Egg
An egg is an organic vessel that an animal grows to carry a fertilized egg cell, a zygote. Inside that vessel, an embryo is incubated until it can survive on its own. Then the animal hatches. The largest egg ever recorded came from a whale shark and measured 30 by 14 by 9 centimeters. The smallest known bird egg comes from the bee hummingbird and weighs half of a gram, roughly 0.02 ounces. Between those two extremes lies a vast catalogue of shapes, shells, and survival strategies. How does a single structure stretch across arthropods, fish, reptiles, birds, and even a handful of mammals? Why are some eggs jellylike and shell-less while others are armored in calcium carbonate? And how did this thing on breakfast menus worldwide also become a symbol of life, healing, and rebirth?
Most fish spawn eggs that are fertilized externally, the male inseminating them after the female lays them. Large fish can produce over 100 million eggs in a single spawning, then leave them to develop without parental care. The eggs of fish and amphibians, the anamniotes, are jellylike. They have no shell and would dry out in the air, which is why even air-breathing amphibians lay their eggs in water. The Coast foam-nest treefrog, Chiromantis xerampelina, instead deposits its eggs in protective foam. When larvae hatch, they often carry the remains of the yolk in a yolk sac that nourishes them for a few days as they learn to swim. Once the yolk is gone, a critical point arrives. They must learn to hunt and feed, or they die. Amniotes broke this dependence on water entirely. They are air-breathing vertebrates with complex eggs containing an amniotic membrane, the feature that gives the group Amniota its name. Forming such an egg requires internal conception, and the shell isolates the developing embryo from the mother. Reptile eggs are leathery for snakes and most lizards, while turtles have a calcareous shell. These shells survive in the air and absorb water from the environment, swelling as the fetus grows. For turtles, tuatara, and most lizards, the temperature of the surroundings determines the sex of the embryo. That single innovation, the shelled egg, let vertebrates carry their young onto dry land.
Bird eggs carry a hard shell of calcium carbonate bound with a 5 percent organic matrix. This surface prevents the contents from drying out, limits mechanical damage, and blocks microbes, all while letting gas pass through. Shell thickness ranges from paper thin up to 2.7 in ostriches, and the shell typically makes up 11 of an egg's weight. The variety in appearance is striking. Cormorant eggs are rough and chalky, tinamou eggs are shiny, duck eggs are oily and waterproof, cassowary eggs are heavily pitted, and jacana eggs look lacquered. Tiny pores let the embryo breathe, exchanging oxygen, carbon dioxide, and water with the outside world. A domestic hen's egg has around 7000 pores, and pore size runs inversely to the incubation period. Some shells carry a coating of vaterite spherules, a rare polymorph of calcium carbonate. In the Greater Ani, Crotophaga major, this vaterite coating is thought to act as a shock absorber, protecting the calcite shell from fracture when eggs collide in the nest. The shape varies too. The brown boobook lays a nearly spherical shell, the maleo egg is highly ellipsoidal, and the least sandpiper egg is far more conical. That shape forms as the egg moves through the final part of the oviduct. Cliff-nesting birds often lay highly conical eggs, which roll in a tight circle rather than off the ledge, a trait likely shaped by natural selection. Many hole-nesting birds, by contrast, lay nearly spherical eggs.
The default color of an avian egg is white, the color of the calcium carbonate the shell is made from. But some birds, mainly passerines, produce colored eggs. The color comes from pigments laid over that white base. Biliverdin and its zinc chelate, along with bilirubin, give a green or blue ground color, while protoporphyrin IX produces reds and browns as a ground color or as spotting. The same oviduct shell gland that builds the shell secretes these colors. Where soil holds too little calcium, the shell may thin, especially in a circle around the broad end. Protoporphyrin speckling compensates and increases as available calcium falls, and later eggs in a clutch are more spotted than early ones as the female's pigment glands deplete. Color carries a deeper genetic story. Within the common cuckoo lineage, the color of individual eggs appears inherited through the mother only, suggesting the pigment gene sits on the sex-determining W chromosome. For avian hosts of brood parasites, selection pushes the host to evolve distinctive colorations so foreign eggs can be spotted and rejected. The parasite then evolves better mimics, producing an evolutionary arms race between host and parasite. In the common guillemot, which nests in large groups, each female's eggs carry very different markings, making it easier to identify her own among the crowded cliff ledges.
Many animals feed on eggs. The principal predators of the black oystercatcher's eggs include raccoons, skunk, mink, river and sea otters, gulls, crows, and foxes. The stoat, Mustela erminea, and the long-tailed weasel, M. frenata, steal ducks' eggs. Snakes of the genera Dasypeltis and Elachistodon specialize entirely in eating eggs. Some birds turn other birds' nests into nurseries. Brood parasitism happens when one species lays its eggs in the nest of another, an uncommon behavior practiced by roughly 1 percent of bird species as obligate parasites. In some cases the host's own eggs are removed or eaten by the female, or expelled by her chick. The brood parasites include the cowbird, the black-headed duck, the cuckoo-finch, and three Old World cuckoo species. That parasitic pressure helps explain why a host species would invest so heavily in eggs that no rival could convincingly fake.
More than half the proteins in egg yolk are phosphoglycoproteins, equivalent to milk proteins in mammals or storage proteins in plant seeds. The yolk feeds the growing embryo, and its major precursor is the polypeptide vitellogenin, which occurs in all egg-laying animals and is synthesized in the liver. The insect form is called vitellin. Surrounding the yolk in the egg white is the albumen, a clear hydrophilic layer that serves as a water reservoir. Its predominant protein is ovalbumin, more than half the proteins by mass, though its role is unknown. Ovomucoid, making up 11 percent of the albumen, is the primary egg allergen. Scientists sort vertebrate eggs by how much yolk they carry. Eggs with little yolk are microlecithal, medium ones are mesolecithal, and large ones with a concentrated yolk are macrolecithal. In microlecithal eggs, the yolk is evenly spread, so cleavage divides the egg into cells of fairly similar size. These small eggs can be produced in huge numbers and are the norm where egg mortality is high, as in bivalves and marine arthropods. In placental mammals, where the mother nourishes the embryo throughout the fetal period, the egg holds little if any yolk. Macrolecithal eggs work differently. Their yolk is so large that cell division cannot split the mass, so the fetus develops as a plate on top of the yolk and only envelops it later. These big-yolked eggs appear only in cephalopods, cartilaginous fish, bony fish, reptiles, birds, and monotreme mammals. The eggs of the coelacanths can reach 9 centimeters across, and the young complete full development in the uterus, living on that copious yolk.
Platypuses and two genera of echidna, the spiny anteaters, are Australian monotremes, the only living order of egg-laying mammal. Their eggs are macrolecithal, very much like those of reptiles, and early extinct mammal species also laid eggs, probably the ancestral state. Marsupials lay no eggs, yet their eggs are still macrolecithal, merely small, developing inside the female without forming a placenta. Their young are born at a very early stage and can be classified as a larva in the biological sense. Placental mammals use two types of placenta, the yolk sac and the chorioallantoic. In humans, the first nutrient source is a yolk sac placenta, replaced by a chorioallantoic placenta at around four weeks. Around the eighth week, the yolk sac is absorbed into the umbilical cord, and the fetus finishes developing inside the uterus on nutrients from the mother. The French biologist Thierry Lodé proposed a scheme that divides reproduction by how much development happens before young leave the adult's body. It runs from ovuliparity, where the female spawns unfertilized eggs, through ovoviviparity, where the zygote stays in the body without feeding interactions, to hemotrophic viviparity, where a placenta channels nutrients from the mother's blood. In some sharks and in the black salamander Salamandra atra, embryos feed by consuming other ova or sibling embryos, a form of intra-uterine cannibalism.
By a wide margin, the egg most often eaten by people is the chicken egg, typically unfertilized, though duck, roe, and caviar are also popular choices. Under the Kashrut, the set of Jewish dietary laws, eggs are considered pareve, neither meat nor dairy, despite being an animal product, and may be mixed with milk or kosher meat. The chicken egg also became a tool of medicine. In 1931, Alice Miles Woodruff and Ernest William Goodpasture at Vanderbilt University discovered that the rickettsia and viruses behind many diseases will grow in chicken embryos. That finding enabled vaccines against influenza, chicken pox, smallpox, yellow fever, typhus, and Rocky mountain spotted fever, among others. Beyond food and medicine, eggs carry symbolic weight, often standing for life and rebirth, healing and protection, and sometimes appearing in creation myths. Christians view Easter eggs as symbolic of the resurrection of Jesus Christ. The same object also serves as a weapon of nuisance. Throwing raw eggs at houses, cars, or people, known as egging in English-speaking countries, is a minor form of vandalism that can damage vehicle paint and cause serious eye injury. Collecting wild birds' eggs was once a popular hobby, including among European Australians, with the embryo removed before the shell was stored. The practice is now banned in many jurisdictions because it can threaten rare species. In the United Kingdom it is prohibited by the Protection of Birds Act 1954 and the Wildlife and Countryside Act 1981. Those early collections survive as a record. The Australian Museum holds about 20,000 registered clutches, and scientists treat such collections as good natural-history data, because the collectors' notes have helped them understand how birds nest.
Common questions
What is an egg in biology?
An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell, a zygote. Inside the vessel an embryo is incubated until it can survive on its own, at which point the animal hatches. In other kingdoms, similar reproductive structures are termed spores, seeds, or egg cells.
What is the largest egg and the smallest egg?
The largest recorded egg came from a whale shark and measured 30 by 14 by 9 centimeters. The ostrich egg, at 1.5 kilograms and up to 17.8 by 14 centimeters, is the largest egg of any living bird, while the bee hummingbird lays the smallest known bird egg at half of a gram, around 0.02 ounces.
What are bird eggshells made of?
Bird eggshells are made of calcium carbonate with a 5 percent organic matrix. The shell prevents the contents from drying out, limits mechanical damage, and protects against microbes while allowing gas exchange. It can range from paper thin up to 2.7 in ostriches and typically forms 11 of the egg's weight.
Why are some bird eggs colored?
Egg color comes from pigments deposited over the white calcium carbonate base. Biliverdin and bilirubin give green or blue ground colors, while protoporphyrin IX produces reds and browns as ground color or spotting. Passerines mainly lay colored eggs, and protoporphyrin speckling also increases where soil calcium is low.
Which mammals lay eggs?
Platypuses and two genera of echidna, the spiny anteaters, are Australian monotremes and the only living order of egg-laying mammal. Their eggs are macrolecithal, very much like those of reptiles. Early extinct mammal species also laid eggs, which was probably the ancestral state.
How are eggs used to make vaccines?
Many vaccines for infectious diseases are produced in fertile chicken eggs. In 1931, Alice Miles Woodruff and Ernest William Goodpasture at Vanderbilt University found that the rickettsia and viruses behind many diseases will grow in chicken embryos. This enabled vaccines against influenza, chicken pox, smallpox, yellow fever, typhus, and Rocky mountain spotted fever.
Why is collecting wild bird eggs banned?
Collecting the eggs of wild birds is now banned by many jurisdictions because the practice can threaten rare species. In the United Kingdom it is prohibited by the Protection of Birds Act 1954 and the Wildlife and Countryside Act 1981. Early collections have since come to museums, where scientists treat the collectors' notes as good natural-history data.
All sources
112 references cited across the entry
- 1webWhale Shark – Cartilaginous FishSeaWorld Parks & Entertainment
- 2bookBiology of BirdsD.R. Khanna — Discovery Publishing House — 2005
- 3bookThe Biology of LampreysM. W. Hardisty et al. — Academic Press Inc. — 1971
- 4bookSharks of the World: An annotated and illustrated catalogue of shark species known to dateLeonard J. V. Compagno — Food and Agriculture Organization of the United Nations — 1984
- 5bookLivebearing FishesScott Peter — Tetra Press — 1997
- 6bookGeneral and Comparative Endocrinology: An Integrative ApproachA. M. Schreiber — CRC Press — 2023
- 7bookEcology of Marine FishJanet T. Duffy-Anderson et al. — Elsevier — 2024
- 8bookMeat Eaters: Raptors, Sharks, and CrocodilesThe Rosen Publishing Group, Inc — 2011
- 9bookFish ReproductionRobert A. Patzner — CRC Press — 2008
- 10bookRed Sea SharksJeremy Stafford-Deitsch — Trident Press Ltd — 1999
- 11journalMechanistic basis of adaptive maternal effects: egg jelly water balance mediates embryonic adaptation to acidity in Rana arvalisLongfei Shu et al. — November 2015
- 12bookThe Bare Bones: An Unconventional Evolutionary History of the SkeletonMatthew F. Bonnan — Indiana University Press — 2016
- 13bookThe Central Nervous System of Cartilaginous Fishes: Structure and Functional CorrelationsW. J. A. J. Smeets et al. — Springer Science & Business Media — 2012
- 14bookBiology and Ecology of FishesJames S. Diana et al. — John Wiley & Sons — 2023
- 15bookThe Ancestor's Tale: A Pilgrimage to the Dawn of EvolutionRichard Dawkins — Houghton Mifflin Harcourt — 2004
- 16bookAmniote Origins: Completing the Transition to LandMary J. Packard et al. — Elsevier — 1997
- 17bookReproduction in Mammals: The Female PerspectiveVirginia Hayssen et al. — JHU Press — 2017
- 18bookReptile Medicine and SurgeryDougls R. Mader — Elsevier Health Sciences — 2005
- 19bookDiseases and Pathology of ReptilesElliott R. Jacobson et al. — CRC Press — 2021
- 20journalUnderstanding the evolution of viviparity using intraspecific variation in reproductive mode and transitional forms of pregnancyCamilla M. Whittington et al. — June 2022
- 21journalThe first dinosaur egg was softMark A. Norell et al. — 2020
- 22journalReview of the monotreme fossil record and comparison of palaeontological and molecular dataA. M. Musser — December 2003
- 23bookThe Maryland Master Naturalist's HandbookJoy Shindler Rafey — JHU Press — 2025
- 24journalThe double-yolked egg: from the 'miracle of packaging' to nature's 'mistake'Attila Salamon et al. — 19 March 2020
- 25journalElongated shape and unusual eggshell microstructure enable first confirmed hatching of avian twinsKrzysztof Damaziak et al. — July 2025
- 26journalLife History of the Andean Condor in EcuadorJuan Sebastián Restrepo-Cardona et al. — 2024
- 27journalBreeding dispersal and demography of wild and hand-reared grey partridges Perdix perdix in FinlandAhti Putaala et al. — September 1998
- 28bookEgg: A Dozen OvaturesLizzie Stark — W. W. Norton & Company — 2023
- 29bookAdvances in Agriculture Research and ApplicationScholarlyEditions — 2012
- 30bookAncient and Modern Approaches to the Problem of Relativism: A Study of Husserl, Locke, and PlatoMatthew K. Davis — Springer Nature — 2023
- 31journalThe evolution of biomineralization in metazoansM. Debiais-Thibaud et al. — 2023
- 32bookOrnithologyFrank B. Gill — Macmillan — 2007
- 33bookBird-life: A Guide to the Study of Our Common BirdsFrank Michler Chapman — D. Appleton and Company=1919 — 1919
- 34bookEssentials of Avian Medicine and SurgeryJohn Wiley & Sons — 2008
- 35webThe Parts of the Egg
- 36journalA rare mineral, vaterite, acts as a shock absorber in the eggshell of a communally nesting birdJ. P. Portugal et al. — 2018
- 37journalAvian egg shape: Form, function, and evolutionMary Caswell Stoddard et al. — 23 June 2017
- 38newsWhy Are Bird Eggs Egg-Shaped? An EggsplainerEd Yong — 22 June 2017
- 39journalThe Mathematics of Egg ShapeYutaka Nishiyama — 2012
- 40newsThe chemistry of eggshell coloursCarol Rääbus — Australian Broadcasting Corporation — 18 February 2018
- 41journalTo Prevent Oxidative Stress, What about Protoporphyrin IX, Biliverdin, and Bilirubin?Ana Martínez et al. — 23 August 2023
- 42bookBirdpedia: A Brief Compendium of Avian LoreChristopher W. Leahy — Princeton University Press — 2021
- 43journalSurface texture heterogeneity in maculated bird eggshellsMarie R. G. Attard et al. — 12 July 2023
- 44journalSandgrouse: models of adaptive compromiseC. L. Maclean — 1985
- 45journalAll around the egg: consistency of spottiness and colouration across an avian eggshellPiotr Minias et al. — July 2024
- 46bookThe Nesting Season: Cuckoos, Cuckolds, and the Invention of MonogamyBernd Heinrich — Harvard University Press — 2010
- 47bookEgg Quality: Current Problems and Recent AdvancesS.E. Solomon — Butterworths — 1987
- 48journalWhy are birds' eggs speckled?Andrew G. Gosler et al. — 2005
- 49journalThe evolution of egg colour and patterning in birdsR. M. Kilner — 2006
- 50journalAncient origin and maternal inheritance of blue cuckoo eggsFrode Fossøy et al. — January 2016
- 51webWhy are eggs different colors?Paul Schattenberg — April 2025
- 52journalRobin's egg blue: does egg color influence male parental care?Philina A. English et al. — May 2011
- 53journalParents, predators, parasites, and the evolution of eggshell colour in open nesting birdsDaniel Hanley et al. — 2013
- 54journalColour, vision and coevolution in avian brood parasitismMary Caswell Stoddard et al. — 5 July 2017
- 55journalThe chemical basis of a signal of individual identity: shell pigment concentrations track the unique appearance of Common Murre eggsMark E. Hauber et al. — April 2019
- 56journalBlack Oystercatcher Habitat Selection, Reproductive Success, and Their Relationship with Glaucous-Winged GullsK. Vermeer et al. — 1992
- 57bookKodiak Island Wildlife: Biology and Behavior of the wild animals of Alaska's Emerald IsleRobin Barefield — Publication Consultants — 2021
- 58journalBlack Oystercatcher (Haematopus bachmani) Conservation Action PlanDavid F. Tessler et al. — February 2010
- 59journalPredation by ermine and long-tailed weasels on duck eggsJ. P. Fleskes — 1988
- 60journalSnake predators of bird eggs: a review and bibliographyJ. M. Barends et al. — 2022
- 61journalEgg Destruction and Egg Removal by Avian Brood Parasites: Adaptiveness and ConsequencesBrian D. Peer — Oxford University Press — January 2006
- 62bookEvolution of the Vertebrates – A History of Backboned Animals Through TimeH. E. Colbert et al. — John Wiley & Sons inc — 1991
- 63bookReproductive ToxicologyM. Basavarajappa et al. — CRC Press — 2016
- 64bookInvertebrate HistologyJohn Wiley & Sons — 2021
- 65bookSea Fishes Of The Mediterranean Including Marine InvertebratesLawson Wood — Bloomsbury Publishing — 2015
- 66bookArthropods-Are They Beneficial for Mankind?G. N. Susanto — Books on Demand — 2021
- 67bookAn Introduction to the InvertebratesJanet Moore — Cambridge University Press — 2006
- 68bookFather Nature: The Science of Paternal PotentialJames K. Rilling — MIT Press — 2024
- 69encyclopediaMale ReproductionBernard Jégou — Academic Press — 2018
- 70bookComparative Vertebrate ReproductionJulian Lombardi — Springer Science & Business Media — 2012
- 71web11.3: Sexual Reproduction18 December 2021
- 72bookAnimal Behaviour: An Evolutionary PerspectivePeter M. Kappeler — Springer Nature — 2022
- 73bookDevelopment and Reproduction in Humans and Animal Model SpeciesWerner A. Mueller et al. — Springer — 2015
- 74journalAnimal egg as evolutionary innovation: a solution to the 'embryonic hourglass' puzzleS. A. Newman — 2011
- 75journalThe evolution of egg yolk proteinsB. M. Byrne et al. — 1989
- 76bookAmniote Origins: Completing the Transition to LandElsevier — 1997
- 77bookBioactive egg compoundsV. Lechevalier et al. — Springer Berlin Heidelberg — 2007
- 78journalOvomucoid (the most important Egg white allergen) as a cause of severe Egg allergy: A reviewT. Ahmed et al. — 2024
- 79bookClinical Anatomy and Physiology for Veterinary TechniciansThomas P. Colville et al. — Elsevier Health Sciences — 2015
- 80bookUnderstanding ReproductionGiuseppe Fusco et al. — Cambridge University Press — 2023
- 81bookThe Vertebrate BodyA. S. Romer et al. — Saunders — 1985
- 82bookHuman Life Before BirthFrank Dye — CRC Press — 2000
- 83journalQualitative shift to indirect development in the parasitic sea anemone Edwardsiella lineataA. M. Reitzel et al. — 2006
- 84bookInvertebrate ZoologyR. D. Barns — W. B. Saunders Company — 1968
- 85bookMammalian DevelopmentManju Yadav — Discovery Publishing House — 2008
- 86bookAnalysis of Vertebrate StructureM. Hildebrand et al. — John Wiley & Sons, Inc — 2001
- 87journalHagfish developmentA. Gorbman — June 1997
- 88conferenceThe Biology of Cephalopods1977
- 89bookAmniote Origins-Completing the Transition to LandJ. R. Stewart — Academic Press — 1997
- 90journalEvidence for lecithotrophic viviparity in the living coelacanthH. W. Fricke et al. — October 1992
- 91journalOviparity or viviparity? That is the question…Lodé T. — 2012
- 92bookVertebrate endocrinologyDavid O. Norris et al. — Academic Press — 2013
- 93journalEvolution and morphogenesis of the placenta in sharksWilliam C. Hamlett — 1989
- 94journalMorphogenesis of extraembryonic membranes and placentation inMabuya mabouya (Squamata, Scincidae)Adriana Jerez et al. — November 2003
- 95bookVertebrate Endocrinology: Fundamentals and Biomedical ConsiderationsIvor M. D. Jackson — Elsevier — 2012
- 96webhisto-, hemo-Online Etymology Dictionary
- 97bookEggs in CookeryOxford Symposium — 2007
- 98bookMicrobial Food Safety: A Food Systems ApproachCharlene Wolf-Hall et al. — CABI — 2017
- 99bookThe Cambridge World History of FoodWilliam J. Stadelman — Cambridge University Press — 2000
- 100webJewish Dietary Laws (Kashrut): Overview of Laws & RegulationsJewish Virtual Library — 17 January 2013
- 101webHow Influenza (Flu) Vaccines Are MadeCDC — 30 September 2024
- 102journalThe Cultivation of Vaccine and Other Viruses in the Chorio-Allantoic Membrane of Chick EmbryosE. W. Goodpasture et al. — 9 Oct 1931
- 103bookA Shot in the Arm: How Science, Engineering, and Supply Chains Converged to Vaccinate the WorldYossi Sheffi — MIT CTL Media — 2021
- 104webThe Ancient Art of Decorating Eggs Folklife TodayStephanie Hall — 6 April 2017
- 105newsEaster Eggs: History, Origin, Symbolism And Traditions (PHOTOS)Jahnabi Barooah — 2 April 2012
- 106bookTreat or Trick? Halloween in a Globalising WorldJonas Frykman — Cambridge Scholars Publishing — 2008
- 107newsBeyond a yolk: a brief history of egging as a political protestChitra Ramaswamy — 5 October 2015
- 108webCollecting bird eggsAnita Mcinnes — 9 March 2017
- 110webEgg specimensMichael Sladek — 29 April 2009
- 112newsThe Lost Victorian Art of Egg CollectingKate Golembiewski — 25 March 2016