seashore, and are known as "Mermaids' purses." The egg of the Port Jackson shark (Cestracion) is of enormous size, pear-shaped, and provided with a spiral flange extending along the whole length of the capsule. In the Chimaera the egg is long, more or less spindle-shaped, and produced on each side into a broad flange having a fringed edge, so that the whole bears a close resemblance to a long leaf, broad and notched at one end, pointed at the other. This likeness to the seaweed among which it rests is doubtless a protective device, akin to that of protectively coloured birds' eggs.
Among the bony fishes the eggs generally take the form of small spheres, enclosed within a tough membrane or capsule. But they present many important differences, being in some fishes heavy and remaining at the bottom of the water, in other light and floating on the surface. While in some species they are distributed separately, in others they adhere together in masses. The eggs of the salmon, for example, are heavy, hard and smooth, and deposited separately in a trough dug by the parent and afterwards covered to prevent them from being carried away by the stream. In the perch they are adhesive and form long band-like masses of spawn adhering to water-plants. In the gobies the egg is spindle-shaped, and attached by one end by means of a network of fibres, resembling rootlets; while in the smelt the egg is loosely suspended by a membrane formed by the peeling off of a part of the outer sheath of the capsule. The eggs of the garfish (Belone vulgaris) and of the flying-fish of the genus Exocoetus, attach themselves to foreign objects, or to one another, by means of threads or cords developed at opposite poles of the egg.
Among a number of fishes the eggs float at the surface of the sea, often in enormous masses, when they are carried about at the mercy of tides and currents. An idea of the size which such masses attain may be gathered from the fact that the spawn of the angler-fish, Lophius piscatorius, takes the form of a sheet from 2 to 3 ft. wide, and 30 ft. long. Another remarkable feature of these floating eggs is their transparency, inasmuch as they are extremely difficult to see, and hence they probably escape the rapacious maws of spawn-eating animals. The cod tribe and flat-fishes lay floating eggs of this description.
The maximum number of eggs laid by fishes varies greatly, some species laying relatively few, others an enormous number. But in all cases the number increases with the weight and age of the fish. Thus it has been calculated that the number laid by the salmon is roughly about 1000 to every pound weight of the fish, a 15 lb salmon laying 15,000 eggs. The sturgeon lays about 7,000,000; the herring 50,000; the turbot 14,311,000; the sole 134,000; the perch 280,000. Briefly, the number is greatest where the risks of destruction are greatest.
The eggs of the degenerate fishes known as the lampreys and hag-fishes are remarkable for the fact that in the latter they are large in size, cylindrical in shape, and provided at each end with hooklets whereby they adhere one to another; while in the lampreys they are extremely small and embedded in a jelly.
Molluscs. - Among the Mollusca, Crustacea and Insecta yolk-stored eggs of very remarkable forms are commonly produced.
In variety, in this connexion, the Mollusca must perhaps be given the first place. This diversity, indeed, is strikingly illustrated by the eggs of the Cephalopoda. In the squids (Loligo), for example, the eggs are enclosed in long cylindrical cases, of which there are several hundreds, attached by one end to a common centre; the whole series looking strangely like a rough mop-head. Each case, in such a cluster, contains about 250 eggs, or about 40,000 in all. By way of contrast the eggs of the true cuttle-fish (Sepia) are deposited separately, each enclosed in a tough, black, pear-shaped capsule which is fastened by a stalk to fronds of sea-weed or other object. They appear to be extruded at short intervals, till the full complement is laid, the whole forming a cluster looking like a bunch of grapes. The octopus differs yet again in this matter, its eggs being very small, berry-like, and attached to a stalk which runs through the centre of the mass.
The eggs of the univalve Mollusca are hardly less varied in the shapes they take. In the common British Purpura lapillus they resemble delicate pink grains of rice set on stalks; in Busycon they are disk-shaped, and attached to a band nearly 3 ft. long. The eggs of the shell-bearing slugs (Testacella) are large, and have the outer coat so elastic that if dropped on a stone floor they will rebound several inches; while some of the snails (Bulimus) lay eggs having a white calcareous and slightly iridescent shell, in size and shape closely resembling the egg of the pigeon. Some are even larger than the egg of the wood-pigeon. The beautiful violet-snail (Ianthina) - a marine species - carries its eggs on the under side of a gelatinous raft. No less remarkable are the eggs of the whelk; since, like those of the squids, they are not laid separately but enveloped in capsules, and these to the number of many hundreds form the large, ball-like masses so commonly met with on the seashore. When the eggs in these capsules hatch, the crowd of embryos proceed to establish an internecine warfare, devouring one another till only the strongest survives!
With the Mollusca, as with other groups of animals, where the eggs are exposed to great risks they are small, produced in great numbers, and give rise to larvae. This is well illustrated by the common oyster which annually disperses about 60,000,000 eggs. But where the risk of destruction is slight, the eggs are large and produce young differing from the parent only in size, as in the case of the pigeon-like eggs of Bulimus.
Crustaceans. - Among the higher Crustacea, as a rule, the eggs are carried by the female, attached to special appendages on the under side of the body. But in some - Squillas - they are deposited in burrows. Generally they are relatively small so that the young which emerge therefrom differ markedly in appearance from the parents, but in deep-sea and freshwater species the eggs are large, when the young, on emerging, differ but little from the adults in appearance.
Insects, etc. - The eggs of insects though minute, are also remarkable for the great variety of form which they present, while they are frequently objects of great beauty owing to the sculptured markings of the shell. They are generally laid in clusters, either on the ground, on the leaves of plants, or in the water. Some of the gnats (Culex) lay them on the water. Cylindrical in shape they are packed closely together, set on end, the whole mass forming a kind of floating raft. Frequently, as in the case of the stick and leaf insect, the eggs are enclosed in capsules of very elaborate shapes and highly ornamented.
As to the rest of the Invertebrata - above the Protozoa the eggs are laid in water, or in damp places. In the former case they are as a rule small, and give rise to larvae; while eggs hatched on land are sometimes enclosed in capsules, "cocoons," as in the case of the earthworm, where this capsule is filled with a milky white fluid, of a highly nutritious character, on which the embryos feed.
Among some invertebrates two different kinds of eggs are laid by the same individual. The water-flea, Daphnia (a crustacean), lays two kinds of eggs known as "summer" and "winter" eggs. The summer eggs are carried by the female in a "brood-pouch" on the back. The "winter" eggs, produced at the approach of winter, differ markedly in appearance from the summer eggs, being larger, darker in colour, thicker shelled, and enclosed in a capsule formed from the shell or carapace, of the parent's body. "Winter eggs," however, may be produced in the height of summer. While the "summer eggs" are unfertilized, the winter eggs are fertilized by the male, and possess the remarkable power of lying dormant for months or even years before they develop. The production of these two kinds of eggs is a device to overcome the cold of winter, or the drying up of the pools in which the species lives, during the heat of the summer. The power of resistance which such eggs possess may be seen in the fact that a sample of mud which had been kept dry for ten years still contained living eggs. In deep water where neither drought nor winter cold can seriously affect the Daphnias, they propagate all the year round by unfertilized "summer" eggs.
Bibliography. - For further details on this subject the following authors should be consulted: - Mammals: F. E. Beddard, "Remarks on the Ovary of Echidna," Proc. Roy. Phys. Soc. Edin. vol. viii. (1885); W. H. Caldwell, "The Embryology of Monotremata and Marsupialia," Phil. Trans. Roy. Soc. vol. 178 (1887); E. B. Poulton, "The Structures connected with the Ovarian Ovum of the Marsupialia and Monotremata," Quart. Journ. Micros. Sci. vol. xxiv. (1884). Birds, Systematic: - H. Seebohm, Coloured Figures of the Eggs of British Birds (1896); A. Newton, Ootheca Wooleyana (1907); E. Oates, Cat. Birds' Eggs Brit. Mus. (appearing), vols. i.-iv. published. General: - A. Newton, Dictionary of Birds (1896). Colouring matter: - Newbegin, Colour in Nature (1898). Reptiles and Amphibia: - H. Gadow, "Reptiles," Camb. Nat. Hist. (1901); G. A. Boulenger, "The Tailless Batrachians of Europe," Ray Soc. (1896). Fishes: - Bridge and Boulenger, "Fishes, Ascidians, etc.," Camb. Nat. Hist. (1904); B. Dean, Fishes Living and Fossil (1895); J. T. Cunningham, Marketable Marine Fishes (1896). Invertebrate: - G. H. Carpenter, Insects. Their Structure and Life (1899); L. C. Miall, A History of Aquatic Insects (1895); T. R. R. Stebbing, Crustacea, Internat. Sci. series (1893); M. C. Cooke, "Mollusca," Camb. Nat. Hist. (1906). For further references to the above and other Invertebrate groups see various text-books on Entomology, Zoology.
(W. P. P.)
Note - this article incorporates content from Encyclopaedia Britannica, Eleventh Edition, (1910-1911)